About the author Peter Schouten is an acclaimed wildlife artist who has a passion for all things feathered, furred and scaled – from both present and past He delights in painting creatures that either cannot be or have not been photographed, due to extinction or rarity He aims to draw attention to the unfortunate plight of many of these creatures and to emphasise the need for urgent conservation He recently completed work on Feathered Dinosaurs: The Origin of Birds – a collection of images which challenges all of our preconceived notions of those truly colossal animals of the past – the dinosaurs His spectacular paintings are keenly collected and have been widely exhibited at major galleries and museums around the world FullCOV_GlidingMammals_FNL copy.indd Stephen Jackson About the artist Illustrated by Peter Schouten Stephen Jackson is a behavioural and environmental ecologist who has studied Australian mammals in the wild and in captivity over the last 20 years He has worked in a number of different roles including field ecologist, zookeeper, curator, government regulator, part-time lecturer and wildlife consultant He has published numerous scientific articles and four books as a result of his research, with several other books nearing completion One of his books, Australian Mammals: Biology and Captive Management, was awarded the prestigious Whitley Medal for the best natural history book from the Royal Zoological Society of New South Wales Gliding Mammals of the World T he world’s gliding mammals are an extraordinary group of animals that have the ability to glide from tree to tree with seemingly effortless grace There are more than 60 species of gliding mammals including the flying squirrels from Asia, Europe and North America, the scaly-tailed flying squirrels from central Africa and the gliding possums of Australia and New Guinea But the most spectacular of all are the colugos – or so called flying lemurs – that occur throughout South-East Asia and the Philippines Animals that glide from tree to tree descend at an angle of less than 45 degrees to the horizontal, while those that parachute descend at an angle greater than 45 degrees Gliding is achieved by deflecting air flowing past well-developed gliding membranes, or patagia, which form an effective airfoil that allows the animal to travel the greatest possible horizontal distance with the least loss in height The flying squirrels and scaly-tailed flying squirrels even have special cartilaginous spurs that extend either from the wrist or elbow, respectively, to help support the gliding membrane Gliding Mammals of the World provides, for the first time, a synthesis of all that is known about the biology of these intriguing mammals It includes a brief description of each species, together with a distribution map and a beautiful full-colour painting An introduction outlines the origins and biogeography of each group of gliding mammals and examines the incredible adaptations that allow them to launch themselves and glide from tree to tree Stephen Jackson Illustrated by Peter Schouten Gliding Mammals of the World 7/06/12 11:23 AM Gliding Mammals of the World This page has been intentionally left blank Gliding Mammals of the World Stephen Jackson Illustrated by Peter Schouten © 2012 Text: Stephen Jackson; Illustrations: Peter Schouten All rights reserved Except under the conditions described in the Australian Copyright Act 1968 and subsequent amendments, 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, duplicating or otherwise, without the prior permission of the copyright owner Contact CSIRO PUBLISHING for all permission requests National Library of Australia Cataloguing-in-Publication entry: Jackson, Stephen Gliding mammals of the world / by Stephen Jackson ; illustrated by Peter Schouten 9780643092600 (hbk.) 9780643104051 (epdf) 9780643104068 (epub) Includes bibliographical references and index Gliders (Mammals) Mammals – Flight 599.369 Published by CSIRO PUBLISHING 150 Oxford Street (PO Box 1139) Collingwood VIC 3066 Australia Telephone: Local call: Fax: Email: Web site: +61 9662 7666 1300 788 000 (Australia only) +61 9662 7555 publishing.sales@csiro.au www.publish.csiro.au Front cover: Squirrel Glider Front flap: Feathertail Glider Back flap: Red-cheeked Flying Squirrel Back cover: Whiskered Flying Squirrel Original artworks are available from www.studioschouten.com.au Set in Perpetua 11.5/14 Cover design by Alicia Freile, Tango Media Text design by James Kelly Typeset by Oryx Publishing Pty Ltd Printed in China by 1010 Printing International Ltd CSIRO PUBLISHING publishes and distributes scientific, technical and health science books, magazines and journals from Australia to a worldwide audience and conducts these activities autonomously from the research activities of the Commonwealth Scientific and Industrial Research Organisation (CSIRO) The views expressed in this publication are those of the author(s) and not necessarily represent those of, and should not be attributed to, the publisher or CSIRO The copyright owner shall not be liable for technical or other errors or omissions contained herein The reader/user accepts all risks and responsibility for losses, damages, costs and other consequences resulting directly or indirectly from using this information Original print edition: The paper this book is printed on is in accordance with the rules of the Forest Stewardship Council® The FSC® promotes environmentally responsible, socially beneficial and economically viable management of the world’s forests Contents Preface Acknowledgements List of species Introduction vi viii x Gliding Marsupials 23 Colugos 47 Flying Squirrels 57 Scaly-tailed Flying Squirrels 163 Appendix 1: Subspecies 180 Appendix 2: Gliding mammal localities 182 Appendix 3: IUCN Red List Categories 189 Glossary 190 References 192 Index 214 v Preface This book explores the origins, distribution and biology of the world’s gliding mammals It aims to reveal – for the first time – the extraordinary beauty, behaviour, ecology and origins of a wonderfully diverse and intriguing group of mammals that are united, not by their evolutionary history, but by their ability to glide Many of the 65 species of gliding mammals discussed here are only poorly known – even the most basic information on the biology and distribution of many species has not been adequately recorded I hope, therefore, that this book will stimulate further research and conservation of these spectacular animals Because the most significant aspect that links this group of animals together is gliding, I have included in the introductory chapter a detailed examinantion of the adaptations and behaviour associated with gliding It looks at the behaviour of these animals during the preparation, aerial descent and landing phases of a glide, and includes a comparison of the gliding efficiency of the different groups In writing this book, exhaustive efforts were made to find every available published reference on the world’s gliding mammals, although not every reference was used This included extensive internet searches and visits to libraries at the Australian Museum (Sydney, Australia), the National Museum of Natural History (Smithsonian Institution, Washington DC, USA), the Natural History Museum (London, England) and the Naturalis Museum (Leiden, Netherlands) The vernacular and scientific names used here follow a major revision of the taxonomy of all gliding mammals which was undertaken by myself and Dr Richard Thorington (Smithsonian Institution, Washington DC) and published as ‘Gliding Mammals: Taxonomy of Living and Extinct Species’ in Smithsonian Contributions to Zoology (2012) The designation of many subspecies as species within the giant flying squirrels of the genus Petaurista remains to be confirmed, and there is an urgent need to review their validity A colour painting by the internationally renowned artist Peter Schouten accompanies the account of each species, revealing its distinctive features and the wonderful diversity in size, colour and shape of the various gliding mammals For a number of species the paintings are the first ever depictions in any form This fact also highlights the urgent need for further field research on these often little understood mammals The paintings used in this publication were derived from numerous photos of live specimens (where available) as well as museum specimens which the artist and author took of every species (and most subspecies) of gliding mammal from the museums mentioned above and the American Museum of Natural History (New York, USA) The photos included views of the upper and lower surfaces of the whole skins as well as close up views of the front and side of the head Some of the distribution maps in the species accounts were collated by adding maps from the regional or country level so that the entire distribution for each species is shown on the one map Where subspecies are recognised within a species, the maps endeavour where possible to include the approximate distribution of each vi Gliding Mammals of the World subspecies There may be some inaccuracies in the sources of the maps and the scale used; however, they are based on the best available information at this time The measurements provided for each species offer an important aid in identification and were derived from information associated with museum specimens and the available literature in books and other published information The measurements given are as follows: HB the length of the head and body from the tip of the snout to the cloaca (or anus) along the ventral surface; TL the length of the tail from the cloaca or anus to the last bone in the tail tip; HF the length of the hind foot from the heel to the base of the claw of the longest toe; M the body mass; this provides a good indicator of general size and assists in broadly categorising the different groups of gliding mammals The appendices include a list of the gliding mammals found at specific locations around the world The aim of this list is to allow those interested to find out which species are located within their area, be they a tourist wishing to see these species or a scientist wishing to undertake further research It also helps to highlight the regions that are gliding mammal ‘hot spots’ and therefore should be given particular priority for the conservation of their habitat There is also a glossary that explains some of the technical terms used and an appendix that details the more technical aspects of gliding for those who might want to explore the mechanics of gliding in greater detail Owing to the breadth of information used and the difficulty with which this information has been obtained, an extensive reference list has been included which has been the source of information used in this book I hope that this will serve to stimulate further research into this group of often poorly studied animals Every effort has been made to ensure the accuracy of the information used within this book by making exhaustive reference to both published and unpublished literature Readers are encouraged to make use of the primary literature by referring to the references at the end of this book Given the still unstable nature of the taxonomy for some species and the lack of information available for a number of species there are no doubt errors within the text that will be revealed in due course It is also recognised that some errors from the literature may have been continued One of the motivations for writing this book was to highlight the need for further research to expand the knowledge of these mammals and also to highlight inconsistencies in the literature I encourage future researchers to look at species or groups across their distribution, rather than to one country, wherever possible in order to give a broader perspective and hopefully resolve some of the issues Dr Stephen Jackson February 2012 Preface vii Acknowledgements This work would not have been possible without significant assistance from a number of people First, I am truly grateful to Peter Schouten for coming on board with this project and the enthusiasm and dedication he has shown in creating the most extraordinary paintings and drawings My sincere thanks go to Richard Thorington, who provided abundant advice on flying squirrels and assisted me greatly before, during and after my visits to the National Museum of Natural History (Smithsonian Institution) in Washington DC Thanks to James Whatton who organised x-rays of scalytailed flying squirrel forearms and answered my queries Thanks also to the curators of the different museums for assisting me during my visits to take photos of the gliding mammal skins, from which the paintings of each species were completed These included Richard Thorington, Linda Gordon and Kris Helgen (Smithsonian Institution, Washington DC), Eileen Westwig (American Museum of Natural History, New York), Roberto Miguez (Natural History Museum, London) and Hein van Grouw (Nationaal Natuurhistorisch Museum, Naturalis Museum, Leiden) Colin Groves from the Australian National University in Canberra supported the concept of this project from the beginning and assisted greatly in providing references and shedding light on various aspects of the taxonomy of fossil and extant gliding mammals Momchil Atanassov provided significant information on the citations of fossil gliding mammals and Christopher Beard answered various questions on the taxonomy of fossil dermopterans Many thanks to Joanne Burden, Peter Stevens, Ian Renard, and Richard and Caroline Travers for translating several important manuscripts Thanks also to Yoshinari Kawamura and Tatsuo Oshida who provided a number of references that were difficult to obtain Eric Sargis provided important information and support during the writing of this text with respect to the Dermoptera Many thanks to Davide Molone for providing accommodation and useful discussions during one of my visits to the Natural History Museum in London Thanks also to Paul Andrew and Dion Hobcroft for helping with photos and information on several species Many thanks also to Anthea Gentry who provided valuable information on the history of the Arrow-tailed Flying Squirrel John Scheibe is gratefully acknowledged for providing valuable footage of flying squirrels, several important references and valuable advice Motokazu Ando also provided numerous references on the different species of Japanese flying squirrels and unpublished information, which has been gratefully received Important information on the scaly-tailed flying squirrels was provided by Michael Hoffman, which is much appreciated Ken Aplin provided important information on the taxonomy of the Feathertail Glider and the Greater Glider.Various colleagues also read over chapters or sections of this book depending on their areas of expertise, which has greatly helped the accuracy of this book These colleagues include: Ken Aplin, Douglas viii Gliding Mammals of the World Boyer, Greg Byrnes, Anthea Gentry, Ross Goldingay, Kris Helgen, Graeme Huxley, Norman Lim, Tatsuo Oshida, Richard Rowe, John Scheibe, Anja Schunke, Richard Thorington and Peter Zahler Several libraries and their associated staff were very helpful in bringing the enormous literature that this project required together These include Carol Gokce, Paul Cooper, Eliza Walsh, Kirsten Marshall, John Rose and Emma Solway from the Natural History Museum in London, who provided many of the references and assisted me during my first visit to the library Nicola Gamba, Paul Cooper, Lisa Di Tommaso, Samantha Gare, Nadja Noel, Kamila Reekie, John Rose and Angela Thresher assisted me during my second visit to the Natural History Museum Thanks also to Therese Nouaille-Degorce and Evelyne Bremond-Hoslet from the Bibliothèque Centrale du Museum National d’Histoire Naturelle in Paris for providing a number of valuable references Great thanks also to the staff at the libraries of the National Museum of Natural History (Smithsonian Institution, Washington DC) including Martha Rosen, Leslie Overstreet, Daria Wingreen-Mason and Kirstin van der Veen, who helped me enormously in finding and copying references for this project Thanks also to the staff of the National Museum of Natural History Naturalis in Leiden, including Tom Gilissen, Marianne van der Wal and Agnes Bavelaar for all their help Many thanks also to the Australian Museum and staff including Fiona Simpson, Anina Hainsworth, Fran Smith and Leone Lemmer Thanks also to Rose Bollen and Leonie Cash from the Museum Victoria library for their help in providing access to some historical images My gratitude is also extended to Nick Alexander and CSIRO Publishing for their great support of this project Finally a sincere thank you to Kerstin, Olivia and James for all their encouragement and entertainment during the writing of this book This book is dedicated to my mother, Dorothy ‘Jill’ Jackson, who passed away before this book was finished, and my father who both encouraged this project from the beginning and continue to inspire Acknowledgements ix McKay GM (2008) Greater Glider Petauroides volans In The Mammals of Australia 3rd edn (Eds S Van Dyck and R Strahan) pp 240–242 Reed New Holland, Sydney Mackowski CM (1986) Distribution, habitat and status of Yellowbellied Gliders Petaurus australis Shaw (Marsupialia: Petauridae) in north-eastern New South Wales Australian Mammalogy 9, 141–144 Mackowski CM (1988) Characteristic of eucalypts incised for sap by the Yellow-bellied Glider Petaurus australis Shaw (Marsupialia: Petauridae), in north-eastern New South Wales Australian Mammalogy 11, 5–13 Malekian M, Cooper SJB and Carthew SM (2006) An extension to the known distribution of the Squirrel Glider 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36 Biswamoyopterus, 61 biswasi, 4, 61, 72 Black Flying Squirrel, 66 Chinese Giant Flying Squirrel, 19, 134 Complex-toothed Flying Squirrel, 19, 160 Cynocephalidae, 52–54 Cynocephalus, volans, 4, 52, 184–185 Dermoptera, 2, 48 Diprotodontia, x Dwarf Scaly-tailed Flying Squirrel, 6, 167, 174 Eoglaucomys, 61, fimbriatus, 4, 74, 185, 187 Eomys, quercyi, 14 Eupetaurus, 4, 6, 59–61 sp., 78, 185–186 cinereus, 76, 185 Feathertail Glider, 2, 5, 10, 24, 26–28, 30 214 Gliding Mammals of the World Galeopterus, variegatus, 4, 54, 182–184 Glaucomys, 59 sabrinus, 4, 80, 187–188 volans, 59, 61, 82, 187–188 Glirulus, japonicus, 13 lissiensis, 13 Gray-headed Giant Flying Squirrel, 116, 134 Grey-cheeked Flying Squirrel, 92 Greater Glider, 2, 4–5, 7–10, 15, 24, 25, 26, 27, 29, 44 Hagen’s Flying Squirrel, 6, 144, 146 Hainan Giant Flying Squirrel, 120 Hairy-footed Flying Squirrel, 6, 70, Hodgson’s Giant Flying Squirrel, 7, 19, 126, 128 Hose’s Pygmy Flying Squirrel, 106, 108, 110 Hylopetes, 6, 58, 61, alboniger, 84, 183–186 bartelsi, 86, 183 nigripes, 4, 88, 183–184 phayrei, 90, 183–186 platyurus, 92, 182–185 sagitta, 62, 94, 183 sipora, 96, 183 spadiceus, 98, 182–185 winstoni, 100, 184 Indian Giant Flying Squirrel, 61, 121, 132 Idiurus, 6, macrotis, 4, 176, 188 zenkeri, 178, 188 Indochinese Flying Squirrel, 6, 90 Iomys, 61, 100 horsfieldi, 4, 102, 182, 183, 184, 185 sipora, 104, 183 Japanese Dormouse, 13 Japanese Flying Squirrel, 2, 60, 61, 154 Japanese Giant Flying Squirrel, 18, 60, 124 Javanese Flying Squirrel, 102, 104, 158 Kashmir Flying Squirrel, 19, 74 Lesser Pygmy Flying Squirrel, 106, 108 Long-eared Scaly-tailed Flying Squirrel, 6, 176, 179 Lord Derby’s Scaly-tailed Flying Squirrel, 5, 10, 165, 168, 170 Mahogany Glider, 9, 12, 15, 19, 40 Malayan Colugo, 50–52, 54 Mentawai Flying Squirrel, 104 Mindanao Flying Squirrel, 6, 138, 148 Miopetaurista, 59 Namdapha Flying Squirrel, 61, 72 North Chinese Flying Squirrel, 19, 59, 64 Northern Flying Squirrel, 9, 10, 14, 16, 20, 59, 80, 82, 180 Northern Glider, 16, 17, 19, 32 Pakistan Woolly Flying Squirrel, 76, 78 Palawan Flying Squirrel, 88, 138 Particolored Flying Squirrel, 84, 90 Pel’s Scaly-tailed Flying Squirrel, 6, 166, 168, 172, 179 Petauroides, 5, 6, 29, ayamaruensis, 25 volans, 4, 44, 182 Petauridae, 2, 24, 32–42 Petaurillus, emiliae, 4, 6, 61, 106, 183 hosei, 108, 182–183 kinlochii, 110, 184 Petaurista, 6, 59–62, albiventer, 112, 185–187 alborufus, 114, 183, 185–186, caniceps, 116, 183, 185–186, elegans, 118, 182– 186, hainana, 120, 186 lena, 122, 186 leucogenys, 4, 124, 186, magnificus, 126, 185–186, nobilis, 128, 185 petaurista, 63, 130, 181–186, philippensis, 132, 183, 185–186, xanthotis, 134, 186 yunanensis, 136, 183–186 Petauristodon, Petaurus, 2, 4, 5, 6, 16, 24, 25, 26, 27, 29, 38, 44 abidi, 32, 182 australis, 34, 182 biacensis, 19, 36, 182 breviceps, 4, 38, 182 gracilis, 40, 182 kholsi, 36 norfolcensis, 26, 42, 182 Petinomys, 6, 58, 61, 142, 148, crinitus, 138, 184 fuscocapillus, 140, 185 genibarbis, 4, 142, 182–184 hageni, 144, 183–184 lugens, 146, 183–184 mindanensis, 148, 184 setosus, 150, 182–185, vordermanni, 152, 182–185 Philippine Colugo, 48, 50–52, 54 Pseudocheiridae, 2, 24, 44 Pteromyini, 58 Pteromys, 59, 61, 165, 166, 167 momonga, 154, volans, 4, 63, 156, 185–186, Pteromyscus, 61 pulverulentus, 4, 158, 182–185 Pygmy flying squirrels, 6, Pygmy Scaly-tailed Flying Squirrel, 6, 176, 178 Red and White Giant Flying Squirrel, 19, 114 Red-cheeked Flying Squirrel, 58, 90, 92, 94, 96, 98 Red Giant Flying Squirrel, 61, 63, 68, 112, 121–122, 130, 132, 181 Rodentia, Sciuridae, 2, 58, 64–160, 164 Selangor Pygmy Flying Squirrel, 108, 110 Siberian Flying Squirrel, 5, 20, 60–61, 63, 156 Siberut Flying Squirrel, 6, 146 Sipora Flying Squirrel, 96 Smoky Flying Squirrel, 102, 158 Southern Flying Squirrel, 9, 14, 16, 20–21, 59, 62, 80, 82, 180 Spotted Giant Flying Squirrel, 60, 61, 116, 118 Squirrel Glider, 12, 25–27, 29, 38, 40, 42 Sugar Glider, 6, 9, 11–12, 18–19, 25, 28, 32, 36, 38, 40, 42 Sumatran Flying Squirrel, 100 Taiwan Giant Flying Squirrel, 7, 60, 122, 130 Temminck’s Flying Squirrel, 6, 150 Thomas’s Flying Squirrel, 68 Tibetan Woolly Flying Squirrel, 78 Travancore Flying Squirrel, 140 Trogopterus, 61, 72, xanthipes, 4, 160, 186 Volaticotherium, antiquus, 14, 15 Vordermann’s Flying Squirrel, 6, 94, 98, 152 Whiskered Flying Squirrel, 6, 142 White-bellied Giant Flying Squirrel, 112 Yellow-bellied Glider, 7, 9, 12, 25, 27, 30, 34 Yunnan Giant Flying Squirrel, 136 Index 215 ... anus) along the ventral surface; TL the length of the tail from the cloaca or anus to the last bone in the tail tip; HF the length of the hind foot from the heel to the base of the claw of the longest... of the taxonomic group considered In the gliding possums of the genus Petaurus, the plagiopatagium extends from the joint of the second and third bones in its fifth digit of the front paw to the. .. changing the position of the limbs and the tension of the muscular gliding membrane A left turn is accomplished by lowering the left Gliding Mammals of the World A Malayan Colugo often launches itself