Gliding Mammals of the World

Gliding

Mammals

of the

World

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

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Front cover: Squirrel Glider

Front flap: Feathertail Glider

Back flap: Red-cheeked Flying Squirrel

Back cover: Whiskered Flying Squirrel

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Contents

Preface

Acknowledgements

List of species

1 Introduction

2 Gliding Marsupials

3 Colugos

4 Flying Squirrels

5 Scaly-tailed Flying Squirrels

Appendix 1: Subspecies

Appendix 2: Gliding mammal localities

Appendix 3: IUCN Red List Categories

Glossary

References

Index

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 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

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 scaly-tailed 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 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.

List of species

DIPROTODONTIA

Acrobatidae

Petauridae

Pseudocheiridae

DERMOPTERA

Cynocephalidae

RODENTIA

Sciuridae, Pteromyini

Anomaluridae

Introduction

The world of gliding

The world’s gliding mammals are a diverse group of animals that have the unusual ability to glide from tree to tree with seemingly little effort. They do this by launching from the upper branches or trunk of a tree and spreading out their specially adapted gliding membranes, which stretch from the sides of their body between their fore and hind limbs. This allows them to glide silently through the night air for a considerable distance – some species are able to glide for more than 100 metres. During these glides they can twist and turn around obstacles to make a safe landing on a target tree without the need to come to the ground.

There are currently 65 recognised species of gliding mammals from six different families. There are three families of gliding marsupials that live in Australia, New Guinea and surrounding islands. These families include the Feathertail Glider (Family Acrobatidae), the gliding possums of the genus Petaurus (Family Petauridae), and the Greater Glider (Family Pseudocheiridae). However, by far the greatest diversity of gliding mammals occurs in the Order Rodentia, where they are represented by the flying squirrels belonging to the Family Sciuridae and the unrelated scaly-tailed flying squirrels of the Family Anomaluridae. The Sciuridae includes all the tree and ground squirrels with some 51 genera and 278 species in total. Of these, the flying squirrels comprise 15 genera and 49 species, and are found throughout Asia, Europe and North America. The family of scaly-tailed flying squirrels that live in central and western Africa has seven species, although one species does not glide. Gliding reaches its most spectacular and efficient in the two species of colugos, also known as flying lemurs, of the Order Dermoptera, which occur throughout South-East Asia.

Animals that glide between trees descend at an angle less than 45° to the horizontal, while those that parachute descend at an angle greater than 45°. Gliding is achieved by deflecting air flowing past a well-developed gliding membrane, or patagium, on each side of the body. These membranes convert the animal’s body into an effective airfoil, allowing it to travel the greatest possible horizontal distance with the minimum loss of 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 their gliding membranes.

In addition to mammals and birds, gliding has evolved independently as a form of locomotion in several groups of arboreal and even aquatic vertebrates. These include flying fish of the Family Exocoetidae (from which we get the name Exocet missile), which have either two enlarged pectoral fins or even four enlarged fins (both pectoral and pelvic) to act as wings. These modifications allow them to make glides of over 60 metres above the water to escape from predators and potentially save energy.

The flying snakes of the genus Chrysopelea (and possibly the genus Dendrelaphis) of the Family Colubridae have developed a form of gliding by flattening and broadening their body like a ribbon through the use of hinged ventral scales, and by drawing in the belly so that it forms a concave surface when they leap out of trees.

Flying geckos of the genus Ptychozoon, fringed geckos of the genus Luperosaurus, and house geckos of the genus Cosymbotus (Family Gekkonidae) jump from tree to tree aided by webbed feet, flaps or folds of skin along the lateral body wall, and dorso-ventrally flattened tails that increase their horizontal surface area. Among the reptiles, however, the development of gliding reaches its pinnacle within the gliding lizards of the genus Draco (Family Agamidae). The ribs of these species are greatly elongated to create a large gliding surface, which folds against the sides of the body when not in use. When these lizards jump from a tree, they spread their ribs to stretch out the gliding membrane and can make glides of over 30 metres.

The Japanese Flying Squirrel weighs about 200 grams and is capable of gliding up to 100 metres.

Non-mammal vertebrates that have evolved the ability to glide include (from the top) the fying lizard (Draco), the fying gecko (Ptychozoon), the fying snake (Chrysopelea), the fying fsh (Exocoetidae) and the fying frog (Rhacophorus).

A variety species of frogs from three families are known to parachute or potentially glide. These include various species of flying frogs from the genus Rhacophorus and some of the whipping frogs of the genus Polypedates (Family Rhacophoridae) from South-East Asia. Several species of South American frogs of the Family Hylidae have extensive webbing including the Gliding Leaf Frog (Agalychnis spurrelli), Fringe-limbed Treefrog (Hyla miliaria) and Rabb’s Fringe-limbed Treefrog (Ecnomiohyla rabborum) and may also have the ability to glide. Various other frogs are known or thought to undertake controlled aerial descent including Hyperolius castaneus (Family Hyperoliidae) that occurs in tropical and subtropical forests of central Africa. The most developed frogs typically possess enlarged toes that have well developed webbing between them, with some species from the old world even having flaps of skin on the forelimbs and hind limbs to help trap air during descent. While the frogs with the most highly developed webbing may be able to truly glide, most species are more accurately described as parachuters. Nonetheless, the aerial control of these frogs is often so well developed that they can even make banked 180° turns.

Gliding adaptations

The two things (which may be related) that link every species of gliding mammal together are their nocturnal behaviour and their ability to glide. Despite the diversity of their origins, the different groups of gliding mammals show a remarkable degree of convergence in relation to their gliding adaptations and behaviour. But how do these animals undertake this extraordinary method of locomotion and how far can they glide?

The patagia, or gliding membranes, of mammals consists of skin