Wildlife of madagascar 2016 by Angel C

WILDGuides

Published by Princeton University Press, 41 William Street, Princeton, New Jersey 08540 In the United Kingdom: Princeton University Press, 6 Oxford Street, Woodstock, Oxfordshire OX20 1TR nathist.press.princeton.edu Requests for permission to reproduce material from this work should be sent to Permissions, Princeton University Press First published 2016 Copyright ÂŠ 2016 Princeton University Press Copyright in the photographs remains with the individual photographers. All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording, or otherwise, without the prior permission of the publishers. British Library Cataloging-in-Publication Data is available Library of Congress Control Number 2016930334 ISBN 978-0-691-16171-6 Production and design by WILDGuides Ltd., Old Basing, Hampshire UK. Printed in China

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Ken: To my wife Keith: For my Dad and Mom, who taught me to love nature. And to my wife Yi-fang and son Joshua, who allow me the time to enjoy it.

Contents A brief introduction to Madagascar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 About this book . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Map of Madagascar showing biogeographic zones and most visited natural sites . . . . . . 10 Biogeographic zones of Madagascar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 How to use this book . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 MAMMALS

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CHEIROGALEIDAE: Mouse lemurs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 Dwarf lemurs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 Giant mouse and fork-marked lemurs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 LEPILEMURIDAE: Sportive lemurs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 LEMURIDAE: Bamboo lemurs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 Ring-tailed Lemur . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 Northern brown lemurs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46 Widespread brown lemurs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48 Distinctive Eulemur species of the north and west . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50 Ruffed lemurs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52 Woolly lemurs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54 INDRIIDAE: Western sifakas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56 Eastern sifakas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58 Local sifakas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60 Indri . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62 DAUBENTONIIDAE: Aye-aye . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64 EUPLERIDAE: Malagasy carnivorans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66 Fosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68 CHIROPTERA: Bats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70 TENRECIDAE: Tenrecs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74 NESOMYIDAE: Rodents . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78 CETACEA: Marine mammals

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BIRDS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82 Ducks and grebes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84 Gamebirds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86 Herons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88 White egrets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92 Compact herons and Hamerkop . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94 Larger raptors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98 Smaller raptors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100 Shy birds of the forest floor . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102 Rails and jacana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104 Plovers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106 Smaller, plainer sandpipers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110 Larger, more distinctive sandpipers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112 Endemic pigeons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114 Aquatic oddities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114 Terns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116 Birds of the sea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118 Doves and pigeons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120 Endemic pigeons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122 Parrots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124 Cuckoos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 126 Couas of the dry west & southwest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 128 Couas of the wet east & north . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130 Owls . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132 Nightjars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134 Aerial feeders . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136 Kingfishers and bee-eater . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138 Hoopoe and roller . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140 More common rainforest ground-rollers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142 Scarcer rainforest ground-rollers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144 Long-tailed Ground-roller and Cuckoo-roller . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146 Asities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148 Common open-country birds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150 Widespread songbirds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152 Robin-like birds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154 Warblers that live close to the ground . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 156 Forest canopy warblers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160 Bernierids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162 Sunbirds and white-eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 164 Common and widespread vangas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166 Typical vangas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168 Strange vangas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 170 Large vangas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 172 Dark-headed vangas and cuckoo-shrike . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 174 Mid-sized dark birds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176 Weavers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178

REPTILES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180 Tortoises and terrapins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182 Sea turtles and crocodile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184 Tiny leaf chameleons of the north . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 186 Widespread medium to large leaf chameleons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 188 The largest chameleons on earth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190 Common chameleons of degraded areas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 192 Eastern rainforest chameleons: small . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 194 Eastern rainforest chameleons: medium to large . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196 Chameleons of the north . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 200 Chameleons of the west . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 204 Swifts I: west and southwest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 208 Swifts II: rock-dwelling swifts of the southwest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210 Plated lizards I: species without strong stripes on back . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212 Plated lizards II: species with strong stripes on back . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 214 Skinks I . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216 Widespread day geckos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220 Day geckos of the north . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222 Day geckos of the west, south and centre . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 224 Dwarf geckos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 226 Small nocturnal geckos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 228 Large nocturnal geckos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 230 Leaf-tailed geckos I: leaf-like and dry bark-like species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232 Leaf-tailed geckos II: mossy bark-like species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 234 Malagasy boas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 238 Distinctive snakes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 240 Common and widespread snakes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 242 Rainforest snakes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 244 FROGS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 246 Ridged frog and reed frogs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 248 Distinctive terrestrial microhylids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 250 Arboreal microhylids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 252 Mantellas I: species with orange . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 254 Mantellas II: green-and-black species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 256 Small to medium bright-eyed frogs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 258 Medium to large bright-eyed frogs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 262 Various arboreal mantellids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 266 Various terrestrial mantellids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 268 Bridge frogs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 270 Madagascar frogs: plain species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 274 Madagascar frogs: more distinctive species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 278

INVERTEBRATES

Invertebrates: distinctive species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 282 Butterflies: swallowtails . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 286 Butterflies: pierids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 292 Butterflies: metalmarks and lycanids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 396 Butterflies: orange nymphalids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 398 Butterflies: various nymphalids (emperors, commodore and pansy) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 304 Butterflies: pansies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 306 Butterflies: black-and-white nymphalids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 308 Butterflies: satyrs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 312 Butterflies: skippers, and moths . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 314 PLANTS

Iconic plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 316 Baobabs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 320

Glossary of terms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Further reading and useful resources . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acknowledgements and photo credits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

324 327 328 332

A brief introduction to Madagascar Madagascar is so different from the rest of the world that it is sometimes called ‘The Eighth Continent’. Not only does it have the high level of endemism (species not found elsewhere) that is typical of an island, but it also boasts remarkable diversity, which for some groups approaches that more typical of a whole continent. Madagascar is the land of lemurs, a radiation of our own primate order that evolved into exhilarating diversity on this island. These endearing creatures are the ambassadors for Malagasy nature; many people who have no idea where this island is located immediately recognize the Ring-tailed Lemur. Madagascar is also a land of fabulous birds, ancient reptile lineages, and six of the world’s nine species of baobabs. All naturalists find Madagascar fascinating as a treasure-trove of biodiversity and a ‘laboratory of evolution’, much like the Galápagos but on a grander scale. And for travelling naturalists, Madagascar is a ‘must-visit’ place. Although Madagascar has long been known for birds and mammals, its reptiles, amphibians, insects and plants are just as unique.

Madagascar is a land of distinctive and odd creatures, a whole different track of evolution from the rest of the world. A good example of this is the Mossy Leaf-tailed Gecko, a master of disguise whose mossy fringes and colour changing abilities allow it to blend into the tree trunks where it sleeps during the day.

About this book This is the first field guide that has attempted to cover the whole range of diversity of this ‘mini-continent’ in one book. It obviously cannot be comprehensive, but rather aims to cover the species and groups that are most likely to be encountered in the most frequently visited sites. Madagascar’s most popular natural sites are listed on page 10, ranked by the approximate number of visitors each receives every year. As an example of the way that species selection for this book was weighted towards the most frequently visited sites, an especially common or interesting species that is found only at the island’s most popular natural site of Andasibe might be included, whereas a species that is found only in Marojejy National Park is more likely to be excluded since that site is much less visited, albeit wonderful. A few exceptionally interesting species that can be found only in sites that are off the main tourist routes have been included in the hope of inspiring people to visit new sites. Also included are some especially interesting species that do occur within the most-visited sites, but which are hard to find. These are included to inspire visitors to seek them out with the help of skilled guides. Focused birders will locate a few species not covered in this book; keen mammal enthusiasts might visit remote forests and find lemurs that, similarly, have not been included; and fanatical herpetologists who spend their nights stalking the forest and digging in the leaflitter will find many reptiles and amphibians that are not featured. However, this book aims to cover the vast majority of the birds, mammals, reptiles and butterflies (plus a few other arthropods and plants) that a casual visitor or a general naturalist will see. Visitors with a strong interest in one particular group will want to bring a thorough reference that covers it, but will still find this guide valuable for its broad coverage of other wildlife. Finally, naturalists who might never travel to Madagascar should find this guide a fascinating exhibition of the island’s wild riches. To some degree, this book reflects the mammal and bird bias of the average visitor, covering those groups more thoroughly than others. But at the same time, visitors are encouraged to broaden their horizons. It is hoped that the extensive coverage of Madagascar’s truly incredible reptiles, amphibians and insects (butterflies in particular) will make these seem more accessible. One of the countless odd and unique facets of Madagascar is that you are required to hire a local guide to visit almost any natural site. While this might be frustrating for some ‘lone wolf ’ naturalists, it also has great benefits. Most Malagasy local guides are passionate and knowledgeable, and their expertise will enhance your trip and increase the number of species that you see. The information in the ‘where to see’ section will be useful in planning your trip, especially if you have key targets. Once you arrive in Madagascar, you can refer to this book when directing the efforts of your local guides. Most local guides specialize in lemurs, but there are many fascinating birds, reptiles and frogs that they probably know, but would not focus on finding unless you mentioned your interest.

Map of Madagascar showing biogeographic zones and most-visited natural sites This map shows the distribution of Madagascar's remaining natural habitats. The major division is between the wet east, where rainforest is the natural habitat, and the dry west, with deciduous forest. The central High Plateau is virtually deforested. For more information on biogeographic zones, see pages 12–13.

Madagascar’s most-visited natural sites Rank GENERAL TOURISTS 1 Andasibe area (Andasibe-Mantadia NP, etc.) 2 RN7 south: Toliara / Isalo NP 3 4 5 6 7

Nosy Be Far north: Ankarana and Amber Mountain NPs, etc. Tsingy de Bemaraha NP St. Marie Island Berenty / Fort Dauphin

Masoala NP

9 10 11 12 13

Kirindy Forest Ankarafantsika NP Ranomafana NP Marojejy NP Anjozorobe area (Anjozorobe-Angavo forest)

SERIOUS NATURALISTS Andasibe area (Andasibe-Mantadia NP, etc.) Ranomafana NP Isalo NP Berenty / Fort Dauphin Kirindy Forest Tsingy de Bemaraha NP Nosy Be Far north: Ankarana and Amber Mountain NPs, etc. Masoala NP Ankarafantsika NP Marojejy NP Toliara / Anakao / Ifaty Anjozorobe area (Anjozorobe-Angavo forest)

BIRDERS Andasibe area (Andasibe-Mantadia NP, etc.) RN7 south: Toliara / Ifaty / Isalo area / ZombitseVohibasia NP Ranomafana NP Ankarafantsika NP / Betsiboka Delta (Mahajanga) Masoala NP Berenty Anjozorobe area (Anjozorobe-Angavo forest) Far north: Ankarana and Amber Mountain NPs, etc. Marojejy NP Kirindy Forest Tsingy de Bemaraha NP Nosy Be St. Marie Island

Key to the sites 1 2 3 4 5 6 7 8 9 10

Amber Mountain NP Anakao Andasibe-Mantadia NP Anjozorobe-Angavo forest Ankarafantsika NP Ankarana NP Berenty Reserve Betsiboka Delta (Mahajanga) Fort Dauphin (Tôlanaro) Ifaty

11 12 13 14 15 16 17 18 19 20

Isalo NP Kirindy Forest Marojejy NP Masoala NP Nosy Be Ranomafana NP Sainte Marie Island Tsingy de Bemaraha NP Toliara (Tulear) Zombitse-Vohibasia NP

45°E

KEY

50°E

Diego-Suarez

Montane scrubland High-altitude Rainforest

Mid-altitude Rainforest

N 6

Lowland Rainforest Tapia (Uapaca) Woodland Evergreen Coastal Forest Deciduous – seasonally dry (western)

Deciduous – dry (Spiny forest and scrub) Deciduous Coastal Forest

15°S

Marsh

15°S 14

Mahajanga

Mangrove

1,500 1,000 500 200 0

Toamasina 4 18

20°S

Morondava

Antananarivo

20°S

E Co Biogeographic zones see pages 12–13 N W C E S Co

11 20 10 19

North West Central East South Coast

Toliara

25°S 45°E

100 150 km 25°S

Tôlanaro 50°E

Biogeographic zones of Madagascar

NDiego-Suarez

The zones presented here are reflected in the range icons that accompany each species account.

East [E]

l Predominant natural habitat is humid forest (‘rainforest’),

Mahajanga

N N

Madagascar’s most species-rich habitat. N l Rainforests stay lush and green year-round. C Toamasina They are watered by moisture off the Indian Ocean C that is precipitated by the eastern escarpment. Antananarivo C W Trees very tall in lowlands, but shorter in C Morondava higher-elevation forest. E Co l Species distribution within eastern rainforest is S complex. Many lemurs and frogs, and some reptiles, have small and localized distributions, often within S S specific elevations. N Toliara l Vast deforestation; much of eastern zone now covered C in scrubby secondary forest and artificial savannah. SW S Tôlanaro Lower-elevation forests have been most drastically impacted. l The highest mountains support open heath and grassland above treeline. These habitats are species-poor. l Eastern marsh is one of Madagascar’s most special and most threatened habitats. It supports several endemic birds and many frogs.

West [W]

l The western two thirds of Madagascar, comprising the central highlands and the

western lowlands, are much drier than the east, lying in the rain shadow of the eastern mountains. l Originally covered in dry deciduous forest, which has lower diversity than rainforest, but is still rich, with many endemics. Lemurs and reptiles are especially rich in the west, while frog diversity is low. l Most of the west is now savannah in which mango and palms are prominent. Large areas have also been converted to rice cultivation. l Most (although not all) of the trees in the western forest are deciduous: they lose their leaves during the dry season. The abundant leaf-litter and more open character of this habitat make it very different from eastern rainforest. l The west holds extensive wetland habitat: marshes, lakes and wide rivers. Most have been degraded by humans.

North [N]

l The north is essentially a mix of the eastern and western zones. Here the island tapers to

a point, while general climatic trends are also disrupted by Tsaratanana, Madagascar’s highest mountain massif. The Sambirano region around Nosy Be is the only area on the west coast that sees almost as much rainfall as parts of the eastern zone. l Some parts of the north, such as Amber Mountain and Lokobe National Parks (NPs), hold rainforest. This forest lacks the diversity of eastern rainforest, but does support many localized endemics. l Much of the north, such as Ankarana NP, holds dry forest like that of the western zone. Large stretches are also covered in human-created savannah habitat.

Centre [C]

l Mostly above 800 m (2,600’) elevation.

l Before humans arrived, Madagascar’s central highlands probably supported a mix of

savannah and forest. But this is the part of Madagascar that has been most heavily impacted by human activity and is now dominated by arid grasslands and eroded gullies. l Another feature of the centre is vast areas of rice cultivation. Rice paddies do support some birds and frogs. l Forest persists in a few places, mainly in valleys where it is shielded from burning. Such forest resembles eastern rainforest, although it is drier and supports fewer species.

Southwest [SW]

l The southwest is the driest part of Madagascar. Its aridity has given rise to the island’s

most distinctive habitat, the spiny forest. There is also some gallery forest and wetland.

l Least diverse of Madagascar’s zones, but supports many endemic species. The plants are

fascinating, with most endemic not just to Madagascar, but to this zone.

Coast [Co]

l Madagascar’s 10,000 kilometres (6,200 miles) of coastline encompass mangroves

(mainly in the west and north), mudflats, rocky and sandy beaches, and salt marshes.

l Supports many species of birds, along with sea turtles and marine mammals.

Coral reefs are beyond the scope of this book, but Madagascar’s reefs are world-class.

Human-modified grasslands, rice paddies and villages South of Ambositra, Central Zone

Hamerkop p. 96 Mascarene Martin p. 136

Spotted Blue Swallowtail p. 288

Jewelled Chameleon p. 192

Black Kite p. 100

Common Myna p. 176

Madagascar Fody p. 178

Mascarene Ridged Frog p. 248

Cattle Egret p. 92

Mid-elevation eastern rainforest Vohiparara Sector, Ranomafana NP, Eastern Zone

Rufous Mouse Lemur p. 32

Giraffe-necked Weevil p. 284

Comet Moth p. 314

Trees become more stunted at higher elevations; lush year-round.

Stripe-throated Jery p. 160

Satanic Leaf-tailed Gecko p. 234

Elongate Ancient Leaf Chameleon p. 188

Low-elevation eastern rainforest Masoala NP, Eastern Zone

Helmet Vanga p. 172

White-fronted Brown Lemur p. 46

Common Leaf-tailed Gecko p. 236

Cream-lined Swallowtail p. 286

Trees grow very tall; dense understorey; lush year-round.

Speckled Day Gecko p. 222

Red Ruffed Lemur p. 52

Scaly Ground-roller p. 144

Lowland Streaked Tenrec p. 76

Sambirano rainforest, coastal mudflats, and sandy beach Lokobe NP, Northern Zone Northern rainforest is very similar to eastern rainforest, but has lower species diversity.

Madagascar Paradise Flycatcher p. 152

Orange-backed Mantella p. 254

Red-legged Plated Lizard p. 212

Lesser Frigatebird p. 118

Sanfordâ&#x20AC;&#x2122;s Brown Lemur p. 46

Lesser Crested / Roseate Terns p. 116

The west coast is generally richer than the cyclone-scoured east coast.

Green Sea Turtle p. 184

Bottlenose dolphin p. 80

Western dry forest and freshwater marsh Lake Ravelobe, Ankarafantsika NP, Western Zone

Coquerelâ&#x20AC;&#x2122;s Sifaka p. 56

Fosa p. 68

White-faced Whistling-duck p. 84

Madagascar Swordtail p. 288

Western dry forest is desiccated in the dry season, but lush in the wet season, although the understorey is often open. Very similar dry forest can also be found in the northern zone.

Leaf-nosed snake p. 240

Schlegel’s Asity p. 148

Cuvier’s Madagascar Swift p. 208

Western marshes are Madagascar’s richest for birds, although they are poorer in terms of endemics than the eastern marshes.

Grey Heron p. 88

Spiny forest Parc Mosa, near Ifaty, Southwestern Zone Madagascarâ&#x20AC;&#x2122;s most visually distinctive habitat; almost barren in the dry season, but sees a wet season flush of green. Octopus trees p. 318 are found only in the spiny forest.

Grey-brown Mouse Lemur p. 32

Madagascar Giant Swallowtail p. 290

Long-tailed Ground-roller p. 146

Baobabs pp. 320â&#x20AC;&#x201C;323 are found throughout the southwest, west, and north, but are especially prominent here.

White-lined Chameleon p. 204

Madagascar Keeled Plated Lizard p. 214

How to use this book Headings Closely related and/or similar species are grouped together onto page-spreads. Each such spread has a heading. The headings in the mammal and butterfly sections include scientific family names, as family differences are fundamental to understanding these groups. Scientific family names are less useful for other groups, and are therefore not included. SECTION INTRODUCTIONS There is a two-page introductory spread for each of the four major vertebrate groups. These include information on the way to approach identification within a given group (e.g. mammal identification should be approached in a very different manner than frog identification).

Names Each species account starts with a common name. With good reason, biologists prefer scientific names for their greater precision and lesser variability. But casual naturalists generally find common names easier to understand and process, so this guide includes them for all species. In some cases, as with birds and mammals, common names are fairly standardized. The source for bird names was The Birds of Africa: Volume VIII, for lemur names was Lemurs of Madagascar, and for other mammals was Mammals of Madagascar. Finding standardized common names for reptiles, amphibians, insects and plants was more challenging. For reptiles and amphibians, the main source used was Complete Guide to Scientific and Common Names of Reptiles and Amphibians of the World. For butterflies, Field Guide to Butterflies of South Africa was followed for the species and genera covered in that book, the other names being those in general use. See Further reading (page 327) for more details of all of these volumes. Other species have one or more well established common names, often from the pet trade, in which case these were used. For many species, there is simply no established common name, or the existing names are misleading or otherwise inadequate, in which case the only option was to invent new ones (marked with asterisks). All naturalists are encouraged to pay attention to both the common and scientific names, and to realize that most Malagasy local guides will not know common names for reptiles or amphibians. The scientific names are included and, where relevant, the subspecies name if it is endemic to Madagascar. In some cases, a whole subgenus, genus, cluster of genera, or family is treated with a single account. As well as the common and scientific names, most accounts also include Malagasy and French common names. It is hoped that these will be useful both to non-English-speaking visitors and to Malagasy guides and researchers. They can also be helpful in interacting with Malagasy people who are not directly involved in tourism, but whose goodwill is essential to preserving natural places. Malagasy and French mammal and bird names are mainly from the same sources as English common names. Malagasy reptile names come from the 26

Malagasy edition of Amphibians and Reptiles of Madagascar. Other names come directly from Malagasy researchers and naturalists. Widespread species tend to have many different Malagasy names, depending on the region, but it is beyond the scope of this book to include them all. There is also wide overlap in the usage of Malagasy names, with one name often used for many different species. To avoid confusion, the use of the same Malagasy name for more than one species is generally avoided in this book. As a result, the preferred Malagasy name for certain species in a given location may not be included, or may be applied to a different species.

Icons STATUS Each account features several icons. The numbers correspond to the adjacent photographic plate, allowing easy correlation of the text and the photos. There are also icons for species that are endemic or introduced, and a coloured bar for species that are considered globally threatened (Critically Endangered, Endangered, Vulnerable) or near threatened on the International Union for Conservation of Nature (IUCN) Red List (see www.iucnredlist.org for more information). The icons and codes are summarized in the panel to the right. DISTRIBUTION N NC Each species account includes a S distribution map that shows the N broad biogeographical zones where N that species occurs. There are six N C zones (East, Southwest, West, Centre, C C North and Coastal), each of which W C is subdivided into three sections E Co S (northern, central and southern). S S These sections are shaded to indicate N C relative abundance as shown in SW S the key. These are not true range maps, but rather a shorthand way of representing approximate distributions. Those interested in detailed information about the ranges of Malagasy species should look to more in-depth references to individual groups. However, for even the most casual visitor, quick reference to range is one of the best ways to narrow down a species that you are trying to identify.

KEY TO ICONS IN SPECIES ACCOUNTS Status E Endemic Family – the species and the whole family to which it belongs are found only on Madagascar E Endemic – a species found only on Madagascar B Breeding endemic – only breeds on Madagascar R Regional Endemic – found only in the Malagasy region e Endemic subspecies – subspecies in Madagascar is (or are) endemic to the island I Introduced – a species introduced to Madagascar by humans IUCN Red List Status CRITICALLY ENDANGERED ENDANGERED VULNERABLE NEAR THREATENED

Distribution ■ common to fairly common ■ uncommon to rare Other icons and codes ♀ female ♂ male br breeding nb non-breeding im immature ju juvenile M F

Malagasy name French name 27

Boxes DIMENSIONS A grey box contains the species’ dimensions. The default dimension, unless noted otherwise, is the total length from the tip of the nose to the tip of the tail. For many reptiles, the distance from the tip of the nose (snout) to the vent (at the base of the tail), the snout to vent length [SVL] is more useful; these instances are marked. For butterflies, wingspan [WS] is the most useful measurement. When males and females are significantly different in size, separate dimensions for the sexes have been included. For mammals, weights are also given. WHERE TO SEE This box aims to go beyond abstract ideas of range to give you a substantial idea of where you might find a given species. It might include tips on the best sites for that species, and what times of day and season are best for a search. In mentioning the best sites for each species, a deliberate distinction is made between national parks proper (e.g. Andasibe-Mantadia NP) and areas that include a national park (e.g. Andasibe area). Finding certain species requires entering a national park, while others can be found both in and around a given park, including disturbed secondary habitat, community forests, and smaller satellite reserves run by community associations or NGOs.

Main Text DISTRIBUTION: This section goes beyond the information in the range icon to provide more details of where a species is found, both worldwide and within Madagascar. Terms such as ‘west’ and ‘north’ used in this section don’t necessarily correspond to the names of the biogeographic zones that form the basis for the maps. For example, ‘west coast’ might refer to the entire west coast of the island, which includes portions of the southwestern and northern zones in addition to the coast of the western zone proper. The distribution section also contains information about what habitat a species prefers. IDENTIFICATION: This section isn’t intended as a detailed description of each species, but rather a quick condensation of the best ways to recognize it and separate it from similar species. Differences between males and females are mentioned here if useful. In some cases, range and habitat information that is immediately relevant to identification is also included. VOICE: If a species is vocal, its most frequent and distinctive vocalizations are described. BEHAVIOUR: This variable section includes some further details about how a species lives: whether it’s social or solitary, nocturnal or diurnal, arboreal or terrestrial. When they are known, details of diet and reproduction are also often included here.

Inset Boxes Inset boxes are interspersed throughout the book; these usually highlight something interesting or unusual about a species or group covered by that particular page spread. There are also a few boxes that mention similar species that aren’t covered in the main text. This kind of information is also included in the identification section of the main text in some instances. 28

Photos The photos on each plate are scaled to illustrate the real size relationship between the species. Sometimes smaller photos of a different scale, such as birds in flight, are included, but these cases should be obvious. Photos included within boxes are at a different scale from the others. For most species in which males and females are very different, there are captioned photos illustrating the two. In some cases where only one photo is included, but the individual shown can be reliably sexed, this is captioned. Most of the photos were taken by the authors. Photos by other photographers are credited on pages 328â&#x20AC;&#x201C;331.

Visiting Madagascar

Seasoned international travellers with lots of time and the ability to be flexible can backpack around Madagascar and see most of its wonders. But most people will prefer to visit the country with the help of a tour company. Many international companies run natural history or birding tours to Madagascar. Both of the authors work for Tropical Birding (www.tropicalbirding.com), which offers birding, natural history and photography tours all over the world. Another way to visit Madagascar is with the help of a Malagasy tour operator. Two such companies that are reliable and professional are 8th Continent Expeditions (www.8thcontinentexpeditions.com) and Za Tour (www.zatours-madagascar.com).

Green Bright-eyed Frog p. 260

Mammals About 240 species of mammal have been described from Madagascar, but the basic inventory of its mammal species is ongoing: dozens of new species have been described in recent decades. l One-third of Madagascar’s mammals are considered to be either Critically Endangered or Endangered on the latest IUCN Red List (see page 27). l There are eight orders of mammal represented on Madagascar. The most important groups are illustrated below.

Lemurs (pages 32–65). A massive radiation of primates that is endemic to the island. There are five living families, plus a further three that have become extinct.

Tenrecs (pages 74–77). Family endemic to Madagascar, apart from a couple of otter-shrews found on the African mainland. Tenrecs have radiated into incredible diversity, ranging from shrew, hedgehog and mole, to otter-like in form.

Rodents (pages 78–79). Madagascar supports an endemic sub-family of rodents, whose most remarkable member is the rabbit-like Giant Jumping Rat.

Bats (pages 70–73). These flying mammals have colonized the island with relative ease. About 80% of the species are endemic.

Malagasy Carnivorans (pages 66–69). These predators comprise an endemic family of ten species, that vary from puma-like to mongoose-like in form.

Whales and dolphins (pages 80–81). The warm waters around Madagascar are relatively poor in marine mammals, but a few species are frequently sighted.

l Each of Madagascar’s major terrestrial mammal

groups: tenrecs, lemurs, rodents and carnivores, seems to have originated from a single colonizing event, most likely animals that rafted their way onto the island. For example, the whole range of lemurs, from tiny mouse lemurs up to the extinct ‘gorilla’ lemurs, are likely to have evolved from a single common ancestor. l Several species of mammal appear to have reached Madagascar with human assistance, namely: Asian Musk Shrew Suncus murinus, Small Indian Civet Viverricula indica, Black Rat Rattus rattus and Brown Rat Rattus norvegicus, House Mouse Mus musculus and Bush Pig Potamochoerus larvatus. Of these, the rats and mice are by far the most frequently seen. l Lemurs are Madagascar’s most celebrated biological treasure. Fifteen percent of the world's primate species and subspecies, 20% of its primate genera, and one third of its primate families, are endemic to this island. Lemurs form one of the most prominent voices in the Malagasy forest. Most species are vocal and produce many different calls. l The majority of Madagascar's mammals are nocturnal. Three of its five lemur families consist of nocturnal species, while bats, tenrecs and rodents are either mainly or entirely nocturnal. The Malagasy carnivorans vary from being mainly diurnal to entirely nocturnal. l Local guides in all protected areas invariably prioritize lemurs, and are very good at finding their quarry. Seeing most tenrecs, rodents, carnivorans and bats is a matter of luck and the amount of time spent in the forest, particularly on night walks. l Most lemur species time their breeding for the flush of food that becomes available during the wet season, so most eastern species give birth between September and November, and western species slightly later. One of the reasons to visit Madagascar between October and February is to see lemur babies. l The identification of most Malagasy mammals is made easier by reference to their ranges. Confusing groups, like woolly lemurs and sportive lemurs, often have only a single species in any one location.

Indri p. 62 is the largest living lemur

Cheirogaleidae: MOUSE LEMURS

These tiny nocturnal lemurs include the smallest living primate in the world: Madame Berthe’s Mouse Lemur Microcebus berthae. They closely resemble the galagos, or 'bushbabies', of Africa’. Modern taxonomy has seen the number of recognized species increase dramatically from two to 18, and more species may yet be described. Although mouse lemurs are occasionally found sleeping during the day (resembling tiny fur balls), most sightings are during the night, usually in the form of a pair of eyes bounding about the forest at remarkable speed. There is some variation in size and colour (from grey to rufous), but all mouse lemurs look very similar, especially with a typical night walk view. According to current information, most can be identified simply by where you are, so refer to the locality and species table on this page. E

Mouse lemurs Microcebus spp. [18 species]

M Tsidy Microcèbe

DISTRIBUTION: Found throughout the island in forest Length: 23–31 cm | 9–12" and some human-modified habitats such as scrubby Weight: 30–110 g | 1–4 oz secondary forest and plantations. WHERE TO SEE: Almost any ID: Tiny size and near-constant activity separate them Malagasy forest, and many from most other lemurs. When seen at close range, the degraded areas, will support at long tail is conspicuous. See giant mouse lemurs (page 36) least one species. for separation from those species. Most areas of the west hold two species: Grey Mouse Lemur 1b plus one other species (see below). Identification in these cases can be very diﬃcult, but compared with Grey Mouse Lemur, Golden-brown Mouse Lemur 1f has warmer brown colour on its back and a longer tail; Grey-brown Mouse Lemur 1e has a rufous stripe along its back, and a more contrasting pale grey-and-rufous facial pattern; and Madame Berthe’s Mouse Lemur is smaller, with more rufous coloration. VOICE: High-pitched, squeaky and rather rodent-like. BEHAVIOUR: Feed mainly on fruit, supplemented with invertebrates and tree gum. Species that are well studied have been shown to give birth to 1–3 young from September to May. WHICH MOUSE LEMUR AM I LOOKING AT? The most frequently visited sites in Madagascar and the mouse lemur species that can be expected there: Amber Mountain NP Andasibe-Mantadia NP Andringitra NP Anjozorobe-Angavo forest Ankarana NP Ankarafansika NP

1a 1d 1b 1f

Berenty

1b 1e

Isalo NP Kirindy Forest

Marojejy NP Masoala NP Nosy Be / Lokobe NP Nosy Mangabe Ranomafana NP Tsingy de Bemaraha NP Toliara / Ifaty Zombitse-Vohibasia NP

1c 1a 1b 1e 1b

Montagne d’Ambre Mouse Lemur Goodman’s Mouse Lemur Rufous Mouse Lemur Goodman’s Mouse Lemur Tavaratra Mouse Lemur Grey Mouse Lemur Golden-brown Mouse Lemur Grey Mouse Lemur Grey-brown Mouse Lemur Grey Mouse Lemur Grey Mouse Lemur Madame Berthe’s Mouse Lemur uncertain; Mittermeier’s Mouse Lemur? undescribed species Nosy Be Mouse Lemur uncertain; Anjiahely Mouse Lemur? Rufous Mouse Lemur Peters’ Mouse Lemur Grey Mouse Lemur Grey-brown Mouse Lemur Grey Mouse Lemur

M. arnholdi M. lehilahytsara M. rufus M. lehilahytsara M. tavaratra M. murinus M. ravelobensis M. murinus M. griseorufus M. murinus M. murinus M. berthae M. mittermeieri? M. sp. nova M. mamiratra M. macarthurii? M. rufus M. myoxinus M. murinus M. griseorufus M. murinus

Mouse lemurs are fond of licking tree gum, or bananas in the case of the famous Rufous Mouse Lemurs of Ranomafana!

1a 1c

Mouse lemur of uncertain species from Nosy Mangabe 1e

During the day mouse lemurs sleep in tree holes and dense vegetation

Cheirogaleidae: DWARF LEMURS

Dwarf lemurs are small nocturnal lemurs in the same family as mouse lemurs. They are remarkable as the only primates known to go into a hibernation-like torpor during the dry season (approximately May to September), during which they live off the reserves of fat stored in their tails. Dwarf lemur taxonomy is still imperfectly understood, and there are several undescribed species. There seems to be little overlap in the range of most species, so tentative identifications can be based on the location of a sighting. Refer to the locality and species table on this page. E

Dwarf lemurs Cheirogaleus spp. [7+ species]

Tsidihy, Hataka, Matavirambo F Cheirogale

DISTRIBUTION: Found almost throughout in forest and Length: 40–55 cm | 16–22" some human-modified habitats such as secondary forest Weight: 0·1–0·5 kg | 0·3–1·1 lb and plantations. Generally absent from the High Plateau, WHERE TO SEE: Amber and scarce in the southwest. Mountain, Masoala, and ID: Larger and slower-moving than mouse lemurs. Ankarafantsika NPs, Kirindy Smaller and more horizontal than sportive lemurs. Forest, and the Andasibe area Superficially similar to fork-marked lemurs, but with smaller are some of the easiest places to ears, no fork mark on the back, slower movements, and much find a dwarf lemur. less vocal. Separation of individual dwarf lemur species can be diﬃcult, with range the best guide. There are three widespread groups. The western and northern medius complex (including Fat-tailed Dwarf Lemur 1c ) is fairly distinctive: cold grey above and white below, with no warm tones. The eastern and northern crossleyi complex (including Montagne d'Ambre Dwarf Lemur 1b ) has broad black eye rings and shows a considerable amount rufous-brown on the head, as well as on the body in some individuals. The eastern major complex (including Greater Dwarf Lemur 1a ) has intermediate coloration, with colder brown on the head and body, and less black on the face than the crossleyi complex. VOICE: Generally quiet. Calls that have been noted include screams during territorial confrontations, a rising squeak, and a high “kak-kak-kak…” similar to the call of the diurnal Broad-billed Roller (page 140). BEHAVIOUR: Move slowly through the trees, with a mainly horizontal posture. Give birth during the wet season to 1–4 young, with 2–3 being the most common. Diet consists of fruit, nectar, leaves, buds, and insects. WHICH DWARF LEMUR AM I LOOKING AT? The most frequently visited sites in Madagascar and the dwarf lemur species that can be expected there: Amber Mountain NP Andasibe-Mantadia NP Andringitra NP Anjozorobe-Angavo forest Ankarana NP Ankarafantsika NP Kirindy Forest Marojejy NP Masoala NP / Nosy Mangabe Ranomafana NP Sainte Marie Island Tsingy de Bemaraha NP Zombitse-Vohibasia NP

1a 1a

Montagne d’Ambre Dwarf Lemur Crossley’s (or Furry-eared) Dwarf Lemur undescribed species – crossleyi complex Crossley’s (or Furry-eared) Dwarf Lemur Sibree’s Dwarf Lemur undescribed species – medius complex possible undescribed species – medius complex Fat-tailed Dwarf Lemur possible undescribed species – crossleyi complex Greater Dwarf Lemur undescribed species – crossleyi complex Greater Dwarf Lemur Fat-tailed Dwarf Lemur Fat-tailed Dwarf Lemur

C. andysabini C. crossleyi C. sp. nova C. crossleyi C. sibreei C. sp. nova C. sp. nova? C. medius C. sp. nova? C. major C. sp. nova C. major C. medius C. medius

Species in the major group have colder tones and a smaller black 'mask' than those in the crossleyi group. This is a Greater Dwarf Lemur from Maroantsetra.

Species in the crossleyi group show some warm rufous tones. This individual is a Montagne d'Ambre Dwarf Lemur.

Species in the medius group (including Fat-tailed Dwarf Lemur) are cold grey on back, with no warm tones. This individual is from a possible undescribed species in Ankarafantsika NP.

Cheirogaleidae: GIANT MOUSE AND FORK-MARKED LEMURS

Giant mouse and fork-marked lemurs are also members of the nocturnal Cheirogaleidae family. In general, they are seen much less frequently than mouse or dwarf lemurs. E

Giant mouse lemurs Mirza coquereli / Mirza zaza

DISTRIBUTION: Coquerel’s Giant Mouse Lemur M. coquereli 1a is found in the southern half of western Madagascar, in dry forest. Northern Giant Mouse Lemur M. zaza 1b is found in forest fragments and plantations in the northwest. ID: Combination of size, large ears, and active habits will normally identify them. The long, reddish tail is dark towards the end. VOICE: Makes a quiet “swiiick” call, especially at the start of the wet season. BEHAVIOUR: Nocturnal and typically solitary. Move actively, although not as frenetically as typical mouse lemurs (page 32). Diet is catholic: fruit, flowers, tree gum, invertebrates, and even small vertebrates. Give birth to twins. E

M Tsiba Microcèbe Géant

ENDANGERED Length: Weight:

54–57 cm | 21–22" 0·3 kg | 0·7 lb

WHERE TO SEE: Although uncommon Coquerel’s Giant Mouse Lemur can be seen during night walks in Kirindy Forest. Also occurs in Zombitse and Tsingy de Bemaraha NPs. Northern Giant Mouse Lemur does not occur in any frequently visited protected areas, but can be seen in plantations around the town of Ambanja.

Tanta Phaner

Fork-marked lemurs Phaner spp. [4 species]

DISTRIBUTION: Pale Fork-marked Lemur P. pallescens 2a is found in the west, Sambirano P. parienti (not illustrated) and Montagne d’Ambre Fork-marked Lemurs P. electromontis 2b in the north, and Masoala Forkmarked Lemur P. furcifer (not illustrated) in the northeast. Fork-marked lemurs are found both in forest and some disturbed habitats like plantations. ID: If seen, the fork marks on the back of the head are diagnostic. VOICE: One of the most vocal groups of nocturnal lemurs. The most common call is a piercing “wheet” much like the ‘laser gun’ call of Crested Coua (page 126). Also give chattering and squirrel-like calls. BEHAVIOUR: Move faster than all other lemurs except the mouse lemurs (page 32). Often found in small groups. Specialize in eating tree sap and gum. 2b

ENDANGERED / VULNERABLE Length: Weight:

52–66 cm | 20–26" 0·3 kg | 0·7 lb

WHERE TO SEE: Pale Forkmarked Lemur is common during night walks in Kirindy Forest, and also occurs in Zombitse and Tsingy de Bemaraha NPs. The rarely seen Masoala Fork-marked Lemur is found around Masoala. Sambirano Fork-marked Lemur occurs in the northwest, and is sometimes seen around Ambanja. Montagne d’Ambre Fork-marked Lemur is occasionally seen on Amber Mountain.

Large ears and long bushy tail

Dark line on back which forks 2a on the head 2a

Sometimes sits horizonally, but at other times is more vertical

Lepilemuridae: SPORTIVE LEMURS

Sportive lemurs are classified as an entirely separate family. They are rather chunky, with big eyes and ears, and have a vertical posture. These nocturnal lemurs are often seen during the day, roosting in tree cavities or dense tangles. At night, they move about with impressive leaps, while retaining their vertical posture. This is another group, like the mouse lemurs, where the number of recognized species (currently 26) has increased dramatically in recent years, and new species may yet be described. These species look very similar, especially at night, but show some variation in size, colour, prominence of the ears, and other traits. However, it is unusual for two species to coexist, and so most can be identified by location (see table below). E

Sportive lemurs Lepilemur spp. [26 species]

DISTRIBUTION: Throughout the island, in forested habitats, including spiny forest. ID: Similar in size to dwarf lemurs (page 34), but have a vertical rather than horizontal posture. They also tend to sit most of the time, making occasional leaps, rather than constantly moving about in a tree. Often confused with woolly lemurs (page 54), which are also nocturnal and regularly seen in habitual sleeping sites during the day. However, woolly lemurs have less prominent ears and have pale stripes on the back of their thighs; they are also more likely to be found during the day in a family huddle than the more solitary sportive lemurs. VOICE: Quite vocal, especially at the beginning of the wet season. Calls are emphatic grunts, squeals, and screams. BEHAVIOUR: During the day, they sleep in tree cavities and forks, and dense tangles. Local guides often know the whereabouts of sportive lemur sleeping sites and are able to show them to visitors. Feed on leaves, nectar, and fruit. The breeding habits of most species are poorly known, but they are thought to give birth to a single young at the beginning of the wet season.

Hataka, Boengy, Apongy F Lépilémur

Length: Weight:

43–64 cm | 17–25" 0·5–1·2 kg | 1–2·6 lb

WHERE TO SEE: Night walks in forest almost anywhere on the island can result in sightings.

1g Small-toothed Sportive Lemur from Ranomafana NP, one of the darker species

WHICH SPORTIVE LEMUR AM I LOOKING AT? The sportive lemur species that occur at the most frequently visited sites in Madagascar: Amber Mountain NP Andasibe-Mantadia NP Andringitra NP Anjozorobe-Angavo forest Ankarana NP Ankarafansika NP Berenty Isalo NP Kirindy Forest Marojejy NP Masoala NP Nosy Be / Lokobe NP Ranomafana NP Tsingy de Bemaraha NP Toliara / Ifaty Zombitse-Vohibasia NP

1a 1g 1a 1f 1c 1d 1d

1e 1g

Ankarana Sportive Lemur Weasel Sportive Lemur Small-toothed Sportive Lemur Weasel Sportive Lemur Ankarana Sportive Lemur Milne-Edwards’ Sportive Lemur White-footed Sportive Lemur Red-tailed Sportive Lemur Red-tailed Sportive Lemur Seal’s Sportive Lemur Masoala Sportive Lemur Nosy Be Sportive Lemur Small-toothed Sportive Lemur Bemaraha Sportive Lemur Uncertain; Petter’s Sportive Lemur? Zombitse Sportive Lemur

L. ankaranensis L. mustelinus L. microdon L. mustelinus L. ankaranensis L. edwardsi L. leucopus L. ruﬁcaudatus L. ruﬁcaudatus L. seali L. scottorum L. tymerlachsonorum L. microdon L. randrianasoloi L. petteri? L. hubbardorum

Occasionally has a horizonal posture 1c

Long, bushy tail. Some species of sportive lemur show a dark line on the back

1e 1f

Usually sleeps in tree holes or tangles during the day

Lemuridae: BAMBOO LEMURS I

Bamboo lemurs are members of the ‘true lemur’ family, all of which are active during the day. As their name suggests, bamboo lemurs eat bamboo and are usually found in stands of it. Three of the six Malagasy species are covered here. E

Greater Bamboo Lemur Prolemur simus

DISTRIBUTION: Formerly widespread, but now restricted to a few patches of rainforest in the central and southern parts of eastern Madagascar. ID: Much larger than other bamboo lemurs. Has a broad face and white ear-tufts. VOICE: Gives quiet “chuck” notes and a short, rising, raspy trill. BEHAVIOUR: Like Golden Bamboo Lemur, eats mainly Giant Bamboo Cathariostachys madagascariensis, although it prefers the pith inside the canes, leaving impressively mangled bamboo stalks in areas where it has fed. One baby is born in October or November.

Varibolo, Godroka Grand Hapalémur

CRITICALLY ENDANGERED Length: Weight:

85–90 cm | 2·9' 2·2–2·5 kg | 4·9–5·5 lb 1

WHERE TO SEE: Ranomafana NP is famous for this species, but the small population that is normally seen by visitors is under threat and may soon vanish, for reasons that are poorly understood. Another place to see this species is Torotorofotsy, a site near Mantadia NP that is managed by Association Mitsinjo. Entrance tickets and a local guide can be arranged at their office near Andasibe.

Broad face and white ear-tufts

The main food is the pith inside Giant Bamboo stems

Lemuridae: BAMBOO LEMURS II 1

Grey Bamboo Lemur Hapalemur griseus

M F

Bokombolo, Kotrika Petit Hapalémur Gris

DISTRIBUTION: Found in rainforest with bamboo VULNERABLE throughout the central and southern parts of eastern Length: 67 cm | 2·2' Madagascar. Also found in dry forest in the west, although Weight: 0·7–0·8 kg | 1·5–1·9 lb rarely seen by visitors there. WHERE TO SEE: Easiest to ID: Reminiscent of a sportive lemur, but active during the see in Andasibe-Mantadia NP, day, and their ears and eyes are much smaller. The very with tame individuals often similar Northern Bamboo Lemur Hapalemur occidentalis along the main trail in the (not illustrated) is found in the north and northeast, and is Analamazaotra section of the occasionally seen in Marojejy and Masoala NPs. park. Also fairly common in VOICE: Not very vocal. Sometimes gives a quiet grunting Ranomafana NP. Occasionally similar to that given by brown lemurs, and a plaintive squeal. seen in Tsingy de Bemaraha NP. BEHAVIOUR: Sometimes feeds sluggishly for long periods, but highly active at other times, making impressive leaps. Usually alone or in small groups. Feeds on bamboo and other plant material. One infant is born at the beginning of the wet season. E

Golden Bamboo Lemur Hapalemur aureus

DISTRIBUTION: Very small range in the rainforest of the southeast. ID: The golden colour of this species distinguishes it from both the smaller Grey Bamboo Lemur and the larger Greater Bamboo Lemur (page 40). VOICE: Vocal. Gives a quiet grunting similar to that of brown lemurs. The territorial call is a complicated clucking and whistling. BEHAVIOUR: Usually seen in small groups, feeding placidly within a thick stand of Giant Bamboo Cathariostachys madagascariensis. One infant is born in November or December.

Bokombolomena Hapalémur Doré

CRITICALLY ENDANGERED Length: Weight:

70–80 cm | 2·3–2·6' 1·3–1·7 kg | 2·9–3·7 lb

WHERE TO SEE: Ranomafana NP, which exists in large part due to the discovery of this species there in 1985. Local guides are skilled at tracking this flagship species.

Golden Bamboo Lemur has an amazing ability to process the potent poison cyanide, which is found in large quantities in its favourite food: the young shoots and leaves of Giant Bamboo.

1 The race around Andasibe (right) is rich brown in colour; the race in the west and southeast (below) is paler and greyer

Young shoots of Giant Bamboo are a favourite food

Lemuridae: MAKY

This handsome member of the ‘true lemur’ family is the best-known Malagasy creature, the face of the island’s biodiversity. It is also the best-studied of the island’s lemurs, and even gives its name to a popular souvenir shop. E

Ring-tailed Lemur Lemur catta

DISTRIBUTION: Found in the southern third of Madagascar in a range of habitats, from spiny forest up to the high mountains of Andringitra. Despite its extensive range, its actual distribution is quite fragmented. Like most lemurs, it is threatened by forest clearance for charcoal, pasture and agriculture; hunting for food; and capture as pets. ID: Black eye patches and a ringed tail make it unmistakable. VOICE: Plaintive hoots and raucous chattering, often given in concert by groups. BEHAVIOUR: Lives in mixed-sex groups of up to 30, in which females are dominant. Typically, a single infant is born in September. Diet consists of a large variety of plant material, plus small animals.

Maky Maki

ENDANGERED Length: Weight:

95–110 cm | 3–3·5' 2·2–3·5 kg | 4·8–7·7 lb

WHERE TO SEE: Berenty is famous for this species. It can also be found in Isalo and Andringitra NPs, as well as in the small community-run Parc Anja near Ambalavao along the RN7 highway.

Although females are socially dominant, male Ring-tailed Lemurs still vie for the attention of females. They stage ‘jumping fights’ in which they slash at each other with their upper canines. They also engage in ‘stink fights’ in which they ritually waft scent at each other.

Lemur feet are leathery, with long toes

Commonly seen in trees The most terrestrial lemur

Vocal

Black nose and rings around the eyes

Lemuridae: NORTHERN BROWN LEMURS

Brown lemurs are a large genus (Eulemur) within the ‘true lemur’ family, and include some of the lemur species most frequently seen by visitors. There are 12 species, of which eight are covered in this guide. Brown lemurs are medium-sized, vocal, and generally found in groups. Unusually for lemurs, the male and female of most species have obviously different coloration. They are generally diurnal, but can also be active at night. Fruit is their most important food, although other plant parts and small animals are also eaten. This is another group in which the number of species has increased in recent years, due to the recognition of several species that were formerly considered subspecies of Brown Lemur. E

Sanford’s Brown Lemur Eulemur sanfordi

Ankomba, Beharavoaka F Lémur de Sanford

DISTRIBUTION: Found in both rainforest and dry forest ENDANGERED in the far north. Length: 88–95 cm | 3' ID: The male has white tufts around the face, while the Weight: 1·8–1·9 kg | 4 lb female is quite plain. Occurs alongside the similarly sized WHERE TO SEE: Not hard to Crowned Lemur (page 50), but has a very different facial find in Amber Mountain and pattern. Ankarana NPs. VOICE: Makes a guttural grunting, often imitated by local guides, and also gives a squealing call. BEHAVIOUR: Active during both day and night, usually in small groups. Young are born in September or early October. Feeds on fruit, leaves, and flowers. Often found in mixed groups with Crowned Lemur. E

White-fronted Brown Lemur Eulemur albifrons

DISTRIBUTION: Rainforest in the northeast, with distribution centred on Maroantsetra. ID: The male has a mostly white head and is unmistakable, although the female is drabber. This is the only brown lemur likely to be seen in most of the sites where they occur. VOICE: Similar to Sanford’s Brown Lemur. BEHAVIOUR: Similar to Sanford’s Brown Lemur.

M Varikosy Lémur à Front Blanc

ENDANGERED Length: Weight:

89–96 cm | 2·9–3·2' ~ 2 kg | 4·4 lb

WHERE TO SEE: Fairly common on Nosy Mangabe, on Masoala, and in Marojejy NP.

1♀

1♂

Male (right) shows pale tufts on head; female (left) is plainer

2♀

2♂

Male (right) shows a mostly white head; female (left) has a dark head

Lemuridae: WIDESPREAD BROWN LEMURS 1

(Common) Brown Lemur Eulemur fulvus

M Varika Lémur Brun

DISTRIBUTION: Common in rainforest in the central NEAR THREATENED and northern parts of the east, dry forest in the northwest, Length: 84–101 cm | 2·7–3·3' and forest fragments on the High Plateau. Introduced to Weight: 1·7–2·1 kg | 3·7–4·6 lb Mayotte. WHERE TO SEE: ID: A medium-sized brownish lemur with a blackish face. Common around Andasibe. Male and female look similar. In the east, occurs with Less common but still easy to Red-bellied Lemur, which is darker red-brown overall, see in Ankarafantsika NP. with a black tail, and white marks in front of the eyes of males. In Ankarafantsika NP, occurs alongside Mongoose Lemur (page 50), which is greyer overall, and has different male and female facial patterns, both of which are different from Brown Lemur. VOICE: Fairly vocal. Most distinctive call is a mechanical rasping wail that is often given in concert by a whole group at the same time. Also gives a pig-like grunt which is often imitated by local guides. BEHAVIOUR: Found in small to medium-sized groups which feed on fruits, leaves and flowers, often remaining in one small area; capable of rapid movement through the forest. E

DISTRIBUTION: Fairly common though local both in eastern rainforest and western dry forest, in the southern half of Madagascar. ID: Male has a bright rufous patch on top of the head and a greyish back. Female is brownish with dull rufous on the head. In the east, occurs alongside Red-bellied Lemur, which is darker red-brown overall, with a black tail, and a different facial pattern. In Tsingy de Bemaraha NP, replaced by the very similar Rufous Brown Lemur Eulemur rufus (not illustrated). VOICE: Similar to Brown Lemur. BEHAVIOUR: Similar to Brown Lemur. E

M Gidro Lémur à Front Roux Méridional

Red-fronted Brown Lemur Eulemur rufifrons

Red-bellied Lemur Eulemur rubriventer

NEAR THREATENED Length: Weight:

85–103 cm | 2·7–3·4' average 2·3 kg | 5 lb

WHERE TO SEE: Ranomafana, often along the main road through the national park. Also common in Kirindy Forest. Red-fronted Brown Lemur and Red-collared Brown Lemur Eulemur collaris (not illustrated) have both been introduced to Berenty, where they hybridize.

M Tongona, Barimaso Lémur à Ventre Rouge

DISTRIBUTION: Widespread in eastern rainforest, VULNERABLE although absent from Masoala Peninsula. Length: 78–93 cm | 2·6–3·1' ID: Male has a reddish-brown body and bold white Weight: 1·6–2·4 kg | 3·5–5·3 lb ‘teardrop’ marks on its face. Female has a pale belly and WHERE TO SEE: Ranomafana less distinct ‘teardrops’. Both sexes have a black tail, and NP, including along the main are more reddish in overall coloration than other brown trails. Also occurs in Andasibelemurs. Mantadia NP, but much less VOICE: Rising, quizzical “whooey” that sounds like some common there. calls of Common Myna (page 176). Also gives quiet grunts and a series of pure, whistled notes. BEHAVIOUR: Similar to Brown Lemur, although generally more retiring and diﬃcult to see. 48

1 Medium-sized brownish lemur with mostly blackish face

2♀

2♂ Male grey on back with rufous on top of head; female brownish

3♀ Female has pale belly

3♂ Male shows white ‘teardrops’

Lemuridae: DISTINCTIVE EULEMUR SPECIES OF THE NORTH AND WEST 1

Black Lemur Eulemur macaco

M Komba Lémur Macaco

DISTRIBUTION: Found in a small area of northern VULNERABLE Madagascar in primary and secondary Sambirano forest, Length: 90–110 cm | 2·9–3·6' timber plantations, and areas of mixed cultivation and Weight: 1·8–1·9 kg | 4 lb forest. Seems more tolerant of disturbance than many WHERE TO SEE: In Lokobe lemurs. NP on Nosy Be, or on a tourist ID: All-black male is unique. The female with its overall excursion to Nosy Komba, brown coloration and white facial tufts is also unmistakable where a tame group of lemurs within its range. seems to subsist mainly on VOICE: Most common call is a distressed squeal, often given by handouts. a group in concert. BEHAVIOUR: Much like other Eulemur species. Feeds mainly on fruit, and is active both day and night. A single young is born between September and November. E

Ankomba Fiaka Lémur Couronné

Crowned Lemur Eulemur coronatus

DISTRIBUTION: The most northerly of Madagascar's ENDANGERED lemurs, almost reaching Cap d’Ambre at the island’s Length: 75–85 cm | 2·5–2·8' northern tip. Uses both moist and dry forest, including Weight: 1·1–1·3 kg | 2·4–2·9 lb degraded forest. WHERE TO SEE: Fairly common ID: Male is rufous-grey, and female pale grey. Both show and conspicuous in Amber a diagnostic crown patch, which is black in the male and Mountain and Ankarana NPs. rufous in the female. Often around the camping sites VOICE: Gives a distressed squeal, very similar to that of in both national parks. Black Lemur. BEHAVIOUR: Much like other Eulemur species. Found in small groups up to about 15 individuals. Often associates with Sanford’s Brown Lemur (page 46). Eats mainly fruit, and is active both day and night. Young are born between mid-September and October. E

Mongoose Lemur Eulemur mongoz

DISTRIBUTION: Dry forest in the northern part of western Madagascar. Also introduced to the Comoros. ID: Found alongside the similar Brown Lemur (page 48) in Ankarafantsika NP. Mongoose Lemur is paler and greyer overall, and shows strong white patches on the snout, whereas Brown Lemur has an all-black face. VOICE: Fairly quiet. Gives a long, pig-like snore, and a rasping hiss. BEHAVIOUR: Generally similar to other Eulemur species, although found in smaller groups and generally less conspicuous. Young are born in October.

M Dredrika Lémur Mongoz

CRITICALLY ENDANGERED Length: Weight:

75–83 cm | 2·5–2·7' 1·1–1·6 kg | 2·4–3·5 lb

WHERE TO SEE: Fairly common in Ankarafantsika NP. Watch the mango trees around the park headquarters, both during the day and at night.

1♂ 1♀

Both sexes have tufts on the sides of the head. All-black male very different from female

2♂

2♀

Male is rufous-grey, with black on top of the head; female is pale grey, with a rufous ‘crown’ 3♂

3♀

Male grey, with rufous on the face; female mostly grey, with white below

Lemuridae: RUFFED LEMURS 1

Black-and-white Ruffed Lemur Varecia variegata

DISTRIBUTION: Lowland and middle-elevation rainforest in the east. Its range is patchy and fragmented. ID: Large, long-tailed, black-and-white lemur. Similar in size and coloration to Indri (page 62), but that species has a very short tail. VOICE: Extremely loud and distinctive: aggressive and almost agonized screaming, often given by a whole group in concert, and answered by neighbouring groups. BEHAVIOUR: Generally found high in the canopy. Capable of tremendous leaps, but they often do not move much for long periods. Normally found in small groups. Feeds mainly on fruit, often hanging by its back legs to feed. Female gives birth to twins, which are placed in a nest when young. Ruffed lemurs seem to be unique among primates in this nesting behaviour. E

Red Ruffed Lemur Varecia rubra

DISTRIBUTION: Primary rainforest on the Masoala Peninsula, in the northeast. ID: Large size and red colour render it unmistakable. VOICE: Similar to Black-and-white Ruffed Lemur: very loud and vocal. BEHAVIOUR: Similar to Black-and-white Ruffed Lemur. Female gives birth to 2–3 young between September and early November.

M Varikandana Vari Noir-et-blanc

CRITICALLY ENDANGERED Length: Weight:

1·1–1·2 m | 3·5–3·9' 3·1–3·6 kg | 6·8–7·9 lb

WHERE TO SEE: Can usually be found in the Mantadia sector of Andasibe-Mantadia NP with some effort. Also in Ranomafana NP, where it can often be heard on the farther (southeastern) reaches of the main trail system, but is difficult to see due to the precipitous terrain. M

Varimena, Varignena F Vari Roux

CRITICALLY ENDANGERED Length: Weight:

1–1·2 m | 3·3–3·9' 3·3–3·6 kg | 7·3–7·9 lb

WHERE TO SEE: Fairly common and conspicuous in and around Masoala NP. Sometimes in forest along the beach near the tourist lodges, but more commonly in the adjacent primary forest.

There is a large amount of variation in the pattern of black and white on the Black-and-white Ruffed Lemur. There are three different subspecies described, which some experts consider full species: ‘Variegated’ Varecia variegata, ‘Southern’ V. editorum, and ‘Northern’ V. subcincta. ‘Southern’ can be seen in Ranomafana and Andasibe-Mantadia NPs, while ‘Northern’ may be encountered on Nosy Mangabe. ‘Variegated’ is best seen in Betampona Special Nature Reserve, well off the normal tourist circuit. ‘Northern’ shows the most black, with an all-black rump and half-black hindlegs. ‘Southern’ has a white rump, and a smaller patch of black on the hindlegs. ‘Variegated’ is similar to Southern, but with a white band along its back. 1

'Northernâ&#x20AC;&#x2122; Black-and-white Ruffed Lemur (below) has much black on its back half

The 'red panda' of Madagascar!

Ruffed lemurs are extremely loud and vocal, and often hang upside down

Indriidae: WOOLLY LEMURS

Woolly lemurs are members of the family Indriidae, which also includes the sifakas (pages 56–61) and the Indri (page 62). All Indriidae are characterized by vertical clinging and leaping behaviour, resting upright against a support, and moving through the forest in impressive leaps of up to 10 m (33 feet). Woolly lemurs are quite different from the rest of the family, being much smaller, and nocturnal rather than diurnal. They are often pointed out at day roosts by local guides in national parks. As in many other groups of lemur, there has been a flurry of new species recognized in recent decades. There are now nine described woolly lemurs, and that number may grow further. Several species are very local and will be seen only by visitors heading to remote parts of the island. All woolly lemurs are similar, with only subtle differences between them; fortunately, most locations have only a single species, which can be identified by reference to the table below. E

Woolly lemurs

Avahi spp. [9 species]

Fotsife, Avahy, Ampongy F Avahi

DISTRIBUTION: Found in forest, including secondary ENDANGERED / forest. Six of the nine species are found in the eastern VULNERABLE rainforest, one in the Sambirano forest of the north, and Length: 48–70 cm | 19–28" two in the dry forest of the west. Weight: 0·9–1·4 kg | 2–3 lb ID: Superficially similar in size and shape to sportive lemurs (page 38). The easiest way to distinguish a woolly WHERE TO SEE: Eastern Woolly Lemur is often found lemur is normally the white stripes on the back of the thighs. at day roosts or at night in Woolly lemurs found at a day roost are often in huddles of multiple family members, whereas this is uncommon in sportive the Andasibe area. Western Woolly Lemur is frequent in lemurs. Dwarf lemurs (page 34) are smaller, have a more Ankarafantsika NP. Peyriéras’ horizontal posture, and tend to crawl rather than leap when on Woolly Lemur is occasionaly the move. found in Ranomafana NP. VOICE: A high-pitched, grunted “avahi”, which gives the genus its name. Also produces a long, descending series of quavering whistles. BEHAVIOUR: Generally sluggish, perhaps due to its nutrient-poor diet of leaves. Groups sleep huddled together during the day. The single young is born in August or September. WHICH WOOLLY LEMUR AM I LOOKING AT? The woolly lemur species that occur at the most frequently visited sites in Madagascar: Andasibe-Mantadia NP Andohahela NP Andringitra NP Anjozorobe-Angavo forest Ankarafansika NP Marojejy NP Masoala NP Ranomafana NP Tsingy de Bemaraha NP

1b 1c 1b 1a

Eastern Woolly Lemur Southern Woolly Lemur Peyriéras’ Woolly Lemur Eastern Woolly Lemur Western Woolly Lemur Eastern Woolly Lemur Masoala Woolly Lemur Peyriéras' Woolly Lemur Bemaraha Woolly Lemur

A. laniger A. meridionalis A. peyrierasi A. laniger A. occidentalis A. laniger A. mooreorum A. peyrierasi A. cleesei

Nocturnal lemurs can be easily spotted at night by their conspicuous eye shine

Typical daytime sighting of a 'ball' of half-asleep Eastern Woolly Lemurs

Vertical posture is typical of all Indriidae, including woolly lemurs

Smaller ears than sportive lemurs; conspicuous white stripe on thighs

Indriidae: WESTERN SIFAKAS

Sifakas are among the biggest, best-known, and most engaging of the island’s lemurs. Seeing all nine species is something of an accomplishment for lemur-watchers, as the species are scattered all around the island, and some of them are very rare and local. Although all species are similar in diet and behaviour, the sifakas in the eastern rainforest tend to live at lower densities, in smaller groups with larger home ranges. E

Decken’s Sifaka Propithecus deckenii

DISTRIBUTION: Dry forest in the central part of western Madagascar. ID: A large white sifaka with a black face. VOICE: Explosive “see-fak” similar to Verreaux’s Sifaka. BEHAVIOUR: Similar to Verreaux’s Sifaka. Crowned Sifaka (page 60) has a small range, wedged between those of Decken’s and Coquerel’s Sifakas.

Coquerel’s Sifaka Propithecus coquereli

DISTRIBUTION: Dry forest in the northwest. ID: A large chestnut, black and white lemur. VOICE: Explosive “see-fak” similar to Verreaux’s Sifaka. BEHAVIOUR: Found in groups of 3–10. Diet and most behaviours similar to other sifakas. Single young born in June or July. E

Verreaux’s Sifaka Propithecus verreauxi

DISTRIBUTION: Spiny forest and other kinds of dry forest in the southwest. ID: A large, mostly white lemur with a dark brown crown. Occasionally shows dark grey on limbs and back. Unmistakable within its range. VOICE: The harsh “see-fak” alarm call gives this group of animals its common name. BEHAVIOUR: Found in groups from 2–14, averaging 5–6. They move through trees by making often-spectacular leaps, and sometimes also along the the the ground with an extraordinary ‘dancing’ motion. The diet consists of leaves, fruit, and flowers. The single young is born in July or August.

M Sifaka Fotsy* Propithèque de von der Decken

ENDANGERED Length: Weight:

0·9–1·1 m | 3–3·6' 3–4·5 kg | 6·6–9·9 lb

WHERE TO SEE: Easy to see at Tsingy de Bemaraha NP. Also common and tame in the remote Beanka Forest, north of Bemaraha.

M Sifaka Mena Sandry* Propithèque de Coquerel

ENDANGERED Length: Weight:

0·9–1·1 m | 3–3·6' 3·7–4·3 kg | 8·1–9·5 lb

WHERE TO SEE: Ankarafantsika NP, often around the headquarters.

M Sifaka Propithèque de Verreaux

ENDANGERED Length: Weight:

0·9–1·1 m | 2·9–3·6' 3–3·5 kg | 6·6–7·7 lb

WHERE TO SEE: This is one of the lemurs for which Berenty is famous, and that is where most ‘dancing lemur’ photos have been taken. Also easy to find in Kirindy Forest, and occurs in Zombitse and Isalo NPs, although it can be more elusive in those parks.

Despite jumping powerfully between octopus trees (page 318) in the spiny forest, Verreaux's Sifakas seem to avoid damage to their hands and feet.

Whitest of the western sifakas 3

Mostly white; dark brown crown

Dark brown marks on limbs; white head, black face 1

All sifakas are capable of incredible leaps (above); sifakas 'dance' to cross treeless areas (below). 3

Indriidae: EASTERN SIFAKAS

Sifaka coloration The nine species of sifaka vary from all-white (Silky) to all-black (Perrier’s), these two extremes living in fairly close geographical proximity to each other in northern Madagascar. There is also considerable colour variation within some species. The taxonomic status of the sifakas is still controversial, and some experts recognize fewer than nine species. E

Silky Sifaka Propithecus candidus

DISTRIBUTION: Rainforest on and around the Marojejy Massif of the northeast. ID: The whitest of the sifakas, with a black face, which in some individuals is mottled with pink. VOICE: Similar to Milne-Edwards’ Sifaka. BEHAVIOUR: Diet consists of leaves, fruit, and seeds. A single infant is born in June or July.

Milne-Edwards’ Sifaka Propithecus edwardsi

DISTRIBUTION: Middle- and high-elevation rainforest in the southern part of eastern Madagascar. ID: A large, dark sifaka that cannot be confused with anything else in its range. VOICE: Not very vocal, but makes an explosive sneezing “simpona”, which gives it its Malagasy name, and also a series of grunts and a plaintive whistle. BEHAVIOUR: Social habits and diet similar to other sifakas. Young are born in June and July. E

Diademed Sifaka Propithecus diadema

DISTRIBUTION: Rainforest in the north-central part of eastern Madagascar. ID: A large grey, gold, and white sifaka. VOICE: Similar to Milne-Edwards’ Sifaka. BEHAVIOUR: Similar to other sifakas.

M Simpona Fotsy Propithèque Soyeux

CRITICALLY ENDANGERED Length: Weight:

0·9–1·1 m | 3·1–3·4' 5–6 kg | 11–13 lb

WHERE TO SEE: In Marojejy NP. Usually not too difficult to find around Camp Marojejia, the middle of the park’s three camps. This rare beauty is worth the hike.

M Simpona Mainty* Propithèque de Milne-Edwards

ENDANGERED Length: Weight:

0·8–1·0 m | 2·6–3·3' 5·0–6·5 kg | 11–14 lb

WHERE TO SEE: With some effort, can be found in Ranomafana NP, either on the main trails or the Vohiparara Trail.

M Simpona Propithèque à Diadème

CRITICALLY ENDANGERED Length: Weight:

0·9–1·1 m | 3·1–3·4' 6–8·5 kg | 13–19 lb

WHERE TO SEE: AndasibeMantadia NP. The reintroduced sifakas in the Analamazaotra part of the park are the easiest to see.

Can show pink on face 1

Whitest sifaka

Mostly black 3

Most colourful sifaka

Indriidae: LOCAL SIFAKAS 1

Perrier’s Sifaka Propithecus perrieri

DISTRIBUTION: Small area of dry forest in the far north. ID: An unmistakable all-black sifaka. VOICE: Gives an explosive sneezing call. BEHAVIOUR: Similar to other sifakas, although groups are smaller: from 2–6 individuals.

M Ankomba Joby Propithèque de Perrier

CRITICALLY ENDANGERED Length: Weight:

85–90 cm | 2·9' 4·3–5·0 kg | 9·5–11 lb

WHERE TO SEE: Requires a special visit to the Andrafiamena protected area, run by the NGO FANAMBY. For accommodation, there is a community campsite and a newly constructed lodge called ‘Black Lemur Camp’. Also found in the Analamerana Special Reserve, administered by Madagascar National Parks.

Tattersall’s (Golden-crowned) Sifaka Propithecus tattersalli

DISTRIBUTION: Small area of dry forest in the far north. ID: A mostly white sifaka with gold on top of the head, which gives this species its alternative English name. VOICE: Various calls include a “see-fak” call similar to that of Verreaux’s Sifaka (page 56). BEHAVIOUR: Found in groups of 3–10. Young are born in July and August.

M Ankomba Malandy Propithèque de Tattersall

CRITICALLY ENDANGERED Length: Weight:

87–94 cm | 2·8–3·1' average 3·5 kg | 7·7 lb

WHERE TO SEE: Will require a visit to the remote Daraina region of the far north. There is a protected area run by FANAMBY where this sifaka can easily be found. Visitors can bring their own camping gear or stay at the Camp Tattersalli tented camp. Visits should be coordinated through FANAMBY.

Crowned Sifaka Propithecus coronatus

DISTRIBUTION: Small area of dry forest in the northwest. ID: Separated from Decken’s Sifaka (page 56) by its alldark head. BEHAVIOUR: Not well known, but generally similar to other sifakas.

M Sifaka Mainty Loha* Propithèque Couronné

ENDANGERED Length: Weight:

0·9–1 m | 2·8–3·3' 3·5–4·3 kg | 7·7–9·5 lb

WHERE TO SEE: One accessible site is the tiny Antrema Forest, near the lighthouse a few kilometres north of Katsepy, and across the Betsiboka River from Mahajanga. Another site is the Bombetoka-Belemboka Forest, near the village of Boanamary, south of Mahajanga. This site is managed by FANAMBY, and visits should be coordinated through them. 2 A Tattersall's Sifaka demonstrating the astounding leaping abilities typical of all the sifakas.

Blackest sifaka All-dark head 2

Tufty ears and gold crown

All sifakas have long, powerful hind legs

Indriidae: INDRI

The Indri is one of Madagascar’s best-known creatures and is classified as the sole member of its genus within the sifaka family. E

Indri Indri indri

DISTRIBUTION: The central and northern portions of the eastern rainforest. Absent from the Masoala Peninsula. ID: This is the heaviest living lemur. Although its colour is variable, the Indri’s size, short tail, and bushy ear-tufts make it unmistakable. VOICE: One of the world’s great natural sounds. Groups start calling with a breathless hooting, and then break into long, slurred, remarkably loud notes, almost deafening at close range. Indris are most vocal during the morning and in the wet season, but may call at any time, even at night. BEHAVIOUR: Lives in small groups of 2–6, with females seemingly the dominant individuals. Like other members of the sifaka family, it generally sits in a vertical position, but is capable of incredibly powerful leaps through the forest. Eats leaves, along with some fruit, flowers, seeds, bark, and a daily dose of soil. Female gives birth every two or three years to a single infant, born in May or June.

Babakoto F Indri

CRITICALLY ENDANGERED Length: Weight:

69–77 cm | 2·2–2·5' 6–9·5 kg | 13–21 lb

WHERE TO SEE: The flagship species of Andasibe-Mantadia NP. The habituated Indris of the Analamazaotra portion of the park are usually easy to find. Although the Indris in Mantadia are often shy, that sector of the park offers the wonderful experience of hearing several groups counter-sing across a rugged landscape. The black form of Indri can be found in the Anjozorobe-Angavo forest, accessed via Saha Forest Camp.

Extinct Megafauna The Indri, as the largest living lemur, is a reminder of Madagascar’s lost megafauna. At least 17 extinct lemur species have been identified from subfossils. All of these were larger than the Indri, some much larger. There were three ‘koala lemurs’, weighing as much as 85 kg (187 lb); a whole family of ‘sloth lemurs’, many of which climbed through trees upside down, in sloth fashion, and probably weighed almost 200 kg (441 lb); and even a giant aye-aye. Human activity contributed to the extinction of these giant lemurs, although they may have survived until well after the first Europeans visited Madagascar. The Malagasy name for Indri, Babakoto, roughly translates as ‘father of man’

Very short tail

Extremely vocal

Daubentoniidae: AYE-AYE

This remarkable creature is one of the world’s most bizarre animals. Its strangest features are its perpetually growing incisor teeth and its thin, elongated middle fingers, which are used to extract larvae from dead wood. Although it was sometimes considered to be a rodent in the past, recent genetic studies have placed it firmly in the lemurs. It forms its own family (one of Madagascar’s five lemur families), whose exact relationship to the other lemurs is still uncertain. An extinct giant aye-aye is known from subfossils. E

Aye-aye Daubentonia madagascariensis

M F

Hay-Hay Aye-aye

DISTRIBUTION: Perhaps the most widely distributed ENDANGERED lemur. Found in most of the forested environments on the Length: 74–90 cm | 2·4–2·9' island, although absent from the southwestern spiny forest. Weight: 2·5 kg | 5·5 lb ID: The large eyes and ears, and grizzled black fur are distinctive. Larger than other nocturnal lemurs. VOICE: Rarely heard “aye-aye-aye” that gives the species its name. BEHAVIOUR: Nocturnal and generally solitary. One of its most important foods is the nut of the Ramy tree Canarium madagascariense, but it feeds on a variety of larvae, seeds, nectar, and fruit. Signs of Aye-aye feeding, such as gnawed nuts and holes in trees, are seen much more often than the animals themselves. Nests are often used for sleeping during the day. The birth of a single young does not seem to be seasonal; the female gives birth every 2–3 years. WHERE TO SEE: A remarkably shy and elusive nocturnal creature. Although Aye-aye is present in most national parks, it is very rarely seen. The best chance is on Ile Roger, or ‘Aye-aye Island’, which hosts an introduced population. This island is located off Mananara-Nord, in the northeast. Visits can be arranged by the owner of Chez Robert in Mananara-Nord, who owns the island. Nosy Mangabe, off Maroantsetra, was formerly good for Aye-aye, but the current ban on night walks in national parks has made it virtually impossible to find there. Some visitors have found Aye-aye on night walks around the town of Maroantseta.

Nightlife of the Eighth Continent A visit to Madagascar would not be complete without at least one night walk in dry forest and another in rainforest. This is the best way to see a remarkably high proportion of the island’s wildlife. About half of the lemurs, several birds, and many geckos, snakes, and frogs are best sought at night, and chameleons are easiest to find when they sleep on conspicuous low tree limbs. Good local guides have flashlights, but keen visitors should bring a headlamp, and perhaps a more powerful light. Most mammals are easily spotted by their ‘eyeshine’ and local guides are remarkably good at finding amphibians and reptiles. Night walks inside the national protected area system have been prohibited for the last few years and visitors are now restricted to walks along main roads, in peripheral zones, or in community forests and other alternative reserves. 64

Huge ears; mix of black and white hairs

Long, bushy tail

The incisors grow constantly, like those of a rodent (above); the middle finger is almost skeletal (below)

Eupleridae: MALAGASY CARNIVORANS

There are 11 species of Malagasy carnivorans, of which the four most likely to be seen by visitors are covered here. Sighting even one of these four species takes some luck, as these are scarce and retiring creatures. E

Ring-tailed Vontsira Galidia elegans

DISTRIBUTION: Found in most of the island’s rainforest, and in Sambirano forest. Scarce in the western dry forest, and absent from the spiny forest. ID: Dark reddish coloration and long, ringed tail easily identify it: formerly called Ring-tailed Mongoose. VOICE: The most frequently heard call is a high-pitched whistle. Also gives a low-pitched, quiet chattering. BEHAVIOUR: Generally diurnal, and found alone or in small groups, on the ground or in trees. Its diverse diet includes small mammals, birds, reptiles, amphibians, fish and invertebrates. Young are born between November and January. E

M Vontsira Vontsira à Queue Annelée

Length: Weight:

55–69 cm | 22–27" 0·8–1·1 kg | 1·7–2·4 lb

WHERE TO SEE: Fairly tame individuals can be seen around the campsites in Ankarana and Amber Mountain NPs. Also seen regularly in eastern rainforest, especially Ranomafana NP, and in western dry forest in Tsingy de Bemaraha NP.

Northern Bokiboky Mungotictis decemlineata

M Bokiboky Mungotictis

DISTRIBUTION: A small area of western dry forest ENDANGERED centred on Morondava. Length: 45–56 cm | 18–22" ID: Mongoose-like shape, with long nose and bushy tail. Weight: 0·4–0·5 kg | 0·9–1·2 lb Eight to ten fine lines on the back and sides: formerly WHERE TO SEE: Fairly common called Narrow-striped Mongoose. in Kirindy Forest, often in the VOICE: Quiet “tok tok tok” calls, given by members of group to camp itself. stay in touch. BEHAVIOUR: Generally diurnal and social, found in groups of up to 12. Most often on the ground, but also climbs trees. Eats mainly insects, but also other invertebrates, small mammals, and reptiles. The single young is born between October and December. E

Spotted Fanaloka Fossa fossana

DISTRIBUTION: Mainly in eastern rainforest, although also occurs in drier forest in the north. ID: Heavily spotted and striped on the back. Quite similar to the introduced Small Indian Civet Viverricula indica (not illustrated), but the civet is larger, with thicker legs, a longer body, and a tail with complete rings. VOICE: Not frequently heard, but does make a contact call described as “coq-coq”. BEHAVIOUR: A shy, nocturnal species that is found singly or in pairs, usually on the ground or low in trees. Feeds on small mammals, reptiles, frogs and invertebrates. The single young is born during the wet season. 66

M Fanaloka Fanaloka Tacheté

VULNERABLE Length: Weight:

62–71 cm | 24–28" < 1·9 kg | 4·2 lb

WHERE TO SEE: Extremely shy. Formerly habituated at ‘Belle Vue’ along the main trails at Ranomafana NP, but the current night walk ban precludes seeing this species there. Occasionally spotted elsewhere in Ranomafana NP, in Ankarana NP, and in other parks, but seeing this animal requires luck.

1 Northern individuals (pictured) very red; darker in the east; black legs and belly in the west

Long and low to ground; complete bands on tail 2

Fine lines on back; pointed face

Rows of spots on back; incomplete bands on tail

Eupleridae: FOSA 1

Fosa Cryptoprocta ferox

M Fosa Cryptoprocte

DISTRIBUTION: Although rarely seen, the Fosa is found VULNERABLE in virtually all of the intact forest on the island, and will Length: 1·4–1·7 m | 4·6–5·6' even venture out of forest into agricultural areas and Weight: 5–10 kg | 11–22 lb villages during the night. WHERE TO SEE: Generally hard ID: This long-tailed, heavy-jawed, puma-like carnivoran is to fi nd. Kirindy Forest is the best the largest living indigenous land mammal in Madagascar, location. Sightings there are and is unlikely to be mistaken for anything else. likely between May and October, VOICE: Most common call is a quiet, ominous, grunted “meow”. and almost guaranteed during During the mating season, the male gives a low roar, while the the November mating season. female makes a wild, higher-pitched howling. Also regularly seen in Ankarana, BEHAVIOUR: Active both day and night. Equally at home on Andasibe-Mantadia, and the ground and high in trees, the Fosa is the feared predator of Zombitse-Vohibasia NPs, most Madagascar, preferring mammals, including the largest lemurs often during November, when pairs may mate for hours high in and the introduced Bush Pig Potamochoerus larvatus (not the trees. illustrated). It will also eat smaller carnivorans, Giant Jumping Rat, tenrecs, reptiles, birds, amphibians, invertebrates and domestic animals. Between two and four young are born during the wet season. Con-fosa-ing Two animals are often called ‘fosa’ in Malagasy: the Fosa proper (alternately ‘fosa varika’), and the Spotted Fanaloka (page 66). Adding to the confusion, the scientific name of the Spotted Fanaloka is Fossa fossana due to some confusion when this species was described. The rare Falanouc Eupleres goudotii (not illustrated) may also be referred to as 'fosa', so care is needed when when discussing carnivoran sightings, especially with local guides who are helping you to search for these creatures. Male Fosas have the largest penis bone in relation to their size of any land mammal, and pairs engage in snarling bouts of mating that can last for hours.

Shaped like a stretchedout mountain lion

Equally at home in trees and on the ground

Powerful feet and jaws

Chiroptera I: MEGABATS

There are at least 45 species of bat in Madagascar, of which nearly 80% are endemic. Most Malagasy bat lineages originated from Africa, but at least one Asian group is also represented. Many of these species are virtually impossible to distinguish based on typical night-time views in flight. The few species that are distinctive even in flight, or that are frequently seen when roosting, are the ones covered here. For most bats, wingspan (WS) is given as well as length. E

Madagascar Straw-coloured Fruit Bat

Eidolon dupreanum

DISTRIBUTION: Throughout Madagascar, in a variety of habitats. ID: Large, but smaller than Madagascar Flying Fox. Shows duller buff coloration, a narrower snout, and narrow whitish wings in flight. VOICE: High-pitched squeals and chatters. BEHAVIOUR: Forms roosts during the day, which often contain hundreds of individuals. At night, ventures out to feed on fruit, nectar and pollen.

VULNERABLE Length: WS: Weight:

WHERE TO SEE: At night may be seen in many places. Well-known day roosts include Raphia palms in Tsimbazaza Zoo, Tana, caves in Ankarana NP, and cliffs in Tsingy de Bemaraha NP.

Madagascar Flying Fox Pteropus rufus

M Angavo Roussette Paillée de Madagascar

19–22 cm | 7–9" 75–95 cm | 2·5–3·1' 0·2–0·3 kg | 0·6–0·8 lb

Narrower muzzle than flying fox M Fanihy Renard Volant de Madagascar

DISTRIBUTION: Found throughout, mainly in forest, VULNERABLE although day roost sites are often outside of forest, for Length: 24–27 cm | 9–11" example in Eucalyptus plantations. WS: 1–1·3 m | 3·3–4·1' ID: A huge bat with a wingspan of about 1 m (3·3'), looking Weight: 0·5–0·8 kg | 1·1–1·7 lb almost pterodactyl-like in flight. The Madagascar Strawcoloured Fruit Bat is similar but smaller, with a more pointed and less fox-like face, and whitish wings, visible in flight. VOICE: High-pitched chattering. BEHAVIOUR: Forms roosts during the day that can contain thousands of individuals. At night, ventures out to feed on fruit, masticating the pulp, then spitting it out after extracting the juice. WHERE TO SEE: There is a large roost at Berenty. Also not difficult to see on and around Nosy Mangabe, on Masoala, and on Nosy Tanikely near Nosy Be.

Broad muzzle

Dull buffy coloration

Bright buffy coloration

Chiroptera II: SMALLER BATS 1

Mauritian Tomb Bat Taphozous mauritianus

M Ramanavy Taphien de Maurice

DISTRIBUTION: Found throughout much of the Length: 10–11 cm | 4" Afrotropics and the Malagasy region. Widespread in Weight: 30g | 1·1 oz many habitats, but seemingly absent from the far north. WHERE TO SEE: Can usually ID: Bicoloured: grey on the back and white below. be found roosting on trees Distinctly upright roosting posture, often supporting around the headquarters at the body with folded wings. Ankarafantsika NP. VOICE: Quiet chatter. BEHAVIOUR: During the day, sleeps in small groups on tree trunks, cliffs, and man-made structures. Stays alert, and may fly off if disturbed. Feeds on insects at night. E

Commerson’s Leaf-nosed Bat Hipposideros commersoni

DISTRIBUTION: Occurs widely in forested habitats. ID: The relatively large size and bizarre ‘leaf-nose’ are distinctive. Trident bats Triaenops spp. and Paratriaenops spp. (not illustrated) are similar, but much less frequently seen by casual visitors. Trident bats can be recognized by the three fleshy spikes protruding from the top of the nose. BEHAVIOUR: Roosts in small groups in caves and hollow trees. At night, hunts insects and perhaps small frogs.

M Valo Orona Phyllorhine de Commerson

NEAR THREATENED Length: WS: Weight:

11–15 cm | 4–6" 48–67 cm | 19–22" 40–80g | 1·4–2·8 oz

WHERE TO SEE: Can occasionally be found during the day or on night walks in Kirindy Forest, Nosy Mangabe, and Ankarana, Ankarafantsika, and Tsingy de Bemaraha NPs.

The caves in Ankarana and Tsingy de Bemaraha NPs are good for several bat species, including the Madagascar Free-tailed Bat Otomops madagascariensis 3 .

Small nose and rounded ears 2

Large nose and large, pointed ears

Grizzled grey back

Tenrecidae: TENRECS I

Tenrecs are among Madagascar’s most engaging creatures, but are unfortunately difficult to see. There are 32 described species, the majority of which are shrew tenrecs Microgale spp., which are rarely observed by non-specialists. The largest and most frequently encountered tenrecs are covered here. E

Greater Hedgehog Tenrec Setifer setosus

M Sokina Grand Tenrec-hérisson

DISTRIBUTION: Throughout Madagascar in most Length: 16–23 cm | 6–9" habitats, sometimes even in large towns. Weight: 0·2–0·3 kg | 0·4–0·7 lb ID: A spiny tenrec that is similar to Lesser Hedgehog WHERE TO SEE: Not Tenrec, but is larger and has a more pointed snout. uncommon during the wet Smaller than Common Tenrec (page 76), and covered in season, and can be found spines rather than coarse fur. during night walks in most of VOICE: Various grunts and squeaks. the popular protected areas. BEHAVIOUR: Nocturnal. Forages on the ground for invertebrates and fruit, but can climb into vegetation. When threatened, it rolls into a ball. During the dry season, aestivates in tree hollows. One to four young are born early in the wet season. E

Lesser Hedgehog Tenrec Echinops telfairi

DISTRIBUTION: Southwestern and central-western Madagascsar, in spiny forest and dry forest. ID: A spiny tenrec that is similar to Greater Hedgehog Tenrec, but is smaller, with a less pointed snout. BEHAVIOUR: Nocturnal. Found both in trees and on the ground. During the dry season, aestivates in a hollow tree. Feeds on invertebrates and fruit. When threatened, rolls into a ball. One to ten young are born in the wet season.

Length: Weight:

14–18 cm | 5·5–7" 110–230g | 4–8 oz

WHERE TO SEE: Night walks during the wet season in Berenty, Kirindy Forest, and around Ifaty.

Hedgehog tenrec defensive posture 2

M Tambotriky Petit Tenrec-hérisson

Smaller than Common Tenrec (page 76), and has spines

Snout slightly shorter than that of Greater Hedgehog Tenrec

Tenrecidae: TENRECS II

Lowland Streaked Tenrec Hemicentetes semispinosus

M Sora Tenrec Zébré des Terres Basses

DISTRIBUTION: Eastern Madagascar, in lowland and Length: 13–19 cm | 5–8" mid-elevation rainforest, and adjacent human-modified Weight: 90–220g | 3–8 oz areas. WHERE TO SEE: Common at ID: Covered in spines. Distinctive pattern of black and night in the villages adjacent cream stripes along the body. Highland Streaked Tenrec the main cluster of lodges on Hemicentetes nigriceps (not illustrated) is similar and can be the Masoala Peninsula. Also found in Andringitra NP; it has broader and paler stripes, and is occasionally found around less spiny. Andasibe and Ranomafana. Most sightings are in the wet VOICE: Makes quiet sounds by vibrating the spines. season. BEHAVIOUR: Mainly nocturnal. Roots through leaf-litter to find invertebrates. When threatened, it erects the spines and twitches towards an antagonist. Two to 11 young are born in the wet season. E

Common Tenrec Tenrec ecaudatus

M Trandraka Hérisson Malgache

DISTRIBUTION: Throughout Madagascar in open Length: 29–40 cm | 11–16" grassland, forest, and edge habitat. Introduced to the Weight: 1–2 kg | 2·2–4·4 lb Comoros, Seychelles, and Mascarenes. WHERE TO SEE: Scarce due to ID: The largest tenrec. Adults have shaggy, uniformly hunting, but possible in most brownish fur. Young have stripes and resemble Lowland protected areas, especially at Streaked Tenrec, but will usually be with their mother. night during the wet season. BEHAVIOUR: Largely nocturnal and solitary. Roots in leaf-litter and soil to find invertebrates, small vertebrates and fruit. Aestivates during the dry season, and the young are born in December or January.

The Common Tenrec is one of the world’s most prolific mammals, with a brood size up to 32! The female has a remarkable 17 pairs of nipples, while the retractable penis of the male is almost as long as its whole body.

1 Lowland Streaked Tenrecs can communicate with an ultrasonic sound produced by quivering a special set of spines on their backs!

The largest tenrec

Nesomyidae: RODENTS

There are 30 species of rodent known from Madagascar. Of these, 27 comprise a subfamily that is endemic to the island, while the other three are introduced species. Most Malagasy rodents are difficult to find; the introduced rodents are seen more often than any indigenous species. Covered here are the rodents that are most likely to be spotted by visiting naturalists. E

Red forest rats Nesomys spp. [3 species]

M Voalavomena Rat Rouge Forestier*

DISTRIBUTION: Mainly eastern rainforest: Eastern Red Length: 31–41 cm | 12–16" Forest Rat N. rufus 1b at middle and high elevations Weight: 125–250 g | 4·4–8·8 oz and Lowland Red Forest Rat N. audeberti 1c at lower elevations. The rare Western Red Forest Rat N. lambertoni 1a is found locally in dry forest in the west. ID: Always have redder tones to their fur than other rats. BEHAVIOUR: Diurnal and exclusively terrestrial, feeding on seeds and fruits on the forest floor. WHERE TO SEE: Eastern Red Forest Rat occurs in Ranomafana, Andasibe-Mantadia and Marojejy NPs. Lowland Red Forest Rat can be seen in the lower parts of Ranomafana NP, and in Masoala NP. The uncommon Western Red Forest Rat is restricted to Tsingy de Bemaraha NP.

Tuft-tailed rats Eliurus spp. [12 species]

Sokitralina, Voalavon’ala F Rat à Queue Touffue*

DISTRIBUTION: Found in forest throughout. Length: 22–38 cm | 9–15" Most species are endemic to eastern rainforest, with a few Weight: 20–125g | 1–4·4 oz extending to the north, and two restricted to the west. ID: Long tail (longer than body), with a conspicuous tuft towards the end. BEHAVIOUR: All species are nocturnal, and most are strictly or mainly arboreal. WHERE TO SEE: Western Tuft-tailed Rat E. myoxinus 2a is the species most frequently seen during night walks in Kirindy Forest and Ankarafantsika NP. Tuft-tailed rats are also occasionally seen in eastern rainforest parks. Look for eyeshine in the forest, but much fainter than that of nocturnal lemurs.

Giant Jumping Rat Hypogeomys antimena

M Vositsy Rat Géant Sauteur

DISTRIBUTION: Small area of western dry forest north ENDANGERED of Morondava. Length: 54–59 cm | 21–23" ID: This large and unmistakable species is the flagship Weight: average 1·2 kg | 2·6 lb rodent of Madagascar. It is more like a rabbit than a typical rat: grey-brown in colour and with big ears. BEHAVIOUR: Nocturnal. Walks on forest floor, feeding on plant material. One or two young are born in the wet season. WHERE TO SEE: Can be found only in Kirindy Forest, often in the camp itself. Becomes active about three hours after sunset, later than most nocturnal mammals. Western Big-footed Mouse Macrotarsomys bastardi (not illustrated) is an endemic rodent that is occasionally seen in western forests, particularly Ankarafantsika NP and Kirindy Forest. It looks and acts like a gerbil, hopping along the ground in a manner resembling a mini kangaroo.

Introduced Rodents Black Rat Rattus rattus is found throughout, sometimes even in the forest. Brown (or Norway) Rat R. norvegicus (not illustrated) is more local, mainly found in towns. House Mouse Mus musculus (not illustrated) is found throughout, in association with humans, as well as in marshy areas.

Dark reddish-brown

Black Rat

The reddest of the three species

Dark brown, with a bushy tail 3

Rabbit-like ears

Cetacea: MARINE MAMMALS

Madagascar is not especially rich in marine mammals, but it is a renowned place to see Humpback Whale, and dolphins are sometimes sighted. 1

Humpback Whale Megaptera novaeangliae

DISTRIBUTION: Found in oceans almost worldwide. One population that spends the summer in cold Antarctic waters migrates thousands of kilometres every year, visiting the Madagascar coast to breed and calve during the austral winter, from May to December. The Bay of Antongil is used in July and August as a major breeding and calving site. ID: A huge marine mammal with a humped back, short stubby dorsal fin and distinctive long flippers that are mostly white. Sightings of other whales are rare in Malagasy waters. VOICE: Renowned for its complex, ever-changing undersea song. BEHAVIOUR: One of the most demonstrative whales due to its frequent breaching, tail and flipper slapping, and bubble-feeding behaviours. Feeds on krill and small fish. The female gives birth to a single calf. 2

Length: Weight:

12–16 m | 39–52' 20,000–36,000 kg | 50,000–80,000 lb

WHERE TO SEE: Possible anywhere around the coast, especially to the east between May and December. St. Marie Island hosts a large whalewatching industry, with July to September being the best time. The Bay of Antongil (bounded by the Masoala Peninsula) is less developed for whalewatching, but sightings are also dependable there.

M Feso Grand Dauphin / Grand Dauphin de L’Océan Indien

Common / Indo-Pacific Bottlenose Dolphin Tursiops truncatus / aduncus

M Trozona Baleine à Bosse

DISTRIBUTION: Found in warm and temperate oceans Length: 2–4 m | 7–13' worldwide. Occurs all around Madagascar’s coast. These Weight: 150–650 kg | two species were recently split, and the identity of the 330–1,430 lb dolphins in Malagasy waters remains to be determined. WHERE TO SEE: Possible ID: A large, chunky, generally uniformly greyish dolphin anywhere around the coast, with a sturdy, bottle-like beak. This classic, familiar but most frequently seen in the species accounts for the vast majority of dolphin sightings Nosy Be area. around Madagascar. Spinner Dolphin Stenella longirostris (not illustrated) is occasionally seen, usually far offshore; it averages smaller and slimmer, with a much longer beak; and often spins when it leaps out of the water. VOICE: Uses clicks to echolocate prey. BEHAVIOUR: A social animal that lives in pods, normally containing up to 15 individuals, but sometimes comprising hundreds. Active and playful, and makes spectacular leaps out of the water. Feeds mainly on fish, krill, and squid. Females give birth to a single calf.

Very demonstrative, often breaching or slapping tail or flippers on the surface

Each individual has distinctive tail markings

Short, stubby dorsal fin

1 2

Dolphins often leap out of the water

Birds Almost 300 species of bird have been recorded in Madagascar. Of those, about 260 occur regularly, 100 are endemic, and another 30 are Malagasy regional endemics. Four bird families are endemic to Madagascar, and a further two endemic to the Malagasy region. l Birds have a much greater ability to cross water barriers than groups like reptiles and amphibians. The modern Malagasy avifauna is the result of many ‘colonization events’, from Africa, Asia and probably even Australasia. Some of these events were ancient, while others were much more recent. Only the recently extinct elephant birds pre-dated the breakup of Gondwanaland; all other bird lineages have colonized the island subsequently. l Several birds have reached Madagascar with human assistance, namely Feral Pigeon (page 120), Common Myna (page 176) (unfortunately now the island’s most common bird), House Crow Corvus splendens (not illustrated: so far restricted to Toamasina) and House Sparrow Passer domesticus (not illustrated: still not widespread). l One of the notable things about Malagasy birds is their relative silence compared to birds elsewhere. Few species have particularly complex or musical songs, and many species simply do not vocalize very often. Lemurs and frogs are usually the predominant voices in Malagasy forest. l Birds are generally common and conspicuous in Madagascar. Unfortunately, there are vast deforested stretches of the country with only a handful of native species, plus the ever-present introduced Common Myna. Although rainforest birding is always difficult, Madagascar’s rainforest birds tend to be more approachable and less shy than rainforest birds elsewhere in the world. To find retiring forest specialties like mesites and groundrollers, the best strategy is to enlist the services of guides who specialize in birding. l Some Malagasy birds breed year-round, but most do so during the wet season. As such, they are most vocal between October and December in the rainforest, and slightly later in the dry forest. l Migration is inconspicuous on Madagascar, even compared to mainland Africa. There are a few species, such as Madagascar Pratincole (page 114), Madagascar Cuckoo (page 126) and Broad-billed Roller (page 140) that breed on the island then migrate to the mainland during Madagascar’s dry season. Madagascar also hosts some birds from Europe and Asia, predominantly shorebirds, that migrate south to escape the northern winter. l Bird identification on Madagascar is generally straightforward, as there is not the diverse range of drab birds found in most of the world’s tropics. Couas, like these Giant Couas, are members of the cuckoo family.

l Shorebirds and terns can

be difficult to identify, but their diversity is relatively low in Madagascar, reducing the chances of confusion. l Among forest birds, the only tricky ones are the warbler-like birds, most of which are best found and identified by song. l Sadly, Madagascar’s biggest and perhaps best birds have already become extinct. There were at least five species of elephant bird, one of which, Aepyornis maximus, was the largest bird that the world has ever known. It stood about three metres (ten feet) tall and weighed as much as 400 kg (880 lb). Its enormous eggs were larger even than those of the largest dinosaurs, and the largest single cell known to biology. l The arrival of humans seems to have doomed the elephant birds, along with all of the larger mammals such as pygmy hippopotamuses and ‘gorilla’ lemurs, although they probably survived until well after Madagascar’s first contact with Europe. Sturdy fragments of elephant bird eggs can still be found on the beaches of the southwest, a sad reminder of Madagascar’s lost biological riches.

Endemic bird families: Madagascar

Mesites (page 102).

Asities (page 148).

Bernierids (page 162).

Ground-rollers (pages 142–147). Colourful birds that can easily be overlooked by those who are not prepared to search for them.

Strange, primitive, rail-like, ground-dwelling birds.

A family of cryptic species, the existence of which has only recently been recognized.

A small family of gem-like birds. Also a primitive group.

Endemic bird families: Malagasy Region

Cuckoo-roller (page 146).

One species, found on both Madagascar and the Comoros. It is a strange and enigmatic bird whose loud calls often dominate the rather quiet Malagasy forest.

Vangas (pages 166–175). Madagascar’s counterpart of Darwin’s finches, this family has close ties to some African groups and has evolved to exhibit an incredible diversity on Madagascar. 83

DUCKS and GREBES 1

Little Grebe Tachybaptus ruficollis

DISTRIBUTION: Europe, Afrotropics, and much of Asia. Fairly common though local throughout Madagascar on freshwater lakes and ponds, and occasionally rivers. ID: Much smaller than any duck, with a pointed bill. Bright chestnut and dark grey in breeding plumage 1br and dull brownish at other times 1nb . The less common but also widespread, although Vulnerable, Madagascar Grebe T. pelzelnii is smaller, and generally prefers ponds and lakes in more forested environments. In breeding plumage, it has a diagnostic white stripe behind the eye and a pale throat. Very similar to Little Grebe in non-breeding plumage. VOICE: A loud, rattling trill sometimes reveals its presence on a pond before it is seen. BEHAVIOUR: Usually in pairs. Dives to catch fish.

Length:

M Vivy Grèbe Castagneux

23–29 cm | 9–11"

WHERE TO SEE: Generally uncommon, but can be seen on almost any lake or pond. Even in Tana at Lake Alarobia.

Madagascar Grebe has a pale throat and a white stripe behind the eye. 2

Red-billed Duck (Teal) Anas erythrorhyncha

M Menamolotra Canard à Bec Rouge

DISTRIBUTION: Eastern and southern Afrotropics. Length: 43–48 cm | 17–19" Locally common throughout Madagascar on freshwater WHERE TO SEE: Fairly common wetlands, especially marshes and shallow lakes. in Tana, especially at Lake Will use flooded rice paddies, but not as commonly as Alarobia. Often seen on ponds White-faced Whistling-duck. around Isalo and Anjozorobe. ID: Much slimmer than whistling-ducks, with red bill, pale cheeks and dark cap, and a swifter flight. BEHAVIOUR: Feeds on aquatic plants and invertebrates by dabbling. Forms flocks, especially when not breeding.

White-faced Whistling-duck Dendrocygna viduata

M Tsiriry Dendrocygne Veuf

DISTRIBUTION: Central and South America and the Length: 44–48 cm | 17–19" Afrotropics. Locally common throughout Madagascar WHERE TO SEE: Common on freshwater wetlands, including lakes, ponds and rice around Tana, especially at Lake paddies. The most common duck in Madagascar. Alarobia. Also common in the ID: Mostly rufous and dark brown, with a white face and west, especially Ankarafantsika finely barred flanks The superficially similar Fulvous and Tsingy de Bemaraha NPs, Whistling-duck Dendrocygna bicolor (not illustrated) is and around Maroantsetra. golden-brown, lacks the white face, and is much less common. VOICE: Distinctive, insistent, three-part whistle, most often given by birds in flight, sometimes at night. BEHAVIOUR: Social; frequently in flocks. In some areas, roosts during the day and feeds on rice paddies at night to avoid persecution. Feeds mostly on aquatic plants. 84

1nb

1br 1br Duck-like, but smaller than any duck

Mostly brown, with a red bill 2

Colourful, with a white face and dark bill

GAMEBIRDS

Madagascar Partridge

Margaroperdix madagarensis

DISTRIBUTION: Endemic to Madagascar, and introduced to Mascarenes. Widespread but uncommon and local in grassland, scrub and other open habitats. ID: The beautiful rufous and grey, speckled and spotted male is unmistakable; female much more drab; its larger size and more powerful flight distinguish it from Madagascar Buttonquail. BEHAVIOUR: Generally shy, although it will occasionally walk onto roads and into other open areas, especially at dawn and dusk. Most often seen when flushed into explosive and startling flight. R

Length:

24–26 cm | 9–10"

Where to see: Widespread but never common. Sometimes in the grasslands around Isalo, Anjozorobe and Tsingy de Bemaraha NP. Occasionally even along the runway at the Ivato Airport in Tana.

Madagascar Buttonquail Turnix nigricollis

DISTRIBUTION: A regional endemic found on Madagascar and Réunion. Found in most habitats except rainforest interior and wetlands. Most common in grassland, dry scrub, spiny forest and western dry forest. ID: Tiny size and quail-like shape are distinctive. Male is pale brown, while the female is much more colourful, with a black breast patch and a rufous side to the neck. VOICE: Quiet, low-pitched, pulsing “huhuhuhuhu…”. BEHAVIOUR: Runs on the ground. Often feeds by rotating and kicking away the leaf-litter, leaving clean circles of bare earth. The male incubates the eggs and cares for young.

M Tsipoy Perdrix Malgache

M Kibobo Turnix Malgache

Length:

14–16 cm | 6"

WHERE TO SEE: Possible anywhere, but most common in western dry forest (as at Ankarafantsika NP and Kirindy Forest) and spiny forest.

Although they look like quails, buttonquails are in a completely different family whose closest relatives are still not certain. They have recently been considered to be relatives of the cranes, but may actually be most closely related to gulls and sandpipers. Buttonquail feeding circles 3

Helmeted Guineafowl Numida meleagris

M Akanga Pintade de Numidie

DISTRIBUTION: Most of the Afrotropics. Length: 55–60 cm | 22–24" Thinly distributed throughout Madagascar in a wide Where to see: Generally variety of habitats, including savannah, scrub, forest edge, uncommon, but can be seen forest and marsh. Probably colonized Madagascar without almost anywhere. Frequently human assistance. sighted around Ankarafantsika ID: A large, distinctive and familiar bird with a colourful and Tsingy de Bemaraha NPs, bare head and copious small white spots on the body. Berenty, and the lodges on the VOICE: A loud series of harsh clucks that speeds up then slows Masoala Peninsula. down: a classic sound of the African bush. BEHAVIOUR: Highly social and usually found in flocks. Roosts in tall trees at night. Feeds on seeds, small fruits and invertebrates. The Malagasy name for guineafowl (Akanga) seems to have come from Kiswahili (Kanga). 86

1♂ 1♀

Male colourful 2♀

Female colourful

Female brownish 2♂

Male brownish

Heavily spotted

HERONS I

Grey Heron Ardea cinerea firasa

DISTRIBUTION: Most of Europe, Africa and Asia. In Madagascar mostly along the west coast, in a variety of both salt and freshwater wetlands. Uncommon in the interior, sometimes on rice paddies. ID: Tall, long-necked bird with grey wings and pale head and underparts. Larger and paler than Purple Heron (page 90) and slightly smaller and paler than Madagascar Heron. VOICE: A loud and grumpy “graaak” mainly given in flight. BEHAVIOUR: Usually solitary. Stands motionlessly or slowly stalks prey, which includes fish and almost any other animal.

Length:

0·9–1·0 m | 3’

WHERE TO SEE: Most easily found in the west, as in wetlands around Morondava and Toliara.

Madagascar (Humblot's) Heron Ardea humbloti

DISTRIBUTION: Rare and local in Madagascar and the Comoros. Mainly coastal, although very rare on most of the east coast. Uses both salt and freshwater wetlands. ID: Large size, massive bill and dark grey colour. Darker and more uniform than Grey Heron. VOICE: Similar to Grey Heron. BEHAVIOUR: Similar to Grey Heron; sometimes shows hostility to that closely related species.

M Vanokasira Héron Cendré

M Vorompasika Héron de Humblot

ENDANGERED Length:

1·0–1·1 m | 3·3–3·6’

WHERE TO SEE: Most common in the west, such as wetlands in and around Ankarafantsika NP and coastal flats around Toliara. Occasionally seen at Lake Alarobia in Tana.

Madagascar Heron (left) with Grey Heron (right); on average, Madagascar Heron is slightly larger

Pale grey and white, with black marks on neck and head

Dark grey

HERONS II

Purple Heron Ardea purpurea madagascariensis

M Vanomena Héron Pourpré

DISTRIBUTION: Much of Europe, Africa and Asia. Length: 78–86 cm | 2·6–2·8’ Found throughout Madagascar in small numbers, WHERE TO SEE: More preferring freshwater marshes with dense reedbeds. widespread than the previous ID: Medium-sized heron that is mostly slate grey, with a two herons. Can be seen in heavily striped neck. Smaller than Grey and Madagascar small numbers in wetland Herons (page 88). habitat almost anywhere on the VOICE: Similar to Grey and Madagascar Herons. island. BEHAVIOUR: Mainly solitary, waiting motionlessly before stabbing prey, which comprise a wide range of fish, invertebrates, frogs and lizards. 2

Black Heron Egretta ardesiaca

M F

Lombokohomana Aigrette Ardoisée

DISTRIBUTION: Afrotropics. Locally fairly common Length: 50–60 cm | 1·7–2' throughout Madagascar, mainly in freshwater wetlands. WHERE TO SEE: Fairly common Often feeds in rice paddies. around Tana, and throughout ID: Small and compact all-black heron with bright yellow much of the island. feet. Distinguished from dark morph Little Egret (page 92) by shorter neck and bill, dark throat, and lack of any white in the wing. BEHAVIOUR: Fairly social, and often seen in groups, which form tight packs in flight. Sometimes uses an ‘umbrella’ feeding technique with wings spread over hunched head. This either reduces glare, attracts prey, or both. Preys mainly upon small fish. 2

Black Herons adopting the 'umbrella' feeding technique

Complex pattern on head and neck; favours thick vegetation

Rich coloration visible in flight

Dark and compact

Yellow feet

WHITE EGRETS 1

Cattle Egret Bubulcus ibis

M Vorompotsy Héron Garde-boeufs

DISTRIBUTION: Almost worldwide. Common in Length: 48–54 cm | 19–21" Madagascar in many habitats including dry fields, WHERE TO SEE: Almost cultivated areas and wetland shores. This is the least anywhere. aquatic of Madagascar’s herons and egrets. ID: Smaller and more compact than Little Egret, and much more terrestrial. Also identified by yellow bill, unicoloured legs and buff patches on the head and breast of breeding birds. BEHAVIOUR: Usually in groups, often with livestock. Insects are its main food, so it benefits from the disturbance created by cattle, which flush grasshoppers and other small animals. 2

Little Egret Egretta garzetta dimorpha

M Vanofotsy Aigrette Garzette

DISTRIBUTION: Much of the Old World. Throughout Length: 55–65 cm | 22–26" Madagascar in a variety of salt and freshwater wetlands. WHERE TO SEE: Common at Most common along the muddy shores of the west coast. all coastal sites, and present Dark morph birds generally predominate along the coast throughout the island. while light morph birds are more common inland. The race dimorpha of Little Egret that ocurs in Madagascar is sometimes split as Dimorphic Egret. ID: Light and dark morphs are very different. Light morph can be distinguished from Great Egret by its smaller size, dark bill and legs, and yellow feet. Dark morph differentiated from Black Heron (page 90) by longer bill and neck, and usually a white throat and white patch in the wing. Slaty-grey intermediate birds look unlike other Malagasy egrets. BEHAVIOUR: Can be solitary or in groups. Feeds more actively than most egrets and herons, sometimes dashing around frantically. Fish are its primary food. 3

Great Egret Ardea alba

DISTRIBUTION: Much of the world. Common throughout Madagascar in fresh and saltwater wetlands, including rice paddies. ID: The largest white egret, with dark legs and feet. Never shows head plumes. In flight, wingbeats are slow and long legs trail behind. BEHAVIOUR: Forages alone or in groups, primarily eating fish. Like most egrets and herons, roosts communally with other species.

Size is one of the quickest ways of telling the white egrets apart

Length:

M Vanofotsy Be Grande Aigrette

85–95 cm | 2·8–3·1’

WHERE TO SEE: Common around Tana and throughout the island.

3 2

Facial skin turns bright green when breeding

intermediate

dark morph 3

Dark feet and yellow bill

dark morph

light morph

COMPACT HERONS and HAMERKOP

COMPACT HERONS I 1

Squacco Heron Ardeola ralloides

Mpiandrivoditatatra F Crabier Chevelu

DISTRIBUTION: Europe, Middle East and Africa. Fairly Length: 42–46 cm | 17–18" common throughout Madagascar in freshwater wetlands. WHERE TO SEE: Commonly Sometimes in rice paddies. seen in most wetlands. Large ID: Compact shape, bicoloured bill, and mix of brown breeding colony in Tana at Lake and white plumage. Streaky in non-breeding plumage. Alarobia. The Madagascar Pond Heron A. idae 3 is rare and Endangered, but has been recorded throughout most of Madagascar. It is all-white in breeding plumage and dark brown (rather than buff ) and white in non-breeding plumage. BEHAVIOUR: Secretive and usually solitary. Feeds mainly on fish and insects, which it catches by waiting motionlessly, then striking. e

Striated Heron Butorides striata rutenbergi

Vorompantsika F Héron Strié

DISTRIBUTION: Old World tropics and South America. Length: 40–44 cm | 16–17" Fairly common, though inconspicuous, throughout WHERE TO SEE: Can be found Madagascar in salt and freshwater wetlands. in almost any wetland. ID: Small, compact and dark, with streaks on the breast and a blackish cap. VOICE: Gives a harsh “quaak” call in flight. BEHAVIOUR: Mainly solitary, standing motionless for long periods, waiting for prey such as fish to come close. Easily missed, as it blends into its surroundings.

3br

Lake Alarobia in Tana is a good place to see the rare Madagascar Pond Heron.

1nb

Buff and brown plumage

1br

2im Greenish-grey and brown plumage

Ple b pi wi He

COMPACT HERONS II and HAMERKOP 1

Black-crowned Night Heron Nycticorax nycticorax

M Goaka Bihoreau Gris

DISTRIBUTION: Much of the Americas, Europe, Length: 54–60 cm | 21–24" Asia, and Africa. Locally fairly common throughout WHERE TO SEE: Breeding Madagascar in salt and freshwater wetlands. Needs thick colony at Lake Alarobia in Tana. marsh or swamp areas for roosting. Sightings possible in most ID: Adults are sharply tricoloured: black, grey and white. wetlands. Brown juveniles could be could be confused with Striated or Squacco Herons (page 94), but are larger, with bold spotting on the back. VOICE: Grumpy “goak” given mainly in flight. BEHAVIOUR: Nocturnal. Away from breeding colonies, usually seen roosting motionlessly in thick vegetation during the day, or flying out to feed at dusk. Diverse diet of small animals. Although superficially like a heron, the Hamerkop is taxonomically very different, so much so that it is placed in its own family. e

Hamerkop Scopus umbretta tenuirostris

DISTRIBUTION: Afrotropics. Occurs throughout Madagascar in small numbers in shallow wetlands, including rice paddies. ID: All-brown colour and hammer-headed shape are unmistakable. BEHAVIOUR: Usually solitary or in pairs. Eats amphibians, small fish, invertebrates, and other small animals. Builds huge nests in trees that rank among the largest in the world, and can resemble a pile of material left by a flood.

M Takatra Ombrette Africaine

Length:

50–58 cm | 20–23"

WHERE TO SEE: Possible in rice paddies or other wetlands almost anywhere. Watch especially along the RN7 highway between Fianarantsoa and Antsirabe.

1im

1ju Often seen in flight at dusk and dawn

1ju

2 Hamerkop flies high and sometimes soars

LARGER RAPTORS

Madagascar Buzzard Buteo brachypterus

DISTRIBUTION: Fairly common throughout Madagascar, except in treeless areas. Will feed in open areas as long as there are some trees nearby. ID: A variable brown-and-white raptor, with narrower wings than those of Madagascar Harrier-hawk. Black Kite (page 100) shows a forked tail and less pale coloration. VOICE: Whiny, slightly descending whistle. BEHAVIOUR: Frequently soars, and often perches conspicuously. Hunts birds, small mammals, reptiles and amphibians. E

Madagascar Harrier-hawk Polyboroides radiatus

Length:

M Hindry Buse Malgache

48–51 cm | 19–20"

WHERE TO SEE: Common in many places, although perhaps easiest to find in the west.

M Fihiaka Gymnogène Malgache

DISTRIBUTION: Uncommon throughout the island, Length: 68 cm | 2·3’ except in treeless areas. WHERE TO SEE: Possible ID: Big, long-legged, and broad-winged. The grey-andalmost anywhere in the black adult is distinctive. When excited, the bare yellow proximity of forest, but skin on the face turns pink. The brown immature could be easiest to see in the west, confused with Madagascar Buzzard, but can be recognized such as around Morondava, in by its broad wings, long legs, bare facial skin, and pale head. Ankarafantsika NP, and in the VOICE: Gives a whistle similar to that of Madagascar Buzzard, spiny forest around Toliara. but longer and higher-pitched. BEHAVIOUR: Frequently soars. Unique highly flexible knee joint allows it to insert its legs into crevices and cavities to extract prey. Its varied diet includes eggs, nestling and adult birds, lemurs, rodents, tenrecs, reptiles and amphibians.

Madagascar Fish Eagle Haliaeetus vociferoides

M Ankoay Pygargue Malgache

DISTRIBUTION: Rare and local along the west coast of CRITICALLY ENDANGERED Madagascar, sometimes up to 100 km (60 miles) inland. Length: 70–80 cm | 2·3–2·7’ Uses both saltwater and freshwater wetlands with adjacent WHERE TO SEE: Lake Ravelobe forest (including mangrove and gallery forest). in Ankarafantsika NP and the ID: Large brown eagle with broad wings and a short white Manambolo River in Tsingy de tail. Immature is buff and brown, and lacks white tail of Bemaraha NP. Sightings are also the adult. Generally rare, and unlikely to be seen except in a possible at the Tsiribihina River few select locations. north of Morondava and around VOICE: Very vocal and has an evocative call more like that Nosy Be. of a gull than a raptor. A ringing “whee weet-weet-weet”. The male has a higher-pitched voice than the female. BEHAVIOUR: Usually seen in pairs. Sometimes soars. Feeds almost exclusively on fish, which are caught by swooping down and plucking them from just below the water’s surface. Madagascar Fish Eagle is Critically Endangered – only about 120 breeding pairs remain.

Brown and white, with feathered face

3 1

2im Long legs and bare facial skin

Madagascar's largest raptor

SMALLER RAPTORS 1

DISTRIBUTION: Most of Europe, Asia, Australasia and Africa. Common throughout Madagascar in most habitats except dense forest. Often seen around towns. ID: Dark brown, medium-sized raptor with a distinctive forked tail. VOICE: Quavering whistles, often given in flight. BEHAVIOUR: Usually seen on the wing, often soaring. Feeds mostly by scavenging, but also hunts small animals. R

Frances’s Sparrowhawk

Length:

50–58 cm | 20–23"

WHERE TO SEE: The most common raptor in Madagascar. Visitors will see it many times while travelling around the country.

Accipiter francesiae francesiae

Papango Milan Noir

Black (Yellow-billed) Kite Milvus migrans

M Firasamadinika Épervier de Frances

DISTRIBUTION: Madagascar and the Comoros. Length: 30–35 cm | 12–14" Found throughout the island in almost any habitat with WHERE TO SEE: Throughout, woody vegetation; fairly common but inconspicuous. but most easily found in the ID: Typical sparrowhawk shape: long tail and short, more open forest of the west. rounded wings. The grey back and pale underparts of the male are distinctive. Female distinguished from uncommon Banded Kestrel Falco zoniventris (not illustrated) by slim shape, longer legs, and fine barring underneath. Madagascar Sparrowhawk A. madagascariensis (not illustrated) can be extremely similar, but is rarely seen. Henst’s Goshawk A. henstii (not illustrated) is much larger, and also much less frequently seen. VOICE: Single raspy “kreek” often repeated over and over. BEHAVIOUR: Fairly shy, secretive and solitary. Often perches inside the forest, where it hunts for reptiles, birds and insects. R

Madagascar Kestrel Falco newtoni newtoni

M Hitsikitsika Faucon de Newton, Crécerelle Malgache

DISTRIBUTION: Madagascar and Seychelles (Aldabra). Length: 25–30 cm | 10–12" Common in small numbers throughout Madagascar in WHERE TO SEE: Common a variety of open habitats. Often nests on buildings in throughout the island. Nests on human settlements. buildings in Tana. ID: Quite similar to the world’s widespread kestrel species, American Falco sparverius and Common Kestrels F. tinnunculus (neither illustrated), one of which is likely to be familiar to most visitors. Long tail, pointed wings, and variable buffy colour are distinctive. VOICE: A quick series of high-pitched and squeaky notes. BEHAVIOUR: Often perches conspicuously on poles, wires and branches. Capable of quick and powerful flight, and also hovers skilfully. Hunts lizards and insects. There are other falcons on Madagascar (not illustrated): the rare Banded Kestrel, the scarce resident Peregrine Falcon F. peregrinus; and two migratory species, Sooty Falcon F. concolor and Eleonora’s Falcon F. eleonorae, the majority of which arrive in December and depart by May, a period during which few tourists visit the island.

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1 Madagascar's most common and widespread raptor

2♂

2♀

3♀

pale morph

dark morph

In flight, Madagascar Cuckoo (page 126) looks very similar to a small sparrowhawk or falcon. However, the cuckoo is slimmer than any raptor, and has more pointed wings than a sparrowhawk. 3♂

pale morph

SHY BIRDS OF THE FOREST FLOOR

Madagascar Crested Ibis Lophotibis cristata

M Akohonala Ibis Huppé Malgache

DISTRIBUTION: Uncommon and local in extensive NEAR THREATENED patches of rainforest or western dry forest, though most Length: 50 cm | 20" common in eastern rainforest. Prefers wet areas within WHERE TO SEE: Finding this forest, such as swamps and stream banks. shy bird takes some luck, and ID: Diﬃcult to see, but the brown body, white wings, pale usually the help of a skilled crest and red bare facial skin make this bird unmistakable. local guide. Andasibe-Mantadia, Often only glimpsed in dense forest as a blur of bright white Amber Mountain and Masoala wings flying away. NPs are the best sites. VOICE: Very vocal and loud, especially at dusk and sometimes during the night. Gives a variety of yelping and whining notes. Calls often become quite repetitive, emphatic and plaintive, especially when a pair is calling together. BEHAVIOUR: Very shy. Usually found in pairs. Feeds on invertebrates and small frogs and reptiles on the forest floor. Roosts and nests high in trees. Mesites constitute a family that is found only in Madagascar. They are unusual, primitive birds, so different from anything else that they may deserve to be placed in their own order; they are therefore of considerable taxonomic significance. E

White-breasted Mesite Mesitornis variegata

DISTRIBUTION: A rare endemic whose presence has been confirmed in only a few scattered sites, most of which support dry forest habitat. ID: Strange rail-like bird with spotted underparts and striped face. Unlikely to be seen to be seen unless you are looking for it. VOICE: Usually found after being heard: a loud, elaborate duet of rasping and piping notes. This duet has a whirling, frantic, up-and-down sound to it. When heard up close, it attracts the attention even of forest visitors who are not looking for birds. BEHAVIOUR: Pairs and family groups walk on the forest floor with heads bobbing and tails wagging. When severely startled, flies into mid-storey of forest, perches with tail cocked up and head down, and remains motionless until the threat has passed. There are two other mesite species (neither illustrated). Brown Mesite Mesitornis unicolor is found in the eastern rainforest, and can be seen with some effort and the help of a guide in Ranomafana and Masoala NPs. Subdesert Mesite Monias benschi is restricted to spiny forest, and can be seen near Ifaty.

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M Agolinala Mésite Variée

VULNERABLE Length:

31 cm | 12"

WHERE TO SEE: With the help of a good local guide, usually not difficult to find in Ankarafantsika NP or Kirindy Forest. Also possible but less common in Ankarana NP.

Builds a very large stick nest

White wings

Walks along forest floor

RAILS and JACANA

R e

White-throated Rail Dryolimnas cuvieri cuvieri

M Tsikoza Râle de Cuvier

DISTRIBUTION: Madagascar and Seychelles (Aldabra). Length: 31 cm | 12" Fairly common although inconspicuous throughout WHERE TO SEE: Common but Madagascar in most wetland habitats, including rice shy. Andasibe-Mantadia and paddies, moist forest and scrub, and mangroves. Ankarafantsika NPs and Berenty ID: Shape, rich brown head and underparts, and are some of the best sites. white throat are distinctive. The juvenile shows duller coloration, but still has a white throat. VOICE: A series of loud ascending cries, often given in duet. Also makes deep, grunting calls. BEHAVIOUR: Usually found in pairs. Fairly shy, but will sometimes walk into the open, often flicking its tail up as it walks. Diet is mainly invertebrates. 2

Common Moorhen Gallinula chloropus

M Akohondrano Gallinule Poule-d’eau

DISTRIBUTION: Found throughout much of the Old Length: 30–36 cm | 12–14" World. Locally fairly common across Madagascar, mostly WHERE TO SEE: Easily seen in freshwater marshes. at Lake Alarobia in Tana, and ID: Grey, rail-like bird with a mostly red bill and yellow in Andasibe-Mantadia and legs. Immature browner with a dark bill and legs. Ankarafantsika NPs. VOICE: Creaking and hooting calls which are distinctive once you know them. BEHAVIOUR: Feeds while walking across wetland vegetation, or swimming like a duck. Eats a wide variety of plant and animal matter. E

Madagascar Jacana Actophilornis albinucha

M Mokimbo Jacana Malgache

DISTRIBUTION: Scarce and local, mainly in western NEAR THREATENED Madagascar, in freshwater wetlands with extensive floating Length: 30 cm | 12" vegetation, especially water lilies. WHERE TO SEE: Wetlands in ID: Adults are black, white and chestnut, with a blue bill. and around Ankarafantsika NP Juveniles are pale on the head and underparts and have and Morondava. a grey bill. The structure of jacanas is distinctive, and in laboured flight the huge feet trail behind. VOICE: A series of short trills. BEHAVIOUR: Usually seen singly or in pairs, and occasionally in small groups. Uses its enormous feet to walk across wetland vegetation. 3

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Usually walks but can swim like a duck

2im Equally comfortable walking or swimming

3ju

Walks on top of marshy vegetation

PLOVERS I

Three-banded Plover

M Monditra Pluvier à Triple Collier

Charadrius tricollaris bifrontatus

DISTRIBUTION: Afrotropics. Uncommon throughout Length: 18 cm | 7" Madagascar on freshwater mudflats, wetland verges and WHERE TO SEE: Perhaps rice paddies. easiest in Ankarafantsika NP, but ID: The two black bands on the chest, one white band watch for it on rice paddies and around the head, and red-based bill are diagnostic. If you other wetlands throughout the see a plover in the interior, it is probably this species. country. VOICE: High-pitched, three-part “tsii-tsii-tsii”. BEHAVIOUR: A typical plover that feeds by alternating stretches of running and stopping, plucking invertebrates from the surface. Some scientists treat the Madagascar Three-banded Plover as a full species, C. bifrontatus, adding yet another avian endemic to the island's impressive tally. e

White-fronted Plover

Charadrius marginatus tenellus

M Fotsihandrina Pluvier à Front Blanc

DISTRIBUTION: Sub-Saharan Africa. Fairly common Length: 16–17 cm | 6–7" on sandy beaches and mudflats all along the coast of WHERE TO SEE: Watch for it on Madagascar. Less common inland, occurring mostly along any coastal beach or mudflat. rivers. ID: Small and pale. Can be separated from Greater Sand Plover (page 108) by its smaller size and pale collar. VOICE: Quiet “whiit”. BEHAVIOUR: Like other plovers, feeds by alternating periods of running and stopping, picking invertebrates from the surface. Often found on sandy beaches frequented by sunbathing tourists

2ju

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Two bands on the chest and one on the head

PLOVERS II 1

Greater Sand Plover Charadrius leschenaultii

Fandiafasikabe F Gravelot

DISTRIBUTION: Breeds in Asia. Winters on coasts of Length: 22–25 cm | 9–10" Africa, Arabia, India, Southeast Asia, and Australia. WHERE TO SEE: Common Found on mudflats and sandy beaches all around the coast on most tidal flats, including of Madagascar. Occurs year-round, but more common those around Diego-Suarez, from October to April. Maroantsetra, and Toliara. ID: In Madagascar, seen mainly in non-breeding plumage, which is nondescript: pale grey-brown on the back, and with white underparts. Compared to White-fronted Plover (page 106), larger, longer-legged, bigger-billed, and lacks a pale collar. The very similar but less common Lesser Sand Plover C. mongolus (not illustrated) has smaller bill and shorter, darker legs, but can be very hard to separate from Greater Sand Plover. BEHAVIOUR: Like other plovers, feeds by alternating stretches of running and stopping, gleaning invertebrates from the surface. 2

Grey (Black-bellied) Plover Pluvialis squatarola

Vikivikivolombatolalaka F Pluvier Argenté

DISTRIBUTION: Circumpolar, breeding in Arctic tundra. Length: 27–31 cm | 11–12" Winters along most of the world’s coastlines away from WHERE TO SEE: Easily seen the far north. In Madagascar, found on mudflats and in in much the same places as other coastal habitats all around the coast year-round, but Greater Sand Plover. most common from October to April. ID: A large plover with long legs and a heavy bill. In Madagascar, seen mostly in plain grey non-breeding plumage. Black ‘wingpits’ in flight are distinctive. Larger than Greater Sand Plover, with a checkered pattern on the back. VOICE: Plaintive, whistled “peeeowee”, frequently given in flight. BEHAVIOUR: Often in loose groups. Like other plovers, feeds with a run-stop action and picks invertebrates from the surface.

2br 2nb

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2nb

1nb

SMALLER, PLAINER SANDPIPERS 1

Common Sandpiper Actitis hypoleucos

M Vikiviky Chevalier Guignette

DISTRIBUTION: Occurs almost throughout the Old Length: 19–21 cm | 7–8" World, either as a breeder, migrant, or wintering bird. WHERE TO SEE: Possible in Non-breeding birds are fairly common throughout almost any wetland. Madagascar, mainly from October to March. Uses a wide variety of salt and freshwater wetlands. ID: Short-legged, medium-sized shorebird. Constantly bobs its tail, and flies in a distinctive weak and fluttering manner. VOICE: High-pitched, three-part “tsii-tsii-tsii”, very like call of Three-banded Plover (page 106). BEHAVIOUR: Usually solitary. Diet mainly invertebrates. 2

Curlew Sandpiper Calidris ferruginea

DISTRIBUTION: Breeds in Arctic tundra, mainly in the Old World. Winters in Africa, south Asia and Australia. In Madagascar, found mainly on mudflats along the west coast, although it can occur anywhere along the coast or inland. Present year-round, but largest numbers occur between September and May. ID: Mid-sized shorebird with a long, drooping bill. Red underparts of breeding plumage are distinctive, but in Madagascar seen mostly in grey-and-white non-breeding plumage. BEHAVIOUR: Often found in flocks. Strong, fast flight. Feeds by probing mudflats with its long bill for invertebrates. 3

Sanderling Calidris alba

M Vikivikimena Bécasseau Cocorli

NEAR THREATENED Length:

18–23 cm | 7–9"

WHERE TO SEE: Fairly common in coastal areas with mudflats, including Toliara and Maroantsetra.

M Agolibe Bécasseau Sanderling

DISTRIBUTION: Breeds throughout much of the Arctic Length: 19–21 cm | 7–8" tundra, and winters along most of the world’s coastlines WHERE TO SEE: Possible away from the far north. Found all the way around the on almost any sandy beach, coast of Madagascar, on sandy beaches and flats; occurs including popular tourist areas year-round, but the largest numbers are present between like Nosy Be and St. Marie September and April. Island. ID: Pale, short-billed sandpiper in non-breeding plumage. Brighter breeding plumage rarely seen in Madagascar. Much more likely to be seen along sandy beaches than Curlew Sandpiper. BEHAVIOUR: Usually feeds and roosts in flocks. Follows the surf, rushing down to forage when a wave retreats, then scurrying back up the beach when the next wave arrives. Feeds mainly on invertebrates. 2

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Shorebirds are often seen in flocks

'Flicking' flight with white wing stripe

2nb

Fast flight with white wing stripe and white rump

3nb

3br

Fast flight with white wing stripe and dark rump

LARGER, MORE DISTINCTIVE SANDPIPERS 1

Ruddy Turnstone Arenaria interpres

M Viko Tournepierre à Collier

DISTRIBUTION: Breeds on the Arctic tundra. Winters Length: 21–25 cm | 8–10" along much of the world’s coastline, generally at moderate WHERE TO SEE: Small numbers latitudes in the northern hemisphere. Uses a variety of found in most coastal areas, habitats all along the coast of Madagascar, although it including Toliara, St. Marie Island, prefers rocky shores and reefs. Some remain year-round, Nosy Be and Diego-Suarez. but largest numbers occur from August to April. ID: Distinctive, squat, orange-legged shorebird. In breeding plumage, boldly marked with black, chestnut and white; otherwise more subdued brown and white. VOICE: Short, low-pitched trills. BEHAVIOUR: Often in flocks. Searches for invertebrates by turning over small rocks and other debris with its bill, or by probing into the substrate. 2

Common Greenshank Tringa stagnatilis

M Vikivikipotaka Chevalier Aboyeur

DISTRIBUTION: Breeds in northern Eurasia. Winters Length: 30–34 cm | 12–13" in southern Eurasia, Africa, and Australia. Found WHERE TO SEE: Fairly common throughout Madagascar in a wide variety of wetlands, in most coastal areas, including although most common on the coast. Some can be Diego-Suarez, Mahajanga and found year-round, but largest numbers occur between Toliara. September and April. ID: Tall grey-and-white shorebird with a slightly upturned bill. VOICE: Distinctive three-part “tew-tew-tew”, most often given in flight. BEHAVIOUR: Often in small groups. Feeds actively in shallow water or along water’s edge. Eats invertebrates, small fish and tadpoles. 3

Whimbrel Numenius phaeopus

M Koikoika Courlis Corlieu

DISTRIBUTION: Breeds in tundra all around the Arctic. Length: 40–43 cm | 16–17" Winters along much of the world’s coastline, although WHERE TO SEE: Common in in the northern hemisphere only at moderate latitudes. most coastal areas, including Fairly common in a variety of habitats around the coast Diego-Suarez, St. Marie Island, of Madagascar. Some remain year-round, but largest Nosy Be and Toliara. numbers occur from August to April. ID: Big brown shorebird with a long, downcurved bill. VOICE: Rapid, multi-note call with a bubbly, haunting quality. BEHAVIOUR: Often in flocks. Like many shorebirds that occur in mangrove areas, perches in mangrove trees at high tide, when the mudflats are covered. Feeds mainly on crabs by both picking them from the surface and probing into the substrate. A mixed flock of waders: Whimbrel, Common Greenshank, Grey Plover and Little Stint

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1br

2nb

White stripe on back; slightly upcurved bill

White stripe on back; downcurved bill

AQUATIC ODDITIES

Madagascar Pratincole Glareola ocularis

M Vorombato Glaréole Malgache

DISTRIBUTION: Breeds exclusively in Madagascar, and VULNERABLE spends the non-breeding season (May to August) in east Length: 25 cm | 10" Africa. Generally scarce and local, although it uses a variety WHERE TO SEE: Common of different habitats: rocky coast, rivers with exposed rocks on rocks around the Masoala and sand banks, and short grassland. Ranges widely when Peninsula and Nosy Mangabe, feeding and can be see in flight almost anywhere. plus the Manambolo River in ID: This elegant brown, buff and white bird looks halfway between a tern and a shorebird when perched. The belly is white, Tsingy de Bemaraha NP. and there are white patches on the rump and behind the eye. VOICE: Often the first clue to the presence of pratincoles. High-pitched “vik” call, often doubled, tripled, or given in excited chorus by a flock of birds. Pratincoles call frequently when perched, and incessantly in flight. BEHAVIOUR: Often in pairs or flocks. Perches on seaside rock formations, sand banks or rocks in rivers, or on man-made structures such as bridges. Feeds on insects, which are caught on the wing. Its bounding and swooping flight is graceful and powerful. Sometimes flies very high. 2

Crab-plover Dromas ardeola

M Tsakaranta Drome Ardéole

DISTRIBUTION: Western Indian Ocean and Red Sea Length: 33–38 cm | 13–15" coast. Scarce and local non-breeding visitor to Madagascar, WHERE TO SEE: Often seen in although some occur year-round. Uses most coastal Toliara Harbour, on the island of habitats, although it prefers sandy beaches and mudflats. Nosy Ve (southwest of Toliara) ID: An unmistakable large, black-and-white shorebird that and around Nosy Be. is so different from other shorebirds that it is placed in its own family. VOICE: Simple yelps and cries, given mainly in flight. BEHAVIOUR: Usually seen in flocks, sometimes containing dozens of birds. Feeds much like a typical plover, walking along the shore, pausing frequently, and scanning for food. As the name suggests, eats mainly crabs, but also takes other crustaceans.

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Strong black-and-white pattern particularly noticeable in flight

White rump

Rufous underwing

Big, unmistakable shorebird

TERNS 1

Greater Crested (Swift) Tern Thalasseus bergii

Trobakamavovava F Sterne Huppée

DISTRIBUTION: Coast of southern and eastern Africa, Length: 46–49 cm | 18–19" Arabia, India, Southeast Asia and Australia. Breeds only WHERE TO SEE: Possible at locally in Madagascar, but can be seen in small numbers most coastal sites, including anywhere along the coast. Toliara, Mahajanga and Nosy Be. ID: Among Madagascar’s common tern species, this is the largest. Can sometimes be identified at long distances simply by its graceful and powerful flight. When seen at close range, its heavier yellow bill and darker grey back separate it from Lesser Crested Tern. The globally widespread Caspian Tern Hydroprogne caspia (not illustrated) is uncommon in Madagascar, seen mainly along the coast. It is larger, bulkier, shorter-tailed, and has a heavy orange bill. VOICE: Various low-pitched rasping calls, often given in pairs. BEHAVIOUR: A typical tern: hunts on the wing, making shallow plunge-dives into the water to catch fish and other small aquatic animals. Flocks roost on sandbars and rocks, often with other species of terns 2

Lesser Crested Tern Thalasseus bengalensis

M F

Trobakamadinika Sterne Voyageuse

DISTRIBUTION: Similar to Greater Crested Tern, Length: 35–37 cm | 14–15" although also present along the coast of northern Africa. WHERE TO SEE: More common Also more migratory and localized as a breeder; not than Greater Crested Tern, and known to breed in Madagascar. Fairly common all likewise possible at most coastal around the coast as a non-breeding visitor, mainly from sites. November to June, although some remain year-round. ID: Smaller, slimmer and paler than Greater Crested Tern, with a smaller, slimmer, orange bill. Flight is graceful, but not as powerful as Greater Crested Tern. VOICE: Rasping and piping calls, higher-pitched than those of Greater Crested Tern. BEHAVIOUR: A typical tern. More likely to be seen far from shore than Greater Crested Tern. 3

Roseate Tern Sterna dougallii

M Varevaka Sterne de Dougall

DISTRIBUTION: Wide but patchy distribution across Length: 33–38 cm | 13–15" the world’s oceans. Breeds on islands off the Malagasy WHERE TO SEE: Roosting coast. Not commonly seen in most places, but possible on offshore islets around the anywhere, both along the coast and at sea. Masoala Peninsula. ID: Much slimmer than Lesser Crested Tern, with a proportionally longer tail. Also lacks crest and has a black tip to the bill. Common Tern S. hirundo (not illustrated) is a frequent non-breeding visitor to the Malagasy coast. It is less common than Roseate Tern around Masoala, but more common elsewhere. Identification can be diﬃcult, but in all plumages, Common Tern is shorter-tailed, darker grey on the back, and flies with deeper wingbeats than Roseate Tern. VOICE: Two-part “skidgit” call is distinctive. Also gives simpler, lower-pitched calls. BEHAVIOUR: A typical tern in most respects. Flies with shallow wingbeats, unlike the strong, rowing wingbeats of crested terns. Feeds on small fish, exclusively offshore.

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1 3

All three of these tern species show a black forehead in breeding plumage, and a white forehead in nonbreeding plumage.

BIRDS OF THE SEA 1

Lesser Frigatebird Fregata ariel

Bomomandry Frégate Ariel

DISTRIBUTION: Patchily distributed around Pacific, Length: 70–82 cm | 28–32" Indian and Atlantic Oceans. Not confirmed as a breeder in WHERE TO SEE: Frequent Madagascar, but frequently seen along the coast, especially around Nosy Be, especially on in the north and west. Nosy Tanikely. ID: Big black bird with long wings and a jagged shape. Female has a white band across the chest, and male has an inflatable red throat pouch. Great Frigatebird Fregata minor (not illustrated) also occurs but is less common. From a distance it is very diﬃcult to separate the two species, but with a good view Lesser Frigatebird can be identified by the white 'wingpit' markings which are absent in Great Frigatebird. VOICE: Silent at sea. BEHAVIOUR: Spends most of its time on the open sea. Huge wings and long tail allow graceful flight. Feeds mainly on squid and flying fish. Unlike other frigatebirds, it does not regularly pirate food from other birds. 2

White-tailed Tropicbird Phaethon lepturus

DISTRIBUTION: Distributed patchily in tropical seas worldwide. Breeds on rocky coastlines and feeds far out at sea. In Madagascar, it breeds between Nosy Be and DiegoSuarez and is rarely seen elsewhere. ID: Very white and tern-like (page 116) but with longer tail streamers. Tropicbirds fly in a distinctive way: the wings are held straight and the wingbeats are shallow, powering strong and steady flight. VOICE: Quiet “chik”, often given in series. BEHAVIOUR: Usually solitary, or in small loose groups around breeding colonies. Strong and elegant flight, sometimes very high. Feeds by plunge-diving for fish and squid. 3

Red-tailed Tropicbird Phaethon rubricauda

DISTRIBUTION: Warm stretches of Indian and Pacific Oceans. The sole Malagasy breeding colony is on Nosy Ve, off the southwest coast. Occasionally seen from the coast elsewhere. ID: Larger and heavier than White-tailed Tropicbird, with all-white wings, and thin, red tail streamers. VOICE: Croaking calls given around breeding colony. BEHAVIOUR: Flight powerful, but not as elegant as that of White-tailed Tropicbird; somewhat pigeon-like. Feeds mainly on fish, which it catches by diving. Nosy Ve was only recently colonized by Red-tailed Tropicbirds, probably in the late 1970s. This vulnerable colony has been protected by a local fady (taboo). 118

M Kafotsy Phaéton à Bec Jaune

Length:

45–50 cm | 18–20" (without tail strea mers, which are 30–50 cm long)

WHERE TO SEE: Around Nosy Be, especially Nosy Tanikely. Near Diego-Suarez at places often visited on tourist excursions: the ‘Three Bays’ and Nosy Diego on the Emerald Sea.

M Vorompano Phaéton à Brins Rouges

Length:

60–70 cm | 24–28" (without tail streamers, which average 40 cm long)

WHERE TO SEE: Likely to be seen only on Nosy Ve, a small island just off Anakao, and southwest of Toliara.

Long, jagged wings

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Long, forked tail 1â&#x2122;&#x201A;

Yellow bill Black slashes on upperwing

2 White tail streamers 3 Red bill

Thin red tail streamers

DOVES and PIGEONS

Madagascar Turtle Dove Streptopelia picturata

M Domohina Pigeon Malgache

DISTRIBUTION: Throughout the Malagasy region. Fairly Length: 28 cm | 11" common though inconspicuous in all habitats that have WHERE TO SEE: Common in some woody vegetation. most natural and semi-natural ID: A large brown pigeon that is distinctive if not habitats, from spiny forest to colourful. Obvious white corners to the tail in flight. rainforest. Can even be found VOICE: A low-pitched rising “hwooo” or “ho-hwoooo”, repeated in the more vegetated parts of about every 1·5 seconds. The wings flap noisily at takeoff. Tana. BEHAVIOUR: Solitary or in small groups. Shy, and often flushes off roads and paths. Walks along the ground searching for seeds and secondary foods such as fruits, flowers and insects. Fast, powerful flight is generally low, through or just above forest. e

Namaqua Dove Oena capensis aliena

M Katoto Tourtelette Masquée

DISTRIBUTION: Sub-Saharan Africa and Arabian Length: 28 cm | 11" Peninsula. Found in most of Madagascar, although most WHERE TO SEE: Easily seen common in the west, south and far north. Prefers open in the dry west and north, as woodland, scrub, cultivated areas and spiny forest. around Toliara, Morondava, and Rarely found in pure grassland. Ankarafantsika and Ankarana ID: A tiny-bodied, long-tailed dove. The male has a black NPs. face and a red and yellow bill, while the female is more drab. VOICE: A rising “hoowooo”, repeated about every two seconds. Similar to Madagascar Turtle Dove, but much higher-pitched. BEHAVIOUR: Solitary, or in pairs or small groups. Walks on the ground, feeding on seeds. I

Feral Pigeon (Rock Dove) Columba livia

M F

Voromailala Pigeon Biset

DISTRIBUTION: Originally Eurasia and North Africa, Length: 32–34 cm | 13" but widely introduced and found locally in small numbers WHERE TO SEE: Found in most throughout Madagascar, in close association with humans. towns. ID: Extremely variable, ranging from all-white to allblack, but generally grey with green iridescence on the neck and dark bars across the wings. The variation in plumages often gives flocks a ‘patchwork’ appearance. Its association with humans is usually suﬃcient for identification. VOICE: Low-pitched cooing. BEHAVIOUR: Usually in flocks. Roosts and nests mainly in man-made structures. Walks on the ground searching for seeds and scraps left by humans.

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Female Namaqua Dove is plainer than the male

1 White band on end of tail

2♂

2♀

ENDEMIC PIGEONS

Madagascar Green Pigeon Treron australis

M Voronadabo Colombar Maïtsou

DISTRIBUTION: Comoros (Mohéli) and Madagascar. Length: 32 cm | 13" Erratic and perhaps semi-nomadic throughout much of WHERE TO SEE: Easiest to see the island, but rarely above 1,000 m (3,300'), and absent in the west. Usually common from the High Plateau. Uses a variety of woodland and around the headquarters at forest, although usually not closed-canopy forest. Ankarafantsika NP. ID: A large, mostly green pigeon. Distinctive even in Also frequent around Toliara. flight, when green body and black wings with white wingbars are evident. Usually flies higher than the similarly sized, all-brown Madagascar Turtle Dove. VOICE: A rather maniacal, crackling mix of trills, cackles and hoots. BEHAVIOUR: Often in small groups. Arboreal species that feeds mainly on fruit, especially figs. In the east, sometimes found in fruiting trees alongside Madagascar Blue Pigeons. E

Madagascar Blue Pigeon

Alectroenas madagascariensis

DISTRIBUTION: Uncommon, mainly in rainforest in eastern and northern Madagascar. ID: Dark blue, with red bare skin around the eyes and a red tail. In flight, looks shorter-tailed and darker than other pigeons and doves. VOICE: Virtually silent, unusually for a pigeon. BEHAVIOUR: Often in small flocks. An arboreal species that feeds exclusively on fruits. Perches on snags and tall trees.

Length:

M Finengo Founingo Bleu

28 cm | 11"

WHERE TO SEE: Forest edges in and around Andasibe-Mantadia NP. Also quite common in Anjozorobe-Angavo forest. Less common but possible in Ranomafana and Amber Mountain NPs.

Tale of two pigeons Madagascar’s two native pigeons exemplify the varied origins of its biological riches. Green pigeons are found in both Africa and Asia, but the Madagascar Green Pigeon is closely related to the African Green Pigeon Treron calvus, and it seems clear that Madagascar was colonized from nearby Africa in this case. Blue pigeons on the other hand are found only in the Malagasy region. Their closest relative seems to be the Cloven-feathered Dove Drepanoptila holosericea, found on the island of New Caledonia, 1,200 km east of Australia in the Pacific Ocean. Despite having managed to arrive from Oceania and thrive on the remote opposite side of the Indian Ocean, blue pigeons never successfully colonized the African mainland.

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PARROTS

Vasa parrots have one of the strangest breeding systems among birds. Males are among a small group of birds that develop a phallus and mating takes place over protracted periods – as opposed to the 'blink-and-you-miss-it' copulation typical of most birds. Mating can be highly social, with large and demonstrative groups of birds gathering around a courting and mating pair. The larger females are dominant, and both males and females have multiple partners. One male will bring food to the nests of several females. R e

Lesser Vasa Parrot Coracopsis nigra

M Bolokikely Perroquet Noir

DISTRIBUTION: Comoros and Madagascar. Fairly Length: 35 cm | 14" common in all types of forest throughout the island. WHERE TO SEE: Generally Generally more common in eastern rainforest than common, but perhaps most Greater Vasa Parrot. conspicuous in western dry ID: Vasa parrots are distinctive as a group but separation forest, as at Ankarafantsika NP of the two species is diﬃcult. Lesser Vasa Parrot is 20–25% or Berenty. smaller than Greater Vasa Parrot, but size can be hard to judge. Lesser Vasa Parrot has a smaller head and bill. Details of the bare parts are also useful: Lesser Vasa Parrot has less bare skin around the eye, a crooked line of bare skin at the back of the upper mandible, and a darker bill; Greater Vasa Parrot has extensive bare skin around the eye, a straight line of bare skin at the back of the upper mandible, and a very pale bill. VOICE: Highly vocal, with a wide repertoire. The most distinctive vocalization is the song, a series of melodic whistles that first descend then ascend. Also gives simpler and rougher calls. BEHAVIOUR: Often in groups. Nests in cavities. Feeds on fruit.

Greater Vasa Parrot Coracopsis vasa

M Bolokibe Perroquet Vaza

DISTRIBUTION: Comoros and Madagascar. Found Length: 50 cm | 20" primarily in forest, but more tolerant of human-modified WHERE TO SEE: Easiest to areas than Lesser Vasa Parrot. see in western dry forest, as at ID: See Lesser Vasa Parrot. The female Greater Vasa Parrot Ankarafantsika and Tsingy de becomes completely or partially bald when rearing chicks, Beharaha NPs or Berenty. so any parrot with a bare head is a Greater Vasa Parrot. VOICE: Calls generally louder and harsher than those of Lesser Vasa Parrot. BEHAVIOUR: Similar to Lesser Vasa Parrot. E

Grey-headed Lovebird Agapornis canus

M Sarivazo Inséparable à Tête Grise

DISTRIBUTION: A Madagascar endemic that has been Length: 15 cm | 6" introduced to the Comoros. Common in the north, west, WHERE TO SEE: Common and south in a variety of open habitats, both natural and in most dry lowland sites, human-modified. Uncommon in the east in deforested including Ankarafantsika NP, areas. Absent from the High Plateau, but escaped cage Kirindy Forest, the Toliara area birds can be seen anywhere. and Berenty. ID: A tiny, short-tailed parrot with bright green plumage. VOICE: Short, dry “prrrtt”, mainly given in flight. BEHAVIOUR: A highly sociable bird that is always found in pairs or flocks. Flight is very fast and direct. Nests in cavities. Feeds on seeds and is considered a pest in some agricultural areas. 124

Lesser Vasa Parrot: smaller bill, more 'in front' of head; less bare skin around eye;

Greater Vasa Parrot: larger bill, in line with head; more bare skin around eye;

bare skin at the back of the upper mandible is jagged;

bare skin at the back of the upper mandible is straight;

Can resemble a raptor in flight

2 Vasa parrots show iridescent highlights in bright light

Very fast and direct flight

3♂

3♀

3♂

CUCKOOS

R e

Madagascar Coucal Centropus toulou toulou

M Toloho Coucal Toulou

DISTRIBUTION: Seychelles (Aldabra) and Madagascar. Length: 45–50 cm | 18–20" Fairly common although inconspicuous in any habitat WHERE TO SEE: Almost with thick cover, including scrub, marsh and forest. anywhere, even in Tana. ID: A large, rufous-and-black bird with a heavy bill. The scientific, Malagasy and Floppy flight with long tail and short wings is distinctive. VOICE: Liquid, descending series of hoots, often likened to water French names all reflect the coucal’s ‘water bottle’ call. glugging out of a bottle. BEHAVIOUR: Sometimes perches up, but more often creeps through the undergrowth, almost like a reptile. Eats almost any animal it can catch and swallow.

Madagascar (Lesser) Cuckoo Cuculus rochii

M Taotaonkafa Coucou Malgache

DISTRIBUTION: Breeds exclusively on Madagascar, from Length: 28 cm | 11" August to April, when it is often heard, although can be WHERE TO SEE: Widespread hard to spot. Most migrate to Africa after breeding. Found and common, but easiest to in all habitats with a woody component. see in the more open western ID: When perched, grey colour, barred underparts forest, as at Ankarafantsika NP, and horizontal posture are distinctive. Strong flight, Kirindy Forest, and in the spiny long tail and pointed wings make it look very like a falcon or forest around Ifaty and Toliara. sparrowhawk when flying. VOICE: A whistled, descending, three- or four-note “hew hew tew tew”. It sings incessantly day and night from October to January – one of the most frequently heard and distinctive natural sounds of Madagascar. BEHAVIOUR: Generally shy, but often sings from a prominent perch. A brood parasite that lays its eggs in the nests of many species of smaller birds. Eats insects, mainly caterpillars. E

Crested Coua Coua cristata

M Tivoka Coua Huppé

DISTRIBUTION: The most widespread coua, found in Length: 42 cm | 17" forest throughout Madagascar. It is most common in dry WHERE TO SEE: Easiest in dry forest, and tolerates degraded forest. forest – Berenty, around Toliara, ID: Floppy crest and arboreal habits usually identify it Ankarafantsika and Ankarana easily. Mostly grey, with a buff chest, white belly and NPs and Kirindy Forest. colourful two-tone bare skin around the eye. VOICE: Highly vocal, with many different calls. The most common is a doubled, wooden “touk-touk”. Also gives a long series of piercing ‘laser gun’ calls. BEHAVIOUR: This is the coua most likely to be seen bouncing around high in trees. More sociable than other couas, often in groups. Eats small animals and fruit. 1

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Coucals act almost like mammals, creeping through the undergrowth and running along the ground.

1nb

1br

3 Colourful bare facial skin typical of couas

COUAS of the DRY WEST & SOUTHWEST

Couas are members of the cuckoo family. They form a genus, or perhaps even a sub-family of birds that is found only in Madagascar. Their bold patterns of bare facial skin and exuberant treebouncing or ground-strutting behaviours make them favourites with many visitors. E

Red-capped Coua Coua ruficeps

M Aliotse Coua à Tête Rousse

DISTRIBUTION: Two distinctive subspecies: one with a Length: 42 cm | 17" red cap, and one with an olive-brown cap. ‘Red-capped’ WHERE TO SEE: Common is in western Madagascar in dry forest. ‘Green-capped’ is in Ankarafantsika NP. For the found in spiny forest and other dry woody habitats in the ‘Green-capped’ subspecies, try southwest. the Arboretum d’Antsokay in ID: Tall, slim, long-tailed coua. The red-capped subspecies Toliara. overlaps in range with Coquerel’s Coua, but is separated by its red cap, longer legs and slimmer shape. VOICE: Vocal, with several different calls. Most distinctive is series of higher rising notes followed by lower pitched notes: “quee-quee-quee-hm-hm-hm”. Also makes agitated scratchy calls and a simple low-pitched note. BEHAVIOUR: Walks along the forest floor feeding on insects. Often calls from higher perches. E

Coquerel’s Coua Coua coquereli

M Gory Coua de Coquerel

DISTRIBUTION: Uncommon to fairly common in Length: 42 cm | 17" western and far northern Madagascar in dry forest; rare WHERE TO SEE: Fairly and local in the southwest. common in Kirindy Forest and ID: Relatively small, compact, short-legged, brownish Ankarafantsika and Zombitsecoua with a rufous belly and bright blue bare facial skin. Vohibasia NPs. Giant Coua is very similar coloration, but is much larger. VOICE: Song is a single lower note followed by several higher notes: “chwee-quee-quee-quee”. Call comprises two notes: the first high and rising, and the second raspy and low-pitched. BEHAVIOUR: The most terrestrial of the western couas: walks along the forest floor feeding on insects. E

Giant Coua Coua gigas

M Lejabe Coua Géant

DISTRIBUTION: Uncommon in dry forest in much of the Length: 62 cm | 24" south and west, although absent from the northwest. WHERE TO SEE: Easy to see at ID: Well-named, as this is an enormous coua. Like Berenty and Kirindy Forest. Also Coquerel's Coua, has brownish coloration, rufous belly present in Zombitse-Vohibasia and blue facial skin, but is much larger and also shows a NP, although more shy. pink patch behind the eye. VOICE: Loud, resonant, ringing calls. Some are similar to the calls of Coquerel’s Coua, but are usually lower-pitched and louder. BEHAVIOUR: Walks on the forest floor and feeds on insects and small vertebrates. Sometimes goes high into trees to roost or call.

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Often seen on paths, but shies away at the approach of people.

The subspecies in the southwest has an olive cap, and is sometimes split as 'Green-capped Coua' CouaÂ olivaceiceps.

Blue bare skin on face

Similar to Coquerel's Coua, but much larger

Blue and pink bare skin on face

COUAS of the WET EAST & NORTH

Red-fronted Coua Coua reynaudii

M Taitoaka Coua de Reynaud

DISTRIBUTION: Uncommon to locally fairly common in Length: 40 cm | 16" eastern rainforest. WHERE TO SEE: With some ID: Dark grey-green coua with a red forehead. Smaller effort, can usually be found than Red-breasted Coua, with grey rather than dark red in Ranomafana, Andasibeunderparts. Mantadia and Masoala NPs. VOICE: Most frequent and distinctive vocalization is a very long, mechanical rattle, unlike the calls of other couas. BEHAVIOUR: A shy species, found alone or in pairs. Walks on the ground, sometimes along trails, or creeps through thick tangles, sometimes well above the forest floor. Insects are its main food. E

Red-breasted Coua Coua serriana

M Koa Coua de Serre

DISTRIBUTION: Uncommon to locally common in Length: 42 cm | 17" the lush rainforests in the northern half of eastern WHERE TO SEE: Masoala and Madagascar. Marojejy NPs. Also present in ID: Dark coua with red underparts. Andasibe-Mantadia NP, but VOICE: Song is a loud, whistled “koaaaaa”, often followed much harder to find. by a much quieter, low-pitched grumble. Distinctive call is an abrupt “chik-wit”. BEHAVIOUR: Shy species that is found alone or in pairs. Walks on the forest floor, feeding on fruit, insects and small vertebrates. E

Blue Coua Coua caerulea

DISTRIBUTION: Fairly common in eastern rainforest and in some patches of Sambirano forest in the north. ID: A large, dark blue coua, one of the most spectacular Malagasy birds when seen in good light. VOICE: Growls, rasps and explosive short trills. Also gives a more melodic, descending series of notes: “kwee-kwe-kwe-kwe-kwe…”. BEHAVIOUR: An arboreal species, usually found singly or in pairs. More likely to make long flights than most couas, gliding on its short wings and steering with its huge tail. Eats fruit, large invertebrates and reptiles. Last of the giant couas Giant and Blue Couas are big, but Madagascar used to have even larger couas. There are two huge extinct species known from subfossils, plus the Snail-eating Coua C. delalandei that was found on the island of St. Marie until its extinction in the 19th century.

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Length:

M Mariha Coua Bleu

50 cm | 20”

WHERE TO SEE: Usually not difficult to find in Ranomafana, Andasibe-Mantadia and Masoala NPs.

Blue Coua is often found in the canopy, and is seen in flight more often than other couas.

Multi-coloured bare facial skin

Blue bare facial skin

Large and blue

OWLS

White-browed Owl Ninox superciliaris

M Tovotovoka Ninoxe à Sourcils Blancs

DISTRIBUTION: Found in forest and woodland Length: 30 cm | 12" throughout much of the island. Prefers clearings, forest WHERE TO SEE: Local guides edge and scrub. Most common in the southwest and at Berenty and Zombitsenortheast, although still quite local even in those parts of Vohibasia NP often know the Madagascar. location of a day roost. Fairly ID: Medium-sized brown owl with a barred belly. common in Masoala and Kirindy Larger size, dark eyes and barring on the belly separate it from Forest; may be heard at night Madagascar Scops Owl. Darker than Barn Owl, and less likely to from the lodges at those sites. be found in a village or open grassland. VOICE: Distinctive vocalizations are given frequently, including during the day: a rising “hwoooo” and a series of yelps. BEHAVIOUR: Pairs often roost together in a dense tangle of vegetation. Hunts at night, watching from a perch, then swooping down on insects and small vertebrates. 2

Barn Owl Tyto alba

M Tararaka Effraie des Clochers

DISTRIBUTION: Found throughout most of the world, Length: 30–35 cm | 12–14" including all of Madagascar. More common in towns, WHERE TO SEE: Nests in many farms and deforested areas than in natural habitats. Malagasy villages, and even in ID: Medium-sized, grey, buff and white owl with a heartcentral Tana. Sometimes seen at shaped face. a day roost at Tsimbazaza Zoo. VOICE: Unnerving raspy shriek, probably the reason that many Malagasy people consider this owl to be a bad omen. BEHAVIOUR: Hunts small animals, especially mammals. In some cities and villages, subsists mainly on introduced rats and mice. Nests in cavities in rocks, trees and man-made structures. E

Madagascar Scops Owl Otus rutilus

DISTRIBUTION: Fairly common throughout Madagascar Length: in most wooded habitats, from primary forest to heavily disturbed scrub. Two subspecies are recognized: 'western' O. r. madagascariensis (west, south and central) and 'eastern' O. r. rutilus (east). 'Eastern' has a red 3a and a grey-brown morph 3b , 'western' a grey 3c and grey-brown morph 3d . These are sometimes split as full species: Torotoroka Scops Owl in the west and Malagasy (or Rainforest) Scops Owl in the east. ID: Madagascar’s only small owl, which sports distinctive ear-tufts. VOICE: A plaintive series of several hoots that become first louder then softer. In the east, the notes tend to be very pure, while in the west they are slightly trilled. BEHAVIOUR: Roosts in dense tangles of vegetation, often in pairs. Hunts insects at night. WHERE TO SEE: Local guides in Berenty, Kirindy Forest, and Zombitse, Ankarafantsika and Andasibe-Mantadia NPs often know the location of a day roost. At night, can be found in almost any forest.

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M Torotoroka Petit-duc Malgache

22–24 cm | 9" 3d

Often seen roosting in tree cavities and dense tangles during the day

Often found roosting during the day

Heart-shaped face

NIGHTJARS

Madagascar Nightjar

Caprimulgus madagascariensis madagascariensis

M Matoriandro Engoulevent Malgache

DISTRIBUTION: Madagascar and Seychelles (Aldabra). Length: 21 cm | 8" Common throughout Madagascar in most habitats except WHERE TO SEE: Very common closed-canopy forest and treeless grassland. and familiar bird that can be ID: The only nightjar likely to be seen in most of heard and seen almost anywhere Madagascar, usually in graceful, bounding flight, when its on the island at dusk, even on long-tailed, crooked-winged shape is distinctive. Shows buildings in central Tana. Guides white in the wings and tail, and is not found inside thick forest, at Arboretum d’Antsokay in unlike Collared Nightjar. Toliara can sometimes point out VOICE: A highly distinctive call that is one of the most evocative roosting birds. sounds of sounds of Madagascar: a single introductory note followed by an accelerating trill: “tuk t-r-r-r-r-r-r-r-r-r-r”. Most vocal at dusk. BEHAVIOUR: Roosts on the ground during the day, and becomes active at dusk. Feeds on the wing, swooping across open areas and catching flying insects. The Malagasy name for Madagascar Nightjar (Matoriandro) means ‘day sleeper’. E

Collared Nightjar Gactornis enarratus

M Tataroala Engoulevent à Nuque Rousse

DISTRIBUTION: Rare and local in undisturbed eastern Length: 24 cm | 9" rainforest. WHERE TO SEE: AndasibeID: Unlikely to be seen except by those who are Mantadia NP. Among the particularly looking for it. If there is any doubt, the scaly feats regularly performed by back and rufous collar should clinch the identification. Madagascar’s amazing local Madagascar Nightjar does not roost on the floor of thick guides, the ability consistently rainforest. In flight, Collared Nightjar lacks white in the wings to find this cryptic species on and tail. the forest floor during the day is particularly impressive. VOICE: Unusually among nightjars, the song is unknown. BEHAVIOUR: Single birds or pairs roost on or near the forest floor during the day. They are active at night, when they hunt for insects on the wing, inside the forest. Nests in a small palm or fern, up to two metres above the ground. The Collared Nightjar is an unusual and mysterious bird, seemingly more primitive than most other nightjars. Its atypical behaviours include feeding inside the forest and nesting in small trees. The fact that its voice is still unknown is quite amazing.

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Nightjars have bristles around their bill which serve to funnel insects into their mouth when they are feeding aerially at night.

Nightjars often sit in open areas, such as on roads, at night.

Shows white in the wings

Cryptic, but sometimes spotted roosting on the ground during the day.

Often roosts in pairs

AERIAL FEEDERS

African Palm Swift Cypsiurus parvus gracilis

M Manaviandro Martinet des Palmes

DISTRIBUTION: Most of Afrotropics. Fairly common Length: 16 cm | 6" in open habitats below 1,100 m (3,600') in much of WHERE TO SEE: Almost Madagascar, often in association with palms, especially anywhere in the lowlands Bismarck Palm Bismarckia nobilis, Coconut Palm Cocos where there are palms nearby. nucifera and Hyphaene spp. Absent from the High Plateau. ID: Spindly, long-tailed swift with agile but weak flight. Smaller, slimmer, and paler grey than African Black Swift. VOICE: Weak, rattling trills given in flight. BEHAVIOUR: Like all swifts, normally seen only in flight. It eats, drinks and mates on the wing, and perches only near the nests, inside palm trees. e

African Black Swift Apus barbatus balstoni

Sidintsidinamainty F Martinet du Cap

DISTRIBUTION: Scattered areas of sub-Saharan Africa. Length: 16–18 cm | 6–7" The two subspecies in the Malagasy region may prove to WHERE TO SEE: Ranges be full species. Occurs throughout Madagascar, though widely and can be seen almost erratically; perhaps semi-migratory. Often feeds over wet anywhere. Can usually be areas like rice paddies, lakes and the ocean. spotted in the skies over Tana. ID: Big, dark, powerful swift. Much stronger flier than African Palm Swift, and tends to fly higher. VOICE: High-pitched trill given in flight. BEHAVIOUR: As is typical for swifts, usually seen only on the wing, often in flocks, and even seems capable of sleeping in flight! Eats flying insects. R

Mascarene Martin Phedina borbonica

M Sidintsidina Hirondelle des Mascareignes

DISTRIBUTION: Breeds on Madagascar, Réunion and Length: 14 cm | 5·5" Mauritius. Common in Madagascar year-round, but some WHERE TO SEE: When in regularly reach mainland Africa, suggesting migratory Madagascar, look up! Very behaviour. Widespread in many habitats, although gathers common. especially over wetlands. Readily nests on man-made structures, and can be seen in towns. ID: This is the default swallow in Madagascar. Although superficially similar to swifts, swallows are not in a closely related family. Flight more relaxed than a swift. Wings are more crooked, and less sickle-shaped. Unlike swifts, martins are often seen perched. Brown back and pale, dark-streaked underparts are unlike those of any swift. The Brown-throated Martin Riparia paludicola (not illustrated) is uncommon, mainly in the highlands: it is smaller than Mascarene Martin, has plain underparts, and is a much weaker flier. VOICE: Short sizzling call given mainly in flight. BEHAVIOUR: Often in small flocks. Insect prey is caught in flight. 3

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Long, thin tail

Much heavier than African Palm Swift, with much stronger flight

Malagasy subspecies are sometimes split as a full species: Madagascar Black Swift A.Â balstoni

Broader wings and more squared off tail than swifts

KINGFISHERS and BEE-EATER

Madagascar Malachite Kingfisher

Corythornis vintsioides vintsioides

M Vintsirano Martin-pêcheur Vintsi

DISTRIBUTION: Comoros and Madagascar. Fairly Length: 15 cm | 6" common in small numbers throughout Madagascar in WHERE TO SEE: Can be seen almost any kind of wetland habitat, including marshes, almost anywhere, even on rice rivers, lakes, rice paddies, mangroves and seashore. paddies. Present in Tana, such as ID: Colour, shape and habits all distinctive: a chunky little at Lake Alarobia. bird with a brilliant blue back and rufous underparts. VOICE: High-pitched “tsip”. BEHAVIOUR: Perches motionlessly, watching for prey, which it swoops down to snatch. Eats a wide variety of small animals including fish, insects and crustaceans. E

Madagascar Pygmy Kingfisher Corythornis madagascariensis

M Vintsiala Martin-pêcheur Malgache

DISTRIBUTION: Rainforest and dry forest below 1,200 m Length: 14 cm | 5·5" (4,000'). Absent from most of centre and southwest. Most WHERE TO SEE: Present at common in eastern rainforest, probably due to a higher most forest sites, but most abundance of prey, particularly frogs. common at Ranomafana and ID: Unmistakable jewel of a bird with an orange bill and Andasibe-Mantadia NPs. back, and white underparts. Despite its bright colour, can be surprisingly hard to spot in the gloomy forest interior. VOICE: Single note that is similar to that of Madagascar Malachite Kingfisher, but slightly lowerpitched, stronger, and more penetrating. BEHAVIOUR: This is a forest kingfisher, unlike the aquatic Madagascar Malachite Kingfisher. Like the Madagascar Malachite Kingfisher, it hunts from a perch. Eats insects and small vertebrates, especially frogs and small reptiles. Digs a nest cavity into an embankment, which can be along a dry streambed or beside a road. 3

Olive (Madagascar) Bee-eater Merops superciliosus

M Kiriokirioke Guêpier Malgache

DISTRIBUTION: Eastern and southern Africa, Comoros Length: 23–31 cm | 9–12" and Madagascar. Common throughout the island in WHERE TO SEE: Almost virtually all types of habitat. Nests in colonies in earthen anywhere, in open habitat. banks, often along rivers and streams. ID: Green colour and elongated central tail feathers are distinctive, even in flight. VOICE: A short rolling trill, often given in flight: “k-r-r-r-ree” which gives the bird its Malagasy name. BEHAVIOUR: Perches conspicuously on branches, snags or wires, often in small groups. Catches insect prey in flight.

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1 Always near water

Always in forest, sometimes near water

Colourful and conspicuous

ROLLER and HOOPOE 1

Broad-billed Roller Eurystomus glaucurus

Vorombaratra F Rolle Violet

DISTRIBUTION: Breeds throughout Madagascar in open Length: 32 cm | 12·5" forest, forest edge, woodland and savannah. Not usually WHERE TO SEE: Generally found in closed-canopy forest or in treeless areas. Present common and conspicuous, mainly between October and March, migrating to Africa but especially prominent during the dry season. around the Ankarafantsika NP ID: A large, colourful bird with a yellow bill, although can headquarters and Isalo. look dull and dark at a distance. Flies powerfully and can easily be mistaken for a falcon, although the roller has a yellow bill, light blue tail, and less smooth, more jerky flight action. VOICE: Loud series of complaining “kachak-kachak-kachak” notes that can be short or long and frequently repeated. Often sounds quite agitated, especially when a pair is calling together. BEHAVIOUR: Single birds or pairs spend long periods sitting high on prominent perches. Also flies around for extended periods, especially at dawn and dusk. Feeds mainly on insects, caught on the wing. E

Madagascar Hoopoe Upupa marginata

M Takodara Huppe Malgache

DISTRIBUTION: Fairly common throughout much of Length: 32 cm | 12·5" Madagascar, although scarce or absent in the east. Uses WHERE TO SEE: Western natural woodland and forest, as well as human-modified dry forest, as around habitats. Berenty, Kirindy Forest, and ID: Hoopoes are among the most unusual-looking and Ankarafantsika and Ankarana distinctive of all birds, and are readily identified by their NPs. peachy head and breast, prominent crest, and broad, black-andwhite wings that makes them look like huge butterflies in flight. VOICE: Low trill, very different from that of the world’s other hoopoes. BEHAVIOUR: Usually seen alone or in pairs. Walks along the ground with a bobbing gait, feeding on insects. Uses trees for perching (especially when singing), and nests in cavities. One of countless surprising things about Madagascar is that it lacks woodpeckers, despite being close to the African mainland, where woodpeckers are common. Madagascar Hoopoe, Nuthatch Vanga (page 170) and some tetrakas (page 162) are among the birds that extract invertebrates from dead wood and bark, filling the woodpecker niche on Madagascar. 1

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Boldly marked black-and-white wings and tail.

MORE COMMON RAINFOREST GROUND-ROLLERS

Ground-rollers form a family of birds found only on Madagascar. They are among the most beautiful and remarkable of the island’s birds. Most species that are elusive and local have not been covered in this book. However, an exception was made with ground-rollers, due to the attractive and unique nature of the group, and in the hope that including them will inspire more visitors to seek them out. The best way to find ground-rollers is to enlist the help of guides who specialize in birds. Ground-rollers are burrow nesters. When they fly through the rainforest, they seem surprisingly large, an impression that is enhanced by their noisy wingbeats. E

Pitta-like Ground-roller Atelornis pittoides

M Tsakoaka Brachyptérolle Pittoïde

DISTRIBUTION: Eastern rainforest. This is the most Length: 27 cm | 11" adaptable ground-roller, found in different kinds of forest WHERE TO SEE: This is the from low elevation up to 2,000 m (6,500'). ground-roller most likely to ID: Distinctive bird with a dazzling array of colours: be seen without expressly white, rufous, green and blue. Do not miss the spangles searching for it. Fairly common behind the eyes. in Ranomafana and AndasibeVOICE: Like all the rainforest ground-rollers, gives a “boop” Mantadia NPs. Possible but less song that can easily be imitated by humans. This vocalization common in Amber Mountain NP. Like all the ground-rollers, is quiet but penetrating, and repeated at intervals of several best searched for when vocal, seconds. from late September to early BEHAVIOUR: A shy and elusive bird, usually seen singly. Walks January. on the ground, sometimes on paths, dashing and jumping acrobatically to catch its prey, including insects, chameleons and frogs. Often wags its long tail. Will ascend to the mid-storey of the forest, especially to call. E

Rufous-headed Ground-roller Atelornis crossleyi

DISTRIBUTION: Uncommon in middle- and highelevation rainforest in eastern Madagascar. Prefers very moist forest with lots of bamboo, moss and fallen trees. ID: Smallest ground-roller. Shape and back colour similar to Pitta-like Ground-roller, but the rufous head and underparts are diagnostic. If flushed, the tail is uniformly coloured, unlike the brown-and-blue, two-toned tail of Pitta-like Ground-roller. Both species show a white blaze in the wing, both on the ground and in flight. VOICE: Gives a “boop” similar to that of Pitta-like Groundroller, but slightly lower-pitched and more disyllabic. BEHAVIOUR: Similar to Pitta-like Ground-roller. Less likely to call from a high perch; generally remains low to the ground.

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M Voromboka Brachyptérolle de Crossley

NEAR THREATENED Length:

25 cm | 10"

WHERE TO SEE: The Vohiparara Trail in Ranomafana NP. But even here, seeing this bird is difficult and likely to take some time. Other places that hold this bird include the higher sections of Andasibe-Mantadia NP and the Anjozorobe-Angavo forest.

The most widespread and frequently encountered ground-roller

Shy and difficult to see

SCARCER RAINFOREST GROUND-ROLLERS 1

Short-legged Ground-roller Brachypteracias leptosomus

M Fandikalalana Brachyptérolle Leptosome

DISTRIBUTION: Locally uncommon to rare in eastern VULNERABLE rainforest, in the most moist microhabitats. Prefers low Length: 38 cm | 15" and middle elevations. WHERE TO SEE: In Masoala NP, ID: The largest ground-roller, with a heavy chest and big this species is common enough square head. White tips to the tail feathers are diagnostic that it may be encountered by in flight. chance. It can also be found in VOICE: Typical ground-roller “boop”, higher-pitched than Andasibe-Mantadia NP with a that of Pitta-like or Rufous-headed Ground-rollers (page 142). dedicated search and the help Sometimes gives a quick rolling series of notes without pausing. of a guide. Occasionally found BEHAVIOUR: Unusual among ground-rollers in terms of in Ranomafana NP. behaviour, as it spends most of its time in trees. Nests in trees, either in a natural cavity or a burrow excavated into the roots of a big epiphyte. Forages for insects, crabs, frogs, chameleons, geckos and other reptiles. Often found in pairs. E

Scaly Ground-roller Geobiastes squamiger

M Fangadiovy Brachyptérolle Écaillé

DISTRIBUTION: Locally uncommon to rare in rainforest VULNERABLE of eastern Madagascar. Found in lower and middle Length: 30–32 cm | 12" elevations, especially in damp valleys, often along streams. WHERE TO SEE: Easiest to ID: One of the most spectacular of all Malagasy birds, find in Masoala and Marojejy with rufous, green and blue highlights, and fine scaly NPs. It can also be found in the markings. Pale blue tips to the tail feathers are distinctive Mantadia sector of Andasibein flight, although not as striking as the white tail tips of Mantadia NP with a dedicated Short-legged Ground-roller. search and the help of a VOICE: Song is a series of “boop” notes like other groundguide. Occasionally found in rollers, but the individual notes are lower-pitched and longer Ranomafana NP. than those of other species. Seems to be less vocal than other rainforest ground-rollers. BEHAVIOUR: Similar to Rufous-headed Ground-roller (page 142). Often leaves forest floor to call from a low perch. Feeds mainly on invertebrates, plus some small vertebrates. 2

144

Like New World antpittas and Old World pittas, ground-rollers are sometimes seen on forest trails.

Purple highlights on the back of the head

The most arboreal ground-roller

Complex and beautiful patterning

LONG-TAILED GROUND-ROLLER and CUCKOO-ROLLER

Long-tailed Ground-roller

Uratelornis chimaera

M Tolohoranto Brachyptérolle à Longue Queue

DISTRIBUTION: A small portion of the spiny forest along the southwest coast, north of Toliara. ID: One of Madagascar’s most distinctive birds, with a long tail, powder blue wing patch and boldly marked head. VOICE: Less vocal than rainforest ground-rollers. Gives a variable series of “toop” calls which are delivered faster than in the other ground-rollers. BEHAVIOUR: Secretive and elusive despite the open nature of its spiny forest habitat. Behaves much like a roadrunner (distinctive New World birds), running on its strong legs, and cocking up its long tail. Eats insects.

VULNERABLE Length:

47–52 cm | 19–20"

WHERE TO SEE: Not found anywhere on the circuit of sites frequently visited by general naturalists. Birders who are keen to find this, and other spiny forest specialties, visit the small private forest reserves near Ifaty, north of Toliara. There, skilled local guides assist in locating this shy species.

Cuckoo-roller is another of Madagascar’s odd avian treasures, and one of the world’s most enigmatic birds. It is so different from other birds that it is placed in its own family, which, in turn, may even deserve to be considered a distinct order. What this means is that it has been evolving independently for a long time. R

DISTRIBUTION: Comoros and Madagascar. Found throughout Madagascar in all forest types. Sometimes found in heavily degraded habitats that still contain trees. Less common in the spiny forest than in other forest types, and most common in eastern rainforest. ID: A large, distinctive bird with an oddly reptilian aspect when seen up close. Very different male and female plumages. Brown female could be mistaken for a raptor when perched, but has a hook-less bill and a different pattern from any raptor. The male has a dark wings with iridescent highlights and blue-grey underparts; it is unlike any other bird. Could also be mistaken for a raptor in flight, but can usually be recognized by its incessant call. VOICE: One of the most obvious and distinctive Malagasy natural sounds: an incessantly-given, three-part, whistled, descending “pheeeew teeeew teeew”. This may be the only bird sound to be heard on a hot afternoon in the forest. This song is given both when perched, and more often in flight. Male makes an elaborate flight display that includes calls, gliding, flapping and rolling. BEHAVIOUR: Perches motionlessly in the canopy for long periods. Frequently flies over the forest, alternating periods of flapping and soaring. Nests in cavities. Eats a range of small animals, although especially fond of chameleons. 146

Cuckoo-roller Leptosomus discolor

Vorondreo F Courol

Length:

50 cm | 20"

WHERE TO SEE: Widespread but easiest to see in the western dry forest, as in Ankarafantsika, Zombitse and Tsingy de Bemaraha NPs, and Kirindy Forest.

2♀

2♂ Most often seen in flight; soars like a raptor

2â&#x2122;&#x20AC;

Off-centre positioning of the eye gives the Cuckoo-roller an oddly reptilian appearance.

2â&#x2122;&#x201A;

ASITIES

Asities are yet another family of birds found only in Madagascar, the closest relatives to which are the broadbills of Asia. Breeding males sport caruncles, bizarre patches of bright blue and/or green skin on their face. These caruncles grow at the start of the breeding season but fade away during the non-breeding season. Asities have a unique method of constructing their hanging nests. They first weave an orb then poke a hole into it to create an entrance, rather than building the entrance into the nest. E

Velvet Asity Philepitta castanea

DISTRIBUTION: Uncommon in eastern rainforest, especially at middle elevations. ID: The black male with green caruncles when breeding or yellow fringes when not breeding 1a is unmistakable. The female is a drab bird, but can be identified by its round shape and short tail. VOICE: Most common call consists of very high-pitched notes, almost like a rusty hinge. BEHAVIOUR: Lives mainly in forest mid-storey. Perches motionlessly for long periods and often allows close approach. Feeds on fruit. Unusual breeding system in which males have leks where they compete to attract the attention of females. Display behaviours include opening the wings, displaying the bright caruncle, and sometimes even performing somersaults. E

Schlegel’s Asity Philepitta schlegeli

Length:

M Asity Philépitte Veloutée

16 cm | 6"

WHERE TO SEE: AndasibeMantadia and Ranomafana NPs.

M Soisoy Philépitte de Schlegel

DISTRIBUTION: Rare and local, mainly in western dry NEAR THREATENED forest, but also in Sambirano forest in the north. Length: 13 cm | 5" ID: Breeding males’ multi-coloured caruncles render WHERE TO SEE: Ankarafantsika them unmistakable. Non-breeding males, females and and Tsingy de Bemaraha NPs. immature birds are fairly drab, but within their range can be identified by their round shape and short tail. VOICE: Extremely high-pitched “sit” notes usually delivered in series. BEHAVIOUR: Similar to Velvet Asity, although more shy, and more often found in canopy. Also leks, but has a different display from Velvet Asity; lekking behaviour may involve drooping the wings, fluﬃng up the breast feathers, cocking the tail and singing. E

Common Sunbird Asity Neodrepanis coruscans

DISTRIBUTION: Fairly common, although hard to see, in middle- and high-elevation eastern rainforest. ID: One of Madagascar’s smallest birds. Although the breeding male is distinctive, having a blue caruncle, the female and non-breeding male could be confused with a sunbird (page 164), but are smaller, shorter-tailed and have a more curved bill. VOICE: Short series of extremely high-pitched “seep” notes. BEHAVIOUR: Usually seen singly. Generally stays in the canopy, but will descend to lower flowering trees. Feeds mainly on nectar. 148

M Zafindrasity Philépitte Souimanga

Length:

11 cm | 4"

WHERE TO SEE: Not rare, but mostly confined to the high canopy – so be prepared to get a stiff neck when searching for it! Best looked for in Ranomafana and AndasibeMantadia NPs.

1♀

1♂

2♀

2♂

3♀

3♂

In breeding plumage, male asities develop colourful caruncles

COMMON OPEN-COUNTRY BIRDS

Madagascar Lark Mirafra hova

DISTRIBUTION: One of the most common and widespread Malagasy birds, found throughout in open habitat. ID: A small, short-tailed, streaky bird with a pale bill. Larks can be confusing, but this is the only small terrestrial brown bird in Madagascar. Madagascar Cisticola (page 158) is somewhat similar but smaller, and only occasionally walks on the ground. VOICE: Mixture of drawn-out and musical notes, and shorter, chatty notes. Sings in flight. BEHAVIOUR: Walks on the ground, feeding on seeds and insects. E

Madagascar Wagtail Motacilla flaviventris

M Sorohitra Alouette Malgache

Length:

13 cm | 5"

WHERE TO SEE: Common throughout most of Madagascar. This is one of few native birds that manages to thrive in completely deforested places, like the desolate grasslands of the High Plateau.

M Triotrio Bergeronnette Malgache

DISTRIBUTION: Throughout the island in open habitats, Length: 19 cm | 7·5" often near water. Especially common on the High Plateau, WHERE TO SEE: Common in often in towns. Tana and throughout most of ID: A small, long-tailed bird with a grey back, the island. yellow belly and a black 'necklace' across the chest. VOICE: Normal call is a two-part “trio-trio”. The song includes more musical, whistled notes. BEHAVIOUR: Continually walks around and wags its tail. Feeds on invertebrates. Some English common names and even scientific names of birds are ‘onomatopoeic’, created in imitation of a call made by the bird. But a much higher proportion of Malagasy names are delightfully onomatopoeic. The wagtail is a good example: ‘Triotrio’ in imitation of the wagtail’s scratchy two-part call. 3

Pied Crow Corvus albus

M Goaika Corbeau Pie

DISTRIBUTION: Sub-Saharan Africa. Common in small Length: 46–50 cm | 18–20" numbers throughout Madagascar in almost any open WHERE TO SEE: Conspicuous habitat. Generally less common on the High Plateau. and numerous, especially in dry, ID: Large black-and-white bird. Can soar and look raptoropen habitat in the west. like at a distance, but no raptor has its pied patterning. VOICE: Rough “crrrraw”, similar to that given by most crows. BEHAVIOUR: Often in pairs or small groups. Frequently seen flying, often quite high. Highly adaptable, and feeds on almost anything; will eat many plant materials, prey on small animals, and scavenge. 3

150

Often mobbed by smaller birds like Crested Drongo (page 176)

Sings while performing a display flight

Usually seen on the ground or perched on man-made structures

Looks raptor-like in flight, but is strikingly black-and-white

WIDESPREAD SONGBIRDS

Madagascar Bulbul Hypsipetes madagascariensis

M Horovana Bulbul Malgache

DISTRIBUTION: Seychelles (Aldabra), Glorieuses, Length: 24 cm | 9·5" Comoros and Madagascar. Found throughout Madagascar WHERE TO SEE: Almost in almost any habitat with woody vegetation. anywhere, including Tana. ID: Slim, medium-sized brown-black bird. Separated from all other species by its orange bill. VOICE: A complex series of rattling calls. Also gives a cat-like whining. BEHAVIOUR: Often in pairs or groups, which are active and conspicuous. Feeds on fruit and insects. The Madagascar Bulbul belongs to a genus that is found in Asia and the Malagasy region, but not on mainland Africa, a clear example of a Malagasy species with Asian origins. R

Madagascar Paradise Flycatcher Terpsiphone mutata mutata

M Tsingitry Gobe-Mouche de Paradis de Madagascar

DISTRIBUTION: Comoros and Madagascar. Fairly Length: 18 cm | 7" common throughout Madagascar in woodland and forest (long–tailed males habitat, including areas heavily modified by humans. up to 30 cm | 12") ID: Long-tailed males are easy to recognize, and occur WHERE TO SEE: Common at all as white or rufous morphs. The shorter-tailed all-rufous forest sites, although perhaps female is slightly less obvious, but still distinctive. easiest to see in western dry All birds show blue bare skin around the eye. forest. VOICE: The call is a short, rough “shree-shreet”. The song is an exuberant and bubbling series of pure whistles. BEHAVIOUR: Usually seen singly or in pairs, moving restlessly through the middle storey. Feeds mainly on insects. E

Madagascar Mannikin Lonchura nana

DISTRIBUTION: Fairly common although inconspicuous throughout Madagascar in almost any open habitat. ID: Tiny brown bird with a black bib. VOICE: Short whistled “tsweet”, most frequently given in flight and often in chorus by a flock. BEHAVIOUR: Highly social, and usually in active flocks of up to 20. Feeds exclusively on seeds, especially those of grasses.

Length:

M Tsiporitika Capucin Malgache

9 cm | 3·5"

WHERE TO SEE: Common at most sites, but can be inconspicuous and is easily overlooked.

152

Almost always found in flocks

2♂

Male white morph. Can show a white or black back 2♀

Orange bill

Tiny, social birds

2♂

Immature male rufous morph. Tail is elongated, but shorter than full adult plumage, which takes 3–5 years to attain

ROBIN-LIKE BIRDS

Madagascar Magpie-robin

Copsychus albospecularis

M Fitatrala Shama Malgache

DISTRIBUTION: Fairly common throughout Madagascar Length: 18 cm | 7" in all types of forest. Although absent from completely WHERE TO SEE: Common in deforested areas, it is found in forest edge and secondary all forest sites, although most habitats. easily seen in dry forest. Look ID: Black-and-white male is distinctive. Brown-and-white for black-bellied males in female is less obvious, but can still be identified by white wing Masoala and Marojejy NPs. patch. The males in the northeastern rainforest have a black belly. VOICE: The song is a complex, variable series of thin whistles and lower notes, sometimes incorporating mimicry of other birds. BEHAVIOUR: Usually seen singly or in pairs, on the ground or in the understorey. Hops rather than walks along the ground. Feeds mainly on insects and spiders. E

Forest Rock Thrush Monticola sharpei

DISTRIBUTION: Patchily distributed and generally uncommon in eastern rainforest and in rocky areas of the west. There is also an isolated population in the rainforest of Amber Mountain. ID: Rufous-and-grey male is distinctive. Brown female lacks the white patch in the wing of Madagascar Magpierobin, and is larger than female African Stonechat. Western populations are paler than eastern and northern ones, and were formerly split as Benson's Rock Thrush M. bensoni 2a . VOICE: Astoundingly variable series of mournful-sounding whistles. Often mimics other birds within its song. BEHAVIOUR: Eastern and northern birds are found in the understorey of forest, often adjacent to rocky areas. Western birds live on exposed rocks. Eats insects and fruit. WHERE TO SEE: The Isalo area, including the vicinity of some of the lodges. Also often seen along the main highway through Ranomafana NP, and in Tsingy de Bemaraha and Amber Mountain NPs.

African Stonechat Saxicola torquatus

M Androbaka Monticole de Forêt

Length:

16 cm | 6"

Beachgoers who visit Anakao, south of Toliara, should see the Littoral Rock Thrush M. imerinus, a localized bird of the southwest coast.

M Fitatra Tarier d’Afrique

DISTRIBUTION: Sub-Saharan Africa, Comoros, and Length: 14 cm | 5·5" Madagascar. Occurs widely in brushy habitat and marshes. WHERE TO SEE: Widespread One of the few birds that occurs on top of the highest and common. Can be seen on mountains. the deforested High Plateau. ID: The distinctive male is black-and-white, with a rufous chest patch. The brown female is less obvious, but is the only brown robin-like bird likely to be seen perching in open country. VOICE: Harsh “chak” calls and complex, musical song. BEHAVIOUR: Single birds and pairs perch conspicuously on exposed twigs. Often flicks wings and tail. Eats insects and spiders. 154

1♂

Males have black belly and tail in the northeast. Birds away from the east have a big white patch in the wing and white in the tail.

1♀ 1

2a♂

2♀

2♂

The isolated subspecies of Forest Rock Thrush on Amber Mountain (M. s. erythronotus) is distinctive, and was long considered a full species, Amber Mountain Rock Thrush. Males have a rufous-brown, rather than grey, back.

3♂

3ju

Brownish juvenile and female are similar

WARBLERS THAT LIVE CLOSE TO THE GROUND I

R e

Madagascar Brush Warbler Nesillas typica

M Poretaka Nésille Malgache

DISTRIBUTION: Comoros and Madagascar. Common Length: 17–18 cm | 7" in forest undergrowth and brushy habitats in most of WHERE TO SEE: Common in Madagascar except the southwest, where it is replaced by many places, including around the similar but paler Subdesert Brush Warbler. Andasibe and Ranomafana, ID: Surprisingly large, long-tailed grey-brown bird. although its skulking habits can Its size and skulking habits usually serve to identify it, make it difficult to see. even when it is not seen well. See Subdesert Brush Warbler and Madagascar Swamp Warbler (page 158) for separation from those similar species. VOICE: Dry ‘ticking’ call, like playing with the tines of a comb. BEHAVIOUR: Stays low in thick undergrowth. Usually alone or in pairs. Eats small insects and spiders.

Subdesert Brush Warbler Nesillas lantzii

Poratakatanoravolo F Nésille de Lantz

DISTRIBUTION: Common throughout the southwest in a Length: 17 cm | 7" variety of dry, brushy habitats, from spiny forest to village WHERE TO SEE: Common gardens. in the southwest, including ID: Like a bleached-out Madagascar Brush Warbler. Berenty and the Toliara area. There is little overlap between these two species, so they can normally be identified by range. VOICE: Similar to Madagascar Brush Warbler, but the ‘ticking’ call is higher-pitched and drier. BEHAVIOUR: Like Madagascar Brush Warbler.

156

Long bill and tail, and rather drab colours

Shape like a Madagascar Brush Warbler, but much paler

WARBLERS THAT LIVE CLOSE TO THE GROUND II

Madagascar Cisticola Cisticola cherina

M Tsintsina Cisticole Malgache

DISTRIBUTION: Glorieuses, Seychelles, and Madagascar. Length: 12 cm | 5" Common throughout Madagascar in open habitats like WHERE TO SEE: Almost grasslands, marshes, and cultivated areas. anywhere except thick forest. ID: The only small, brown, warbler-like bird to be found in open habitats. In flight, the white tips to the tail are distinctive. VOICE: Dry “tsip”, usually repeated several times, given in a display flight. BEHAVIOUR: Often perches up conspicuously on grass and shrubs, but can be skulking. Eats insects and spiders. E

Madagascar Swamp Warbler Acrocephalus newtoni

DISTRIBUTION: Locally fairly common throughout Madagascar in thickly vegetated wetlands, including papyrus and mangrove swamps. ID: Similar to Madagascar Brush Warbler (page 156). A large warbler in a soggy marsh is probably a Madagascar Swamp Warbler, but the two species overlap in wet scrub. Madagascar Swamp Warbler can be distinguished by its striped throat, greyer colour, thicker bill and very different voice. VOICE: Complex, musical, chortling song. Harsh“chak” calls. BEHAVIOUR: Lone birds and pairs hop about in marsh vegetation. Sometimes perches in the open to sing.

M Vorombararata Rousserolle de Newton

Length:

18 cm | 7"

WHERE TO SEE: Almost any wetland, including Lake Alarobia in Tana.

Sometimes looks quite grey

158

Sings in flight

Pale tips and black band on tail

Long bill and tail, striped throat and breast, and dull plumage

FOREST CANOPY WARBLERS

Common Jery Neomixis tenella

M Jijikely Petite Éroesse

DISTRIBUTION: Fairly common throughout Madagascar Length: 10 cm | 4" wherever there are trees or brush. WHERE TO SEE: Common ID: Greenish colour and thin, pointed bill typical of jeries. everywhere, but especially easy Very similar to Stripe-throated Jery, but distinguished by to see in the spiny forest of the grey back of the head, pale bill and smaller size. southwest, where they perch VOICE: Bouncy, accelerating series of several buzzy notes. conspicuously and sing. BEHAVIOUR: Active little bird that often sings from the top of the tallest available vegetation, whether bushes, tall trees, or dead snags. Joins flocks with other jeries, white-eyes, and vangas. Eats mainly insects. E

Stripe-throated Jery Neomixis striatigula

DISTRIBUTION: Found in all types of forest, though strangely absent from the northwest and most of the centre and far north. Fairly common in much of the east and southwest ID: See Common Jery for separation from that species. VOICE: Accelerating series of notes: more musical, structured and longer than the song of Common Jery. BEHAVIOUR: Canopy species that often often joins mixed flocks. Like Common Jery, sings from prominent perch. Eats insects. The behaviour of some of the cryptic brown-and-green Malagasy warblers is among the strangest natural phenomena on an island that is famous for the unusual. The Stripe-throated Jery (right) and the Rand’s Warbler (left) belong to different bird families, but sing almost the same song, and take turns singing side-by-side on the same tall trees and snags during the breeding season. Both species must benefit from this ‘double coverage’ of their territory’s auditory space. E

Length:

12 cm | 5"

WHERE TO SEE: Consistently perched up and singing around the Ranomafana NP headquarters. Also easy to see at Andasibe-Mantadia NP and in the spiny forest. 2 3

Rand’s Warbler Randia pseudozosterops

DISTRIBUTION: Uncommon to fairly common in eastern rainforest. ID: Separated from Stripe-throated Jery by broad eyebrow, greyer colour, and less pointed bill. VOICE: Very similar to Stripe-throated Jery, but less varied. A musical, accelerating trill. BEHAVIOUR: A canopy species that is most often seen when perched up on a snag or tall tree to sing. Feeds by creeping along branches and checking on the underside.

M Kimitsy Grande Éroesse

Length:

M Biritsy Randie Malgache

12 cm | 5"

WHERE TO SEE: Reliable at Ranomafana NP, including at the headquarters alongside Stripe-throated Jery. Also fairly common in Andasibe-Mantadia NP.

Rand’s Warbler is a member of the Bernieridae family (see page 162), which was recognized only in 2010, and is endemic to Madagascar. Despite their profound genetic distinctiveness, and despite being described, the members of this family were overlooked by taxonomists for many years, due to their superficial similarity to birds within other families. 160

Eastern birds are darkest and dullest

Southwestern birds are pale below

Western and northern birds are brightest

Strong, dark bill; prominent stripe over the eye

Southwestern birds are paler

Grey back and pale underparts; broad stripe over the eye

Creeps along branches

BERNIERIDS

This guide covers the most widespread and easily seen warbler-like birds of Madagascar. However there are others, including Green Jery Neomixis viridis, Madagascar Yellowbrow (or Yellow-browed Oxylabes) Crossleyia xanthophrys and Grey-crowned Tetraka (or Grey-crowned Greenbul) Bernieria cinereiceps. Although these are not Madagascar’s most colourful birds, they do have interesting behaviours, and provide some evidence of the course of evolution on this isolated island. E

Long-billed Tetraka Bernieria madagascariensis

DISTRIBUTION: Found throughout most of Madagascar in all types of forest. Generally absent from spiny forest, although it occurs locally in this habitat north of Toliara. ID: A relatively large, long-tailed, olive-green bird with a large bill. Male is bigger overall than female, and has a very long bill. Female is smaller-billed, and can look similar to Spectacled Tetraka, but is larger and has a weaker eye ring. VOICE: Call is a short, rough “brrrt”. Song is a series of musical whistles. BEHAVIOUR: An active understorey and mid-storey species that often flocks with other species. Probes for arthropods, sometimes creeping along tree trunks and limbs like a woodpecker.

Length:

18–20 cm | 7–8"

WHERE TO SEE: Fairly common in all forested sites except in the southwest.

Feeds by creeping along trunks, sometimes even vertical ones

Madagascar's tetrakas were formerly known as 'greenbuls' before their true status as an endemic family was recognized. E

M Tretraka Tétraka à Bec Long

Spectacled Tetraka Xanthormixis zosterops

M Farifotramavo Tétraka à Bec Court

DISTRIBUTION: Fairly common in eastern rainforest, Length: 16 cm | 6" plus some patches of northern Sambirano forest, and the WHERE TO SEE: One of the isolated rainforest on Amber Mountain. most common forest birds ID: See Long-billed Tetraka for separation from that in Ranomafana, Andasibesomewhat similar species. The Spectacled Tetrakas in the Mantadia, and Masoala NPs. northeast, such as at Masoala and Marojejy, are quite dark, while those on Amber Mountain are pale and greyish. VOICE: Piercing, high-pitched “tsit” calls, which sometimes run into a short series: very different from any call of Long-billed Tetraka. BEHAVIOUR: Active and social bird of the understorey and mid-storey, where it is a core member of mixed flocks. Primarily eats insects. E

White-throated Oxylabes Oxylabes madagascariensis

M Farifotra Mena Oxylabe à Gorge Blanche

DISTRIBUTION: Uncommon to fairly common in eastern Length: 17 cm | 7" rainforest, plus some patches of northern Sambirano WHERE TO SEE: Ranomafana forest, and the isolated rainforest on Amber Mountain. and Andasibe-Mantadia NPs. ID: Distinctive, unlike most other Bernierids, having a brown back, rufous cap and underparts, and bright white throat. VOICE: Call is a sizzling, descending trill. Song is a musical, oscillating series of cheerful notes. BEHAVIOUR: Pairs and small groups found on or near the ground. Hops on the ground and readily climbs low branches, especially when vocalizing. Feeds on insects and spiders. 162

1â&#x2122;&#x201A; 1â&#x2122;&#x20AC;

Male has a very large bill

Broken yellow eye-ring

An understorey skulker

Bright white throat

SUNBIRDS and WHITE-EYE

Souimanga Sunbird Cinnyris souimanga

M Soimangakely Souimanga Malgache

DISTRIBUTION: Seychelles (Aldabra), Glorieuses, and Length: 10 cm | 4" Madagascar. Fairly common throughout Madagascar in WHERE TO SEE: Almost virtually all habitats with some woody vegetation. anywhere, including Tana. ID: Breeding male is distinctive, with a green head and back, yellow belly and multi-coloured breast. Female and non-breeding male distinguished from Madagascar Green Sunbird by smaller bill and lack of strong streaks on underparts. Female could also be confused with female Common Sunbird Asity (page 148), but is longer-tailed and has a longer straighter bill. VOICE: One of the most commonly heard bird songs on the island, a joyous jumble of notes. Call is a short, high-pitched whine. BEHAVIOUR: Pugnacious bird that is constantly active. Uses its long bill to extract nectar from flowers, and to catch small insects and spiders. R

Madagascar Green Sunbird Cinnyris notatus

M Soimangabe Souimanga Angaladian

DISTRIBUTION: Comoros and Madagascar. Found Length: 14 cm | 5·5" throughout the island in most habitats with a woody WHERE TO SEE: Fairly component. Generally less common than Souimanga common throughout, including Sunbird. Tana. Perhaps most easily ID: Male looks all-black in poor light, although in found in western forest, as at good light it shows a green head and back with purple Ankarafantsika NP. highlights. Female distinguished from Souimanga Sunbird by its thicker bill and heavily streaked underparts. VOICE: Most common vocalization is a sparrow-like “cheep”. BEHAVIOUR: Similar to Souimanga Sunbird, but its larger bill allows it to extract nectar from bigger flowers. R

Madagascar White-eye Zosterops maderaspatanus

M Fotsy Maso Zostérops Malgache

DISTRIBUTION: Seychelles, Comoros, Glorieuses, Length: 12 cm | 5" Europa, and Madagascar. Found throughout Madagascar WHERE TO SEE: Quite common in habitats with some woody vegetation. at most sites. ID: Somewhat like a jery (page 160), but has bright yellow throat and undertail, and a white ring around the eye. VOICE: Extremely vocal. The song is a warbling chatter. Calls a variety of simple notes and phrases. BEHAVIOUR: A hyperactive bird that is usually found in flocks, which often contain jeries, newtonias, and sunbirds. 3

164

1♂

transitioning to breeding plumage

1♂

1♀ Heavy bill

2♂

Bold white ring around the eye

2♀

COMMON and WIDESPREAD VANGAS

Vangas comprise another bird family that is endemic to the Malagasy region. They are the Malagasy counterpart to the Darwin’s finches of Galápagos, both astounding examples of groups of birds that colonized new parts of the world, then evolved to fill a wide variety of previously vacant niches. There are warbler-like vangas, shrike-like vangas, babbler-like vangas and even a nuthatch vanga! This single family of 21 species contains almost the same range of diversity of body sizes, bill shapes and feeding feeding strategies of the whole world's songbird radiation, containing more than 5,000 species. Vangas are closely related to the helmetshrikes of Africa, and some authorities now place them in the same family. E

Common Newtonia Newtonia brunneicauda

M Tretreka Newtonie Commune

DISTRIBUTION: Fairly common in all types of forest, Length: 12 cm | 5" including heavily disturbed and secondary forest, WHERE TO SEE: Easily seen at throughout Madagascar. any site with forest. ID: Small, thin-billed dull greyish vanga. Angry-looking pale eye separates it from all potential confusion species like jeries and female Red-tailed Vanga. Vocalizations also distinctively different from similar species. VOICE: Jerky, syncopated series of harsh calls. Often sounds like “gidyup-gidyup-gidyup”. BEHAVIOUR: Highly active and feisty bird. Joins flocks with many other species of birds including other vangas, jeries, Madagascar Paradise Flycatcher, and Souimanga Sunbird. Dark Newtonia Newtonia amphichroa (not illustrated) is a species of eastern rainforest, with an isolated population on Amber Mountain; it is darker than Common Newtonia and has a dark eye.

Red-tailed Vanga Calicalicus madagascariensis

M Totokarasoka Calicalic Malgache

DISTRIBUTION: Uncommon to fairly common Length: 13–14 cm | 5" throughout in all types of forest. WHERE TO SEE: Fairly common ID: Red, grey, and black male is distinctive. Grey-brown in all forested sites. female can be confused with Common Newtonia, but is larger, with a bigger bill, and a dark eye. VOICE: Loud ‘wolf whistle’ with some regional variation. Also gives various calls, including a harsh “krrrrrit”. BEHAVIOUR: Similar to the chickadees or tits which are familiar to most visitors from the northern hemisphere. Active, pugnacious birds that often form the core of forest bird flocks, and whose calls lead the group. Eats invertebrates. E

Chabert Vanga Leptopterus chabert

M Patsatsatra Artamie Chabert

DISTRIBUTION: Fairly common throughout Madagascar Length: 14 cm | 5·5" in all forest types, mainly below 1,300 m (4,300'). This is WHERE TO SEE: Present also the only vanga likely to be found in heavily modified throughout, but especially habitats like plantations and artificial savannah. easy to see in the west, such as ID: Blue-black and white, with bright blue eye ring. around Toliara and Morondava, VOICE: Harsh repeated “chak” call mainly given in flight. and Ankarafantsika NP. BEHAVIOUR: Acts more like a finch than a vanga; usually seen in small, active flocks, which fly high. Feeds on insects both in trees and on the wing. 166

Pale eye and thin bill

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Dark eye and red tail

White patches in tail

TYPICAL VANGAS

The three species shown here are perhaps the most ‘average’ of the vangas in terms of size and shape. R

Cyanolanius madagascarinus madagascarinus

Patsatsatrala Artamie Azure

Blue Vanga

Length:

16 cm | 6"

DISTRIBUTION: Uncommon to fairly common WHERE TO SEE: Found in all throughout most of Madagascar in all types of forest, forest sites, although perhaps although largely absent from spiny forest. easiest to see in dry forest, ID: Blue back and pale underparts are diagnostic. such as at Kirindy Forest and The female is duller. Has the same overall pattern as Ankarana and Ankarafantsika Chabert Vanga (page 166), but with blue, rather than blue-black, NPs. on the back. VOICE: Harsh rolling “trrrt-trrrt-trrrt”. BEHAVIOUR: Often in flocks with vangas and other birds. Usually in the upper-storey of the forest. Feeds on invertebrates and fruit. E

White-headed Vanga Artamella viridis

M Voromasiaka Artamie à Tête Blanche

DISTRIBUTION: Uncommon throughout Madagascar in Length: 20 cm | 8" all types of forest, including edge habitat and secondary WHERE TO SEE: Found in all forest. forest sites, although perhaps ID: Shares pied plumage with Sickle-billed (page 172) easiest to see in dry forest, and Chabert (page 166) Vangas, but is much smaller and such as at Ankarana and shorter-billed than Sickle-billed, and larger, and palerAnkarafantsika NPs. headed than Chabert Vanga. VOICE: The calls are harsh, grating notes. The song comprises various low, penetrating whistles. BEHAVIOUR: Often joins mixed flocks. Usually in the upper-storey of the forest. Feeds on invertebrates. In the west, large numbers of White-headed and Sickle-billed Vangas sometimes roost together, and even nest together; it is as if the smaller, stubby-billed White-headed Vangas feel safe with their bruising ‘older brothers’! E

Rufous Vanga Schetba rufa

M Vangamena Artamie Rousse

DISTRIBUTION: Found in both eastern rainforest and Length: 20 cm | 8" western dry forest, although more common in western WHERE TO SEE: Most easily forest. Absent from most of the spiny forest and from the found in the western dry far north. forest, as at Kirindy Forest, and ID: Bright rufous back, black head, and white underparts Zombitse and Ankarafantsika are distinctive. Male shows black onto the breast, while NPs. female has a pale breast. VOICE: Most common and distinctive vocalization is a long, melodic, descending trill. Also gives a clear, three-part “whit-whit-tew”, which pairs sometimes combine with the descending trill for a beautiful duet. BEHAVIOUR: Found mainly in the forest mid-storey. Often sits motionlessly for long periods. Joins flocks less frequently than most other vangas. Eats invertebrates. 168

1♀

Usually seen in the canopy

2♀

1♂

2♂

Female has dusky markings on head and underparts

3♀

3♂

Rufous back

STRANGE VANGAS

Serikalambo Vanga de Ward

Ward’s Vanga Pseudobias wardi

DISTRIBUTION: Fairly common in eastern rainforest, Length: 15 cm | 6" mainly at middle elevations. Particularly favours forest WHERE TO SEE: Ranomafana edge. and Andasibe areas. Look ID: An odd and unique little bird with black-and-white especially along the main road plumage and blue bare skin around the eye. Much more through Ranomafana, and likely to be taken for a flycatcher than a vanga. Some male behind the Feon’ny Ala Hotel Madagascar Paradise Flycatchers (page 152) are black-andnear Andasibe. white, but they are much larger, longer-tailed, and lack the black chest band of Ward’s Vanga. VOICE: Short, dry trill. BEHAVIOUR: Found singly or in pairs in the upper-storey of the forest. Sometimes in flocks with other species. Feeds by catching insects on the wing, much like a flycatcher. E

Nuthatch Vanga Hypositta corallirostris

Voronakodidina F Vanga Sitelle

DISTRIBUTION: Rare and local in eastern rainforest Length: 13 cm | 5" below 1,000 m (3,300'). WHERE TO SEE: Most readily ID: The blue plumage, coral-red bill and behaviour are found in vanga flocks in distinctive. Male is brighter than the female. Andasibe-Mantadia NP. Also VOICE: Short, very high-pitched trill. Very easy to possible in Masoala NP. overlook if you are not specifically listening for it. BEHAVIOUR: Usually found in flocks of vangas and other birds. This unusual little vanga seems to have evolved to fill the niche of birds like nuthatches and woodpeckers, which are absent from Madagascar. It creeps up tree trunks in search of insects. E

Crossley’s Vanga Mystacornis crossleyi

DISTRIBUTION: Uncommon in eastern rainforest, mainly below 1,300 m (4,300'). ID: Behaviour and bold black, grey and white facial markings identify this bird. Very different from other vangas. VOICE: Piercing, high-pitched whistle, which is often repeated several times. BEHAVIOUR: Walks around on forest floor in singles or pairs. Pairs display by circling each other with tails cocked up. Feeds on insects and spiders. Crossley’s Vanga was formerly known as ‘Crossley’s Babbler’, while Ward’s Vanga was known as ‘Ward’s Flycatcher’. It was only recently that genetic analysis revealed that these babbler- and flycatcherlike species are actually vangas.

170

Length:

M Talapiotanty Vanga de Crossley

16 cm | 6"

WHERE TO SEE: Ranomafana and Andasibe-Mantadia NPs. Also present although uncommon in Masoala NP.

Blue eye ring like a paradise flycatcher; canopy-dwelling 2♀

2♂

Surprisingly woodpecker-like in behaviour

3♀

3♂

A shy bird of the forest floor

LARGE VANGAS

Hook-billed Vanga Vanga curvirostris

M Vangasoratra Vanga Écorcheur

DISTRIBUTION: Fairly common in small numbers Length: 25–29 cm | 10–11" throughout the island in all forest types. Tolerates WHERE TO SEE: Easiest to secondary forest and degraded habitat not used by many find in western dry forest, as other vanga species. at Berenty, Kirindy Forest, and ID: Big, black-and-white, shrike-like vanga. Ankarafantsika NP. VOICE: Drawn-out, hollow whistle. BEHAVIOUR: Slowly moves through the mid-storey of forest. Usually seen alone or in pairs, and rarely in mixed species flocks. This large vanga is a major predator both of invertebrates and a range of vertebrates, including reptiles, amphibians, small birds, and even baby mouse lemurs. E

Helmet Vanga Euryceros prevostii

DISTRIBUTION: Rare and local in rainforest in northeast Madagascar. ID: The huge blue bill alone identifies this bird. VOICE: A quiet bird that sometimes gives a soft, musical, descending trill. BEHAVIOUR: Ranges mainly in mid-storey of forest, sometimes perching motionlessly for long periods. Often joins flocks of other large vangas. Feeds on invertebrates and small vertebrates like geckos and frogs.

Sickle-billed Vanga Falculea palliata

M Siketribe Eurycère de Prévost

VULNERABLE Length:

29 cm | 11·5"

WHERE TO SEE: Masoala is the most accessible site. Easiest to find September–December, when local guides have often located nests. Marojejy NP is also excellent, for those willing to hike. Does occur in the Andasibe area, though rarely in areas that are easily accessible.

M Voronjaza Faculie Mantelée

DISTRIBUTION: Fairly common and conspicuous in all Length: 32 cm | 12·5" types of dry forest and woodland in northern, western, WHERE TO SEE: Fairly common and southwestern Madagascar, including secondary forest and quite conspicuous at and savannah. Especially fond of areas with baobabs. Ankarana and Ankarafantsika ID: Large, black-and-white, curve-billed bird. NPs, Kirindy Forest, and Berenty. Floppy flight on broad wings is distinctive. VOICE: Arresting combinations of cackles, rasps, and mournful wails. BEHAVIOUR: Social bird that is usually found in groups. Adept at probing for invertebrates with its huge bill.

Some of the vocalizations of Sickle-billed Vanga are uncannily like the wails of a child, giving rise to its Malagasy name, which means ‘baby bird’. 172

Hook-billed and Helmet Vangas have hefty bills, and are serious predators in the Malagasy forest.

One of Madagascar's most soughtafter birds

Floppy flight

DARK-HEADED VANGAS and CUCKOO-SHRIKE

DISTRIBUTION: Recorded throughout Madagascar in all forest types. Fairly common in eastern rainforest, but very rare elsewhere. ID: Rather similar to Pollen’s Vanga but with a much smaller bill. Also reminiscent of Madagascar Cuckooshrike, but has rufous underparts. VOICE: Distinctive song is a three-part whistle: “whit-wer-whiit”. BEHAVIOUR: Singles or pairs often join mixed feeding flocks. Generally found in the upper-storey of the forest. Feeds on invertebrates. E

Tylas Vanga Tylas eduardi

Length:

21 cm | 8"

WHERE TO SEE: Ranomafana and Andasibe-Mantadia NPs.

Tylas Vanga has been placed in other families, including the oriole family, but recent genetic studies show that it should be placed firmly within the vangas. M

Pollen’s Vanga Xenopirostris polleni

DISTRIBUTION: Uncommon and local in eastern rainforest, although seemingly absent from the north, such as Masoala and Marojejy NPs. ID: Underparts white in male and buffy in female; can look similar to Tylas Vanga, but is always bigger, chunkier, and with a much larger bill. VOICE: Single, penetrating ‘laser gun’ whistle. BEHAVIOUR: Slow-moving species that sometimes joins mixed flocks. Feeds on invertebrates.

Mokazavona Vanga Tylas

Vangamaintiloha Vanga de Pollen

NEAR THREATENED Length:

24 cm | 9·5"

WHERE TO SEE: Ranomafana NP, especially the Vohiparara Trail. 2a

Two other vangas in Madagascar are similar to Pollen’s Vanga, although their ranges do not overlap. The Endangered Van Dam’s Vanga Xenopirostris damii (not illustrated) is found in western dry forest, mainly within Ankarafantsika NP. The more common Lafresnaye’s Vanga X. xenopirostris 2a is endemic to the southwestern spiny forest and can be seen around Toliara, especially at Ifaty. Female Lafresnaye's Vanga

R e

Madagascar Cuckoo-shrike Coracina cinerea

M Vorondavenona Échenilleur Malgache

DISTRIBUTION: Comoros and Madagascar. Fairly Length: 24 cm | 9·5" common throughout Madagascar in all types of forest, WHERE TO SEE: Fairly although unobtrusive. common in almost all forested ID: Although in a completely different family, Madagascar sites, including Ranomafana, Cuckoo-shrike is somewhat similar to the two darkAndasibe-Mantadia, Ankarana, headed vangas above. The grey (rather than white or and Tsingy de Bemaraha NPs. rufous) underparts separate it from both, while its much smaller bill also distinguishes it from Pollen’s Vanga. VOICE: A quiet and easily overlooked series of soft notes. Also a short, explosive call. BEHAVIOUR: Often joins feeding flocks, especially with larger vangas. Like those species, moves slowly through the upper-storey of the forest. Feeds mainly on invertebrates.

174

Buffy underparts

2♀

2♂ Can be very white below

3♀

3♂ Grey underparts

MID-SIZED DARK BIRDS

R e

Crested Drongo Dicrurus forficatus forficatus

M Railovy Drongo Malgache

DISTRIBUTION: Comoros (Anjouan) and Madagascar. Length: 26 cm | 10" Common throughout most of Madagascar in almost any WHERE TO SEE: Very common habitat with woody vegetation, from open savannah to bird that can be seen at all forest gardens to dense forest. sites, and even around hotels on ID: Glossy black bird with a towering crest. Impossible to Nosy Be or St. Marie Island. mistake for anything else. VOICE: Wide variety of whistles, rasps, clicks, and other kinds of notes, including mimicry. Often delivered in long, flowing series. BEHAVIOUR: A bold, aggressive bird that does not hesitate to harass Pied Crows, raptors, and other large birds. Often perches in the open. Frequent member of mixed flocks. Eats invertebrates, many of which are insects taken on the wing.

Madagascar Starling Saroglossa aurata

M Vorontain’omby Étourneau Malgache

DISTRIBUTION: Most of Madagascar, although largely Length: 20 cm | 8" absent from the southwest. Most common in eastern WHERE TO SEE: Ranomafana, rainforest, although also found in western dry forest, plus Andasibe-Mantadia, and a variety of more open and edge habitats. Ankarana NPs are good places ID: Nondescript slim brownish bird. Could be confused to look, especially on snags with Madagascar Bulbul (page 152), but has a shorter tail adjacent to the main roads and lacks that species’ orange bill. Big white patches in the wing through these parks. Sometimes and tail are distinctive in flight. common and at other times scarce. VOICE: Quiet, simple, liquid calls that are easily missed. BEHAVIOUR: Usually in the forest canopy, often on exposed branches and snags, although will descend to the ground to feed. A social species, often found in flocks, although rarely joins mixed flocks. Diverse diet includes fruit, invertebrates, and seeds. I

Common Myna Acridotheres tristis

M Maritaina Martin Triste

DISTRIBUTION: Native to much of southern Asia, and Length: 25 cm | 10" widely introduced elsewhere, including Madagascar. WHERE TO SEE: Virtually Abundant throughout the island, usually close to human omnipresent in humanactivity and in disturbed habitat. modified areas. One of the most ID: A large, dark bird with conspicuous yellow bare skin common and conspicuous birds behind the eye. Shows large white wing patches and a on the High Plateau. white tip to the tail in flight. VOICE: Very noisy, with varied and rather unpleasant vocalizations that usually have an aggressive tone. BEHAVIOUR: A bold, conspicuous bird that perches on man-made structures and feeds mainly on the ground, striding along and snatching up a wide range of animals and plants. Often in flocks. Common Myna is an aggressive non-native species that probably has a negative effect on cavitynesting native species by out-competing them for nest sites. 176

Perches conspicuously in the open

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This introduced species may now be Madagascar's most common bird. It if often seen on man-made structures.

WEAVERS

Nelicourvi Weaver Ploceus nelicourvi

M Fodisaina Tisserin Nelicourvi

DISTRIBUTION: Fairly common in eastern rainforest, Length: 15 cm | 6" some northern Sambirano forest, and the isolated WHERE TO SEE: Ranomafana rainforest on Amber Mountain. and Andasibe areas. Often ID: The combination of yellow on head and brick-red nests near the restaurant at undertail is distinctive. Male has a mostly black head. the Feon’ny Ala Hotel near VOICE: Buzzy “tzz-chhh-chhh-chhh”. Andasibe. BEHAVIOUR: Unlike most weavers, nests solitarily, rather than in colonies. Its beautiful and distinctive nest is a hanging sack with a long entrance tunnel, and is often built over a stream. Frequently joins mixed flocks that include tetrakas. Feeds mainly on insects. E

Sakalava Weaver Ploceus sakalava

M Fodisahy Tisserin Sakalava

DISTRIBUTION: Common in dry western, southern, and Length: 15 cm | 6" northern Madagascar. Found in western dry forest, spiny WHERE TO SEE: A conspicuous forest, and various open habitats, including those heavily resident in the dry lowlands, modified by humans. such as around Toliara, ID: Yellow-headed male is distinctive. Female is very Morondava, and Ankarafantsika similar to a female or non-breeding male fody, but is and Ankarana NPs. larger, with a larger bill, tan-striped face, and a reddish eye ring. VOICE: Chirpy and swizzling notes, often delivered in an exuberant chorus by many birds at a colony. BEHAVIOUR: Nests socially in bustling colonies, unlike Nelicourvi Weaver. The nest is similar to that of Nelicourvi Weaver, though is usually more loosely woven. Often feeds on the ground, searching for seeds and insects. When not breeding, forms large flocks that often include Madagascar Fody. The Sakalavas are the people who populate most of western Madagascar, and are found in much the same places as their namesake weaver. In some places, it is considered a good omen if weavers nest over or near the house of the village mayor. R

Madagascar Fody Foudia madagascariensis

M Fody Foudi Rouge

DISTRIBUTION: Probably originally endemic to Length: 13 cm | 5" Madagascar, but has been introduced to almost every WHERE TO SEE: Common island of the Malagasy region. Found in almost any habitat almost everywhere, including except closed-canopy forest. downtown Tana. ID: The bright red plumage of breeding male is distinctive. Female and non-breeding male are quite drab (see Sakalava Weaver). The rarer Forest Fody F. omissa (not illustrated) is bigger than Madagascar Fody but breeding male has a dark green belly (though beware that male Madagascar Fody coming into breeding plumage can match this pattern). Females of the two species are extremely similar. Forest Fody is found in the interior of eastern rainforest. VOICE: High-pitched “tsit” given singly, or in a quick series. BEHAVIOUR: Male is bold and conspicuous when breeding. Non-breeding birds form flocks. Eats seeds, which are either gathered on the ground or directly from plants. 178

1♀

1♂

Solitary nester

2♀

2♂

Colonial nester

3♀

Females and nonbreeeding males look very similar

3♂

In October and November, most males transition into breeding plumage

3♂

Reptiles l Almost 400 species of reptile have been described from Madagascar, of which more than

90% are endemic.

l There are four orders of reptiles in the world: crocodiles, tuatara, lizards and snakes, and

turtles. All of these except New Zealand’s tuatara are found in Madagascar. Thirteen families are represented. l The majority of Malagasy reptiles are diurnal, but about 85 species are nocturnal, including most geckos and many snakes. Some reptiles can be active both night and day, and even some strongly nocturnal species can occasionally be active by day. Nocturnal reptiles generally have vertical ‘slit’ pupils while diurnal ones have round pupils, although there are exceptions. l Reptiles are remarkably common and conspicuous in Madagascar, one of the easiest places on Earth for a visiting naturalist to amass an impressive reptile list. Many species can be seen on exposed rocks or tree trunks. Chameleons can be hard to spot during the day, but sleep on low-hanging branches, making them easier to find at night. Plated lizards and skinks often reveal themselves by the noise of their movement through the leaf-litter. Poking around dead wood and taking night walks will increase your chances of finding more secretive and nocturnal reptiles. l Many reptiles can be seen year-round in Madagascar, but the wet season is the best time, particularly for snakes and chameleons. During the dry season, you might spot only a few snakes, and fail to see any leaf chameleons. l Most Malagasy reptiles lay eggs, although a few give birth to live young. The eggs are buried in the ground or hidden in crevices. Most hatch after 1–3 months, but the eggs of certain species, like Parson’s Chameleon hatch only after more than a year. l The identification of many reptiles in Madagascar is not difficult. The first thing to consider is range: similar species often do not overlap geographically. Secondly, basic colours and patterns distinguish most species. Colour can be misleading or non-diagnostic for chameleons, where structure is more useful. For herpetologists, characteristics of the scales provide the most definitive means of identifying most species. l This guide covers about 30% of Madagascar’s reptiles, including most of the species likely to be seen in frequently visited sites. Refer to Amphibians and Reptiles of Madagascar by Glaw and Vences for thorough coverage (see Further reading (page 327) for details). l In many Malagasy reptiles, males and females are similar. When there is a difference, as in ‘typical’ chameleons and day geckos, males tend to be larger and more colourful. In the case of chameleons, males also have more elaborate structures on their head. Leaf chameleons form an exception to the rule: females are larger than males. l There is no up-to-date standardization of common names for reptiles. Common names novel to this book are marked with asterisks. Herpetologists and most local guides use scientific names exclusively. l In some lizards, total length (TL) is the most useful measurement. In others, the tail length is highly variable, or the tail is often curled up (as in typical chameleons), making the length from the tip of the snout to the vent (SVL) a more useful measurement. For turtles, the carapace length (CL) is given. 180

Panther Chameleon (page 192)

Gold-dust Day Gecko (page 222)

Madagacasar tree boa (page 238)

Radiated Tortoise (page 182)

TORTOISES and TERRAPINS

Madagascar supports several species of terrestrial tortoise and aquatic terrapin, in three different families. Many are rare, local, and in danger of following the island's two giant tortoises into extinction. Included here are the species most likely to be seen by visitors. The standard measurement for tortoises, terrapins, and turtles is carapace length (CL), the length of the curved line that traces the top of the carapace. E

DISTRIBUTION: Dry spiny forest of southwest. ID: The ‘radiated’ pattern on the carapace of the adults is variable but distinctive, though it may be absent on older individuals. This is Madagascar’s most frequently seen tortoise. BEHAVIOUR: Remains hidden much of the time. Sometimes located by its tracks in the sand. Can live well over 100 years.

CRITICALLY ENDANGERED Length:

CL < 40 cm | 16"

WHERE TO SEE: Captive specimens can be seen at many hotels and parks. Sightings in the wild are increasingly rare, but possible around Ifaty (north of Toliara), at Berenty, and elsewhere in the spiny forest.

Spider Tortoise Pyxis arachnoides

DISTRIBUTION: Spiny forest of the southwest, close to the coast. ID: Much smaller than full-grown Radiated Tortoise. ‘Spider’ pattern is quite different from the ‘radiated’ pattern of that species. BEHAVIOUR: Similar to Radiated Tortoise.

Sokatra, Sokake Tortue Rayonnée

Radiated Tortoise Astrochelys radiata

M Tsakafy Tortue-Araignée

CRITICALLY ENDANGERED Length:

CL 12 cm | 5"

WHERE TO SEE: Lucky visitors might find it in the spiny forest, as around Ifaty and Berenty.

The Critically Endangered Flat-tailed Tortoise P. planicauda 2a is very similar to Spider Tortoise, but has a different range, occurring only in the central west. Look for it in Kirindy Forest, mainly during the wet season.

Terrapins [2 families; 3 genera; 4 species]

DISTRIBUTION: One species, Madagascan Big-headed Turtle Erymnochelys madagascariensis 3a , is endemic, while three (Yellowbelly Mud Turtle Pelusios castanoides (not illustrated); East African Black Mud Turtle Pelusios subniger 3b and Marsh Terrapin Pelomedusa subrufa 3c ) are also found in Africa. Three species are mainly found in the west, one (P. subniger) is in the east, and all are rare or absent from the centre of Madagascar. ID: These aquatic animals have a different structure from the tortoises: they are more flattened, and also lack the distinctive markings normally shown by tortoises. BEHAVIOUR: Sometimes seen sunbathing, or just as a head emerging from the water. Occasionally found walking across dry ground, including roads. 182

M Kapidrano Tortue d’Eau Douce

Length: CL < 41 cm | 16" (Madagascan Big-headed Turtle) CL < 25 cm | 10" (other species) WHERE TO SEE: At least one species can be found around most wetlands, but seeing any Malagasy terrapin takes a lot of luck. Sightings are most likely in the west during the wet season.

SEA TURTLES and CROCODILE

Five of the world’s eight species of sea turtles are found along Madagascar’s coast. Only the two covered below are seen frequently. 1

Green Sea Turtle Chelonia mydas

DISTRIBUTION: Tropical and subtropical seas worldwide. Found all around Madagascar’s coast. ID: A huge oceanic turtle. Similar to Hawksbill Sea Turtle, but larger, lacks hooked snout, and adults lack serrations along the edges of the shell. BEHAVIOUR: Mainly ocean-dwelling, coming onshore only to lay eggs. Unlike other ocean turtles, adults are predominantly vegetarian. Frequently hunted and eaten by humans.

Nile Crocodile Crocodylus niloticus

M Fanozaty Tortue Verte

ENDANGERED Length: Weight:

CL 1–1·5 m | 3–5’ < 300 kg | 660 lb

WHERE TO SEE: Most often seen around Nosy Be, sometimes from the ferry between the mainland and the island. Also frequent at the ‘Three Bays’ near Diego-Suarez.

Hawksbill Sea Turtle Eretmochelys imbricata

DISTRIBUTION: Found in tropical seas worldwide. In Madagascar, occurs along most of the coastline. ID: Smaller than Green Sea Turtle, with a hooked snout, and strong serrations along the edges of the shell. BEHAVIOUR: Similar to Green Sea Turtle, but feeds mainly on marine invertebrates and algae. If when snorkelling you observe a turtle eating coral, it is very likely to be this species. 3

M Fanohara Tortue Imbriquée

CRITICALLY ENDANGERED Length: Weight:

CL ·6–1 m | 2–3·3’ < 140 kg | 300 lb

WHERE TO SEE: Often seen around Nosy Be, especially in the coral reefs of the Nosy Tanikely marine reserve.

M Voay Crocodile du Nil

DISTRIBUTION: Throughout sub-Saharan Africa. Length: TL < 4 m | 13’ In Madagascar, most common in the west. Weight: < 1,000 kg | 1 tonne The Malagasy population is estimated at 30,000. WHERE TO SEE: Generally ID: Aquatic habits, streamlined shape, and armoured uncommon, but usually easy to skin identify even small juveniles. The large adults see in Ankarafantsika NP. Can are unmistakable by size alone; they dwarf all other occasionally be spotted from Malagasy reptiles. major road bridges that cross BEHAVIOUR: Feeds on a wide variety of animals, from fish less-disturbed western rivers. to zebu cattle. Often leaves the water to bask. Courtship is elaborate, and the parents are quite attentive to their eggs and young. Crocodilians are the last living non-avian members of the archosaurs, the group of reptiles that includes the dinosaurs, and which dominated the earth before mammals rose to prominence. They have changed little in the last 65 million years. Amazingly, ‘crocs’ are actually more closely related to birds than to other reptiles. 3

184

Rounded snout

Adults have a smooth edge to the shell

Sharp bill

Points along edge of shell

TINY LEAF CHAMELEONS OF THE NORTH

Some of world’s smallest reptiles belong to the endemic genus Brookesia; the smallest less than 3 cm long. There are at least 30 species of leaf chameleon, although new species are still being described. The Elongate Ancient Leaf Chameleon (page 188) has been moved into a new genus: Palleon. Although only nine species are covered here, these are the ones that most visitors are likely to see, especially away from the far north. Leaf chameleons are generally small and brownish. They spend the day walking slowly through the leaf-litter, then climb up into the understorey vegetation to sleep at night. Sightings are much more common in the wet season, especially in the dry western forest. These primitive chameleons lack significant capacity to change colour. Their tails are fairly short and not usually rolled up in classic chameleon fashion. Females are larger than males. E

Minute Leaf Chameleon Brookesia minima

Rakolaka Kely

DISTRIBUTION: Small distribution in the Sambirano ENDANGERED rainforest of the northwest. Length: 3 cm | 1" ID: Tiny size makes it unmistakable within its range. WHERE TO SEE: Wet season in Among the world’s smallest reptiles. Beige above and dark Lokobe NP, on Nosy Be. brown below, with fairly smooth skin. BEHAVIOUR: When threatened by a predator, this species, and most other small leaf chameleons, tuck in their legs and roll to the ground, remaining motionless to 'disappear' into the leaf-litter. E

Montagne d’Ambre Leaf Chameleon

Brookesia tuberculata

DISTRIBUTION: Rainforest on Amber Mountain. ID: Uniformly brown or beige in colour. The legs are sturdier than those of other small leaf chameleons. Tiny size identifies it within Amber Mountain NP. This is another of the world’s smallest reptiles.

VULNERABLE Length:

Northern Madagascar holds the highest diversity of leaf chameleons, with at least five species occurring on Amber Mountain. E

Antongil Leaf Chameleon Brookesia peyrierasi

DISTRIBUTION: Small distribution in the rainforest of the northeast, in the Masoala area. ID: Brown or mottled grey and brown. Tiny size makes it unmistakable within its small range.

Male leaf chameleons generally have a longer tail than the female, although females are larger overall.

3 cm | 1"

WHERE TO SEE: Endemic to Amber Mountain. Usually easy to find there, and very common during the wet season. Often found around the main picnic area and campsite. M

Rakolak’i Nosy Mangabe ENDANGERED

Length:

3–4 cm | 1–1·5"

WHERE TO SEE: Wet season on the island of Nosy Mangabe, between Maroantsetra and Masoala. 3♂

3♀

186

Rakolaka Madinika

Minute Leaf Chameleon was long thought to be Madagascar’s smallest reptile. But in 2012, a new, even smaller species of leaf chameleon (B. micra) was discovered on the island of Nosy Hara near Madagascar’s northern tip, highlighting that much still remains to be discovered about Malagasy reptiles.

Often shows beige above and dark brown below Small spikes along sides of spine. Knobby spikes above the eye

2♂

3♀

Small spikes along sides of spine

WIDESPREAD MEDIUM TO LARGE LEAF CHAMELEONS

In 2013 the ancient leaf chameleons were removed from the genus Brookesia and placed in the new genus Palleon, an addition to Madagascar’s traditional list of three chameleon genera: Brookesia, Calumma and Furcifer. E

Plated Leaf Chameleon Brookesia stumpﬃ

DISTRIBUTION: Widespread in both rainforest and dry forest across the west and north. ID: Fairly large and uniformly pale brown dwarf chameleon. Could be confused with Brygoo’s Leaf Chameleon in Ankarafantsika NP, but Plated Leaf Chameleon is larger and has much smaller spiny plates along its back. E

Brygoo’s Leaf Chameleon Brookesia brygooi

Rakolaka Bory Tandroka

Length:

8–9 cm | 3–3·5"

WHERE TO SEE: Common in Ankarana NP, especially during the wet season. May also be found in Amber Mountain and Ankarafantsika NPs and on Nosy Be.

Rakolaka Tandrefana

DISTRIBUTION: Dry forests of the west. Length: 7–8 cm | 3" ID: Usually pale brown above and dark brown below. WHERE TO SEE: During the Similar to Plated Leaf Chameleon (which has a wet season in Ankarafantsika more northern range although the two overlap at and Tsingy de Bemaraha NPs Ankarafantsika NP), but is smaller, darker, less uniform and Kirindy Forest. Most often in coloration, and has larger spiny plates along its encountered at night, sleeping back. Decary’s Leaf Chameleon B. decaryi (not illustrated) is on low branches. endemic to Ankarafantsika NP. It is very similar to Brygoo’s Leaf Chameleon, but has a fatter body, shorter tail, and less prominent bumps above its eyes. The recently described Bruno’s Leaf Chameleon B. brunoi (not illustrated) is similar to Brygoo’s Leaf Chameleon, but has a discrete range, being found exclusively in Parc Anja, where is it sometimes located by local guides. E

Brown Leaf Chameleon Brookesia superciliaris

DISTRIBUTION: Widespread in eastern rainforest. ID: Large, variably marked with brown and beige, and with large, triangular projections above the eyes. This is the most common, or only, leaf chameleon in much of the eastern rainforest.

Elongate Ancient Leaf Chameleon Palleon (Brookesia) nasus

DISTRIBUTION: Rainforest of the southeast. ID: Distinctively pointed nose. Lacks the big triangular projections above the eye of Brown Leaf Chameleon.

188

Rakolaka Tatsinanana

Length:

6–9 cm | 2–3·5"

WHERE TO SEE: Found most frequently around Andasibe, but also possible at Ranomafana and in other eastern rainforest sites.

Rakolaka Fisa-batana VULNERABLE

Length:

< 9 cm | 3·5"

WHERE TO SEE: Ranomafana is the best site, although it is still uncommon there. Best searched for on the Vohiparara Trail.

1 2

Pale brown

Large cones above the eyes

Small spikes on chin

Small bump on the tip of the snout

Usually pale above and dark below

THE LARGEST CHAMELEONS ON EARTH

Most of Madagascar’s chameleons are in two genera of ‘typical chameleons’: Furcifer and Calumma. There is no simple way to separate them, although Calumma are generally found in rainforest in the north and east, while Furcifer are more widespread and better able to adapt to degraded habitat. They vary from being small (similar in size to large leaf chameleons) up to 70 cm (2'), the largest chameleons on earth. These are the classic chameleons, often colourful, and with the astonishing ability to rapidly change colour. The males of many species sport impressive structures on their head, while the females are more modest. All typical chameleons are arboreal and active during the day, although they are sometimes found walking on the ground to cross open areas. They are frequently spotted sleeping on low branches at night. E

Oustalet’s Chameleon Furcifer oustaleti

M Tarondro Caméléon d’Oustalet

DISTRIBUTION: Occurs throughout the island. Found Length: SVL < 28 cm | 11" inside western dry forest, but largely avoids the interior WHERE TO SEE: One of the of eastern rainforest. Common in degraded areas, even in most common Malagasy towns and gardens. chameleons; possible almost ID: Big rounded casque (rounded helmet-like projection) anywhere, although most on the back of the head of the male. No horns on the nose. common in the west. More than 45 fairly large spikes along the spine. Variable, but usually not colourful: grey, brown, or dull green. Warty Chameleon (page 206) can be extremely similar to Oustalet’s Chameleon and some females cannot be separated. Jewelled Chameleon (page 192) is also widespread and common in many habitats. It averages much smaller, has a less prominent casque, lacks spikes along the spine, and females are always colourful (although highly variable). See Panther Chameleon (page 192) for separation from that species. Furcifer nicosiai (not illustrated) resembles Oustalet’s Chameleon, but is smaller. It can be found in Tsingy de Bemaraha NP and Kirindy Forest. E

Parson’s Chameleon Calumma parsonii

DISTRIBUTION: In the eastern rainforest zone, but not usually inside of thick forest. Prefers forest edges, roadsides, and sometimes even degraded areas. ID: Large, with a big, triangular head. The female is green, while the male can be green, turquoise, or yellow. Both usually show several diagonal dark slashes along the sides. The male has twin horns on its nose. See O’Shaughnessy’s Chameleon (page 198) for distinction from that species.

Tarondro Maitso Be Caméléon de Parson

M F

NEAR THREATENED Length:

SVL ♂ < 30 cm | 12" ♀ < 20 cm | 8"

WHERE TO SEE: Andasibe and Ranomafana areas.

2ju

2♀

190

Female lacks a nose horn

1♀

1♂

Male grey or brown 1♂

Female can be bright green, but is usually duller than the male

Male Oustalet's Chameleons can be up to 70 cm (28") long (including the tail), and females up to 40 cm (16")

2♂

Male has a large, triangular head and a prominent nose horn

COMMON CHAMELEONS OF DEGRADED AREAS

Along with Oustalet’s Chameleon (page 194) and Warty Chameleon (page 206), these are the species most likely to be seen away from prime forest habitats. E

Panther Chameleon Furcifer pardalis

M Sakorikita Caméléon Panthère

DISTRIBUTION: Found in a variety of habitats, mainly Length: SVL ♂ < 25 cm | 10" in the northern third of Madagascar. It does occur inside ♀ < 17 cm | 7" forest, especially western dry forest, but is more often in WHERE TO SEE: Common in degraded areas. the north and northeast. Watch ID: Males have highly variable coloration, both for it around Diego-Suarez, individually and geographically. Colourful individuals can Ankarana NP, Nosy Be, and show a rainbow of colours, and are among the most beautiful Maroantsetra. Frequently seen of chameleons. Male shows a bump on the tip of the nose and crossing the road, even the a small casque, which separate even a dull male from Oustalet’s main northern artery, the RN6. Chameleon. Female Oustalet’s Chameleon can be very similar, although Panther Chameleon usually has more grey-and-orange or grey-and-pinkish (rather than brown or green) coloration, and shows a small bump on the top front of the nose; Oustalet’s Chameleon has a smooth nose. 1♂

Diego-Suarez

1♂

Nosy Be

Ankarana NP

Jewelled Chameleon Furcifer lateralis

DISTRIBUTION: Found in a wide variety of habitats in the southern two-thirds of Madagascar. ID: Medium-sized chameleon with a moderate casque. Male usually green, females are a variety of colours. Both usually show a pale line and three dark circles along the sides. Oustalet’s Chameleon is larger and lacks these flank markings. Panther Chameleon can be similar, but their ranges do not overlap. Very similar to several scarce chameleon species, but full coverage of these is beyond the scope of this guide. A host of Malagasy beliefs and traditions surround chameleons. In some places, chameleons are revered as reincarnations of the ancestors, and it is forbidden to point at them, except with a curled finger. Many people fear these creatures, and the sight of a vazaha (foreigner) touching or approaching them can be shocking. Some people believe that you will be cursed if you touch a chameleon, and will die a death as slow as the chameleon’s gait. 192

1♂

Length:

Tanalahy

SVL ♂ < 14 cm | 5·5" ♀ < 11 cm | 4·5"

WHERE TO SEE: Common in many places. This is the most frequently encountered chameleon in the central highlands, including cities like Tana and Antsirabe. 2♀

Female is variable in colour

1♀

1♂

Marojejy NP 1♂

Ambanja 1♂

Male shows a small nose horn and casque Female usually shows some orangish or pinkish coloration

Amber Mountain NP opposite and above:

Male Panther Chameleons are highly variable, both individually and geographically

2♂

Males are drab, often green, but note the pale stripe and three ovals on the side

2♀

EASTERN RAINFOREST CHAMELEONS: SMALL

Perinet (Band-bellied) Chameleon

Calumma gastrotaenia

DISTRIBUTION: Common in rainforest from central to southern Madagascar. ID: A small, mainly green chameleon without any outstanding structural features. Male shows a bold white line along the sides; female is more muted. See similar Glaw’s Chameleon. E

Glaw's Chameleon Calumma glawi

Length:

Tana Maitso Kely Tatsinanana SVL < 7 cm | 3"

WHERE TO SEE: Often abundant on night walks along the main road through Ranomafana NP. Also occurs around Andasibe. M

Tana Maitso Kelin’i Ranomafana

DISTRIBUTION: Small range in mid-elevation rainforest Length: SVL < 7 cm | 3" in the southeast. WHERE TO SEE: Uncommon ID: Small green chameleon that is very similar to Perinet around Ranomafana NP. Mainly Chameleon. Traits which may be present, and which can found on night walks. identify Glaw’s Chameleon, include pale blue spots on the body; black on the casque of the males; and a larger flap (occipital lobe) at the back of the head, especially on males. E

Canopy Chameleon Furcifer willsii

DISTRIBUTION: Patchy distribution in eastern rainforest. ID: The male has two large horns protruding from its nose, but female lacks horns. Both show only a small casque. Usually green, with some markings overlaid. Often shows pale, yellowish diagonal stripes along the sides. When present, these serve to separate female Canopy Chameleon from the similar Perinet Chameleon. E

Big Nose Chameleon Calumma nasutum

DISTRIBUTION: Widespread in eastern rainforest and forest edge. Also occurs in some places in the Sambirano rainforest of the north. ID: A small chameleon with a warty horn on the nose. Coloration variable but usually dull. Similar to several other less frequently seen species, but full coverage of their separation is beyond the scope of this guide. 4

194

Sometimes called Nose-horned Chameleon

Sakorkita misy Tandroka Roa Length:

SVL < 8 cm | 3"

WHERE TO SEE: Uncommon in Andasibe and Ranomafana.

Tana Kely misy Orona

Length:

SVL < 5 cm | 2"

WHERE TO SEE: Fairly common in most eastern rainforest sites, including Andasibe and Ranomafana.

1♂

2♂

Female has smaller lobes at the back of the head than male

Male has pale lines along the sides

Female nondescript

1♀

3♀

3♂

Large nose horn

Dull coloration

Male has large nose horns (which female lacks). Both sexes often show yellow stripes on the body

Calumma fallax (not illustrated) is similar to Big Nose Chameleon, and both are often seen around Ranomafana. C. fallax has a slightly larger casque and scales that are more heterogenous in size.

EASTERN RAINFOREST CHAMELEONS: MEDIUM to LARGE I

Cryptic (Blue-legged) Chameleon

Calumma crypticum

Tarondro Pentina Manga

DISTRIBUTION: Middle- and high-elevation rainforest, Length: SVL < 12 cm | 5" mainly in the east. WHERE TO SEE: Fairly common ID: Large flaps (occipital lobes) at the back of the head. in Ranomafana NP. Sleeping Male has a single modest nose horn. Often colourful, individuals are often located at marked with blue, green, and rich brown. Often has blue night along the main road. markings on its legs, hence its alternate common name. Short-horned Chameleon is very similar: the male Cryptic Chameleon has a slightly longer nose, but females may be inseparable. E

Short-horned Chameleon

Calumma brevicorne

DISTRIBUTION: Patchy distribution in middle-elevation rainforest of the east. ID: Structure very similar to Cryptic Chameleon. Male Short-horned Chameleon shows a slightly longer brick-red nose horn. Both sexes are generally less colourful than Cryptic Chameleon, more brown-and-grey, though females may be inseparable.

Length:

Tarondro Be Sofina SVL 11–17 cm | 4·5–7"

WHERE TO SEE: Fairly common around Andasibe.

Chameleons are famous for changing colour to match their environment. However, camouflage is just one of the ways they use this ability, since they also make colour changes as a visual display of their mood, or in response to cues like changing light level or temperature.

2♀

196

Females of both of these species have only a small bump on the nose

1♂

Male has a small horn on the nose

1♀

2♂

Male has a reddish horn on the nose 2♂

EASTERN RAINFOREST CHAMELEONS: MEDIUM to LARGE II

Two-banded (Belted) Chameleon

Furcifer balteatus

DISTRIBUTION: Middle-elevation rainforest and degraded habitat in the southeast. ID: Large, usually bright green chameleon. Male shows twin horns on the nose. Can almost always be identified by the long, pale slash that runs along the side, from behind the top of the head down to the hind legs.

O’Shaughnessy’s Chameleon

Tanakely Mitsipibatana ENDANGERED

Length:

SVL ♂ < 18 cm | 7" ♀ < 15 cm | 6"

WHERE TO SEE: Ranomafana area. Most common in the degraded areas around the village of Ranomafana. M

Tarondro Maitso Be misy Sofina

Calumma oshaughnessyi

DISTRIBUTION: Eastern rainforest of the southeast. ID: Large green chameleon. Can be similar to smaller individuals of Parson’s Chameleon (page 190). Both species have triangular heads, but O’Shaughnessy’s Chameleon has flaps of skin at the back of the head (occipital lobes) that are absent in Parson’s Chameleon. O’Shaughnessy’s Chameleon also shows heterogeneous scales along its sides, with a regular pattern of much larger scales interspersed among smaller ones, whereas Parson’s Chameleon has scales all of roughly the same size.

2♀

198

VULNERABLE Length:

SVL < 18 cm | 7"

WHERE TO SEE: Ranomafana NP. Frequently found at night along the main road, often sleeping quite high in the forest.

The scales on the sides vary significantly in size

1â&#x2122;&#x201A;

Male has twin horns on the nose; female lacks prominent nose horns

2â&#x2122;&#x201A;

Male has a large triangular head and prominent twin horns on the nose

CHAMELEONS OF THE NORTH I

Amber Mountain Chameleon Calumma amber

DISTRIBUTION: Endemic to the rainforest of Amber Mountain. ID: Within its range, the big ‘elephant ears’ at the back of the head are distinctive. Arthur’s Chameleon is larger, usually green, and has a large, triangular head. Similar to Short-horned and Cryptic Chameleons (page 196), but does not overlap in range. E

Arthur’s Chameleon* Calumma ambreense

DISTRIBUTION: Found in mid-elevation rainforest in the north, on Amber Mountain and perhaps elsewhere. ID: Very similar to the closely related O’Shaughnessy’s Chameleon (page 198), but found in the far north instead of the southeast. Big triangular head distinctive within its small range.

Tarondro Marevaka

NEAR THREATENED Length:

SVL 7–11 cm | 3–4·5"

WHERE TO SEE: Can usually be located during the day along the trails in Amber Mountain NP.

Tarondro Marevaka

NEAR THREATENED Length:

SVL < ~ 18 cm | 7"

WHERE TO SEE: Fairly easy to find in Amber Mountain NP during the wet season; more difficult in the dry season. 2♀

2♀

Female lacks nose horn, and can show yellowish coloration

200

1â&#x2122;&#x201A;

Male has a small nose horn; female lacks a nose horn, and is usually green

2â&#x2122;&#x201A;

Male has a smaller nose horn than does the closely related O'Shaughnessy's Chameleon, which only occurs in the southeast; the ranges of the two species do not overlap

CHAMELEONS OF THE NORTH II

DISTRIBUTION: Forest and adjacent human-modified habitats in the north. Probably a complex that contains multiple undescribed species. ID: Male shows two large horns on the nose, but female lacks horns. Overall colour green or yellow-green, and often has white lips. Male has a pale line along the side, while female shows two pale spots.

M Tanakelin’ny Avaratra Andrefana

Petter’s Chameleon Furcifer petteri complex

Northern Blue-nosed / Boettger’s Chameleon Calumma linotum / boettgeri complex

VULNERABLE Length:

SVL ♂ < 9 cm | 3·5" ♀ < 7 cm | 3"

WHERE TO SEE: Ankarana and Amber Mountain NPs and the surrounding areas. Often found in scrubby habitat in the town of Joffreville. M

Tana Kely Mitsingoloka

DISTRIBUTION: Rainforest and secondary habitats in Length: SVL < 6 cm | 2·5" the north and northeast. C. linotum is apparently found WHERE TO SEE: C. linotum is only on Amber Mountain, while the C. boettgeri complex fairly common in and around (containing multiple undescribed species) is more Amber Mountain NP. C. boettgeri widespread. is possible around Nosy Be and ID: Small chameleon with a small casque, but a big horn Marojejy NP, often in secondary on the nose of both sexes. Variable in colour, but usually shows habitat. bands on the tail. C. linotum usually has a bright blue nose horn. Big Nose Chameleon (page 194) overlaps in range, and is very similar, but lacks the small flap at the back of the head (occipital lobe) shown by Boettger’s Chameleon.

2♂

202

Both sexes have a small casque and a big nose horn

1♂

1♀

Female lacks a nose horn

2♂

2♀

CHAMELEONS OF THE WEST I

White-lined (Antimena) Chameleon Furcifer antimena

DISTRIBUTION: A small area of spiny forest in the southwest. ID: Green and brown, with male brighter than female. Usually shows a white line and three dark circles on the side. Within its range, identified by the large warty nose horn of the male, and the much smaller one of the female.

Tarondro Ben’ny Atsimo Andrefana VULNERABLE

Length:

WHERE TO SEE: Spiny forest around Toliara and Ifaty, especially during the wet season. M

Rhinoceros Chameleon Furcifer rhinoceratus

DISTRIBUTION: Restricted to a small area of dry forest in the northwest. ID: Coloration variable but usually dull. Within its range, identified by the huge warty horn on the nose of the male, and the much smaller one on the female.

2♂

204

SVL ♂ < 17 cm | 7" ♀ < 10 cm | 4"

Sakorikita Malemy Orona

VULNERABLE Length:

SVL ♂ < 14 cm | 5·5" ♀ < 7 cm | 3"

WHERE TO SEE: Uncommon in Ankarafantsika NP, seen mainly during the wet season.

Male has a chainsaw-like projection on the nose

1â&#x2122;&#x201A;

Male has a large casque, spikes along the spine and a nose horn; these are less well developed in the female

CHAMELEONS OF THE WEST II

Warty Chameleon Furcifer verrucosus

DISTRIBUTION: Widespread in dry forest and disturbed habitat across most of western and southwestern Madagascar. Present, but less common, in parts of the central highlands. ID: Large and generally dull-coloured chameleon that can easily be mistaken for Oustalet’s Chameleon (page 190). Male Warty Chameleon has larger spikes on the throat and along the spine; it has fewer than 40 spikes on the spine, whereas Oustalet's Chameleon has more than 45. Female generally lacks spikes along the spine, unlike female Oustalet’s Chameleon which has prominent spikes, although some may be indistinguishable. E

Labord’s Chameleon Furcifer labordi

DISTRIBUTION: Wide but patchy distribution in western dry forest. ID: Structure similar to White-lined Chameleon (page 204), but does not overlap in range. Male has white lips and white lines along the sides. Female has green bars along the sides, and some show blue spotting.

M Sakorikita Be Caméléon Verruqueux

Length:

SVL ♂< 27 cm | 11" ♀< 10 cm | 4"

WHERE TO SEE: Common in much of the west and southwest. Often in towns and gardens. One reliable spot is the Arboretum d’Antsokay near Toliara. Also occurs in Parc Anja, alongside Oustalet’s Chameleon.

Tarondro Tsara Tarehy VULNERABLE

Length:

SVL ♂ < 14 cm | 5·5" ♀ < 8 cm | 3"

WHERE TO SEE: Kirindy Forest, mainly during the wet season.

Uniquely among reptiles, all adult Labord’s Chameleons die at the end of the wet season, after they have bred. The breeding population the following year is comprised entirely of individuals that have hatched out of the previous generation’s eggs – a breeding cycle analogous to that of an annual plant! 2♀

2♂

206

1â&#x2122;&#x20AC;

Female Warty Chameleon can be indistinguishable from Oustalet's Chameleon Male Warty Chameleons can be up to 57 cm (22") long; not quite as large as the largest Oustalet's Chameleons, but still impressive

Warty Chameleon is the most common chameleon in the arid southwest 1â&#x2122;&#x201A;

SWIFTS I: WEST AND SOUTHWEST

Madagascar’s (reptilian) swifts are something of a mystery, as they are closely related to iguanids, found only in the Americas, and on some Pacific islands. They probably originate from the ancient time when Madagascar was connected to South America via Antarctica. Unlike the iguanids, Madagascar’s swifts are found only in dry habitats, not in rainforest. It was only recently recognized that Madagascar’s seven swift species constitute their own regionally endemic family, the Opluridae: previously they were considered part of the Igaunidae. ‘Swifts’ is a common name shared by two very different groups: iguana-like lizards from Madagascar and a widespread family of birds (see page 136). E

DISTRIBUTION: Found on the ground in spiny forest, dry forest, and degraded habitat across the west and southwest. ID: Variable, but usually boldly marked. Boasts a ‘third eye’, a dark spot on the top of the head which contains photoreceptors. BEHAVIOUR: Generally stands motionlessly, but is capable of dashing with great speed. E

Three-eyed Lizard Chalarodon madagascariensis

Dangalia

Length: Average TL ~ 20 cm | 8" WHERE TO SEE: Common in spiny forest in the southwest. Also found in some western dry forest sites including Kirindy Forest and Zombitse-Vohibasia NP.

Merrem’s Madagascar Swift Oplurus cyclurus

Androngovato misy Sanga

DISTRIBUTION: Found mainly on tree trunks in dry Length: TL < 25 cm | 10" forest of the southwest and west. Most common in the WHERE TO SEE: Fairly common southwest. in the spiny forest of the ID: The extremely spiny tail separates it from Dumeril’s southwest, for example around Madagascar Swift and Grandidier’s Madagascar Swift Toliara and Ifaty. (both page 210). In some places occurs alongside Cuvier’s Madagascar Swift, although that species generally prefers moister habitats. Merrem’s can be separated by its narrow (rather than broad) black collar, often bordered in white. A close look at the tail also reveals that there are no small rows of scales between the big spiny scales, unlike in Cuvier’s Madagascar Swift. R

Cuvier’s Madagascar Swift Oplurus cuvieri

DISTRIBUTION: Madagascar and Grande Comore. Found on tree trunks and on the ground in western dry forest and degraded habitat. ID: Similar to Merrem’s Madagascar Swift, but usually larger, with a much broader black collar, and has small scales between the large spines on its tail.

208

Length:

Androngon-kazo TL < 37 cm | 14·5"

WHERE TO SEE: Very common and conspicuous in Ankarafantsika NP, including around the park headquarters. Also common in Tsingy de Bemaraha NP and Kirindy Forest.

Narrow black 'collar'

The '3rd eye' on the back of the head actually contains photoreceptors!

These two species can be separated by a careful look at their tails. There are small rows of scales between the big spiny scales on Cuvier's 3 , but no such scales on Merrem's 2 . Broad black 'collar'

SWIFTS II: ROCK-DWELLING SWIFTS OF THE SOUTHWEST 1

Dumeril’s Madagascar Swift Oplurus quadrimaculatus

Androngovato Atsimo Atsinanana

DISTRIBUTION: Classic habitat is rocky, arid areas of the southwest, but occurs in a variety of habitat from sea level up to 2,000 m (6,500') in southwest and south-central Madagascar. ID: A large lizard that lacks the enlarged tail spines of Merrem’s Madagascar and Cuvier’s Madagascar Swifts (page 208). Very similar to Grandidier’s Madagascar Swift, but that species usually shows a single pale line along the middle of its back, whereas Dumeril’s Madagascar Swift shows either an unlined back, or two pale lines along the sides (rather than the centre) of the back. In the far south, such as around Fort Dauphin and Berenty, confusion is possible with the very similar Marked Madagascar Swift O. saxicola 1a , but that species never shows stripes on the sides of the back. BEHAVIOUR: Often sits on rocks. Can be surprisingly tame.

Length:

WHERE TO SEE: Common and conspicuous around Isalo, including the tourist lodges. Also common in Parc Anja.

1a Marked Madagascar Swift Oplurus saxicola

Grandidier’s Madagascar Swift

Oplurus grandidieri

DISTRIBUTION: Found in south-central Madagascar. Lives on rocks, often within extensive grassland areas. ID: Similar to Dumeril’s Madagascar Swift, but usually shows a pale line along the middle of the back, rather than thin stripes along the sides of the back or no stripes at all. The tail is very fat at its base, then quickly tapers. BEHAVIOUR: Like other rock-dwelling Madagascar swifts.

210

TL < 39 cm | 15"

Length:

Androngovato Be TL < 35 cm | 14"

WHERE TO SEE: Parc Anja, where it is common alongside Dumeril’s Madagascar Swift.

Pale stripes along the sides of the back

Pale stripe along the centre of the back

PLATED LIZARDS I: SPECIES WITHOUT STRONG STRIPES ON BACK

Plated lizards comprise a family that is found across Africa and Madagascar. Of the 19 Malagasy species, the eight that are most frequently seen by visitors are covered. These large lizards are diurnal and conspicuous. Except for the rarely seen green species (see box below), all are ground-dwelling, and can often be located by their noisy movements in the leaf-litter. E

Tsingy Plated Lizard Zonosaurus tsingy

DISTRIBUTION: Dry forest with tsingy rock formations in a small area of the far north. ID: Base colour is variable, but often includes attractive shades of blue, orange and yellow. There are small pale speckles across the body. Always lacks strong stripes along the back. Zonosaurus subunicolor (not illustrated) is similar but occurs farther south, such as on Nosy Be and in Marojejy NP. BEHAVIOUR: Often sits on tsingy rock formations.

Androngovaton’ ny Tsingy

Length:

SVL < 8·5 cm | 3·5"

WHERE TO SEE: Fairly common in Ankarana NP. 1♂

Breeding males can be colourful

Red-legged Plated Lizard Zonosaurus rufipes

DISTRIBUTION: Rainforest in a small area of the north. ID: Mostly brown, with reddish legs, and lots of pale speckles. Unlike most other plated lizards, has stripes on the throat. Lines along the sides of the back are either weak or absent.

Brygoo’s Plated Lizard Zonosaurus brygooi

DISTRIBUTION: Lowland rainforest along the northern part of the east coast. ID: Base colour is brown, with pale and dark speckles. There are lines along the sides of the back, but they are much weaker than those on Madagascar Plated Lizard (see page 214).

Androngon’i Lokobe

NEAR THREATENED Length:

SVL < 9 cm | 3.5"

WHERE TO SEE: Common in Lokobe NP. Also possible in other forests on Nosy Be, and in Marojejy NP. M

Androngo Vato Fotsy Hatoka Length:

SVL < 8 cm | 3"

WHERE TO SEE: Not uncommon on the Masoala Peninsula. Also possible on Nosy Mangabe.

Herpetological Holy Grail Some of Madagascar’s most sought-after but hardest-to-see reptiles are the green plated lizards. They were first described in the late 19th century, then ‘disappeared’ for 100 years before being seen again by scientists. There are two species: Zonosaurus boettgeri, known from a handful of sites including Lokobe and Masoala NPs in the north; and Z. maramaintso, known from Tsingy de Bemaraha NP. Unlike other plated lizards, these green species live in trees, often high in the canopy. Count yourself extremely lucky if you manage to spot one! Plated Lizards versus Skinks Plated lizards resemble skinks (pages 216–219), and are often mistaken for them, even by knowledgeable guides. Plated lizards are generally larger; have bodies that are more squared-off rather than rounded; have slightly larger and squarer scales; and diagnostically, have a sharp furrow all along the bottom of the side of their body (a ‘lateral longitudinal fold’) whereas skinks have such a line only between their head and front legs. 212

Usually dark grey-brown with light spots

Reddish legs and flanks; distinctive stripes on throat

Weak lines along edges of back

PLATED LIZARDS II: SPECIES WITH STRONG STRIPES ON BACK

Madagascar Plated Lizard Zonosaurus madagascariensis

M Bekaratsaka Zonosaure de Madagascar

DISTRIBUTION: Found in many habitats including Length: Average SVL 13 cm | 5" heavily human-modified ones, throughout most of eastern WHERE TO SEE: Common and northern Madagascar, with an isolated population in throughout much of the east the central west. Mostly absent from rainforest interior. and north. ID: For separation from the similar Western Plated Lizard, see that species account. On Amber Mountain, Madagascar Plated Lizard is replaced by the similar Meier’s Plated Lizard Zonosaurus haraldmeieri (not illustrated), while the two species hybridize in Ankarana NP. The similar Bronze Plated Lizard Z. aeneus (not illustrated) overlaps in the central and southern parts of the east. It averages smaller, with a dot-and-dash pattern rather than continuous stripes along the sides of the back. E

Ornate Plated Lizard Zonosaurus ornatus

DISTRIBUTION: Forest edge and open montane habitat in central and central-eastern Madagascar. ID: A beautiful and distinctive species with five stripes along the back.

Length:

Marabe

SVL < 13 cm | 5"

WHERE TO SEE: Forest edges around Ranomafana NP and Anjozorobe. Also in the high mountains of Andringitra NP. M

Western Plated Lizard Zonosaurus laticaudatus

Androngoben’i Fiherenana

DISTRIBUTION: Found in dry forest, plantations, and Length: Average SVL 14 cm | 5.5" degraded habitats, mainly in the west, although also in the WHERE TO SEE: Common Ambanja region of the north, and around Fort Dauphin in around the headquarters the far south. of Ankarafantsika NP and in ID: Very similar to Madagascar Plated Lizard, but there Kirindy Forest. is little overlap in range. Where they do occur together, around Ambanja, Western Plated Lizard can be recognized by the fact that the pale stripes along the sides of its back are very broad just behind the head, then quickly taper. E

Karsten’s Plated Lizard Zonosaurus karsteni

DISTRIBUTION: Patchy distribution in western dry forest and secondary habitats. ID: Has fine black markings on the head, unlike the similar Western Plated Lizard, which averages larger and has more tapering pale stripes on the sides of the back. E

Androngon’i Fiherenana

Length: Average SVL 13 cm | 5" WHERE TO SEE: Fairly common in Kirindy Forest and around the Tsingy de Bemaraha.

Madagascar Keeled Plated Lizard

Sitry

Tracheloptychus madagascariensis

DISTRIBUTION: Dry forest in the southwest. ID: Pattern of three pale stripes along the back is distinctive. At Berenty, the somewhat similar Three-lined Plated Lizard Zonosaurus trilineatus (not illustrated) also occurs. It is more boldly marked, with stripes that extend onto the head. 214

Length:

SVL < 7 cm | 3"

WHERE TO SEE: Fairly common in forest throughout much of the southwest. One good place is the Arboretum d’Antsokay near Toliara.

Stripes along sides of back fade towards back legs

Can show orange or red below, especially on throat

Distinctive pattern of stripes and spots on the side

Stripes along sides of back start broad, then narrow but stay strong

Throat sometimes red

Large spots on sides. Narrow stripes along sides of back

5 Scales along back have 'keels' unlike normal plated lizards

Complex pattern

SKINKS I

Skinks are a family of lizards that occurs almost throughout the world. Although over 70 species are found in Madagascar, most live in leaf-litter or burrows, and are seldom seen. Many have reduced limbs and elongated bodies, and are snake-like. The Trachylepis skinks (which were formerly placed in the genus Mabuya) are familiar lizards that are diurnal and conspicuous, found on the ground, on rocks and on dead tree trunks. E

DISTRIBUTION: Found throughout most of the island. Seems to be more common than Elegant Skink in montane and rainforest-edge habitats. ID: Bold striped pattern. Can be very similar to Elegant Skink. R

Elegant Skink Trachylepis elegans

DISTRIBUTION: Most of the island in open habitats. Never inside true forest. ID: Definitive identification may be impossible, but classic features for Elegant Skink are an orange or red spot on the side of the neck a complex pattern of spots and stripes. Very similar to Gravenhorst’s Skink; the true relationship between these two species has yet to be resolved. E

Tandrefana Skink* Trachylepis tandrefana

DISTRIBUTION: Western dry forest. ID: Highly variable. Some individuals similar to Goldspotted, Elegant, or Gravenhorst’s Skinks, but rarely as boldly marked. Definitive identification may require a detailed analysis of the scales. 2

216

Gravenhorst’s Skink Trachylepis gravenhorstii

Length:

Androngo Matetika SVL < 9 cm | 3·5"

WHERE TO SEE: Very common. Look for it in degraded habitats adjacent to eastern rainforest.

Androngo Mena Hatoka

Length:

SVL < 6 cm | 2·5"

WHERE TO SEE: Common. Fond of hot, dry places such as Diego-Suarez, Morondava, and Toliara.

Katsatsaka Tandrefana

Length:

SVL 5–6 cm | 2–2·5"

WHERE TO SEE: Tsingy de Bemaraha NP and Kirindy Forest.

Simpler back pattern. Small or no white spots on sides

Complex pattern on back. White spots on sides. Orange neck marking.

Lacks bold stripes. Highly variable - some much darker than this individual

SKINKS II

Gold-spotted Skink Trachylepis aureopunctata

DISTRIBUTION: Forested and rocky areas of the west and southwest. ID: Distinctive pale spots on the front half of the body. BEHAVIOUR: More secretive than the other Trachylepis skinks included in this book. Often hides in the crevices of dead trees. R

Comoro Skink Trachylepis comorensis

DISTRIBUTION: Mainly a species of the Comoros; in Madagascar found only on Nosy Tanikely, where it is very common. ID: A large brown skink with pale speckles. Madagascar Plated Lizard (page 214) is also common on Nosy Tanikely, but it shows bold white stripes along the side of the back. Also see Plated Lizards versus Skinks on page 212 for structural differences between skinks and plated lizards.

Katsatsaka Pentina Fotsy

Length:

SVL < 8 cm | 3"

WHERE TO SEE: Kirindy Forest and the spiny forest around Toliara and Ifaty.

Katsatsaka Kaomoriana

Length:

SVL 10–11 cm | 4–4·5"

WHERE TO SEE: The small island of Nosy Tanikely, a popular snorkeling destination adjacent Nosy Be.

Secretive skinks several genera [~ 60 species]

DISTRIBUTION: Found in all the island’s habitats. The eastern rainforest holds the highest diversity. ID: Very variable in size, but snout to vent length (SVL) for most is 3–10 cm (1·2–4"). All have relatively smaller limbs and more elongated bodies than the Trachylepis skinks, giving them a snake-like appearance. Some lack limbs altogether, but differ from snakes in having eyelids, ear openings and being able to shed their tail as a defence mechanism. BEHAVIOUR: Most live in leaf-litter, although some of the more snake-like species burrow into the ground. 3

Voeltzkowia rubrocaudata is one of the snake-like skinks

218

Length:

SVL 2–23 cm | 0·8–9"

WHERE TO SEE: It takes luck or some digging in the leaf-litter to find any of these skinks. Common Madagascar Skink Madascincus melanopleura (not illustrated) is sometimes seen at Nosy Mangabe; Ranomafana NP is good for a couple of Amphiglossus species.

Conspicuous spotting

Gold-spotted Skink has distinctive pale spots on the front half of the body, usually forming seven or eight rows. Trachylepis vato (not illustrated) is very similar but has nine rows of white spots on the front half of the body; it is found mainly in the central part of the southern half of Madagascar but overlaps with Gold-spotted Skink in the far south.

Only on Nosy Tanikely

Amphiglossus frontoparietalis is typical of the secretive skinks that have long bodies and small limbs, but retain basic lizard structure.

WIDESPREAD DAY GECKOS

This delightful group of geckos is brightly coloured and diurnal. They are restricted to the Indian Ocean region, with the majority endemic to Madagascar. Most species inhabit trees and man-made structures: the walls of your hotel may be the best place to seek out the local species of day gecko. Other favoured haunts include banana trees and Traveller’s Palm (page 316). Males are larger and more colourful than females. E

Giant Day Gecko Phelsuma grandis

Fingoko Be*

DISTRIBUTION: Endemic to Madagascar, but introduced Length: TL 22–30 cm | 9–12” to the Mascarene Islands. Found from the northern tip of WHERE TO SEE: Quite common Madagascar, south to Marojejy and Ambanja. Uses many in the north, including in towns habitats, from rainforest to heavily human-modified areas. and villages. ID: The largest day gecko, with bright green coloration, and often red spots on the head and/or back. There is always a red stripe between the nostril and the eye, but this stripe ends at the eye rather than continuing onto the neck as in the closely related Madagascar Day Gecko Phelsuma madagascariensis (not illustrated). These two species also have little overlap in range: Giant Day Gecko is in the north, south to Masoala, while Madagascar Day Gecko is in the east, south of Masoala. Giant, Koch's and Madagascar Day Geckos are all similar and closely related, and were considered to be a single species until recently. For the most part, they have discrete ranges. Giant Day Gecko is common and frequently seen in the north. Koch’s Day Gecko is fairly common in much of the west. Madagascar Day Gecko is less frequently seen but occasionally spotted around Andasibe and St. Marie Island. E

Koch’s Day Gecko Phelsuma kochi

DISTRIBUTION: Found throughout most of the west, both in forest and human-modified habitat. ID: Similar to Giant Day Gecko, but duller-green, and averages smaller. Little or no overlap in range.

Peacock Day Gecko Phelsuma quadriocellata

DISTRIBUTION: Forest and human-modified habitats in the east and far northwest. ID: Green, with a blue-edged black spot behind the front legs. Lined Day Gecko can show a similar black spot, but it is never edged in blue. 220

Length:

TL 22–31 cm | 9–12"

WHERE TO SEE: Check the buildings at headquarters in Ankarafantsika NP. Also found in Tsingy de Bemaraha NP and Kirindy Forest.

Lined Day Gecko Phelsuma lineata

DISTRIBUTION: Very widespread in many different habitats, but absent from the southwest and most of the west. ID: Mostly green, with a black line along the side. The very similar Phelsuma dorsivittata (not illustrated) is found in the north, mainly on Amber Mountain. Small Lined Day Gecko (page 222) is similar, but smaller. E

Fingoko Tandrefana*

Length:

Atsatsatra Maitso TL 11–15 cm | 4–6"

WHERE TO SEE: Common in and around most eastern protected areas, including Andasibe and Ranomafana. Also on the High Plateau, and can be found in Tana. M

Length:

Atsatsam-bakoana TL 13 cm | 5"

WHERE TO SEE: Fairly common in and around most eastern protected areas, e.g. Andasibe and Ranomafana. The ‘Belle Vue’ viewpoint in Ranomafana NP is a reliable locality.

Bright green, usually with red spots

Paler coloration than Giant Day Gecko

Variable, but always shows dark lines along the sides

Black spot surrounded by blue, behind front legs

DAY GECKOS OF THE NORTH

Gold-dust Day Gecko Phelsuma laticauda

Atsatsa-maitso Tavaratra

DISTRIBUTION: Madagascar, and some other Indian Length: TL < 13 cm | 5" Ocean islands, to which it may have been introduced, as it WHERE TO SEE: Common in has been to Hawaii and Florida. In Madagascar, common much of the far north including in rainforest openings, plantations, and villages in the far Nosy Be and Sambava. northwest and northeast. ID: A beautiful gecko with blue rings around the eyes and a sprinkle of gold across the back. Three red bars on the head, and three on the back. Lacks the strong line along the side shown by Lined (page 220) and Small Lined Day Geckos. E

Speckled Day Gecko Phelsuma guttata

DISTRIBUTION: Rainforest and forest edge in the northeast. Usually in sun-lit places, often on Pandanus and Traveller’s Palm (page 316). ID: Three dark ‘V’-shaped marks on the throat are distinctive. Madagascar and Giant Day Geckos (page 220) can be similar, but are larger. R

Atsatsa-maitso Tatsinanana Length:

TL < 13 cm | 5"

WHERE TO SEE: Not hard to find on Nosy Mangabe. Also in Marojejy NP.

Aldabra Day Gecko Phelsuma abbotti

e DISTRIBUTION: The Seychelles, and northern and

Length:

Atsatsatra

TL 13–15 cm | 5–6"

western Madagascar. Found in villages, plantations, and WHERE TO SEE: Nosy Be, other human-modified habitats. Favours sun-lit tree trunks. around Ambanja, and in Tsingy ID: A relatively plain green or bluish species. Similar to de Bemaraha NP. Speckled Day Gecko, but has minimal overlap in range. If in doubt, can be identified by the dark line along the spine, a maximum of one or two ‘V’-shaped markings on the throat (three in Speckled), and the lack of the bold red dots on the back that are usually shown by Speckled Day Gecko.

Small Lined Day Gecko* Phelsuma pusilla

DISTRIBUTION: Hot east coast lowlands, often in and around towns and villages. Fond of banana and palm trees. ID: Similar pattern to Lined Day Gecko (page 220), but smaller. 1

222

Atsatsatra Maitsom-bakoana Length:

TL < 8·5 cm | 3·5"

WHERE TO SEE: Toamasina, St. Marie Island, and Maroantsetra.

Broad tail. Three red bars on head and back. Fine gold spots 4

Three dark Vs on throat and sides of neck. Usually red spots on back

Dark stripe through eye and along centre of back; often shows one or two â&#x20AC;&#x2DC;Vâ&#x20AC;&#x2122;-shaped marks on throat and sides of neck Like a miniature Lined Day Gecko

DAY GECKOS OF THE WEST, SOUTH AND CENTRE

Modest Day Gecko Phelsuma modesta

DISTRIBUTION: Dry forest, scrub and man-made structures, mainly in the southwest. ID: Pale greenish base colour, often with twin red lines on the back and faded-looking dark stripes along the sides. Looks washed-out and dull compared to many other day geckos. Thicktail Day Gecko can also be pale, but it never shows any red on the back or a darker stripe along the sides. E

Atsatsa-maitso Tandrefana

Length:

TL < 12 cm | 5"

WHERE TO SEE: Fairly common in southwestern towns and villages including Ifaty and Toliara.

Thicktail Day Gecko Phelsuma mutabilis

Atsatsatra Fotsikibo

DISTRIBUTION: Found in forest, scrub, and on manLength: TL < 10 cm | 4" made structures in the dry and hot west and southwest. WHERE TO SEE: Around ID: Capable of remarkable changes in colour, from brown Berenty, Toliara, and Zombitseto very pale grey. The tail can be pale blue 2a , green Vohibasia NP. 2b , brown or pale grey 2c . Although highly variable and changeable, all of its colour combinations are different from those of most other day geckos in its range. Very similar to Bora’s Day Gecko Phelsuma borai (not illustrated), which is endemic to Tsingy de Bemaraha NP, and Gould’s Day Gecko P. gouldi (not illustrated), which is known only from Parc Anja, and differs from both of these species in fine details of the scales. E

Standing’s Day Gecko Phelsuma standingi

DISTRIBUTION: Small area of spiny forest and dry forest in the southwest. Frequently seen on baobabs (pages 320–323). ID: Large, beautiful, and distinctive species with green, blue, and greyish base colour overlaid with complex dark markings. E

224

M Atsatsabe Gecko Diurne de Standing

VULNERABLE Length:

TL 21–28 cm | 8–11"

WHERE TO SEE: Often pointed out by guides in ZombitseVohibasia NP. Also in spiny forest around Ifaty.

Barbour’s Day Gecko Phelsuma barbouri

DISTRIBUTION: Found on the ground and on rocks in the central and southern mountains. This is the most terrestrial day gecko. ID: Variable coloration, with pale green or brownish base colour and darker brown stripes on the head and back. Other day geckos are absent from its habitat. 1

Length:

Atsatsam-bato TL < 15 cm | 6"

WHERE TO SEE: The mountains of Andringitra NP. Also present in less-disturbed rocky areas along the RN7 highway between Tana and Antsirabe.

Modest Day Gecko is a variable species: males are brightest; some females very dull

Often shows blue tail

Typically looks 'washed-out' 1 2b

Can be greenish 4

Some are very dull

Beautiful, complex, pattern Terrestrial, highland species

DWARF GECKOS

This large group of small geckos is found mainly in Africa and Madagascar. Dwarf geckos are diurnal and, although cryptic, can often be spotted on rocks, on tree trunks and elsewhere in the forest, and sometimes on man-made structures. They are capable of changing colour, although they generally maintain sombre shades of grey, brown and olive-green. As with chameleons, sleeping dwarf geckos can sometimes be found at night, on low vegetation within the forest. 1

Dwarf geckos Lygodactylus spp. [22+ species]

DISTRIBUTION: Found throughout Madagascar in most habitats, including spiny forest, rainforest, western dry forest, tsingy, and rocky high-elevation grassland. Some species readily use man-made structures. ID: Diurnal habits and small size usually identify them. Pattern variable, but never colourful, always with a base colour of grey or brown. BEHAVIOUR: Diurnal.

Antsiantsy

Length: SVL 1·5–4 cm | 0·5–1·5" WHERE TO SEE: Ankarafantsika, Zombitse, and Lokobe NPs and the town of Antsirabe are among the easier places to spot a dwarf gecko. Although they are common in many places, it takes sharp eyes to spot these small, cryptic geckos.

Definitive identification of dwarf geckos is difficult and requires detailed analysis of the scales. Furthermore, the classification of the Malagasy dwarf geckos is incomplete, and many species remain to be described. As such, this genus is covered without reference to individual species, except for tentative identifications in the photo captions.

226

Spiny Forest Dwarf Gecko Lygodactylus tuberosus Parc Mosa, Ifaty

Madagascar Dwarf Gecko Lygodactylus madagascariensis Lokobe NP

Robust Dwarf Gecko Lygodactylus pictus Lalatsara Forest

Grandidierâ&#x20AC;&#x2122;s Dwarf Gecko Lygodactylus tolampyae Zombitse-Vohibasia NP Variable, but can be boldly marked Rainforest Dwarf Gecko Lygodactylus miops Mitsinjo Reserve, Andasibe Cryptic: rainforest species; can show greenish tones

SMALL NOCTURNAL GECKOS 1

House geckos Hemidactylus spp. [3 species]

Atsatsaka an-Trano* F Gecko de Maison

DISTRIBUTION: Introduced widely by humans, and Length: TL 7–19 cm | 3–7·5" found throughout the warmer parts of the world. WHERE TO SEE: Can be found Probably at least one species occurred on Madagascar on almost any human structure. before people arrived. Found mainly in and around Especially abundant in hotter, man-made structures, although sometimes also in scrub lower-lying areas. and secondary forest away from human habitation. ID: Generally small to medium-sized. The upper range of the Grey’s House Gecko H. mercatorius 1c is the most length measurements applies only to Flathead House Gecko widespread and common H. platycephalus, which is restricted to the north. House Geckos species. are highly variable in colour, from extremely pale to dark grey Common House Gecko with strong black markings. Vertical pupils and many bumps H. frenatus 1b is found (tubercules) on the body and tail. mainly in the north, west, and southwest. VOICE: The voice of Common House Gecko H. frenatus is a Flathead House Gecko familiar sound of human civilization. Its most frequent call is a H. platycephalus 1a occurs in quick, fading series of 5–10 “tok” calls. the north. BEHAVIOUR: Generally nocturnal, although sometimes active during the day in shaded areas. Often hunt insects around lights at night. R

Fish-scale geckos Geckolepis spp. [~ 3 species]

Matahotrandro misy Kiran-Trondro*

DISTRIBUTION: Endemic to Madagascar and the Length: TL 11–14 cm | 4–5·5" Comoros. Found throughout most of Madagascar in WHERE TO SEE: Possible in forest, secondary habitat and plantations. Sometimes on and around almost any forest, man-made structures. but hard to find. Seemingly ID: Among Madagascar’s most bizarre creatures. more common in the north, as The fish-like scales are unique, but even more remarkable around Nosy Be and Ankarana is these geckos' ability to shed almost their entire skin as an and Ankarafantsika NPs. escape mechanism if grasped by a predator, regenerating it within a few weeks. Given current knowledge, distinction of the three Malagasy species – Peters' Spotted Fish-scale Gecko G. maculata 2a , Many-scaled Fish-scale Gecko G. polylepis (not illustrated) and Grandidier's Fish-scale Gecko G. typica (not illustrated) – is diﬃcult. BEHAVIOUR: Nocturnal, hiding during the day in nooks and crannies in dead trees, roots and Traveller’s Palm (page 316) trees. R

Madagascar Clawless Gecko Ebenavia inunguis

Matahotrandro Matrokila

DISTRIBUTION: Comoros, Mauritius, and Madagascar, Length: TL 8 cm | 3" where it is found in primary and secondary rainforest in WHERE TO SEE: Scarce but the east and north. Found only occasionally on man-made possible on night walks in structures. almost any rainforest. ID: Somewhat like a house gecko, but has a more pointed nose and a thinner and more elongated body. Also much less likely to be seen on man-made structures. BEHAVIOUR: Mainly nocturnal, though sometimes active during the day. 228

Flathead House Gecko is larger than the other Malagasy house geckos

Common House Gecko has a few rows of small bumps on back 1c

Grey's House Gecko has several rows of large bumps on back. Tail often black-striped 2a

Long nose and body 3

'Fish scales' make this gecko unmistakable

LARGE NOCTURNAL GECKOS I

There are at least 15 species of ground gecko in Madagascar, but many are rare and local. The three most commonly seen species are covered here. E

Madagascar Ground Gecko Paroedura picta

DISTRIBUTION: Lives mostly on the ground in spiny forest and dry western dry forest. ID: All ground geckos have huge eyes with a vertical pupil. Madagascar Ground Gecko has a beautiful pattern of oblique slashes along the sides. Bumps (tubercules) are scattered across the body. BEHAVIOUR: Nocturnal. E

Graceful Big-headed Gecko Paroedura gracilis

DISTRIBUTION: Rainforest in central and northern Madagascar. Found on the ground and in forest understorey. ID: Variable blotchy pattern. The only ground gecko in most of the eastern rainforest. BEHAVIOUR: Nocturnal.

Matahotrandro Mandady

Length:

TL < 24 cm | 9Âˇ5"

WHERE TO SEE: Widespread and fairly common in the west and south. Can be abundant at night in Kirindy Forest. Also possible on night walks around Ifaty and Berenty. M

Mocquardâ&#x20AC;&#x2122;s Ground Gecko Paroedura bastardi

DISTRIBUTION: On the ground and in the understorey of spiny forest and dry western dry forest. Sometimes found on man-made structures. ID: Variable, and probably a complex of species that will be split in the future. Never shows the oblique slashes of Madagascar Ground Gecko. Large bumps (tubercules) on the body are arranged into rows. BEHAVIOUR: Nocturnal. E

Length:

Matahotrandro Telo Tsipika TL < 23 cm | 9"

WHERE TO SEE: Uncommon in Kirindy Forest; sometimes in the camp bungalows. Also possible during night walks around Ifaty and Berenty.

Matahotrandro Maintibe

Length:

TL < 12 cm | 5"

WHERE TO SEE: Uncommon but possible at night around Andasibe, Masoala, and Amber Mountain.

Vertical pupil typical of nocturnal Malagasy reptiles

230

Slashes along the sides. Smaller bumps on body, not in rows. 2

Large bumps on body, arranged into rows

White tail tip

LARGE NOCTURNAL GECKOS II

Madagascar velvet geckos Blaesodactylus spp. [4 species]

Atsatsaka

DISTRIBUTION: Found mainly in dry forest in the Length: TL < 30 cm | 12" west and north, although also occurs in northeastern WHERE TO SEE: Easiest at rainforest. Occasionally on man-made structures. Ankarana NP, Kirindy Forest and ID: Usually large; can be impressively huge. Colour Ifaty, although widespread. See variable but often similar to house geckos (page 228). table below. Madagascar velvet geckos are larger, with a wider head and thicker body. BEHAVIOUR: Nocturnal. During the day, hides in deep cracks and crevices. WHICH MADAGASCAR VELVET GECKO AM I LOOKING AT? The most frequently visited sites in Madagascar that support a species of Madagascar velvet gecko: Ankarana NP Ankarafansika NP Kirindy Forest Marojejy NP Nosy Mangabe Tsingy de Bemaraha NP Toliara / Ifaty Zombitse-Vohibasia NP

232

1c 1a 1b

1b 1b 1b

Northern Madagascar Velvet Gecko* Ambonihazo Madagascar Velvet Gecko* Sakalava Madagascar Velvet Gecko Antongil Madagascar Velvet Gecko Antongil Madagascar Velvet Gecko Sakalava Madagascar Velvet Gecko Sakalava Madagascar Velvet Gecko Sakalava Madagascar Velvet Gecko

B. boivini B. ambonihazo B. sakalava B. antongilensis B. antongilensis B. sakalava B. sakalava B. sakalava

Northern Madagascar Velvet Gecko is the largest species

Ambonihazo Madagascar Velvet Gecko was formally described in 2011

Sakalava Madagascar Velvet Geckos, like other velvet geckos, usually hide in crevices during the day.

LEAF-TAILED GECKOS I: LEAF-LIKE AND DRY BARK-LIKE SPECIES

Leaf-tailed geckos are found in trees, and active at night. Local guides often know the locations of sleeping individuals. Otherwise, you will have to find one at night when they are moving around. Some species give loud, hissing distress calls when disturbed. There are at least 14 species, of which the seven most commonly seen are covered here. Some of the current species may be split into multiple species in the future. E

Satanic Leaf-tailed Gecko Uroplatus phantasticus

DISTRIBUTION: Eastern rainforest in the southern two thirds of Madagascar. ID: Colour variable, but always remarkably like a dead leaf. Spikes above the eyes makes it look rather sinister. Distinguished from the very similar Spearpoint Leaftailed Geckos by its relatively smaller head and much longer tail.

Taharaikitra Madinika

Length:

WHERE TO SEE: With luck, can be found at night around Ranomafana, sometimes along the main road through the national park. Also around Andasibe, though less frequently seen there.

Spearpoint leaf-tailed geckos

Uroplatus ebenaui / finiavana complex

DISTRIBUTION: This complex of leaf-tailed geckos is found mainly in the understorey of rainforest in the north. It seems to contain several undescribed species. Uroplatus finiavana 2a , which is endemic to Amber Mountain, was described in 2011. ID: Similar to Satanic Leaf-tailed Gecko, but with a smaller tail and larger head.

Lined Leaf–tailed Gecko Uroplatus lineatus

DISTRIBUTION: Northern part of the eastern rainforest. Often in stands of bamboo. ID: A distinctive pale yellow colour with faint dark stripes.

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Taharaikitra Kely

VULNERABLE / NEAR-THREATENED Length:

TL 6–8 cm | 2·5–3"

WHERE TO SEE: With some searching, U. finiavana 2a can usually be seen at night around Amber Mountain, while U. ebenaui 2b is occasionally found on Nosy Be.

Günther’s Leaf-tailed Gecko Uroplatus guentheri

DISTRIBUTION: Patchy distribution in dry forest of the west. ID: Variable coloration, but usually shows lines along the back, and resembles dry bark. In some places, occurs alongside Henkel’s Leaf-tailed Gecko (page 236), but is much smaller, lacks ‘mossy’ fringes, and has spikes around the eye.

TL 9 cm | 3·5"

Tahafisaka Mitsipika ENDANGERED

Length:

TL 10–13 cm | 4–5"

WHERE TO SEE: Possible, although rare, in Ankarafantsika and Tsingy de Bemaraha NPs and Kirindy Forest. Most likely during the wet season. M

Tahafisaka Mavomavo

Length:

TL 16–21 cm | 6–8"

WHERE TO SEE: A very lucky find on night walks around Marojejy and Masoala NPs.

The French name for leaf-tailed geckos is ‘uroplates’.

Leaf-like tail

Spearpoint-like tail

4 Can have bold lines or can be quite pale

E LEAF-TAILED GECKOS II: MOSSY BARK-LIKE SPECIES

On an island of marvels, leaf-tailed geckos rank among Madagascar’s most amazing creatures. They are some of the best-camouflaged creatures on Earth. Not only does their anatomy resemble their environment, but they also have the capacity to change their colour to perfect their disguise. When the mossy-looking species flatten themselves onto a tree trunk and change their colour to match the background, they become almost invisible (see page 8). E

Mossy Leaf-tailed Gecko Uroplatus sikorae

DISTRIBUTION: Mid-elevation rainforest in eastern and northern Madagascar. ID: Mid-sized gecko with fringes that make it look like mossy bark (see page 8). Colour highly variable. Can be similar to Common/Giant Leaf-tailed Geckos, but always smaller, and without those species’ distinctive eye pattern. The very similar Uroplatus sameiti (not illustrated) is found in low-elevation rainforest. Henkel’s Leaf-tailed Gecko can be very similar, although it is usually larger, and the two species overlap only in the north. BEHAVIOUR: Nocturnal; sleeps on tree trunks during the day. E

Henkel’s Leaf-tailed Gecko Uroplatus henkeli

DISTRIBUTION: Patchy distribution in Sambirano forest of the north and dry forest of the west. ID: Larger than Mossy Leaf-tailed Gecko. Similar to Giant Leaf-tailed Gecko, but smaller, with a different eye pattern: pale yellow with irregular red zigzags instead of curved red lines. BEHAVIOUR: Similar to Mossy Leaf-tailed Gecko.

Tandrekitra Maro Loko

Length:

WHERE TO SEE: Quite common in Amber Mountain NP, often on trees around the ticket office. Also possible in most eastern rainforest sites including Marojejy, Andasibe and Ranomafana.

Tahafisaka Razam-Boay VULNERABLE

Length:

Uroplatus fimbriatus / giganteus

236

TL < 26 cm | 10"

WHERE TO SEE: Nosy Be, especially Lokobe community forest (distinct from the national park). Less frequently sighted in Ankarafantsika and Tsingy de Bemaraha NPs.

Common / Giant Leaf-tailed Gecko

DISTRIBUTION: Rainforest in eastern and northern Madagascar. ID: Huge geckos – the second largest on Earth – with a distinctive eye pattern: pale base colour with dark curved lines on either side of the pupil. BEHAVIOUR: Sleeps on a tree trunk during the day, usually with its head pointing down. If threatened, it can make a surprising display in which it opens its mouth and shows its bright red tongue. At night, moves around on smaller branches.

TL < 19 cm | 7·5"

Length:

Tahafisaka

TL 24–32 cm | 9–13"

WHERE TO SEE: Common Leaf-tailed Gecko U. fimbriatus 3a is easy to see on Nosy Mangabe. Also possible at many other eastern rainforest sites, including Ranomafana NP. Giant Leaf-tailed Gecko U. giganteus 3b is uncommon in Amber Mountain NP. It also seems to be the large leaf-tailed gecko that occurs in Marojejy NP, but the taxononomy and separation of these species is unclear.

Eye details are useful for separating leaf-tailed geckos

1 brown or yellow

whitish with dark brown lines

2 pale with wavy red lines 1

3a beige with reddish-brown lines Mid-sized 'mossy' species

3b 3a

Intermediate between Mossy and Giant Leaf-tailed Geckos in many respects

MALAGASY BOAS

These large snakes are one of Madagascar’s biogeographical mysteries. Their exact affinities have long been debated, but recent studies suggest that their closest relative is the very different Calabar Boa Calabaria reinhardtii (formerly called ‘python’) of Africa. They form a family that is endemic to Madagascar (Sanziniidae), although this is often still considered a subfamily of Boiidae. E

Madagascar tree boas

Sanzinia madagascariensis / volontany

M Manditra Boa des Forêts de Madagascar

DISTRIBUTION: Throughout most of the island, though Length: TL < 2 m | 6' scarce in the central highlands. Found in forest, secondary WHERE TO SEE: One of the habitat, and plantations. Western populations belong to most widespread Malagasy S. volontany 1a , while eastern populations belong to snakes. Possible in almost any S. madagascariensis 1b . These species are both often protected area, and in many treated as subspecies of S. madagascariensis. heavily human-modified areas ID: Large snake with a distinctive structure. The triangular head as well. features deep grooves on the upper lip, which probably contain thermal receptors used in nocturnal hunting. Young boas are small, but are still identifiable by their structure. S. volontany is brownish, while S. madagascariensis is greenish. BEHAVIOUR: Generally found in trees during the day and on the ground at night. Hunt mainly mammals (rats, tenrecs and small lemurs), but also birds and amphibians. Give birth to 1–19 live young. E

Madagascar Ground Boa Acrantophis madagascariensis

M Do Boa de Madagascar

DISTRIBUTION: Inhabits various types of forest, mainly Length: TL < 3·2 m | 10.5’ in the northern third of Madagascar, although also in the WHERE TO SEE: With luck, can central-west. be found in Ankarafantsika, ID: A large snake with intricate dark patterning on a pale Ankarana, Masoala and Tsingy brown base. Lacks the upper lip grooves of Madagascar de Bemaraha NPs. Tree Boas. Larger than Duméril’s Ground Boa, and very little overlap in range. BEHAVIOUR: Nocturnal and ground-dwelling. Usually hides during the day. Hunts for mammals, from tenrecs up to the largest lemurs and birds. Gives birth to 2–6 large live young. E

Duméril’s Ground Boa Acrantophis dumerili

M Fagnano Boa de Duméril

DISTRIBUTION: Found in various kinds of habitat, from Length: TL < 3 m | 10’ spiny forest to open savannah, mainly in the southern half WHERE TO SEE: With luck or of Madagascar. Absent from the eastern rainforest. persistence, can be found in ID: Very similar in colour to Madagascar Ground Boa, and around Toliara and Ifaty, but its range barely overlaps that species, its average size Zombitse-Vohibasia NP, Parc is smaller, and the dark markings on its flanks are more Anja, and Berenty. distinctly ‘vertebra’-shaped. Lacks the distinctive upper lip grooves of Madagascar tree boas. BEHAVIOUR: Nocturnal and ground-dwelling. Gives birth to 6–13 live young, much smaller than those of Madagascar Ground Boa. 238

Diamondshaped marks with white centres along sides

Tree boas are told from ground boas by grooves on upper lip

1b 2 1a

Brownish base colour

Greenish base colour

Ground boa lacks grooves on upper lip

Pattern like Madagascar Ground Boa, but smaller; little overlap in range

DISTINCTIVE SNAKES

Madagascar’s snakes, other than the boas, are in the widespread Lamprophiidae family. All but one species (Mimophis mahfalensis) belong to a regionally endemic subfamily (Pseudoxyrhophiinae). Madagascar has no dangerously venomous snakes, but the hognose, leaf-nosed and cat-eyed snakes, and members of the genus Ithycyphus (not illustrated), do seem to possess mild venom, albeit with fangs that are at the back of the mouth, making it very difficult to inject venom into humans. E

Giant Hognose Snake Leioheterodon madagascariensis

Menarana

DISTRIBUTION: Common throughout in many different Length: TL < 1·5 m | 5’ habitats, including those heavily modified by humans. WHERE TO SEE: Most visitors Introduced on Grande Comore. will see this common snake ID: Colour and structure are distinctive: bold black-andsomewhere in Madagascar. yellow pattern, and an up-turned ‘hog nose’ which is used for digging. BEHAVIOUR: Diurnal and ground-dwelling. Often basks in the sun, for example along roads and paths. Diet includes a range of vertebrates and the eggs of Madagascar swift lizards (pages 208–211). E

Blonde Hognose Snake Leioheterodon modestus

DISTRIBUTION: Patchy distribution throughout much of the island, although absent from the far north and the eastern rainforest. Found mainly in dry habitats, both forest and human-modified ones. ID: Shares the ‘hog nose’ of Giant Hognose Snake, but is much more uniform beige in colour. BEHAVIOUR: Similar to Giant Hognose Snake. E

Length:

WHERE TO SEE: Best sought in western dry forest, especially Ankarafantsika NP.

Length:

TL < 1·3 m | 4’

Parastenophis / Phisalixella / Lycodryas spp. [12 species]

240

Filoala

WHERE TO SEE: Rarely seen, though present in almost any forest. Perhaps easiest to see in western dry forest as found at Ankarafantsika and Tsingy de Bemaraha NPs.

Madagascar tree snakes

DISTRIBUTION: Throughout Madagascar. Most species are restricted to forest, but some are also found in montane habitat, and in heavily human-modified areas. ID: Short snouts, and big eyes with vertical pupils. Leafnosed snakes also have vertical pupils, but have odd facial projections. Madagascar cat-eyed snakes (page 242) also have vertical pupils, but are found mainly on the ground and they have thicker bodies than most Madagascar tree snakes. BEHAVIOUR: Nocturnal and arboreal.

Mandipotsy TL < 1·2 m | 4’

Leaf-nosed snakes Langaha spp. [3 species]

DISTRIBUTION: Generally rare, but found in forest throughout most of Madagascar. ID: Another bizarre and unmistakable Malagasy animal: the males have a spear-like projection on their nose, while the females have a club-like projection. L. madagascariensis is the only leaf-nosed snake that is encountered frequently. BEHAVIOUR: Mainly diurnal and arboreal. E

Length:

Fandrefiala

Largest species TL < 1·5 m | 5’ Most species TL < 1 m | 3·3’

WHERE TO SEE: Generally uncommon, but night walks in the rainforest or western dry forest during the wet season give some chance.

Distinctive bold pattern

Aptly named, blonde!

3â&#x2122;&#x201A;

Male's nose is spear-like; female's is club-like

3â&#x2122;&#x20AC;

Langaha madagascariensis Parastenophis betsileanus

Lycodryas gaimardi

Lycodryas granuliceps

COMMON AND WIDESPREAD SNAKES

Along with the hognose snakes (page 240) and Madagascar tree boas (page 238), these are the most common and widespread snakes in Madagascar. E

Madagascar cat-eyed snakes Madagascarophis spp. [4 species]

Lapata

DISTRIBUTION: Two species are confined mainly Length: TL < 1 m | 3·3’ to the arid southwest, and one to the far north, while WHERE TO SEE: Quite M. colubrinus 1a occurs throughout the rest of the common, and seen by most country in many habitats, and is probably the most visitors who go on night walks. common Malagasy snake. Often around water. ID: Variable, but always show a pale base colour, with fine dashed dark markings superimposed throughout. The eyes have a vertical pupil (hence 'cat-eyed'). Madagascar tree snakes (page 240) also have vertical pupils, but are found mainly in trees. BEHAVIOUR: Active from dusk to dawn. Feed on small reptiles, amphibians and birds. When threatened, they bite readily, so are best treated with caution. E

Striped Madagascar Garter Snake*

Thamnosophis lateralis

DISTRIBUTION: Found throughout Madagascar in open and often wet habitats. Most common on the High Plateau and in the east. ID: Boldly marked with black and white or yellow. Goldcollared Snake (page 244) also has stripes, but shows a red belly, and usually a collar. BEHAVIOUR: Terrestrial and diurnal. Feeds on frogs. E

Length:

Maroandavaka TL < 1·1 m | 3·6’

WHERE TO SEE: Easiest to find in the dry west, as around Toliara and Morondava.

Length:

Bibimora

TL < 1 m | 3·3’

WHERE TO SEE: Common in the west and southwest, even in towns and villages.

Four-striped Snake Dromicodryas quadrilineatus

DISTRIBUTION: Found in forest and human-modified habitat in the northern half of Madagascar. ID: Pattern of four dark stripes is distinctive. BEHAVIOUR: Terrestrial and diurnal. 242

Mahafaly Sand Snake Mimophis mahfalensis

DISTRIBUTION: Absent only from the east. Uses almost all types of habitat except eastern rainforest. ID: Most individuals have intricate and distinctive buff and brown markings. Male usually distinctly striped, while female is plainer. BEHAVIOUR: Mainly terrestrial and diurnal. Prey items include other reptiles and amphibians. E

TL < 0·8 m | 2·5’

WHERE TO SEE: Common in many places; even possible to see it in Tana.

Bernier’s Striped Snake Dromicodryas bernieri

DISTRIBUTION: Found throughout in most habitats, although absent from the interior of eastern rainforest. ID: One stripe along the back, and two on the sides. Similar pattern to Striped Madagascar Garter Snake, but generally more brown-and-buff, rather than black-andyellow or black-and-white. BEHAVIOUR: Terrestrial and diurnal. E

Length:

Bibindrano

Length:

Marolongo TL < 1·2 m | 4’

WHERE TO SEE: One of the most common snakes in the north.

'Cat eyes' are unique among grounddwelling Malagasy snakes

Blackish stripes

Brownish stripes

Four dark stripes on back

The Mahafaly Sand Snake belongs to a different sub-family and genus from the other snakes of Madagascar; it probably reached Madagascar from Africa much more recently than the other species

Can be quite plain or boldly marked; complex markings on top of head

RAINFOREST SNAKES

These are some of the most common snakes that are found exclusively or primarily in rainforest. All are diurnal except the Gluttonous Bighead Snake. Note that Madagascar cat-eyed snakes and Four-striped Snake (page 242) also occur inside rainforest. E

Gold-collared Snake* Liophidium rhodogaster

DISTRIBUTION: Rainforest in the east and north. ID: Striped pattern above, with a reddish belly. Usually shows a pale collar. BEHAVIOUR: Diurnal and terrestrial. E

Collared Bright Snake Liophidium torquatum

DISTRIBUTION: Mainly in eastern and northern rainforest, but also locally in dry western forest. ID: Brown or grey, often with a darker head and a black collar. The upper lip is white, and the pale belly has black speckles. BEHAVIOUR: Diurnal and terrestrial. E

Tompontany Menakibo

Length:

TL < 60 cm | 2’

WHERE TO SEE: One of Ranomafana NP’s most common snakes. M

Tompontany Miovaloko

Length:

TL < 70 cm | 2·3’

WHERE TO SEE: Shy and inconspicuous, but possible in most forest sites, including Ranomafana, Amber Mountain, Lokobe, and Ankarafantsika NPs.

Forest Madagascar Garter Snake

Bibilava Volamena

Thamnosophis infrasignatus

DISTRIBUTION: Eastern rainforest and secondary habitats. Length: TL < 1 m | 3’ ID: A plain brownish snake with faint yellowish WHERE TO SEE: Andasibe and lines down the back and black marks behind the eye. Ranomafana areas. Coloration can be similar to Madagascar cat-eyed snakes (page 242) but those species have distinctive ‘cat eyes’ with vertical pupils. BEHAVIOUR: Diurnal and terrestrial. Known to eat frogs and small chameleons. E

Trail Madagascar Garter Snake*

Thamnosophis epistibes

DISTRIBUTION: Eastern rainforest and secondary habitat. Often near streams. ID: Similar pattern, but paler and not as boldly marked as Striped Madagascar Garter Snake (page 242), which has a perference for open areas. BEHAVIOUR: Diurnal and terrestrial. Hunts frogs. E

244

Length:

TL < 92 cm | 3’

WHERE TO SEE: Andasibe and Ranomafana areas.

Gluttonous Bighead Snake Compsophis laphystius

DISTRIBUTION: Mid-elevation eastern rainforest. One of the species most likely to be seen at night in the rainforest, usually along small streams. ID: Yellow-brown with faint dark lines. BEHAVIOUR: Nocturnal, and both arboreal and terrestrial. Feeds on frogs and their eggs.

Bibilava Mavo

Length:

Bibimaimbo Vangavanga TL < 62 cm | 2’

WHERE TO SEE: Night walks in Ranomafana, Andasibe, and Marojejy areas.

Stripes on back; pale collar; pink belly

Dark line through eye and along spine; collar lacking in some; black spots on belly

2 3

Black line behind eye; yellow stripes on back

Yellowish with dark brown stripes

Yellow base colour; fine dark stripes

Frogs l More than 300 species of frog have been described

from Madagascar, but DNA barcoding has revealed more than 500 species on the island, so at least 200 remain to be formally described! l Almost 10% of the world’s frogs are endemic to Madagascar, an extraordinary richness of amphibians for an island that makes up only 0·4% of the land area of the globe. l There are three orders of amphibian in the world: frogs, salamanders and caecilians. Only frogs are found in Madagascar. Six families are represented: Mantellidae (pages 254–281). Endemic to Madagascar and Mayotte. The largest Malagasy frog family. Diverse members include colourful mantellas, terrestrial Madagascar frogs, and arboreal bright-eyed frogs.

Microhylidae (pages 250–253). Worldwide tropical family. Madagascar has three endemic subfamilies that include some of its most interesting frogs – beautiful rain frogs, the tiny stump-toed frogs and bizarre Tomato Frog.

Golden Mantella

Marbled Rain Frog

Green Bright-eyed Frog

Stump-toed frog Stumffia pygmaea

Madagascar fringed frog

246

Tomato Frog

Hyperoliidae (page 248). A mainly African family represented in Madagascar by the reed frogs.

Starry Night Reed Frog

Ptychadenidae (page 248). Mainly African family represented in Madagascar by one species: the Mascarene Ridged Frog, the most common frog on the island.

Bufonidae. Madagascar’s only toad is the Asian Toad Duttaphrynus melanostictus (not illustrated), a recently introduced invasive species whose effects on Malagasy wildlife may be disastrous. Dicroglossidae. The Indian Bullfrog Hoplobatrachus tigerinus (not illustrated), probably another recent introduction.

l The origins of Malagasy frogs can be traced to seven separate colonization events. The first was some 60 million years ago, while the most recent was within the last couple of years (the Asian Toad). l Most frogs are vocal, and voice is one of the best ways to find and identify them. But their small size, frequently concealed calling locations, and the ventriloquial quality of their voices can make them bafflingly difficult to see. l Most frogs are not active during the day and are best searched for at night. However, there are some exceptions, including the mantellas (see pages 254–265) and some of the Madagascar frogs (see pages 274–281). l Most frogs call only during the wet season, and many disappear entirely during the dry season, possibly retreating to burrows and becoming torpid. This is especially true in the dry western parts of the island. In the eastern rainforest, some species are active and vocal year-round. But even there, a night walk on a dry night can be quiet, while there can be a cacophony of frog voices on a rainy night. The dry season timing of the main May to October tourist season means that many visitors leave the island without seeing many frogs: the frogs come into their own after the tourists have left. l Although only adult frogs are covered in this guide, frog reproduction is fascinating. All Malagasy frogs lay eggs, which in most species develop into aquatic tadpoles, then adult frogs. But within these basic parameters, they use an incredible diversity of reproductive strategies: at least 11 of the world’s 30 known modes of frog reproduction are used by Malagasy frogs. l Frog identification can be challenging. There is tremendous colour variation within many species, such that details of colour may be virtually useless for identification. For casual observers, the things to focus on are: – Size and shape (although beware of juveniles) – The quality of the skin (is it smooth, or bumpy and/or ridged?) – The call (if heard) – The general colour pattern l In most Malagasy frogs, females are larger than males. In the following species accounts, male and female dimensions are given separately for species with a large size difference. l This guide covers less than 20% of Madagascar’s described frogs, but the species covered here will account for the majority of sightings in frequently visited sites. Refer to Amphibians and Reptiles of Madagascar by Glaw and Vences for thorough coverage (see Further reading on page 327 for details). l There is no up-to-date standardization of common names for frogs, but this guide includes common names for the benefit of those who find scientific names off-putting. Where there is a logical existing common name, that one is used. In some cases, it was necessary to invent a new common name, or to replace or modify one that is a misnomer. These new common names are marked with asterisks. Be aware that herpetologists and most local guides use scientific names exclusively. l Frog diversity is very low in the dry southwest, and fairly low in most of the west. The eastern rainforest is remarkably rich in frogs. Within this biome, the rarely visited Tsaratanana massif of the north supports the highest diversity.

247

RIDGED FROG AND REED FROGS

Mascarene Ridged Frog Ptychadena mascareniensis

Sahona anTanimbary

DISTRIBUTION: Across Africa, Madagascar, and other Length: < 5·5 cm | 2·2" Indian Ocean islands. This is the most common and WHERE TO SEE: Day or night, widespread frog in Madagascar. It is found around almost almost anywhere! Often in rice any kind of stagnant water, although it is absent from paddies. rainforest interior. ID: Pointed nose. Raised ridges along the back. Colour extremely variable: the base colour is normally brown, with a paler stripe along the middle of the back. This stripe can be yellow or lime green. Distinguished from the Indian Bullfrog Hoplobatrachus tigerinus (not illustrated) by its smaller size and straight ridges as opposed to discontinuous bumps along the back. VOICE: Loud, harsh calls that are often repeated quickly. One of the call types resembles the bleat of an angry goat. BEHAVIOUR: Lives on the ground, in or near water. Active both day and night. E

Starry-night Reed Frog Heterixalus alboguttatus

DISTRIBUTION: Swamps and rice fields in the southern third of eastern Madagascar. Not found inside forest habitats. Often found in bananas trees. ID: Female sports an amazing ‘starry night’ pattern. Male is often plainer; some are plain buff. Members of this genus can always be recognized by their nearly triangular pupils. VOICE: Slow, rattling trill. BEHAVIOUR: Arboreal and nocturnal. Sleeps next to bodies of water during the day, sometimes in the open. E

Length:

♀ 3 cm | 1·2" ♂ s maller

WHERE TO SEE: At night around the village of Ranomafana. Good local guides know places to search for this species.

Betsileo Reed Frog Heterixalus betsileo

DISTRIBUTION: Swamps, rice fields, and secondary Length: < 3 cm | 1·2" forest in central Madagascar. Occurs both around forested WHERE TO SEE: At night in the areas on the east slope and on the High Plateau. Andasibe and Anjozorobe areas. ID: Distinctive triangular pupil shape identifies it as a Also found in highland towns Heterixalus species. Base colour varies from pale green such as Fianarantsoa. to yellow, often with paler stripes along the sides of the back and fine dark speckles. Can resemble Green Bright-eyed Frog (page 260) in colour, but that species has red-and-blue eyes and rounded pupils. VOICE: Single loud, rasping note, regularly repeated. BEHAVIOUR: Similar to Starry-night Reed Frog. 2♀

Heterixalus frogs have a distinctive pupil shape

248

2â&#x2122;&#x20AC;

Colour highly variable; often a pale stripe down centre of back; ridges on back

Females are unmistakable

2â&#x2122;&#x201A;

Usually speckled, and often has yellow lines on sides of back

Male much duller than female

DISTINCTIVE TERRESTRIAL MICROHYLIDS

DISTRIBUTION: Mainly around the Bay of Antongil, in the northeast. Seems to prefer human-modified habitat. ID: Large all-red frog. Redder than the closely related and possibly conspecific Sambava Tomato Frog Dyscophus guineti (not illustrated), which is typically orange. VOICE: An unusual series of hollow, low-pitched notes. BEHAVIOUR: Nocturnal and terrestrial. E

Tomato Frog Dyscophus antongilii

Sangongogno

NEAR THREATENED Length:

♂ 6–6·5 cm | 2·5" ♀ 8·5–10·5 cm | 3·4–4·1"

WHERE TO SEE: Wet habitat around the town of Maroantsetra.

Web-foot Frog Paradoxophyla palmata

DISTRIBUTION: Eastern rainforest and adjacent Length: 1·9–2·6 cm | 0·7–1" disturbed habitat. WHERE TO SEE: In the ID: Small frog with a bizarre shape: a small triangular Andasibe and Ranomafana head on a fat body. areas during and just after rain. VOICE: Very cricket-like. Mainly at night. BEHAVIOUR: Both terrestrial and aquatic. Breeds explosively when there is rain, and can suddenly become common, hopping around on roads and other open areas. Rather shy, and capable of long leaps to escape from danger. E

Marbled Rain Frog Scaphiophryne marmorata

DISTRIBUTION: Mid-elevation eastern rainforest and adjacent secondary habitat. ID: Beautiful frog with a fat body, marbled green-andbrown colour pattern, and very bumpy skin. Some other members of this genus are similar, but less frequently encountered. VOICE: Hollow trill. Not frequently heard. BEHAVIOUR: Nocturnal and terrestrial. Breeds explosively in the rains. E

VULNERABLE Length:

3·2–4·4 cm | 1·3–1·7"

WHERE TO SEE: At night around Andasibe after recent rain, when it can sometimes be found hopping around everywhere.

Stump-toed frogs Stumpﬃa [16 species]

DISTRIBUTION: Mainly in eastern and northern rainforest, although some species are found in the west and centre, and in human-modified habitats. ID: Small to tiny in size. Can often be tentatively identified by their size and reduced digits, but beware juveniles of other frog species. VOICE: Most species have a single high-pitched chirping call that is frequently repeated at short intervals. BEHAVIOUR: Terrestrial. Active during the day; most vocal at dusk.

Length:

1–2 cm | 0·4–0·8"

WHERE TO SEE: Easily overlooked due to their small size. S. pygmaea 4a is common in Lokobe NP. Stumpﬃa of undescribed species are fairly common on Masoala. A couple of species occur in the Andasibe and Ranomafana areas. Can be seen during the day.

All Stumpffia frogs are small, and S. pygmaea 4a and S. tridactyla are tiny. They are among the world’s smallest vertebrates, only about one centimetre (less than half an inch) long. 250

1â&#x2122;&#x20AC;

Females red; males yellowish 2

Pointed head

Wide body

Undescribed species from Andasibe

ARBOREAL MICROHYLIDS

Boulenger’s Giant Treefrog Platypelis grandis

DISTRIBUTION: Eastern and northern rainforest. Usually seen in or adjacent to tree holes. ID: One of the world’s largest arboreal frogs. As with most tree-dwelling frogs, has large tips to the toes. Grainy-looking back, with many large bumps. Adult is brown, with irregular paler markings. Juvenile starts bright green, then darkens to olive, then brown. VOICE: Atypical for a frog; sounds more like a bird or mammal. A very loud single note that sounds like chopping wood. Sometimes calls during the day. BEHAVIOUR: Arboreal and nocturnal.

Length:

♂ 5–9 cm | 2–3·5" ♀ 4–6 cm | 1·6–2·4"

WHERE TO SEE: Possible in most rainforest sites including Ranomafana, Andasibe, Nosy Mangabe and Amber Mountain. Can be seen asleep in tree holes during the day, or active nearby at night.

In 90% of frog species, females are larger than males. This enables the 'amplexus' mating embrace in which the male sits on the female’s back. Boulenger's Giant Treefrog is one of the exceptions: males are larger, which seems to give them advantage in territorial battles. E

Mahanoro Tree-hole Frog* Plethodontohyla notosticta

DISTRIBUTION: Eastern rainforest. Found in tree holes and the axils of tree ferns and Pandanus (page 316). ID: A rather wide, squat frog that is likely to be seen in holes in trees. Brownish colour, with a fine white line running along the junction of the side and the back. Around Andasibe, some individuals have a green back. Climbing Tree-hole Frog Plethodontohyla mihanika (not illustrated) has longer hindlimbs. VOICE: A simple hoot that is repeated over and over. BEHAVIOUR: Arboreal. Calls mainly at night. E

WHERE TO SEE: Possible in most eastern forest sites including Ranomafana, Andasibe and Masoala. During the day often found sitting in tree holes; at night found by call.

Length:

3–4 cm | 1·2–1·6"

WHERE TO SEE: Can be found during the day in AndasibeMantadia and Ranomafana NPs.

Barbour’s Climbing Frog* Platypelis barbouri

DISTRIBUTION: Eastern rainforest between Marojejy and Andasibe. ID: A small frog covered in big bumps. Variable but generally pale brown colour, often with darker markings interspersed. VOICE: Single high-pitched “peet” repeated about every three seconds. BEHAVIOUR: Nocturnal and arboreal. 252

3·5–4·2 cm | 1·4–1·7"

Ambatoharanana Climbing Frog* Platypelis tuberifera

DISTRIBUTION: Found in Pandanus (page 316) in the eastern rainforest. ID: Orange or buff, often with a thin pale line down the spine and a dark line behind the eye. The skin is smooth. Other Platypelis and Cophyla frogs can be similar, but are smaller and not usually found in Pandanus. VOICE: Plaintive, rising whistle. Repeated every several seconds. BEHAVIOUR: Nocturnal and arboreal. E

Length:

Length: ♂ 1·9–2·3 cm | 0·8–0·9" WHERE TO SEE: At night around Andasibe.

1ju

Juveniles start bright green, then transition to brown

Adults are brown 2

2ju

White lines along edges of back

Buffy Pandanusdwelling frog

Small frog with big bumps

MANTELLAS I: SPECIES WITH ORANGE

All the frogs covered on this and subsequent pages are in the Mantellidae family, endemic to Madagascar and Mayotte. The most famous of these are the mantellas: colourful, toxic frogs that resemble the unrelated poison dart frogs of the Neotropics, a striking example of convergent evolution. Even local guides who know little about frogs will seek out mantellas, so most visitors will see at least one species. All mantellas are active during the day, and most live on the ground. E

Baron’s Mantella Mantella baroni

Anakova

DISTRIBUTION: In rainforest and adjacent secondary Length: 2·2–3 cm | 0·8–1·2" habitat in the central and southern portions of eastern WHERE TO SEE: In the Madagascar. Prefers stream banks and other wet areas. Mantadia sector of AndasibeID: A Beautiful multi-coloured species. Madagascar Mantadia NP, and Ranomafana Mantella Mantella madagascariensis (not illustrated) is NP, mainly from November to similar, but much less common. They occur together in March. the Vohiparara region of Ranomafana NP, but Baron’s Mantella is more abundant even there. The two species are best identified by the pattern on the underside of their hindlegs: Madagascar Mantella shows red all the way up the legs, with orange patches interspersed, while Baron’s Mantella has red only up to the thighs, and usually shows no orange. Madagascar Mantella also shows a horseshoe-shaped blue mark on the throat whereas Baron’s Mantella shows either no mark or a blue spot. VOICE: Metallic single click. Often repeated. BEHAVIOUR: Active and vocal during the day; sometimes calls at night. E

Orange-backed Mantella Mantella ebenaui

DISTRIBUTION: Various types of forest and secondary habitat in the northern third of Madagascar. ID: Orange and black, with blue highlights. VOICE: High-pitched double click. Frequently repeated. BEHAVIOUR: Active and vocal day and night. E

WHERE TO SEE: Abundant on Nosy Be and Nosy Komba. Also possible on St. Marie Island, around Maroantsetra, and in Ankarafantsika NP.

Length:

1·8–2·6 cm | 0·7–1"

WHERE TO SEE: Tsingy de Bemaraha NP and Kirindy Forest.

Golden Mantella Mantella aurantiaca

DISTRIBUTION: Tiny range in central-eastern Madagascar. Prefers marsh edges and wet forest. ID: Unmistakable orange frog: one of Madagascar’s ‘flagship’ conservation species. VOICE: High-pitched clicks. Sometimes doubled. BEHAVIOUR: Active and vocal during the day, especially in the morning. 254

1·8–2·6 cm | 0·7–1"

Betsileo Mantella Mantella betsileo

DISTRIBUTION: In wet areas within western dry forest and secondary habitat. ID: Virtually identical to Orange-backed Mantella, but they do not overlap in range. VOICE: Double click, like Orange-backed Mantella. BEHAVIOUR: Active and vocal day and night. E

Length:

Sahona Mena

CRITICALLY ENDANGERED Length:

1·9–2·4 cm | 0·8–0·9"

WHERE TO SEE: Torotorofotsy Marsh, northwest of Andasibe village, from November to January.

Red thighs without orange markings interspersed separate this frog from the similar but much less common Madagascar Mantella

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Electric blue markings below

Orange back 3

Virtually identical to Orangebacked Mantella

The site for Golden Mantella at Torotorofotsy Marsh is managed by the NGO Mitsinjo. Entrance permits and a local guide can be arranged at their office just south of Andasibe.

MANTELLAS II: GREEN-AND-BLACK SPECIES

Climbing Mantella Mantella laevigata

DISTRIBUTION: Eastern rainforest in the north of NEAR THREATENED Madagascar. Unlike most mantellas, it climbs trees, and Length: 2·2–2·9 cm | 0·9–1·1" can be found anywhere from the forest floor up into the WHERE TO SEE: Nosy Mangabe mid-storey. is the classic site. Can also be ID: Shares black-and-green coloration with Guibé’s found in Marojejy NP. Mantella, but has a much more clearly defined pattern: mostly black, with a broad band of green or yellow-green on the back. VOICE: Double clicks. Frequently repeated. BEHAVIOUR: Active during the day. Breeds in water-filled tree holes. E

Blue-legged Mantella Mantella expectata

DISTRIBUTION: Found along streams and other wet areas within the Isalo sandstone massif of the southwest. ID: Yellow back, black sides and blue or blue-grey legs: unmistakable within its tiny range. VOICE: Rattling double click.

2–2·6 cm | 0·8–1"

WHERE TO SEE: Can be found in Isalo NP during the wet season, from November to early January. Best sought in the flooded, grassy areas at the entrance and outlet of canyons.

Length:

2·7–2·8 cm | 1·1"

WHERE TO SEE: Marojejy NP, especially around Camp Marojejia, the middle of the park’s three camps.

Green Mantella Mantella viridis

DISTRIBUTION: Tiny distribution in both dry and humid forest at the northern tip of Madagascar. Found in wet areas, for example along streams. ID: Pale green, with black along the sides. VOICE: Metallic double clicks. BEHAVIOUR: Active day and night.

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Length:

Guibé’s Mantella Mantella nigricans

DISTRIBUTION: Eastern rainforest in the north of Madagascar. Found along streams. ID: Limbs mostly fluorescent green, and the back colour varies between mostly black and mostly green. VOICE: Metallic single “tick” like a tiny hammer striking metal. BEHAVIOUR: Active during the day. E

ENDANGERED

ENDANGERED Length:

2·2–3 cm | 0·9–1·2"

WHERE TO SEE: Near Diego-Suarez at Montagne des Français, which can be accessed via the Jungle Park lodge. Common along the little stream that runs through that camp. Also possible in Amber Mountain NP.

Streams running through eastern rainforest are Madagascar's richest habitat for frogs

SMALL TO MEDIUM BRIGHT-EYED FROGS I

One of the largest genera within the Mantellidae family is Boophis, the ‘bright-eyed frogs’: classic tree frogs with enlarged tips to their toes, large eyes, and nocturnal habits. E

Central Bright-eyed Frog Boophis rappiodes

DISTRIBUTION: Eastern rainforest and forest edge. ID: Small and colourful. Green base colour; yellow stripes that start on the nose, then fade along the sides of the body; and reddish dots and blotches. Lacks dark brown markings of similar Bott’s Bright-eyed Frog. VOICE: Quiet double click. BEHAVIOUR: Nocturnal. Calling males usually sit in the forest understorey along rivers and streams. E

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♂ 2–2·5 cm | 0·8–1" ♀ 3–3·4 cm | 1·2–1·3"

WHERE TO SEE: One of the more common frogs on rainy nights around Andasibe and Ranomafana.

Bott’s Bright-eyed Frog* Boophis bottae

DISTRIBUTION: Eastern rainforest and forest edge. ID: Very similar to Central Bright-eyed Frog, but usually has some dark brown coloration on the back. VOICE: A short trill followed by a series of clicks that are similar to those given by Central Bright-eyed Frog. BEHAVIOUR: Similar to Central Bright-eyed Frog. Male calls from fairly low down within forest.

Length:

Length: ♂ 2·1–2·4 cm | 0·8–0·9" ♀ 3·5 cm | 1·4" WHERE TO SEE: Like Central Bright-eyed Frog, one of the more common frogs on rainy nights around Andasibe and Ranomafana.

Little or no dark brown coloration

Very similar to Central Bright-eyed Frog, but shows some brown: often a dark bar between the eyes

SMALL TO MEDIUM BRIGHT-EYED FROGS II

Fiery Bright-eyed Frog* Boophis pyrrhus

DISTRIBUTION: Around slow-moving water in eastern Length: ♂ 2·6–3·2 cm | 1–1·3" rainforest and forest edge. ♀ 3·7 cm | 1·5" ID: Unique coloration: red or orange base colour with red WHERE TO SEE: One of the dots and a darker 'hourglass' pattern on the back. most common frogs at night VOICE: Series of more than ten harsh, metallic notes that around Andasibe. become more intense towards the end. BEHAVIOUR: Nocturnal. Male calls from fairly low down within forest. E

Green Bright-eyed Frog Boophis viridis

DISTRIBUTION: Around slow-moving water in eastern Length: 3–3·5 cm | 1·2–1·4" rainforest and along forest edge. WHERE TO SEE: One of the ID: Pale green, often with tiny red spots. more common frogs on rainy VOICE: Harsh, high-pitched croaking notes repeated in nights around Andasibe. either long or short series. BEHAVIOUR: Nocturnal. Male calls from fairly low down within forest. E

Red-eyed Bright-eyed Frog Boophis luteus

DISTRIBUTION: Forest canopy in eastern rainforest and adjacent degraded habitats. Isolated populations also exist in riparian forest in the centre and south. ID: Bright green, with the outer part of the iris red. VOICE: A conspicuous part of the nocturnal soundscape. Loud and fast series of notes that has been likened to an American police car siren, or to a quick repetition of the French numbers “vingt-sept, vingt-huit”. BEHAVIOUR: Nocturnal. Males call from high perches along streams, making it diﬃcult to find.

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Length:

♂ 3·5–4 cm | 1·4–1·6" ♀ 5·1 cm | 2"

WHERE TO SEE: Night walks around Andasibe and Ranomafana. Easy to hear, but difficult to see. Most often seen when it is washed out of the canopy on nights of heavy rain.

Orange or red with dark red dots

Pale green with red spots

Bright green, with crimson in the eyes

MEDIUM TO LARGE BRIGHT-EYED FROGS I

Western Bright-eyed Frog* Boophis doulioti

DISTRIBUTION: Found around almost any non-flowing water in the west and southwest. Often on banana plants and inside man-made structures. ID: The only widespread western species of bright-eyed frog. The base colour is brown; some individuals have two yellow bands along the sides of the back. Almost identical to Duméril’s Bright-eyed Frog, but they do not overlap in range. VOICE: Often-repeated yelps, and occasional trills. BEHAVIOUR: Nocturnal. E

Length: ♂ 3·5–4·2 cm | 1·4–1·7" ♀ 4·1–5 cm | 1·6–2" WHERE TO SEE: Ankarafantsika NP, often in the camp bathrooms. Also possible in most western towns.

Duméril’s Bright-eyed Frog Boophis tephraeomystax

DISTRIBUTION: Uses the same habitats as Length: ♂ 3·5–4·2 cm | 1·4–1·7" Western Bright-eyed Frog, but in the east and north. ♀ 4·1–5 cm | 1·6–2" Frequently found in rice paddies, villages, and other WHERE TO SEE: Found in and human-modified habitats. around all the major eastern ID: Almost identical to Western Bright-eyed Frog, but and northern rainforest sites. with a different range. VOICE: Yelps and trills, similar to those of Western Bright-eyed Frog, but simpler. BEHAVIOUR: Nocturnal. E

Madagascar Bright-eyed Frog Boophis madagascariensis

DISTRIBUTION: Found along slow-flowing streams Length: ♂ 5–6·5 cm | 2–2·6" through forest and secondary habitat in eastern and ♀ 7–8 cm | 2·8–3·2" central Madagascar. WHERE TO SEE: One of the ID: Fairly large, plain frog with spikes on the elbows more common frogs on night and heels. Colour varies between pale brown and rich walks around Andasibe and reddish-brown. Some similar species occur in the north, Ranomafana. and Boophis roseipalmatus (not illustrated) is often seen on Amber Mountain. VOICE: Odd, complex combinations of clucks, rattles, and clicks. BEHAVIOUR: Nocturnal. Male usually calls from eye-level or below along streams.

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Can be reddish-brown, dull brown or pale brown and shades in between

Large spikes on heels

MEDIUM TO LARGE BRIGHT-EYED FROGS II

Goudot’s Bright-eyed Frog Boophis goudoti

DISTRIBUTION: Found along streams both on the High Plateau and in the central part of the eastern rainforest belt. Uses both forested and heavily degraded habitats. ID: Large. Brownish base coloration, with variable darker markings. Lacks the white lip of White-lipped Bright-eyed Frog, and the elbow spike of Madagascar Bright-eyed Frog (page 262). VOICE: Loud, harsh rattles and grunts. Similar to White-lipped Bright-eyed Frog, but slightly lower-pitched and harsher. BEHAVIOUR: Nocturnal. Male calls from or close to water. E

Length:

♂ 5–7 cm | 2–2·8" ♀ 7·5–10 cm | 3–3·9"

WHERE TO SEE: Most likely away from protected areas, at night around towns like Moramanga and Ambositra. Might show up on your plate if you order ‘cuisse de nymph’ in a restaurant!

White-lipped Bright-eyed Frog Boophis albilabris

DISTRIBUTION: Along streams in eastern and northern Length: ♂ < 7·3 cm | 2·9" rainforest and gallery forest. Generally found in the forest ♀ < 8·1 cm | 3·2" mid-storey. WHERE TO SEE: At night ID: Large. Usually green, although some individuals are around Ranomafana and brown, but always has a white upper lip. Green individuals Masoala. are much larger than other green species of bright-eyed frog. Brown individuals can resemble Madagascar Bright-eyed Frog (page 262) but do not have elbow or heel spikes. VOICE: Loud, harsh rattles and grunts. BEHAVIOUR: Nocturnal. Larger Boophis species inflate themselves when threatened, perhaps to make it more difficult for a snake to swallow them!

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1 Generally large and brownish but colour variable

Usually green (see opposite) but can be brownish

Lacks spikes on heels

VARIOUS ARBOREAL MANTELLIDS

Pandanus frogs Guibemantis subgenus

Pandanusicola spp. [9+ species]

DISTRIBUTION: Found in Pandanus (page 316) plants within the rainforest. Most species are restricted to the east, although some also occur in the northern and central parts of Madagascar. This complex is still poorly understood, and many species remain to be described or split. ID: Usually found in Pandanus. Ambatoharanana Climbing Frog (page 252) and other Platypelis species also live in Pandanus, but have flatter bodies and different coloration. Pandanus frogs are extremely variable: the most common colours are green or pale yellow, with darker brown markings. The skin is smooth, and looks almost gelatinous. VOICE: Series of notes with the quality of pebbles being clicked together. Some species give these notes in quick series, while others deliver them more slowly. BEHAVIOUR: Sometimes calls during the day, but most active and vocal at night. E

2·3–3·3 cm | 0·9–1·3"

WHERE TO SEE: Almost any rainforest site, including Amber Mountain, Marojejy, Masoala, Andasibe-Mantadia, and Ranomafana NPs.

Probable Guibemantis variabilis from Mantadia

Madagascar fringed frogs* Spinomantis [4 fringed species]

DISTRIBUTION: Along streams in eastern rainforest. Usually in the mid-storey, often quite high. ID: Masters of disguise, clad in spines, fringes, and cryptic coloration that resembles moss; similar to the camouflage of leaf-tailed geckos (pages 234–237). VOICE: Loud, metallic calls given in series, almost like a small hammer beating on metal. BEHAVIOUR: Nocturnal and arboreal. E

Length:

Sahonavakoana

Length:

3·1–5·1 cm | 1·2–2"

WHERE TO SEE: Night walks around Andasibe, Ranomafana, and Masoala. Sometimes local guides will point out well-camouflaged individuals sleeping on tree trunks during the day.

Pointy frogs* Blommersia spp. [10 species]

DISTRIBUTION: Most species are eastern, but a few are Length: 1·4–2·6 cm | 0·6–1" found in the north, west, and centre. Habitats range from 3b open wetlands to forest edge. ID: Small frogs with a rather long, pointed nose. Most species are brownish and/or greyish, but extremely variable. Often a dark ‘mask’ behind the eye. VOICE: Sharp, raspy clicks. Given in series by most species. BEHAVIOUR: Call mainly at night, but sometimes also during the day. Often on vegetation in the lower part of the forest, but can Blommersia domerguei also be found hopping on the ground. WHERE TO SEE: Blommersia blommersae 3a is the species that is most frequently seen, especially at night around Andsibe and Ranomafana. B. wittei (not illustrated) is fairly common in much of the north and west, usually in human-modified habitat, including villages. B. domerguei 3b is fairly common on the High Plateau and in higher parts of the eastern zone.

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Guibemantis liber from Amber Mountain NP

Guibemantis pulcher from near Ranomafana

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Spinomantis aglavei is one of the most frequently seen Madagascar fringed frogs 3a

Blommersia blommersae

VARIOUS TERRESTRIAL MANTELLIDS

Finding frogs In Madagascar, you ‘bump into’ frogs much less often than mammals, birds, and reptiles. Although some species are active during the day, most are nocturnal. Even the species that are day-active may be missed by observers who are more attuned to the movement of birds or mammals in the canopy than to jumping frogs at their feet. Frog sightings are also highly rain-dependent, especially in dry habitats, but even in the eastern rainforest. The peak season for frogs is the November to March wet season, after most visitors have left. Doing night walks with skilled local guides who know frog calls and microhabitats will greatly increase the number of species that you see. Night walks are currently not allowed within national parks but can be taken in areas close by (for example along the main road at Ranomafana NP) or in adjacent non-MNP reserves (such as the ones near Andasibe that are run by Mitsinjo and the local community association). E

Madagascar Bullfrog Laliostoma labrosum

DISTRIBUTION: Most of Madagascar, apart from the eastern rainforest and most of the High Plateau. ID: A medium-sized and very broad frog. Variable in colour, grey or brown, with some darker markings. VOICE: Fast trill of rough notes that lasts about two seconds. BEHAVIOUR: Terrestrial. Calls at night around water. Breeds explosively in temporary ponds after heavy rains. E

Length: ♂ 4·2–4·8 cm | 1·7–1·9" ♀ 6–8 cm | 2·4–3·2" WHERE TO SEE: Can be common in the north, west and southwest after heavy rain. Look for it around Ankarana, Ankarafantsika and ZombitseVohibasia NPs, Kirindy Forest, Toliara and Berenty.

Madagascar jumping frogs Aglyptodactylus spp. [3+ species]

DISTRIBUTION: Found on the floor of eastern, northern, and western forest. ID: Brown to buff base colour, some with darker markings interspersed. The distinctive black markings in front of and behind the eyes, and adjacent to the hindlimbs, are often enough to identify them as Madagascar jumping frogs. Males turn yellow during breeding season. VOICE: Croaked notes given in series. BEHAVIOUR: Terrestrial and largely diurnal. Often seen as they make impressive leaps across the forest floor. Explosive breeders.

Length:

3·5–6 cm | 1·4–2·4"

WHERE TO SEE: At least one species occurs in most protected areas. Aglyptodactylus madagascariensis 2a is the common species in the east and north, while A. securifer 2b is common in the north and west. The large, rare, A. laticeps (not illustrated) can be found in Kirindy Forest.

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Black marks on both sides of the eyes and adjacent to the hindlimbs

Can be plain or boldly marked

For a long time, there was thought to be only one species of Madagascar jumping frog. However, recent investigations have shown that this genus actually contains multiple species. In particular, Aglyptodactylus madagascariensis seems to represent a complex of several species.

Breeding male yellow below

BRIDGE FROGS I

In general, bridge frogs are earth-coloured, with bumps and/or ridges on their backs. They are mainly terrestrial, although some species call at night from low perches above the ground. Most species are active and vocal during the day, so these frogs are frequently encountered during daytime forest walks. Identifying bridge frogs can be difficult. Call is one of the most important ID clues. This section can only touch on this diverse and complicated group, which often confuses even amphibian experts. The new name ‘bridge frogs’ is used here, as the former name ‘Madagascar frogs’ refers now only to members of the genus Mantidactylus. E

Boulenger’s Bridge Frog Gephyromantis boulengeri

DISTRIBUTION: Eastern rainforest and secondary habitats. ID: A small, plain, brown frog, with bumps, but no ridges, on the back. Variable in colour; can have a pale stripe on the back. VOICE: A characteristic voice of eastern rainforest: a quick series of several “tsek” notes. E

2·5–3 cm | 1–1·2"

WHERE TO SEE: Common during the day on Nosy Mangabe and St. Marie Island, and around Andasibe and Masoala.

Webb’s Bridge Frog Gephyromantis webbi

DISTRIBUTION: Small area of eastern rainforest around the Bay of Antongil. ID: A beautiful but cryptic species with colours matching those of its mossy rock habitat. VOICE: Series of several harsh “tcheck” notes.

Length:

ENDANGERED Length:

♂ 2·5 cm | 1" ♀ 3·1 cm | 1·2"

WHERE TO SEE: Common during the day on rocks along small streams on Nosy Mangabe and Masoala.

Massif Bridge Frog Gephyromantis pseudoasper

DISTRIBUTION: Both Sambirano and eastern rainforest Length: 3·3 cm | 1·3" in the far north. WHERE TO SEE: Lokobe, ID: A small brownish frog with bumpy skin. Can be Ankarana, Amber Mountain, superficially similar to Boulenger’s Bridge Frog, but their and Marojejy NPs. ranges are largely discrete, and Massif Bridge Frog is paler below, often with some yellow. VOICE: Typical call is three quick series of three notes (nine notes in total). Boulenger's and Webb's Bridge Frogs can be found on the island of Nosy Mangabe, adjacent to Masoala.

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Brownish and bumpy; can be plain or boldly marked

Mossy-looking coloration

Pale underneath, often with yellowish coloration, unlike Boulenger's Madagascar Frog which is darker below

BRIDGE FROGS II

Rough Bridge Frog* Gephyromantis asper

DISTRIBUTION: Around streams in eastern rainforest. ID: Very rough skin: bumps throughout, ridges on the back, and spikes above the eyes and on the heels. Highly variable in colour, from buff to dark brown. VOICE: Very quick series of 2–4 “tik” notes, often repeated several times. E

2·7–3·1 cm | 1–1·2"

WHERE TO SEE: During the day in Ranomafana and AndasibeMantadia NPs.

Engraved Bridge Frog* Gephyromantis sculpturatus

DISTRIBUTION: Mid-elevation eastern rainforest. ID: Very similar to Beige Bridge Frog, which this species replaces at middle elevations. VOICE: Similar to Beige Bridge Frog, but notes are harsher. E

Length:

Length: ♂ 3·8–4·3 cm | 1·5–1·7" WHERE TO SEE: Ranomafana NP. Also present around Andasibe, although less common.

Beige Bridge Frog* Gephyromantis luteus

DISTRIBUTION: Low-elevation eastern rainforest and secondary habitat. ID: Usually pale brown, with some darker markings. The skin is fairly smooth, but there are ridges on the back. A pair of black-edged ridges runs from behind the eyes, nearly converging on the back. VOICE: Long series of up to 21 fairly musical notes.

Length: ♂ 3·6–4·3 cm | 1·4–1·7" ♀ 4·1–4·7 cm | 1·6–1·9" WHERE TO SEE: One of the most common frogs on Nosy Mangabe and Masoala.

The bridge frogs and Madagascar frogs are the ‘LBJs’ (‘little brown jobs’, a colloquial term for confusingly cryptic brownish birds) of the frog world! 3

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Very rough skin

Very similar to Beige Bridge Frog, but generally found at higher elevations 3

Black-edged ridges on back

MADAGASCAR FROGS: PLAIN SPECIES I

Mantidactylus is another of Madagascar’s largest frog genera. Like Gephyromantis, they are mainly ground-dwelling, and many species are vocal during the day. To a casual observer the two genera look very similar, although Mantidactylus tend to breed in streams and are usually more closely associated with water. They are ‘classic’ aquatic frogs, whereas Gephyromantis are more delicate and terrestrial. Many Mantidactylus are plain and earth-coloured, while others are slightly more colourful and distinctive. Some plainer species are covered here and on pages 276–277, while more distinctive ones follow. E

Ivohimanitra Madagascar Frog Mantidactylus majori

DISTRIBUTION: Along streams in rainforest in southeastern Madagascar. ID: A medium-sized frog, with fairly smooth skin. Sharply bicoloured: dark above and pale below. VOICE: Quick series of about ten creaky notes. E

Length:

4·1–4·7 cm | 1·6–1·9"

WHERE TO SEE: Commonly encountered in Ranomafana NP, even during the day.

Grandidier’s Madagascar Frog Mantidactylus grandidieri

Radaka

DISTRIBUTION: Deep and slow-flowing portions of Length: 7·5–10·8 cm | 3–4·3" eastern rainforest streams, north to the Masoala area. WHERE TO SEE: At night in ID: A large brown frog, usually with pale spotting, eastern rainforest, including identifiable by its size and habitat. Probably a complex Ranomafana, Andasibe, and that contains multiple species. Mantidactylus guttulatus Masoala areas. (not illustrated) is very similar, but has a more northern distribution, from the Marojejy area north. Goudot's Brighteyed Frog (page 264) also lives along streams, and can be superficially similar, but is smaller and slimmer, and has the enlarged tips to the toes that are typical of bright-eyed frogs. 2

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A large, stream-dwelling frog

Pointed nose; often sits on snags along stream edges

1 2

2ju

Identifying juvenile frogs can be very confusing, as they are smaller and often a different colour and pattern from the adults.

MADAGASCAR FROGS: PLAIN SPECIES II

Fort / Mocquard’s Madagascar Frog Mantidactylus femoralis / mocquardi

DISTRIBUTION: These two ‘species’ seem actually to form Length: ♂ 3·1–3·7 cm | 1·2–1·5" a poorly understood complex of species. Members of this ♀ 4·5–5·5 cm | 1·8–2·2" group are common along rainforest streams in eastern and WHERE TO SEE: Possible along northern Madagascar, and locally in wet refuges in the west. almost any rainforest stream, ID: Highly variable, but generally mid-sized, brown frogs, even during the day. ranging from buff to very dark brown, usually with darker markings interspersed; some individuals have a pale stripe along the spine. VOICE: Varies across the complex, but in general, give grumpy, unmusical notes in a series of variable length. E

Warty Madagascar Frog Mantidactylus ulcerosus

DISTRIBUTION: Found in the northwest in a variety of wet habitats, including highly human-modified ones. ID: Covered in large ‘warts’, which form ridges on the back. Plain brown in colour. Can be very similar to Betsileo Madagascar Frog, but seemingly little overlap in range. VOICE: A quick series of several creaky, trilled notes. E

Length:

3·2–4·1 cm | 1·2–1·6"

WHERE TO SEE: Ankarafantsika NP and the Nosy Be area. Can be seen during the day.

Betsileo Madagascar Frog Mantidactylus betsileanus

DISTRIBUTION: Along streams and swampy areas in and Length: ♂ 2·3–2·8 cm | 0·9–1·1" adjacent to eastern rainforest. Seemingly restricted to the ♀ 2·8–3·5 cm | 1·1–1·4" central and southern part of the rainforest belt, but very WHERE TO SEE: Andasibe and similar undescribed species occur farther north. Ranomafana areas, even during ID: Actually a complex of species whose definitive the day, although more vocal identification is diﬃcult. Small, with bumps and ridges at night. on the back. Generally pale brown with darker markings, with some having a paler stripe along the spine. Usually has a white spot on the tip of the nose. VOICE: Very fast, hollow trill that lasts about three seconds.

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Variable; often but not always has a pale stripe on the back

Warty ridges on back

Highly variable; usually has a white tip to the nose, with some having a pale stripe all along the spine

MADAGASCAR FROGS: MORE DISTINCTIVE SPECIES I

Charlotte’s Madagascar Frog* Mantidactylus charlotteae

DISTRIBUTION: Near streams in low-elevation rainforest Length: 2·2–3·2 cm | 0·9–1·3" in the northeast. WHERE TO SEE: Common in the ID: This species and the other two shown here share northeast, on the islands of St. the same basic colour pattern: dark brown (almost Marie and Nosy Mangabe, and in black) on the sides, pale brown on the back, and a white Marojejy and Masoala NPs. stripe under the eye. Compared with White-whiskered Madagascar Frog, Charlotte's Madagascar Frog is usually smaller, and the white stripe below the eye fades before reaching the nostril. VOICE: Quick series of creaky notes, more complex than those of White-whiskered Madagascar Frog, and delivered more slowly. BEHAVIOUR: Active and vocal during the day. E

White-whiskered Madagascar Frog* Mantidactylus melanopleura

DISTRIBUTION: In the vicinity of streams within eastern rainforest. ID: This is the largest and most boldly marked of the three species shown here, and the only one with a white stripe that reaches its nostril. VOICE: Long series of “tak” notes, delivered quickly. BEHAVIOUR: Active and vocal during the day. E

Length:

3–4·1 cm | 1·2–1·6"

WHERE TO SEE: During the day at most eastern rainforest sites including Ranomafana, Andasibe, Masoala and Marojejy.

Central Madagascar Frog Mantidactylus opiparis

DISTRIBUTION: In the vicinity of streams within eastern Length: 2·4–3·3 cm | 0·9–1·3" rainforest. WHERE TO SEE: During the ID: Similar to Charlotte’s and White-whiskered Madagascar day at most eastern rainforest Frogs, but less boldly patterned; in particular the white stripe sites including Ranomafana, below the eye is usually blurred. Often shows a diamondAndasibe, and Marojejy. Usually shaped marking within the pale brown patch on the back less common than White(like White-whiskered Madagascar Frog, but unlike Charlotte’s whiskered Madagascar Frog. Madagascar Frog). VOICE: Very similar to White-whiskered Madagascar Frog, but the series is delivered much more slowly. BEHAVIOUR: Active and vocal during the day. 2

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1ju

Juvenile, probably Charlotte's Madagascar Frog White stripe fades before nostril

White stripe reaches nostril

Indistinct white line below eye

MADAGASCAR FROGS: MORE DISTINCTIVE SPECIES II

Folohy Madagascar Frog Mantidactylus argenteus

DISTRIBUTION: Along streams in eastern rainforest. ID: Smooth skin is green with brown markings. Silvery below. Looks more like a pandanus frog (page 266) than the other Madagascar frogs. VOICE: Series of 6–15 quiet “tick” notes, slightly fading at the end. BEHAVIOUR: The only Madagascar frog that is largely arboreal. Calls from trees along rainforest streams during the day. E

280

♂ 2·7 cm | 1·1" ♀ 3·1 cm | 1·2"

WHERE TO SEE: During the day in the Mantadia sector of Andasibe-Mantadia NP.

Duméril’s Madagascar Frog Mantidactylus lugubris complex

DISTRIBUTION: A complex of species that occurs across the eastern rainforest and locally elsewhere. Found along streams, usually sitting on rocks very close to the water. ID: Skin smoother than most Madagascar frogs, although there are some bumps. Colour highly variable, but beautiful, and matches their rocky streamside habitat. Typically shows black, brown and olive bands across the back. VOICE: A single short, rasping trill. BEHAVIOUR: Vocal at night. 2

Length:

3–4·5 cm | 1·2–1·8"

WHERE TO SEE: Andasibe and Ranomafana areas. Possible during the day but easier at night.

Mostly arboreal

Smooth skin, bands across back

INVERTEBRATES: DISTINCTIVE SPECIES I

The vast majority of living things are invertebrates, with arthropods alone representing over 75% of Earth’s biodiversity. Madagascar supports well over 100,000 species of invertebrate, most of which are endemic. This represents an extraordinarily high proportion of the world’s 1·3 million or so described animal species, and serves to demonstrate again the importance of Madagascar as a repository of biodiversity. A handful of the more conspicuous and exceptional invertebrates is covered below. I

Giant African Land Snail Achatina fulica

DISTRIBUTION: Found throughout the island except in very dry environments. Occurs in most habitats, from gardens to primary forest. Unfortunately, this introduced species is the most common and conspicuous of the island’s almost 700 terrestrial snails, most of which are endemic. ID: A large snail that is usually brown, although highly variable in colour. E

M Sifotra Escargot Géant Africain

Length: < 20 cm | 8" Weight: < 1·5 kg | 3·3 lb (Average size much smaller) WHERE TO SEE: Almost anywhere.

Giant pill millipedes

Zoosphaerium / Sphaeromimus / Microsphaerotherium spp. DISTRIBUTION: There are about 40 species of giant pill millipede found all around Madagascar, mainly in forest. They belong to an ancient family that is also found in southern India and Sri Lanka, and which was split apart by the breakup of Gondwanaland. ID: They look like giant pillbugs or woodlice, but are in a completely different subphylum of organisms. Variable in colour; most commonly dark brown, buff or dark green.

Length:

Tainkintana < 10 cm | 4"

WHERE TO SEE: Possible in most forest sites, but most often seen in eastern and northern rainforest.

‘Normal’ millipedes are also abundant and conspicuous in Malagasy forests. Some are quite beautiful, sporting yellow bands or bright red colours

282

Rolls up in defence, like a pillbug or woodlouse

Often shows brown coloration and banding

INVERTEBRATES: DISTINCTIVE SPECIES II 1

Golden orb-web spiders Nephila spp.

DISTRIBUTION: Spiders of this genus are found throughout the world in warm climates. Occur across Madagascar, although most common in the east. Often in close association with humans, sometimes building webs on electric lines and poles. ID: Black body with yellow markings and red-and-black-striped legs. Females dwarf males. BEHAVIOUR: Constructs huge webs with golden strands. E

Length (legspan):

Hala

♂ 1–2 cm | 0·4–0·8" ♀< 20 cm | 8"

WHERE TO SEE: Very common and widespread, often in towns.

Giraffe-necked Weevil Trachelophorus giraﬀa

M Gisagisa Scarabée Girafe

DISTRIBUTION: Found mainly along forest edge and Length: ♂ ~ 2·5 cm | 1" in secondary habitat. The Malagasy species is the most WHERE TO SEE: Fairly famous, but other very long-necked weevils occur common in the Andasibe elsewhere in the world. and Ranomafana areas. Often ID: Incredible and unmistakable long-necked weevil with pointed out by local guides. a red back. Males have much longer necks than females. BEHAVIOUR: Males fight with their extremely long necks, while females' more modest necks assist in making their favoured food plants into leaf rolls, in which they lay eggs. E

Flatid Leaf Insect Phromnia rosea

DISTRIBUTION: Found in forest throughout Madagascar. ID: Usually found in large aggregations. Adults are red and flat, while the nymphs look like white flowers due to a waxy substance that they extrude to deter predators.

Length:

~ 2 cm | 0·8"

WHERE TO SEE: Common in most protected areas.

3ju

284

Both adults (opposite) and nymphs form groups and look almost like flowers at a distance!

1♂

1♀

2♂

BUTTERFLIES: SWALLOWTAILS I

There are about 300 butterflies described from Madagascar, about three quarters of which are endemic. This section includes the most widespread and obvious species and groups. It will allow you to identify most of the butterflies that you see on a typical nature tour, at least to genus. There is no standardization of common names for butterflies; common names novel to this guide are marked with an asterisk. Wingspan (WS) is the measurement given for butterflies. 1

Citrus Swallowtail Papilio demodocus

DISTRIBUTION: Sub-Saharan Africa. One of Madagascar’s most common butterflies. Found in all open habitats except high mountains. ID: Large and distinctive black-and-cream butterfly with elaborate markings. Lacks 'swallow tails'. BEHAVIOUR: Some of its food plants are cultivated citrus trees. e

WS 10–13 cm | 4–5"

WHERE TO SEE: Can be found in almost any garden or village. Common even in Tana.

Length:

WS 8–11 cm | 3–4·5"

WHERE TO SEE: Generally uncommon. Masoala is a good site.

Cream-lined Swallowtail Papilio delalandei

DISTRIBUTION: Found in rainforest and dry forest. ID: Distinctive swallowtail with dark base coloration and cream bands through the wings, which are visible both above and below. The largely deforested High Plateau does not hold many butterflies, but does play host to the common and familiar Citrus Swallowtail.

286

Length:

M Lolomboasary Papillon de Vinson

Mocker Swallowtail Papilio dardanus meriones

DISTRIBUTION: Sub-Saharan Africa. Found in forest and edge habitat throughout Madagascar. ID: Black-and-cream buterfly with less complex pattern than Citrus Swallowtail, and pale-tipped 'swallow tails'. In mainland Africa, the female mimics unpalatable species such as African Monarch (page 298), but on Madagascar, there is no mimicry and both sexes are similar. BEHAVIOUR: Typical swallowtail: flighty and rarely lands for long. E

Length:

WS ~ 8–11 cm | 3–4·5"

WHERE TO SEE: Most common in eastern rainforest, such as around Ranomafana and Andasibe.

Lacks tails 2

Cream base colour

BUTTERFLIES: SWALLOWTAILS II

Spotted Blue Swallowtail* Papilio epiphorbas

DISTRIBUTION: Madagascar and the Comoros. Widespread and common, both in forest and humanmodified habitat, although absent from the dry southwest. ID: A dark swallowtail with blue markings on the top of the wings. Similar to Banded Blue Swallowtail, but has discrete blue spots on each wing instead of blue lines. through both wings. E

288

Length:

WS ~ 8–11 cm | 3–4·5"

WHERE TO SEE: Common in most rainforest sites, including Andasibe.

Madagascar Swordtail* Graphium evombar

DISTRIBUTION: Comoros and Madagascar, in rainforest and western dry forest. ID: Intricate patterning and very long tails. The pale parts of the wings range from pale yellow to pale blue. BEHAVIOUR: Very fast flier. Sometimes lands on mud.

WS ~ 8–11 cm | 3–4·5"

WHERE TO SEE: Especially common in the west and on the High Plateau. Can be seen in gardens in Tana.

Banded Blue Swallowtail Papilio oribazus

DISTRIBUTION: Found in and around rainforest. ID: See Spotted Blue Swallowtail. Most common in rainforest, whereas Spotted Blue Swallowtail prefers drier and more open habitats. R

Length:

WS ~ 7–8 cm | 3"

WHERE TO SEE: Perhaps easiest to see in western forest, as in Ankarafantsika NP and Kirindy Forest.

Plain below

Occasionally lacks tails

2 3

Very long tails

BUTTERFLIES: SWALLOWTAILS III

Green Lady* Graphium cyrnus

DISTRIBUTION: Found throughout Madagascar in all types of forest. ID: Handsome tailless swallowtail. Black and pale green above; dark brown and pale green below, with variable red highlights.

290

WS ~ 6–7 cm | 2·5–3"

WHERE TO SEE: Possible in most forest sites. Frequent in Andasibe-Mantadia NP.

Madagascar Giant Swallowtail Pharmacophagus antenor

DISTRIBUTION: Found in open habitats, including towns and gardens. Absent from the east, and most common in the west and southwest. ID: This huge swallowtail can be identified simply by its size. The base colour is black, with pale yellow spots, and some red highlights. The forewings are extremely long. BEHAVIOUR: Wanders widely, and often flies high.

Length:

Length: WS 12–14 cm | 4·5–5·5" WHERE TO SEE: Fairly common around Toliara, Ifaty, and Morondava.

Most often seen in flight

Lacks tails

Huge, dark swallowtail

BUTTERFLIES: PIERIDS I

Madagascar Orange Tip* Colotis evanthe

DISTRIBUTION: Madagascar and the Comoros. Found both in forest and open, human-modified areas. ID: Mostly white. Male has bright orange markings on the tip of the upperside of the forewing; the forewing tips of the female are duller. After landing, closes wings, when only a speckled white pattern is visible below. BEHAVIOUR: Flies weakly and usually stays close to the ground. 2

Length:

WS ~ 3–4 cm | 1–1·5"

WHERE TO SEE: Especially common in the west and north.

Grass yellows Eurema spp. [4 species]

DISTRIBUTION: Members of this genus are found Length: WS 3–4 cm | 1–1·5" throughout much of the world in open habitats, including WHERE TO SEE: Almost those that are human-modified. anywhere. ID: Small, yellow butterflies, with black edges to the upperside of the wings. Separation of the four Malagasy species can be tricky. Broad-bordered Grass Yellow Eurema brigitta 2b is the most common and widespread species. It has broad dark borders to the forewing (and often the hindwing) above, and is dull yellow with broad greyish or pinkish borders and simple speckling below. Malagasy Grass Yellow E. floricola 2a has complex spots below, including a roughly ‘T’-shaped mark near the centre of the hindwing. Angled Grass Yellow E. desjardinsii 2c can be very similar to Malagasy Grass Yellow, but shows an angled rather than evenly rounded hindwing. Marsh Grass Yellow E. hapale (not illustrated) can also be very similar to the preceding two species, but is generally paler, and shows linear (not ‘T’-shaped) outlined marks near the centre of the hindwing and the front of the forewing below. BEHAVIOUR: Fairly weak flight, usually close to the ground. 3

Migrants Catopsilia spp. [2 species]

Length: WS 5–7 cm | 2–2·5" DISTRIBUTION: C. thauruma 3a is a regional endemic, while C. florella 3b has a wide Old World distribution. WHERE TO SEE: Most towns Found in all open habitats. and gardens. ID: Male is greenish-white, while the female is yellowish. Whiter individuals can be mistaken for whites (page 294), but black markings not obvious. The two Malagasy species are hard to distinguish. C. florella is slightly smaller than C. thaurumaand has more pointed forewings. C. thauruma is more common; often observed around villages. BEHAVIOUR: Strong fliers. 3a♀

292

Probable Catopsilia ﬂorella

1♂

1♀

Grass yellows are yellow with black wing edges above

3a♂

Males greenish-white

3b♀

Females yellow

BUTTERFLIES: PIERIDS II

African Wood White Leptosia alcesta sylvicola

DISTRIBUTION: Afrotropical forest. ID: Mostly plain white, with faint speckles below, and one black spot on the forewing. BEHAVIOUR: Very weak flight gives rise to its colloquial name 'Flip-flop'. E

WS 3–4 cm | 1–1·5"

WHERE TO SEE: Present in most forest sites, but inconspicuous.

Grandidier’s Caper White* Belenois grandidieri

DISTRIBUTION: Dry forest and adjacent secondary habitat. Absent from the east. ID: Delicate black tracery and scattered hints of yellow below. E

Length:

WS ~ 4 cm | 1·5"

WHERE TO SEE: Fairly common in western forest, as in Kirindy Forest and around Toliara.

Madagascar Dotted Border* Mylothris phileris

DISTRIBUTION: Forest throughout Madagascar. Length: WS ~ 5 cm | 2" ID: White, with black spots along the edges of the wings, WHERE TO SEE: Fairly common and a variable orange blaze at the base of the hindwing in most forest sites. In rainforest, occurs alongside Albatross White Appias sabina (not illustrated). That species shows smaller spots along the hindwing, and more wedge-shaped marks along the edge of the forewing. BEHAVIOUR: More languid flight, and more often in the forest canopy than other pierids. E

Yellow-painted Caper White* Belenois helcida

DISTRIBUTION: Rainforest, mainly in the east and north. ID: Broad black edges to both wings. Orange blaze at the base of the forewing below.

294

Length:

WS ~ 4–5 cm | 1·5–2"

WHERE TO SEE: Rainforest sites such as Andasibe.

Black spot and black edge on forewing above 2

White with speckles below 2

Lacy pattern below 3

Dots along wing edges

Pattern similar on top and bottom of wings

BUTTERFLIES: METALMARKS and LYCANIDS

Madagascar metalmarks Saribia spp. [4 species]

DISTRIBUTION: Rainforest, mainly in the east and north. ID: Intricate, distinctive pattern of zig-zags, bars and eye spots below. The plainer topside is rarely seen, as the wings are usually held closed when perched. There are multiple tails at the back of the hindwing. BEHAVIOUR: Flies low, inside of the forest. 2

Length:

WS ~ 4 cm | 1·5"

WHERE TO SEE: Amber Mountain, Andasibe-Mantadia, and Ranomafana NPs.

Blues / hairstreaks 16 genera [~ 46 species]

DISTRIBUTION: Various species found throughout Length: WS mostly 2–3 cm | 1" Madagascar in most habitats. WHERE TO SEE: Blues are ID: Large group of small butterflies. Most are blue or very common, even in Tana; purple above, and pale grey below. Many have ‘false eye’ hairstreaks are harder to find. spots on their hindwing, while some have thin tails, or ‘false antennae’. After landing, the hindwings are waved in circles, enhancing the impression of a false head to misdirect the attacks of predators. BEHAVIOUR: Most species have low, weak, fluttery flight. 2

Common Smoky Blue Euchrysops malathana 2

296

Hairstreak Hemiolaus cobaltina

Clover Blue Zizina antanossa 2

Brown Playboy Deudorix antalus

Definitively identifying Madagascar metalmarks usually requires dissection and/or DNA sequencing!

Probable Saribia perroti from Ranomafana NP

Hairstreak, probably Deudorix batikeli (female)

Sooty Blue Zizeeria knysna

Possible Saribia tepahi from Amber Mountain NP Common Zebra Blue Leptotes pirithous

Gaika Blue Zizula hylax

BUTTERFLIES: ORANGE NYMPHALIDS I

The Nymphalidae or ‘brush-footed butterfly’ family contains some of the most colourful and conspicuous Malagasy butterflies. 1

African Monarch Danaus chrysippus

DISTRIBUTION: Widespread Old World species. Found throughout Madagascar except in the high mountains. ID: Large and distinctive orange, black-and-white butterfly. Mimicked by female Common Diadem (page 310); see that species for separation. BEHAVIOUR: Flies strongly, but with frequent glides. E

298

Length:

WS ~ 6–7 cm | 2·5"

WHERE TO SEE: Fairly common in small numbers in most forest sites.

Madagascar Beauty* Salamis anteva

DISTRIBUTION: Forest throughout. ID: Beautiful orange, black, and white pattern above, but usually lands with wings closed, when only the cryptic leaf-like underside of the wings is visible. BEHAVIOUR: Floppy flight.

WS 5–7·5 cm | 2–3"

WHERE TO SEE: Almost anywhere.

Madagascar Forest Nymph* Aterica rabena

DISTRIBUTION: Forest throughout. ID: Distinctive pattern: largely dark brown forewing with white markings, and largely orange hindwing. BEHAVIOUR: Flies low within forest, and frequently perches on the ground with wings open. E

Length:

WS ~ 6 cm | 2·5"

WHERE TO SEE: Most conspicuous in western forest, as in Ankarafantsika and Tsingy de Bemaraha NPs.

1 2

BUTTERFLIES: ORANGE NYMPHALIDS II 1

Common Leopard Phalanta phalantha

DISTRIBUTION: Open areas throughout much of the Length: WS 4–5 cm | 1·5–2" Old World, including Madagascar. WHERE TO SEE: Almost ID: Mid-sized orange butterfly with copious black anywhere. markings. Very similar to Madagascar Leopard, but prefers open areas (Madagascar Leopard is always in forest). The lines along the edges of the wings are thin and wavy in Common Leopard, and thicker and straighter in Madagascar Leopard. BEHAVIOUR: Strong, fast flier. E

Madagascar Leopard* Phalanta madagascariensis

DISTRIBUTION: Forest throughout. ID: Mid-sized orange butterfly with black markings. Very similar to Common Leopard, but has thicker and straighter black lines at the edges of its wings, and is found in forest, whereas Common Leopard prefers open areas. 3

300

WS ~ 4–5 cm | 1·5–2"

WHERE TO SEE: Less common than Common Leopard, although possible in most forest sites.

Common Joker Byblia anvatara

DISTRIBUTION: Throughout most of Afrotropics. Found in almost any open habitat, especially humancreated ones. Absent only from high mountains and rainforest interior. ID: Distinctive and beautiful pattern of orange, buff, and brown. BEHAVIOUR: The erratic flight is rather moth-like.

Length:

WS 4–5 cm | 1·5–2"

WHERE TO SEE: Almost anywhere, including Tana.

Lines along edges of wings are thick and fairly straight

Complex patterning above and below

Lines along edges of wings are thin and wavy

BUTTERFLIES: ORANGE NYMPHALIDS III 1

Polka Dot Pardopsis punctatissima

DISTRIBUTION: From South Africa to Ethiopia, and throughout Madagascar. Prefers open habitats created by humans, such as pastures. ID: Similar in shape to acraeas, but small, widely spaced, black ‘polka dots’ are unmistakable. BEHAVIOUR: Flight is weak and low to ground. 2

Length:

WS 3–4 cm | 1–1·5"

WHERE TO SEE: Possible almost anywhere, although inconspicuous.

Tree nymphs Sevenia spp. [3 species]

DISTRIBUTION: An Afrotropical genus. All three Length: WS ~ 6 cm | 2·5" Malagasy species are endemic, found in forests. WHERE TO SEE: Present, ID: Dull orange above. Underside of hindwing is fuzzy although inconspicuous, in grey with darker spots and bars. Sevenia howensis (not most forest sites. illustrated) is dark and heavily marked on the underside of the hindwing; S. madagascariensis (not illustrated) is pale and faintly marked; S. amazoula 2a is intermediate, with a pale base colour but fairly bold markings. BEHAVIOUR: Slow, gliding flight. Often land in trees and hang upside down. 3

Acraeas Acraea spp. [18 species]

DISTRIBUTION: Large Old World tropical genus. Ten species are endemic to Madagascar. Most species are restricted to forest, but some inhabit open areas. ID: A diverse group of long-winged butterflies. All show red or orange, with black markings. Many have partially transparent wings. BEHAVIOUR: Distinctive slow flight with frequent glides. Often in the mid-storey and canopy of forest.

302

Acraea turna Ifaty

Length:

WS 4–7 cm | 1·5–3"

WHERE TO SEE: Multiple species occur in all forest sites, and a few are usually common.

Sevenia amazoula 3â&#x2122;&#x20AC;

Acraea dammii Lokobe NP

Acraea ranavalona Lokobe NP

BUTTERFLIES: VARIOUS NYMPHALIDS (EMPERORS, COMMODORE and PANSY) 1

Emperors Charaxes spp. [9 species]

DISTRIBUTION: A large Old World genus. All of the nine Length: WS 5–10 cm | 2–4" Malagasy species are found exclusively in forest, and eight WHERE TO SEE: Not often of them are endemic. observed, but at least one ID: Pointed forewing, and one or two tails on the species is found in most forest hindwing. A variable group, but all species show a jagged, sites. complex pattern below, visible when the wings are closed after landing. The upperwing has a black pattern on a base colour of white, beige, or orange. BEHAVIOUR: Distinctive flight alternates powerful wingbeats and quick glides on flat wings. Found mainly in forest canopy. Territorial; will defend food sources. R

Madagascar Brown Pansy* Junonia goudotii

DISTRIBUTION: Found in Madagascar, the Comoros, and Mauritius. Present in many habitats, from humancreated savannah to forest edge. ID: Distinctive brown and orange pattern above. BEHAVIOUR: All pansies slowly open and close their wings when perched. E

Length:

WS 5 cm | 2"

WHERE TO SEE: Possible almost anywhere, though in small numbers.

Madagascar Commodore* Precis andremiaja

DISTRIBUTION: Rainforest edges and adjacent scrub. ID: Above, male has a white stripe, and female a broader orange one. Below, both sexes are cryptic and leaf-like. BEHAVIOUR: Slowly opens and closes wings after landing.

Length:

WS ~ 6 cm | 2·5"

WHERE TO SEE: Common in many places such as around Andasibe and Masoala.

Club-tailed Charaxes Charaxes zoolina Zombitse-Vohibasia NP

304

Charaxes antamboulou Zombitse-Vohibasia NP

Charaxes andara Tsingy de Bemahara

Female (below); male has narrower white bands through both wings

3â&#x2122;&#x20AC;

BUTTERFLIES: PANSIES

Brilliant Blue Junonia rhadama

DISTRIBUTION: Mascarenes, Comoros, Seychelles, and Madagascar. Found at forest edge, and in a variety of open and human–modified habitats. ID: Shows more blue than other blue pansies, especially in the forewing. Male brighter than female. e

WS ~ 4–5 cm | 1·5–2"

WHERE TO SEE: Widespread and common, even in the gardens of Tana.

Yellow Pansy Junonia hierta paris

DISTRIBUTION: Old World tropics. Found throughout Madagascar in open and human-modified habitats, and forest edge. Absent from rainforest and the High Plateau. ID: Distinctive pattern of rich buff and dark brown with dark blue or purple spots above. Male has brighter colours than the more subdued female. 3

Length:

WS 4–5 cm | 1·5–2"

WHERE TO SEE: Almost anywhere in drier habitat.

Blue Pansy Junonia oenone

DISTRIBUTION: Afrotropics. Throughout Madagascar Length: WS 4–5 cm | 1·5–2" in most open habitats except high mountains and the WHERE TO SEE: Almost arid southwest. anywhere. ID: Large blue patch on the hindwing above, bigger and brighter on male. Female has several ‘eye spots’, and can resemble the much less common Eyed Pansy Junonia orithya (not illustrated), but always shows blue at the wing centre, rather than towards the edges of the wings. BEHAVIOUR: The male is aggressively territorial.

Underwing pattern subdued, but mirrors pattern above

306

1♂

1♀

Bolder pattern below than other pansies 2♂

2♀

Large buff patches above 3♂

3♀

BUTTERFLIES: BLACK-AND-WHITE NYMPHALIDS I

Clouded Mother-of-Pearl Protogoniomorpha anacardii duprei

DISTRIBUTION: Much of Afrotropics. Widespread in forest and secondary habitat in Madagascar. ID: Pattern above distinctive, even in flight: white with black borders. Female has broader dark edges than male. Below, cryptic and leaf-like. BEHAVIOUR: Bounding, floppy flight. Generally holds wings closed when perched, but sometimes opens and closes them slowly. E

308

Length:

WS ~ 7–9 cm | 3–3·5"

WHERE TO SEE: Small numbers in rainforest sites such as Andasibe, Ranomafana, and Amber Mountain NPs.

African Map Butterfly Cyrestis camillus elegans

DISTRIBUTION: Afrotropics. In Madagascar, found in rainforest and adjacent secondary habitat. ID: Beautiful, unmistakable species with a white base colour and vaguely cartographic black and yellowish markings. BEHAVIOUR: Visits muddy spots, as along roads.

WS 6–7·5 cm | 2–3"

WHERE TO SEE: Most common in rainforest, for example around Amber Mountain and Andasibe.

Madagascar Diadem Hypolimnas dexithea

DISTRIBUTION: Eastern and northern rainforest, and locally in western dry forest. ID: Unmistakable; one of Madagascar’s most spectacular butterflies. A complex pattern of white, rufous-brown and black. BEHAVIOUR: Usually stays high in forest canopy. A strong and high flier. e

Length:

WS 4–5 cm | 1·5–2"

WHERE TO SEE: Generally uncommon. Lokobe NP is one good site.

BUTTERFLIES: BLACK-AND-WHITE NYMPHALIDS II

Spotted Sailer Neptis saclava saclava

DISTRIBUTION: Widespread in Afrotropics. Found in both primary forest and disturbed habitats in Madagascar. ID: See similar Madagascar Sailer. They can occur together, but Spotted Sailer is more likely in secondary habitat. BEHAVIOUR: Like Madagascar Sailer. E

Length:

WS 4–5 cm | 1·5–2"

WHERE TO SEE: Fairly common and widespread.

Madagascar Sailer* Neptis kikideli

DISTRIBUTION: Found in forest throughout Madagascar. Length: WS ~ 4–5 cm | 1·5–2" ID: Sailers have a dark grey base colour and a broad band WHERE TO SEE: Fairly common of white across both wings. They are smaller than other in most forest sites. pied butterflies. Madagascar Sailer is very similar to Spotted Sailer, but has a broader band of white on both wings, and a subtly different pattern of white spots on the forewing. BEHAVIOUR: Slow flight with frequent glides. Usually lands with wings open. e

False Chief Pseudacraea lucretia apaturoides

DISTRIBUTION: Afrotropics. Found in rainforest in Madagascar. ID: Bold black-and-white pattern with veins running through the white patches. Imerina Chief Pseudoacraea imerina (not illustrated) has similar shape and overall colour, but has bluish spots rather than a pattern of veins. BEHAVIOUR: Unhurried flight. Often glides with wings open. 4

Length:

WS 6–8 cm | 2·5–3"

WHERE TO SEE: Quite common. Good areas include Andasibe, Ranomafana, and Anjozorobe.

Common Diadem Hypolimnas misippus

DISTRIBUTION: Worldwide tropical distribution. Length: WS 6–8 cm | 2·5–3" Occurs widely in Madagascar, although absent from WHERE TO SEE: Widespread the High Plateau and the southwest. Found in humanbut generally uncommon. modified habitats. Often near coastal towns. ID: The male has a distinctive pattern: a black base colour, with large white blotches and purplish highlights. The female mimics the unpalatable African Monarch (page 298) with remarkable accuracy, but has a heavier build and subtly different shape: a slightly more wavy trailing edge to the hindwing and a slightly more curvaceous trailing edge to the forewing. 4♂

310

4♀

4♂

4♀

BUTTERFLIES: SATYRS 1

Evening Brown Melanitis leda

DISTRIBUTION: Old World tropics and Australia. Found Length: WS 6–7 cm | 3" throughout Madagascar in open habitats and forest edge WHERE TO SEE: Widespread but absent from high mountains. but uncommon and ID: A large brown butterfly, with different dry and wet inconspicuous. season forms: conspicuous ‘eye-spots’ in the wet season, and few or none in the dry season. Larger than Madagascar and Indian Ocean satyrs. BEHAVIOUR: Active at dusk or on cloudy days. Reluctant to fly, and flies weakly. E

Madagascar satyrs* Strabena spp. [37 species]

DISTRIBUTION: A large, endemic genus. Various species can be found all around the island. Most live in forest, but some occur in secondary habitat and grasslands, more so than in the Indian Ocean satyrs. ID: Generally small and brown. Similar to Indian Ocean satyrs, but separated by the pattern on the forewing. Both groups show an orange eyespot ring with a smaller black iris inside. Madagascar satyrs usually have the main eyespot towards the tip rather than the centre of the forewing, and it contains two bluish pupils. In Indian Ocean satyrs, the main eyespot is closer to the centre of the forewing, and has only one whitish pupil inside the black iris. BEHAVIOUR: Fairly weak, skipping flight, low to the ground. These are among the few butterflies to be found in the understorey of thick forest. Feed on nectar. R

WS ~ 4–6 cm | 1·5–2·5"

WHERE TO SEE: In general, much less commonly seen than Indian Ocean satyrs. 2

Madagascar and Indian Ocean satyr forewing eyespots

Indian Ocean satyrs* Heteropsis spp. [64 species]

DISTRIBUTION: A large, regionally endemic genus. Various species are found all around the island. Most live in forest, but a few occur in secondary habitat and grasslands. ID: See Madagascar satyrs. BEHAVIOUR: Like Madagascar satyrs, but feed mainly on fruit. 3

312

Length:

WS ~ 4–6 cm | 1·5–2·5"

WHERE TO SEE: Can be found in most places, although not usually in gardens or towns.

Heteropsis turbans Ranomafana NP

Heteropsis vola Andasibe

Wet season form very plain

1 Strabena tamatavae High Plateau

Dry season form with eye spots Strabena triophthalma Anjozorobe-Angavo forest

Heteropsis difficilis Ranomafana NP

Heteropsis subsimilis Amber Mountain NP

Heteropsis narcissus Masoala

BUTTERFLIES: SKIPPERS, AND MOTHS

Comet Moth Argema mittrei

Papillon Comète de Madagascar

DISTRIBUTION: Endemic to Malagasy rainforest. ID: Huge spectacular yellow silk moth with large eye-spots and long tails. BEHAVIOUR: Only lives for a few days. E

Length:

Madagascar Sunset Moth Chrysiridia rhipheus

DISTRIBUTION: Throughout most of Madagascar in rainforest, western dry forest, and secondary habitats. ID: A large and beautiful multi-coloured and multi-tailed moth that is often mistaken for a butterfly. BEHAVIOUR: Flies during the day, unlike most moths. 3

Some skippers flat-winged: Eagris sabadius Amber Mountain NP

Length:

Lolo Andriana

WS 7–9 cm | 3–3·5"

WHERE TO SEE: Most likely in eastern rainforest sites such as Andasibe and Masoala.

Coeliades rama Kirindy Forest

Length:

Most skippers plain: Parnara naso Diego-Suarez

WS 2–7 cm | 1–3"

WHERE TO SEE: Widespread, but generally inconspicuous.

Offset wing posture unique to skippers: probable Fulda coroller Diego-Suarez

Some skippers quite colourful

314

Skippers Hesperiidae [56 species]

DISTRIBUTION: Large, worldwide family that is well represented in Madagascar. Most species are found in forest, but some use secondary habitats and grassland. ID: Moth-like butterflies, with thick bodies and large eyes. Some are fairly colourful, but most are brown, grey, or dull orange. BEHAVIOUR: Fast, bounding flight gives the family its common name. 3♂

WS ~ 15–20 cm | 6–8"

WHERE TO SEE: Ranomafana area.

Skippers show a variety of postures when at rest: some species hold their wings closed, others open, and some partially spread.

ICONIC PLANTS I

Madagascar supports over 11,000 species of plant, most of which are endemic. It boasts eight endemic and near-endemic families and over 300 endemic genera. Six of the world’s nine species of baobab are found only on the island. It supports about 850 species of orchid, and more than twice the diversity of palms than the continent of Africa. Although this guide is primarily focused on animals, a few of the most conspicuous and distinctive plants are highlighted here. E

Traveller’s Palm Ravenala madagascariensis

M Ravinala Arbre du Voyageur

DISTRIBUTION: Originally endemic to Madagascar, but Height: < 20 m | 65’ introduced around the warmer parts of the world as an WHERE TO SEE: Gardens all ornamental species. Thrives in human-modified areas and around the world! forest edges. There are whole ‘forests’ of Traveller’s Palm in some disturbed areas. ID: The mature trees are crowned by a broad fan of palm-like fronds. Its iconic shape is featured on the Air Madagascar logo and in many other places. The Traveller’s Palm was so named because of the water that accumulates at the base of its huge leaves, a godsend for travellers who found themselves without water. It is not actually a palm, but a member of the bird-of-paradise (Strelitziaceae) family. Ruffed lemurs (page 52) seem to be this tree’s main natural pollinator. 2

Pandanus Pandanus spp.

Vakoana Baquois

DISTRIBUTION: Almost 100 of the world’s 700 Pandanus Height: < 20 m | 65’ species are endemic to Madagascar. On the island, they WHERE TO SEE: Any Malagasy are found in all types of forest, from spiny forest to forest, from spiny forest to rainforest. Many species thrive in swamps. rainforest. ID: Some species are shrubs, while other can be tall trees. Most have exposed supporting roots at their base. The leaves are long, thin and often pointed. Pandanus is sometimes called ‘pandanus palm’ or 'screw pine' but is neither a palm nor a pine, belonging instead to the Pandanaceae family.

Traveller's Palm can dominate vast areas of savannah-like habitat

316

Iconic fan shape

Most species have a spiny trunk (above) and exposed supporting roots (below)

ICONIC PLANTS II 1

Octopus tree Didiereaceae [11 species]

M Sogno Arbre Pieuvre

DISTRIBUTION: Formerly thought to comprise a Height: < 15 m | 50’ Malagasy endemic family, although some continental WHERE TO SEE: One of the African species have recently been placed in this family. dominant tree families in the In Madagascar, there are four genera and 11 species, all spiny forest. found in the spiny forest of the southwest. ID: One of the most striking components of Madagascar's unique spiny forest. All species have spines and small leaves that grow close to the main trunk. The 'classic' species, including the widespread Didierea madagascariensis, have a profusion of tentacle-like trunks reaching toward the sky. 2

Pachypodium Pachypodium spp.

DISTRIBUTION: Most of the world’s 25 or so species are endemic to Madagasacar. They grow mainly in dry and rocky places, and thrive in the southwest. ID: Succulent plants with variable, but generally unusual structure. The trunks of some species have a swollen base, while others have a bottle-like shape. Many are covered in spines, which serve to condense fog that runs down and waters the plant.

Height:

Vontake < 6 m | 20’

WHERE TO SEE: Southwestern spiny forest, for example around Toliara and Ifaty. Also prominent around Isalo.

Some pachypodiums have bottle-shaped trunks

318

Didierea madagascariensis

Pachypodium rosulatum

BAOBABS I

Six of the world’s nine baobabs are endemic to Madagascar. They are found in dry parts of the southwest, west and north, and lose their leaves during the dry season. E

Western Baobab* Adansonia za

DISTRIBUTION: Dry forest of the west and southwest. Found at elevations up to 800 m (2,600’). The most widespread Malagasy baobab. ID: Large, with grey bark, fairly straight trunk and rounded crown. Fruit is variable in shape, but is always somewhat oval and never round. See Grandidier's and Northwestern Baobabs for separation from those species.

M Za Baobab Za*

NEAR THREATENED Height:

< 30 m | 100’

WHERE TO SEE: Berenty, Toliara area, Zombitse-Vohibasia NP, and Kirindy Forest.

No living Malagasy animal is capable of dispersing baobab seeds; it is likely that the ‘gorilla lemurs’, prior to their extinction, were the main dispersers of the seeds of these huge trees. E

Grandidier’s Baobab Adansonia grandidieri

DISTRIBUTION: Fairly small distribution in the southern part of the west. ID: Huge, stately, columnar baobab with pale grey bark. Similar to Western Baobab, but has a different shape: a long, bare trunk with a flat array of straight branches at the top, whereas Western has branches starting just above the middle of the tree, and some of these branches hang down irregularly. Grandidier’s Baobab has an oval-shaped fruit, more rounded than the elongated oval fruit of Western Baobab.

M Reniala Baobab de Grandidier

ENDANGERED Height: Diameter:

< 30 m | 100' < 3 m | 10’

WHERE TO SEE: Around Morondava, most famously at the ‘Allée des Baobabs’ northeast of town.

The Malagasy name for Grandidier’s Baobab is ‘Reniala’, meaning ‘mother of the forest'. E

Northwestern Baobab* Adansonia madagascariensis

DISTRIBUTION: Dry forest of the northwest and north. ID: Medium-sized baobab with smooth grey bark. Often tall and thin. Best separated from Western Baobab by its round rather than oval-shaped fruit. It is difficult to age baobabs, but some African specimens have proven to be well over 1,000 years old.

320

M Bozy Baobab du Nord-ouest*

NEAR THREATENED Height: Diameter:

< 20 m | 65' < 3 m | 10’

WHERE TO SEE: Ankarafantsika and Ankarana NPs, and around Diego-Suarez.

Grey bark and straight trunk 3

Tall and majestic, with smooth grey bark Often very tall and thin

BAOBABS II

Diego-Suarez Baobab* Adansonia suarezensis

DISTRIBUTION: Tiny range in dry forest around DiegoSuarez, in the far north. ID: A beautiful baobab with burgundy bark, and often a ‘T’- or ‘Y’-shaped form, with heavy branches spreading from near the top of the tree. Elongated oval fruits. Perrier’s Baobab Adansonia perrieri (not illustrated) is a rare northern species that can be seen in Ankarana National Park. E

M Bozy Diego* Baobab de Diego-Suarez*

ENDANGERED Height: Diameter:

WHERE TO SEE: On the slopes of Montagne des Français, along the road between Diego-Suarez and the beach town of Ramena.

Fony Baobab* Adansonia rubrostipa

DISTRIBUTION: Found in dry forest below 300 m (1,000’) elevation, from central western Madagascar to the northern part of the southwest. ID: The smallest and most distinctive baobab: reddish bark, round fruit, serrated leaves and usually a bottleshaped trunk.

< 25 m | 80' < 2 m | 7’

M Fony Baobab Fony*

NEAR THREATENED Height:

< 20 m | 65’

WHERE TO SEE: Common around Toliara and Ifaty.

The famous baobab in Mahajanga, in the middle of the roundabout (traffic circle) where the town’s main road reaches the coast, is actually an African Baobab Adansonia digitata, not a Malagasy species.

2 2

322

Baobabs bloom at the beginning of the rainy season and have large and spectacular flowers.

Baobabs have hard fruits covered in fine hair. They vary in shape from round to oval depending on the species.

Maroon bark

Reddish bark and bottle shape

Glossary of terms aestivate Afrotropics

arboreal arthropod bare parts call carrion caruncle casque CL Comoros

dabble / dabbling

diurnal dry season

epiphyte Europa gallery forest Glorieuses 324

to go into a state of torpor similar to hibernation ecozone that includes sub-Saharan Africa and all the Indian Ocean islands of the Malagasy region. Sometimes also considered to include the southern part of the Arabian Peninsula and small parts of Iran and Pakistan living in trees invertebrates with exoskeletons, segmented bodies and jointed appendages. They include the insects, spiders and crustaceans, and make up the phylum Arthropoda parts of a bird not covered in feathers, namely legs, bill and any exposed skin around the eye and the base of the bill shorter and simpler bird vocalizations that are used mainly for communication between birds: to stay in touch, warn of predators, or to share food sources (see also song) flesh of dead animals an odd type of bare part shown by asities, an endemic Malagasy bird family hard, helmet-like projection on the head of some chameleons carapace length: the standard length measurement used for turtles, terrapins and tortoises islands lying in the Indian Ocean northwest of Madagascar. Geographically, Mayotte is part of the Comoros, although politically it is part of France, whereas the other Comoran islands comprise their own country a feeding strategy in which the bill is moved around in shallow water. Often used to refer to ‘dabbling ducks’ which feed on the surface of the water as opposed to ‘diving ducks’ which dive to feed active during the day the portion of the year that is cooler, with lower rainfall. Generally between April and October in the east. The rains are shorter, more sporadic, and usually arrive later in the west. In the southwest they can fail altogether a plant that grows on another plant, but which does not parasitize it small island off the coast of southern Madagascar. Politically part of France, although claimed by Madagascar band of forest that grows along a watercourse, usually in dry country French possession consisting of a group of tiny islands northwest of Madagascar. Also known as the Glorioso Islands

Gondwanaland high plateau

invertebrate Kirindy Forest

Malagasy Region Mascarenes Mayotte MNP Neotropical nocturnal NP occipital lobe Old World onomatopoeia Pandanus primary (forest) rainforest

riparian (forest)

the southern of two supercontinents formed by the breakup of Pangea. It contained present-day South America, Antarctica, Arabia, Africa, Australia and peninsular India elevated part of central Madagascar comprising the contiguous area with an elevation above 800 m (2,600’). Divided from the northern Madagscar highlands by an area of lowlands. Often referred to by its French name: ‘Hauts-Plateaux’ animal lacking a backbone privately owned forest reserve that is part of a former logging concession. Properly referred to as Kirindy CNFEREF, and not to be confused with Kirindy-Mitea National Park, which lies south of Morondava Madagascar, the Comoros (including Mayotte), the Seychelles, the Mascarenes, a few remote outlying islands, and the surrounding waters the islands of Réunion, Mauritius and Rodrigues, which lie east of Madagascar. Réunion is part of France, while Mauritius and Rodrigues make up their own country island lying northwest of Madagascar. Geographically part of the Comoros, but politically part of France Madagascar National Parks, the Malagasy government agency responsible for the management of the national protected area system the tropics of the New World: southern Mexico, the Carribean, central America and much of South America active at night national park a flap of skin at the back of the head, possessed by some chameleons Europe, Africa and Asia the formation of a word that imitates a natural sound. Some bird names are onomatopoeic, reflecting their voice (e.g. cuckoo) A genus of palm-like plants. Sometimes called ‘screw pine’ or ‘screw palm’ old forest that has never been cut or suffered significant disturbance. Sometimes also called ‘virgin forest’ or ‘old-growth forest’ forest with high rainfall, in which most trees stay green and retain their leaves year-round. This term is sometimes reserved for lowland humid forest, but in this book is used for all humid forest, from sea-level up to the treeline along the banks of a river or stream 325

Sambirano forest

forest type in far northern Madagascar, which resembles eastern rainforest. Some of this forest is on the west coast, providing an exception to the general rule of Madagascar’s east being wet while its centre and west are dry secondary (forest / habitat) forest or other habitat that has regenerated after the destruction of the original, primary forest or other habitat Seychelles a nation of 115 islands that lies northeast of Madagascar SNR Strict Nature Reserve, a type of protected area that is manged by Madagascar National Parks (MNP) song a longer and more complex bird vocalization whose primary purposes are to attract a mate and defend a territory (see also call) spiny forest arid, scrubby habitat found in southwestern Madagascar. Sometimes called ‘spiny thicket’ or ‘spiny desert’. It is not arid enough to be truly considered desert, but lacks the closed canopy of true forest. Prominent vegetation includes baobabs and octopus trees Sub-Saharan Africa the portion of the African continent that lies south of the Sahara Desert SVL snout-to-vent length. This measurement is often used for reptiles, many of whose tails are variable in length, often curled up, and/or prone to being shed taxonomy the scientific study of naming species and classifying the relationships between them terrestrial dwelling on or in the ground TL total length. For most animals, this is the length from the tip of the nose to the tip of the tail tsingy Malagasy word used to describe a landscape of eroded limestone pinnacles underparts the part of an animal between the chin and the vent, including the breast and belly western dry forest the predominant type of forest in western Madagascar. Also found locally in the north. Much more open than rainforest, and seasonally very dry. This is a deciduous forest: meaning that many trees lose their leaves during the dry season wet season (rainy season) the portion of the year that is warmer, with more rainfall. Generally between November and March in the east. The rains are shorter, more sporadic, and usually arrive later in the west. In the southwest they can fail altogether wingbar(s) on birds, one or more distinct bands across the coverts, at the top of the wing WS wingspan (used as a measurement for butterflies and bats) 326

Further reading and useful resources Mammals Les Carnivora de Madagascar (2012) by Steven M. Goodman. Association Vahatra. Les Chauves-souris de Madagascar (2011) by Steven M. Goodman. Association Vahatra. Lemurs of Madagascar (2010) by Russell A. Mittermeier et al. Conservation International. Mammals of Madagascar (2007) by Nick Garbutt. Yale University Press. Les Petits Mammifères de Madagascar (2011) by Voahangy Soarimalala and Steven M. Goodman. Association Vahatra.

Birds The Birds of Africa: Volume VIII (2013) by Roger Safford and Frank Hawkins. Christopher Helm. Birds of Madagascar (1998) by Pete Morris and Frank Hawkins. Yale University Press. Birds of the Indian Ocean Islands (2013) by Ian Sinclair and Olivier Lagrand. Struik Nature. Guide to the Birds of Madagascar (1990) by Olivier Langrand. Yale University Press.

Reptiles and amphibians Les Amphibiens des Zones Arides de l’Ouest et du Sud de Madagascar (2014) by Franco Andreone, Gonçalo M. Rosa, and Achille P. Raselimanana. Association Vahatra. Complete Guide to Scientific and Common Names of Reptiles and Amphibians of the World (1996) by Norman Frank and Erica Ramus. A Field Guide to the Amphibians and Reptiles of Madagascar (2007) by Frank Glaw and Miguel Vences.

Butterflies Butterflies of South Africa (2012) by Steve Woodhall. Struik Nature.

Plants Baobabs de Madagascar (2012) by Andry Petignat. Arboretum d’Antsokay. Guide des Plantes Succulentes du Sud-ouest de Madagascar (2009) by Andry Petignat and Blaise Cooke. Arboretum d’Antsokay.

General Les Animaux et Ecosystèmes de l’Holocène Disparus de Madagascar (2013) by Steven M. Goodman and William L. Jungers. Association Vahatra. Atlas of Selected Land Vertebrates of Madagascar (2014) edited by Steven M. Goodmand and Marie Jeanne Raherilalao. Association Vahatra. The Natural History of Madagascar (2007) edited by Steven M. Goodman and Jonathan P. Benstead. University of Chicago Press.

Online resource

The IUCN Red List of Threatened Species. www.iucnredlist.org 327

Acknowledgements and photo credits Malagasy local guides and biologists have been instrumental in our exploration of their country and learning about its wildlife. Some of the most important of these have been RAJERIARISON Emile, RAJERISOA Luc, RALAHY Andriamasy (Ndrema), RATSISAKANANA BESOA Maurice, RAZAFIMANANTSOA Angeluc and TAREHA Sylvain Christian. Steve Goodman reviewed the bird and mammal sections and provided invaluable input. The assistance and expert review of Devin Edmonds, Gonçalo M. Rosa and Mark Scherz was essential in allowing us to cover reptiles and amphibians. Likewise, butterflies could not have been so thoroughly or accurately covered without the help of David Lees. Josh Engel, Iain Campbell, Charley Hesse, Christian Boix, Derek Schuurman, Nivo Ravelojaona and Ian Sinclair provided various forms of help and advice over the years. Janet Behrens and Chris and Judith Gibson reviewed and edited the entire manuscript. This book would not have been as attractive or complete without the support of many contributing photographers, to whom we extend our sincere thanks. All the photographers are listed in the following photo credits section. We enjoyed the company of many different Tropical Birding clients while guiding tours in Madagascar. Several of these have allowed the use of their photos in the book. Robert Kirk of Princeton Univeristy Press believed in the concept for this guide. Rob Still, Rachel Still and Andy Swash of WILDGuides put the book together. Their patience, long hours, and desire to make everybody involved in the project satisfied with the end product is much appreciated.

PHOTO CREDITS Photographs not taken by Ken Behrens are gratefully acknowledged and each image is listed, together with the photographer’s initials, as follows: Nick Athanas (antpitta.com) [NA]; Keith Barnes [KB]; Steve Blain (steveblain.blogspot.com) [SB]; Iain Campbell (iaincampbell.smugmug.com) [IC]; Roger and Liz Charlwood (worldwildlifeimages.com) [RLC]; Leslie Clapp [LC]; Greg and Yvonne Dean (worldwildlifeimages.com) [GYD]; Devin Edmonds [DE]; Charley Hesse [CH]; Andrew Hollander [AH]; Rob Hutchinson [RH]; Chris Krog [CK]; Markus Lagerqvist (pbase.com/lagerqvist) [ML]; David Lees [DL]; Sandrine Martinez (flickr.com/photos/msandrine) [SM]; Pete Morris (birdquest-tours.com) [PM]; Gonçalo M. Rosa [GR]; Daniel Rosenberg (flickr.com/photos/cowyeow) [DR]; Rob Schell (flickr.com/photos/robschellphotography) [RS]; Dubi Shapiro (pbase.com/ dubisha) [DS]; Dave Smallshire (flickr.com/photos/davesmalls) [DSm]; Tom Stephenson [TS]; John Sullivan (wildherps.com) [JS]; Kristian Svensson (flickr.com/photos/macronyx) [KS]; Andy and Gill Swash (worldwildlifeimages.com) [AGS]; Warwick Tarboton (warwicktarboton.co.za) [WT]; Nigel Voaden (flickr.com/photos/nvoaden) [NV] and John Wilkinson [JW]. 328

Images sourced via the photographic agencies Agami (agami.nl), Naturepl (naturepl.com) or Shutterstock (shutterstock.com) are credited in full. Images reproduced under the terms of the Creative Commons Attribution-ShareAlike 3.0 Unported license, or the GNU Free Documentation License, Version 1.2 or any later version published by the Free Software Foundation, are also credited in full (these are indicated by “/CC” after the photographer’s name in the list); in this respect we would particularly like to acknowledge Frank Vassen (flickr.com/photos/42244964@N03) who provided many images. Images that are in the public domain are credited to the source followed by “(public domain)”.

Introduction 16 Rufous Mouse Lemur [Ryan M. Bolton/ Shutterstock]; Giraffe-necked Weevil [KB]; Comet Moth [KB]. 18 Landscape [Frank Vassen/CC (Flickr)]. 18 Cream-lined Swallowtail [SB]; White-fronted Brown Lemur [Frank Vassen/CC (Flickr)]. 19 Speckled Day Gecko [GR]. 21 Green Sea Turtle [Wouter Hagens/CC]; Bottlenose dolphin [NASAs (public domain)]; Lesser Frigatebird [RLC]. 22 Landscape [KB]. 22 Coquerel’s Sifaka [KB]. 23 Leaf-nosed snake [GR]. 24 Landscape [KB]. 29 Green Bright-eyed Frog [reptiles4all/ Shutterstock].

Mammals 33 Mouse lemur (1c) [Frank Vassen/CC (Flickr)]. 34 Dwarf lemur (1c) [NA]. 37 Coquerel’s Giant Mouse Lemur (1a) [SM]; Northern Giant Mouse Lemur (1b) [Russell Mittermeier/CC]. 38 Small-toothed Sportive Lemur [KB]. 39 Milne-Edwards' Sportive Lemur (1f) [SB]; White-footed Sportive Lemur (1c) [KB]. 40 Greater Bamboo Lemur (feeding) [KB]. 44 Ring-tailed Lemur (main photo) [SB]. 45 Ring-tailed Lemur (in tree) [KB]; (group on ground) [KB]. 47 White-fronted Brown Lemur (female) [KB]. 49 Red-bellied Lemur (male) [KB]. 51 Mongoose Lemur (male) [CH]; (female) [David C. Azor/Shutterstock]. 52 Black-and-white Ruffed Lemur [LC].

53 ‘Northern’ Black-and-white Ruffed Lemur [Frank Vassen/CC (Flickr)]. 55 Masoala Woolly Lemur (1a) [ML]; Eastern Woolly Lemur (1b) (with baby) [KB]. 57 Verreaux’s Sifaka (dancing) [IC]; Coquerel’s Sifaka [KB]. 59 Diademed Sifaka [RH]. 63 Indri (hanging) [KB]. 64 Aye-aye [javarman/Shutterstock]. 67 Ring-tailed Vontsira [DS]; Spotted Fanaloka [DSm]. 68 Fosa [SB]. 70 Madagascar Flying Fox [KB]. 71 Madagascar Flying Fox [KB]. 73 Mauritian Tomb Bat (two together) [Frank Vassen/CC (Flickr)]; Commerson’s Leaf-nosed Bat [JW]. 76 Common Tenrec [SB]. 79 Eastern Red Forest Rat (1b) [SB]; Black Rat [CSIRO/CC]; Western Tuft-tailed Rat [Nick Garbutt/ Naturepl]. 80 Bottlenose Dolphin [Truncatus (public domain)]. 81 Bottlenose Dolphin [NASAs (public domain)].

Birds 83 Scaly Ground-roller [AGS]; Blue Vanga [KB]. 85 Red-billed Duck (main photo) [CH]; White-faced Whistling-duck (main photo) [JW]. 87 Madagascar Partridge (female) [NA]. 89 Grey Heron (main photo) [KB]; Madagascar Heron (main photo) [DS]. 90 Black Herons [KB]. 329

91 Purple Heron (main photo) [NV], (flight) [KB]; Black Heron (main photo) [WT]. 92 Great Egret (inset) [KB]. 94 Madagascar Pond Heron [SB]. 95 Squacco Heron (main photo) [JW]. 96 Black-crowned Night Herons [KB]. 97 Black-crowned Night Heron (adult) [KB]. 99 Madagascar Buzzard (main photo) [NV]; Madagascar Harrier-hawk (adult) [AGS]. 101 Black Kite (main photo) [KB]; Frances’s Sparrowhawk (perched female) [KB]. 103 Madagascar Crested Ibis (both photos of bird) [RH]. 105 Common Moorhen (main photo) [KB]. 109 Greater Sand Plover (main photo) [AGS], (flight) [Daniele Occhiato/Agami]. 113 Whimbrel (main photo) [AGS], (flight) [Dave Montreuil/Shutterstock]. 115 Crab-plover [Adrin Shamsudin/ Shutterstock]. 118 Red-tailed Tropicbird [KB]. 119 Lesser Frigatebird (male and female) [RLC]. 121 Feral Pigeon (main photo) [KB]. 127 Madagascar Coucal (inset) [TS]. 130 Blue Coua [AGS]. 133 White-browed Owl (right) [SB]; Barn Owl (main photo) [CK]. 135 Madagascar Nightjar (main photo) [DS]; Collared Nightjar (main photo) [Frank Vassen/CC (Flickr)]. 137 Mascarene Martin (perched) [NV]. 139 Madagascar Malachite Kingfisher (left) [DS]. 141 Broad-billed Roller (left) [DS], (right) [AGS]. 143 Pitta-like Ground-roller [DS]; Rufous-headed Ground-roller [NA]. 145 Scaly Ground-roller [DS]. 147 Long-tailed Ground-roller (main photo) [KB]. 149 Schlegel’s Asity (male) [JW]; Common Sunbird Asity (female) [CH], (male) [DS]. 151 Pied Crow (main photo) [Ralph Griebenow/ Shutterstock]. 153 Madagascar Bulbul [KB]. 155 ‘Amber Mountain’ Forest Rock Thrush [DS]. 156 Subdesert Brush Warbler [AGS]. 330

161 Rand’s Warbler [PM]. 163 Long-billed Tetraka (male) [KB]; Spectacled Tetraka [AGS]; White-throated Oxylabes [PM]. 165 Souimanga Sunbird (breeding male) [DS], (transitional male) [KB], (female) [KB]; Madagascar Green Sunbird (male) [TS], (female) [KS]. 167 Red-tailed Vanga (male) [DS]; Chabert Vanga (main photo) [DS]. 169 Blue Vanga (female) [NA], (male) [DS]; White-headed Vanga (female) [NA], (male) [GYD]. 171 Nuthatch Vanga (female) [DS]; Crossley’s Vanga (male) [DS]. 175 Tylas Vanga (right) [KB]; Pollen’s Vanga (female) [DS]; Madagascar Cuckoo-shrike (female) [NA]. 177 Madagascar Starling (male) [CH], (female) [NA]. 179 Sakalava Weaver (female) [KB].

Reptiles 181 Madagascar tree boa [KB]. 183 Flat-tailed Tortoise (2a) Frank Vassen/CC (Flickr)]; Madagascan Big-headed Turtle (3a) [Ryan M. Bolton/Shutterstock]. 185 Green Sea Turtle [Bthv/CC]; Hawksbill Sea Turtle [LC]; Nile Crocodile [KB]. 187 Minute Leaf Chameleon [GR]. 189 Brygoo’s Leaf Chameleon [GR]; Elongate Ancient Leaf Chameleon [Axel Strauss/CC]. 191 Parson’s Chameleon [KB]. 192 Jewelled Chameleon [IC]. 193 Jewelled Chameleon (male) [GR], (female) [KB]. 194 Big Nose Chameleon [Frank Vassen/CC (Flickr)]. 195 Perinet Chameleon (male) [Gavinevans/CC]; Big Nose Chameleon [CH]. 197 Cryptic Chameleon (male) [Frank Vassen/ CC (Flickr)]; Short-horned Chameleon (male) [Frank Vassen/CC (Flickr)]. 204 Rhinoceros Chameleon [SM].

206 Labord’s Chameleon (male and female) [Frank Vassen/CC (Flickr)]. 209 Cuvier’s Madagascar Swift (tail detail) [KB]. 210 Marked Madagascar Swift [SB]. 218 Voeltzkowia rubrocaudata [GR]. 219 Gold-spotted Skink [AH]. 221 Koch’s Day Gecko [KB]; Lined Day Gecko [KB]. 223 Small Lined Day Gecko [GR]. 224 Modest Day Geckos [Gail Hampshire/CC (Flickr)]. 227 Grandidier’s Day Gecko [KB]. 229 Madagascar Clawless Gecko (top photo) [Frank Vassen/CC (Flickr)]. 231 Mocquard’s Ground Gecko [DR]. 233 Ambonihazo Madagascar Velvet Gecko [CH]. 235 Satanic Leaf-tailed Gecko (main photo) [Frank Vassen/CC (Flickr)]; Spearpoint Leaf-tailed Gecko (inset) [Alex Telford/CC]; Günther’s Leaf-tailed Gecko [Frank Vassen/ CC (Flickr)]; Lined Leaf-tailed Gecko [Frank Vassen/CC (Flickr)]. 239 Duméril’s Ground Boa [KB]. 241 Giant Hognose Snake [Frank Vassen/CC (Flickr)]; Leaf-nosed snake (male) [GR]; Parastenophis betsileanus [Frank Vassen/ CC (Flickr)]; Lycodryas gaimardi [GR]. 243 Striped Madagascar Garter Snake (left) [GR], (right) [CH]; Bernier’s Striped Snake (left) [GR]. 245 Trail Madagascar Garter Snake [CH]; Gluttonous Bighead Snake [GR].

Frogs 253 Boulenger’s Giant Treefrog (adult) [RS], (juvenile) [DE]. 257 Blue-legged Mantella [GR]; Guibé’s Mantella [GR]. 261 Red-eyed Bright-eyed Frog [DE]. 264 White-lipped Bright-eyed Frog [Axel Strauss/CC]. 265 Goudot’s Bright-eyed Frog (main photo) [DE], (inset) [IC]; White-lipped Bright-eyed Frog [Frank Vassen/CC (Flickr)].

266 Probable Guibemantis variabilis [Frank Vassen/CC (Flickr)]. 267 Guibemantis pulcher [Brian Gratwicke/CC]. 269 Madagascar Bullfrog (both images) [JS]. 271 Boulenger’s Bridge Frog (right) [KB]; Webb’s Bridge Frog (main photo) [Frank Vassen/CC (Flickr)]. 272 Beige Bridge Frog [KB]. 275 Ivohimanitra Madagascar Frog (right) [Frank Vassen/CC (Flickr)]. 279 Charlotte’s Madagascar Frog [RS]; Central Madagascar Frog [RS]. 280 Duméril’s Madagascar Frog [Ryan M. Bolton/Shutterstock]. 281 Folohy Madagascar Frog [DE].

Invertebrates 283 Giant African Land Snail [Forest & Kim Starr/CC]. 285 Giraffe-necked Weevil [Frank Vassen/CC (Flickr)]; Golden orb-web spiders [KB]. 287 Mocker Swallowtail (right) [Dave Rogers/ CC]; Cream-lined Swallowtail (right) [SB]. 289 Blue-banded Swallowtail (inset) [Bernard Dupont/CC]. 291 Madagascar Giant Swallowtail [KS]. 293 Eurema brigitta [Alslutsky/Shutterstock]. 295 African Wood White (left) [Charles J. Sharp/ CC]. 305 Charaxes antamboulou [KB]. 309 Clouded Mother-of-Pearl (right) [DL]; Madagascar Diadem (left) [KB], (right) [Michel-Georges Bernard/CC]. 311 False Chief (right) [DL]. 315 Comet Moth [KB]; Madagascar Sunset Moth (left) [DL], (right) [Frank Vassen/CC (Flickr)].

Plants 317 Traveller’s Palm [CEphoto, Uwe Aranas/CC]. 322 Fony Baobab (flower) [Axel Strauss/ CC (commons.wikimedia.org/wiki/ File:Adansonia_rubrostipa_02.jpg)]. 323 Fony Baobab [KB]. 331

Index This index includes the English and scientific names of all the species included in this book. Bold text is used to highlight a main species account. Grey text is used to indicate 'groups' of species which have a main species account. Regular text is used for species, or 'groups' of species, that are not subject to a full account. Italicized numbers relate to pages where a photograph may be found, for those species that do not have a full account. A Accipiter francesiae ............................................................... 100 — henstii .......................................................................................... 100 — madagascariensis ............................................................ 100 Achatina fulica .......................................................................... 282 Acraea ............................................................................................... 302 Acraea ................................................................................................. 302 — dammii ...................................................................................... 303 — ranavalona ............................................................................. 303 — turna ............................................................................................ 302 Acrantophis dumerili .......................................................... 238 — madagascariensis ............................................................ 238 Acridotheres tristis ................................................................. 176 Acrocephalus newtoni ........................................................ 158 Actitis hypoleucos ................................................................... 110 Actophilornis albinucha ................................................... 104 Adansonia digitata ................................................................ 322 — grandidieri ............................................................................. 320 — madagascariensis ............................................................ 320 — perrieri ....................................................................................... 322 — rubrostipa ................................................................................ 322 — suarezensis ............................................................................. 322 — za ..................................................................................................... 320 Aepyornis maximus ................................................................. 83 Agapornis canus ....................................................................... 124 Aglyptodactylus ........................................................................ 268 — laticeps ........................................................................................ 268 — madagascariensis ............................................................ 269 — securifer ..................................................................................... 268 Alectroenas madagascariensis ................................... 122 Amphiglossus frontoparietalis .................................... 219 Anas erythrorhyncha ............................................................. 84 Appias sabina .............................................................................. 294 Apus balstoni ............................................................................... 136 — barbatus .................................................................................... 136 Ardea alba ......................................................................................... 92 — cinerea ........................................................................................... 88 — humbloti ...................................................................................... 88 — purpurea ...................................................................................... 90 Ardeola idae .................................................................................... 94 — ralloides ........................................................................................ 94 332

Arenaria interpres .................................................................. 112 Argema mittrei .......................................................................... 314 Artamella viridis ...................................................................... 168 Asity, Common Sunbird ............................................. 148 —, Schlegel’s .............................................................................. 148 —, Velvet ........................................................................................ 148 Astrochelys radiata ................................................................ 182 Atelornis crossleyi ................................................................... 142 — pittoides ..................................................................................... 142 Aterica rabena ............................................................................ 298 Avahi ....................................................................................................... 54 — cleesei .............................................................................................. 54 — laniger ............................................................................................ 54 — meridionalis ............................................................................. 54 — mooreorum ............................................................................... 54 — occidentalis ............................................................................... 54 — peyrierasi .................................................................................... 54 Aye-aye ................................................................................................ 64 B Babbler, Crossley’s (see Crossley’s Vanga) .......................................... 170 Bamboo lemur ........................................................................... 40 Bamboo, Giant .................................................................... 40, 42 Baobab, African ..................................................................... 322 —, Diego-Suarez ................................................................... 322 —, Fony ........................................................................................... 322 —, Grandidier’s ..................................................................... 320 —, Northwestern .................................................................. 320 —, Perrier’s ................................................................................... 322 —, Western .................................................................................. 320 Bat, Commerson’s Leaf-nosed ................................ 72 —, Madagascar Free-tailed ............................................ 72 —, Madagascar Straw-coloured Fruit ............ 70 —, Mauritian Tomb ............................................................. 72 —, trident .......................................................................................... 72 Beauty, Madagascar .......................................................... 298 Bee-eater, Madagascar (or Olive) ..................... 138 Belenois grandidieri .............................................................. 294 — helcida ........................................................................................ 294 Bernieria cinereiceps ............................................................ 162 — madagascariensis ............................................................ 162

Bernieria, Long-billed (see Long-billed Tetraka) ................................ 162 Bighead Snake, Gluttonous ..................................... 244 Bird, Elephant ............................................................................... 83 Bismarckia nobilis .................................................................. 136 Blaesodactylus ............................................................................ 232 — ambonihazo .......................................................................... 233 — antongilensis ........................................................................ 232 — boivini ......................................................................................... 232 — sakalava .................................................................................... 233 Blommersia ................................................................................... 266 — blommersae ........................................................................... 267 — domerguei ............................................................................... 266 — wittei ............................................................................................ 266 Blue ........................................................................................ 296, 306 Blue, Brilliant .......................................................................... 306 —, Clover ....................................................................................... 296 —, Common Smoky ........................................................... 296 —, Common Zebra ............................................................. 297 —, Gaika .......................................................................................... 297 —, Sooty .......................................................................................... 297 Boa, Calabar ............................................................................... 238 —, Duméril’s Ground ..................................................... 238 —, Madagascar Ground ............................................... 238 —, Madagascar tree .......................................................... 238 Bokiboky, Northern ............................................................. 66 Boophis albilabris ................................................................... 264 — bottae ........................................................................................... 258 — doulioti ....................................................................................... 262 — goudoti ....................................................................................... 264 — luteus ........................................................................................... 260 — madagascariensis ............................................................ 262 — pyrrhus ....................................................................................... 260 — rappiodes .................................................................................. 258 — roseipalmatus ...................................................................... 262 — tephraeomystax ................................................................. 262 — viridis .......................................................................................... 260 Border, Madagascar Dotted .................................... 294 Brachypteracias leptosomus ......................................... 144 Bridge frog .................................................................................. 270 Bright Snake, Collared .................................................. 244 Bright-eyed frog ................................................................... 258 Brookesia brunoi ...................................................................... 188 — brygooi ....................................................................................... 188 — decaryi ........................................................................................ 188 — micra ............................................................................................ 187 — minima ...................................................................................... 186 — nasus ............................................................................................ 188 — peyrierasi ................................................................................. 186 — stumpffi ...................................................................................... 188 — superciliaris ........................................................................... 188 — tuberculata ............................................................................. 186 Brown, Evening ..................................................................... 312

Brush Warbler, Madagascar ................................... 156 —, Subdesert ............................................................................. 156 Bubulcus ibis ................................................................................... 92 Bulbul, Madagascar .......................................................... 152 Bullfrog, Indian ......................................................... 246, 248 —, Madagascar ....................................................................... 268 Buteo brachypterus ................................................................... 98 Butorides striata .......................................................................... 94 Butterfly, African Map .................................................. 308 Buttonquail, Madagascar .............................................. 86 Buzzard, Madagascar ........................................................ 98 Byblia anvatara ......................................................................... 300 C Calabaria reinhardtii .......................................................... 238 Calicalicus madagascariensis ..................................... 166 Calidris alba ................................................................................. 110 — ferruginea ................................................................................ 110 Calumma amber ...................................................................... 200 — ambreense ............................................................................... 200 — boettgeri .................................................................................... 202 — brevicorne ............................................................................... 196 — crypticum ................................................................................ 196 — gastrotaenia .......................................................................... 194 — glawi ............................................................................................. 194 — linotum ...................................................................................... 202 — nasutum .................................................................................... 194 — oshaughnessyi ..................................................................... 198 — parsonii ...................................................................................... 190 Canarium madagascariense ........................................... 64 Caper White, Grandidier’s ....................................... 294 —, Yellow-painted .............................................................. 294 Caprimulgus madagascariensis ............................... 134 Cat-eyed snake, Madagascar .................................. 242 Cathariostachys madagascariensis ................. 40, 42 Catopsilia ........................................................................................ 292 — florella ............................................................................ 292, 293 — thauruma ................................................................................ 293 Centropus toulou ..................................................................... 126 Chalarodon madagascariensis .................................. 208 Chameleon, Amber Mountain ............................ 200 —, Antimena ............................................................................. 204 —, Antongil Leaf .................................................................. 186 —, Arthur’s ................................................................................. 200 —, Band-bellied .................................................................... 194 —, Belted ....................................................................................... 198 —, Big Nose ................................................................................ 194 —, Blue-legged ........................................................................ 196 —, Boettger’s ............................................................................. 202 —, Brown Leaf ........................................................................ 188 —, Bruno’s Leaf ........................................................................ 188 —, Brygoo’s Leaf ................................................................... 188 —, Canopy ................................................................................... 194 333

Chameleon, Cryptic ......................................................... 196 —, Decary’s Leaf ..................................................................... 188 —, Elongate Ancient Leaf .......................................... 188 —, Glaw’s ....................................................................................... 194 —, Jewelled .................................................................................. 192 —, Labord’s ................................................................................. 206 —, Minute Leaf ....................................................................... 186 —, Montagne d’Ambre Leaf .................................... 186 —, Northern Blue-nosed ............................................ 202 —, Nose-horned (see Big Nose Chameleon) ............................. 194 —, O’Shaughnessy’s .......................................................... 198 —, Oustalet’s ............................................................................. 190 —, Panther ................................................................................... 192 —, Parson’s .................................................................................. 190 —, Perinet ..................................................................................... 194 —, Petter’s ..................................................................................... 202 —, Plated Leaf ......................................................................... 188 —, Rhinoceros ......................................................................... 204 —, Short-horned .................................................................. 196 —, Two-banded ...................................................................... 198 —, Warty ........................................................................................ 206 —, White-lined ....................................................................... 204 Charadrius bifrontatus ...................................................... 106 — leschenaultii .......................................................................... 108 — marginatus ............................................................................. 106 — mongolus .................................................................................. 108 — tricollaris .................................................................................. 106 Charaxes .......................................................................................... 304 — andara ........................................................................................ 305 — antamboulou ....................................................................... 305 — zoolina ........................................................................................ 304 Charaxes, Club-tailed ....................................................... 304 Cheirogaleus .................................................................................... 34 — andysabini ................................................................................. 35 — crossleyi ........................................................................................ 34 — major .............................................................................................. 35 — medius ........................................................................................... 35 — sibreei ............................................................................................. 34 Chelonia mydas ........................................................................ 184 Chief, False .................................................................................. 310 —, Imerina .................................................................................... 310 Chrysiridia rhipheus ............................................................ 314 Cinnyris notatus ....................................................................... 164 — souimanga .............................................................................. 164 Cisticola cherina ...................................................................... 158 Cisticola, Madagascar .................................................... 158 Civet, Small Indian ........................................................ 31, 66 Clawless Gecko, Madagascar ................................. 228 Climbing Frog, Ambatoharanana ................... 252 —, Barbour’s .............................................................................. 252 Cocos nucifera ............................................................................ 136 334

Coeliades rama .......................................................................... 314 Colotis evanthe .......................................................................... 292 Columba livia ............................................................................. 120 Commodore, Madagascar ......................................... 304 Compsophis laphystius ...................................................... 244 Cophyla ............................................................................................. 252 Copsychus albospecularis ................................................ 154 Coracina cinerea ...................................................................... 174 Coracopsis nigra ....................................................................... 124 — vasa ............................................................................................... 124 Corvus albus ................................................................................ 150 — splendens ..................................................................................... 82 Corythornis madagascariensis .................................. 138 — vintsioides ............................................................................... 138 Coua caerulea ............................................................................ 130 — coquereli ................................................................................... 128 — cristata ....................................................................................... 126 — delalandei ................................................................................ 130 — gigas .............................................................................................. 128 — olivaceiceps ............................................................................ 129 — reynaudii .................................................................................. 130 — ruficeps ....................................................................................... 128 — serriana ..................................................................................... 130 Coua, Blue .................................................................................... 130 —, Coquerel’s ........................................................................... 128 —, Crested .................................................................................... 126 —, Giant ......................................................................................... 128 —, Green-capped ................................................................... 129 —, Red-breasted .................................................................... 130 —, Red-capped ....................................................................... 128 —, Red-fronted ...................................................................... 130 —, Snail-eating ......................................................................... 130 Coucal, Madagscar ............................................................ 126 Crab-plover ................................................................................ 114 Crested Ibis, Madagascar ........................................... 102 Crested Tern, Greater ..................................................... 116 —, Lesser ........................................................................................ 116 Crocodile, Nile ....................................................................... 184 Crocodylus niloticus ............................................................. 184 Crossleyia xanthophrys ..................................................... 162 Crow, House .................................................................................. 82 —, Pied ............................................................................................. 150 Cryptoprocta ferox .................................................................... 68 Cuckoo-roller ........................................................................... 146 Cuckoo-shrike, Madagascar ................................... 174 Cuckoo, Madagascar (or Lesser) ....................... 126 Cuculus rochii ............................................................................. 126 Cyanolanius madagascarinus .................................... 168 Cypsiurus parvus .................................................................... 136 Cyrestis camillus ...................................................................... 308

D Danaus chrysippus ................................................................ 298 Daubentonia madagascariensis .................................. 64 Dendrocygna bicolor .............................................................. 84 — viduata .......................................................................................... 84 Deudorix antalus .................................................................... 296 — batikeli ........................................................................................ 297 Diadem, Common ............................................................. 310 —, Madagascar ....................................................................... 308 Dicrurus forficatus ................................................................ 176 Didierea madagascariensis ........................................... 319 Didiereaceae .............................................................................. 318 Dolphin, Common Bottlenose ............................... 80 —, Indo-Pacific Bottlenose ......................................... 80 —, Spinner ....................................................................................... 80 Dot, Polka ..................................................................................... 302 Dotted Border, Madagascar .................................... 294 Dove, Cloven-feathered ................................................. 122 —, Madagascar Turtle .................................................... 120 —, Namaqua .............................................................................. 120 —, Rock ........................................................................................... 120 Drepanoptila holosericea ................................................ 122 Dromas ardeola ........................................................................ 114 Dromicodryas bernieri ...................................................... 242 — quadrilineatus .................................................................... 242 Drongo, Crested .................................................................... 176 Dryolimnas cuvieri ............................................................... 104 Duck, Fulvous Whistling- .............................................. 84 —, Red-billed .............................................................................. 84 —, White-faced Whistling- ......................................... 84 Duttaphrynus melanostictus ....................................... 246 Dwarf gecko ............................................................................... 226 Dwarf lemur ................................................................................. 34 Dyscophus antongilii ........................................................... 250 — guineti ......................................................................................... 250 E Eagle, Madagascar Fish ................................................... 98 Eagris sabadius ......................................................................... 314 Ebenavia inunguis ................................................................. 228 Echinops telfairi ........................................................................... 74 Egret, Cattle .................................................................................. 92 —, Dimorphic (see Little Egret) ............................ 92 —, Great ............................................................................................. 92 —, Little ............................................................................................. 92 Egretta ardesiaca ........................................................................ 90 — dimorpha .................................................................................... 92 — garzetta ......................................................................................... 92 Eidolon dupreanum ................................................................. 70 Eliurus ................................................................................................... 78 — myoxinus .................................................................................... 79

Emperor .......................................................................................... 304 Eretmochelys imbricata .................................................... 184 Erymnochelys madagascariensis ............................. 183 Eucalyptus ......................................................................................... 70 Euchrysops malathana ...................................................... 296 Eulemur albifrons ...................................................................... 46 — collaris ........................................................................................... 48 — coronatus .................................................................................... 50 — fulvus .............................................................................................. 48 — macaco .......................................................................................... 50 — mongoz ......................................................................................... 50 — rubriventer ................................................................................ 48 — rufifrons ....................................................................................... 48 — rufus ................................................................................................ 48 — sanfordi ......................................................................................... 46 Eupleres goudotii ........................................................................ 68 Eurema .............................................................................................. 292 — brigitta ........................................................................................ 293 — desjardinsii ............................................................................. 293 — floricola ..................................................................................... 293 — hapale .......................................................................................... 292 Euryceros prevostii ................................................................. 172 Eurystomus glaucurus ....................................................... 140 F Falanouc ............................................................................................. 68 Falco concolor ............................................................................. 100 — eleonorae .................................................................................. 100 — newtoni ...................................................................................... 100 — peregrinus ............................................................................... 100 — sparverius ................................................................................ 100 — tinnunculus ........................................................................... 100 — zoniventris .............................................................................. 100 Falcon, Eleonora’s .................................................................. 100 —, Peregrine ............................................................................... 100 —, Sooty .......................................................................................... 100 Falculea palliata ....................................................................... 172 Fanaloka, Spotted .................................................................. 66 Fish Eagle, Madagascar ................................................... 98 Flycatcher, Madagascar Paradise ...................... 152 —, Ward’s (see Ward’s Vanga) ............................... 170 Flying Fox, Madagascar .................................................. 70 Fody, Forest ................................................................................. 178 —, Madagascar ....................................................................... 178 Forest Nymph, Madagascar .................................... 298 Forest rat ........................................................................................... 78 Fork-marked lemur ............................................................. 36 Fosa ......................................................................................................... 68 Fossa fossana .................................................................................. 66 Foudia madagascariensis ............................................... 178 — omissa ......................................................................................... 178

335

Fox, Madagascar Flying .................................................. 70 Fregata ariel .................................................................................. 118 — minor ........................................................................................... 118 Frigatebird, Greater ........................................................... 118 —, Lesser ........................................................................................ 118 Fringed frog, Madagascar ......................................... 266 Frog, Ambatoharanana Climbing ................... 252 —, Barbour’s Climbing ................................................. 252 —, Beige Bridge ..................................................................... 272 —, Betsileo Madagascar .............................................. 276 —, Betsileo Reed .................................................................. 248 —, Bott’s Bright-eyed ...................................................... 258 —, Boulenger’s Bridge ................................................... 270 —, Central Bright-eyed ................................................ 258 —, Central Madagascar ................................................ 278 —, Charlotte’s Madagascar ...................................... 278 —, Climbing Tree-hole .................................................... 252 —, Duméril’s Bright-eyed .......................................... 262 —, Duméril’s Madagascar ......................................... 280 —, Engraved Bridge ......................................................... 272 —, Fiery Bright-eyed ....................................................... 260 —, Folohy Madagascar .................................................. 280 —, Fort Madagascar ......................................................... 276 —, Goudot’s Bright-eyed ............................................ 264 —, Grandidier’s Madagascar ................................. 274 —, Green Bright-eyed .................................................... 260 —, Ivohimanitra Madagascar ............................... 274 —, Madagascar Bright-eyed ................................... 262 —, Madagascar fringed ................................................ 266 —, Madagascar jumping ............................................. 268 —, Mahanoro Tree-hole ............................................... 252 —, Marbled Rain .................................................................. 250 —, Mascarene Ridged ..................................................... 248 —, Massif Bridge .................................................................. 270 —, Mocquard’s Madagascar .................................... 276 —, Pandanus ............................................................................. 266 —, Pointy ....................................................................................... 266 —, Red-eyed Bright-eyed ........................................... 260 —, Rough Bridge .................................................................. 272 —, Sambava Tomato ........................................................... 250 —, Starry-night Reed ...................................................... 248 —, Stump-toed ....................................................................... 250 —, Tomato .................................................................................... 250 —, Warty Madagascar .................................................... 276 —, Web-foot ............................................................................... 250 —, Webb’s Bridge ................................................................. 270 —, Western Bright-eyed .............................................. 262 —, White-lipped Bright-eyed ............................... 264 —, White-whiskered Madagascar .................... 278 Fruit Bat, Madagascar Straw-coloured ........ 70 Fulda coroller .............................................................................. 314

336

Furcifer antimena ................................................................... 204 — balteatus ................................................................................... 198 — labordi ........................................................................................ 206 — lateralis ...................................................................................... 192 — nicosiai ....................................................................................... 190 — oustaleti ..................................................................................... 190 — pardalis ...................................................................................... 192 — petteri .......................................................................................... 202 — rhinoceratus .......................................................................... 204 — verrucosus ............................................................................... 206 — willsii ............................................................................................ 194 G Gactornis enarratus ............................................................. 134 Galidia elegans ............................................................................. 66 Gallinula chloropus ............................................................... 104 Garter Snake, Forest Madagascar .................... 244 —, Striped Madagascar ................................................ 242 —, Trail Madagascar ........................................................ 244 Gecko, Aldabra Day ......................................................... 222 —, Ambonihazo Madagascar Velvet ................ 233 —, Antongil Madagascar Velvet ............................ 232 —, Barbour’s Day ................................................................ 224 —, Bora’s Day ............................................................................. 224 —, Common House ............................................................ 229 —, Common Leaf-tailed ............................................. 236 —, Dwarf ....................................................................................... 226 —, Fish-scale ............................................................................. 228 —, Flathead House ............................................................... 229 —, Giant Day ............................................................................ 220 —, Giant Leaf-tailed ........................................................ 236 —, Gold-dust Day ............................................................... 222 —, Gould’s Day ........................................................................ 224 —, Graceful Big-headed .............................................. 230 —, Grandidier’s Fish-scale ........................................... 228 —, Grandidier’s Dwarf ..................................................... 227 —, Grey’s House ...................................................................... 229 —, Günther’s Leaf-tailed ............................................ 234 —, Henkel’s Leaf-tailed ................................................. 236 —, House ....................................................................................... 228 —, Koch’s Day .......................................................................... 220 —, Lined Day ............................................................................ 220 —, Lined Leaf–tailed ....................................................... 234 —, Madagascar Clawless ............................................. 228 —, Madagascar Day ............................................................ 220 —, Madagascar Dwarf ...................................................... 227 —, Madagascar Ground ............................................... 230 —, Madagascar velvet ..................................................... 232 —, Many-scaled Fish-scale .......................................... 228 —, Mocquard’s Ground ................................................ 230 —, Modest Day ....................................................................... 224 —, Mossy Leaf-tailed ....................................................... 236

Gecko, Northern Madagascar Velvet .............. 232 —, Peacock Day ..................................................................... 220 —, Peters’ Spotted Fish-scale .................................... 229 —, Rainforest Dwarf .......................................................... 227 —, Robust Dwarf ................................................................... 227 —, Sakalava Madagascar Velvet ............................. 233 —, Satanic Leaf-tailed .................................................... 234 —, Small Lined Day .......................................................... 222 —, Spearpoint leaf-tailed ........................................... 234 —, Speckled Day ................................................................... 222 —, Spiny Forest Dwarf ..................................................... 226 —, Standing’s Day ............................................................... 224 —, Thicktail Day ................................................................... 224 Geckolepis ....................................................................................... 228 — maculata .................................................................................. 229 — polylepis .................................................................................... 228 — typica ........................................................................................... 228 Geobiastes squamiger ......................................................... 144 Gephyromantis asper .......................................................... 272 — boulengeri ............................................................................... 270 — luteus ........................................................................................... 272 — pseudoasper .......................................................................... 270 — sculpturatus ........................................................................... 272 — webbi ............................................................................................ 270 Giant mouse lemur ............................................................... 36 Giant Treefrog, Boulenger’s .................................... 252 Glareola ocularis ...................................................................... 114 Goshawk, Henst’s ................................................................... 100 Graphium cyrnus .................................................................... 290 — evombar .................................................................................... 288 Grass yellow ............................................................................... 292 Grass Yellow, Angled ......................................................... 293 —, Broad-bordered ............................................................. 293 —, Malagasy ................................................................................ 293 —, Marsh ........................................................................................ 292 Grebe, Little .................................................................................. 84 —, Madagascar ............................................................................ 84 Green Pigeon, African .................................................... 122 —, Madagascar ....................................................................... 122 Greenbul, Grey-crowned ............................................. 162 —, Long-billed (see Long-billed Tetraka) .................................... 162 —, Spectacled (see Spectacled Tetraka) ....................................... 162 Greenshank, Common .................................................. 112 Ground Boa, Duméril’s ................................................ 238 —, Madagascar ....................................................................... 238 Ground-roller, Long-tailed ...................................... 146 —, Pitta-like ............................................................................... 142 —, Rufous-headed .............................................................. 142 —, Scaly ........................................................................................... 144 —, Short-legged ..................................................................... 144

Guibemantis ................................................................................. 266 — liber ............................................................................................... 267 — pulcher ........................................................................................ 267 — variabilis .................................................................................. 266 Guineafowl, Helmeted ..................................................... 86 H Hairstreak ..................................................................................... 296 Haliaeetus vociferoides ......................................................... 98 Hamerkop ....................................................................................... 96 Hapalemur aureus .................................................................... 42 — griseus ............................................................................................ 42 — occidentalis ............................................................................... 42 Harrier-hawk, Madagascar ......................................... 98 Hedgehog Tenrec, Greater ........................................... 74 —, Lesser ........................................................................................... 74 Hemicentetes nigriceps ......................................................... 76 — semispinosus ............................................................................ 76 Hemidactylus .............................................................................. 228 — frenatus ..................................................................................... 229 — mercatorius ........................................................................... 229 — platycephalus ....................................................................... 229 Hemiolaus cobaltina ............................................................ 296 Heron, Black ................................................................................. 90 —, Black-crowned Night ............................................... 96 —, Grey ............................................................................................... 88 —, Humblot’s ............................................................................... 88 —, Madagascar .......................................................................... 88 —, Madagascar Pond ............................................................ 94 —, Purple ......................................................................................... 90 —, Squacco ..................................................................................... 94 —, Striated ...................................................................................... 94 Hesperiidae ................................................................................. 314 Heterixalus alboguttatus ................................................. 248 — betsileo ........................................................................................ 248 Heteropsis ........................................................................................ 312 — difficilis ...................................................................................... 313 — narcissus ................................................................................... 313 — subsimilis ................................................................................. 313 — turbans ....................................................................................... 312 — vola ................................................................................................ 312 Hipposideros commersoni ................................................. 72 Hognose Snake, Blonde ............................................... 240 —, Giant ......................................................................................... 240 Hoopoe, Madagascar ...................................................... 140 Hoplobatrachus tigerinus ................................... 246, 248 House gecko ............................................................................... 228 Hyphaene ........................................................................................ 136 Hypogeomys antimena ......................................................... 78 Hypolimnas dexithea .......................................................... 308 — misippus .................................................................................... 310 Hypositta corallirostris ...................................................... 170 Hypsipetes madagascariensis ...................................... 152 337

I Ibis, Madagascar Crested ........................................... 102 Indri ....................................................................................................... 62 Indri indri .......................................................................................... 62 Insect, Flatid Leaf ............................................................... 284 Ithycyphus ...................................................................................... 240 J Jacana, Madagascar .......................................................... 104 Jery, Common .......................................................................... 160 —, Green ......................................................................................... 162 —, Stripe-throated ............................................................. 160 Joker, Common ...................................................................... 300 Jumping frog, Madagascar ....................................... 268 Junonia goudotii ....................................................................... 304 — hierta ........................................................................................... 306 — oenone ........................................................................................ 306 — orithya ........................................................................................ 306 — rhadama ................................................................................... 306 K Kestrel, American .................................................................. 100 —, Banded ..................................................................................... 100 —, Eurasian .................................................................................. 100 —, Madagascar ....................................................................... 100 Kingfisher, Madagascar Malachite ................. 138 —, Madagascar Pygmy .................................................. 138 Kite, Black (Yellow-billed) ........................................ 100 L Lady, Green ................................................................................. 290 Laliostoma labrosum ........................................................... 268 Langaha ............................................................................................ 240 — madagascariensis ............................................................ 241 Lark, Madagascar ................................................................ 150 Leaf Insect, Flatid ............................................................... 284 Leaf-nosed Bat, Commerson’s ................................ 72 Leioheterodon madagascariensis ............................ 240 — modestus .................................................................................. 240 Lemur catta ...................................................................................... 44 Lemur, Anjiahely Mouse ................................................. 33 —, Ankarana Sportive ......................................................... 39 —, Bemaraha Sportive ........................................................ 38 —, Bemaraha Woolly ........................................................... 54 —, Black ............................................................................................. 50 —, Black-and-white Ruffed ........................................ 52 —, Brown ......................................................................................... 48 —, Common Brown ............................................................ 48 —, Coquerel’s Giant Mouse ........................................... 37 —, Crossley’s Dwarf ............................................................... 34 —, Crowned .................................................................................. 50 —, Dwarf .......................................................................................... 34 —, Eastern Woolly ................................................................... 55 —, Fat-tailed Dwarf ............................................................... 35 338

Lemur, Fork-marked .......................................................... 36 —, Furry-eared Dwarf ........................................................ 34 —, Giant mouse ........................................................................ 36 —, Golden Bamboo ............................................................. 42 —, Golden-brown Mouse ................................................ 33 —, Goodman’s Mouse .......................................................... 32 —, Greater Bamboo ............................................................. 40 —, Greater Dwarf ..................................................................... 35 —, Grey Bamboo ..................................................................... 42 —, Grey Mouse ........................................................................... 33 —, Grey-brown Mouse ....................................................... 33 —, Madame Berthe’s Mouse ......................................... 32 —, Masoala Fork-marked ................................................ 36 —, Masoala Sportive ............................................................. 38 —, Masoala Woolly ................................................................ 55 —, Milne-Edwards’ Sportive ........................................ 39 —, Mittermeier’s Mouse .................................................... 32 —, Mongoose ............................................................................... 50 —, Montagne d’Ambre Mouse .................................... 32 —, Montagne d’Ambre Dwarf ..................................... 35 —, Montagne d’Ambre Fork-marked .................. 37 —, Mouse .......................................................................................... 32 —, Northern Bamboo .......................................................... 42 —, Northern Giant Mouse ............................................. 37 —, Nosy Be Mouse .................................................................. 32 —, Nosy Be Sportive ............................................................. 39 —, Pale Fork-marked ........................................................... 37 —, Peters’ Mouse ....................................................................... 32 —, Petter’s Sportive ................................................................. 38 —, Peyriéras’ Woolly ............................................................. 54 —, Red Ruffed ............................................................................. 52 —, Red-bellied ........................................................................... 48 —, Red-collared Brown ..................................................... 48 —, Red-fronted Brown .................................................... 48 —, Red-tailed Sportive ....................................................... 39 —, Ring-tailed ............................................................................ 44 —, Rufous Brown ..................................................................... 48 —, Rufous Mouse ..................................................................... 33 —, Sambirano Fork-marked ......................................... 36 —, Sanford’s Brown ............................................................. 46 —, Seal’s Sportive ...................................................................... 38 —, Sibree’s Dwarf ...................................................................... 34 —, Small-toothed Sportive ............................................. 39 —, Southern Woolly .............................................................. 54 —, Sportive .................................................................................... 38 —, Tavaratra Mouse ............................................................... 33 —, Weasel Sportive ................................................................. 38 —, Western Woolly ................................................................. 55 —, White-footed Sportive ............................................... 39 —, White-fronted Brown .............................................. 46 —, Woolly ........................................................................................ 54 —, Zombitse Sportive .......................................................... 39

Leopard, Common ............................................................ 300 —, Madagascar ....................................................................... 300 Lepilemur ........................................................................................... 38 — ankaranensis ........................................................................... 39 — edwardsi ...................................................................................... 39 — hubbardorum ......................................................................... 39 Lepilemur leucopus .................................................................. 39 — microdon ..................................................................................... 39 — mustelinus ................................................................................. 38 — petteri ............................................................................................. 38 — randrianasoloi ....................................................................... 38 — ruficaudatus ............................................................................ 39 — scottorum ................................................................................... 38 — seali ................................................................................................... 38 — tymerlachsonorum ........................................................... 39 Leptopterus chabert .............................................................. 166 Leptosia alcesta ......................................................................... 294 Leptosomus discolor ............................................................. 146 Leptotes pirithous .................................................................... 297 Liophidium rhodogaster ................................................... 244 — torquatum ............................................................................... 244 Lizard, Bronze Plated ....................................................... 214 —, Brygoo’s Plated ............................................................. 212 —, Green plated ...................................................................... 212 —, Karsten’s Plated ............................................................ 214 —, Madagascar Keeled Plated .............................. 214 —, Madagascar Plated ................................................... 214 —, Meier’s Plated ................................................................... 214 —, Ornate Plated .................................................................. 214 —, Red-legged Plated ..................................................... 212 —, Three-eyed .......................................................................... 208 —, Three-lined Plated ....................................................... 214 —, Tsingy Plated ................................................................... 212 —, Western Plated .............................................................. 214 Lonchura nana .......................................................................... 152 Lophotibis cristata ................................................................. 102 Lovebird, Grey-headed ................................................. 124 Lycodryas ........................................................................................ 240 — gaimardi ................................................................................... 241 — granuliceps ............................................................................. 241 Lygodactylus ................................................................................. 226 — madagascariensis ............................................................ 227 — miops ........................................................................................... 227 — pictus ............................................................................................ 227 — tolampyae ................................................................................ 227 — tuberosus .................................................................................. 226 M Mabuya ............................................................................................. 216 Macrotarsomys bastardi ..................................................... 78 Madagascarophis .................................................................... 242 — colubrinus ............................................................................... 243 Madascincus melanopleura .......................................... 218

Magpie-robin, Madagascar ...................................... 154 Mannikin, Madagascar ................................................. 152 Mantella aurantiaca ............................................................ 254 — baroni .......................................................................................... 254 — betsileo ........................................................................................ 254 — ebenaui ...................................................................................... 254 Mantella expectata ................................................................ 256 — laevigata ................................................................................... 256 — madagascariensis ............................................................ 254 — nigricans ................................................................................... 256 — viridis .......................................................................................... 256 Mantella, Baron’s ................................................................. 254 —, Betsileo .................................................................................. 254 —, Blue-legged ........................................................................ 256 —, Climbing .............................................................................. 256 —, Golden .................................................................................... 254 —, Green ........................................................................................ 256 —, Guibé’s ..................................................................................... 256 —, Madagascar ......................................................................... 254 —, Orange-backed ............................................................. 254 Mantidactylus argenteus ................................................. 280 — betsileanus .............................................................................. 276 — charlotteae .............................................................................. 278 — femoralis ................................................................................... 276 — grandidieri ............................................................................. 274 — guttulatus ................................................................................ 274 — lugubris ...................................................................................... 280 — majori ......................................................................................... 274 — melanopleura ....................................................................... 278 — mocquardi .............................................................................. 276 — opiparis ...................................................................................... 278 — ulcerosus ................................................................................... 276 Map Butterfly, African .................................................. 308 Margaroperdix madagarensis ....................................... 86 Martin, Brown-throated ............................................... 136 —, Mascarene ........................................................................... 136 Megaptera novaeangliae ..................................................... 80 Melanitis leda ............................................................................. 312 Merops superciliosus ............................................................ 138 Mesite, Brown ........................................................................... 102 —, Subdesert ............................................................................... 102 —, White-breasted ............................................................. 102 Mesitornis unicolor ............................................................... 102 — variegata .................................................................................. 102 Metalmark, Madagascar ............................................. 296 Microcebus ........................................................................................ 32 — arnholdi ....................................................................................... 32 — berthae .......................................................................................... 32 — griseorufus ................................................................................. 33 — lehilahytsara ............................................................................ 32 — macarthurii .............................................................................. 33 — mamiratra ................................................................................. 32 339

Microcebus mittermeieri .................................................... 32 — murinus ........................................................................................ 33 — myoxinus .................................................................................... 32 — ravelobensis .............................................................................. 33 — rufus ................................................................................................ 33 — tavaratra ..................................................................................... 33 Microgale ............................................................................................ 74 Microsphaerotherium ......................................................... 282 Migrant ............................................................................................ 292 Millipede, Giant pill ........................................................ 282 Milvus migrans .......................................................................... 100 Mimophis mahfalensis ....................................................... 242 Mirafra hova ................................................................................ 150 Mirza coquereli ............................................................................ 37 — zaza .................................................................................................. 37 Monarch, African ................................................................ 298 Mongoose, Narrow-striped (see Northern Bokiboky) .................................... 66 —, Ring-tailed (see Ring-tailed Vontsira) .................................. 66 Monias benschi .......................................................................... 102 Monticola bensoni .................................................................. 155 — erythronotus ......................................................................... 155 — imerinus .................................................................................... 155 — sharpei ........................................................................................ 154 Moorhen, Common .......................................................... 104 Motacilla flaviventris .......................................................... 150 Moth, Comet ............................................................................. 314 —, Madagascar Sunset .................................................. 314 Mother-of-Pearl, Clouded ........................................ 308 Mouse lemur ................................................................................ 32 Mouse, House ...................................................................... 31, 79 —, Western Big-footed ....................................................... 78 Mungotictis decemlineata ................................................. 66 Mus musculus ....................................................................... 31, 79 Musk Shrew, Asian .................................................................. 31 Mylothris phileris .................................................................... 294 Myna, Common .................................................................... 176 Mystacornis crossleyi ........................................................... 170 N Neodrepanis coruscans ...................................................... 148 Neomixis striatigula ............................................................. 160 — tenella .......................................................................................... 160 — viridis .......................................................................................... 162 Nephila .............................................................................................. 284 Neptis kikideli ............................................................................. 310 — saclava ........................................................................................ 310 Nesillas lantzii ............................................................................ 156 — typica ........................................................................................... 156 Nesomys ............................................................................................... 78 — audeberti ..................................................................................... 79 340

Nesomys lambertoni ................................................................ 79 — rufus ................................................................................................ 79 Newtonia amphichroa ....................................................... 167 — brunneicauda ...................................................................... 166 Newtonia, Common ......................................................... 166 —, Dark ............................................................................................ 167 Night Heron, Black-crowned ................................... 96 Nightjar, Collared ............................................................... 134 —, Madagascar ....................................................................... 134 Ninox superciliaris ................................................................. 132 Numenius phaeopus ............................................................. 112 Numida meleagris ..................................................................... 86 Nycticorax nycticorax ............................................................ 96 Nymph, Madagascar Forest .................................... 298 —, Tree ............................................................................................. 302 O Oena capensis ............................................................................. 120 Oplurus cuvieri ......................................................................... 208 — cyclurus ..................................................................................... 208 — grandidieri ............................................................................. 210 — quadrimaculatus ............................................................. 210 — saxicola ...................................................................................... 210 Orange Tip, Madagascar ............................................ 292 Otomops madagascariensis ............................................. 72 Otus rutilus ................................................................................... 132 Owl, Barn ...................................................................................... 132 —, Madagascar Scops ..................................................... 132 —, Malagasy Scops (see Madagascar Scops Owl) ....................... 132 —, Rainforest Scops (see Madagascar Scops Owl) ....................... 132 —, Torotoroka Scops (see Madagascar Scops Owl) ....................... 132 —, White-browed ................................................................ 132 Oxylabes madagascariensis .......................................... 162 Oxylabes, White-throated ......................................... 162 —, Yellow-browed ................................................................. 162 P Pachypodium ........................................................................... 318 Pachypodium ............................................................................... 318 — rosulatum ................................................................................ 319 Palleon nasus ............................................................................... 188 Palm Swift, African ........................................................... 136 Palm, Bismarck ........................................................................ 136 —, Coconut .................................................................................. 136 —, Raphia ......................................................................................... 70 —, Screw ......................................................................................... 316 —, Traveller’s ............................................................................ 316 Pandanus ........................................................................................ 316 Pandanus ....................................................................................... 316 Pandanusicola ............................................................................ 266

Pansy, Blue ................................................................................... 306 —, Eyed ............................................................................................ 306 —, Madagascar Brown .................................................. 304 —, Yellow ....................................................................................... 306 Papilio dardanus ..................................................................... 286 — delalandei ................................................................................ 286 — demodocus ............................................................................. 286 — epiphorbas .............................................................................. 288 — oribazus .................................................................................... 288 Paradise Flycatcher, Madagascar ...................... 152 Paradoxophyla palmata ................................................... 250 Parastenophis .............................................................................. 240 — betsileanus .............................................................................. 241 Paratriaenops ................................................................................. 72 Pardopsis punctatissima .................................................. 302 Parnara naso ............................................................................... 314 Paroedura bastardi ................................................................ 230 — gracilis ........................................................................................ 230 — picta .............................................................................................. 230 Parrot, Greater Vasa ......................................................... 124 —, Lesser Vasa ......................................................................... 124 Partridge, Madagascar ..................................................... 86 Passer domesticus ...................................................................... 82 Pelomedusa subrufa ............................................................. 183 Pelusios castanoides .............................................................. 182 — subniger ..................................................................................... 183 Phaethon lepturus .................................................................. 118 — rubricauda ............................................................................. 118 Phalanta madagascariensis .......................................... 300 — phalantha ................................................................................ 300 Phaner ................................................................................................... 36 — electromontis ........................................................................... 37 — furcifer ........................................................................................... 36 — pallescens .................................................................................... 37 — parienti ......................................................................................... 36 Pharmacophagus antenor .............................................. 290 Phedina borbonica ................................................................. 136 Phelsuma abbotti .................................................................... 222 — barbouri .................................................................................... 224 — borai ............................................................................................. 224 — dorsivittata ............................................................................ 220 — gouldi ........................................................................................... 224 — grandis ........................................................................................ 220 — guttata ........................................................................................ 222 — kochi ............................................................................................. 220 — laticauda .................................................................................. 222 — lineata ......................................................................................... 220 — madagascariensis ............................................................ 220 — modesta ..................................................................................... 224 — mutabilis .................................................................................. 224 — pusilla .......................................................................................... 222 — quadriocellata ..................................................................... 220 — standingi ................................................................................... 224

Philepitta castanea ................................................................ 148 — schlegeli ...................................................................................... 148 Phisalixella .................................................................................... 240 Phromnia rosea ......................................................................... 284 Pig, Bush ................................................................................... 31, 68 Pigeon, African Green .................................................... 122 —, Feral ........................................................................................... 120 —, Madagascar Blue ........................................................ 122 Pigeon, Madagascar Green ...................................... 122 Pill millipede, giant .......................................................... 282 Pine, Screw .................................................................................. 316 Platypelis barbouri ................................................................ 252 — grandis ........................................................................................ 252 — tuberifera ................................................................................. 252 Playboy, Brown ........................................................................ 296 Plethodontohyla mihanika ............................................ 252 — notosticta ................................................................................. 252 Ploceus nelicourvi ................................................................... 178 — sakalava .................................................................................... 178 Plover, Black-bellied ........................................................ 108 —, Crab- ......................................................................................... 114 —, Greater Sand .................................................................... 108 —, Grey ............................................................................................ 108 —, Lesser Sand ......................................................................... 108 —, (Madagascar) Three-banded ........................ 106 —, White-fronted ................................................................ 106 Pluvialis squatarola .............................................................. 108 Polka Dot ...................................................................................... 302 Polyboroides radiatus ............................................................. 98 Pond Heron, Madagascar ................................................ 94 Potamochoerus larvatus ............................................ 31, 68 Pratincole, Madagascar ................................................ 114 Precis andremiaja ................................................................... 304 Prolemur simus ............................................................................ 40 Propithecus candidus ............................................................. 58 — coquereli ...................................................................................... 56 — coronatus .................................................................................... 60 — deckenii ......................................................................................... 56 — diadema ....................................................................................... 58 — edwardsi ...................................................................................... 58 — perrieri .......................................................................................... 60 — tattersalli ..................................................................................... 60 — verreauxi ..................................................................................... 56 Protogoniomorpha anacardii ..................................... 308 Pseudacraea lucretia ............................................................ 310 — imerina ...................................................................................... 310 Pseudobias wardi .................................................................... 170 Pteropus rufus ............................................................................... 70 Ptychadena mascareniensis .......................................... 248 Pyxis arachnoides ................................................................... 182 — planicauda ............................................................................. 183

341

R Rail, White-throated ....................................................... 104 Rain Frog, Marbled ........................................................... 250 Ramy ...................................................................................................... 64 Randia pseudozosterops ................................................... 160 Raphia ................................................................................................... 70 Rat, Black ................................................................................. 31, 79 —, Brown ................................................................................. 31, 79 —, Eastern Red Forest ......................................................... 79 Rat, Forest ....................................................................................... 78 —, Giant Jumping .................................................................. 78 —, Lowland Red Forest ...................................................... 79 —, Norway .............................................................................. 31, 79 —, red forest ................................................................................. 78 —, tuft-tailed ............................................................................... 78 —, Western Red Forest ....................................................... 79 —, Western Tuft-tailed ....................................................... 78 Rattus norvegicus ............................................................. 31, 79 — rattus ...................................................................................... 31, 79 Ravenala madagascariensis ......................................... 316 Reed Frog, Betsileo ........................................................... 248 —, Starry-night ...................................................................... 248 Riparia paludicola ................................................................. 136 Robin, Madagascar Magpie- .................................. 154 Rock Thrush, Amber Mountain ........................... 155 —, Benson’s ................................................................................. 155 —, Forest ........................................................................................ 154 —, Littoral ..................................................................................... 154 Roller, Broad-billed .......................................................... 140 —, Cuckoo- ................................................................................. 146 —, Long-tailed Ground- ............................................. 146 —, Pitta-like Ground- .................................................... 142 —, Rufous-headed Ground- ................................... 142 —, Scaly Ground- ................................................................ 144 —, Short-legged Ground ............................................ 144 S Sailer, Madagascar ............................................................. 310 —, Spotted .................................................................................... 310 Salamis anteva ........................................................................... 298 Sand Plover, Greater ........................................................ 108 —, Lesser ......................................................................................... 108 Sand Snake, Mahafaly .................................................... 242 Sanderling .................................................................................... 110 Sandpiper, Common ....................................................... 110 —, Curlew ..................................................................................... 110 Sanzinia madagascariensis ........................................... 239 — volontany ................................................................................. 239 Saribia ................................................................................................ 296 — perroti ......................................................................................... 297 — tepahi ........................................................................................... 297 Saroglossa aurata .................................................................... 176 342

Satyr, Indian Ocean .......................................................... 312 —, Madagascar ....................................................................... 312 Saxicola torquatus ................................................................. 154 Scaphiophryne marmorata ........................................... 250 Schetba rufa .................................................................................. 168 Scops Owl, Madagascar ............................................... 132 —, Malagasy (see Madagascar Scops Owl) ....................... 132 —, Rainforest (see Madagascar Scops Owl) ....................... 132 Scops Owl, Torotoroka (see Madagascar Scops Owl) ....................... 132 Scopus umbretta .......................................................................... 96 Sea Turtle, Green ................................................................. 184 —, Hawksbill ............................................................................. 184 Secretive skink ........................................................................ 218 Setifer setosus ................................................................................. 74 Sevenia ............................................................................................... 302 — amazoula ................................................................................. 303 — howensis ................................................................................... 302 — madagascariensis ............................................................ 302 Shrew, Asian Musk .................................................................. 31 Sifaka, Coquerel’s ................................................................... 56 —, Crowned .................................................................................. 60 —, Decken’s .................................................................................... 56 —, Diademed ............................................................................... 58 —, Golden-crowned ............................................................ 60 —, Milne-Edwards’ ............................................................... 58 —, Perrier’s ..................................................................................... 60 —, Silky ............................................................................................... 58 —, Tattersall’s .............................................................................. 60 —, Verreaux’s ............................................................................... 56 Skink, Common Madagascar .................................. 218 —, Comoro .................................................................................. 218 —, Elegant .................................................................................... 216 —, Gold-spotted ................................................................... 218 —, Gravenhorst’s .................................................................. 216 —, secretive ................................................................................. 218 —, Tandrefana ......................................................................... 216 Skipper ............................................................................................. 314 Snail, Giant African Land ......................................... 282 Snake, Bernier’s Striped .............................................. 242 —, Blonde Hognose .......................................................... 240 —, Collared Bright ............................................................ 244 —, Forest Madagascar Garter ............................... 244 —, Four-striped ..................................................................... 242 —, Giant Hognose .............................................................. 240 —, Gluttonous Bighead ................................................ 244 —, Gold-collared .................................................................. 244 —, leaf-nosed ............................................................................ 240 —, Madagascar cat-eyed ............................................. 242

Snake, Madagascar tree ................................................ 240 —, Mahafaly Sand ............................................................... 242 —, Striped Madagascar Garter ............................ 242 —, Trail Madagascar Garter ................................... 244 Sparrow, House ........................................................................... 82 Sparrowhawk, Frances’s ............................................... 100 —, Madagascar ......................................................................... 100 Sphaeromimus ........................................................................... 282 Spider, Golden orb-web ............................................... 284 Spinomantis .................................................................................. 266 — aglavei ......................................................................................... 267 Sportive lemur ........................................................................... 38 Starling, Madagascar ...................................................... 176 Stenella longirostris .................................................................. 80 Sterna dougallii ......................................................................... 116 Stonechat, African ............................................................. 154 Strabena ........................................................................................... 312 — tamatavae ............................................................................... 313 — triophthalma ........................................................................ 313 Streaked Tenrec, Highland ........................................... 76 —, Lowland .................................................................................... 76 Streptopelia picturata .......................................................... 120 Stumpffia ......................................................................................... 250 — pygmaea ................................................................................... 251 — tridactyla ................................................................................. 250 Sunbird Asity, Common ............................................. 148 Sunbird, Madagascar Green ................................... 164 —, Souimanga ......................................................................... 164 Suncus murinus ........................................................................... 31 Swallowtail, Banded Blue .......................................... 288 —, Citrus ....................................................................................... 286 —, Cream-lined ..................................................................... 286 —, Madagascar Giant ..................................................... 290 —, Mocker .................................................................................... 286 —, Spotted Blue ..................................................................... 288 Swamp Warbler, Madagascar ................................ 158 Swift, African Black [bird] ........................................ 136 —, African Palm [bird] ................................................. 136 —, Cuvier’s Madagascar [reptile] ...................... 208 —, Dumeril’s Madagascar [reptile] ................. 210 —, Grandidier’s Madagascar [reptile] ......... 210 —, Madagascar [reptile] ............................................... 208 —, Madagascar Black [bird] .................................... 136 —, Marked Madagascar [reptile] .......................... 210 —, Merrem’s Madagascar [reptile] ................... 208 Swordtail, Madagascar .................................................. 288 T Tachybaptus pelzelnii ............................................................. 84 — ruficollis ....................................................................................... 84 Taphozous mauritianus ....................................................... 72 Teal, Red-billed ......................................................................... 84

Tenrec ecaudatus ........................................................................ 76 Tenrec, Common .................................................................... 76 —, Greater Hedgehog ........................................................ 74 —, Highland Streaked ......................................................... 76 —, Lesser Hedgehog ............................................................ 74 —, Lowland Streaked ......................................................... 76 Tern, Greater Crested ..................................................... 116 Tern, Lesser Crested ......................................................... 116 —, Roseate ................................................................................... 116 —, Swift ........................................................................................... 116 Terpsiphone mutata .............................................................. 152 Terrapin .......................................................................................... 182 —, Marsh ........................................................................................ 183 Tetraka, Grey-crowned .................................................. 162 —, Long-billed ........................................................................ 162 —, Spectacled ........................................................................... 162 Thalasseus bengalensis ...................................................... 116 — bergii ............................................................................................ 116 Thamnosophis epistibes .................................................... 244 — infrasignatus ........................................................................ 244 — lateralis ...................................................................................... 242 Thrush, Amber Mountain Rock ........................... 155 —, Benson’s Rock ................................................................... 155 —, Forest Rock ........................................................................ 154 —, Littoral Rock ...................................................................... 154 Tip, Madagascar Orange ............................................ 292 Toad, Asian ................................................................................... 246 Tomb Bat, Mauritian .......................................................... 72 Tortoise, Flat-tailed ............................................................ 183 —, Radiated ................................................................................ 182 —, Spider ....................................................................................... 182 Trachelophorus giraffa ....................................................... 284 Tracheloptychus madagascariensis ....................... 214 Trachylepis aureopunctata ............................................ 218 — comorensis .............................................................................. 218 — elegans ........................................................................................ 216 — gravenhorstii ........................................................................ 216 — tandrefana .............................................................................. 216 — vato ................................................................................................ 219 Tree boa, Madagascar .................................................... 238 Tree nymph ................................................................................. 302 Tree-hole Frog, Climbing ............................................ 252 —, Mahanoro ............................................................................ 252 Tree, Octopus ........................................................................... 318 Treefrog, Boulenger’s Giant .................................... 252 Treron australis ......................................................................... 122 — calvus ........................................................................................... 122 Triaenops ............................................................................................ 72 Tringa stagnatilis ..................................................................... 112 Tropicbird, Red-tailed ................................................... 118 —, White-tailed ..................................................................... 118 Turnix nigricollis ........................................................................ 86 343

Turnstone, Ruddy ............................................................... 112 Tursiops aduncus ....................................................................... 80 — truncatus ..................................................................................... 80 Turtle Dove, Madagascar ........................................... 120 Turtle, East African Black Mud ............................ 183 —, Green Sea ............................................................................. 184 —, Hawksbill Sea ................................................................. 184 —, Madagascan Big-headed ...................................... 183 —, Yellowbelly Mud ............................................................ 182 Tylas eduardi ............................................................................... 174 Tyto alba .......................................................................................... 132 U Upupa marginata ................................................................... 140 Uratelornis chimaera ........................................................... 146 Uroplatus ebenaui .................................................................. 234 — fimbriatus ............................................................................... 236 — finiavana .................................................................................. 234 — giganteus .................................................................................. 236 — guentheri .................................................................................. 234 — henkeli ........................................................................................ 236 — lineatus ...................................................................................... 234 — phantasticus .......................................................................... 234 — sameiti ........................................................................................ 236 — sikorae ........................................................................................ 236 V Vanga curvirostris .................................................................. 172 Vanga, Blue ................................................................................. 168 —, Chabert .................................................................................. 166 —, Crossley’s ............................................................................. 170 —, Helmet .................................................................................... 172 —, Hook-billed ....................................................................... 172 —, Lafresnaye’s ......................................................................... 174 —, Nuthatch ............................................................................... 170 —, Pollen’s .................................................................................... 174 —, Red-tailed ........................................................................... 166 —, Rufous ..................................................................................... 168 —, Sickle-billed ...................................................................... 172 —, Tylas ........................................................................................... 174 —, Van Dam’s ............................................................................. 174 —, Ward’s ...................................................................................... 170 —, White-headed ................................................................. 168 Varecia rubra ................................................................................. 52 — variegata ..................................................................................... 52 Vasa Parrot, Greater ......................................................... 124 —, Lesser ........................................................................................ 124 Velvet gecko, Madagascar .......................................... 232 Viverricula indica ............................................................. 31, 66 Voeltzkowia rubrocaudata ............................................ 218 Vontsira, Ring-tailed .......................................................... 66

344

W Wagtail, Madagascar ....................................................... 150 —, Madagascar Brush .................................................... 156 —, Madagascar Swamp ................................................. 158 —, Rand’s ....................................................................................... 160 —, Subdesert Brush .......................................................... 156 Weaver, Nelicourvi ............................................................. 178 —, Sakalava ................................................................................. 178 Weevil, Giraffe-necked ................................................. 284 Whale, Humpback ................................................................ 80 Whimbrel ...................................................................................... 112 Whistling-duck, Fulvous ................................................ 84 —, White-faced ......................................................................... 84 White-eye, Madagascar ................................................ 164 White, African Wood ..................................................... 294 —, Albatross ................................................................................ 294 —, Grandidier’s Caper .................................................. 294 —, Yellow-painted Caper ........................................... 294 Wood White, African ..................................................... 294 Woolly lemur ............................................................................... 54 X Xanthormixis zosterops .................................................... 162 Xenopirostris damii .............................................................. 174 — polleni ......................................................................................... 174 — xenopirostris ......................................................................... 174 Y Yellow, Angled Grass ........................................................ 293 —, Broad-bordered Grass ............................................ 293 —, Grass ......................................................................................... 292 —, Malagasy Grass ............................................................... 293 —, Marsh Grass ....................................................................... 292 Yellowbrow, Madagascar ............................................... 162 Z Zizeeria knysna ......................................................................... 297 Zizina antanossa ..................................................................... 296 Zizula hylax .................................................................................. 297 Zonosaurus aeneus ............................................................... 214 — boettgeri .................................................................................... 212 — brygooi ....................................................................................... 212 — haraldmeieri ......................................................................... 214 — karsteni ...................................................................................... 214 — laticaudatus .......................................................................... 214 — madagascariensis ............................................................ 214 — maramaintso ....................................................................... 212 — ornatus ....................................................................................... 214 — rufipes ......................................................................................... 212 — subunicolor ............................................................................ 212 — trilineatus ................................................................................ 214 — tsingy ............................................................................................ 212 Zoosphaerium ............................................................................ 282 Zosterops maderaspatanus ............................................ 164