SECOND EDITION

FIELD GUIDE TO THE

MAMMALS OF SOUTH-EAST ASIA

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9781472934970_Mammals of SE Asia.indb 2 19/02/2019 10:45
 SECOND EDITION

FIELD GUIDE TO THE

MAMMALS OF SOUTH-EAST ASIA

CHARLES M. FRANCIS

Illustrated by Priscilla Barrett, Robin Budden, John Cox, Sandra Doyle,

Brin Edwards, Ray Hutchings, William Oliver, Guy Troughton and Lyn Wells

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 BLOOMSBURY WILDLIFE
Bloomsbury Publishing Plc
50 Bedford Square, London, WC1B 3DP, UK

BLOOMSBURY, BLOOMSBURY WILDLIFE and the Diana logo are trademarks of

Bloomsbury Publishing Plc

First edition published in 2008 by New Holland Publishers (UK) Ltd

This edition published in the United Kingdom 2019

This electronic edition published in 2019 by Bloomsbury Publishing Plc

Copyright © Charles M. Francis, 2019

Cover illustrations © Priscilla Barrett; Robin Budden; Sandra Doyle; Brin Edwards; William Oliver; Lyn Wells, 2019
Half-title page illustrations © Sandra Doyle (top) and Lyn Wells (bottom), 2019
Title page illustration © Lyn Wells, 2019
Plate colour illustrations © Priscilla Barrett 38–48;
Robin Budden 1–3; 72–81;
John Cox 14–17, 28;
Sandra Doyle 5–13, 18 (part), 25;
Brin Edwards 4, 29, 50–54;
Ray Hutchings 63–66;
William Oliver 55–62;
Guy Troughton 18 (part), 19–24, 26–27;
Lyn Wells 30–37, 49, 67–71,
2019

Charles M. Francis has asserted his right under the Copyright, Designs and Patents Act, 1988, to be identified as
Author of this work

For legal purposes the Figure credits on p. 408

constitute an extension of this copyright page

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to in this book. All internet addresses given in this book were correct at the time of going to press. The author
and publisher regret any inconvenience caused if addresses have changed or sites have ceased to exist, but can
accept no responsibility for any such changes

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ISBN: PB: 978-1-4729-3497-0; eBook: 978-1-4729-3498-7; ePDF: 978-1-4729-3499-4

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 CONTENTS

Preface to second edition 7

Acknowledgements 8
Introduction 11
What is a mammal? 12
Classification and naming 12
How to identify mammals 13
Colour plates 15
Species accounts 16
Finding and studying mammals 20
Where to find mammals 26
Conservation of mammals in South-east Asia 28
PLATES 30
SPECIES ACCOUNTS 192
PHOLIDOTA 192
Manidae Pangolins 192
INSECTIVORA 193
Erinaceidae Gymnures 193
Talpidae Moles 195
Soricidae Shrews 198
SCANDENTIA 209
Ptilocercidae Feather-tailed treeshrews 209
Tupaiidae Treeshrews 210
DERMOPTERA 211
Cynocephalidae Colugos 211
CHIROPTERA 212
Pteropodidae Fruit bats 213
Rhinopomatidae Mouse-tailed bats 222
Emballonuridae Sheath-tailed and tomb bats 223
Craseonycteridae Hog-nosed bats 226
Nycteridae Slit-faced bats 226
Megadermatidae False-vampires 227
Rhinolophidae Horseshoe bats 228
Hipposideridae Roundleaf bats 238
Vespertilionidae Common bats 248
Miniopteridae Bent-winged bats 285
Molossidae Free-tailed bats 286
PRIMATES 288
Lorisidae Lorises 289
Cercopithecidae Monkeys 290
Hylobatidae Gibbons 299

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 CARNIVORA 303
Canidae Dogs 303
Ursidae Bears 305
Ailuridae Red pandas 306
Mustelidae Martens, weasels, badgers and otters 307
Viverridae Civets 312
Herpestidae Mongooses 317
Prionodontidae Linsangs 318
Felidae Cats 319
Otariidae Fur seals and sealions 323
Phocidae Seals 324
CETACEA 324
Delphinidae Oceanic dolphins 325
Phocoenidae Porpoises 330
Physeteridae Sperm whales 330
Ziphiidae Beaked whales 331
Balaenopteridae Rorquals or baleen whales 332
SIRENIA 335
Dugongidae Dugong 335
PROBOSCIDEA 335
Elephantidae Elephants 335
PERISSODACTYLA 337
Tapiridae Tapirs 337
Rhinocerotidae Rhinoceroses 337
ARTIODACTYLA 338
Suidae Pigs 339
Tragulidae Mousedeer 340
Moschidae Musk-deer 342
Cervidae Deer 342
Bovidae Cattle, buffalo, antelopes, goats and sheep 347
RODENTIA 351
Sciuridae Squirrels 351
Muridae Rats and mice 372
Platacanthomyidae Pygmy-dormice 395
Cricetidae Voles 395
Spalacidae Bamboo rats 397
Diatomyidae Kha-nyou 398
Hystricidae Porcupines 399
LAGOMORPHA 399
Ochotonidae Pikas 400
Leporidae Hares and rabbits 400
Glossary 402
Selected bibliography 404
Figure credits 408
Index 409

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 PREFACE TO SECOND EDITION

This book is the first comprehensive field guide species; for example, detailed analysis of type
to mammals of mainland South-east Asia, specimens of two species of bats included in the
describing all species of terrestrial and marine first edition has shown that they never existed; the
mammals known to occur in Myanmar, Thailand, names were based on mixed-up or poorly prepared
Laos, Vietnam, Cambodia, Peninsular Malaysia type specimens.
and Singapore. It is intended to help both general Additions to the book also include some very
naturalists and scientists to identify wild mammals distinctive large mammals. Most surprisingly, a
they may encounter, as well as to increase public very distinctive new species of snub-nosed monkey
understanding of the diversity of mammals in the was discovered in northern Myanmar, although it
region, thus encouraging efforts to conserve them. is very rare and localised, and is now considered
In the 10 years since the first edition of this Critically Endangered. Two species of pinnipeds, the
book was published, much new information has Northern Fur Seal and the Spotted Seal, were also
been obtained on mammals in South-east Asia. At documented for the first time in waters off the coast
least 55 new species have been added to the c.470 of Vietnam. There are some papers suggesting
species that were recognised from mainland South- some additional carnivores, including a possible
east Asia in the first edition of the book. Several new ferret-badger and a civet, but there is still
additional species have had name changes, often as some uncertainty about their identity, so they are
a result of research showing that species formerly not included in this edition.
considered widespread are actually multiple Unfortunately, there have also been losses to
species, often with quite restricted ranges. the mammals in the region in this same period.
Many of the newly recognised species are Most dramatically, both species of rhinoceros that
bats (34 newly recognised species) or shrews formerly occurred in South-east Asia are now
(10 additional species), reflecting a large increase extinct in the region. Only small populations of each
in research efforts on these species. In some now persist in Indonesia, and both are considered
cases, these represent new species that had Critically Endangered. Continuing human population
never previously been seen by scientists. In other growth, increasing development of urban areas,
cases, detailed analyses, often aided by genetic expansion of roads, clearing of native forests for
techniques, have helped with the discovery of agriculture – including rice, rubber and oil palm
cryptic species that had previously been confused plantations – uncontrolled hunting and illegal
with closely related similar looking species. In some trade are threatening many other native species
cases, a species is known to be distinct, but the of mammals. Endangered or Critically Endangered
appropriate scientific name remains uncertain. In species include both pangolins, several carnivores,
these cases, the term ‘cf.’ (meaning ‘compare with’) primates, and ungulates. Substantial conservation
is used to indicate the most similar looking species. efforts are needed to prevent these species from
New research has not always resulted in additional also becoming extinct.

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 ACKNOWLEDGEMENTS

This field guide would not have been possible work while preparing Mammals of Borneo. The
without the pioneering efforts of many previous Malaysian Economic Planning Unit and the National
mammalogists working in the region and publishing Parks and Wildlife Department (PERHILITAN) allowed
their work. I have drawn particularly heavily on A me to carry out field work from 1991 to 1996 in
Field Guide to the Mammals of Borneo by Junaidi Peninsular Malaysia. The Wildlife Conservation
Payne and myself, with illustrations by Karen Society (WCS) generously supported my research
Phillipps. I am grateful to Junaidi, Karen and the in Peninsular Malaysia, as well as subsequent field
Sabah Society for allowing me to use some of the work in Laos from 1995 to 1998. I thank the WCS
line drawings from that book as a starting point for Lao staff for assistance during field work in Laos,
this one. I also made extensive use of Corbet and Hill particularly Bill Bleisch, William Robichaud, Rob
(1992) as a starting point for developing the species Tizard, Will Duckworth, Chanhthavy Vongkhamheng,
lists and basic taxonomic information, for checking Khamkhoune Khounboline and Khoonmy. Nik
distribution maps, for references and for tables Aspey and Antonio Guillén-Servent also assisted
of identification characters, although the species me on surveys there. I have also appreciated
lists have since been updated to reflect Wilson opportunities to spend some time in the field with
and Reeder (2005) and other recently published Sara Bumrungsri, Pipat Soisook, and their students
scientific papers. Many other books on mammals in Thailand, with Vu Dinh Thong and colleagues
of the world and of the region were also useful in in Vietnam, and with Faisal Ali Anwarali Khan and
preparing this book, as well as many original papers students in Sarawak. The Senckenberg Museum,
in the scientific literature. A list of some of the most through Joerg Habersetzer, provided me with
important references is included in the bibliography, equipment for recording bat echolocation calls.
as well as a selection of recent papers describing Michael Bradstreet allowed me to take time off my
new species from the region. duties at Bird Studies Canada to survey mammals in
I have been fortunate to have been able to work the region, while Trevor Swerdfager allowed me to
in the field in the region for many years, particularly use some in-kind support for this project from the
on bats. Much of the information in this book on Canadian Wildlife Service of Environment Canada.
bats, as well as some other groups of mammals, I am grateful to the staff of several museums for
is based on field data I collected myself, often allowing me to use their facilities and collections,
with various colleagues. I am very grateful to all and in some cases to collaborate on joint projects.
those who have supported my field work in South- These include Judith Eger and Burton Lim at the
east Asia. D.R. Wells and D. Melville provided my Royal Ontario Museum; Paula Jenkins, Daphne
first introduction to the fascinating fauna of this Hills and the late John E. Hill at the Natural History
region during a season of field work in Peninsular Museum, London (formerly known as the British
Malaysia and Sarawak in 1979. Ken Scriven has Museum (Natural History)); Paul Bates at the
provided ongoing support over the years. The Sabah Harrison Museum; the late Karl Koopman at the
Wildlife Department (originally part of the Sabah American Museum of Natural History; and the late
Forest Department) and the Canadian volunteer Richard D. Thorington at the National Museum of
organisation CUSO supported me during several Natural History at the Smithsonian. I am particularly
years of field work in Sabah, providing logistical grateful to John E. Hill for teaching me the basics of
support and funding. I am particularly grateful to the mammal taxonomy.
late Mahedi Andau, Director of the Sabah Wildlife During preparation of this book, many people
Department, for allowing me to work on mammals generously answered numerous questions about
as well as birds during my stay in Sabah, and to the individual species, provided myself or the artists with
many of his staff who accompanied and assisted me reprints or manuscripts of their work, unpublished
during field work there. I greatly appreciated working information and photographs, or helped in other
with Junaidi Payne and Karen Phillipps during field ways. These include Alexei Abramov, Bruce Banwell,

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 Paul Bates, Isabel Beasley, Bill Bleisch, Alex ranges have changed substantially and for newly
Borisenko, John Burton, Polly Campbell, Michael recognised species in the region.
Carleton, Gabor Csorba, Jennifer Daltry, Yoan I greatly appreciate the efforts of the artists Brin
Dinata, Will Duckworth, K. Fletcher, Angela Frost, Edwards, Sandra Doyle, John Cox, Guy Troughton,
Neil Furey, Thomas Geissmann, Lon Grassman, Robin Budden, Lyn Wells, Priscilla Barrett, William
Colin Groves, Antonio Guillén-Servent, Simon Oliver, Ray Hutchins and John Buerling who
Hedges, Kristofer Helgen, Jeremy Holden, Tigga contributed their time and skills towards producing
Kingston, Andrew Kitchener, Sergei Kruskop, Darrin the colour plates. Karen Phillipps, Alex Borisenko,
Lunde, Tony Lynam, Debbie Martyr, Masaharu Sergei Kruskop and Nico van Strien kindly allowed
Motokawa, Susan Murray, Phil Myers, Tilo Nadler, me to use some of their line drawings in the text,
W.F. Perrin, Le Khac Quyet, Alan Rabinowitz, Scott while the American Museum of Natural History, the
Roberton, Roland and Julia Seitre, Andrew T. Smith, Sabah Society and the Mammalogical Society of
Pipat Soisook, Carly Starr, Ulrike Streicher, Steven Japan gave permission to reproduce a selection of
Swann, Rob Timmins, Joe Walston, Roland Wirth, line drawings from their publications. I am grateful
Fahong Yu and many others. to Jo Hemmings, Sarah Whittley, James Parry,
The distribution maps in the first edition of this Krystyna Mayer, Charlotte Judet and Liz Dittner for
book, as well as some of the ecological information, support in publishing the first edition, and to the
were derived from the Southeast Asian Mammal team at Bloomsbury for helping to see the second
Databank (SAMD), a project of the Instituto Ecologia edition through to fruition, including Katy Roper, Jim
Applicata in Italy with funding support from the Martin, Susan McIntyre and Amanda Harman.
World Conservation Union (IUCN) and many different Finally, I would like to thank my wife, Cecilia Fung,
partners, which were subsequently used as a basis and my daughter, Fiona Francis, for accompanying
for the IUCN Redlist range maps. I thank them for me in the field when possible, for tolerating my
permission to use the maps in this book. Laine Shaw absences for field trips and museum work when
and Andrew Couturier helped to prepare the maps they could not join me, and for putting up with my
in suitable format for publication. Maps have been preoccupation during many weekends and evenings
updated from a variety of sources where known while completing this book.

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 DISCLAIMER AUTHOR CONTACT

Although this book contains much scientific No doubt I have overlooked some published
information, it is not intended as a primary source works with information on mammals in the region,
of new scientific data on mammals in the region. created errors through my mistakes in interpreting
In keeping with the usual format for field guides, the literature or perpetuated previously published
the text does not include references to sources. errors. I would be most grateful to hear from
Information in this guide has been drawn from anybody who finds errors in this book, or who has
a wide variety of sources, including primary and new information on mammals in the region, so that I
secondary publications, as well as unpublished can incorporate this information into future editions.
data from my own field work and that of colleagues. I can be contacted at:
Whenever possible, I have attempted to verify
information from multiple sources, to minimise the National Wildlife Research Centre
risk of errors. Nevertheless, independent information Environment Canada
was not always available, and I may inadvertently Ottawa, Ontario, Canada, K1A 0H3
have replicated published mistakes or introduced E-mail: [email protected]
new ones. As a result, readers interested in citing or [email protected]
technical details should not consider this guide
as a primary source, but instead should seek out Charles M. Francis
original sources of information, some of which are February 2019
listed at the end of the book. This is particularly true
for measurements, which are provided as guides to
approximate sizes of animals and are not necessarily
indicative of actual specimens.

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 INTRODUCTION

The primary aim of this field guide is to provide closely related species, unless the animal has been
a convenient means for both the general public collected and prepared as a museum specimen.
and professional scientists to identify all of the Some species are difficult to identify, even for
mammals currently known to occur in mainland experts, and users should not expect to identify
Asia, including Myanmar (also known as Burma), every mammal they find, especially those that are
Thailand, Lao People’s Democratic Republic only seen in the field.
(generally referred to as Laos in this book), Vietnam, This book describes every species of wild
Cambodia, Peninsular Malaysia and Singapore. mammal presently known to occur in the region,
The hope is that people using this book will be able including smaller offshore islands. It includes
to identify any mammal that they observe well in approximately 520 species of land mammal
this region by looking through the colour plates from South-east Asia. Sea mammals, including
and reading the corresponding text descriptions. the cetaceans (whales, dolphins and porpoises),
In most cases, it should be possible to identify pinnipeds and the Dugong, are also included.
animals to species, but for some mammals it may Because of the nomadic or migratory nature of
only be possible to narrow this down to a group of many marine mammals, the book includes not only

HANOI
MYANMAR
(BURMA)
V

L
A
I

VIENTIANE E
YANGON O T
(RANGOON)
THAILAND N
S
A

Coco Islands BANGKOK

M

CAMBODIA

PHNOM PENH

0 500km

PENINSULAR
MALAYSIA
KUALA LUMPUR

SINGAPORE

Fig. 1 Map of mainland South-east Asia.

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 species that have been confirmed as occurring in of their hair and their hind legs and replaced their
the region, but also those that have been recorded front legs with flippers. They still breathe air, and
nearby and are likely to show up in South-east they give birth to live young and feed them on milk.
Asian waters. The pangolin somewhat resembles a reptile because
New knowledge is being gained every year on of its scales, lack of teeth and long tongue, but the
mammals in this region by researchers working both scales are actually formed from packed hairs, and it
in the field and in museums. Ongoing field surveys has all the remaining features of a mammal.
in new areas, or using new techniques – such as
camera traps for large mammals, harp traps and
clap nets for bats, and various trapping methods CLASSIFICATION AND NAMING
for small mammals – are providing new information
on the distribution and ecology of species, including To help keep track of the large number of animals in
the discovery of many new species. New study the world and to indicate their relationships, animals
techniques, particularly those that incorporate are classified at different levels. Individuals that can
genetic studies combined with re-examination of freely interbreed and produce fertile offspring are
morphological characters, are leading to further considered to belong to one species. Closely related
new discoveries, as well as better understanding of species are put in the same genus. Genera (plural
species limits in many groups. Since the first edition of genus) that share many features are grouped
of the book, at least 55 new species have been in families, which in turn are grouped into orders,
recognised or discovered in the region. However, which are grouped into classes. All mammals are
much more remains to be learned. There is ongoing put in the class Mammalia. Mammals, along with
uncertainty about species limits in many groups, not other vertebrates such as birds (class Aves), reptiles
only in small mammals such as insectivores, rodents (class Reptilia) and fish (class Pisces) are grouped in
and bats, but also in some groups of large mammals the phylum Chordata.
such as muntjacs. As a result, a book like this must Every species of animal known to science has
be considered a work in progress. been designated a scientific name. These names
are usually based on Latin or Greek, and are used
by scientists all over the world regardless of their
WHAT IS A MAMMAL? own language. Scientific names are always written
in italics to distinguish them. The name is composed
Mammals are distinguished from other animals of two parts, indicating the genus and the species.
by several features. Nearly all species, except a If populations of a species in different geographical
few egg-laying mammals in Australia and New areas can be consistently distinguished from one
Guinea, give birth to live young, and all species another by measurements or colour, then they are
feed their young on milk. Most mammals have fur given subspecies names. For example, Callosciurus
or hair, although in some sea mammals, such as is the genus name for several species of similar
whales and dolphins, the hairs are scattered and medium-sized squirrels in South-east Asia.
inconspicuous. All species are warm-blooded and Callosciurus finlaysonii refers specifically to the
share many features of internal anatomy. Most have Variable Squirrel. Callosciurus finlaysonii sinistralis,
four limbs: two hind legs and two front legs, wings sometimes abbreviated to C. f. sinistralis, is the form
or arms. A few mammals somewhat resemble, of Variable Squirrel found in north-west Thailand,
and could be confused with, other types of animal. while C. f. boonsongi is the form found in north-
Bats are sometimes confused with birds because east Thailand. These subspecies differ in colour, but
they can fly, but they are structurally very different, they are thought likely to interbreed in areas where
with wings formed of skin stretched between long their range overlaps other subspecies. In this book,
fingers, fur rather than feathers, and teeth instead subspecies are generally mentioned only if their
of a beak. They also give birth to live young. Whales, colour patterns are very distinctive, to the extent
dolphins and porpoises are often confused with fish, that they might complicate identification, or where
but they are actually mammals that have lost most there is some doubt as to whether they really should

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 be considered full species, rather than subspecies. but this is misleading – all mammals alive today are
Unfortunately, although there are official rules for equally ‘advanced’ in that all are descended from the
selecting scientific names, the names can change same primitive ancestor over the same time period. In
for a number of reasons. The relationships between this respect, the sequence of orders is rather arbitrary
many animals are still not well understood, and and it becomes, as much as anything, a matter of
as a result not everyone agrees on the taxonomy. convenience as long as closely related species are
A common problem is determining relationships presented near one another. Ongoing research is
between populations of closely related animals from leading to many changes in our understanding of
different geographical areas. Some people may taxonomic relationships among mammals, which
consider them to be all the same species and use has led some authors to change the sequence.
the same name for all of them, while others consider However, as much of this research is still ongoing,
each a separate species, using a different name for I have chosen to retain the sequence followed by
each. At the genus level, some authors may consider Corbet and Hill (1992) for orders, families and usually
two species to be in the same genus, while others genera, as this is similar to the arrangement used in
may consider the differences between them so many published works from the region. Within each
large that they should be in separate genera. New genus, I have attempted to arrange species that
research sometimes shows that animals living are most similar to one another close together. In
together in the same area, and formerly considered some cases, the sequence on the colour plates has
all one species, differ in subtle ways that had not been altered, either to present superficially similar
previously been noticed, and are actually more than species together (e.g. the Dugong is presented with
one species. This sometimes leads to problems the cetaceans), or for convenience to fit species
when deciding to which species the original name on plates.
should be applied. New genetic research is helping English names are a particular challenge, because
to clarify many of the relationships, but it is also many larger mammals have multiple names in the
revealing new challenges and puzzles. Depending literature, while most of the smaller mammals (such
on the authors’ viewpoint, or when a work was as bats and rodents) do not have widely used English
written, books may differ in the name used for a names, largely because the scientists who study
particular animal. them use only the scientific name. As a result, it has
The classification and scientific names used been necessary to invent many of the names used
in this field guide usually follow those given in in this book. Duckworth and Pine (2003) proposed a
Wilson and Reeder (2005), except where more set of rules for developing English names, and they
recent research (published or otherwise) provides gave examples of names for many mammals in the
new information (including many newly described Indochinese region. I have attempted to follow the
species). In some cases, where the names in Wilson principles they advocate in choosing names, but have
and Reeder (2005) do not match the names used not always adopted the names they recommended,
in Corbet and Hill (1992), and where there is still particularly if an alternative name is already well
substantial disagreement in the literature, I have established. Since many of the English names in this
used the older names because these may be more book are not widely known, the scientific name is
familiar to people using other reference books. presented along with the English name whenever a
In these cases, the alternative name is usually species is mentioned in the text.
mentioned as well under ‘Taxonomic notes’.
The arrangement of species in this book, like
most other similar books, is based on grouping HOW TO IDENTIFY MAMMALS
closely related species together. Traditionally, orders
are arranged starting with those believed to have The first step in identifying a mammal is to
diverged earliest from the remainder of the mammal determine to which group or family it belongs.
lineages, and families are arranged within orders This usually involves initially determining the order
on a similar basis. This is sometimes thought to be (e.g. primate, rodent or carnivore), then the family
a ranking from most ‘primitive’ to most ‘advanced’, or subfamily (e.g. within primates, is it a langur, a

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 macaque or a gibbon?). The easiest way to do this mammals and bats, if you are working in a region that
is to look through the colour plates until you find has been poorly studied, you may find animals in new
animals that look generally similar in shape to your areas outside the range shown, but this is less likely
animal. In many cases, this will be straightforward for larger mammals. Note also that some species that
(for example, gibbons are fairly easily distinguished look very similar to other species, but possibly with
from other primates by long arms and the absence quite different distributions, may not be illustrated.
of a tail), but in some cases it may be necessary to These are mentioned on the plate captions, usually
check more closely. For example, treeshrews look under the most similar species; in these cases, you
superficially similar to squirrels, even though they should read the main text to determine differences
are not at all closely related. In these cases, it may and the range of the other species.
be useful to check the descriptions of each order or As a final check on a species, you should
family in the text, to find a description of the main consult the main species account in the text, double
diagnostic differences. Some of these differences checking the description, and also the ‘Similar
may be hard to observe in the field (for example, species’ section, which highlights species with which
treeshrews and squirrels have very different teeth), it might be confused. For groups that contain many
but often there are other characters, such as the different species (such as bats and rodents), you
shape of the body or head, that can help. With should also read the section describing the genus.
practice, most animals can be identified fairly readily Some very diverse genera, such as Rhinolophus,
at least to family, based on their overall shape and are further divided into groups of species that
behaviour, and the search reduced. share certain features. These characters are often
Once the family has been determined, the somewhat more technical (e.g. dentition or shape of
colour plates for that family should be examined the noseleaf) but should be checked to ensure that
closely to find the most similar looking illustration. the identification is correct.
If you are not completely sure of the family, then Many of the larger mammals can be identified
check similar species in each family. Always read from a distance, but fieldworkers who have received
the captions opposite the plates. These highlight the the appropriate specialist training will generally
key identification features that tend to be the most capture smaller species for close examination.
reliable. For some species, colour patterns may be Rodents caught in cage traps can usually be identified
very important (for example, squirrels), while for without handling them. If they need to be held, heavy
others, colour may be unreliable and shape may be gloves can be useful to avoid being bitten.
more important (for example, some horseshoe bats). For some of the smaller species, such as bats,
The captions also indicate if there is more variation shrews and some rodents, the teeth need to be
than is apparent in the plates. For example, in the examined to confirm the identity. These can usually
Variable Squirrel, it was possible to illustrate only be seen in a live bat, for example, by gently prying
some of the many variations of this species – in some open the mouth with a toothpick or similar tool. A
areas, this species may have colour patterns that magnifying lens is generally necessary to see the
are different from any of those shown. Size is also smallest teeth.
important for distinguishing many mammals. For The number and arrangement of teeth is
each species, one key measurement for the group is given in many of the genus and family accounts
given on the caption page. Additional measurements as a ‘Dental formula’. This is a shorthand way
are provided in the text. Measurements are based of indicating the number of teeth in one side of
on standard techniques for measuring museum the upper and lower jaws. The teeth are always
specimens (see page 16). However, the apparent given in the order: incisors, canines, premolars
size of a live animal may vary depending on its and molars (see Fig. 7). For example, the dental
posture. If it hunches its back, the body may seem formula for most Myotis bats is: 2/3 1/1 3/3 3/3,
shorter and the tail proportionately longer. indicating 2 incisors, 1 canine, 3 premolars and
Once you have found a likely candidate for the 3 molars on each side of the upper jaw; and
species, check the range maps to determine whether 3 incisors, 1 canine, 3 premolars and 3 molars on
it is likely to occur in your region. For some small each side of the lower jaw, for a total of 38 teeth.

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 Some teeth are very small, and could be easily accommodate some of the species newly reported
overlooked. Occasionally teeth are missing or have since the first edition. Species that are not illustrated
fallen out. Usually, if this happens, there is a gap generally are those that resemble another species
where the tooth used to be, or the two sides of the jaw so closely as to look essentially the same in the
are not symmetrical, but sometimes the result can be illustrations, as well as a few for which no adequate
confusing. To show the patterns of the teeth and their specimens or descriptions were available. For
relative sizes, diagrams are provided of the tooth rows some species, multiple illustrations have been
for many species where they help with identification. provided, either to illustrate differences between
With present knowledge, some species can be sexes or ages, or to illustrate geographic variation
positively identified only after museum preparation if this is particularly striking. Young animals are
of the skull. For the benefit of scientists using this illustrated in only a few cases, mainly for some of
book, some of the diagnostic characters of the skulls the larger animals in which they have very different
of these species are included. In the future, when patterns from adults, or in cases where their colour
more field studies have been done, other distinctive pattern can help to identify the adults.
features may be found to identify these species in life. In the majority of cases, the artists had to base
Details of skulls are also useful for identifying their illustrations on a combination of museum
dead mammals or their remains. Experts can often specimens to determine colour patterns, and
determine the identity of a mammal using only its photographs to determine shape. Only a few had
skull. A comprehensive guide to skulls is beyond seen their subjects in life. While it would have been
the scope of this book, but diagrams of some of desirable to work with more live animals, this was
the main skull types are included in the text, so that logistically not possible. This presented a challenge
readers can determine the group to which a skull for illustrating many species, especially the
belongs. For confirmation, the specimen can be sent smaller ones, because of a lack of photographs.
to a reputable museum for comparison with skulls of Many species have never been photographed,
known species. while for others the available photographs were
Young mammals can present a challenge for not taken at the appropriate angle or in suitable
identification, as they often differ in size and colour lighting conditions to show the key identification
from adults. The young of larger mammals are often characters. In many cases, it was necessary to
seen with adults, but in smaller species, such as base the shape of an animal on another closely
rodents or bats, the young may be trapped alone. related species. Basing the colour on museum
This is a particular problem in the case of rats, specimens also has some limitations, because
because fur colour is important for identification. the colour of bare skin is not preserved, and fur
Young can often be recognised by having fluffier and colours may change over time.
darker fur than adults, and by having fresh, unworn While an effort was made to check these
teeth that have not fully erupted from the gums. illustrations with experts who had seen the
Young bats can usually be recognised by their greyer animals in the field, it is quite likely that some
fur and incompletely formed wing bones – if held illustrations have not perfectly captured some of
to the light, the joints of the wings appear to have the subtle variations among species in shape or
pale bands where the cartilage has not yet turned colour. Of course, no matter how accurate the
to bone. In some cases it may not be possible to illustration, animals in the field are likely to be
identify young mammals with certainty. observed from angles or in poses and lighting that
differ from the illustrations. This should always be
taken into account when using this guide. It is the
COLOUR PLATES reason why it is always important to check the
species accounts as well as the plates, in order
Nearly every mammal included in this book is to determine which characters are most important
illustrated in the 81 colour plates. For the second and reliable for identification purposes.
edition, four of the plates of squirrels have been
redone, and there are seven new plates to

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 SPECIES ACCOUNTS also given for some species, particularly if these can
be helpful for identification. Although it is sometimes
Measurements possible to measure the teeth on a live mammal,
The account for each species starts with a series of most skull measurements can only be taken on
standard measurements. These measurements have museum specimens.
been taken from a wide variety of both primary and Depending on the type of mammal, and
secondary sources, including field measurements, the availability of information, several different
museum specimens and published ranges in various measurements are provided for each mammal. The
books. They are intended mainly as an aid to readers measurements used in this book are as follows.
to determine the approximate size of each species in
relation to other similar species. The measurements Body measurements (Figs 2–4)
are given as a range intended to capture the normal Head-and-body length (HB) Measured from the
variation in reproductively mature adults of each anus to the front of the nose when the animal is
species, but not necessarily the extremes. However, stretched out.
for measurements taken from published sources, Tail (T) Measured from the anus to the tip of the
it has not always been possible to verify their fleshy or bony part of the tail, excluding hairs that
accuracy, and in a few cases, where measurements project beyond the end.
were not available, I have estimated them from
those of similar species. Thus, these measurements, Hind foot (HF) From the heel to the end of the
especially external body measurements, should be longest toe, excluding the claws.
used only as a general guide to the expected size Ear length (E) From the bottom of the external
of each species. opening of the ear to the tip.
When observing mammals in the field, it is Forearm (FA) (in bats) From the outside of the
important to recognise that most measurements elbow to the outside of the wrist in the bent wing.
are taken on captured animals that have been
straightened out (but not stretched) before taking Total length (TL) (in whales and dolphins) From
the measurement. The head and body length, the front of the head to the notch in the tail flukes.
in particular, may appear rather different in a live Shoulder height (SH) Height from the ground to the
animal in a natural posture. Skull measurements are top of the shoulder when the animal is standing.

T HB

HF

Fig. 2 Measurements of a typical mammal.

HB T
E
SH

FA HF

Fig. 3 Measurements of a bat. Fig. 4 Measurements of a large mammal.

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 Skull measurements (Figs 5–7)
Greatest length (gl) The longest distance from the gl
back of the skull to the front of the skull. Maxillary
Condylobasal length (cbl) From the back of the Premaxillary
occipital condyles to the front of the premaxillae.
Condylocanine length (ccl) From the back of the
occipital condyles to the front of the canines.
Maxillary toothrow (mt) The length of the upper
toothrow from the back of the molars to the front cbl
of the canines (excluding the incisors); usually this
is measured to the base of the canine, but for bats Occipital mt
this extends to the front of the curve of the canine; condyle
for rodents, which have no canines, this includes
only the molars and premolars (three teeth in rats,
four in squirrels – the tiny premolar at the front is
excluded).
Upper toothrow (ut) For shrews the relationships
of the teeth are unclear, and the whole toothrow is 0 2
measured.
Fig. 5 Skull measurements of a rodent.
Molar width (M–M) The width across the outside of
the widest upper molars (at the bases).
Canine width (C–C) the width across the outside of
the base of the upper canines.
All measurements are in millimetres (mm) unless
explicitly stated as (m) for metres. Weights are in
grams (g) or kilograms (kg) as specified.
I
C
gl
PM M
mt
ccl
cbl

gl

ut
mt

I
C PM M
C–C M–M

Fig. 6 Skull measurements of a shrew Fig. 7 Skull measurements and dentition of a bat
(Anourosorex squamipes). (Myotis horsfieldii ).

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 Identification although it may vary geographically. It can reliably
Following the measurements, a concise description be measured even for bats that are in the hand, and
is given of the species, including the overall can often help to differentiate species that are very
coloration and emphasising the most important similar in appearance. This information is given in this
identification characters. The text concentrates section, if available, along with information on where
on identification characters visible on the live animal. it was recorded.
Where necessary, this includes dental characters Similar species This section highlights distinctions
that can often be seen in a live animal if it has from other mammals with which the species
been captured. However, more technical characters might be confused. The list is not exhaustive but
such as skull shape or baculum shape are also concentrates on the most similar species, including
mentioned, if these are important for identification, species that may look different in the illustrations
for the benefit of people working in museums or who but sometimes look similar in life. In most cases, for
have found a dead specimen. some of the larger groups such as bats or rodents,
The colour description is intended to comple- this section only describes similar species within a
ment the colour plates, rather than to duplicate genus or smaller group. It is always important to
them. In particular, the text highlights the range of check the genus accounts to make sure the correct
variation in colour, while the plates show the pattern genus has been selected.
of colours. These should be considered together.
Taxonomic notes These are included only for those
Ecology and habitat
species for which there are known issues related
to taxonomy; for example, if the scientific name Basic habitat preferences and a summary of the
differs from that used in Wilson and Reeder (2005) general ecology are given for each species, where
or the first edition of this book, if the species is known. This section includes mainly information that
newly described or newly recognised, or if recent might be helpful in finding or identifying a mammal
research suggests that more than one species may or learning more about its ecology. For many
be represented within a group. species, especially bats, shrews and rodents, very
little is known, and the only information available
Voice A description is provided only if information is the capture site of the few specimens that have
on vocalisations is available and may be helpful been collected. For large mammals, which are
for identification, for example with primates. Calls often much better known, only a summary of the
can be difficult to describe in words, but it is hoped available information has been included.
that the brief descriptions provided will help to The habitat is described in fairly general terms.
differentiate among species and also help readers Many mammals in the region are found only in
to remember sounds once they have been learned. forests, but little information is available on the range
Echolocation This is only applicable to bats. All of forest types that they can use. The descriptions
insectivorous bats emit distinctive high-frequency provided are generally based on habitats where the
echolocation calls. These usually cannot be heard species has been found, but these are not necessarily
by the human ear (most are in the frequency range the preferred habitats. For example, forest species
20–200kHz), but with the aid of a bat detector (see have been caught or seen in highly disturbed areas
page 21 under finding and studying mammals) it is such as recently logged forest or secondary growth,
possible to measure some of the main characteristics and are reported as using such habitats, but little
of these calls. For most bats, echolocation calls vary information is available on whether the animals are
in frequency over time and change with the behaviour breeding successfully in those areas, or whether
of the bat, so that it is hard to define simple diagnostic they are still dependent on remaining patches of
characters. A comprehensive call library is beyond the mature forest. When describing habitats, the term
scope of this book. However, for both horseshoe bats ‘gardens’ includes land with fruit trees, shrubs and
(Rhinolophidae) and roundleaf bats (Hipposideridae) patches of secondary forest, whereas ‘plantations’
the calls have a distinctive constant-frequency refers to cocoa, oil palm and rubber tree plantings,
component that is very consistent within a population, unless otherwise stated.

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 Distribution Status
This section first describes the known distribution After the distribution, the status is listed based on
of the species within mainland South-east Asia. For the IUCN Red List assessments, using information
widespread species this is generally just a list of posted on the IUCN Red List database as of May
countries, but more detail may be given for species 2018. In some cases, additional information is
with a more restricted range. The description is provided on population trends or threats.
supplemented by range maps that are presented Least Concern This implies that current threats to
for most species on the caption pages opposite the species are not sufficiently serious that it is an
the colour plates. Distributional information was immediate conservation concern, although this may
derived from a range of primary and secondary change in the future.
published sources, as well as information provided
by participants in the Southeast Asian Mammal Near Threatened This implies that the species
Databank project in 2006. The range maps has undergone moderate to substantial population
in this book were generated from the original declines, but not yet sufficiently severe to meet the
maps produced during that project with some higher criteria.
adjustments to enhance visibility of small ranges, Vulnerable This usually indicates that the species
to correct errors or to reflect new information since has undergone substantial population declines in
the first edition of the book. The maps from that the recent past and is still under threat. Unless the
project were also used as the basis for range maps threats are reduced or removed, the species is likely
in many species accounts on the IUCN Red List to become Endangered in the near future.
(www.iucnredlist.org), although some may have
Endangered This implies that the species population
been subsequently revised and may differ in detail
is relatively low, has been declining rapidly and
from the maps in this book.
continues to experience threats that could lead to its
It is important to note that the range maps
extinction unless the problems are reversed.
indicate the regions where a species is expected
to occur in suitable habitat based on known Critically Endangered This implies that only a very
observations. Because much of the original habitat small population remains of the species, that the
in many areas has been heavily disturbed or population is still threatened and that immediate
altered, the actual sites where a species is found action is required to prevent extinction in the very
may be much more restricted. On the other hand, near future.
many species, especially of smaller animals, are Extinct A few species that occurred until recently in
still relatively poorly known, and the distribution of the region are presumed to now be Extinct, but are
known records reflects as much the distribution nevertheless described in this book in the hope that
of research projects as that of the mammals they may still persist. This includes a few that are
themselves. There is some variation in the maps considered Extinct in mainland South-east Asia, but
as to whether the distribution shows only known may still persist elsewhere.
localities or is extrapolated to a broader area. For
a few species that formerly occurred in the region Data Deficient This generally means that there is
but are now believed to be extinct, the areas where insufficient information known about the species to
they were most recently known to occur are shown determine its conservation status.
in grey. Orange shading is used for species believed
to occur in the region, but whose range is uncertain.
If a species occurs outside mainland South-east
Asia, this range is described next, starting with the
word ‘Also’.

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 FINDING AND STUDYING MAMMALS Mammals are most easily found in relatively open
areas. Good places to search are along rivers, in
Although mammals are generally more difficult to small open clearings, along wide paths or on disused
observe than birds, there are many different ways to logging roads. Many mammals can be located by the
see and study them. The best approach depends on noise of their movements – crashing of branches
the types of mammal and their preferred habitats. or rustling of leaves. Some species have distinctive
calls, including many primates, some squirrels and
Observing mammals in the field muntjacs. With practice many of these mammals
Many species of mammal can be observed by can be identified based only on the call. Gibbons can
walking quietly through a forest or other suitable be heard and identified from great distances, even
habitat and watching carefully. During the day, when they cannot be seen. Unfortunately very little
monkeys, apes and many squirrels are active in is known about the calls of many of the nocturnal
trees, treeshrews and some squirrels are found animals, such as the flying squirrels.
in low bushes, and larger mammals such as pigs, Sea mammals, such as whales and dolphins, can
deer, mongooses, bears and otters are active on the be found most readily by travelling out to sea in boats.
ground or in the water. One option is to accompany local fishermen when
Many of the larger mammals are quite shy they make offshore trips. The fishermen themselves
and difficult to approach closely. They need to be may be able to provide valuable information on the
stalked very quietly. Luck is required to see some of numbers of animals they encounter, and the best
the wary or less common species. Some animals, locations to find them, to help with planning such a
such as monkeys, can be gradually accustomed to a trip. Unfortunately, many records of cetaceans in the
human observer, but time and patience are required. region have come from animals becoming trapped
This is especially difficult in areas where mammals in fishing nets or washing up on shore.
are actively hunted. Binoculars are very useful for
watching many of the tree-dwelling animals, and Tracks
are often essential to see the diagnostic features of Many large mammals are detected most readily
some of the smaller squirrels. by tracks or signs. Animal tracks can be useful for
Most other mammals are active mainly at night, carrying out surveys, especially of many of the larger
including the flying squirrels, civets, rats, mice and mammals. Carefully counting and measuring tracks
bats. Large terrestrial mammals such as pigs and of some of the larger ungulates can give information
elephants can be active at any time of day. Many on population numbers and age structure.
mammals are less wary at night than during the day, Some species, such as elephants, tapirs, rhinos
making them easier to observe. A good way to locate and bears, can usually be identified fairly easily to
them is by using a headlamp or spotlight. The eyes of species from their footprints. Others, such as large
most nocturnal animals reflect a red or yellowish glow cats, deer and pigs, can sometimes be identified
in the light. One way to survey mammals is to walk to species, but at other times only to one of a
through the forest using a headlamp to find eyeshine, group of closely related species, depending on the
and a brighter light to see the animals better once number of species occurring in the area. The main
they are found, especially if they are high in the trees. characters for identifying tracks are shape and
This is easiest on a well-marked, relatively clear trail, size, although other factors, such as the distance
where one can walk quietly with minimal risk of between footprints and habitat, can be helpful
tripping or falling. A 30- or 50-watt spotlight can be clues. Some care is required in identifying species,
especially useful for observing animals in the tops of as shape varies depending on the hardness of
tall rainforest (especially if used with binoculars), but the ground, and size varies not only with the size
it may be too bright for mammals that are close. A red of the animal, but also with the soil. In wet mud,
lamp (easily constructed by placing red cellophane tracks may become smaller if the sides collapse
over a spotlight) may be less likely to scare away inwards, while in some soils heavy rain may enlarge
the animals, so they can be observed over a longer tracks. Sketches of footprints of selected species
period of time. are included in the text, mostly of the hind foot as

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 it would look on moderately firm ground. A few transform these high-frequency sounds to forms
examples are provided of animals on soft ground, that we can understand. There are several different
where the toes or hoofs are often more spread types of bat detector, but most researchers now use
out and, in the case of pigs, deer and cattle, the detectors that digitise and capture the whole signal.
dew toes may be visible. Illustrations show typical- They can operate either in full spectrum mode
sized adults, but tracks will vary in size for larger (which retains the most information, but creates
or smaller individuals; young animals can have large files) or zero-crossing mode, which extracts
substantially smaller tracks, although they may be some of the key frequency information to make
accompanied by tracks of an adult. smaller files. The technology is developing rapidly,
Other signs, such as broken branches, claw and some relatively low-cost detectors can now
marks and faeces, can also be useful for identifying record directly to a smart phone or computer. Others
some species, but are not discussed in detail in can be programmed to run continuously for one or
this book. more nights, to monitor any bats that fly by.
In this book, echolocation call frequency
Camera traps information is provided only for the horseshoe
In recent years, a very successful method for (Rhinolophidae) and roundleaf (Hipposideridae) bats,
surveying mammals has been developed using because their calls involve a constant-frequency
cameras along with an infrared or motion detector. component that is very specific to each species
The camera is placed in the forest, usually tied to and can even be recorded in the hand. As a result,
a tree trunk, in an area where mammals might be considerable reference data are available. For other
expected, for example along a trail or next to a species, the call structure varies with behaviour and
stream. The camera can be left in place for a few must be recorded from free-flying bats to obtain
days to a few weeks or months, depending on the reliable information. A number of researchers are
amount of activity expected. Digital cameras allow now working on obtaining this information, using
storage of large numbers of images, and provide a methods such as recording bats immediately after
date/time stamp for each image provided the clock releasing them. However, most species have not yet
is set accurately. When the camera is checked, the been recorded, and more information is required
pictures can be downloaded and the camera can before a sufficiently comprehensive library is
be reset or moved to a new location. Care should be available to determine the range of variation in each
taken to put cameras in areas where they are not species and the best characteristics for identifying
likely to be damaged by animals or removed by other each species by its calls.
people using the area.
Capturing mammals
Bat detectors Many of the smaller animals, such as insectivores,
Bats (except most of the fruit bats, Pteropodidae), rodents and bats, need to be captured for positive
have a sophisticated form of sonar or echolocation identification. Trapping and handling mammals
to find their way in the dark. They emit short pulses must be done carefully to avoid accidental injuries
of sound through their mouths or nostrils and listen and ensure the animals can be released in good
for the echo to learn about obstacles or food in health. This is a job for professionals working in
front of them. Each species of bat has a distinctive the field, who have the appropriate experience and
frequency and pattern of its echolocation calls. qualifications. Before setting traps for mammals,
Many species can potentially be identified from it is also important to have appropriate permits,
their echolocation calls without capture, once a call especially for work in a protected area such as a
library has been developed. park or reserve. Research permits may be required,
Although a few species have components of as well as permits to trap any protected species.
their calls that some people can hear, most of the Many of the most commonly used traps for
calls are very high frequency and inaudible to human small mammals such as rats, mice, squirrels and
ears. Bat detectors are special high-frequency treeshrews involve some sort of box or cage with a
microphones combined with electronic devices that trap door that closes when the animal enters (e.g.

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 Fig. 8 A small mammal cage trap set on a vine to catch arboreal rats or squirrels.

Fig. 8). Cage traps made of wire mesh are available guide animals into the traps. Drainage holes should
locally in many hardware stores, while others such be placed in the bottom so that trapped animals
as folding sheet metal box traps (e.g. Sherman or do not drown in heavy rain, and the traps should
Longworth traps) may need to be purchased from be checked frequently. Food can be placed in the
speciality suppliers. These can be set on the ground bottom, both as bait and so that the animals have
or tied to fallen trees, tree branches or lianas. Many something to eat when caught.
different foods are suitable for bait, including oil For bats, one of the most effective capture
palm fruit, salt fish and fruits such as small ripe methods, and safest for the bats, is a harp trap.
bananas. A larger version baited with raw meat or This consists of a metal frame, typically 1.5–2.0m
fish and set on the ground by a stream or a path square, that supports two to four banks of tightly
can be used to catch some mustelids and viverrids. strung, narrow (0.2mm diameter) vertical fishing line
The smallest mammals, such as shrews and (Fig. 9). Bats usually have enough momentum to fly
ground-dwelling mice, can be caught in pit-fall through one or two rows of fishing line before sliding
traps. These consist of any smooth-sided container down in between the rows and into a holding bag
dug into the ground so that the edges are level with with funnel-like plastic sides. These traps should be
the ground. They must be deep enough that the placed across narrow paths or small streams, so
animals cannot jump out and escape. Small fences the bats are channelled through the traps. The traps
made of branches or wire mesh can be useful to should be checked at least once per hour, and they

Fig. 9 A three-bank harp trap suspended in front of a cave entrance. These traps can also be positioned on legs.

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 Fig. 10 Mist nets set between trees in the forest and across a stream, with the bottom picture showing
how bats get caught.

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 should not be set in areas with very large numbers For insectivorous bats, nets should be watched
of bats, for example near the entrance to a large continuously, so that the bats do not chew holes in
colony, unless they are being watched closely. them and escape. Fruit bat teeth have less effective
Fine nets, called mist nets, are also used to catch cutting edges and the bats are thus less likely to
bats, and are particularly effective with frugivorous escape, so for them it may be possible to check
species which have no echolocation. They have the nets less frequently (e.g. every half-hour),
the disadvantage that insectivorous bats can often depending on bat activity. Bats can become quite
detect them with echolocation and avoid them, and tangled in a net, and proper training is required to
those that do hit the net may escape by chewing learn how to remove bats and other animals without
holes in the net. For insectivorous bats, mist nets injury to either the animals or the surveyor.
are most effective if they have very fine threads Another capture method involves a mobile ‘flap
(including monofilament) and if they are placed net’. This is made from a small, lightweight fishing
across streams, near cave entrances or other areas net about 2.5 × 3.0m made from thread 0.1–
where bats concentrate (Fig. 10). Fruit bats can 0.15mm in diameter with mesh about 14–18mm
be captured most effectively by placing nets near on each side. This is tied with stronger supporting
fruiting or flowering trees. In tall lowland rainforest, cords to the ends of two long (4–5m) lightweight
many fruit bats are most abundant in the canopy poles (carbon fibre collapsible fishing poles are
of the forest, and it can be very effective to hang suitable) into a trapezoidal shape. The two poles
nets from tree tops. This can be done by restringing are held by the surveyor, who uses a headlight to
them to be tall and narrow and suspending them watch for bats (potentially recording them at the
from a single rope suspended over a branch, or by same time, using a bat detector) and swings the
using multiple ropes or a series of stacked tall poles. net to scoop up any bats that are encountered (Fig.
Occasionally small flying squirrels or nocturnal birds 11). This requires an area wide enough to swing
may be caught in mist nets as well. the net, but it can be very effective on wide paths

Fig. 11 Using a flap net to catch bats over a stream.

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 or narrow roads through forest, over ponds or Collecting scientific specimens
streams, or near a roost entrance. Of course, care In some cases, such as when carrying out surveys
must be taken not to trip or fall when walking at of small mammals in new areas, or when a potential
night with long poles, and to avoid power cables new species has been captured, it may be desirable
and other potential dangers. to collect a mammal and prepare it as a museum
Bats can also be found at roost by searching specimen. As with capturing mammals, it is
inside caves, hollow trees or other suitable roosting important to ensure that all necessary permits have
sites. Sometimes roosting bats can be captured been obtained before collecting any animals, and
directly by hand, or with an insect net on a long pole. that the specimen(s) can be properly preserved and
This has the added advantage that it provides useful stored in an appropriate museum.
information on roosting sites for each species, Scientific specimens are particularly important
which can help with conservation planning – some for confirmation of the identity of new geographic
bat species may be limited in their habitat use by records or for describing a new species. It may
availability of suitable roost sites. also be desirable to collect one or two specimens
Regardless of the capture technique, care of each species in ecological studies to allow for
should be taken to minimise handling time and to later confirmation of the identity, although this can
release the animal in a suitable habitat and at a sometimes now be done genetically (see page
suitable time of day. Whenever possible, bats should 26). If specimens are collected, in order for them
be released at night in or near their capture or roost to be useful, appropriate documentation must be
sites, so that they can continue feeding or find their recorded on a label attached to the specimen.
roost in safety. If an animal needs to be held for a Relevant documentation includes a unique
little while, it should be provided with food and water reference number, the exact location (preferably
and kept from overheating. Fruit bats can be given a from a GPS) and date of capture. Specimens must
solution of sugar and water before release. also be properly stored and maintained, preferably
in a national museum with appropriate storage
Surveys of mammal remains areas and trained curatorial staff.
In some areas, an effective survey technique is to Before preparing any specimen, it is important
search for the remains of animals, particularly skulls first to record external measurements, as described
or skull bones. Searching inside caves often leads earlier, including, if possible, the body weight, and
to discovery of the remains of many bats that have write those on the label that will be attached to
died inside the cave, and occasionally other species the specimen. Colour photographs should also be
as well. A particularly effective technique is to find taken, preferably with the numbered label in the
the roosting site of an owl and collect the pellets photograph (so the photos can later be matched to
that accumulate under the roost. Owls usually the specimen) as well as a colour reference strip in
swallow their prey whole and then later regurgitate case the colours of the animal change over time. As
the fur and bones from the prey as a pellet. Some museums improve the electronic catalogues of their
researchers collecting owl pellets in Thailand and collections, they may be able to store photographs
Laos have found the remains of large numbers with the catalogues.
of shrews that had rarely been captured in any Specimens may be prepared as dry skins
other way – owls are very effective small mammal and skeletons or preserved in alcohol. Skins are
collectors! One species of shrew (Crocidura hilliana) particularly difficult to prepare and maintain in hot
was first described on the basis of skulls found in humid climates like those in much of South-east
owl pellets. Asia. They must be quickly dried in the field and then
Identification of skulls, especially partial skulls stored in a cool, dry area in a bug-proof cabinet,
(which are often all that remain in an owl pellet) otherwise they will deteriorate rapidly or be eaten
goes beyond the scope of this book, although some by insects. Dry skins tend to be better for studying
diagrams of skulls and toothrows are provided. fur colour and texture, but then the shape of the
Confirmation will generally require comparison of animal is lost. Instructions for preparing scientific
the remains with reference material in museums. study skins are available in a number of museum

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 reference manuals and should be consulted before failure or other equipment problems. The tissue
collecting mammals. Alternatively, if possible, try samples must be carefully labelled with a specimen
to find an opportunity to work with an experienced number that can be matched back to the museum
mammal collector. voucher. This can be written on the vial with an
Specimens to be stored in alcohol should first alcohol-resistant pen or scratched into the vial to
be fixed for a few days, either in a 10% solution of ensure that it does not rub off.
formalin (equivalent to 4% formaldehyde) or else
in 75% ethanol. Formalin is much less pleasant Genetic approaches to identification
to humans and must be handled carefully, but it A new initiative to identify animals using genetic
fixes the animal more thoroughly. If possible, techniques has recently been developed, called
it should be buffered to reduce its acidity and DNA barcoding. This involves sequencing part of
minimise damage to teeth and bones, which can the mitochondrial genome called the Cytochrome
otherwise be eroded. A mixture of 6g dibasic and c Oxidase subunit I (COI) gene. The sequences of
4g monobasic sodium phosphate per litre of 10% this gene are different for nearly every species of
formalin is recommended, but calcium carbonate animal. An international library of animal barcodes
can be used for short-term storage. It is important is being developed and maintained on an Internet
to ensure that the insides of the animal are database (www.boldsystems.org). A recent study
thoroughly fixed, either by injecting some of the on South-east Asian bats has shown that about
solution into the body cavity, or by slitting open the 98% of currently recognised species can be
body cavity to allow the liquid to penetrate. Once separated based on the DNA barcodes. Sequencing
the animal has been fixed (which takes only a day or of other genes may allow further refinements of
two in formalin), it should be transferred to a fresh identification. A researcher can collect a small piece
solution of 70–80% ethanol for long-term storage. of skin, or even a tuft of fur, extract the DNA and
If the animal was fixed in formalin, it should first be sequence it in a lab, and then determine whether it
thoroughly rinsed in water to remove as much of matches a known species in the database. This can
the formalin as possible. All specimens should be be particularly useful to confirm species identity in
stored in a dark area, to minimise the amount of ecological studies of small mammals, where the
fading that takes place. The containers should be objective is to study live animals with minimum
checked periodically to ensure that the alcohol has disturbance. However, it does not completely
not evaporated. eliminate the need for museum specimens,
Regardless of the preparation technique, it is because it is essential that the reference sequences
very important first to collect some tissue samples in the library be matched to known vouchers, so
that can be used for later genetic analyses. To they can be linked to traditional taxonomy. Also, if
maximise the preservation of genetic material, the DNA sequence does not match a known form,
these should be collected as soon as possible after specimens are required to determine whether it
death (and before contact with formalin) and stored may represent a new species. Differences in DNA
in 95–99% ethanol. Suitable tissues from a freshly alone do not necessarily indicate a new species,
dead animal include liver or other internal organs as there is some variation within species, but large
that are particularly rich in DNA. However, if the differences from known sequences may indicate
animal has been dead for several hours or days, something new.
muscle or skin tissue may be more appropriate,
as there are fewer enzymes to break down the
DNA. Tissue samples stored in high concentration WHERE TO FIND MAMMALS
ethanol will keep for several years, especially at
low temperatures, but for long-term storage it is The area covered by this book spans nearly
recommended to transfer them to a museum facility 28 degrees of latitude, from tropical lowland
with ultracold freezers. Ideally, tissue samples from rainforest near the equator in Peninsular Malaysia,
each specimen should be stored in more than one to upper montane habitats 5,000 metres above
institution, to guard against loss in case of freezer sea level at the edge of the Himalayas in North

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 Myanmar. This diversity in habitats leads to a high diversity and an understorey mainly of grasses.
diversity in mammals. Natural grasslands also occur. Some mammals,
The Isthmus of Kra, at about 8°N in peninsular such as elephants, wild cattle, a few deer species
Thailand, represents a significant biogeographical and some carnivores, may be most abundant in
barrier, with many species reaching their northern grasslands or savannas consisting of a mixture
or southern limits there. To the south is the Sunda of grassy areas and trees. This region also has
sub-region, including Peninsular Malaysia as a higher diversity of species adapted for open
well as Sumatra, Java and Borneo and adjacent areas, including many species of rodent, some of
small islands. Many species are shared among which have expanded with increasing agriculture.
these large islands, as the seas between them Areas with increased rainfall support broadleaved
are relatively shallow and the whole Sunda sub- evergreen forests with an increased diversity of
region was connected at various times in the past fruiting trees, thus supporting many species of
when sea levels were much lower. The majority mammal. Montane forests are dominated by
of mammal species in this region are adapted to different tree species with many more oaks and
lowland evergreen rainforest, which is perhaps the beeches and fewer dipterocarps, supporting a
richest and most diverse forest in the region. The different suite of mammal species. A surprisingly
dominant tree species are dipterocarps, but with rich area of diversity, with many endemic species
a very high diversity of other tree species – over (found nowhere else in the world), is in the
800 different species of tree and shrub have been Annamite Mountains that separate Laos and
recorded from a single 50-hectare study plot in Vietnam. Many bats, as well as a number of
Malaysia. In its natural state, the main canopy is rodent species, including the newly discovered
30–40m tall with multiple layers of understorey Kha-nyou, Laonastes aenigmamus, and several
trees and shrubs, and some tall emergent trees species of langur, are found only around forest-
reaching up to 60m or more. covered limestone outcrops, which usually contain
Some of the mammals in this region are large numbers of caves. Wetlands, including
restricted to higher elevations in hill or montane coastal mangrove forests as well as swamps and
forest, where the trees are lower, often moss marshes along large rivers and lakes, are a very
covered, with different species composition. important habitat for many species, including
Other important habitats include swamp forest, some specialised mammals such as otters.
with similar species to the lowland rainforest but Northern Myanmar extends into the Himalayan
generally lower diversity, and coastal mangroves sub-region, with many Himalayan and Chinese
with a number of specialists, some of which have species reaching their southern limits there. Many
more recently moved into gardens and disturbed of these species are montane specialists, found in
areas. Only a few mammal species are specialised alpine rhododendron forests and scrub, or grassy
for open habitats such as rice fields or grasslands; meadows and rock piles above the tree line. Others
some of these mammals are recent invaders from are adapted for drier country, such as that found in
other regions. Of course, these include some of the India, and reach their eastern limits in Myanmar.
most familiar mammals, such as the rats and mice In most countries in the region, mammals are
that invade human houses. most readily found in many of the protected areas
In the Indochinese sub-region north of the such as national parks and wildlife sanctuaries,
peninsula, the highest diversity of mammals is still both because these often contain some of the best
in forested areas, but there is a greater diversity remaining habitats, and also because the animals
of forest types. Deciduous forests, which shed are more likely to be protected from hunting, so
their leaves during the dry season, occur in many are less shy and occur in greater numbers. A
areas, although they were formerly much more variety of parks and wildlife sanctuaries has been
extensive. These range from mixed deciduous established in each country, covering most of the
and broadleaf evergreen forests with a relatively major habitat types, although the facilities and
dense understorey of shrubs and bamboos, to access vary considerably. Local travel guides should
sparse dry dipterocarp forest with limited tree be consulted for further details.

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 CONSERVATION OF MAMMALS IN However, with increasing human populations,
increased availability of firearms, reduced areas
SOUTH-EAST ASIA of suitable habitat, easier access to formerly
Originally, nearly all the region was covered in forest remote areas by road, and particularly with an
of one type or another. At an escalating rate over the increased demand for wildlife for internal and
past century, much of this forest has been cleared international trade, excessive hunting and trapping
for agriculture or urban development. Much of what is now threatening many animals with extinction.
remains has been selectively logged or otherwise At least two species of mammal, Schomburgk’s
degraded. Deer and Kouprey, have apparently already been
The greatest single threat to most mammals hunted to extinction. Both species of rhinoceros
and other wildlife in the region has been the loss have become extinct in mainland South-east Asia
or conversion of their natural habitats, especially since the first edition of this book, and persist
forest. Most of the terrestrial mammals in the region only in small populations in Sumatra, Borneo
are adapted for forest cover and cannot survive in and Java. Pangolins, which until recently were
agricultural or cleared areas. Some species appear relatively common, have undergone a dramatic
to be able to survive in disturbed forests such as population decline of 90% or more since 1990,
selectively logged or otherwise degraded areas, owing to trapping and trading of their skins for
though possibly in reduced numbers, and the long- alleged medicinal values. Large carnivores such
term viability of their populations is still unknown. as tigers and bears, as well as several primates,
Others are rarely found outside relatively intact tall are threatened by a combination of habitat loss
forests; even selective logging can result in loss and hunting. Even smaller mammals such as
of roosting sites or shelter, as well as changes in bats have been locally wiped out from some cave
the food supply through loss of fruiting trees and colonies by excessive trapping. Marine mammals
reduced diversity of insects. such as dolphins and dugongs are also threatened
As a result, most forest species have undoubt- by a combination of factors, including deliberate
edly experienced a decline in their overall population hunting, accidental drowning in fishing nets,
size over the past few decades, the losses being reduction of food supplies as a result of overfishing,
most severe in species that cannot tolerate and degradation of water conditions from pollution.
disturbance. Loss of habitat results in a direct loss The net result is that many mammal species in
of populations – if two-thirds of a forest is gone, the region are now listed as Endangered or Critically
the remaining forest cannot support more than one- Endangered, as indicated in the status section of
third of the original population size of the species. the respective species accounts, with many further
The clearing of forest results in permanent loss of species listed as Near Threatened or Vulnerable.
all the mammals that used to occur there – they In order to protect the tremendous diversity of
cannot simply move somewhere else. In addition, mammals in South-east Asia, several measures
much remaining forest may be fragmented into are required, of which the most important are
small patches. The smallest fragments of forest protecting large areas of suitable habitat, and
may be too small to support viable populations of reducing the amount of hunting, trapping and
many mammals, particularly the larger species. If wildlife trade. Adequately large areas of all the
the fragments are isolated patches, animals may major habitat types need to be set aside in the
not be able to move among areas or recolonise if a form of gazetted parks and reserves, to protect
population becomes locally extinct. not only mammals but also all other aspects of
An additional threat to many mammals, both wildlife diversity. Although many areas have been
large and small, is hunting and trapping both designated in each country in the region, these
for food and for the so-called medicinal trade. are not always large enough, and enforcement
Formerly, when human densities were much lower and management are generally lagging behind.
and when mammal populations were higher with Greater efforts and resources are required to
more extensive intact habitat, hunting and trapping ensure protection of reserves once they are
of some species may have been sustainable. designated.

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 In addition, suitable management practices are unsustainable. This includes strengthening national
required outside reserves. Areas being managed for and international laws to limit wildlife trade, and
forestry should be managed to preserve as much enhancing enforcement capacity and penalties for
as possible of the natural tree diversity. Forestry violating these laws.
practices are required that minimise damage to Education is also a vital component of any
the forest during selective logging. Degraded areas conservation programme. People will protect only
should be allowed to regenerate or, if necessary, those things that they understand and value. In
be replanted with natural species. Priority should be this respect, I hope that this book will play a role
given to restoring areas near existing reserves to in enhancing understanding of the diversity of
expand their area and carrying capacity, as well mammals in the region, and the need for immediate
as the provision of corridors to connect different conservation actions to protect them. While these
reserves, thus increasing the viability of populations mammal species still exist, there remains a chance
within them. to protect this tremendous level of biodiversity so
The other major requirement is to halt, as far as that future generations can observe and enjoy the
possible, trade in wildlife, nearly all of which is now mammals of South-east Asia.

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 Plate 1: GYMNURES AND MOLES
1. MOONRAT 5. KLOSS’S MOLE
Echinosorex gymnurus, p. 194 Euroscaptor klossi, p. 196
HB 320–400. Large; elongate HB 130–136. No visible eyes
muzzle; long shaggy fur; white or ears; enlarged, rounded
on head and variable amounts front feet, large claws; short
on front half of body. club-shaped tail. See text
for differences from other
similar-looking moles occurring
2. SHORT-TAILED GYMNURE elsewhere in region that are not illustrated:
Hylomys suillus, p. 194 E. malayana, E. orlovi, E. kuznetsovi, E. parvidens,
HB 120–140. Elongate muzzle; E. subanura, Parascaptor leucura and Mogera
conspicuous rounded ears; latouchei.
short, nearly hairless tail; short
spines in fur. Fur dark reddish- ? 6. LARGE CHINESE MOLE
brown (2a) to grey-brown (2b). Euroscaptor grandis, p. 196
HB 150. Similar to E. klossi,
3. CHINESE GYMNURE but larger.
Hylomys sinensis, p. 194
HB 110–125. Like H. suillus,
but has longer, thin tail; one
less premolar in lower jaw.
H. megalotis (not illustrated) 7. LONG-TAILED MOLE
from C Laos has longer, thicker Scaptonyx fuscicaudus, p. 198
tail, larger ears, no spines in fur. HB 65–90. No visible eyes
or ears; somewhat enlarged,
4. SLENDER SHREW-MOLE elongate front feet; moderate-
Uropsilus gracilis, p. 195 length scaly tail.
HB 65–90. Elongate snout;
small eyes hidden in fur; small
but visible ears; long scaly tail;
front feet only slightly enlarged.

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 1

2a

2b 3

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 Plate 2: BROWN-TOOTHED SHREWS AND MOLE-SHREW
1. INDOCHINESE SHORT- 5. VAN SUNG’S SHREW
TAILED SHREW Chodsigoa caovansunga, p. 201
Blarinella griselda, p. 199 HB 58–64. Similar to C. parca,
HB 65–80. Small; moderately but tail shorter without tuft
short tail; five unicuspid teeth, of hairs at tip; feet brown;
last one small and hard to see. footpads touching.
B. wardi from N Myanmar (not
illustrated) is similar, but slightly
smaller; relatively smaller third upper unicuspid. 6. LESSER STRIPE-BACKED
SHREW
2. HODGSON’S BROWN- Sorex bedfordiae, p. 198
TOOTHED SHREW HB 50–70. Small; moderately
Episoriculus caudatus, p. 200 long tail; dark stripe on back;
HB 50–75. Small; moderately five unicuspid teeth all similar
long tail; four unicuspids, last in size.
one small and hard to see.
E. baileyi (not illustrated) is 7. MOLE-SHREW
similar, but slightly larger; has Anourosorex squamipes, p. 201
bigger upper incisors. HB 85–110. Tiny eyes; no
visible ears; very short, scaly
3. LONG-TAILED BROWN- tail; front feet not especially
TOOTHED SHREW enlarged.
Episoriculus macrurus, p. 200
HB 59–66. Similar to
E. caudatus, but greyer; tail
much longer than body; dark
above, usually paler below.

4. LOWE’S BROWN-TOOTHED
SHREW
Chodsigoa parca, p. 200
HB 65–85. Small; orange-
brown tips to teeth; only three
unicuspids; tail longer than HB
with tuft of longer hairs at tip;
feet pale; footpads separated.

Note All species on this plate except Anourosorex have reddish-brown or orange tips to incisors and some unicuspids.

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 1

2 3

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 Plate 3: WATER SHREWS AND WHITE-TOOTHED SHREWS
1. HIMALAYAN WATER SHREW 6. VORACIOUS SHREW
Chimarrogale himalayica, p. 202 Crocidura vorax, p. 207
HB 110–130. Long silvery guard HB 55–90. Small to medium-
hairs on back; only slightly paler sized; sharply bicoloured tail.
below than above; tail usually all
dark; feet fringed with short stiff
hairs. C. hantu from Peninsular
Malaysia (not illustrated) is very
similar, but with cusp on upper incisor. 7. KE GO SHREW
Crocidura kegoensis, p. 206
2. STYAN’S WATER SHREW HB 48. Very small; dark mark
Chimarrogale styani, p. 202 on muzzle and dark ears;
HB 80–110. Like C. himalayica, relatively flat skull.
but smaller, underparts silvery
grey, sharply contrasting with
upperparts; tail dark above,
pale below. 8. INDOCHINESE SHREW
Crocidura indochinensis, p. 206
3. WEB-FOOTED WATER HB 59–68. Medium-small;
SHREW medium-length tail with long
Nectogale elegans, p. 202 bristles at base; flat skull.
HB 90–130. Silvery-white
underparts contrast with dark
upperparts; rows of white hairs
along dark tail. 9. SUNDA SHREW
Crocidura cf. monticola, p. 205
4. HOUSE SHREW HB 53–65. Very small;
Suncus murinus, p. 203 uniformly dull grey-brown; tail
HB 90–145. Relatively large; with bristles on basal half.
moderate-length thick tail; four
upper unicuspids. Often found
around houses.
10. SOUTH-EAST ASIAN
SHREW
5. PYGMY WHITE-TOOTHED Crocidura fuliginosa, p. 204
SHREW HB 70–100. Medium-large;
Suncus etruscus, p. 203 sparsely haired tail.
HB 50–56. Very small; four
upper unicuspids. S. malayanus
(not illustrated) is even smaller
(HB 37–51) and restricted to
peninsular Thailand and Malaysia.
Note Species on this page have white tips to all teeth. Crocidura have only three unicuspid teeth, while Suncus have
an extra, very small posterior unicuspid, although it can be hard to see. Many species in the region appear very similar,
so only a few species are illustrated. See text for distinctions from other similar-looking small or medium Crocidura:
C. annamitensis, C. attenuata, C. cranbrooki, C. guy, C. hilliana, C. malayana, C. negligens, C. phanluongi,
C. phuquocensis, C. rapax, C. sapaensis, C. sokolovi, C. tanakae, C. wuchihensis, C. zaitsevi.

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 1

5 6

10

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 Plate 4: TREESHREWS
1. FEATHER-TAILED 4. NORTHERN TREESHREW
TREESHREW Tupaia belangeri, p. 210
Ptilocercus lowii, p. 209 HB 160–230. Pointed muzzle;
HB 134–150. Grey to grey- long tail; pale mark on
brown; long tail with feather-like shoulder; greyish fur.
tip; largely nocturnal.

2. NORTHERN SLENDER- 5. COMMON TREESHREW

TAILED TREESHREW Tupaia glis, p. 210
Dendrogale murina, p. 211 HB 135–205. Pointed muzzle;
HB 115–135. Pointed muzzle; long tail; pale mark on
very thin, lightly haired tail; shoulder; reddish fur.
stripes on face.

3. LESSER TREESHREW
Tupaia minor, p. 211
HB 110–140. Small; pointed
muzzle; pale mark on shoulder;
relatively long, thin tail.

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 1
3

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 Plate 5: FLYING-FOXES AND TAILLESS FRUIT BATS
1. LARGE FLYING-FOX 5. SUNDA TAILLESS FRUIT
Pteropus vampyrus, p. 215 BAT
FA 185–200. Largest bat in Megaerops ecaudatus, p. 218
region; dark back; orange FA 51–58. Brownish fur with
collar; underparts usually dark. paler, grey bases; dark rims
Roosts in large colonies in open to ears; only one pair of lower
trees. incisors; upturned rostrum;
protruding nostrils.
2. INDIAN FLYING-FOX
Pteropus giganteus, p. 216 6. NORTHERN TAILLESS
FA 140–180. Similar to FRUIT BAT
P. vampyrus, but slightly Megaerops niphanae, p. 219
smaller; collar paler; underparts FA 53–61. Like M. ecaudatus,
usually pale. but rostrum more sloping;
nostrils smaller.

3. LYLE’S FLYING-FOX
Pteropus lylei, p. 216 7. WHITE-COLLARED FRUIT
FA 145–160. Similar to BAT
P. vampyrus, but much smaller; Megaerops wetmorei, p. 219
underparts pale or dark; FA 45–51. Small; short
pointed ears. rostrum; dark rims to ears;
white tufts on sides of neck in
adult male.
4. ISLAND FLYING-FOX
Pteropus hypomelanus, p. 216
FA 120–145. Relatively small;
underside varies from dark
brown to pale brown; rounded
ears.

Note Heads of flying-foxes are drawn at a smaller scale than heads of Megaerops.

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 4

1b

2a

1a 2b

6b
7b 6a
7a

9781472934970_Mammals of SE Asia.indb 39 19/02/2019 10:46

 Plate 6: SMALL AND MEDIUM FRUIT BATS
1. GREY FRUIT BAT 6. GREATER SHORT-NOSED
Aethalops alecto, p. 220 FRUIT BAT
FA 42–46. Small; uniformly Cynopterus sphinx, p. 217
dark grey-brown to reddish- FA 65–76. Pale rims to ears;
brown; thick fluffy fur; no tail; pale wing bones; collar orange
one pair of lower incisors. to yellow-orange in adult males
(6a), paler yellow in adult
females (6b) and young; two
2. DUSKY FRUIT BAT pairs of upper and lower incisors.
Penthetor lucasi, p. 218
FA 57–62. Medium-sized; short 7. HORSFIELD’S FRUIT BAT
grey to grey-brown fur; dark Cynopterus horsfieldii, p. 218
edges to ears; one pair of lower FA 68–76. Like C. sphinx, but
incisors. averages heavier; cheek teeth
large and rectangular with extra
cusps in middle.
3. BLACK-CAPPED FRUIT
BAT
Chironax melanocephalus, 8. FOREST SHORT-NOSED
p. 220 FRUIT BAT
FA 43–46. Small; contrasting Cynopterus cf. brachyotis
dark head; yellow tufts on sides ‘Forest’, p. 217
of neck in adults; two pairs of FA 56–63. Smallest Cynopterus
lower incisors. in region; muzzle short; collar of
adult male dark orange.
4. MALAYAN SPOTTED-
WINGED FRUIT BAT 9. SUNDA SHORT-NOSED
Balionycteris seimundi, p. 220 FRUIT BAT
FA 40–45. Small; dark fur; pale Cynopterus cf. brachyotis
pinkish-white spots on face and ‘Sunda’, p. 217
wing joints; one pair of lower FA 59–70. Averages smaller
incisors. than C. sphinx, with shorter
ears; muzzle relatively long;
5. BLANFORD’S FRUIT BAT collar of adult male yellowish-
Sphaerius blanfordi, p. 221 orange.
FA 52–60. No tail; interfemoral
membrane thickly covered with
hair; fur grey or brown with
grey bases; pale rims to ears;
pale wing bones; two pairs of
lower incisors.

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 1a
5a

1b

3a
5b

4a
3b
6b

4b

6a
7

8a

8b 9b

8c
9a

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 Plate 7: DYACOPTERUS, ROUSETTES AND NECTAR BATS
1. DAYAK FRUIT BAT 4. CAVE NECTAR BAT
Dyacopterus spadiceus, p. 220 Eonycteris spelaea, p. 221
FA 77–81. Large; short greyish- FA 62–70. Elongate muzzle;
brown fur; heavy muzzle with small teeth; no claw on second
large rectangular cheek teeth; digit.
only three upper cheek teeth.

2. GEOFFROY’S ROUSETTE 5. GREATER LONG-TONGUED

Rousettus amplexicaudatus, NECTAR BAT
p. 214 Macroglossus sobrinus, p. 222
FA 78–87. Large; elongate FA 44–50. Elongate muzzle;
muzzle; claw on second digit; long tongue; very small teeth;
lower molars relatively circular. no grooves on upper lip or
around nostrils (5b). Frequently
feeds on banana flowers (5c)
3. LESCHENAULT’S
ROUSETTE 6. LESSER LONG-TONGUED
Rousettus leschenaultii, p. 215 NECTAR BAT
FA 75–85. Large; elongate Macroglossus minimus, p. 222
muzzle; claw on second digit; FA 39–44. Like M. sobrinus,
last lower molar elongate. but slightly smaller; distinct
grooves below each nostril and
in middle of upper lip (6b).

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 1a

1b

2b

2a

4b

4a

5c

5a
5b

6a
6b

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 Plate 8: MOUSE-TAILED, HOG-NOSED, SHEATH-TAILED AND
TOMB BATS
1. LESSER MOUSE-TAILED 5. POUCHED TOMB BAT
BAT Saccolaimus saccolaimus,
Rhinopoma hardwickii, p. 223 p. 225
FA 53–64. Long, protruding FA 71–78. Relatively large;
mouse-like tail; short grey- tail protrudes from middle of
brown fur; distinct ridge of interfemoral membrane; dark
skin on tip of muzzle. See back with white speckles;
text for distinctions from R. underside usually white,
microphyllum (not illustrated). sometimes dark brown; translucent wings; no wing
pouch; glandular pouch under chin.
2. KITTI’S HOG-NOSED BAT
Craseonycteris thonglongyai, 6. THEOBALD’S TOMB BAT
p. 226 Taphozous theobaldi, p. 224
FA 22–26. Very small; no bony FA 70–76. Relatively large; chin
tail; pig-like nose with slight furred without throat pouch,
ridge of skin on tip. sometimes with dark ‘beard’ in
male (6c); well-developed wing
pouch (similar to 7e); no fur on
3. LESSER SHEATH-TAILED interfemoral membrane or legs.
BAT
Emballonura monticola, p. 223 7. NAKED-RUMPED TOMB
FA 43–45. Small; dark brown BAT
fur; short tail protrudes Taphozous nudiventris, p. 225
from middle of interfemoral FA 71–80. Relatively large;
membrane; alert posture. no fur on lower back or upper
side of flight membranes; chin
4. LONG-WINGED TOMB BAT naked, with distinct gular pouch
Taphozous longimanus, p. 224 in male (7c) but not female
FA 54–63. Medium-sized; chin (7d); well-developed wing pouch (7e).
naked; gular pouch in male
(4c), lacking in female (4d); 8. BLACK-BEARDED TOMB
extensive fur on interfemoral BAT
membrane and base of wing; Taphozous melanopogon, p. 224
small wing pouch; colour varies FA 60–63. Medium-sized; chin
from brown to blackish. furred without pouch, often with
black ‘beard’ in male (8c); well-
developed wing pouch (8d);
colour varies from grey-brown
to buffy.

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3a

1a

3b

4a

1b

5b 5a

4b

6a

8d

4c 4d

7a

8b
7b

8a
8c

6b 6c 7e
7c 7d

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 Plate 9: FALSE-VAMPIRES AND SLIT-FACED BAT
1. LESSER FALSE-VAMPIRE 3. THONGAREE’S FALSE-
Megaderma spasma, p. 227 VAMPIRE
FA 56–63. No tail; large ears Eudiscoderma thongareeae,
joined on top of head; posterior p. 228
noseleaf convex on sides, FA 64–66.5. Intermediate in
rounded at top; intermediate size; no tail; large ears joined
noseleaf heart-shaped and on top of head; posterior
broad. noseleaf disk-shaped with stiff
central ridge; intermediate noseleaf heart-shaped;
2. GREATER FALSE-VAMPIRE anterior noseleaf broad.
Lyroderma lyra, p. 227
FA 65–72. Large; no tail; 4. MALAYAN SLIT-FACED BAT
large ears joined on top of Nycteris tragata, p. 226
head; posterior noseleaf with FA 46–55. Very long tail with
straight sides and square top; T-shaped tip; large separate
intermediate noseleaf narrow; ears; deep hollow slit in middle
chin protruding. of face fringed with flaps of skin
like a noseleaf.

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2a 3

2b

4b

4a

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 Plate 10: WOOLLY HORSESHOE BATS
1. LESSER WOOLLY 3. TREFOIL HORSESHOE BAT
HORSESHOE BAT Rhinolophus trifoliatus, p. 228
Rhinolophus sedulus, p. 229 FA 45–56. Lateral lappets at
FA 38–45. Lateral lappets base of sella; long, woolly pale
at base of sella; long, woolly greyish fur; yellowish noseleaf;
blackish fur; dark noseleaf; pale brown wing membranes
relatively small. with yellow elbows and knees.

2. GREAT WOOLLY 4. FRANCIS’S WOOLLY

HORSESHOE BAT HORSESHOE BAT
Rhinolophus luctus, p. 230 Rhinolophus francisi, p. 229
FA 62–76. Largest FA 52–55. Lateral lappets at
Rhinolophus; lateral lappets base of sella; long, woolly dark
at base of sella; long, woolly brownish or greyish fur; dark
dark grey fur; dark noseleaf. noseleaf, intermediate size.
R. luctoides (not illustrated) is
very similar; distinguished by genetics and longer
lower toothrow.

Note Diagrams beside each Rhinolophus face show profile of noseleaf. See also diagrams of sella in Fig. 33.

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1a

3a

3b

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 Plate 11: HORSESHOE BATS
1. CONVEX HORSESHOE BAT 6. SHAMEL’S HORSESHOE
Rhinolophus convexus, p. 234 BAT
FA 42–43. Small; tall, pointed Rhinolophus shameli, p. 232
connecting process; rounded FA 44–49. Similar to
lancet. R. coelophyllus, but larger with
lower frequency echolocation
call.

2. ACUMINATE HORSESHOE 7. SIAMESE HORSESHOE BAT

BAT Rhinolophus siamensis, p. 231
Rhinolophus acuminatus, p. 232 FA 38–42. Like R. macrotis but
FA 44–50. Relatively large; smaller with higher frequency
triangular, pointed connecting echolocation calls.
process; short lancet; short fur
uniformly dark grey (2a, 2b) or
orange (2c).
8. BIG-EARED HORSESHOE
3. LEAST HORSESHOE BAT BAT
Rhinolophus pusillus, p. 233 Rhinolophus macrotis, p. 230
FA 33–39. Very small; pointed FA 42–47. Large ears; tall
connecting process; broadly rounded lancet; long broad
pointed lancet; fur colour sella narrowing to rounded
variable. Refer to text for tip with cup-shaped base; tall
distinctions from R. subbadius, connecting process rounded at
R. monticolus and R. shortridgei tip, forming notch.
(not illustrated).
9. MARSHALL’S HORSESHOE
4. GLOSSY HORSESHOE BAT BAT
Rhinolophus refulgens, p. 234 Rhinolophus marshalli, p. 232
FA 37–43. Small; pointed FA 41–48. Very large ears;
connecting process, sometimes short rounded lancet; long
curved forwards; tall, bluntly sella with broad base; narrow
pointed lancet; fur varies connecting process.
from grey to orange. Blyth’s
Horseshoe Bat, R. lepidus, (not 10. KING HORSESHOE BAT
illustrated) is similar but with longer upper canine, Rhinolophus rex, p. 231
does not overlap in range. FA 53–57. Enormous ears;
short rounded lancet; long,
5. CROSLET HORSESHOE BAT very broad sella expanded
Rhinolophus coelophyllus, into cup-shaped base; narrow
p. 232 connecting process.
FA 41–45. Short lancet;
thickened lobes covered with
long hairs enclosing base of
connecting process; broadly
arched connecting process. Note Diagrams beside each Rhinolophus face show
profile of noseleaf. See also diagrams of sella in Fig. 33.

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2c
2a

3b 3a 5

6
4

9
7

8
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 Plate 12: HORSESHOE BATS
1. INTERMEDIATE 4. CHINESE HORSESHOE BAT
HORSESHOE BAT Rhinolophus sinicus, p. 237
Rhinolophus affinis, p. 235 FA 44–55. Like R. thomasi, but
FA 48–54. Large; rounded averages larger; canines twice
connecting process; sella as long as upper premolar; wing
concave in middle; underparts relatively long and pointed.
slightly paler than upperparts.

2. THOMAS’S HORSESHOE 5. PENINSULAR HORSESHOE

BAT BAT
Rhinolophus thomasi, p. 236 Rhinolophus robinsoni, p. 237
FA 42–46. Rounded connecting FA 43–46. Medium-small;
process; lancet short, stubby rounded connecting process;
and with narrow point in middle; sella broad at base, narrowing
underparts only slightly paler abruptly in middle.
than upperparts; canines only
slightly longer than largest premolars. 6. CHASEN’S HORSESHOE
BAT
3. RUFOUS HORSESHOE BAT Rhinolophus chaseni, p. 236
Rhinolophus rouxii, p. 237 FA 43–46. Rounded connecting
FA 44–52. Like R. thomasi, but process; tall, triangular lancet;
averages larger; canines more sella broad at base, narrowing
than 30% longer than upper slightly in middle, rounded at
premolar; wing relatively short tip; underparts only slightly
and rounded. paler than upperparts.

Note Diagrams beside each Rhinolophus face show profile of noseleaf. See also diagrams of sella in Fig. 33.

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3 4

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 Plate 13: HORSESHOE BATS
1. LESSER BROWN 3. THAI HORSESHOE BAT
HORSESHOE BAT Rhinolophus thailandensis,
? Rhinolophus stheno, p. 235 p. 238
FA 43–48. Rounded connecting FA 56–61. Large; long woolly
process; tall, triangular lancet; fur; broadly rounded connecting
sella narrow, pointed at top; process; sella broad at base,
underparts distinctly paler than narrowing abruptly in middle.
upperparts. Indochinese Brown Chiew Kwee’s Horseshoe Bat,
Horseshoe Bat, R. microglobosus (not illustrated), is R. chiewkweeae (not illustrated), is similar but
smaller with higher frequency echolocation call. smaller, more orange fur, found only in Peninsular
Malaysia.
2. MALAYAN HORSESHOE
BAT 4. PEARSON’S HORSESHOE
Rhinolophus malayanus, p. 236 BAT
FA 38–44. Rounded connecting Rhinolophus pearsonii, p. 238
process; tall, triangular lancet; FA 49–55. Medium-sized; long
sella broad, squared at top; woolly fur; broadly rounded
underparts distinctly paler than connecting process; sella broad
upperparts. at base, narrowing abruptly in
middle.

Note Diagrams beside each Rhinolophus face show profile of noseleaf. See also diagrams of sella in Fig. 33.

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 Plate 14: LARGE ROUNDLEAF BATS
1. SHIELD-FACED 4. GREAT ROUNDLEAF BAT
ROUNDLEAF BAT Hipposideros armiger, p. 245
Hipposideros lylei, p. 246 FA 85–103. Very large;
FA 73–84. Large; two lateral relatively small anterior
leaflets; greatly enlarged noseleaf; narrow posterior
‘shield’ behind noseleaf in noseleaf; four lateral leaflets,
male; small ridges in female outer one very narrow; adult
(similar to 3b); posterior males have swollen area behind
noseleaf connected to anterior leaf at sides. noseleaf (4a); smaller or lacking in females (4b).
Griffin’s Roundleaf Bat, H. griffini, from Vietnam (not
2. SHIELD-NOSED illustrated) is similar but smaller (FA 83–90) with
ROUNDLEAF BAT higher frequency echolocation calls.
Hipposideros scutinares, p. 246
FA 78–83. Similar to H. lylei, 5. PENDLEBURY’S
but noseleaf slightly broader; ROUNDLEAF BAT
‘shield’ of male smaller Hipposideros pendleburyi, p. 245
(2a); female with very small FA 75–81. Similar to
shield (2b); lower frequency H. armiger, but smaller; males
echolocation calls. with swollen area behind
posterior noseleaf.
3. SWINHOE’S ROUNDLEAF
BAT 6. INTERMEDIATE
Hipposideros swinhoei, p. 246 ROUNDLEAF BAT
FA 81–89. Similar to H. lylei, Hipposideros cf. larvatus, p. 243
but averages larger; ‘shield’ FA 51–67. Medium-sized;
of male averages smaller three lateral leaflets; posterior
(3a); female with small ridges noseleaf broader than anterior
instead of shield (3b); posterior noseleaf. Includes several
noseleaf not joined to anterior leaf at sides. superficially similar species –
see text. Halong Roundleaf Bat, H. alongensis
(FA 66–76) in Vietnam, and Indian Roundleaf Bat,
H. lankadiva (FA 89–94) in Myanmar, have a
similar-shaped noseleaf with three lateral leaflets,
but are much larger (not illustrated).

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2a

3a 3b

4a 5

6a
4b
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 Plate 15: LARGE AND MEDIUM ROUNDLEAF BATS
1. BOONSONG’S ROUNDLEAF 5. ANNAMITE ROUNDLEAF
BAT BAT
Hipposideros lekaguli, p. 244 Hipposideros rotalis, p. 242
FA 71–79. Large; three lateral FA 45–49. Very large ears;
leaflets; posterior noseleaf enlarged, partly unpigmented
formed into three backward- noseleaf that covers muzzle;
facing pockets; intermediate internarial septum forms large
noseleaf with raised triangular disc that hides nostrils; deep
ridge in middle. notch in anterior noseleaf; one pair of lateral leaflets.

2. DIADEM ROUNDLEAF BAT 6. KHAOKHOUAY ROUNDLEAF

Hipposideros diadema, p. 244 BAT
FA 76–87. Large; dark fur with Hipposideros khaokhouayensis,
pale patches on shoulders and p. 243
edges of wings; three or four FA 46–49. Similar to H. rotalis,
lateral leaflets; females usually but internarial disc smaller,
more orange and buff (2b) than does not cover nostrils; lacks
males (2a). lateral leaflets.

3. RIDLEY’S ROUNDLEAF BAT 7. FAWN ROUNDLEAF BAT

Hipposideros ridleyi, p. 242 Hipposideros cervinus, p. 243
FA 47–51. Large ears; dark fur; FA 44–52. Two lateral leaflets;
very large, dark noseleaf that intermediate noseleaf narrower
covers muzzle; enlarged disc than posterior noseleaf; ears
on internarial septum that hides relatively narrow and triangular;
nostrils; no lateral leaflets. tail relatively short.

4. SMALL-DISC ROUNDLEAF 8. CANTOR’S ROUNDLEAF

BAT BAT
Hipposideros orbiculus, p. 242 Hipposideros galeritus, p. 243
FA 46–49. Similar to H. ridleyi, FA 45–50. Two lateral leaflets;
but slightly smaller ears and intermediate noseleaf wider
noseleaf; internarial disc than posterior noseleaf; ears
smaller, not completely covering very broad; tail relatively long.
nostrils.

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3b

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4

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 Plate 16: SMALL ROUNDLEAF BATS
1. GREATER BICOLOURED 4. DAYAK ROUNDLEAF BAT
ROUNDLEAF BAT Hipposideros dyacorum, p. 241
Hipposideros bicolor, p. 239 FA 38–43. No lateral leaflets;
FA 45–49. Fairly large ears; dark noseleaf; ears relatively
no lateral leaflets; relatively short and triangular; fur on
straight-sided internarial underparts dark.
septum. Echolocation call
c.131kHz. See text for
distinctions from H. nequam (not illustrated). 5. ASHY ROUNDLEAF BAT
Hipposideros cf. cineraceus,
2. LESSER BICOLOURED p. 240
ROUNDLEAF BAT (5a) widespread species;
Hipposideros atrox, p. 240 (5b) Malaysian species
FA 41–46. Similar to H. bicolor, FA 33–42. Small; short ears;
but averages shorter forearm; no lateral leaflets. Widespread
noseleaf darker and slightly species has internarial septum
curled at edges; echolocation swollen in middle, and higher frequency echolocation
call c.142kHz. call. Malaysian species has septum swollen only at
base. Myanmar Roundleaf Bat, H. einnaythu (not
3. LARGE-EARED illustrated), is similar, but slightly larger with dark
ROUNDLEAF BAT noseleaf, and one pair of narrow lateral leaflets.
Hipposideros pomona, p. 240
FA 38–45. Similar to H. bicolor,
but slightly smaller with larger
ears. Fur colour varies from
brown and white (3a) to all
orange (3b). Note that other
species are also sometimes orange.

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 Plate 17: SMALL ROUNDLEAF BATS
1. THAI ROUNDLEAF BAT 4. ASIAN TAILLESS
Hipposideros halophyllus, p. 241 ROUNDLEAF BAT
FA 35–38. Small; no lateral Coelops frithii, p. 247
leaflets; internarial septum with FA 34–44. No tail; ears small and
two kidney-shaped lobes in rounded, lacking ridges; anterior
centre that are usually darker noseleaf divided into two lobes;
than rest of noseleaf. lappets protruding from beneath
noseleaf long and narrow.
2. LEAST ROUNDLEAF BAT
Hipposideros doriae, p. 242 5. MALAYSIAN TAILLESS
FA 34–37. Small; dark fur; ROUNDLEAF BAT
no lateral leaflets; posterior Coelops robinsoni, p. 247
noseleaf lacks supporting septa. FA 34–37. Like C. frithii, but
averages smaller; lappets
under noseleaf are broad and
rounded.
3. COMMON TRIDENT BAT
Aselliscus stoliczkanus, p. 247
FA 39–45. Small; posterior
noseleaf raised into three
points; triangular pad in middle
of noseleaf; two lateral leaflets.
Dong Bac Trident Bat,
A. dongbacana (not illustrated),
is similar with slightly longer canines; only known
from N Vietnam.

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5b
5a

4a 4b

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 Plate 18: MYOTIS
1. HAIRY-FACED MYOTIS 5. PETERS’S MYOTIS
Myotis annectans, p. 254 Myotis ater, p. 252
FA 45–49. Medium-large; feet FA 34–39. Like M. muricola,
not enlarged; fur with dark but averages larger; fur darker;
bases and pale tips; often middle upper premolar smaller,
has orange patch on belly; largely intruded in toothrow.
middle upper premolars usually
missing.
6. THICK-THUMBED MYOTIS
2. LARGE INDOCHINESE Myotis rosseti, p. 254
MYOTIS FA 29–31. Small; greyish-
Myotis indochinensis, p. 253 brown; thickened pad at base
FA 43–46. Medium-large; of thumb (6b); only two upper
relatively small feet; dark brown and lower premolars.
fur; middle upper premolars
present, but small and intruded
in toothrow. See text for 7. RIDLEY’S MYOTIS
distinctions from M. montivagus and M. federatus Myotis ridleyi, p. 255
(not illustrated). FA 27–32. Small; heavy body;
dark grey-brown fur; only two
3. ASIAN WHISKERED upper and lower premolars.
MYOTIS
Myotis muricola, p. 251
FA 33–37. Medium-small; feet
small; wing membrane inserted 8. BLACK-AND-ORANGE
at bases of toes (3b); middle WOOLLY MYOTIS
upper premolar slightly intruded Myotis rufoniger, p. 250
in toothrow; underparts FA 45–56. Moderately large;
with buffy to pale brown tips to fur. See text for orange fur; black and orange
distinctions from M. annatessae (not illustrated). wings; black rims to ears.
Herman’s Woolly Myotis,
4. EURASIAN WHISKERED M. hermani (not illustrated),
MYOTIS is similar but larger (FA 56–60).
Myotis mystacinus, p. 252
FA 34–37. Like M. muricola, 9. HODGSON’S WOOLLY
but paler with whitish tips to fur MYOTIS
on underparts. Myotis formosus, p. 251
FA 45–53. Moderately large;
yellowish fur; black and yellow
wings; no black rims to ears.

Note Myotis can be distinguished from other bats by ear and tragus shape, with narrow tragus angled slightly forwards
and tapering to tip. Most species have three upper and lower premolars on each side.

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3b

2
1

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6b

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 Plate 19: MYOTIS
1. RICKETT’S MYOTIS 5. HORSFIELD’S MYOTIS
Myotis pilosus, p. 256 Myotis horsfieldii, p. 255
FA 53–56. Large; greatly FA 35–38. Feet enlarged;
enlarged feet with large claws; wing membrane inserted
wing membrane inserted at on side of foot (5b); middle
ankle; short fur. upper premolars only slightly
displaced inwards; long
canines; low forehead.
2. INDOCHINESE WATER
MYOTIS 6. CHINESE MYOTIS
Myotis laniger, p. 257 Myotis chinensis, p. 250
FA 31–36. Small; feet slightly FA 65–69. Largest Myotis;
enlarged with membrane large ears; well-developed
inserted on side of foot (2b); middle premolar.
long narrow ears; high forehead;
moderately small canines.
See text for distinctions from M. annamiticus and
M. phanluongi (not illustrated). 7. SZECHUAN MYOTIS
Myotis altarium, p. 250
3. SMALL-TOOTHED MYOTIS FA 43–46. Moderately large;
Myotis siligorensis, p. 256 long ears; foot quite long; wing
FA 30–35. Small; light brown membrane inserted at base of
fur; feet slightly enlarged; toe; middle premolars small but
high forehead; middle upper within toothrow.
premolar not reduced; small
canines.

4. HASSELT’S MYOTIS
Myotis hasseltii, p. 255
FA 38–43. Feet enlarged; wing
membrane inserted at base
of foot (4b); middle premolars
small, intruded inwards.

Note Myotis can be distinguished from other bats by ear and tragus shape, with narrow tragus angled slightly forwards
and tapering to tip. Most species have three upper and lower premolars on each side.

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2a

4a

4b

5a

7 5b

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 Plate 20: PIPISTRELLES
1. JAVAN PIPISTRELLE 5. NARROW-WINGED
Pipistrellus javanicus, p. 259 PIPISTRELLE
FA 30–36. Dark brown fur with Pipistrellus stenopterus, p. 261
darker bases; tragus narrow FA 37–42. Relatively large; fur
with rounded tip; anterior upper short, reddish-brown to dark
premolar slightly displaced brown; fifth finger short;
inwards; forehead slightly forehead sloping or slightly
domed; canine with small domed.
secondary cusp; male with medium-length penis.
6. COROMANDEL
2. JAPANESE PIPISTRELLE PIPISTRELLE
Pipistrellus abramus, p. 260 Pipistrellus coromandra, p. 260
FA 29–33. Similar to FA 26–34. Fur dark brown
P. javanicus, but fur greyish- to reddish-brown; forehead
brown, male with very long sloping evenly; canine with
penis. distinct secondary cusp.

7. LEAST PIPISTRELLE
3. KELAART’S PIPISTRELLE Pipistrellus tenuis, p. 261
Pipistrellus ceylonicus, p. 261 FA 25–31. Small; fur mid-
FA 33–42. Relatively large; brown to dark brown; ear
fur varies from grey-brown to relatively short; tragus narrow;
red-brown; forehead sloping or skull with narrow rostrum,
slightly domed. evenly sloping profile.

4. NARROW-WINGED BROWN
BAT
Philetor brachypterus, p. 263
FA 30–36. Dark brown fur;
fifth finger very short; head
flattened; ears large and
broadly triangular; short tragus;
only one upper premolar;
complex genitalia.

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2 3

4 5

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 Plate 21: PIPISTRELLES
1. MOUNT POPA PIPISTRELLE 4. CADORNA’S PIPISTRELLE
Pipistrellus paterculus, p. 260 Hypsugo cadornae, p. 265
FA 27–34. Fur uniform brown FA 33–37. Reddish-brown fur;
above, sandy brown tips below; broad ears; short, broad tragus;
tragus relatively short; male anterior upper premolar very
with extremely long penis. small.

2. BIG-EARED PIPISTRELLE 5. CHOCOLATE PIPISTRELLE

Hypsugo macrotis, p. 264 Falsistrellus affinis, p. 265
FA 33–34. Large ears; reddish- FA 36–38. Long, soft dark
brown fur; partially translucent fur with frosted tips; pale
whitish wings; anterior upper underparts; broad, rounded
premolar very small, displaced ears; tragus broad, pointed in
inwards; forehead sloping. front; narrow muzzle.

3. CHINESE PIPISTRELLE
Hypsugo pulveratus, p. 263
FA 32–37. Dark fur with pale
tips; short narrow tragus;
anterior upper premolar
moderate size, displaced
inwards; forehead domed.
See text for descriptions of
Hypsugo lophurus, H dolichodon and H. joffrei
(not illustrated).

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2a
3

2b

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 Plate 22: MIXED VESPERTILIONID BATS
1. INDOCHINESE THICK- 5. DORIA’S FALSE-SEROTINE
THUMBED PIPISTRELLE Hesperoptenus doriae, p. 269
Glischropus bucephalus, p. 263 FA 38–41. Blackish-brown fur;
FA 32–36. Slightly shaggy rounded profile; only one upper
fur; pink pad on sole of foot premolar; first upper incisor
(1b); triangular, pink thickened large and conical, second
pad at base of thumb (1c); very small, but not displaced
second upper incisor displaced inwards.
outwards. Sunda Thick-thumbed Pipistrelle,
G. tylopus (not illustrated), is smaller, has more 6. TICKELL’S FALSE-
sloping forehead, and does not overlap in range. SEROTINE
Hesperoptenus tickelli, p. 270
2. MAINLAND GREATER FA 49–56. Light yellowish-
BAMBOO BAT brown fur and wing bones;
Tylonycteris malayana, p. 271 dark membranes; first upper
FA 26–30. Very flat skull; incisor large and conical;
flattened broad body; rounded second incisor small, displaced
grey or pinkish pads on foot inwards.
(2b) and base of thumb (2c); fur
sleek and smooth, dark brown 7. TOMES’S FALSE-
with frosted pale tips; thickened skin at base of ear. SEROTINE
Tonkin Greater Bamboo Bat, T. tonkinensis, from N. Hesperoptenus tomesi, p. 269
Vietnam and Laos (not illustrated) looks very similar FA 50–53. Large; dark
but is genetically distinct. blackish-brown fur; rounded
forehead; first upper incisor
3. MAINLAND LESSER large and conical, second
BAMBOO BAT incisor small, displaced
Tylonycteris fulvida, p. 271 inwards.
FA 24–28. Like T. malayana,
but smaller; fur shorter and 8. DISC-FOOTED BAT
fluffier, usually more reddish; Eudiscopus denticulus, p. 257
skin at base of ear thinner. FA 35–38. Myotis-shaped
ears; pale fur; flattened skull;
4. LEAST FALSE-SEROTINE enlarged disc-like pads on hind
Hesperoptenus blanfordi, p. 269 feet.
FA 24–29. Short forearm, but
heavy body; smooth, glossy
dark brown fur; thickened dark 9. GREAT EVENING BAT
pad at base of thumb; sloping Ia io, p. 269
skull profile making head FA 71–79. Very large; broad,
pointed; second upper incisor triangular ears; two upper
displaced inwards. premolars, anterior very small.

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 Plate 23: MIXED VESPERTILIONID BATS
1. HARLEQUIN BAT 6. ASIAN SEROTINE
Scotomanes ornatus, p. 270 Eptesicus pachyomus, p. 267
FA 54–61. Orange fur with FA 53–57. Large; broad muzzle
prominent white marks on top with sloping profile; ears
of head, shoulders and middle narrow, somewhat triangular at
of back; only one upper incisor top; narrow tragus; two upper
on each side. incisors, but only one upper
premolar on each side.
2. FAR-EASTERN
BARBASTELLE 7. ASIAN NOCTULE
Barbastella sp., p. 258 Nyctalus cf. labiata, p. 262
FA 38–43. Ears broad, facing FA 49–58. Uniform brown to
forwards, joined at base over dark brown fur; broad ears;
forehead. short tragus; narrow wings with
short fifth finger; two upper
incisors on each side.
3. GOLDEN-COLLARED BAT
Thainycteris aureocollaris, 8. LESSER ASIAN HOUSE BAT
p. 266 Scotophilus kuhlii, p. 270
FA 47–51. Fur with black FA 45–52. Short, yellowish-
bases, buff to orange tips; brown fur; long, narrow tragus
yellowish fur forms collar under bent forwards; only one upper
chin. incisor and one upper premolar
on each side.
4. BENOM GILDED
PIPISTRELLE 9. GREATER ASIAN HOUSE
Arielulus societatis, p. 266 BAT
FA 35–38. Fur black with Scotophilus heathii, p. 271
reddish-orange tips on the FA 57–69. Like S. kuhli, but
back, golden yellow tips on larger with longer forearm; fur
underparts and head; pale rims varies from yellow-brown to
to ears and tragus; posterior dark orange-brown.
upper molar small.

5. BLACK GILDED
PIPISTRELLE
Arielulus circumdatus, p. 266
FA 40–44. Like A. societatis,
but larger; posterior upper
molar larger with distinct
second cusp.

Note See text for identification of Eptesicus pachyotis (FA 38–39), Cassistrellus dimissus (FA 38–42) and Cassistrellus
yokdonensis (FA c.48).

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 Plate 24: TUBE-NOSED BATS
1. HUTTON’S TUBE-NOSED BAT 6. PENINSULAR TUBE-NOSED
Murina huttoni, p. 275 BAT
FA 32–37. Fur orange-brown Murina peninsularis, p. 273
with dark bases; long ears; both FA 34–41. Back fur orange to
premolars large; upper molars reddish-brown, usually with
without indentation; lower molars small grey bases; underside
with well-developed posterior paler, often with orange
section. Annamite Tube-nosed tinge; dentition similar to
Bat, M. annamitica (not illustrated), has similar M. cyclotis. Guillén’s Tube-nosed Bat, M. guilleni
dentition, but is more orange and smaller (FA 29–34). (not illustrated), in peninsular Thailand is similar but
smaller (FA 32–36); underside usually greyer.
2. FEA’S TUBE-NOSED BAT
Murina feae, p. 276 7. ROZENDAAL’S TUBE-
FA 27–34. Upperparts greyish- NOSED BAT
brown fur with four alternating Murina rozendaali, p. 276
dark-light bands; underparts FA 28–32. Upperparts with
greyish-white with dark bases; dark bases, shiny golden tips;
anterior upper premolar small. underparts white to base; both
upper premolars similar in size;
3. BROWN TUBE-NOSED BAT canine long.
Murina suilla, p. 276
FA 27–32. Upperparts brown to 8. GILDED TUBE-NOSED BAT
orange-brown with grey bases; Murina eleryi, p. 277
underparts greyish-white FA 27–31. Hair of upperparts
with darker bases; anterior alternating dark-pale bands
upper premolar small; canine with golden tips; underparts
relatively short. with long black bases, white
tips; anterior upper premolar
4. ROUND-EARED TUBE- small; canines short. Bala Tube-
NOSED BAT nosed Bat, M. balaensis (not illustrated), similar, but
Murina cyclotis, p. 272 restricted to peninsular Thailand and Malaysia.
FA 29–34. Back fur orange
to reddish-brown, with greyer 9. GREATER TUBE-NOSED
base; both premolars large; BAT
upper molars with indentation Murina leucogaster, p. 279
on outer edge; lower molars FA c.42. Large; upperparts
with reduced posterior section. reddish-brown with dark bases;
underparts yellowish-white;
5. BRONZED TUBE-NOSED anterior upper premolar small,
BAT about half height of posterior
Murina aenea, p. 275 premolar.
FA 34–38. Fur with dark brown
bases, tips shiny golden or
bronze on upperparts, buff on Note Tube-nosed bats, including Hairy-winged Bat (see
underparts; dentition similar to Pl. 26), are distinguished from other bats by enlarged
that of M. cyclotis. tubular nostrils, and somewhat rounded ears with a long,
pointed tragus. See text for diagrams of teeth.

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5

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 Plate 25: TUBE-NOSED BATS
1. HARRISON’S TUBE-NOSED 6. LORELIE’S TUBE-NOSED
BAT BAT
Murina harrisoni, p. 275 Murina lorelieae, p. 279
FA 34–40. Medium to large; FA 33–35.5. Upperparts
upperparts uniform reddish- largely reddish-brown,
brown to base; underparts hairs with dark grey bases;
white; large anterior premolars underparts greyish white
and canines; lower molars with with dark grey bases. Canine
well-developed posterior section. relatively short; anterior upper premolar about
¾ height of posterior premolar.
2. FIONA’S TUBE-NOSED BAT
Murina fionae, p. 274 7. GOLDEN-HAIRED TUBE-
FA 34–38.5. Long fur; upper- NOSED BAT
parts with buff bases and Murina chrysochaetes, p. 278
orange-brown tips; underparts FA 28–30. Fur of upperparts
uniformly pale, whitish or buff banded black, yellow and
orange. Large anterior premolars brown; guard hairs with golden
and canines; lower molars with tips; fur of underparts black with
reduced posterior section. gold tips, whiter on chin.

3. WALSTON’S TUBE-NOSED 8. DA LAT TUBE-NOSED BAT

BAT Murina harpioloides, p. 278
Murina walstoni, p. 277 FA 29–30. Upperparts dark
FA 30–35. Upperpart hairs brown, with long orange-gold
brown to orange-brown with tipped guard hairs; underside
white bases; underparts all dark bases, whitish tips. Anterior
white. Wing and tail membranes upper premolar small.
largely without hair. Anterior
upper premolar small. 9. EVEN-TOOTHED TUBE-
NOSED BAT
4. BEELZEBUB TUBE-NOSED Harpiola isodon, p. 280
BAT FA 31–36. Upperparts long fur,
Murina beelzebub, p. 277 banded dark brown and orange
FA 34–38. Overall very dark; with gold tips; underparts dark
hairs black with a grey band, bases with orange-brown tips.
grey frosting; underparts dark Anterior upper premolar larger
bases, pale tips. Anterior upper than posterior premolar.
premolar small.

5. KONTUM TUBE-NOSED
BAT
Murina kontumensis, p. 279
FA c.32. Upperparts brownish
grey, with long yellow-tipped Note Tube-nosed bats, including Hairy-winged Bat (see
guard hairs; underparts Pl. 26), are distinguished from other bats by enlarged
brownish grey. Long ears. tubular nostrils, and somewhat rounded ears with a long,
Anterior upper premolar small. pointed tragus. See text for diagrams of teeth.

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 Plate 26: BENT-WINGED BATS AND HAIRY-WINGED BAT
1. LARGE BENT-WINGED BAT 4. MEDIUM BENT-WINGED
Miniopterus magnater, p. 285 BAT
FA 47–53. Third finger of wing Miniopterus medius, p. 286
with last bone about three times FA 38–44. Intermediate in size
longer than second last; short, between M. fuliginosus and
broad ear with short tragus; fur M. pusillus, especially skull
typically dark brown to blackish, measurements.
but sometimes with reddish
patches or all reddish. 5. HAIRY-WINGED BAT
Harpiocephalus harpia, p. 280
2. SMALL BENT-WINGED BAT FA 45–51. Similar to Murina,
Miniopterus pusillus, p. 286 but larger with greatly reduced
FA 40–43. Like M. magnater, third upper molar; fur varies
but much smaller; relatively from bright orange with dark
short, narrow skull. bases to reddish-brown with
grey bases; extensive hairs
on tail membrane and bases
of wings.
3. EASTERN BENT-WINGED
BAT
Miniopterus fuliginosus, p. 285
FA 42–46. Like M. magnater,
but somewhat smaller.

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 Plate 27: KERIVOULA AND PHONISCUS
1. CLEAR-WINGED WOOLLY 6. WHITEHEAD’S WOOLLY
BAT BAT
Kerivoula pellucida, p. 283 Kerivoula whiteheadi, p. 284
FA 29–32. Light grey-brown FA 28–29. Fur of upperparts
to orange-brown fur with paler light brown with dark bases;
bases; long, narrow ears; underparts whitish; elongate
translucent pale wings; very premolars.
high, domed forehead.
7. LEAST WOOLLY BAT
2. PAINTED WOOLLY BAT Kerivoula minuta, p. 284
Kerivoula picta, p. 283 FA 25–29. Very small; long
FA 32–39. Light orange fur; fluffy fur orange-brown with
wings with distinctive black and dark bases; small ears; rounded
orange pattern. premolars.

8. SMALL WOOLLY BAT

3. KRAU WOOLLY BAT Kerivoula intermedia, p. 283
Kerivoula krauensis, p. 283 FA 26–31. Very similar to
FA 29–33. Small; dark K. minuta, but slightly larger,
blackish-brown fur with narrow, especially skull and body mass.
shiny golden tips on back, pale
tips below.

9. LESSER GROOVE-
4. HARDWICKE’S WOOLLY TOOTHED BAT
BAT Phoniscus atrox, p. 285
Kerivoula hardwickii, p. 282 FA 31–35. Fur with four bands
FA 29–34. Grey-brown to brown of colour ending with golden
fur; fairly large ears. See text for tips; tragus white; canine long
distinctions from K. titania, with groove on outside; middle
K. depressa, K. furva and upper premolar rounded.
K. dongduongana (not illustrated).
10. GREATER GROOVE-
5. PAPILLOSE WOOLLY BAT TOOTHED BAT
Kerivoula papillosa, p. 281 Phoniscus jagorii, p. 284
FA 39–49. Large; domed FA 35–42. Similar to P. atrox,
forehead; orange-brown fur. but larger; middle upper
K. lenis (not illustrated) is similar premolar elongate.
but smaller; K. kachinensis (not
illustrated) has a much flatter
skull (see text).

Note Kerivoula and Phoniscus are distinguished from other bats by funnel-shaped ears, long, narrow tragus and three
well-developed upper and lower premolars on each side.

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7

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 Plate 28: FREE-TAILED BATS
1. LA TOUCHE’S FREE- 4. ASIAN WRINKLE-LIPPED
TAILED BAT BAT
Tadarida latouchei, p. 286 Chaerephon plicatus, p. 286
FA 53–55. Moderately FA 40–50. Moderately small;
large; short dense fur; thick ears joined on top of head by
protruding tail; ears joined by narrow band of skin at base
narrow fold of skin at base. (4b); underside pale; two upper
premolars.
2. SUNDA FREE-TAILED BAT
Mops mops, p. 287 5. WROUGHTON’S FREE-
FA 41–46. Medium-sized; thick TAILED BAT
dark fur above and below; ears Otomops wroughtoni, p. 288
joined by narrow band of skin FA 63–67. Large; pale marks
(2b); only one upper premolar. on back of neck; ears long,
joined for much of length by
fold of skin.
3. JOHORE WRINKLE-LIPPED
BAT 6. NAKED BAT
Chaerephon johorensis, p. 287 Cheiromeles torquatus, p. 288
FA 44–49. Medium-sized; ears FA 74–86. The world’s heaviest
joined by broad band of skin insectivorous bat; body largely
that extends backwards on lacking fur; dark grey body skin;
top of head to form a pocket thick, protruding tail; ears large,
(3b); underside pale; two upper well separated.
premolars.

Note All free-tailed bats have a tail that protrudes beyond the tail membrane, and narrow wings.

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2b

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 Plate 29: PANGOLINS, COLUGO AND LORISES
1. SUNDA PANGOLIN 4. SUNDA SLOW LORIS
Manis javanica, p. 192 Nycticebus coucang, p. 289
HB 400–650. Scaly; front claws HB 260–300. Large eyes; no
50% longer than hind claws; tail; colour varies from grey-
long tail with 30 or more scales brown to orange-brown; dark
along its side; small ears. stripe on back branches to
connect to ears and eyes.

2. CHINESE PANGOLIN 5. ASIAN SLOW LORIS

Manis pentadactyla, p. 192 Nycticebus bengalensis, p. 289
HB 400–580. Scaly; front claws HB 300–380. Grey-brown to
twice as long as hind claws; orange-brown; dark stripe on
conspicuous ear lobes; short back ends on back of nape.
tail with <20 scales along its
side.

3. SUNDA COLUGO 6. PYGMY LORIS

Galeopterus variegatus, p. 211 Nycticebus pygmaeus, p. 289
HB 330–420. Grey (3a) or HB 210–290. Small; relatively
reddish (3b) with complex large ears; usually orange-
pattern of pale and dark marks; brown; dorsal stripe varies from
extensive gliding membrane narrow and inconspicuous to
encloses tail. broad.

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 Plate 30: PRESBYTIS AND TRACHYPITHECUS LANGURS
1. PALE-THIGHED LANGUR 4. ANNAMESE SILVERED
Presbytis siamensis, p. 290 LANGUR
HB 430–690. Upperparts vary Trachypithecus margarita, p. 292
from greyish to brown; tail, feet HB 500–600. Fur medium to
and hands dark; pale below; dark grey with pale frosting;
whitish patch on outer thigh. contrasting darker feet and
Infants with ‘cruciform’ pattern arms; paler underparts; dark
– pale except for dark stripe up face except for pinkish rings
back and across arms. around eyes. Infants orange.

2. BANDED LANGUR 5. INDOCHINESE SILVERED

Presbytis femoralis, p. 290 LANGUR
HB 460–590. Upperparts dark Trachypithecus germaini, p. 291
brown to blackish; underside HB 490–570. Dark grey above
pale. Infants ‘cruciform’. P. f. and below; long, pale hairs
femoralis (2a) very dark above, around face; dark face. Infants
sometimes with reddish tone; orange.
sharply demarcated white areas
on underparts. P. f. robinsoni (2b) dark brown to
blackish above; indistinctly greyish below, with
white insides of thighs.

3. SUNDAIC SILVERED
LANGUR
Trachypithecus cristatus, p. 291
HB 415–540. Fur long, dark
grey above and below, with
silvery tips; dark face. Infants
with bright orange fur.

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 Plate 31: TRACHYPITHECUS LANGURS
1. CAPPED LANGUR 4. INDOCHINESE GREY
Trachypithecus pileatus, p. 294 LANGUR
HB 500–700. Distinct black cap Trachypichecus crepusculus,
on top of head; dark face; grey p. 293
above; underparts and facial HB 520–620. Upperparts mid
hairs contrasting pale yellowish to light grey, often frosted;
(1a) or orange (1b). underparts contrastingly pale;
arms and legs same colour as
2. SHORTRIDGE’S LANGUR back; feet and hands usually darker; complete pale
Trachypithecus shortridgei, rings around eyes.
p. 295
HB 600–750. Silvery grey 5. TENASSERIM LANGUR
above and below; dark feet Trachypithecus barbei, p. 292
and end of tail; dark face; pale HB 500–700. Fur dark greyish-
orange-yellow eyes. black without pale frosting; grey
tail; dark face with pale mouth;
3. PHAYRE’S LANGUR pale rings around eyes.
Trachypithecus phayrei, p. 293
HB 520–620. Upperparts
uniformly brown to grey; 6. DUSKY LANGUR
underparts vary from Trachypithecus obscurus, p. 292
contrastingly paler to similar to HB 500–700. Body dark grey or
back; hands and feet usually greyish-brown with paler crown,
same colour as back; face dark legs and tail; dark face with
with pale skin on lips; white marks around eyes contrasting pale marks around
form incomplete eye-ring. mouth; pale rings around eyes,
often incomplete.

Note Infants of all Trachypithecus spp. are yellowish to orange, as in Dusky Langur.

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 Plate 32: FRANCOIS’S LANGUR AND RELATIVES
1. FRANCOIS’S LANGUR 4. DELACOUR’S LANGUR
Trachypithecus francoisi, p. 293 Trachypithecus delacouri, p. 294
HB 470–630. Largely black; HB 570–730. Largely black
white stripe from side of mouth body; indistinct white stripe on
to edges of ears. Infants pale face; large white patch on thigh
orange; immatures may be and rump.
black with pale orange head.

2. CAT BA LANGUR 5. HATINH LANGUR

Trachypithecus poliocephalus, Trachypithecus hatinhensis,
p. 293 p. 294
HB 490–590. Dark brownish- HB 500–660. Similar to
black body; pale yellowish to T. francoisi, but white facial
orange head and shoulders; stripes continue behind ear to
pale area on rump. back of head; immatures may
have pale forehead, similar to
3. INDOCHINESE BLACK T. laotum.
LANGUR
Trachypithecus ebenus, p. 294 6. LAO LANGUR
HB 600–700. All glossy black Trachypithecus laotum, p. 294
above and below. HB 460–530. Like T. francoisi,
but white forehead.

Note In all species, infants are orange; as they change into adult patterns, the head may have more extensive pale
areas than an adult’s and may resemble that of other species.

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 Plate 33: SNUB-NOSED MONKEYS
1. TONKIN SNUB-NOSED 2. MYANMAR SNUB-NOSED
MONKEY MONKEY
Rhinopithecus avunculus, Rhinopithecus strykeri, p. 296
p. 296 HB c.555. Flattened face and
HB 510–620. Flattened face nose with thick pinkish lips,
with pale eye-rings; thick pale pink bare skin on face and
pinkish lips; black back; white around eyes. Fur black all over,
to brownish-white head and sometimes slightly browner
underparts; long, white-tipped below, except white hairs on
tail. chin and ears; long, dark tail.

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 Plate 34: DOUCS
1. GREY-SHANKED DOUC 3. BLACK-SHANKED DOUC
Pygathrix cinerea, p. 296 Pygathrix nigripes, p. 295
HB 610–760. Arms pale grey; HB 610–760. Black legs; blue-
legs medium to dark grey; grey facial skin; black on sides
yellow-brown face skin; white of face.
on sides of face.

2. RED-SHANKED DOUC
Pygathrix nemaeus, p. 295
HB 610–760. Reddish lower
legs; black shoulders; white
lower arms; yellow-brown face
skin; white on sides of face.

Note All doucs are distinguished by a white rump and a very long white tail with a tassel at the end. Hybrids have
sometimes been reported, and may show intermediate features.

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 Plate 35: MACAQUES
1. RHESUS MACAQUE 4. NORTHERN PIG-TAILED
Macaca mulatta, p. 298 MACAQUE
HB 470–585. Medium build; Macaca leonina, p. 297
medium-length tail; rump and HB 470–585. Short tail, often
upper thigh reddish; hairs on held curved back; narrow
crown directed backwards. diagonal red lines on face;
brown to golden-brown back;
whitish face.
2. LONG-TAILED MACAQUE
Macaca fascicularis, p. 298 5. SOUTHERN PIG-TAILED
HB 450–550. Medium build; MACAQUE
long tail; upperparts uniformly Macaca nemestrina, p. 297
grey-brown or reddish-brown; HB 470–585. Short tail, often
limited overlap in range with curled forwards; back and
M. mulatta. crown usually dark brown or
blackish; body varies from grey-
3. ASSAMESE MACAQUE brown to orange-brown.
Macaca assamensis, p. 298
HB 510–735. Thickset, heavy 6. STUMP-TAILED MACAQUE
build; medium-length tail; Macaca arctoides, p. 297
greyish hindquarters; hairs on HB 485–635. Very heavy build;
crown parted in middle. very short tail usually held down
and not visible; long shaggy
fur; facial skin dark pink to
bright red.

Note In all macaques, adult males are distinctly larger and heavier than females.

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 Plate 36: HYLOBATES GIBBONS AND SIAMANG
1. WHITE-HANDED GIBBON 3. PILEATED GIBBON
Hylobates lar, p. 300 Hylobates pileatus, p. 300
HB 450–600. Fur of both HB 470–600. Adult male (3b)
sexes varies from brown (1a) to black with white hands and
blonde (1b) to black (1c); hands face; female (3a) greyish-white
and feet contrasting white; with black on belly and top of
white ring around face. head; young all greyish-white.

2. AGILE GIBBON 4. SIAMANG

Hylobates agilis, p. 300 Symphalangus syndactylus,
HB 450–650. Hands same p. 299
colour as feet; white on HB 750–900. Much larger than
forehead does not form other gibbons; all black; both
complete ring around face. sexes have throat pouch that
Usually black (2b), but inflates when calling; very loud,
occasionally blonde (2a). distinctive call.

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1a

1b

2a

3b 2b

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 Plate 37: CRESTED GIBBONS AND HOOLOCK GIBBON
1. BUFF-CHEEKED GIBBON 4. WESTERN BLACK-
Nomascus gabriellae, p. 302 CRESTED GIBBON
HB c.500. Male (1b) largely Nomascus concolor, p. 301
black with short buff to reddish HB c.500. Male (4b) all black;
patches on cheeks that do not female (4a) buffy with black
reach ears; brownish patch crown, extensive black on
on chest. Female (1a) largely underside. Eastern Black-
buffy-brown to orange-brown; crested Gibbon, N. nasutus (not
black only on crown; no white on face. illustrated), in NE Vietnam very similar, but female
lacks black on underparts.
2. SOUTHERN WHITE-
CHEEKED GIBBON 5. HOOLOCK GIBBON
Nomascus siki, p. 302 Hoolock hoolock, p. 301
HB c.500. Both sexes similar HB 450–650. Male (5b)
to N. leucogenys, however largely black; white eyebrows
male (shown) has white cheek separated (as shown) or joined;
patches extending only to lower white whiskers on chin. Female
edges of ears, but touching (5a) varies from buffy to brown;
mouth at bottom; relatively short hair. darker brown on underparts
and cheeks; white eyebrows and thin white line
3. NORTHERN WHITE- around eyes and face.
CHEEKED GIBBON
Nomascus leucogenys, p. 302
HB c.500. Male (3b) all black
with white patches on cheeks
that extend to tops of ears, but
do not touch corner of mouth.
Female (3a) buffy with black
crown, white ring on face.

Note All crested gibbons (Nomascus spp.) have longer hairs in the middle of the head that form a distinct crest.

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1a 1b

4a

3a

5a

5b

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 Plate 38: WILD DOGS
1. GOLDEN JACKAL 3. RACCOON DOG
Canis aureus, p. 303 Nyctereutes procyonoides,
HB 600–800. Golden-brown to p. 304
greyish-brown; dark hairs on HB 500–680. Yellowish-brown
back form saddle; pointed ears; to grey-brown with extensive
tail with dark tip. black hairs; black mask on
face; relatively short tail.

2. RED FOX 4. DHOLE

Vulpes vulpes, p. 304 Cuon alpinus, p. 305
HB 550–700. Slender body; HB 800–1,050. Large; reddish-
pointed face; pointed ears; brown with pale areas below;
usually reddish-brown, but long, bushy tail mostly or all
sometimes dark brown; tail black; rounded ears with white
bushy. centres.

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 Plate 39: RED PANDA AND BEARS
1. RED PANDA 3. SUN BEAR
Ailurus fulgens, p. 306 Helarctos malayanus, p. 305
HB 510–630. Reddish fur; HB 1.1–1.4m. Short fur; small
white and dark markings on rounded ears; buffy U- or
face; long, bushy banded tail. V-shaped mark on chest; pale
muzzle; holds head low.

2. ASIAN BLACK BEAR

Ursus thibetanus, p. 306
HB 1.2–1.5m. Large; long
shaggy hair; large erect ears;
white V on chest; muzzle
largely dark.

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 Plate 40: MARTENS AND BADGERS
1. YELLOW-THROATED 4. LARGE-TOOTHED FERRET-
MARTEN BADGER
Martes flavigula, p. 307 Melogale personata, p. 310
HB 450–650. Long, slender HB 330–390. Similar to
body with long dark tail; dark M. moschata, but white stripe
stripe on side of neck bordering usually extends down back;
yellowish chin, throat and tail averages longer with more
chest. Considerable geographic extensive white tip; typically
variation in colour: back varies from pale yellowish- less black on face; larger teeth.
brown with darker head (as shown) to uniformly
rich reddish-brown. 5. GREATER HOG BADGER
Arctonyx collaris, p. 310
2. STONE MARTEN HB 550–700. Large; long pig-
Martes foina, p. 307 like snout; dark stripes on face
HB 400–540. Brown with variable in extent, black or dark
reddish, yellowish or grey brown; short tail; body colour
tinge; throat white or yellowish varies from greyish to brown.
with variable dark patches; tail
relatively short.

3. SMALL-TOOTHED FERRET-
BADGER
Melogale moschata, p. 310
HB 330–380. Black-and-white
head pattern; body fur varies
from light brown to greyish;
medium-length tail with only
tip white; white stripe on top of
head does not extend down back; small teeth.

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 Plate 41: WEASELS
1. TONKIN WEASEL 4. YELLOW-BELLIED WEASEL
Mustela tonkinensis, p. 308 Mustela kathiah, p. 308
HB c.200. Small; relatively HB 200–290. Dark brown
short tail; brown fur with white upperparts; contrasting
underparts. yellowish or light orange
underparts; tail coloured as
back; no mask.

2. MALAY WEASEL 5. STRIPE-BACKED WEASEL

Mustela nudipes, p. 308 Mustela strigidorsa, p. 309
HB 300–360. Body golden- HB 275–325. White stripe
brown to bright orange; head down middle of back; body
and tip of tail white; long colour dark brown to reddish-
bushy tail. brown; pale on throat and
centre of belly; bushy tail.

3. SIBERIAN WEASEL
Mustela sibirica, p. 308
HB 250–390. Dark mask
on face; body colour highly
variable, ranging from golden-
brown to reddish to dark brown;
underparts similar except for
pale chin.

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 Plate 42: OTTERS
1. EURASIAN OTTER 3. HAIRY-NOSED OTTER
Lutra lutra, p. 311 Lutra sumatrana, p. 311
HB 550–720. Fur dense with HB 500–800. Upperparts
longer, paler guard hairs giving including sides of neck dark
grizzled effect; W-shaped brown; irregular white patches
border between hair on face on chin and throat; tail rounded
and bare skin on tip of nose; tip in cross-section; tip of nose
of tail rounded in cross-section. covered with hair.

2. ORIENTAL SMALL- 4. SMOOTH OTTER

CLAWED OTTER Lutrogale perspicillata, p. 312
Amblonyx cinereus, p. 312 HB 650–750. Pale throat and
HB 360–550. Small; dark sides of neck; smooth shiny
brown to greyish-brown with fur; tail somewhat flattened in
pale throat and sides of neck; cross-section; straight border
small claws. between bare skin on nose and
fur on face.

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 Plate 43: CIVETS
1. SMALL INDIAN CIVET 3. LARGE-SPOTTED CIVET
Viverricula indica, p. 313 Viverra megaspila, p. 313
HB 530–640. Relatively slender HB 720–850. Black stripe down
with short legs; numerous dark back; irregular pattern of large
spots forming rows along back dark spots on sides; tail largely
and sides; tail with 6–9 discrete black with incomplete white
dark bands, pale tip. bands at base; bold black-and-
white throat pattern.
2. MALAY CIVET
Viverra tangalunga, p. 313 4. LARGE INDIAN CIVET
HB 610–670. Black stripe down Viverra zibetha, p. 313
back; tail with about 15 black HB 750–850. Black stripe down
bands joined on top; black-and- back does not continue on tail;
white markings on throat; many sides irregularly mottled grey
small spots on sides. and black; tail with five or six
broad black bands separated
by complete white bands; bold
black-and-white throat pattern.

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 Plate 44: PALM CIVETS AND LINSANGS
1. MASKED PALM CIVET 4. SPOTTED LINSANG
Paguma larvata, p. 315 Prionodon pardicolor, p. 319
HB 510–760. Colour highly HB 350–370. Long and slender
variable from light brown to with long tail; irregular pattern
dark brown or reddish; no dark of black splotches on body that
spots or stripes; dark mask on usually do not form dark bands
face; white cheeks; broad white except on neck; tail with eight
stripe from forehead to neck in or nine complete dark bands.
most of region (1a); narrow white stripe in Malaysia
(1b) and some northern populations. 5. BANDED LINSANG
Prionodon linsang, p. 319
2. COMMON PALM CIVET HB 350–450. Like Spotted
Paradoxurus hermaphroditus, Linsang, P. pardicolor, but black
p. 314 splotches form dark bands
HB 420–500. Colour varies across back; tail usually with
from olive-brown to grey-brown seven complete dark bands.
or sometimes reddish; three
distinct black stripes on back
with extensive black spots on
sides sometimes forming additional stripes; dark
mask on face with white behind eyes.

3. SMALL-TOOTHED PALM
CIVET
Arctogalidia trivirgata, p. 316
HB 440–530. Colour varies
from olive-brown to greyish-
brown; three dark stripes on
back; no spots on sides; long
tail; face blackish, usually with
thin white stripe on forehead.

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1a

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 Plate 45: BINTURONG, BANDED CIVETS AND OTTER CIVET
1. BINTURONG 3. BANDED CIVET
Arctictis binturong, p. 315 Hemigalus derbyanus, p. 316
HB 650–950. Fur long and HB 450–560. Usually buffy-
shaggy, blackish with pale grey brown, but sometimes light grey
tips, especially on head; tail or dark orange; dark brown or
long with thick fur at base; ears black bands across back; no
round with long tufts of hair spots on legs; tail banded at
at tips. base, all dark on distal half.

2. OWSTON’S CIVET 4. OTTER CIVET

Chrotogale owstoni, p. 316 Cynogale bennettii, p. 316
HB 510–630. Greyish-white to HB 575–680. Broad muzzle
buffy-brown; dark bands across with white lips, very long white
back; small black spots on legs; whiskers and white spot above
tail banded at base, all dark on eye; short tail; brown to grey-
distal half. brown body.

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 Plate 46: MONGOOSES
1. JAVAN MONGOOSE 3. SHORT-TAILED
Urva javanica, p. 317 MONGOOSE
HB 320–420. Medium Urva brachyura, p. 317
mongoose, but males much HB 380–445. Dark blackish-
larger than females; fur finely brown with fine buff or orange
speckled buff and dark brown, speckling; relatively short,
with strong reddish cast, tapered tail.
especially on head; long tail.
Small Indian Mongoose, U. auropunctata, in 4. CRAB-EATING MONGOOSE
W Myanmar and India (not illustrated) is similar but Urva urva, p. 318
smaller, lacking any reddish tinge to fur. HB 440–480. Large; long,
shaggy brownish to greyish
2. INDIAN GREY MONGOOSE grizzled fur; conspicuous pale
Urva edwardsii, p. 318 stripe on sides of neck; pale
HB 370–480. Grizzled grey; chin; relatively short but
long tail; no white stripe on bushy tail.
shoulder. Likely no longer
occurs in region.

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 Plate 47: LARGE CATS
1. MAINLAND CLOUDED 3. TIGER
LEOPARD Panthera tigris, p. 320
Neofelis nebulosa, p. 320 HB 1,700–2,300. Very large;
HB 650–950. Irregular cloud- variable pattern of black stripes
like dark splotches on pale on orange coat; relatively
coat; long thick tail; largely shorter tail. May now be
arboreal. extirpated from some parts of
range shown on map.
2. LEOPARD
Panthera pardus, p. 320
HB 1,050–1,300. Long tail;
two colour phases: spotted
form (2a) has black rosettes
with brown centres on pale
coat; black form (2b) is all black
with indistinct patterns of black
rosettes.

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2b

2a

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 Plate 48: SMALLER CATS
1. MARBLED CAT 4. FISHING CAT
Pardofelis marmorata, p. 320 Prionailurus viverrinus, p. 322
HB 450–530. Complex marbled HB 720–780. Large; light
pattern of dark splotches and grey-brown coat; black stripes
black lines on back; spots on on head and neck; small spots
legs; stripes on head; long tail. tend to form rows on flanks and
back; relatively short tail.

2. LEOPARD CAT 5. JUNGLE CAT

Prionailurus bengalensis, Felis chaus, p. 323
p. 322 HB 500–650. Back fur varies
HB 400–550. Orange to from ashy grey to yellowish-
yellowish coat; black spots brown; a few black marks on
vary from small (as shown) to legs; tail with dark rings at tip;
large and round to large and pointed ears with short tufts.
irregular; medium-length tail.
6. ASIAN GOLDEN CAT
3. FLAT-HEADED CAT Catopuma temminckii, p. 321
Prionailurus planiceps, p. 322 HB 760–840. Back fur usually
HB 445–505. Greyish-brown unmarked, varies from golden
body with fine black speckling; to tawny to greyish, rarely all
often some orange on head; black; narrow black-and-white
relatively short, unmarked tail; stripes on head; rounded ears;
short rounded ears; flat sloping relatively short tail with white
forehead. underside. In N Myanmar, marbled form may occur
– see text.

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2
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 Plate 49: FUR SEAL AND SPOTTED SEAL
1. NORTHERN FUR SEAL 2. SPOTTED SEAL
Callorhinus ursinus, p. 323 Phoca largha, p. 324
HB up to 2.1m (male), 1.5m (female). Large front HB up to 1.7m (sexes similar). Distinctive body
and hind flippers; relatively small head with raised shape with moderate-sized head. Body covered
forehead and short snout; visible ears. with black spots; relatively small flippers.

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 Plate 50: KILLER WHALES, PILOT WHALE, RISSO’S DOLPHIN
AND MELON-HEADED WHALE
1. PYGMY KILLER WHALE 4. FALSE KILLER WHALE
Feresa attenuata, p. 325 Pseudorca crassidens, p. 325
TL 2.2–2.6m. Dark grey-brown to black back, paler TL 3.2–4.0m. Conical rounded head; slender body;
sides; white patches on lips and belly; tall dorsal fin; conspicuous dorsal fin; largely black; flippers with
rounded head; flippers smoothly rounded in front. hump on leading edge.

2. KILLER WHALE 5. RISSO’S DOLPHIN

Orcinus orca, p. 326 Grampus griseus, p. 326
TL 6.0–9.5m. Largest dolphin; tall, triangular dorsal TL 3.6–4.0m. Domed forehead with crease in
fin in male, smaller and more curved in female; middle; no beak; very prominent dorsal fin;
black with white mark behind eye, pale saddle on grey body.
back; rounded flippers.
6. MELON-HEADED WHALE
3. SHORT-FINNED PILOT WHALE Peponocephala electra, p. 325
Globicephala macrorhynchus, p. 326 TL 2.2–2.7m. Relatively small; dark grey to black;
TL 3.0–5.4m. Generally black; bulbous forehead domed forehead with no beak; tall, curved dorsal
with no beak; low dorsal fin; pale patch on back; fin; pointed flippers.
curved flippers.

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 Plate 51: DOLPHINS
1. FRASER’S DOLPHIN 4. PANTROPICAL SPOTTED DOLPHIN
Lagenodelphis hosei, p. 327 Stenella attenuata, p. 327
TL 2.3–2.7m. Short beak; dark grey above; broad TL 2–2.5m. Long narrow beak; dark cape, broadest
black stripe on side; dorsal fin slightly curved; below dorsal fin; thin black line from flippers to
relatively small flippers. mouth; varies from heavily spotted (as shown) to
nearly unspotted.
2. SPINNER DOLPHIN
Stenella longirostris, p. 328 5. LONG-BEAKED COMMON DOLPHIN
TL 1.3–2.3m. Long narrow beak; low forehead; tall, Delphinus capensis, p. 327
triangular dorsal fin; black back forming relatively TL 1.8–2.5m. Long beak; black back forming V
straight border with grey sides. below dorsal fin; pale yellow or buff patch in front,
grey patch at back forming hourglass pattern.
3. STRIPED DOLPHIN
Stenella coeruleoalba, p. 328
TL 1.8–2.7m. Long distinct beak; tall, curved
dorsal fin; black back; pale grey stripe from eye
to below dorsal fin, and along sides; narrow black
lines behind eye.

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 Plate 52: DUGONG, PORPOISE AND DOLPHINS
1. DUGONG 4. ROUGH-TOOTHED DOLPHIN
Dugong dugon, p. 335 Steno bredanensis, p. 328
TL 2.5–3.5m. Broad rounded head; thick lips; TL 1.8–2.8m. Forehead slopes into slender beak;
paddle-like flippers; no dorsal fin; generally dark grey above with pale underparts and lips;
moves slowly. irregular white spots on sides; relatively tall
dorsal fin.
2. IRRAWADDY DOLPHIN
Orcaella brevirostris, p. 329 5. INDO-PACIFIC HUMPBACKED DOLPHIN
TL 2.0–2.5m. Rounded forehead; no beak; large Sousa chinensis, p. 329
flippers; small dorsal fin low on back; plain colour TL 2.4–2.8m. Long beak; domed forehead; usually
varies from light grey to dark blue-grey. has hump on back with short dorsal fin; relatively
plain colour varies from dark grey to whitish or pink.
3. INDO-PACIFIC FINLESS PORPOISE
Neophocaena phocaenoides, p. 330 6. INDO-PACIFIC BOTTLENOSE DOLPHIN
TL 1.5–2.0m. Small; rounded forehead; no dorsal Tursiops aduncus, p. 329
fin; small flippers; colour varies from light grey (as TL 2.0–2.6m. Moderate length, thick beak;
shown) to very dark grey. somewhat bulbous forehead; large dorsal fin; large
flippers; relatively plain grey colour. See text for
distinctions from Common Bottlenose Dolphin,
T. truncatus (not illustrated).

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 Plate 53: BEAKED WHALES AND SMALL SPERM WHALES
1. CUVIER’S BEAKED WHALE 4. PYGMY SPERM WHALE
Ziphius cavirostris, p. 331 Kogia breviceps, p. 331
TL 5.5–7.5m. Sloping forehead; jaw not arched; TL 2.7–3.4m. Large blunt forehead; low jaw;
colour varies from pale brown to dark reddish- crescent gill-like mark behind eye; small dorsal fin
brown to slate-grey; small dorsal fin. behind middle of back; swims sluggishly when
on surface.
2. BLAINVILLE’S BEAKED WHALE
Mesoplodon densirostris, p. 332 5. DWARF SPERM WHALE
TL 4.4–4.7m. Low forehead with distinct beak; Kogia sima, p. 331
arched lower jaw with prominent tooth in adult TL 2.1–2.7m. Similar to K. breviceps, but averages
male; back dark grey with extensive scarring and smaller; forehead more pointed; dorsal fin taller,
blotching. Longman’s Beaked Whale, Indopacetus usually slightly further forwards.
pacificus (not illustrated), is also dark, but larger,
with bulbous forehead, long beak, relatively large
dorsal fin.

3. GINKGO-TOOTHED BEAKED WHALE

Mesoplodon ginkgodens, p. 332
TL 4.7–5.2m. Prominent beak; arched lower jaw
with small lobed tooth in male.

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 Plate 54: LARGE WHALES
1. GREAT SPERM WHALE 5. FIN WHALE
Physeter macrocephalus, p. 330 Balaenoptera physalus, p. 333
TL 8–12m. Large head, square forehead; wrinkled TL 18–20m. Very large; dark grey to blackish back;
back with low, stepped dorsal fin; broad square- fin visible shortly after blow; lower lip and baleen
ended tail raised on dive, dark underneath; blow white on right side of head, dark on left.
angled forwards and to left.
6. BLUE WHALE
2. HUMPBACK WHALE Balaenoptera musculus, p. 333
Megaptera novaeangliae, p. 334 TL 22–24m. Largest whale; mottled bluish-grey
TL 11–13m. Large flippers with white and black back pattern; broad rounded head; small fin rarely
markings; low, stepped dorsal fin; broad notched visible until whale dives.
tail visible when dives, largely white below.
7. MINKE WHALE
3. SEI WHALE Balaenoptera acutorostrata, p. 334
Balaenoptera borealis, p. 333 TL 7–8m. Smallest rorqual; narrow pointed
TL 12–14m. Dark grey with white scar marks; head; white marks on flippers; blow is relatively
dorsal fin visible at same time as blow; one ridge inconspicuous.
on top of head.

4. BRYDE’S WHALE
Balaenoptera edeni, p. 334
TL 10–12m. Dark grey with white scar marks;
dorsal fin visible at same time as blow; three
prominent ridges on top of head.

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 Plate 55: ELEPHANT AND RHINOCEROSES
1. ASIAN ELEPHANT 3. LESSER ONE-HORNED
Elephas maximus, p. 335 RHINOCEROS
SH 1.5–3.0m. Very large; Rhinoceros sondaicus, p. 338
distinctive shape with long SH 1.6–1.8m. Large; three
trunk; moderately large ears; folds of skin across back;
adult males with large tusks. horizontal fold of skin below
rump; only one horn, which is
moderate size in male, small in
2. ASIAN TWO-HORNED female. Probably now extinct on mainland SE Asia.
RHINOCEROS
Dicerorhinus sumatrensis,
p. 337
SH 1.2–1.3m. Large; only one
fold of skin crosses back; two
horns, but rear horn is small
and often inconspicuous.
Probably now extinct on mainland SE Asia.

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 Plate 56: TAPIR AND PIGS
1. ASIAN TAPIR 3. EURASIAN WILD PIG
Tapirus indicus, p. 337 Sus scrofa, p. 339
SH 900–1,050. Elongate nose; SH 600–800. Elongate snout
short tail. Adult with distinctive with flat tip; colour varies from
black-and-white body pattern blackish to reddish – may
(1a). Young initially with dark be affected by mud colour;
body and horizontal white lacks large bristles on snout.
stripes (1b); gradually acquires Widespread form (3a) has
adult pattern as young grow older. well-developed mane. S. s. vittatus from Peninsular
Malaysia (3b) tends to have shorter mane; sparser,
2. BEARDED PIG whitish fur; white band of fur on muzzle. Babies
Sus barbatus, p. 339 have brown and beige stripes (3c).
SH 700–900. Elongate snout
with flat tip; prominent bristles
on top of snout, under chin
and on cheeks, longer and
more conspicuous in male (2a)
than female (2b). Averages
larger than S. scrofa; colour varies from blackish to
reddish-brown to grey, depending in part on mud
wallow. Babies with brown and beige stripes (2c).

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1a

2b

2a

2c

3b 3c
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 Plate 57: MOUSEDEER AND SMALL MUNTJACS
1. LESSER MOUSEDEER 5. ANNAMITE MUNTJAC
Tragulus kanchil, p. 340 Muntiacus truongsonensis,
SH 200–230. Very small; fur p. 345
relatively uniformly coloured, SH c.400. Small; dark fur with
reddish-brown finely speckled black legs; head with yellowish
with black; brown and white and black markings; male with
markings on throat appear as tiny antlers on short pedicels,
one white line in profile. often hidden by fur. Distinctions
from Roosevelt’s Muntjac, M. rooseveltorum, and
2. SILVER-BACKED Puhoat Muntjac, M. puhoatensis (not illustrated),
MOUSEDEER uncertain; range map includes all three.
Tragulus versicolor, p. 341
SH c.250. Small; back and 6. GONGSHAN MUNTJAC
flanks silvery grey speckled Muntiacus gongshanensis,
with dark; head and shoulders p. 346
orange-brown; throat markings SH 500–550. Medium-sized;
usually show one white line in body mid- to dark brown;
profile. legs, belly, top of tail black;
contrasting white undertail
3. GREATER MOUSEDEER and insides of legs; head
Tragulus napu, p. 340 pale brown with dark marks on pedicels; antlers
SH 300–350. Larger; back fur relatively short.
mottled orange, greyish and
blackish; dark brown and white 7. FEA’S MUNTJAC
throat markings usually show Muntiacus feae, p. 346
two white lines in profile. SH 500–550. Medium-sized;
dark brown sides, speckled
4. LEAF MUNTJAC with yellow; belly usually similar
Muntiacus putaoensis, p. 346 colour to back or paler, but may
SH c.400. Small; reddish; appear dark in shadow; legs
male with tiny antlers on short and top of tail dark; yellowish
pedicels, often hidden by fur. top of head with black stripes on pedicels.

8. RED MUNTJAC
Muntiacus muntjak, p. 344
SH 500–550. Medium-sized;
fur varies from reddish-brown
to reddish-yellow; tail coloured
as back. Male (8a) with long
pedicels, short antlers; head
colour as back, with dark
stripes on pedicels and forehead. Female (8b) lacks
pedicels and antlers (as in other muntjacs). Fawn
(8c) lightly spotted.

Note Only male muntjacs have pedicels and antlers.

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 Plate 58: LARGE-ANTLERED MUNTJAC, TUFTED DEER,
MUSK-DEER AND HOG DEER
1. LARGE-ANTLERED 4. FOREST MUSK-DEER
MUNTJAC Moschus berezovskii, p. 342
Muntiacus vuquangensis, p. 347 SH 500–600. Similar to
SH 650–700. Larger than other M. fuscus, but fur speckled
muntjacs; fur colour yellowish- brown with underside paler
brown to grey-brown; dark lines than back; long white stripes
on forehead; male (1a) with in middle and sides of throat;
large antlers distinctly forked, ears brown at bases, white in
stout, moderately long pedicels; female (1b) lacks middle, black at tip.
antlers or pedicels.
5. HOG DEER
2. TUFTED DEER Axis porcinus, p. 344
Elaphodus cephalophus, p. 344 SH 650–720. Male (5a) like
SH 500–700. Coarse, slightly small version of Sambar, Cervus
shaggy fur, dark brown on body, unicolor; fur light brown to
greyer on neck; black legs; tuft greyish-brown; tail same colour
of hair on top of head; male as back above, white below;
with small antlers often hidden antlers slender with three tines;
by tuft; white marks at base adult female (5b) often retains some spots; young
of ear. (5c) extensively spotted.

3. BLACK MUSK-DEER
Moschus fuscus, p. 342
SH c.600. No antlers; male with
long protruding upper canines;
long hind legs make rump
higher than shoulders; fur dark
brown to blackish above and
below; ears dark.

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1b

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 Plate 59: LARGE DEER
1. SIKA 3. ELD’S DEER
Cervus nippon, p. 343 Cervus eldii, p. 343
SH 650–1,100. Antlers of male SH 1,200–1,300. Brown above,
with five tines (1a); female lacks white belly; antlers of male (3a)
antlers (1b); coat varies from with prominent brow tine that
very dark in winter (1a) to pale forms continuous curve with
brown in summer (1b); belly main branch, several small tines
usually paler; young (1c) heavily at tip of main branch; tail brown
spotted; adults retain a few to many white spots; above; female (3b) paler than C. unicolor.
small white patch on rump around base of tail.
4. SAMBAR
2. SCHOMBURGK’S DEER Cervus unicolor, p. 342
Rucervus schomburgki, p. 343 SH 1,400–1,600. Large; dark
SH 1,050. Uniformly brown fur, brown or greyish-brown above,
slightly redder legs; antlers of dark below; tail black above,
male (2a) with numerous long white below; antlers of adult
tines; female (2b) uniformly male (4a) usually with only
brown. Probably Extinct. three tines; female (4b) usually
darker than females of other deer; young (4c) may
be lightly spotted.

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 Plate 60: WILD CATTLE AND BUFFALO
1. GAUR 3. KOUPREY
Bos gaurus, p. 348 Bos sauveli, p. 348
SH 1,700–1,850. Adult male SH 1,700–1,900. Adult male
(1a) very muscular; high ridge (3a) with very large dewlap;
of muscles on back; hollow horns curved, tips often split;
area on forehead between dark area between horns; rump
horns; body black; lower legs dark. Adult female (3b) grey;
greyish-white or yellowish. horns lyre-shaped. Probably
Adult female (1b) smaller and less muscular; often Extinct.
browner. Newborn calf (1c) yellowish in some
populations, brown in others. 4. WATER BUFFALO
Bubalus arnee, p. 348
2. BANTENG SH 1,600–1,900. Adult (4a)
Bos javanicus, p. 347 with long horns sticking out to
SH 1,500–1,700. Adult male sides, slightly larger in male
(2a) black with white legs and than female; uniformly grey;
white rump; narrow curved pale marks on neck. Calf
horns. Adult female (2b) brown (4b) grey. Wild animals
with white legs and white rump; more muscular and much more wary than
horns closer together, more domestic ones.
upright.

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1b

1c

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2b

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 Plate 61: TAKIN, SHEEP AND SAOLA
1. TAKIN 3. SAOLA
Budorcas taxicolor, p. 349 Pseudoryx nghetinhensis,
SH 700–1,400. Distinctive p. 349
head shape; thick legs. Adult SH 800–900. Slender; long,
male (1a) with thick short horns smooth curved horns in adults
curved backwards; fur thick (short and straighter in young);
and woolly; colour varies from black-and-white marks on face,
greyish-black to dark brown to feet and rump.
yellowish-brown. Adult female (1b) usually greyer;
horns smaller and straighter. Young (1c) greyish,
often with dark line down back.

2. BLUE SHEEP
Pseudois nayaur, p. 350
SH 800–900. Light bluish-grey,
sometimes with browner tinge.
Male (2a) with large curved
horns; black marks on belly and
flanks. Female (2b) with shorter,
straighter horns; fewer black
marks.

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2a
2b

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 Plate 62: SEROWS AND GORALS
1. SOUTHERN SEROW 4. CHINESE GORAL
Capricornis sumatraensis, Naemorhedus griseus, p. 351
p. 350 SH 500–700. Smaller and more
SH 850–940. Short body; long slender than Capricornis; no
legs; body mainly black with mane; short greyish-brown to
black legs; adults with long brown fur; short curved horns;
mane on back of neck, varying thin dark line on back.
from all reddish to dirty white to
all black. 5. RED GORAL
Naemorhedus baileyi, p. 351
2. INDOCHINESE SEROW SH 550–600. Similar to
Capricornis milneedwardsi, N. griseus, but body and legs
p. 350 reddish-brown; thin dark line
SH 850–940. Similar to on back.
C. sumatraensis, but fur with
extensive white bases, giving
overall greyish colour; lower
legs usually contrasting reddish
or whitish; mane dirty white (2a) or dark (2b).

3. RED SEROW
Capricornis rubidus, p. 350
SH 850–950. Similar shape to
other Capricornis; fur reddish
with white on chin and belly;
short reddish mane; black line
down back.

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2a

2b

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 Plate 63: RATUFA AND CALLOSCIURUS SQUIRRELS
1. BLACK GIANT SQUIRREL 5. SUNDA BLACK-BANDED
Ratufa bicolor, p. 353 SQUIRREL
HB 370–405. Very large with Callosciurus nigrovittatus, p. 355
long tail; upperparts and tail HB 170–240. Brownish agouti
black or very dark brown; above; grey belly; black and
underparts vary from buff to buff stripes on sides; tail grey
light orange; cheeks white with with no red tip; sides of head
thin black moustache. and chin both orange-brown.

2. CREAM-COLOURED GIANT 6. IRRAWADDY SQUIRREL

SQUIRREL Callosciurus pygerythrus, p. 355
Ratufa affinis, p. 353 HB 170–200. Upperparts
HB 310–380. Very large; long agouti with grey or brown tinge;
tail; fur varies from uniformly often has pale patch on legs (as
light buff to orange-brown shown); no black or buff bands
above with paler underparts. on flanks; underparts vary from
orange to buff to grey (see text).
3. PREVOST’S SQUIRREL
Callosciurus prevostii, p. 354 7. PHAYRE’S SQUIRREL
HB 200–270. Medium to large; Callosciurus phayrei, p. 355
distinctive pattern of black HB 210–250. Upperparts
upperparts, white sides, orange greyish to brownish agouti; tip
underparts. of tail black; underparts buff to
orange; black stripe on flank.

4. PLANTAIN SQUIRREL
Callosciurus notatus, p. 354
HB 175–225. Brownish agouti
above; orange belly; black and
buff stripes on sides; usually
has reddish tip to tail.

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 Plate 64: CALLOSCIURUS SQUIRRELS
1. GREY-BELLIED SQUIRREL 3. PALLAS’S SQUIRREL
Callosciurus caniceps, p. 357 Callosciurus erythraeus, p. 355
HB 210–240. Geographic HB 200–260. Varies in colour;
and seasonal variation in most forms have upperparts
colour; legs and feet always agouti with grey or brown tinge;
greyish; belly usually grey but reddish on underparts. Some
sometimes with reddish sides. major variations:
(1a, 1b) C. c. caniceps (3a, 3b) C. e. sladeni (C
(mainland part of range). Back yellowish-brown to Myanmar). Highly variable population, with some
bright red-brown in dry season; grey in wet season; individuals largely rufous; some very pale grey
tail tip black. with yellowish feet and underparts; some closely
(1c) C. c. bimaculatus (peninsular Myanmar and resembling typical C. e. erythraeus.
Thailand N of 7°N). Upperparts mostly grey with (3c) C. e. atrodorsalis (W Thailand and S Myanmar).
some red patches; flanks reddish; tail tip black. Lower back with broad black stripe.
(1d) C. c. concolor (Peninsular Malaysia). Back (3d) C. e. erythraeus group (most of range).
greyish agouti with golden tinge; grey flanks and Upperparts, feet and tail agouti, variably tinged with
belly; tail tip not black. reddish; belly reddish except for agouti stripe up
middle.
2. INORNATE SQUIRREL (3e) C. e. flavimanus (S Vietnam, S Laos and
Callosciurus inornatus, p. 357 Cambodia). Upperparts greyish agouti; feet pale,
HB 220–230. Upperparts grey usually yellow-orange; underparts vary from cream
and brown agouti; underside to bright red-brown.
grey; tail relatively plain with
little or no banding, tip usually 4. STRIPE-BELLIED
not dark. Does not overlap in SQUIRREL
range with C. caniceps. Callosciurus quinquestriatus,
p. 357
HB 210–240. Upperparts
agouti; belly white with three
black stripes, one on each side
and one up middle.

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1b

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3b

3c

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 Plate 65: VARIABLE SQUIRREL
1. VARIABLE SQUIRREL (1e, 1f) C. f. bocourti (E and C Thailand). Dark
Callosciurus finlaysonii, p. 356 phase: dark grey to black above, including tail;
HB 210–220. Extreme white below with white head. Pale phase: grey
geographic variation in colour agouti above; white head, underparts and tail.
pattern; no single character (1g) C. f. floweri (SC Thailand). Brown-grey agouti
defining species. In some above; white below; often reddish around head and
areas more than one colour tail; white marks around eye.
phase exists. Colours may (1h) C. f. cinnamomeus (SE Thailand and SW
also vary seasonally. Some of the major colour Cambodia). Largely orange-brown, darker in middle
variations are illustrated here, but additional of back; often with small patches of greyish agouti
combinations exist, with intermediates where on legs and face.
populations meet (see text). (1i) C. f. subsp. (C Laos). Glossy black with large
(1a) C. f. sinistralis (NW Thailand). Greyish upper- patches of dark red.
parts, often with reddish tinge; reddish underparts (1j) C. f. folletti (Phai island, Thailand). Greyish-
and sides of head; red tail with white base. white to beige; pale buffy underparts; tail pale,
(1b, 1c) C. f. menamicus (N Thailand and N Laos). often with black tip.
Dark phase: dark reddish-brown; slightly paler (1k) C. f. boonsongi (NE Thailand). Black fur with
below; slightly darker at base of tail. Pale phase: white rims to ears.
reddish-orange back, belly and tail; greyish agouti (1l) C. f. harmandi (Phu Quoc island, Vietnam).
on sides and legs. Upperparts vary from dark reddish-brown to pale
(1d) C. f. finlaysonii (Si Chang island, Thailand). grey-brown; pale orange below; grey tail.
Largely white with yellow or grey tinge.

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1h

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 Plate 66: STRIPED AND SUNDASCIURUS SQUIRRELS
1. WESTERN STRIPED 5. INDOCHINESE GROUND
SQUIRREL SQUIRREL
Tamiops mcclellandii, p. 358 Menetes berdmorei, p. 361
HB 110–125. Small; white HB 180–210. Medium-sized
tufts on tips of ears; dark stripe with moderate-length tail; buff
down middle of back with and dark stripes on sides;
two pale stripes and two dark typically pale stripe above
stripes on each side; outer pale and below broad dark stripe;
stripe typically wider and paler than inner stripe, sometimes additional dark stripe above; lower pale
contiguous with cheek stripe; outer dark stripes stripe may be indistinct.
vary from black to brown.
6. LOW’S SQUIRREL
2. CAMBODIAN STRIPED Sundasciurus lowii, p. 359
SQUIRREL HB 130–150. Dark brown to
Tamiops rodolphii, p. 358 reddish-brown upperparts; pale
HB 105–125. Similar to eye-ring; relatively short bushy
T. mcclellandii, but all four pale tail; belly fur uniformly white
stripes typically similar in width; or buff.
inner pair often tinged yellow-
orange; central black stripe with 7. SLENDER SQUIRREL
thin brown line up middle. Sundasciurus tenuis, p. 359
HB 115–155. Olive-brown
3. SWINHOE’S STRIPED upperparts; buff marks around
SQUIRREL eye and on nose; long slender
Tamiops swinhoei, p. 358 tail; belly fur white or buff with
HB 120–155. Similar to grey bases.
T. mcclellandii, but inner pale
stripes dull, similar in colour to 8. HORSE-TAILED SQUIRREL
head; dark stripes vary from Sundasciurus hippurus, p. 359
black to dark brown; outer HB 210–290. Bushy black tail;
pale stripes do not connect to face stripes; belly reddish-brown upperparts;
hairs grey. contrasting grey head and
thighs; reddish-orange
4. EASTERN STRIPED underparts.
SQUIRREL
Tamiops maritimus, p. 358
HB 100–145. Similar to
T. swinhoei, but dark stripes
generally less conspicuous;
belly hairs grey with whiter tips.

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5b

6
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 Plate 67: LONG-NOSED AND GROUND SQUIRRELS
1. PERNY’S LONG-NOSED 5. RED-HIPPED SQUIRREL
SQUIRREL Dremomys pyrrhomerus, p. 360
Dremomys pernyi, p. 360 HB c.215. Like D. rufigenis, but
HB 180–210. Upperparts hips and thighs reddish-orange;
brownish agouti; orange spot muzzle averages slightly more
behind ear; greyish underparts; elongate.
elongate muzzle; often reddish
around base of tail.
6. THREE-STRIPED GROUND
2. ORANGE-BELLIED SQUIRREL
SQUIRREL Lariscus insignis, p. 361
Dremomys lokriah, p. 360 HB 170–230. Upperparts
HB 170–200. Upperparts dark brown to reddish-brown;
greyish to brownish agouti; three black stripes on back;
underparts mixture of orange, underparts white, sometimes
orange-buff and grey; buff spot with buff tinge.
behind ear; some populations
have dark line down middle of back; muzzle often 7. SHREW-FACED GROUND
less pointed than that of D. pernyi. SQUIRREL
Rhinosciurus laticaudatus,
3. RED-THROATED SQUIRREL p. 361
Dremomys gularis, p. 360 HB 195–233. Elongate muzzle;
HB c.220. Like D. rufigenis, upperparts dark reddish agouti
but throat also red; underparts with no stripes; tail relatively
darker greyish. short, often grizzled grey;
underparts buffy-white.

4. RED-CHEEKED SQUIRREL
Dremomys rufigenis, p. 360
HB 170–210. Reddish cheeks,
face and underside of tail;
brownish agouti above; belly fur
with grey bases, white tips.

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3

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 Plate 68: GIANT FLYING SQUIRRELS
1. RED GIANT FLYING 3. YUNNAN GIANT FLYING
SQUIRREL SQUIRREL
Petaurista petaurista, p. 362 Petaurista yunanensis, p. 363
HB 400–520. All forms have HB 400–500. Upperparts dull
distinct black tip to tail. (1a) reddish-brown; back hairs
P. p. melanotus (Peninsular lightly frosted with extensive
Malaysia). Bright orange-brown dull grey tips; tail dark brown
body; black feet and eye-ring. to black.
(1b) P. p. taylori (S Myanmar, W Thailand). Dark
reddish-brown with light speckles on head and 4. CHINDWIN GIANT FLYING
back. (1c) P. p. candidula (N Thailand, Myanmar). SQUIRREL
Reddish or orange-brown with extensive white or Petaurista sybilla, p. 364
grey frosting on back and head; greyish tail. HB 350–400. Smaller than
other Petaurista. Largely
2. INDIAN GIANT FLYING orange-brown, mottled with
SQUIRREL dark grey and buff on back; feet
Petaurista philippensis, p. 363 and tail without black.
HB 400–490. Variable in
colour; tail usually all dark.
(2a) P. p. annamensis (Vietnam).
Dark brown to dark red with
extensive pale frosting; tail
brownish-black to glossy black. (2b) P. p. lylei
(Thailand, E Myanmar). Upperparts dark grey-
brown with extensive grey frosting; tail dark brown;
underside buff. (2c) P. p. cineraceus (W Myanmar).
Upperparts and tail dark grey with extensive
frosting; tail sometimes has a darker tip; underside
greyish-white.

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1b

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2a

2b

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 Plate 69: LARGE AND MEDIUM FLYING SQUIRRELS
1. LESSER GIANT FLYING 3. BLACK FLYING SQUIRREL
SQUIRREL Aeromys tephromelas, p. 364
Petaurista elegans, p. 364 HB 355–426. Large; upperparts
HB 340–365. (1a) P. e. punctata and head dark brownish-black;
(Malay Peninsula). Upperparts underparts sparsely haired,
reddish mixed with black; dark grey-brown.
extensive large white spots on
back; tail black; underside buff.
(1b) P. e. marica (rest of mainland range).

2. LAO GIANT FLYING

SQUIRREL
Biswamoyopterus laoensis,
p. 365
HB 455. Upperparts dark brown
with white frosting; underparts
pale orange.

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 Plate 70: LARGE AND MEDIUM FLYING SQUIRRELS
1. PHAYRE’S FLYING 4. HAIRY-FOOTED FLYING
SQUIRREL SQUIRREL
Hylopetes phayrei, p. 368 Belomys pearsonii, p. 365
HB 145–195. Back dark with HB 190–200. Medium;
grey-brown to buffy-brown tips; upperparts with dark bases,
tail varies from buffy-brown to mottled grey and red tips;
dark brown, rounded on top, underparts creamy white; tail
flat below; underparts creamy reddish-brown, rounded above,
white; face greyish-white. flattened below; tufts of hair behind ear.

2. PARTICOLOURED FLYING 5. HORSFIELD’S FLYING

SQUIRREL SQUIRREL
Hylopetes alboniger, p. 367 Iomys horsfieldii, p. 366
HB 175–225. Similar to HB 165–230. Medium;
H. phayrei, but larger; colour upperparts grey-brown to red-
varies from extensively white- brown with pale tips; orange
frosted (as shown) to much edge to gliding membrane;
darker. tail rounded, orange-brown;
underside white, sometimes with orange tinge.
3. SMOKY FLYING SQUIRREL
Pteromyscus pulverulentus,
p. 365
HB 220–290. Medium-large;
upperparts smoky brownish-
grey with fine white frosting;
white below with grey tinge;
pale grey on cheeks; tail narrow
and flattened.

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 Plate 71: SMALL FLYING SQUIRRELS
1. GREY-CHEEKED FLYING 4. TEMMINCK’S FLYING
SQUIRREL SQUIRREL
Hylopetes platyurus, p. 367 Petinomys setosus, p. 366
HB 115–135. Medium-small; HB 105–125. Small; upperparts
upperparts black with red tips; black with whitish tips to hairs;
tail flat with narrow base, has underparts white with grey
white patch at base; underparts bases; dark ring around eye;
and cheeks greyish-white. tail relatively flat, dark with grey
base; tail with white tip in northern populations.
2. RED-CHEEKED FLYING
SQUIRREL 5. VORDERMANN’S FLYING
Hylopetes spadiceus, p. 367 SQUIRREL
HB 157–184. Upperparts Petinomys vordermanni, p. 366
black with red tips; tail flat with HB 95–120. Small; upperparts
narrow base, has orange patch dark with reddish tips;
at base; underparts and cheeks reddish tail and crown; buffy
buffy-white with reddish tinge. underparts; buffy cheeks with
dark eye-ring; tail flat below,
3. WHISKERED FLYING slightly rounded above.
SQUIRREL
Petinomys genibarbis, p. 366 6. MALAYSIAN PYGMY
HB 160–180. Back dark grey FLYING SQUIRREL
with red-brown tips; lower back Petaurillus kinlochii, p. 368
contrasting pinkish-brown; HB 80–90. Very small;
tuft of whiskers on cheek upperparts dark grey with buff
behind eye. streaks; underparts white with
grey bases; tail flat and feathery
with white tip.

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 Plate 72: TYPICAL RATS
1. HOUSE RAT 4. PACIFIC RAT
Rattus rattus, p. 372 Rattus exulans, p. 374
HB 105–215. Underparts HB 90–140. Small; coarse
buffy-brown with grey bases, spiny fur; greyish-brown above;
sometimes creamy white; long white with grey bases below;
guard hairs on lower back; all-dark tail.
large, well-striated footpads;
dark tail; climbs well.
5. RICEFIELD RAT
2. MALAYSIAN WOOD RAT Rattus argentiventer, p. 373
Rattus tiomanicus, p. 373 HB 140–210. Speckled brown
HB 140–190. Underparts fur; silvery grey underparts;
pure white or creamy white; dark brown tail; small unridged
upperparts sleek with footpads; often has orange tuft
relatively short guard hairs; in front of ear.
well-developed footpads for
climbing; dark tail. 6. OSGOOD’S RAT
Rattus osgoodi, p. 376
3. LESSER RICEFIELD RAT HB 125–170. Thick soft fur;
Rattus sakeratensis, p. 375 dark brown above; dark grey
HB 120–180. Medium-small; below; dark tail; dark feet.
narrow feet with small unridged
footpads; tail shorter than HB;
fur varies from grey-brown
to red-brown. See text for
distinctions from Losea Rat,
Rattus losea (not illustrated).

Note Most Rattus have all-dark tails (except some R. norvegicus), fur with numerous soft spines and elongate footpads.

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 Plate 73: LARGE RATS
1. NORWAY RAT 5. MÜLLER’S RAT
Rattus norvegicus, p. 374 Sundamys muelleri, p. 380
HB 160–265. Large; shaggy HB 210–280. Very large; long
fur; tail dark above, slightly dark tail; coarse shaggy fur
paler below, sometimes with with long guard hairs but no
pale splotches; pale feet; poorly conspicuous spines; white or
developed footpads; relatively pale grey underparts.
small ears and eyes; usually
lives around towns. 6. MILLARD’S GIANT RAT
Dacnomys millardi, p. 385
2. ANNANDALE’S RAT HB 230–290. Very large;
Rattus annandalei, p. 376 thick short fur without spines;
HB 145–220. Grey-brown upperparts brown flecked with
back; white or pale yellow buff; underparts pale; long
below; soft shaggy fur with dark tail.
inconspicuous spines.
7. GREATER BANDICOOT RAT
Bandicota indica, p. 376
3. WHITE-FOOTED HB 190–330. Very large; dark
INDOCHINESE RAT above and below, including feet;
Rattus nitidus, p. 375 long coarse fur; broad front
HB 160–180. Relatively soft incisors; tail shorter than HB.
fur; belly white with grey bases;
feet white, narrow with well-
developed ridged footpads for 8. SAVILE’S BANDICOOT RAT
climbing. Bandicota savilei, p. 377
HB 150–240. Moderately
4. INDOCHINESE FOREST RAT shaggy fur, varies from
Rattus andamanensis, p. 374 brownish-grey to red-brown
HB 155–200. Conspicuous above, grey-buff below; tail
long black guard hairs all over shorter than HB, sometimes
back; creamy white underparts, with white tip. See text
sometimes with orange patch for distinctions from Lesser Bandicoot Rat,
on chest; large footpads; large B. bengalensis (not illustrated).
ears; long tail.

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 Plate 74: B
 ERYLMYS, TREE, SOFT-FURRED AND LIMESTONE
RATS
1. MANIPUR RAT 6. CRUMP’S SOFT-FURRED
Berylmys manipulus, p. 379 RAT
HB 135–185. Smooth stiff Diomys crumpi, p. 387
fur, steel grey above; sharply HB 100–135. Short silky fur;
separated pure white below; dark blackish-grey to brownish-
distal half to two-thirds of tail grey above; white with grey
white; feet white above; front bases below; tail sharply
teeth broad, pale yellow or bicoloured, dark above, pale
white; smaller than other Berylmys. below; feet white.

2. BERDMORE’S RAT 7. POPA SOFT-FURRED RAT

Berylmys berdmorei, p. 378 Millardia kathleenae, p. 387
HB 190–260. Similar to HB 130–165. Short soft fur;
B. manipulus, but slightly larger; light brown to grey above; tail
tail shorter than HB, dark above, distinctly shorter than HB, dark
mottled pale below. above, pale below; only four
footpads.

3. BOWERS’S RAT 8. LAO LIMESTONE RAT

Berylmys bowersi, p. 378 Saxatilomys paulinae, p. 379
HB 235–300. Very large; similar HB 145–164. Relatively small;
colour to other Berylmys; tail shiny greyish-black above; dark
varies geographically from all grey below with light frosting;
dark to dark at base, white long tail, dark above, indistinctly
on distal half. See text for paler below; large bulbous
distinctions from Mackenzie’s footpads.
Rat, B. mackenziei (not illustrated).
9. TONKIN LIMESTONE RAT
4. MALAYAN WOOLLY TREE Tonkinomys daovantieni, p. 379
RAT HB 185–215. Moderately large;
Pithecheir parvus, p. 389 shaggy, dark greyish-black
HB 150–180. Long, soft above, medium-grey below;
reddish-brown fur with grey usually has white mark on
bases, lacking stiff spines; forehead and chest; short tail,
white underparts; short face; dark above, white below and at
long smooth tail. tip; large bulbous footpads.

5. GREY TREE RAT

Lenothrix canus, p. 389
HB 165–220. Soft woolly fur;
grey to grey-brown above; pale
below; tail dark at base, white
at tip; many separate cusps
on teeth.

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 Plate 75: NIVIVENTERS AND TREE RAT
1. CONFUCIAN NIVIVENTER 5. SMOKE-BELLIED
Niviventer confucianus, p. 381 NIVIVENTER
HB 125–170. Long soft fur Niviventer eha, p. 383
yellowish or greyish-brown; HB 110–130. Orange-brown
usually lacks spines or long above; dark marks only around
guard hairs; pure white below; eye; fur of underparts grey with
tail dark above, pale below. white tips; long tail dark above,
slightly paler below.
2. INDOMALAYAN
NIVIVENTER 6. LIMESTONE NIVIVENTER
Niviventer fulvescens, p. 382 Niviventer hinpoon, p. 382
HB 130–170. Yellowish-brown HB 120–160. Upperparts
to reddish-brown; extensive largely grey with extensive
dark brown spines and long spines; underparts buff with
guard hairs; pure white or grey bases; tail same length as
yellowish-white below; long HB, dark above, pale below.
bicoloured tail. Indochinese Mountain Niviventer,
N. tenaster (not illustrated), is similar, but averages 7. DARK-TAILED NIVIVENTER
more yellowish-brown with longer ears. Niviventer cremoriventer, p. 381
HB 130–165. Reddish-brown
3. CAMERON HIGHLANDS to orange-brown above; long
NIVIVENTER guard hairs and many spines;
Niviventer cameroni, p. 381 white below; feet with large
HB 130–170. Bright reddish- footpads; long dark tail with
brown above with many spines tuft at tip. Indochinese Tree
and long guard hairs; white Rat, Chiromyscus langbianis (not illustrated), looks
below; very long tail dark above, similar, but is more yellowish-grey.
white below and at tip.
8. THOMAS’S TREE RAT
4. BRAHMAN NIVIVENTER Chiromyscus thomasi, p. 384
Niviventer brahma, p. 383 HB 145–180. Orange-brown
HB 135–145. Long soft fur; above, usually brighter on sides
orange-brown to yellow-brown of face; dark mark around eye;
above; indistinct dark marks pure white below; first hind
around eye and across muzzle; toe with nail instead of claw.
underparts grey with white Fea’s Tree Rat, C. chiropus (not
tips; very long tail dark above, illustrated), is smaller, without black mask, more
slightly paler below. conspicuous black guard hairs.

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 Plate 76: LEOPOLDAMYS AND MAXOMYS RATS
1. LONG-TAILED GIANT RAT 5. RED SPINY MAXOMYS
Leopoldamys sabanus, p. 385 Maxomys surifer, p. 388
HB 200–275. Large; smooth HB 160–210. Upperparts
fur; brown on back; brighter orange- or reddish-brown with
orange on sides; creamy white extensive stiff spines; white
below; very long tail dark above, below, often with orange under
pale below and at tip. Herbert’s neck or on insides of front legs;
Giant Rat, L. herberti (not tail dark above, white below.
illustrated), found north of the peninsula, is similar,
but less orange. 6. WHITEHEAD’S MAXOMYS
Maxomys whiteheadi, p. 388
2. EDWARD’S GIANT RAT HB 105–150. Small; dark
Leopoldamys edwardsi, p. 386 brown to red-brown above; grey
HB 210–280. Very large; plain to orange-grey below; many
brown above without orange soft spines; bicoloured tail
sides; white below; long dark shorter than HB.
tail. Millet’s Giant Rat, L. milleti
(not illustrated), is similar, but 7. MALAYAN MOUNTAIN
dark blackish-brown above. MAXOMYS
Sundaic Mountain Rat, L. ciliatus (not illustrated), Maxomys inas, p. 388
from Peninsular Malaysia is best distinguished HB 125–160. Medium size;
by range. spiny fur; upperparts reddish-
brown; underparts with
3. NEILL’S RAT grey bases, red-brown tips;
Leopoldamys neilli, p. 386 bicoloured tail slightly longer
HB 200–235. Similar to than HB.
L. edwardsi, but smaller;
upperparts mottled black and 8. INDOCHINESE MAXOMYS
brown; tail dark above, white Maxomys moi, p. 389
below and at tip. HB 140–215. Bright orange
upperparts; fur soft, without
4. RAJAH MAXOMYS spines; sharply demarcated
Maxomys rajah, p. 387 white below; bicoloured tail;
HB 165–225. Upperparts only five footpads.
brown, sometimes tinged
orange; white below, often
with brown streak in middle;
fur with extensive spines; tail
shorter than HB, dark above,
pale below.

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 Plate 77: TYPICAL AND VANDELEURIA MICE
1. ASIAN HOUSE MOUSE 5. SHORTRIDGE’S MOUSE
Mus musculus, p. 390 Mus shortridgei, p. 392
HB 65–90. Dark grey-brown HB 95–120. Fur with many
above; only slightly paler below; stiff spines; grey-brown above;
soft fur; tail all dark, slightly greyish-white below; tail short,
longer than HB. dark grey above, pinkish below.

2. LITTLE INDIAN FIELD 6. RICEFIELD MOUSE

MOUSE Mus caroli, p. 391
Mus booduga, p. 392 HB 65–85. Small; rich brown
HB 60–75. Small; pointed to brownish-grey above; white
rostrum; light brown above; or white with grey bases below;
white below; short bicoloured tail as long as HB, dark above,
tail; incisors curved backwards. white below; incisors orange,
curved forwards.
3. COOK’S MOUSE
Mus cookii, p. 392 7. INDOCHINESE SHREWLIKE
HB 80–105. Relatively large; MOUSE
brown to grey-brown above; Mus pahari, p. 392
pale grey with dark grey bases HB 75–105. Long, shrew-like
below; bicoloured tail slightly nose; small eyes and ears;
shorter than HB. spiny fur; bluish-grey to brown-
grey above; white with grey
4. FAWN-COLOURED MOUSE bases below; tail as long as HB,
Mus cervicolor, p. 391 all grey.
HB 70–95. Brownish-grey
above; white to pale grey below; 8. ASIAN LONG-TAILED
tail short, mid-grey above, paler CLIMBING MOUSE
below; pale buff incisors angled Vandeleuria oleracea, p. 393
forwards. Fragile-tailed Mouse, HB 55–85. Small; soft fur;
M. fragilicauda (not illustrated), extremely long, all-brown tail;
looks very similar but has longer tail with loose skin. orange- to pinkish-brown above,
blending to white below; nail
instead of claw on outer toe.

Note Mus can be distinguished from small rats by having all footpads round.

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 Plate 78: MARMOSET-RATS, MICE AND PYGMY-DORMOUSE
1. GREATER MARMOSET-RAT 5. ASIAN HARVEST MOUSE
Hapalomys longicaudatus, Micromys erythrotis, p. 393
p. 390 HB 46–68. Very small with
HB 150–170. Soft woolly fur; rounded head; small ears; grey-
grey-brown above, orange on brown above, silvery below; thin
flanks, white below; short face; tail slightly longer than HB.
nail instead of claw on hind toe;
ears with tufts of long brown
hairs; long dark tail. 6. SOUTH CHINA WOOD
MOUSE
2. LESSER MARMOSET-RAT Apodemus draco, p. 394
Hapalomys delacouri, p. 390 HB 80–105. Soft fur; yellow-
HB 100–135. Similar to brown to orange-brown, grey
H. longicaudatus, but smaller; bases above; contrasting white
orange-brown above, without below; long dark tail paler
contrasting flank colour. below; large ears. Large-eared
Wood Mouse, A. latronum (not illustrated), is larger
with a longer toothrow.
3. INDOMALAYAN PENCIL-
TAILED TREE-MOUSE 7. SOFT-FURRED PYGMY-
Chiropodomys gliroides, p. 393 DORMOUSE
HB 70–105. Small; nail on first Typhlomys cinereus, p. 395
toe; soft thick fur; red-brown to HB 70–100. Soft fur; dark
grey-brown above; white below; brown-grey above, buff below;
long tail with tuft of fur at tip. long thin tail; narrow feet; cusps
on molars merged into ridges.
4. VERNAY’S CLIMBING
MOUSE
Vernaya fulva, p. 394
HB 60–80. Small; very long,
dark tail; yellow-brown above;
grey with buff tips below; all
toes with claws, not nails.

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 Plate 79: VOLES
1. FORREST’S MOUNTAIN 3. PÈRE DAVID’S VOLE
VOLE Eothenomys melanogaster,
Neodon forresti, p. 396 p. 396
HB 95–115. Short rounded HB 90–100. Dark brown to
ears; short bicoloured tail; very blackish above; slate grey
dark brown above; pale brown below, sometimes with buff
below. tinge; tail <40% of HB.

2. CLARKE’S VOLE 4. KACHIN VOLE

Microtus clarkei, p. 396 Eothenomys cachinus, p. 396
HB 105–120. Dark brown HB 110–125. Bright tawny-
above; slate grey with silvery brown above; grey with buff tips
tips below; tail 50% of HB, below; tail 40–50% of HB.
sparsely covered with short
hairs.

Note Voles can be distinguished from mice by their short rounded ears and the triangular cusps on their molars.

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 Plate 80: BAMBOO RATS AND KHA-NYOU
1. CHINESE BAMBOO RAT 4. LESSER BAMBOO RAT
Rhizomys sinensis, p. 397 Cannomys badius, p. 397
HB 230–450. Medium to very HB 150–265. Small; soft
large; thick soft fur; buffy-brown dense fur reddish-brown with
to reddish-brown with grey grey bases; often with white
bases; short tail (<25% of HB); patch on forehead; ears largely
posterior footpads separate. hidden; short tail.

2. INDOMALAYAN BAMBOO 5. KHA-NYOU

RAT Laonastes aenigmamus, p. 398
Rhizomys sumatrensis, p. 398 HB 210–290. Rat-like head;
HB 280–480. Medium to very bushy, squirrel-like tail; dark
large; sparse coarse hair; pale greyish-black with variable grey
brownish-grey; medium-length frosting.
tail (35–50% of HB); posterior
footpads joined together.

3. HOARY BAMBOO RAT

Rhizomys pruinosus, p. 397
HB 260–350. Medium-sized;
dark brown to grey-brown fur
frosted with white; two separate
footpads; medium-length tail
(35–40% of HB).

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 Plate 81: PORCUPINES, HARES, RABBIT AND PIKA
1. MALAYAN PORCUPINE 5. CHINESE HARE
Hystrix brachyura, p. 399 Lepus sinensis, p. 401
HB 590–720. Largely black HB 350–450. Similar to
with white bands on quills; L. peguensis, but tail brown on
sometimes with slight crest on top; ears sometimes lack black
neck; tail short with rattle-like tips; incisors with V-shaped
quills. groove.

2. BRUSH-TAILED 6. ANNAMITE STRIPED

PORCUPINE RABBIT
Atherurus macrourus, p. 399 Nesolagus timminsi, p. 401
HB 380–520. Greyish-brown HB 350–400. Pale brown to
to brown; long spines on back; brownish-grey above; broad
medium-long scaly tail with tuft black or dark brown stripes on
of rattle-like quills at tip. sides of back and across lower
back; orange rump; relatively
3. LONG-TAILED PORCUPINE short ears.
Trichys fasciculata, p. 399
HB 375–435. Resembles large 7. MOUPIN PIKA
rat; relatively short, flattened Ochotona thibetana, p. 400
spines; long tail with brush of HB 140–180. Small; rounded
flattened spines at tip. ears; no tail; yellow-brown to
dark brown above; light grey to
greyish-yellow below; pale buff
4. BURMESE HARE patch behind ears. Forrest’s
Lepus peguensis, p. 400 Pika, O. forresti (not illustrated),
HB 360–500. Long ears with is similar, but with darker underparts and grey
black tips; mottled brown patch behind ears.
above; reddish-orange patch
on nape; tail black on top, grey
on sides, white below; incisors
with Y-shaped grooves.

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 Order PHOLIDOTA
Family MANIDAE PANGOLINS
Pangolins are readily recognised by their very distinctive scales, which cover the entire upperparts including
the tail. The jaws are elongate and lack teeth (Fig. 12). When they walk, they curl their front claws
inwards, producing a very distinctive footprint (Fig. 13). The infant travels clinging to the upper side of the
base of its mother’s tail. The two species in Asia are similar, distinguished by differences in tail length,
conspicuousness of the ears and relative size of the front claws. All pangolins are threatened by illegal
wildlife trade, especially because of false beliefs about the medicinal values of their scales. Throughout the
region, illegal trade has been very heavy, and populations have declined dramatically, such that both Asian
species are now considered Critically Endangered, indicating a high risk of extinction.

SUNDA PANGOLIN the ground. Tail wrapped around body when animal
Manis javanica PLATE 29 is disturbed, to protect the non-scaly underparts.
Measurements HB 400–650, T 350–560 External ear lobes inconspicuous, reduced to a
(80–90% of HB), HF 61–97. Wt up to 7kg small ridge; scales on top of head extend nearly
Identification Distinctive brownish, scaly mammal, to nostrils; front claws less than 50% longer than
with long claws. Head and tail long and tapered, hind claws; prehensile tail relatively long, with
held below level of body when animal is travelling on about 30 scales along edge and a well-developed,
smooth, glandular pad on underside to grip around
branches. Similar species Chinese Pangolin,
M. pentadactyla, has relatively large, conspicuous
ears, scales on head not reaching nostrils, longer
front claws and shorter tail with fewer than
20 scales along edge.
Ecology and habitat Usually nocturnal, sleeping
during the day in underground burrows. Food
consists exclusively of ants and termites taken from
nests in trees, on the ground or below ground.
0 50 Insect nests are opened with the strongly clawed
feet and the contents licked up with the long, sticky
Fig. 12 Pangolin skull. tongue. Occurs in tall and secondary forests as well
as cultivated areas, including gardens. Most often
seen on roads at night, where it is slow-moving
and conspicuous, although the eyes reflect very
little torchlight.
Distribution SE Asia: S Myanmar, Thailand, S Laos,
Vietnam, Cambodia and Peninsular Malaysia. Also
Sumatra, Java, Borneo and adjacent small islands.
Status Critically Endangered. Populations have
declined more than 90% in some parts of range,
F due to illegal trapping and hunting.

H CHINESE PANGOLIN
Manis pentadactyla PLATE 29
0 50 Measurements HB 400–580, T 250–380 (50–
70% of HB)
Fig. 13 Footprints of Pangolin. Identification Upperparts, including tail, covered

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 with large brown scales. Scales on top of head termites. Known from tall and secondary forests
extend only partway to nostrils; front feet with very and cultivated areas, including gardens. In Laos,
long claws, about twice as long as hind claws; may be found at higher altitudes than M. javanica,
external ear lobes well developed and conspicuous; although both species may have overlapped in
tail relatively short with 16–19 scales along edge, some areas.
and only small pad under tip. Similar species Distribution SE Asia: N Myanmar, N Thailand, Laos
Sunda Pangolin, M. javanica, has smaller ears, and Vietnam. Also E Nepal, E Bangladesh, NE India,
scales on head extending to nostrils, shorter front C and S China and Taiwan.
claws and longer, more flexible tail. Status Critically Endangered. Populations have
Ecology and habitat Mainly nocturnal, with declined dramatically due to illegal trapping and
similar habits to M. javanica. Feeds mainly on hunting.

Order INSECTIVORA
Gymnures, moles and shrews
The order Insectivora includes a mixture of small mammals with several common characters including large
numbers of teeth, usually relatively undifferentiated, with rounded or pointed tops (e.g. Fig. 14, 16, 17 or
18). In this respect, they differ from the rodents, which they superficially resemble: rodents have chisel-like
front teeth (incisors), with a long diastema (gap) behind them, and cheek teeth with complex ridges on the
surface (see e.g. Fig. 65 or 69). Most insectivores typically have a more pointed muzzle than rodents, and
their front feet typically each have five long digits with sharp claws, while in rodents the inner digit on each
front foot is short, with a flat nail instead of a sharp claw.
Recent research suggests that the order Insectivora is an artificial group, and the different families in this
group are not necessarily one another’s closest relatives. Instead, similarities may be due to retention of traits
that characterised the primitive ancestors of all current mammals. Some scientists have suggested splitting
them into several different orders. Among the families in the region, the family Erinaceidae is sometimes
placed in the order Erinaceomorpha, while the families Talpidae and Soricidae are sometimes placed in
the order Soricomorpha. However, there is not yet full agreement on how the different families should be
grouped, so to avoid confusion they are retained together in this book.

Family ERINACEIDAE GYMNURES

Four species of gymnure occur in South-east Asia.
The Moonrat is very distinctive, with its large size
and black-and-white coloration. The three smaller
gymnures are somewhat similar to each other, but
can be distinguished by tail length and ear size. They
are distinguished from shrews by their generally
larger size and larger, more conspicuous external
ears. The skull has a well-developed zygomatic
arch, unlike that of shrews (Fig. 14). Most species
have 44 teeth (3 incisors, 1 canine, 4 premolars
and 3 molars on each side of both the upper and
lower jaw), although H. sinensis has one fewer
premolar on each jaw. The cheek teeth are relatively
rectangular with low cusps.

Fig. 14 Skull of Hylomys suillus.

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 MOONRAT SHORT-TAILED GYMNURE
Echinosorex gymnurus PLATE 1 Hylomys suillus PLATE 1
Measurements HB 320–400, T 205–290, Measurements HB 120–140, T 19–30 (14–24%
HF 65–75. Wt 870–1,100g of HB), HF 20–23, E 15–20. Wt 50–70g.
Identification Large insectivore with elongate Skull: gl 33.5–35.6
muzzle. Front part of body white to greyish- Identification Resembles a large shrew with an
white; remainder of body largely black, often with elongate muzzle, but with conspicuous rounded
extensive pale grey frosting. Fur with long, coarse ears and a distinctive short, slender, nearly hairless
guard hairs, appearing rather shaggy; soft, woolly tail. Upperparts dark brown to reddish-brown to
underfur. White variable in extent, usually including grey-brown. Grey tinge due to grey base to hairs;
head and at least part of back, often extending in underparts light grey to greyish-white, the hairs with
a triangular wedge down lower back. Usually has grey base and white tip. Fur with some flattened
dark marks around eye, which may form a dark spines. Skull has four upper and lower premolars
mask. In Borneo, most individuals are all white on each side.
with scattered black hairs, but white animals are Similar species Chinese Gymnure, H. sinensis, has
rare on the mainland. Tail long, scaly, with very longer tail; Large-eared Gymnure, H. megalotis, has
short hairs, dark at base, whitish or pinkish near longer, thicker tail, larger ears.
tip. Has a distinct pungent odour with strong Ecology and habitat Occurs mainly in hill or
ammonia content, different from the musky smell montane forest with dense undergrowth, but
of carnivores. Footprints are relatively rounded, with sometimes in lowlands. Mainly terrestrial, though
five distinct toes on all feet (Fig. 15a). sometimes climbs low bushes. Active both day
Ecology and habitat Nocturnal and terrestrial. and night. Feeds on arthropods and earthworms.
Roosts in hollow logs, under roots of trees, or in Shelters in nests of dead leaves made in hollows in
abandoned burrows during the day. Does not dig the ground, or under rocks.
its own burrows. Occurs in lowland forest, including Distribution SE Asia: Myanmar, Laos, Vietnam,
mature forest, as well as some secondary forest, Cambodia, Thailand and Peninsular Malaysia. Also
plantation and mangrove. Prefers damp areas, SW China (Yunnan), Sumatra, Java, Borneo and
often entering water. Feeds on earthworms, frogs, some small islands.
crustaceans, insects and other small animals. Status Least Concern.
Distribution SE Asia: peninsular Myanmar, Thailand
and Malaysia. Also Sumatra and Borneo. CHINESE GYMNURE
Status Least Concern, but probably declining owing Hylomys sinensis PLATE 1
to loss of intact forest. Measurements HB 110–125, T 63–73 (50–63%
of HB), HF 24–27, E 18–19. Skull: gl 31.1–33.6
Identification Small gymnure with thin, scaly tail
a b
of medium length. Upperparts mottled dark brown
with a slightly reddish tinge; underparts paler and
greyish, the hairs with grey base and white to buffy
tip. Fur with some flattened spines. Tail pigmented
F above, pale below. Skull has three lower and three
F to four upper premolars on each side. Taxonomic
notes Species sometimes placed in genus
Neotetracus. Similar species Other gymnures
H
H have four upper and lower premolars; Short-tailed
Gymnure, H. suillus, has much shorter tail; Large-
0 50 eared Gymnure, H. megalotis, has thicker, longer
tail, larger ears.
Fig. 15 Footprints of Echinosorex gymnurus (a) Ecology and habitat Mainly or entirely terrestrial.
and a typical treeshrew, Tupaia sp. (b). Found in cool damp forests with fallen logs and

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 rocks for cover. Appears to be found at higher buff or buff and black tip; lacks spines. Nose very
altitudes than H. suillus where ranges overlap, but elongate. Ears large and rounded. Tail thick and
also in lowland forest elsewhere. Tolerant of some long. Teeth relatively robust with four upper and
forest disturbance. lower premolars on each side. Taxonomic notes
Distribution SE Asia: N Myanmar and N Vietnam. First described in 2002. Similar species Other
Also S China (Sichuan and Yunnan). gymnures have shorter tail, smaller ears and some
Status Least Concern. spines in their pelage.
Ecology and habitat Occurs in limestone karst
LARGE-EARED GYMNURE areas.
Hylomys megalotis NOT ILLUSTRATED Distribution SE Asia: Central Laos.
Measurements HB 115–135, T 83–91 (65–75% Status Data Deficient; known only from the
of HB), HF 20–21.5, E 21–23. Skull: gl 36.4–39.2 Khammouane limestone area. Might be expected to
Identification Overall colour grey to brownish-grey; inhabit other limestone areas in the region.
individual hairs long, soft, with grey base with

Family TALPIDAE MOLES

Moles spend much of their time underground and have adaptations for digging, including enlarged front
feet. Their eyes are greatly reduced and they rely instead on their senses of smell and touch to locate food.
They can sometimes be located by raised tunnels, as they often dig close to the surface. The South-east
Asian species are currently separated into five genera, two of which (Uropsilus and Scaptonyx) are highly
distinctive, and two of which (Euroscaptor and Parascaptor) are very similar to each other.

Genus Uropsilus The moles in this genus parts of legs and arms, as well as hands and feet,
superficially resemble shrews more closely than nearly hairless and scaly; claws curved but short
they do moles, with front feet not especially enlarged and weak. One lower incisor, four upper and lower
and externally visible ears. Readily distinguished premolars. Similar species All other genera of
from true shrews in region by very thin, elongate moles have enlarged front feet with strong claws;
snout and tiny eyes that are largely hidden in fur. shrews have small but conspicuous eyes, and no
Dental formula varies among species in the genus: shrews in the region have such a long, thin snout
2/(1–2) 1/1 (3–4)/(3–4) 3/3. Up to four species and long tail.
currently recognised, all occurring in C and S China, Ecology and habitat Found in hill forest at altitudes
but only one is known from South-east Asia. The of 1,200–4,500m. Unlike other moles, believed to
other species can be distinguished by differences behave more like a shrew, foraging mainly in leaf
in dental formula, but these can be difficult to litter rather than digging underground. Presumably
determine as some teeth, even when present, are feeds mainly on invertebrates.
very small and inconspicuous. Distribution SE Asia: N Myanmar. Also adjacent
areas of China (Sichuan and W Yunnan).
SLENDER SHREW-MOLE Status Least Concern.
Uropsilus gracilis PLATE 1
Measurements HB 65–90, T 60–80 (90–100% Genus Euroscaptor These moles closely resemble
of HB) the European moles, Talpa, and have been
Identification Small, shrew-like animal with a very sometimes placed in that genus. They have greatly
long, thin snout formed from two elongate tubular enlarged front feet, wider than they are long, turned
nostrils with a groove along the top. External ear sideways with long claws. The nose narrows to a
small, but large enough to stick out beyond fur. Eyes small, skinny tip; eyes are tiny, completely hidden
tiny and hidden in fur. Fur soft, upperparts dark in the fur; no externally visible ears. Skull elongate
brown to greyish-brown, underparts contrasting with narrow zygomatic arches (Fig. 16). Dental
slate-grey, sometimes with paler tip. Tail long formula is usually 3/3 1/1 4/4 3/3, but sometimes a
and scaly with short, inconspicuous hairs. Lower tooth is missing. The anterior upper three premolars

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 are all small, but anterior lower premolar is larger Ecology and habitat Found in forest and cultivated
than canine. The several species in this genus are areas including tea plantations and vegetable
externally all very similar, with fur colour varying gardens, especially in areas with loose sandy soils.
within and among species, and hence not a reliable Distribution SE Asia: montane areas of N Thailand.
identification character. The distribution of most Likely also occurs in Myanmar or Laos, but no
species is poorly known, as they are rarely trapped published locality records.
and difficult to identify; in some cases, the same Status Least Concern.
specimens have been reported as different species
in different literature sources. Recent research MALAYAN MOLE
based on genetics and morphology has recognised Euroscaptor malayana NOT ILLUSTRATED
several additional species. In general, only one Measurements HB 128–40, T 4.5–8.5 (4.5% of
species has been found in any given locality/ HB), HF 14.5–16.5. Wt 44–71g.
habitat. In N Vietnam, some similar species appear Skull: gl 30.5–32.5
to separate by altitude. Identification Similar to Kloss’s Mole, E. klossi,
with broad-based snout, club-shaped tail, broad
KLOSS’S MOLE palate, relatively large rectangular molars; differing
Euroscaptor klossi PLATE 1 in darker, almost black fur, very short tail, narrow
Measurements HB 130–136, T 9.5–10.5 (7.5% of post-orbital bridge in skull. Only species within its
HB), HF 15. Wt 48–55g. Skull: gl 31.5–33.5 range. Taxonomic notes Formerly considered same
Identification Fur soft and dark grey to brownish- species as Kloss’s Mole, E. klossi, or Himalayan
grey, slightly paler underneath. Enlarged front feet Mole, E. micrura, but genetic and morphological
with long claws. No visible eyes or ears. Tail short studies have shown it is distinct.
and club-shaped (thicker near tip than base) with Ecology and habitat Known only from tea
sparse long hairs. Snout triangular. Molars relatively plantations and adjacent gardens in Cameron
large, rectangular, 8–9mm across. Taxonomic Highlands at altitudes of 1,300–2,000m. Tunnels
notes Previous records from Malaysia and Vietnam underground, but usually near surface, so tunnels
now considered to be other species. are easily visible. Feeds mainly on earthworms as
Similar species Large Chinese Mole, E. grandis, well as arthropods such as insects, and some small
is larger; other species differ in tail length and vertebrates including snakes and lizards.
proportions. Distribution SE Asia: Peninsular Malaysia.
Status Not assessed; known only from Cameron
Highlands where it is locally common, but may
occur elsewhere in highlands of Malaysia.

LARGE CHINESE MOLE

Euroscaptor grandis PLATE 1
Measurements HB 150, T 10, HF 18.
Skull: gl 35.5–37
Identification Similar to Kloss’s Mole, E. klossi, but
larger. Tail short and club-shaped (thicker near tip
than base).
Ecology and habitat Occurs in forested areas in
China.
Distribution SE Asia: likely in NE Myanmar. Also
S China (C Sichuan and W Yunnan).
Status Least Concern, but status in region uncertain
due to limited number of records and taxonomic
Fig. 16 Lateral and ventral views of skull uncertainty.
of Euroscaptor klossi.

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 ORLOV’S MOLE differences in teeth from the nominate form in
Euroscaptor orlovi NOT ILLUSTRATED S Vietnam.
Measurements HB c.133–144, T 14–17 (11% of Ecology and habitat Type specimen was found
HB). Wt 42–77g. Skull: gl 31.0–34.5. on forest floor in red basaltic soil at 800m altitude.
Identification Externally similar to other Euroscaptor Distribution SE Asia: Vietnam.
moles. Distinguished by slender snout, relatively Status Data Deficient; known from several localities
long, thick tail that protrudes well beyond body. in C and S Vietnam, but abundance and trend
Skull has narrow, tapered rostrum, triangular unknown.
molars. Taxonomic notes Newly described in
2016 as distinct from E. longirostris, which is SHORT-TAILED MOLE
now considered to occur only in China. Similar Euroscaptor subanura NOT ILLUSTRATED
species Differs in dental characters and genetics Measurements HB 113–130, T 4–5, HF 13.5–
from Kuznetsov’s Mole, E. kuznetsovi. 15.5. Wt 33–45g. Skull: gl 30.6–32.0
Ecology and habitat Montane forest. Identification Similar to Euroscaptor parvidens, with
Distribution SE Asia: NW Vietnam. narrow palate, but tail is even shorter, ribs are broad,
Status Not assessed; only known from a few last upper premolar (P4) has well developed anterior
records. lobe such that it is more rectangular than triangular.
Taxonomic notes Newly described in 2012.
KUZNETSOV’S MOLE Ecology and habitat Recorded from semi-deciduous
Euroscaptor kuznetsovi NOT ILLUSTRATED forest around limestone hills, generally at lower
Measurements HB 132–136, T 15–17 (11–12.5% elevations (200–1,000m).
of HB) Distribution SE Asia: N Vietnam.
Identification Externally similar to Orlov’s Mole, Status Data Deficient; only a few records, so extent
Euroscaptor orlovi, with relatively long club-shaped of range unknown.
tail and narrow snout. Distinguished by slightly
wider, tapered rostrum in skull; slightly broader, Genus Parascaptor Very similar to Euroscaptor.
triangular molars; first two upper premolars (P1 and Main distinguishing feature is reduced dental
P2) similar in height (P1 is higher than P2 in E. orlovi). formula 3/3 1/1 3/3 3/3, with one fewer small
Taxonomic notes Newly described in 2016; differs premolar and remaining three upper premolars
genetically from other species. relatively larger than in Euroscaptor; also distinct
Ecology and habitat Montane forest. groove in skull behind auditory bullae.
Distribution SE Asia: NW Vietnam. Also S China.
Status Not assessed; very little information from BLYTH’S MOLE
region. Parascaptor leucura NOT ILLUSTRATED
Measurements HB 130–140, Tail c.10% of HB.
SMALL-TOOTHED MOLE Skull: gl 27–31
Euroscaptor parvidens NOT ILLUSTRATED Identification Similar in appearance to Euroscaptor
Measurements HB 135–150, T 5.5–9 (5% of HB), moles, with short, club-shaped tail tipped with long,
HF 13.5–15.5. Wt 37–64g. Skull: gl 30.5–34.5 often white hairs; distinguished by having only three
Identification Externally similar in appearance to relatively larger premolars in each jaw; relatively
other Euroscaptor moles. Distinguished by short tail, wide molars; groove in base of skull behind auditory
largely hidden in fur except a few long protruding bullae.
hairs at tip of tail; slender snout and corresponding Ecology and habitat Montane forest at 1,000–
slender skull (ratio of palatal length to breadth 2,500m altitude.
= 0.53); small, relatively triangular molars; last Distribution SE Asia: N Myanmar and N Laos. Also
upper premolar (P4) with a well-developed cingulum adjacent areas of Assam and S China (Yunnan and
cusp. Taxonomic notes Population in C Vietnam SW Sichuan).
described in 2016 as a distinct subspecies, E. p. Status Least Concern, but poorly known in region
ngoclinhensis, based on slightly smaller size and as there are only a few records.

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 Genus Mogera Generally similar to Euroscaptor Genus Scaptonyx A distinctive mole with forefeet
moles, but one fewer lower incisor, with dental only moderately enlarged, elongate rather than
formula 3/2 1/1 4/4 3/3. Only one species known rounded as in Euroscaptor and Parascaptor. Tail
from region, but several additional species occur in relatively long. Ear openings large but with no
eastern China, Korea, Japan and Taiwan. external pinnae. Eyes minute, covered with a thin
layer of translucent skin.
LA TOUCHE’S MOLE
Mogera latouchei NOT ILLUSTRATED LONG-TAILED MOLE
Measurements HB 103–115, T 12–15 (11–14% Scaptonyx fuscicaudus PLATE 1
of HB), HF 13.5–15.5. Wt 33–52g Measurements HB 65–90, T 30–50 (c.50–60%
Externally similar in appearance to Orlov’s Mole, of HB)
Euroscaptor orlovi, with short, dark grey fur, narrow Identification A distinctive small mole with elongate
snout, thickened cylindrical tail with long hairs that snout, moderately enlarged but still elongate front
protrude well beyond body fur. Distinguished by feet with long flattened claws for digging; hind feet
having only two lower incisors. Skull is slender in smaller but still with large, broad claws; hind and
rostral region, with long, wide openings to auditory forefeet covered with small scales. Tail moderately
bullae. Taxonomic notes Sometimes considered a long, about half length of head and body, covered
subspecies of M. insularis. with scales and sparse long hairs. Fur soft and
Ecology and habitat Has been trapped in farmland velvety, uniform dark slate-grey.
and disturbed forests at 600–1,700m altitude. Ecology and habitat Known from disturbed and
Distribution SE Asia: N Vietnam. Also adjacent primary hill and montane forest at 1,300–4,100m
areas of S China. altitude. Presumably spends much of its time
Status Least Concern; appears to be locally underground, but has been found on the surface.
common, although known from only a few Distribution SE Asia: N Myanmar, N Vietnam. Also
localities. adjacent parts of S China.
Status Least Concern, although few records from
region.

Family SORICIDAE SHREWS

Shrews include the smallest mammals in the world apart from some bats. They have rather short legs,
a sparsely haired tail, short fur and small eyes. Many of the smaller species are difficult to identify.
Even distinguishing among genera can be difficult, unless the animals have been prepared as museum
specimens, as the differences among species and among genera are often based on characters such as
number and size of the teeth.
All shrews have a single enlarged incisor on each upper jaw, followed by two to five small single-
cusped (unicuspid) teeth (canine and premolars), and four larger multicusped teeth (one premolar and three
molars), although the posterior molar is usually small (Figs 17–18, 20–22). The number of unicuspid teeth is
important for identifying genera, but sometimes the posterior ones can be small and hard to see. The colour
of the teeth can also help to distinguish genera; some groups have a reddish-orange tip to some of the teeth,
especially the incisors and some of the unicuspids.

Genus Sorex Small shrews (HB 50–80) with tail Identification Small shrew with long tail. Upperparts
similar length to head and body. Teeth with reddish- mid-brown, hairs with grey base that show through;
orange tip; five unicuspid teeth, all fairly similar in size. dark brown stripe down middle of back; underside
slate-grey with buff, light brown or white tip
LESSER STRIPE-BACKED SHREW to hairs. Teeth with red-orange tip; five small
Sorex bedfordiae PLATE 2 unicuspid teeth. Similar species Greater Stripe-
Measurements HB 50–70, T 45–70 (85–105% of backed Shrew, S. cylindricauda, which is not known
HB), HF 11–15. Skull: gl 17.0–18.6, ut 7.0–7.8 from region but occurs nearby in China, is similar in

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 appearance but larger (HB 70–77, gl 19.5–20.2) and below, dark slate-grey mixed with brown.
with proportionately shorter tail (80% of HB). Relatively small, narrow skull. Third upper unicuspid
Ecology and habitat Montane forest at altitudes of relatively small, only about 25% longer than small
2,000-4,000m in China. fourth unicuspid. Similar species Indochinese
Distribution SE Asia: N Myanmar. Also central Short-tailed Shrew, B. griselda, has larger, broader
S China and E Nepal. skull, third upper unicuspid relatively larger.
Status Least Concern. Ecology and habitat High montane forest up to
2,400m altitude.
Genus Blarinella Small shrews with short tail, Distribution SE Asia: NE Myanmar. Also adjacent
about half body length. Anterior teeth with distinct parts of China (W Yunnan).
reddish-orange tip. Large, well-developed claws. Status Least Concern; appears to be relatively
Five unicuspid teeth, but posterior teeth small and common in appropriate habitat.
compressed inwards, so they are not visible from
the side (Fig. 17). Formerly considered to include INDOCHINESE SHORT-TAILED SHREW
only one species, B. quadraticauda, but recent work Blarinella griselda PLATE 2
has shown that there are three distinct species, with Measurements HB 65–80, T 34–43, HF 11–13.
B. wardi and B. griselda occurring in the region, Skull: gl 20.2–21.0, breadth 9.1–9.4, ut 8.6–9.1,
while B. quadraticauda is confined to China. M–M 5.3
Identification Small, stout-bodied shrew similar
MYANMAR SHORT-TAILED SHREW to B. wardi, with uniformly dark brown fur above
Blarinella wardi NOT ILLUSTRATED and below. Relatively larger, broader skull. Third
Measurements HB 60–75, T 32–40 (about 50% of upper unicuspid not particularly short, about twice
HB), HF 10–14. Skull: cbl 18.4–19.8 (mean 19.1), height of small fourth unicuspid. Similar species
breadth 7.8–9.0 (8.4), ut 7.0–8.4 (8.1), M–M Myanmar Short-tailed Shrew, B. wardi, is smaller
4.5–5.0 (4.8) and has a narrower skull, third upper unicuspid is
Identification Small, stout-bodied shrew with relatively smaller.
relatively short tail, five unicuspid teeth (Fig. 17a); Ecology and habitat Recorded from montane
incisors and at least some unicuspids with reddish- bamboo forest, 1,500–1,700m altitude in
orange tip. Body colour essentially the same above N Vietnam.
Distribution SE Asia: N Vietnam. Also China (SW
Yunnan, Gubei, Hansu).
a Status Least Concern, but poorly known in region.

Genus Episoriculus Medium-small to small

shrews; incisors and some other upper teeth with
reddish-brown tip; only four upper unicuspid teeth,
of which posterior tooth is minute, not visible from
side and often hard to see at all. Posterior molar
relatively well developed, more than half length of
first upper molar. Relatively long tail (80–170% of
b
head and body length). Has often been included in
the genus Soriculus, which is now considered to
include only one species, S. nigrescens. The latter
occurs in the Himalayas and Assam in NE India,
near the edge of the region, and might occur in
N Myanmar. It can be distinguished by a relatively
stout body (HB 80–90), with a short tail (about half
Fig. 17 Ventral view of skulls of Blarinella wardi (a) HB), relatively long front claws (2.5–4mm), and a
and Chodsigoa caovansunga (b). relatively small third lower molar.

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 HODGSON’S BROWN-TOOTHED SHREW of HB), HF 14–17, E 8–10.5. Skull: gl 16.9–18.3,
Episoriculus caudatus PLATE 2 ut 7.2–7.7
Measurements HB 50–75, T 34–66 (60–110% Identification Upperparts grey, with narrow
of HB), HF 11–14.5, E 5–9. Skull: gl 16.7–19.1, brownish tip on some hairs, appearing grey
ut 7.1–8.5 with brown flecks. Tail much longer than body,
Identification Upperparts brown, individual hairs dark above, pale below. Similar species Other
with grey base; underparts similar but sometimes Episoriculus have shorter tail; Van Sung’s Shrew,
with paler tip. Incisors relatively small with small Chodsigoa caovansunga, has only three upper
reddish-brown tip. Taxonomic notes Considerable unicuspid teeth, shorter tail; Lowe’s Brown-toothed
geographic variation in size may indicate that this is Shrew, C. parca, has larger skull.
a complex of species; form in Myanmar is relatively Ecology and habitat Terrestrial, probably feeding
small and dark. Similar species Bailey’s Brown- among dead leaves. Has been found among dwarf
toothed Shrew, E. baileyi, is larger, with relatively bamboo, scrub and grasses in montane forest.
large upper incisor; Long-tailed Brown-toothed Distribution SE Asia: NW Myanmar and N Vietnam.
Shrew, E. macrurus, has much longer tail. Also Nepal, NE India, W and S China.
Ecology and habitat Montane forests at altitudes Status Least Concern.
of 1,800–3,600m.
Distribution SE Asia: N Myanmar, N. Vietnam. Also Genus Chodsigoa Similar to Episoriculus shrews,
S China, India, Nepal. but with only three unicuspids, all well developed
Status Least Concern. (Figs 17, 18). Incisors and some unicuspids with
reddish-brown tip.
BAILEY’S BROWN-TOOTHED SHREW
Episoriculus baileyi NOT ILLUSTRATED LOWE’S BROWN-TOOTHED SHREW
Measurements HB 63–81, T 60–75 (85–110% Chodsigoa parca PLATE 2
of HB), HF 13–14.5, E 7–10. Skull: gl 19–20.6, Measurements HB 65–85, T 70–100, HF 15–20,
ut 8.4–9.3 E 6–8. Skull: gl 19.0–21.0, ut 8.1–9.2
Identification Fur soft and thick; upperparts grey Identification Slate-grey above with brownish
base with dark brown tip, appearing overall brown tinge, slightly paler below. Feet distinctly pale with
flecked with grey; underparts grey base with narrow white skin and short, sparse brown hairs. Tail longer
whitish to light brown tip. Incisors large and robust than head and body; dusky brown above, slightly
with extensive reddish-brown tip. Taxonomic paler below; appears naked, but has moderately
notes Formerly included as a subspecies of long, thin hairs, ending in small tuft at tip of tail.
E. leucops, which is now recognised as occurring
only in Nepal. Specimens from Vietnam are slightly
larger than those from Myanmar. Similar species a
Hodgson’s Brown-toothed Shrew, E. caudatus, has
smaller skull with shorter toothrow, relatively small
upper incisors; Long-tailed Brown-toothed Shrew,
E. macrurus, has much longer tail, greyer body fur,
smaller skull.
Ecology and habitat Montane broadleaved and
coniferous forest. b
Distribution SE Asia: N Myanmar, N Vietnam. Also
Sikkim and Assam in NE India, SW China.
Status Least Concern.

LONG-TAILED BROWN-TOOTHED SHREW

Episoriculus macrurus PLATE 2 Fig. 18 Skull profiles of Chodsigoa parca (a) and
Measurements HB 59–66, T 80–100 (130–150% C. caovansunga (b).

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 similar in length to body; bicoloured, dark above,
pale below; appearing naked but actually covered
with very short hairs; no tuft of hairs at tip of tail.
Pads on hind feet very close together or touching
(Fig. 19b). Skull relatively short with low braincase
(Fig. 18b). Similar species Lowe’s Brown-toothed
Shrew, C. parca, is larger, with longer hairs on tail,
pale feet, pads on feet well separated.
Ecology and habitat Terrestrial. Recorded in
Vietnam from montane forest, including areas of
bamboo, at altitudes of 1,300–2,000m.
Distribution SE Asia: N Vietnam. Also S China.
a b Status Data Deficient; recorded only from type
locality in N Vietnam and one site in China.

Genus Anourosorex Round-bodied shrews with

very short tail, no external ear, tiny eyes, moderately
long foreclaws, teeth without red-brown tip. Skull
with only two unicuspids, very small posterior molar.
Apparently spends much of its time burrowing
like a mole. Only one species in region, but three
other species are found in India, Bhutan and
Fig. 19 Undersides of hind feet of Chodsigoa parca (a) China, some of which were formerly considered the
and C. caovansunga (b). same species.

Two pads in middle of hind foot separated by MOLE-SHREW

approximately width of a pad (Fig. 19a). Skull large Anourosorex squamipes PLATE 2
with tall braincase (Fig. 18a). Similar species Van Measurements HB 85–110, T 10–20, HF 12–16,
Sung’s Shrew, C. caovansunga, is smaller with E 0. Skull: gl 23.3–26.3
lower braincase, shorter hairs on tail, pads on hind Identification Medium-sized shrew with very short
feet touching or nearly so; Episoriculus shrews have tail, no external pinnae, minute eyes. Upperparts
an extra small unicuspid in upper toothrow (though mottled brown and grey, individual hairs with grey
this can be hard to see), shorter hairs on tail. base and dark brown tip; hairs on lower rump with
Ecology and habitat Montane forest at altitudes of shiny tip from a glandular secretion; underparts vary
1,200–2,750m. Has been reported near streams, from brown to light grey, hairs with grey base. Tail
including under rocks or logs. short and scaly, no longer than hind feet. Claws on
Distribution SE Asia: NE Myanmar, N Thailand front feet longer than hind feet, but front feet not
and N Vietnam; may be expected in Laos. Also SW enlarged and hind feet slightly larger than forefeet.
China. Similar species Short-tailed moles (Euroscaptor
Status Least Concern. and Parascaptor) have greatly enlarged front feet
with long, broad claws; Slender Shrew-mole,
VAN SUNG’S SHREW Uropsilus gracilis, also has tiny eyes with relatively
Chodsigoa caovansunga PLATE 2 small feet, but has a long tail.
Measurements HB 58–64, T 51–68, HF 14–16. Ecology and habitat Terrestrial and fossorial,
Skull: gl 17.3–18.4, ut 7.6–8.1 burying into ground and leaf litter. Reported mainly
Identification Upperparts dark slate-grey with from montane forest at altitudes of 1,500–3,100m.
brownish tinge formed from short brown tip on Distribution SE Asia: N Myanmar, N Thailand, Laos
hairs; underparts similar but tip pale brown or and N Vietnam. Also NE India (Assam), China.
light grey. Fore and hind feet overall brownish. Tail Status Least Concern.

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 Genus Chimarrogale Medium-large shrews with Status Least Concern, but poorly known in region.
moderately long tail; dense, dark fur with long
silvery guard hairs on hindquarters; tail thickly MALAYAN WATER SHREW
haired; feet with fringe of short, stiff hairs. Very Chimarrogale hantu NOT ILLUSTRATED
small pinnae. Teeth white without red or brown tip; Measurements HB 95–120, T 85–100 (80–100%
three upper unicuspid teeth. Six species, all in Asia, of HB), HF 21–22. Wt 30g. Skull: gl 25–28,
of which three are known from region. mt 11.4–13.4
Identification Upperparts dark grey-brown, with
HIMALAYAN WATER SHREW scattered silver guard hairs, longest and most
Chimarrogale himalayica PLATE 3 conspicuous on hindquarters; underparts greyish-
Measurements HB 110–130, T 90–100 (80–85% brown. Upper incisors with distinct cusp on inside
of HB), HF 19–22. Skull: gl 25–28, ut 11.1–12.5 edge. Taxonomic notes Sometimes included in
Identification Upperparts dark grey-brown with C. phaeura from Borneo, but distinguished by
conspicuous long silvery guard hairs on hindquarters; deeper braincase (C. hantu 7.7–8.0, C. phaeura
underparts slightly paler greyish-brown, not sharply 7.3–7.5). Similar species Himalayan Water Shrew,
demarcated; tail slender but densely haired, usually C. himalayica, lacks cusps on upper incisors, does
dark above and below; feet fringed with short, not overlap in range.
rather stiff hairs. Similar species Styan’s Water Ecology and habitat Lives in or near small forest
Shrew, C. styani, is slightly smaller and has sharply streams in hill forest.
demarcated pale underparts. Distribution SE Asia: Peninsular Malaysia.
Ecology and habitat Lives in or near small forest Status Near Threatened based on limited range,
streams, feeding on aquatic insects and other loss of habitat including degradation of streams.
invertebrates, but may wander through forest away
from streams. Genus Nectogale Medium-sized shrew; tail
Distribution SE Asia: N Myanmar, N Laos and moderately long with several rows of white hairs;
N Vietnam. Also Nepal, N India (Assam), China and hind feet with fringes of short, stiff hairs on edges
Taiwan. of toes. Teeth white without red or brown tip; three
Status Least Concern, but apparently dependent upper unicuspid teeth. No external pinnae.
on relatively clean mountain streams, and therefore
vulnerable to forest loss and pollution. WEB-FOOTED WATER SHREW
Nectogale elegans PLATE 3
STYAN’S WATER SHREW Measurements HB 90–130, T 90–110
Chimarrogale styani PLATE 3 Identification Upperparts slate-grey, with long
Measurements HB 80–110, T 90–100 silvery guard hairs; underparts silvery-white. Tail
Identification Upperparts dark grey-brown with dark with several rows of stiff white hairs: two
conspicuous silvery guard hairs on hindquarters; rows on either side start at base of tail and curve
underparts contrastingly silvery-grey, sharply under tail to meet on underside for distal half; two
demarcated from colour of upperparts. Tail slender further lateral rows extend along middle third; a
but densely haired, dark above, pale below. Feet dorsal row continues along distal third. Feet have
fringed with short, rather stiff hairs. Similar fringes of short white hairs on edges of toes;
species Himalayan Water Shrew, C. himalayica, round disc-like pads on soles of feet. Snout long
is larger, with underparts darker, not sharply and pointed. No visible pinnae. Similar species
demarcated from upperparts; Web-footed Water Styan’s Water Shrew, Chimarrogale styani, has
Shrew, Nectogale elegans, has distinctive pattern of tail more uniformly haired, dark above and pale
rows of white hairs on a dark tail. below.
Ecology and habitat Lives in or near small forest Ecology and habitat Cold mountain streams;
streams at altitudes of 1,500–3,100m. swims and dives to catch aquatic invertebrates
Distribution SE Asia: NE Myanmar. Also SW and and small fish. Sleeps in burrows in stream banks.
SE China. Reported at 900–2,270m altitude.

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 Distribution SE Asia: N Myanmar. Also Nepal, India Sunda Shrew, Crocidura cf. monticola, is slightly
(Sikkim), Bhutan and China. larger, with only three small upper unicuspid teeth;
Status Least Concern, but susceptible to pollution most other shrews are substantially larger.
or degradation of stream habitats. Ecology and habitat Terrestrial.
Distribution SE Asia: Myanmar, Thailand, Laos,
Genus Suncus Medium-large to very small shrews; Vietnam. Also S Europe, N Africa, Madagascar,
tail short to moderately long, with scattered, long through Middle East to India, S China.
erect hairs; teeth white without red or brown tip; Status Least Concern. Although rarely captured,
four upper unicuspid teeth, although last one can apparently widespread.
be small and hard to see (Fig. 20a). About 19
recognised species worldwide, of which three occur MALAYAN PYGMY SHREW
in the region. Suncus malayanus NOT ILLUSTRATED
Measurements HB 37–51, T 22–34 (c.60% of
HOUSE SHREW HB), HF 6.7–8.2, E 4.4–6.5. Wt 1.1–2.3g.
Suncus murinus PLATE 3 Skull: gl 13.5–14.7, ut 5.6–6.1
Measurements HB 90–145, T 46–86 (50–64% of Identification A very small shrew; upperparts
HB), HF 17–23. Skull: gl 27–37, ut 12.5–16.0 dark brown to blackish; underparts more greyish.
Identification Relatively large, thickset shrew. Entire Scattered long pale hairs in fur. Relatively large
body and tail uniform mid-grey to brownish-grey. ears and feet dark brown to black. Tail thickened
Tail thick, especially at base, narrower at tip. Has a at base and tapering to tip, darker above, paler
musk gland, sometimes visible, on middle of each below; covered with dense short hairs especially
side of the body. Taxonomic notes Considerable on underside, and scattered long hairs on basal
variation throughout range, and may represent a two-thirds. Taxonomic notes Formerly considered
complex of species. Similar species Water shrews, part of S. etruscus, but recent genetic studies
Chimarrogale spp., have long slender tail, different confirm it is distinct. Similar species Sunda Shrew,
habitat; most other shrews are smaller. Crocidura cf. monticola, is slightly larger, with only
Ecology and habitat Usually found in or near three small upper unicuspid teeth; Pygmy White-
houses or in disturbed habitats. toothed Shrew, S. etruscus, averages slightly larger
Distribution SE Asia: Myanmar, Thailand, Laos, and does not overlap in range.
Vietnam, Cambodia and Peninsular Malaysia. Also Ecology and habitat Terrestrial; caught in lowland
Pakistan, India, Nepal, S China, Sumatra, Java, rainforest, as well as hill forest in Cameron
Borneo, Sulawesi and smaller Indonesian islands Highlands, Peninsular Malaysia.
and the Philippines. Distribution SE Asia: peninsular Thailand and
Status Least Concern. Widespread and abundant Malaysia.
in disturbed areas; apparently introduced in some Status Least Concern. Although rarely captured,
areas, including the Philippines. known localities are widely distributed.

PYGMY WHITE-TOOTHED SHREW Genus Crocidura Medium to small shrews; tail

Suncus etruscus PLATE 3 moderate to very long, usually with scattered long
Measurements HB 50–56, T 30–40 (60–75% of hairs or bristles; teeth white without red or brown
HB), HF 7.0–8.0. Wt 1.8–2.5g. Skull: cbl 14.5, tip; only three upper unicuspid teeth (e.g. Figs
ut 5.3–6.4 20b, 21). The genus contains over 180 currently
Identification A very small shrew; upperparts recognised species throughout Africa, Europe and
dark grey with fine whitish grizzling; underparts Asia, of which 20 are known from the region.
somewhat paler. Ears and feet dark brown. Tail Many species are represented by relatively few
slightly thickened, brownish, with dense short specimens, and the taxonomy is still somewhat
hairs and scattered long hairs on basal two-thirds. uncertain. Most species are similar in colour and
Taxonomic notes Formerly included Malayan can be difficult to distinguish without examining
Pygmy Shrew, S. malayanus. Similar species the skulls. Recent molecular studies have found

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 as well as in open disturbed areas. Feeds on insects
and other invertebrates.
a b Distribution SE Asia: Myanmar, Thailand, Laos,
Vietnam, Cambodia and N Peninsular Malaysia. Also
NE India and S China.
Status Least Concern.

MALAYAN SHREW
Crocidura malayana NOT ILLUSTRATED
Measurements HB 77–100, T 56–78, HF 13–16.
Wt 10–14.5g. Skull: gl 22.5–24.5, ut 9.2–10.8
Identification Very similar in external appearance
to C. fuliginosa, with dark brown to blackish fur,
but differs genetically and, at least within area
of overlap in range, has slightly larger skull with
Fig. 20 Right upper tooth row of Suncus murinus (a) and broader rostrum and palate: width across outside
Crocidura fuliginosa (b). of upper unicuspid teeth 2.9–3.4 v 2.4–2.7 in
sympatric populations of C. fuliginosa. Also, middle
several previously unrecognised species that are cusp (mesostyle) of second upper molar is simple
genetically quite distinct but which look very similar. or only very shallowly notched in C. malayana, but
Geographic distribution can help with identification, distinctly notched (bifid) in C. fuliginosa.
as many of the most similar species do not overlap Ecology and habitat Terrestrial. Reported from
in range. The similar species sections in the text both hill and lowland forest, in a variety of forested
below emphasise forms that overlap in geographical habitats.
distribution. Distribution SE Asia: peninsular Thailand and
Malaysia and some small offshore islands.
SOUTH-EAST ASIAN SHREW Status Least Concern; most commonly encountered
Crocidura fuliginosa PLATE 3 Crocidura in Peninsular Malaysia.
Measurements HB 70–100, T 60–90 (65–75% of
HB), HF 15–19. Wt 10–16g. Skull: gl 21.5–25.0, PENINSULAR SHREW
ut 9.2–11.0 Crocidura negligens NOT ILLUSTRATED
Identification Upperparts dark brownish-grey; Measurements HB 75–95, T 55–69 (67–92%
underparts slightly paler. Tail brownish, with few of HB), HF 13.5–15.8. Wt 9.5–12.5g. Skull: gl
or no long hairs. Individuals found in peninsular 20.5–22.3, ut 9.2–9.9
Thailand and Malaysia are distinctly smaller than Identification Genetically closely related to
those elsewhere. Taxonomic notes Formerly C. malayana, but has distinctly smaller skull with
considered to include many forms that are now more rounded, globular braincase. Lingual (inside)
considered separate species; as a result, range edge of large upper premolar tends to be rounded.
listed here is much less extensive than suggested Taxonomic notes Has been confused in the past
in some earlier publications; may still represent with C. attenuata, which is genetically very different,
more than one species. Similar species Malayan and is restricted to north of the peninsula. Similar
Shrew, C. malayana, in area of overlap, is slightly species Malayan Shrew, C. malayana, and South-
larger and broader across teeth; Peninsular Shrew, east Asian Shrew, C. fuliginosa, are larger, with
C. negligens, has more rounded lingual edge to large more angular lingual edge to premolar; Grey Shrew,
upper premolar; Hill’s Shrew, C. hilliana, is slightly C. attenuata, does not overlap in range.
smaller with more robust upper unicuspid teeth (Fig. Ecology and habitat Terrestrial. Reported from
21b); other Crocidura in region are smaller. both primary and secondary forest, but specific
Ecology and habitat Terrestrial. Found in forest, habitat requirements and tolerance to habitat
both primary and degraded, from hills to lowlands, degradation unknown.

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 Distribution SE Asia: peninsular Thailand and
Malaysia, S of the Isthmus of Kra, and a few a
offshore islands.
Status Least Concern, though relatively few records.

SUNDA SHREW
Crocidura cf. monticola PLATE 3 b
Measurements HB 53–65, T 24–37, HF 8.5–11.0.
Wt 3–6g. Skull: gl 15.0–17.5, ut 6.6–7.3
Identification Very small shrew with upperparts
dull grey-brown; underparts paler. Tail paler than
body, with scattered long, bristle-like hairs on basal c
10mm. Braincase relatively high and somewhat
domed. Taxonomic notes Recent genetic analyses
indicate the mainland form is not C. monticola,
which is now considered to be endemic to Java, but
the appropriate name has not yet been determined.
Similar species Malayan Pygmy Shrew, Suncus Fig. 21 Lateral views of skulls of Crocidura attenuata (a),
malayanus, has darker fur, shorter hind foot, an C. hilliana (b) and C. fuliginosa (c).
extra small tooth, smaller skull and shorter hind
foot; other Crocidura within its range are larger. Distribution SE Asia: Myanmar, Thailand, Vietnam,
Ecology and habitat Terrestrial. Recorded from Laos and Cambodia. Distribution somewhat
both lowland and hill forest. uncertain, as some previously published records
Distribution SE Asia: peninsular Thailand and under this name may refer to other species such as
Malaysia. Uncertain whether same species occurs C. tanakae. Also S China.
on Sumatra or Borneo. Status Least Concern.
Status Least Concern (as C. monticola).
TAIWANESE GREY SHREW
ASIAN GREY SHREW Crocidura tanakae NOT ILLUSTRATED
Crocidura attenuata NOT ILLUSTRATED Measurements HB 65–85, T 45–60 (57–69% of
Measurements HB 65–90, T 40–60 (c.70% of HB), HF 12–13.8. Skull: cbl 17.9–20.6, ut 7.9–9.4
HB), HF 13–16. Skull: gl 19.0–22.5, ut 8.2–9.7 Identification Upperparts dark grey with brown
Identification Upperparts dark grey with brown tinge, underparts slightly paler and greyer. Similar
tinge, underparts slightly paler and greyer. Tail dark, to Asian Grey Shrew, C. attenuata, in external
nearly naked except for scattered long hairs (3–5mm) appearance and size. Differs genetically and in some
on basal third. Long hairs also on lower back and features of the skull: area between auditory bullae
limbs. Lingual (inside) edge of large upper premolar relatively flat and broad, compared with narrow and
rounded. Taxonomic notes Specimens formerly ridged in C. attenuata; inner lobe of premolar (talon)
referred to this species from Peninsular Malaysia relatively narrow and rounded, compared with broad
have been subsequently identified as C. negligens, and angular in C. attenuata. Northern populations
which does not apparently overlap in distribution tend to be smaller than southern populations.
with C. attenuata. Similar species South-east Asian Taxonomic notes Recently recognised as occurring
Shrew, C. fuliginosa, tends to have more angular together with C. attenuata in region.
inner edges to large upper premolar; Hill’s Shrew, Ecology and habitat Terrestrial. Grassland,
C. hilliana, has more robust teeth (Fig. 21b); Sokolov’s thickets, secondary forest, bamboo up to 1,800m
Shrew, C. sokolovi, has proportionately longer tail, altitude.
browner fur; Taiwanese Grey Shrew, C. tanakae, is Distribution SE Asia: Vietnam, Laos. Also China,
similar, but differs in skull characters (see below). Taiwan, the Philippines.
Ecology and habitat Terrestrial. Status Least Concern.

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 INDOCHINESE SHREW Distribution SE Asia: N Vietnam.
Crocidura indochinensis PLATE 3 Status Not yet assessed. Only one confirmed
Measurements HB 59–68, T 50–58 (c.85% of specimen.
HB), HF 12–13, E 10–11. Skull: cbl 16.9–17.8,
ut 7.3–8.0 WUCHIH SHREW
Identification Upperparts dark brownish-grey; Crocidura wuchihensis NOT ILLUSTRATED
underparts more greyish. Tail medium in length, Measurements HB 59–65, T 37–42, E 6–9.
nearly naked with thinly scattered short hairs and Skull: cbl 15.7–17.1, ut 6.8–7.3
only a few longer bristles near base. Relatively flat Identification Moderately small shrew. Upperparts
skull. Taxonomic notes Formerly considered part dark greyish-brown, individual hairs with grey base
of C. horsfieldi, which does not occur in region. and brown tip mixed with some longer silvery hairs;
Similar species Wuchih Shrew, C. wuchihensis, underparts slightly paler and greyer. Front and
is smaller; Voracious Shrew, C. vorax, and Chinese hind feet whitish, with pale brown hairs in middle,
Shrew, C. rapax, have distinctly bicoloured tail darker brown on sides. Tail nearly naked, very
covered with short fine hairs. thinly covered with short brown hairs, mixed with
Ecology and habitat Terrestrial. Primary and scattered long bristles on basal half; dark brown
secondary hill forest. above, paler below but not sharply demarcated.
Distribution SE Asia: N Myanmar, N Thailand, Laos Ears pale. Taxonomic notes Formerly confused
and Vietnam. with C. horsfieldi, which is considered to occur only
Status Least Concern, although precise distribution in India. Specimens from N Vietnam are slightly
uncertain as many historic records were confused larger than those from C Vietnam. Similar species
with other species. Indochinese Shrew, C. indochinensis, is larger; Ke
Go Shrew, C. kegoensis, is slightly smaller, with
KE GO SHREW dark mark on muzzle, dark ears, different teeth
Crocidura kegoensis PLATE 3 (Fig. 22).
Measurements HB 48, T 27 (c.55% of HB), HF 10, Ecology and habitat Recorded from montane
E 5. Skull: cbl 14.9, ut 6.5 evergreen forest at 1,300–1,500m altitude.
Identification Very small shrew. Upperparts dark Distribution SE Asia: N and C Vietnam. Also
greyish-brown, individual hairs with grey base and Hainan, China.
brown tip; underparts slightly paler and greyer, Status Data Deficient.
blending on sides with upperpart colours. Dark mark
on muzzle with dark ears. Front and hind feet with a
pale brown skin. Tail brown above, only slightly paler
below, covered with very short brown hairs, mixed
with scattered long bristles on basal half. Ears pale.
Large upper premolar with deeply curved indentation
on posterior edge, molars with deep notches on
inner side (Fig. 22a); anterior cusp of large upper
premolar same height as adjacent unicuspid. Similar b
species Wuchih Shrew, C. wuchihensis, has pale
ears, white skin on top of feet, lacks dark mark on
side of muzzle, shallower notch on large premolar
and molars (Fig. 22b); Zaitsev’s Shrew, C. zaitsevi,
has proportionately longer tail, lacks dark marks on
muzzle; other Crocidura in region are larger; Pygmy
White-toothed Shrew, Suncus etruscus, is smaller,
with an extra unicuspid tooth.
Ecology and habitat Only known record is from Fig. 22 Ventral views of skull of Crocidura kegoensis (a)
lowland rainforest at 200m altitude. and C. wuchihensis (b).

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 HILL’S SHREW smaller on average, with tail indistinctly bicoloured.
Crocidura hilliana NOT ILLUSTRATED Ecology and habitat Terrestrial.
Measurements T 45. Skull: cbl 21.0–23.5, Distribution SE Asia: W, C and NE Thailand, Laos,
ut 8.8–10.2 N. Vietnam. Also India, and S and C China.
Identification Originally described from skull Status Least Concern, though poorly known in
remains, with information on external appearance region and some uncertainty about number of
based on only one preserved specimen, described species.
later. Medium-sized, uniformly dark grey-brown
with slightly paler hands, feet, muzzle and tail. CHINESE SHREW
Tail with scattered long bristle hairs along 85% Crocidura rapax NOT ILLUSTRATED
of its length. Skull moderately large. Upper incisor Measurements HB 55–70, T 38–47 (c.66% of
large, robust, with well-developed posterior cusp, HB), HF 11–13. Skull: cbl 17.5–18.3, ut 7.1–8.1
first upper unicuspid large, nearly as high as Identification Small to medium-small; upperparts
incisor, about twice height and more than twice pale greyish-brown, fur with narrow grey base
crown area of remaining unicuspids (Fig. 21b). and pale brownish tip. Tail indistinctly bicoloured,
Similar species South-east Asian White-toothed dark above, slightly paler below, covered with
Shrew, C. fuliginosa, averages slightly larger, first short, fine hairs with scattered longer bristles on
upper unicuspid relatively small, about half height basal two-thirds. Similar species Voracious Shrew,
of incisor and only slightly higher than remaining C. vorax, is usually slightly larger, with a more
unicuspids; Grey Shrew, C. attenuata, is smaller on distinctly bicoloured tail and pronounced sagittal
average, with slender upper incisor, relatively small crest on the skull; both South-east Asian Shrew,
second and third upper unicuspids (Fig. 21). Other C. fuliginosa, and Grey Shrew, C. attenuata, are
Crocidura in its range are smaller. larger, with darker, greyer fur, and a tail that is only
Ecology and habitat Lowlands in a variety of sparsely haired except for scattered longer bristles
habitats from forest to cultivated lands, including near base; Hill’s Shrew, C. hilliana, is larger.
around limestone karst. Ecology and habitat Terrestrial.
Distribution SE Asia: E Thailand and C Laos. Distribution SE Asia: N Myanmar, Vietnam. Also
Status Data Deficient; likely to be more widespread India and S China.
than currently known. Status Data Deficient; fairly widely distributed
elsewhere and probably Least Concern.
VORACIOUS SHREW
Crocidura vorax PLATE 3 PHAN LUONG’S SHREW
Measurements HB 55–90, T 41–51 (c.70% of Crocidura phanluongi NOT ILLUSTRATED
HB), HF 11–14. Skull: cbl 17.7–20.1, ut 7.2–8.2 Measurements HB 54–66, T 40–48 (61–83% of
Identification Medium-sized; upperparts pale HB), HF 10–12. Skull: cbl 16.8–18.1, ut 7.3–8.2
greyish-brown, fur with dark grey base, then a Identification Small shrew with moderately long
narrow grey band and pale brownish tip, appearing tail. Fur dark grey to grey-brown, slightly paler
speckled. Tail sharply bicoloured, dark above, pale below; tail similar in colour to back, sometimes
below, covered with short, fine hairs that hide markedly paler below, with sparse bristles on
scales, with scattered longer bristles on basal two- proximal two-thirds; feet pale. Posterior lower
thirds. Skull has low but clearly defined sagittal molar (M3) with reduced cusps on posterior lobe
crest. Taxonomic notes Recent research suggests (talonid). Taxonomic notes Newly described
the species with this description in SE Asia may not in 2010. Similar species Indochinese Shrew,
be the same species as C. vorax from China, but C. indochinensis, is similar in size, but averages
appropriate names have not yet been determined. proportionately longer tail; broader interorbital
Similar species C. fuliginosa and C. attenuata region on skull; more complex cusps on lower third
are larger, with darker, greyer fur and a dark tail molar, including larger talonid.
only sparsely haired with scattered longer bristles Ecology and habitat Terrestrial. Has been found
restricted to near base; Chinese Shrew, C. rapax, is in a variety of habitats, including dry dipterocarp,

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 mixed evergreen, bamboo and wetlands, generally PHU QUOC SHREW
in lowlands up to c.450m altitude. Crocidura phuquocensis NOT ILLUSTRATED
Distribution SE Asia: S Vietnam and E Cambodia. Measurements HB 68–72, T 49–59 (69–87% of
Status Least Concern. HB), HF 12–12.5. Skull: cbl 17.6–18.2, ut 7.7–8.0
Identification Medium-sized shrew with moderately
ZAITSEV’S SHREW long tail; fur above and below short, dark grey
Crocidura zaitsevi NOT ILLUSTRATED with slight brown hue; tail similar colour to back,
Measurements HB 48–58, T 33–41 (62–81% of slightly paler below, with sparse bristles on proximal
HB), HF 8–11. Skull: cbl 14.2–15.3, ut 6.2–6.8 60%. Upper incisor slender, posterior cusp around
Identification Very small shrew, with moderately half height of first unicuspid. Taxonomic notes
long tail. Fur grey with a slight brownish tinge, slightly Newly described in 2008. Similar species Only
paler on underside; tail similar colour to body, with species known from Phu Quoc island; Grey Shrew,
long bristles on proximal two-thirds; upperside of feet C. attenuata, is similar in size, but averages slightly
paler than body. Posterior cusp of upper incisor very smaller, especially skull, with more slender rostrum.
large; anterior cusp of large upper premolar much Ecology and habitat Terrestrial. Caught in
shorter than adjacent unicuspid. Taxonomic notes undisturbed dipterocarp forest near sea level.
Newly described in 2007. Similar species Ke Go Distribution SE Asia: Known only from Phu Quoc
Shrew, C. kegoensis, has proportionately shorter tail, island off S Vietnam.
browner fur and dark marks on sides of face; Wuchih Status Data Deficient.
Shrew, C. wuchihensis, averages larger, with broader
rostrum and palate. CRANBROOK’S SHREW
Ecology and habitat Terrestrial. Has been found Crocidura cranbrooki NOT ILLUSTRATED
in tall montane broadleaf forest at altitudes of Measurements HB 65–86, T 65–88 (80–110% of
1,650–2,300m. HB), HF 14–16. Skull: cbl 19.3–20.6, ut 8.6–9.8
Distribution SE Asia: Known only from Ngoc Linh Identification Upperparts dark brown, individual
Mountain, C Vietnam. hairs with grey base and brown tip; underside
Status Data Deficient. somewhat paler, hairs with grey base and buff tip.
Tail long, thin, pale with darker patches; bristle
SOKOLOV’S SHREW hairs on proximal 30%, scale hairs pale, short and
Crocidura sokolovi NOT ILLUSTRATED thin so scales are visible. Taxonomic notes Newly
Measurements HB 70–78, T 65–68 (87–93% of described in 2009. Similar species South-east
HB), HF 14. Skull: cbl 18.8–20.4, ut 8.3–9.2 Asian Shrew, C. fuliginosa, is slightly larger and
Identification Medium-sized shrew with relatively has a thicker tail with dark scale hairs; Asian Grey
long tail; fur long and soft, brown with a grey Shrew, C. attenuata, is greyer with more long hairs
hue, similar above and below; tail brown above, on tail and dark scale hairs.
slightly paler below, enlarged at base with long Ecology and habitat Terrestrial. River valleys and
bristles on proximal third; skull with relatively deep hills at altitudes of 900–1,250m.
rounded braincase in profile, upper incisor relatively Distribution SE Asia: Known only from four localities
slender. Taxonomic notes Newly described in in N Myanmar.
2007. Similar species Grey Shrew, C. attenuata, Status Data Deficient.
is similar but with proportionately shorter tail;
shorter, greyer fur; shallower braincase; larger ANNAMITE SHREW
upper incisor. Crocidura annamitensis NOT ILLUSTRATED
Ecology and habitat Terrestrial. Montane broadleaf Measurements HB 49–58, T 30–33 (c.60% of
evergreen forest and elfin forest at altitudes of HB), HF 9–10. Skull: cbl 14.5–14.8, ut 6.4–6.6
2,300–2,400m. Identification Very small shrew with relatively short
Distribution SE Asia: Known only from Ngoc Linh tail. Upperparts pale brown, individual hairs pale
Mountain, C Vietnam. grey at base, pale brown at tip; underside hairs
Status Data Deficient. pale greyish brown at base, buff at tip, not sharply

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 demarcated from back. Tail dark brown above, pale Ecology and habitat Terrestrial. Found in a variety
brown below. Posterior lower molar has relatively of habitats including mixed evergreen forest, tall
reduced talon (posterior lobe). Taxonomic notes forest, open grassy glades and near streams at
Newly described in 2009. Similar species Other altitudes of 1,400–2,200m.
similar-sized Crocidura shrews in same area have a Distribution SE Asia: N Vietnam.
longer tail or different colour fur. Status Not yet assessed.
Ecology and habitat Terrestrial. Recorded from hill
forest at 920–1,240m. GUY’S SHREW
Distribution SE Asia: Known only from Ha Tinh in Crocidura guy NOT ILLUSTRATED
the Annamite Mountains of C Vietnam. Measurements HB 47–53, T 34–38 (69–77% of
Status Not yet assessed. HB), HF 9–10. Skull: cbl 14.7–14.8, ut 6.4–6.7
Identification Very small with moderately long tail;
SA PA SHREW brownish-grey above, individual hairs with grey
Crocidura sapaensis NOT ILLUSTRATED base and brown tip; silver-grey below, individual
Measurements HB 50–65, T 37–47 (62–84% of hairs with dark grey base and pale tip; no clear
HB). Skull: cbl 15.0–16.5, ut 6.5–7.2 demarcation. Posterior lower molar with somewhat
Identification Small with moderately long tail. Fur reduced talonid. Taxonomic notes Newly described
dark greyish-brown, similar above and below; tail in 2009. Similar species Zaitsev’s Shrew,
dark above, slightly paler below. Skull with relatively C. zaitsevi, is greyer above with darker underparts;
short, rounded rostrum; rounded deep braincase; Kego Shrew, C. kegoensis, and Annamite Shrew,
upper incisor slender, with a relatively small posterior C. annamitensis, are browner above, with darker
cusp less than half height of first unicuspid; lower underparts and relatively shorter tail.
posterior molar has well developed talonid (posterior Ecology and habitat Known only from an area
part of molar), with wide talonid basin. Taxonomic of mixed forest and clearings at an altitude of
notes Newly described in 2013. Similar species 300–700m.
Wuchih Shrew, C. wuchihensis, is similar, but has Distribution SE Asia: only known from Na Hang
a shallower braincase, posterior lower molar with a Nature Reserve, Vietnam.
smaller less developed talonid, with a narrow basin. Status Not yet assessed.

Order SCANDENTIA
Treeshrews
Superficially, treeshrews resemble squirrels, but they differ in many details of anatomy and behaviour. They
have a very long muzzle with a total of 38 teeth, with pointed incisors and several premolars (Fig. 23), unlike
the chisel-like incisors of rodents. All feet have five well-developed digits with claws, which tend to show
well on footprints (Fig. 15b). Treeshrews have been variously classified as primates or insectivores, but they
are now recognised as distinct from both and are placed in their own order, Scandentia.

Family PTILOCERCIDAE FEATHER-TAILED TREESHREWS

Although often placed as a subfamily within the Tupaiidae, this species differs in many morphological and
genetic ways from other treeshrews, and is now considered to be in its own family.

FEATHER-TAILED TREESHREW with a grey base and buffy-white tip. Eyes and
Ptilocercus lowii PLATE 4 ears relatively large. Tail long and naked except
Measurements HB 134–150, T 160–202, for a feather-like tip of long hairs sticking out on
HF 24–30 either side, starting about halfway along the tail,
Identification Upperparts fluffy grey to grey-brown; the first few long hairs black, the remainder white.
underparts pale brownish-grey, the individual hairs Similar species No other mammal in the region

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 has similar feathery tail with naked base. Distribution SE Asia: peninsular Thailand and
Ecology and habitat Nocturnal. Usually arboreal, Malaysia. Also Sumatra, Borneo and some offshore
often travelling on vines. Lowland forest from sea islands.
level to about 1,000m altitude. Also reported from Status Least Concern. Relatively few records, likely
disturbed habitats, including secondary forest and to have declined due to loss of lowland rainforest
plantation. Sometimes travels on the ground. Diet habitat.
mainly insects and other arthropods.

Family TUPAIIDAE TREESHREWS

The Slender-tailed Treeshrew, Dendrogale, is readily distinguished from other species by its narrow tail and
distinctive face pattern. Within the genus Tupaia, three species occur in the region, of which the larger two
are very similar to each other and are distinguished mainly by colour and range.
The Shrew-faced Ground Squirrel, Rhinosciurus laticaudatus (page 361), sometimes resembles a
treeshrew at a distance, but it can be distinguished by its shorter tail, pale underparts, lack of a pale mark
on the neck, the modified digits on the front feet and the very different teeth.

COMMON TREESHREW geographic variation, generally olive-brown in much

Tupaia glis PLATE 4 of range, but in some northern populations there are
Measurements HB 135–205, T 125–195 (80– also reddish forms similar to T. glis. Further studies,
105% of HB), HF 42–49. Wt 90–190g including use of genetics, are needed to determine
Identification Hairs on upperparts banded dark whether additional species are represented.
and pale, appearing finely speckled, with a strong Similar species Common Treeshrew, T. glis, is
reddish tint; underparts buff. Usually has a pale more reddish in areas where range overlaps in
stripe on each shoulder. Female has two pairs of peninsular Thailand, and female has only two pairs
mammae. Taxonomic notes The forms in Sumatra, of mammae.
Java and Borneo are now considered distinct Ecology and habitat Diurnal. Mainly terrestrial and
species. Similar species Northern Treeshrew, in low bushes. Found in a range of forest habitats
T. belangeri, lacks reddish tinge, and has three including evergreen forest, as well as deciduous
pairs of mammae in female. forest, especially near streams and rivers. Tolerant
Ecology and habitat Diurnal. Most often observed of moderate forest disturbance, and found in
active around fallen trees and branches, in low gardens and plantations.
woody vegetation or on the ground. Diet mainly Distribution SE Asia: Myanmar, Thailand, Laos,
insects and other arthropods, and sweet or oily Vietnam and Cambodia. Also Nepal, Bangladesh,
fruits. Occurs in forests as well as in gardens and India and S China.
plantations near forest. Status Least Concern.
Distribution SE Asia: peninsular Thailand and
Malaysia. Also Sumatra, Java and smaller
Indonesian islands.
Status Least Concern, although declining due to
loss of lowland forest.

NORTHERN TREESHREW
Tupaia belangeri PLATE 4
Measurements HB 160–230, T 150–200 (c.100%
of HB), HF 38–45, E 16–17
Identification Hairs on upperparts banded dark and
pale, appearing finely speckled brown. Underparts
0 50
buff. Usually has a pale stripe on each shoulder.
Female has three pairs of mammae. Moderate Fig. 23 Lateral view of skull of Tupaia sp.

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 LESSER TREESHREW NORTHERN SLENDER-TAILED TREESHREW
Tupaia minor PLATE 4 Dendrogale murina PLATE 4
Measurements HB 110–140, T 140–160 (115– Measurements HB 115–135, T 105–130,
130% of HB), E 10–14, HF 25–29. Wt 30–70g HF 26–30, E 10–18. Wt 35–55g
Identification Hairs on upperparts banded light Identification Upperparts brownish-black, speckled
and dark, giving an overall speckled olive-brown with orange-brown. Underparts sharply contrasting
appearance. Underparts buffy, often with a reddish pale buff to pale orange. Muzzle pointed, with buffy-
tinge towards the rear. Upper side of tail darker than orange stripe below and above eye, contrasting with
body. Tail long and relatively thin. Similar species dark mask through eye and darker crown. Tail thickly
Other Tupaia treeshrews in region are larger, with covered with short dark hair. Similar species Lesser
bushier tail; Northern Slender-tailed Treeshrew, Treeshrew, Tupaia minor, has longer, slightly bushier
Dendrogale murina, has relatively shorter, thinner tail, lacks stripes on face, does not overlap in range;
tail, and stripes on face; small tree squirrels have striped squirrels, Tamiops spp., have more rounded
less pointed muzzle. head, different face pattern and paler striped back.
Ecology and habitat Diurnal and mainly arboreal. Ecology and habitat Diurnal. Primarily scansorial
Often seen at 3–8m above ground, sometimes to (climbing), found in trees and bushes near ground,
20m, travelling along lianas or branches of small but may sometimes be in canopy. Most records
trees. Diet includes insects and fruits. Occurs in from evergreen forest, but also recorded from
forests, gardens and plantations. disturbed forest and areas of extensive bamboo.
Distribution SE Asia: peninsular Thailand and Distribution SE Asia: SE Thailand, SE Laos, Vietnam
Malaysia. Also Sumatra and Borneo. and Cambodia. Appears to be patchily distributed
Status Least Concern, although declining due to and relatively poorly known.
loss of lowland forest. Status Least Concern, but declining due to loss
of forest.

Order DERMOPTERA
Family CYNOCEPHALIDAE COLUGOS
The order Dermoptera includes only one living family, with only two known species, one confined to the
southern Philippines and the other found elsewhere in South-east Asia. Colugos, also known as flying
lemurs, cannot truly fly, but instead glide with the aid of a membrane between their legs. The teeth are
highly specialised and unlike those of any other mammal: there are no teeth at the front of the upper jaw,
and the two pairs of lower front teeth look like fine-toothed combs (Fig. 24).

SUNDA COLUGO
Galeopterus variegatus PLATE 29
Measurements HB 330–420, T 180–270,
HF 65–73. Wt 925–1,700g
Identification Usually grey, with extensive black-
and-white markings, but some individuals are tinged
reddish-brown, or are totally reddish-brown with
light buff or orange markings. Most distinctive
feature is the gliding membrane, which extends
between front and hind legs and encloses tail. All
0 50
four legs relatively long with long, sharp claws on
toes which are connected by webs. Small young may Fig. 24 Skull of Sunda Colugo, Galeopterus
be carried with adult, sometimes wrapped in gliding variegatus, with enlarged detail of lower incisor
membrane. Taxonomic notes Formerly included showing comb-like cusps.

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 in genus Cynocephalus. Similar species Flying buds and sap. Occurs in tall and secondary forests,
squirrels have a long bushy tail completely free of gardens and tree plantations.
any enclosing membrane, and much shorter legs. Distribution SE Asia: S Myanmar, S Thailand,
Ecology and habitat Nocturnal, but sometimes Laos, Vietnam, Cambodia and Peninsular Malaysia.
active in the morning and late afternoon. Clings to Also Sumatra, Java, Borneo, Bali and many smaller
the sides of tree trunks or glides between tall trees. Indonesian islands.
May also hang under horizontal branches. Rests in Status Least Concern, although populations
tree holes or crowns of trees. Diet poorly known, declining due to habitat loss and hunting.
but believed to feed on leaves, leaf shoots, flower

Order CHIROPTERA
Bats
Bats are the only mammals capable of true flight. Two other groups of South-east Asian mammals, the
colugos and the flying squirrels, are often said to fly, but they can only glide, and are not capable of powered
flight. They have gliding membranes between their legs, allowing them to travel quite long distances
between tall trees, but they cannot flap them to gain lift. Bats, in contrast, have their forelimbs modified into
wings, with membranes stretched between elongated finger bones (Fig. 25), and powerful chest muscles
to flap their wings.
Bats flying at dusk could potentially be confused with birds. Flying-foxes are similar in size to some
hawks and eagles, while small insectivorous bats are similar in size to small swifts that are often flying
in similar habitats. However, they can be distinguished by the shape of their wings and body, and by their
pattern of flight.
With the exception of a few distinctive species, most bats are difficult to identify visually unless they are
captured and examined closely. In the hand, most species, or at least groups of species, can be distinguished
based on easily visible characteristics and measurements. Fur colour and wing patterns are useful for
identifying some species, though in other groups colour is highly variable and not reliable for identification.
For some species it may be necessary to examine the skull or dentition for confirmation. With care, the teeth
can be examined in a live bat by gently opening the mouth with a toothpick or similar tool. A magnifying
glass can be helpful to see details. The skull itself can only be examined in a museum specimen, but many
features of skull shape are reflected in the outward appearance of the head and, with practice, most species
can be identified in life.
For many of the insectivorous bats, echolocation calls can be another method of identification. Nearly
all bat calls are above the hearing range of humans, so a bat detector is needed to record the calls. Full-
spectrum bat detectors allow the calls to be recorded and then viewed and analysed on a computer to
measure various aspects of their call structure. Reliable identification of bat species by call depends upon
having a comprehensive call library of species found in a particular area, especially because the calls of many
species vary geographically. The frequency and structure of many calls also depends on the behaviour of the
bats. For example, the same species flying in a cluttered environment, such as among trees, or in an open
environment, such as above the canopy, may use different calls. A comprehensive description of calls of
most species is beyond the scope of this book. However, the calls of the horseshoe and roundleaf bats have
a characteristic constant-frequency component that can even be measured in the hand. This component is
often useful for distinguishing very similar-looking species. When information on the calls of these species is
available, it is provided in the species accounts. These must be considered cautiously outside the region where
they were recorded, as substantial geographic variation has been observed in some species.
Juvenile bats are capable of flight before they are fully grown. They tend to be smaller and coloured
differently from adults. Young are generally much duller in colour, grey or grey-brown. If held against a bright
light, the wing joints of immatures appear banded where the cartilage has not yet turned to bone (the adult joint

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 Thumb 1st digit
Ear Forearm Metacarpals

2nd digit
First phalanges

Interfemoral Terminal phalanges

membrane

3rd digit
4th digit
5th digit
Wing membrane

Calcar
Tail Hind foot

Fig. 25 Diagram of a bat showing the naming of finger bones and membranes.

looks like a solid lump). Measurements of juvenile bats are often smaller than those of adults. By the time the
joints are fully ossified and solid, bats are full size, although they may take a bit longer to gain full adult colours.
Bat research has a long history in South-east Asia, but nevertheless much remains to be learned. Many
of the more effective survey techniques, such as harp traps, mist nets and bat detectors, have come into use
only recently (see Figs 9 and 10). Relatively few areas have been subject to intensive surveys, and most new
surveys turn up new distributional records or even previously unknown species. Furthermore, genetic studies
are revealing that many taxa formerly considered to be one species actually represent several species. Much
research remains to be done to resolve these species complexes, not only to determine reliable ways to
distinguish the species based on morphology, but also to determine the appropriate name for each species.
In the 10 years since the first edition of this book was published, at least 34 additional species have been
formally described or recognised as distinct from the region, although three species formerly thought to
occur in the region are no longer considered to be valid species (Myotis oreias, Hypsugo anthonyi and
Paracoelops megalotis).
The species descriptions in this book thus represent the current state of knowledge of bats in the region,
as published at the time of writing, but further changes are anticipated. If there are known taxonomic issues
for a species, these are mentioned in the accounts.
South-east Asian bats are classified into nine families, which can be distinguished by ear shape, muzzle
shape, the presence or absence of a noseleaf and the tail pattern. Within each family, genera can be
distinguished by a variety of characters, such as noseleaf shape, dentition or even colour. In many cases,
identification can be greatly facilitated by first determining the genus characters. To assist with sorting out
the very large number of bats in the region, general accounts of each family and genus are given in addition
to the species accounts.

Family PTEROPODIDAE FRUIT BATS

Fruit bats have large, conspicuous eyes which reflect a dull red eyeshine when a light is shone on them
at night. The ears are relatively small and simple, with a complete unbroken margin. The muzzle appears
rather dog-like, without modifications or leaflets, although the nostrils are well developed and sometimes
slightly tubular. The tail is short or lacking, and the interfemoral membrane is narrow. The second finger is
well developed, independent of the third finger, and usually with a claw at the tip (except Eonycteris). The
teeth have simple cusps, quite unlike the sharp W-shaped cusps of other bats.

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 Many of these features are reflections of their behaviour. Fruit bats lack echolocation (except Rousettus),
and rely instead on eyesight and smell to find their way. Thus, they need large reflective eyes to see well at
night. They feed mainly on nectar and fruits, and often climb through trees using the claws on their wings as
well as their feet to grip branches.
Twelve genera of fruit bats are currently recognised from mainland South-east Asia. For identification,
these can be divided into groups based on morphology and feeding behaviour. The largest group includes
Cynopterus, Dyacopterus, Megaerops, Chironax, Sphaerius, Balionycteris, Penthetor and Aethalops, which
have short, relatively powerful jaws with reduced numbers of large, heavily cusped teeth for feeding on fruit
(Fig. 26b, c, d). The macroglossine bats, which include Macroglossus and Eonycteris, have a long, narrow
muzzle, weak jaws, small teeth and a relatively long tongue for nectar feeding (Fig. 26f, g). Rousettus
(Fig. 26e) and Pteropus (Fig. 26a) are somewhat intermediate in shape, with an elongate muzzle and
moderately well-developed teeth. They may be closer to the ancestral form from which the others evolved.
Dental characters are important for distinguishing genera but must be examined carefully, as some teeth,
particularly the smaller cheek teeth, are sometimes missing or shed in older individuals. This may be due
to deterioration of the teeth associated with the high sugar content of their diets. Thus, it is important to
consider the shapes and sizes of teeth and not just the number.

Genus Rousettus Moderately large bats with a

a
fairly long muzzle, a claw on the second digit and
a well-developed tail. Produce a distinctive clicking
call with the tongue, which is used for echolocation,
presumably mainly to navigate in dark caves. Skull
moderately long with slightly decurved rostrum
(Fig. 26e). Cheek teeth unspecialised with poorly b e
developed cusps. Dental formula 2/2 1/1 3/3 2/3.

GEOFFROY’S ROUSETTE
Rousettus amplexicaudatus PLATE 7 c f
Measurements FA 78–87, T 15–21, E 16.5–19.5.
Wt 55–80g; males average slightly larger than
females d g
Identification Upperparts grey-brown to brown,
darker on top of head; underparts paler grey-
brown. Fur short and sparse except for long pale 0 20
hairs on chin and neck. Adults, especially males,
sometimes have pale yellow tufts of hair on the Fig. 26 Profiles of selected fruit bat skulls:
sides of the neck. Third lower molar nearly circular, Pteropus vampyrus (a), Dyacopterus spadiceus (b),
only slightly longer than wide (Fig. 27a). Similar Cynopterus brachyotis (c), Aethalops alecto (d),
species Leschenault’s Rousette, R. leschenaultii, Rousettus amplexicaudatus (e), Eonycteris spelaea (f)
has a more elongate third lower molar and smaller and Macroglossus minimus (g).
incisors; nectar bats, Eonycteris spp., lack a claw
on the second digit; Dayak Fruit Bat, Dyacopterus attached to the sides of the back, separated by a
spadiceus, is similar in size, but has a much thicker, broad band of fur.
shorter muzzle and fewer, larger cheek teeth (Fig. Ecology and habitat Roosts in caves, sometimes in
28d). Bare-backed Rousette, R. spinalatus, known complete darkness, often in association with Cave
from Borneo and Sumatra, might be expected in Nectar Bat, Eonycteris spelaea. Feeds on fruit as
Peninsular Malaysia; it is distinguished by having well as nectar and pollen.
the wings joined together over the middle of the Distribution SE Asia: S Myanmar, Thailand,
back, unlike other Rousettus, which have wings Vietnam, Laos, Cambodia and Peninsular Malaysia.

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 Also Borneo and most Indonesian islands through
New Guinea, Solomon Islands and the Philippines.
Status Least Concern, but has declined in some a
areas due to excessive hunting.

LESCHENAULT’S ROUSETTE
Rousettus leschenaultii PLATE 7
Measurements FA 75–85, T 10–18, E 18–24.
Wt 60–100g; males heavier than females b
Identification Upperparts grey-brown to buffy-
brown, paler underneath. Fur short and sparse
except for long pale hairs on chin and neck. Third
lower molar elongate, about twice as long as wide
(Fig. 27b). Similar species Geoffroy’s Rousette, Fig. 27 Right lower toothrows of Rousettus
R. amplexicaudatus, is similar in size, but averages amplexicaudatus (a) and R. leschenaulti (b).
slightly darker in colour with a slightly shorter
toothrow, and has a more rounded third lower
molar.
a
Ecology and habitat Like other Rousettus,
produces an audible clicking call when flying, which
is used for echolocation. Roosts in caves, including
wells, mines and artificial caves. Diet mainly flower
nectar, pollen and fruit.
Distribution SE Asia: Myanmar, Thailand, Laos, b
Vietnam, Cambodia and Peninsular Malaysia. Also
Pakistan, India, Nepal, S China, Sumatra and Java.
Status Least Concern.

Genus Pteropus Very large bats, considerably

larger than any other South-east Asian bats, c
including the largest bats in the world. First finger
very long; second finger has a well-developed claw.
Skull large and elongate with an almost tubular
braincase (Fig. 26a). Three upper premolars, but d
anterior very small and often missing in older
individuals. Remaining teeth relatively simple and
unspecialised. Dental formula 2/2 1/1 3/3 2/3.
Four or five species in the region, depending upon
whether P. intermedius is recognised as a distinct Fig. 28 Right lower toothrows of Cynopterus horsfieldii
species. (a, b), C. sphinx (c) and Dyacopterus spadiceus (d).

LARGE FLYING-FOX Sides of head and front of neck mixed reddish-

Pteropus vampyrus PLATE 5 brown and black, blending into blackish-brown
Measurements FA 185–200, T none. Wt 645– underparts. Immatures uniform dull grey-brown.
1,100g Similar species Indian Flying-fox, P. giganteus,
Identification Very large bat, with a wingspread tends to be slightly smaller, with a paler head and
up to 1.5m. Colour somewhat variable, but back pale brown underparts; Lyle’s Flying-fox, P. lylei,
usually black with grey streaking, sharply separated and Island Flying-fox, P. hypomelanus, are both
from orange-brown mantle and back of head. substantially smaller.

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 Ecology and habitat Roosts in large, established colour; muzzle dark, with dark pattern sometimes
colonies on open branches of trees, often in mangrove extending to whole front of head. Underparts vary
or nipah palm. Sometimes flies long distances to feed from buffy-brown to black. Similar species Large
in flowering or fruiting trees. May migrate seasonally Flying-fox, P. vampyrus, is much larger and nearly
to follow fruiting and flowering cycles. Eats both always has dark underparts; Island Flying-fox,
nectar and fruit, including some orchard species such P. hypomelanus, is smaller, with rounded ears.
as rambutans and mangoes. Pollinates the flowers of Ecology and habitat Roosts in large, established
many forest trees, including durian. colonies on open branches of large trees. Most
Distribution SE Asia: lowland areas of S Myanmar, known colonies are in temples or in the middle of
S Thailand, Cambodia, S Vietnam and Peninsular towns or cities. Feeds on fruits and may raid fruit
Malaysia. Possibly Laos. Also Sumatra, Borneo, orchards.
Java, Lesser Sundas and the Philippines. Distribution SE Asia: known only from S Thailand,
Status Near Threatened. Populations in many areas Cambodia and Vietnam.
declining due to over-hunting and loss of habitat. Status Vulnerable: limited range and population
with evidence of substantial recent declines,
INDIAN FLYING-FOX including loss of some colonies, due to excessive
Pteropus giganteus PLATE 5 hunting and persecution.
Measurements FA 140–180, T none
Identification Very large bat. Back black, lightly ISLAND FLYING-FOX
streaked grey; mantle pale yellow-brown; head Pteropus hypomelanus PLATE 5
brown; underparts buffy-brown. Some individuals Measurements FA 120–145, T none, Wt 210g
are darker. Taxonomic notes Some authorities Identification Smallest flying-fox in the region (but
recognise Intermediate Flying-fox, P. intermedius, still much larger than other fruit bats). Lower back
from central Myanmar as a distinct species; it dark blackish-brown, sometimes heavily frosted
resembles P. giganteus, but has browner underparts. with pale tips such that back appears grey; upper
Further analyses are required to determine whether back, neck and back of head pale golden-brown
it is, in fact, a distinct form. Similar species Lyle’s to dark reddish-brown; underside varies from dark
Flying-fox, P. lylei, can be similar in colour, but blackish-brown to buffy-brown. Ears relatively broad
averages smaller and does not overlap in range; and rounded. Similar species Lyle’s Flying-fox,
Large Flying-fox, P. vampyrus, usually has darker P. lylei, is only slightly larger, but its ears are more
underparts, different distribution. pointed.
Ecology and habitat Roosts in large, established Ecology and habitat Roosts mainly on islands,
colonies on open branches of trees, including in in the fronds of coconut palms or amongst the
grounds of temples and in urban areas. May fly branches of trees. Sometimes flies to the mainland
many kilometres in search of food. Eats both nectar to feed.
and fruit, including some orchard species. Pollinates Distribution SE Asia: small islands off coastal
the flowers of many forest trees. Myanmar, Thailand, Vietnam, Cambodia, Malaysia
Distribution SE Asia: known only from W Myanmar. and sometimes adjacent mainland. Also Andamans,
Also Pakistan, Nepal, India and Sri Lanka. Indonesia, the Philippines, New Guinea and the
Status Least Concern, but probably declining due Solomons.
to hunting; need to protect roosts from disturbance. Status Least Concern, but subject to hunting in
some areas, and has probably declined somewhat.
LYLE’S FLYING-FOX
Pteropus lylei PLATE 5 Genus Cynopterus Medium to large bats with short,
Measurements FA 145–160, T none. Wt 390– stout muzzle. Fur is typically yellowish-brown with a
480g reddish or orange collar, best developed in males.
Identification Medium-large flying-fox with back Wing bones and rims to ears are usually contrastingly
and wings dark brown or black; mantle and head white. Most individuals have brown fur with a
buffy to orange-brown, sharply separated from back yellowish or reddish tinge and contrasting whitish

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 wing bones and rims to the ears, although some Status Least Concern, though apparently much less
old individuals have darker rims to the ears, and common than Sunda species.
young bats can be very grey. Skull compact with a
short rostrum (Fig. 26c). Anterior upper premolar SUNDA SHORT-NOSED FRUIT BAT
very small. Upper canines have a slight posterior Cynopterus cf. brachyotis ‘Sunda’ PLATE 6
secondary cusp; cheek teeth fairly stout, generally Measurements FA 59–70, T 8–10, E 14–17.
unspecialised. Dental formula 2/2 1/1 3/3 1/2. Wt 32–42g. Skull: gl 27.2–29.6
Identification Generally brown to yellowish-
FOREST SHORT-NOSED FRUIT BAT brown with an orange collar in adult males, more
Cynopterus cf. brachyotis ‘Forest’ PLATE 6 yellowish collar in females. Ears and wing bones
Measurements FA 56–63, T 9–10, E 13–16. edged in white. Two pairs of lower incisors (Fig.
Wt 24–35g. Skull: gl 26.4–28.8 29a). Taxonomic notes Genetic analyses indicate
Identification Generally brown to yellowish-brown it is distinct from the Forest Short-nosed Fruit
with a brighter collar, dark orange-brown in adult Bat, C. cf. brachyotis ‘Forest’, which co-occurs in
males, more yellowish in females. Immatures greyer Peninsular Malaysia and Borneo. Similar species
with indistinct collar. Ears and wing bones edged in Forest Short-nosed Fruit Bat, C. cf. brachyotis
white. Muzzle relatively short. Two pairs of lower ‘Forest’, averages smaller with a darker orange
incisors (Fig. 29a). Taxonomic notes Genetic collar and shorter muzzle; Greater Short-nosed
analyses indicate that bats formerly referred to as Fruit Bat, C. sphinx, has longer forearm, longer
C. brachyotis belong to several different species, ears and longer skull; Horsfield’s Fruit Bat,
of which at least two species co-occur in mainland C. horsfieldii, is larger and has more rectangular,
South-east Asia. These are here called ‘Forest’ heavily cusped molars (Fig. 28a, b); Dusky Fruit
and ‘Sunda’ C. cf. brachyotis, as it is not yet clear Bat, Penthetor lucasi, has only one pair of lower
which one is actually C. brachyotis. A specimen incisors (Fig. 29b), lacks white edges to the ears
from Myanmar was genetically distinct and may and is greyer; Blanford’s Fruit Bat, Sphaerius
represent a third species in the region, but further blandfordi, has pale rims to the ears and pale wing
study is required to determine its distinguishing bones, but differs in colour, lacks a tail and has a
characteristics. Similar species Sunda Short- thickly furred interfemoral membrane.
nosed Fruit Bat, C. cf. brachyotis ‘Sunda’, is difficult Ecology and habitat Occurs in a wide variety of
to distinguish, but has longer forearm (on average), disturbed habitats, including orchards, plantations
proportionately longer muzzle, and more yellowish- and second growth, where it can be very common.
orange collar. Not usually found in the interior of mature forests.
Ecology and habitat Largely restricted to more Roosts singly or in small groups, usually under the
mature forests, from lowlands to hills, where it leaves of trees, including coconut palms. Feeds on
has been found in both the forest understorey and small fruits, sucking out the juices and soft pulp, as
the canopy. Roosts under large leaves of trees, well as bananas, nectar and pollen.
especially palms or large-leaved trees such as Distribution SE Asia: S Myanmar, S Thailand,
Macaranga. Sometimes modifies leaves into a S Laos, Cambodia, S Vietnam and Peninsular
‘tent’ by biting the mid-rib. Feeds mainly on fruits, Malaysia. Also Java and Borneo.
especially figs. Status Least Concern.
Distribution SE Asia: Peninsular Malaysia. Also
Borneo. GREATER SHORT-NOSED FRUIT BAT
Cynopterus sphinx PLATE 6
a b Measurements FA 65–76, E 18–24. Wt. 40–57g.
Skull: gl 29–33
Identification Upperparts brown to grey-brown;
underparts paler. Collar orange-brown in males,
Fig. 29 Front views of lower canines and incisors of yellowish in females. Ears and wing bones edged
Cynopterus cf. brachyotis (a) and Penthetor lucasii (b). in white. Lower cheek teeth rounded without

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 accessory cusps (Fig. 28c). Similar species Short- incisors about half the length of the inner pair.
nosed fruit bats, Cynopterus spp., are smaller, with Dental formula 2/1 1/1 3/3 1/2.
shorter ears; Horsfield’s Fruit Bat, C. horsfieldii,
has squarer cheek teeth with extra cusps or ridges; DUSKY FRUIT BAT
Blanford’s Fruit Bat, Sphaerius blandfordi, has pale Penthetor lucasi PLATE 6
rims to ears, but differs in colour, lacks a tail and Measurements FA 57–62, T 8–13, E 14–16.5.
has a thickly furred interfemoral membrane. Wt 30–44g
Ecology and habitat Found in disturbed habitats Identification Upperparts dark grey-brown;
and open forests, where it is relatively common. underparts pale buffy-grey. Top of head often
Roosts in trees, under palm fronds, under the leaves distinctly darker down the centre and paler near the
of large epiphytic ferns and occasionally near the eyes. Ears have dark edges. Only one pair of lower
entrances of caves, in rock crevices or under roofs. incisors (Fig. 29b); outer upper incisors shorter than
Feeds on nectar and fruit. inner pair. Similar species Short-nosed fruit bats,
Distribution SE Asia: Myanmar, Laos, Vietnam, Cynopterus spp., are brighter-coloured with two
Cambodia, S Thailand to northern part of Peninsular pairs of lower incisors; tailless fruit bats, Megaerops
Malaysia. Also Sri Lanka, India through S China, spp., lack a tail, have broader, higher nostrils, light
Sumatra, Java, Timor and adjacent islands. brown fur with pale grey base, and small, even
Status Least Concern. upper incisors.
Ecology and habitat Roosts in colonies in rock
HORSFIELD’S FRUIT BAT shelters or caves, sometimes in near-total darkness.
Cynopterus horsfieldii PLATE 6 Eats fruit, which it sometimes carries back to the
Measurements FA 68–76, T 14, E 17–20. cave to eat.
Wt 50–70g. Skull: gl 31–34 Distribution SE Asia: Peninsular Malaysia and
Identification Upperparts grey-brown; underparts extreme S Thailand. Also Borneo, Sumatra and Riau
slightly yellowish-brown; collar dark reddish-brown archipelago.
in adult males, paler in females. Ears and wing Status Least Concern, although uncommon in most
bones edged in white. Cheek teeth broader and areas.
squarer than those of other Cynopterus, with
distinct cusps or ridges on the last lower premolar Genus Megaerops Small to medium-sized bats,
and first lower molar (Fig. 28a, b). Similar species similar to Cynopterus but with a slightly shorter
Greater Short-nosed Fruit Bat, C. sphinx, has nose; usually no visible tail; only one pair of lower
smaller, more rounded cheek teeth; Dayak Fruit Bat, incisors. Skull similar to Cynopterus, but with a
Dyacopterus spadiceus, is larger, lacks the small shorter, higher rostrum (Fig. 30). Canines lack
anterior upper premolar, has more massive teeth secondary cusps. Upper incisors small and roughly
(Fig. 28d) and differs in colour. equal, evenly spaced between the canines. Dental
Ecology and habitat Roosts in trees or under formula 2/1 1/1 3/3 1/2.
banana leaves, which may be modified into a tent
by biting the mid-rib. Also roosts in solution cavities SUNDA TAILLESS FRUIT BAT
in limestone cliffs or caves. Feeds mainly on fruit, Megaerops ecaudatus PLATE 5
tending to eat larger fruit than other Cynopterus. Measurements FA 51–58, T none. Wt 20–38g.
Distribution SE Asia: S Vietnam, Laos, Cambodia, Skull: cbl 24.0–26.2
Thailand and Peninsular Malaysia. Also Borneo, Identification Upperparts yellowish-brown
Sumatra, Java and adjacent islands. to reddish-brown with pale grey base to fur;
Status Least Concern. underparts paler and greyer. Ears with dark edges.
Muzzle short with broad and slightly tubular nostrils.
Genus Penthetor Medium-sized bats, very similar One pair of lower incisors. Interfemoral membrane
to Cynopterus, but coloration generally dark grey- narrow and thinly haired, with no visible tail. Skull
brown; cheek teeth slightly wider and flatter; only with short, high rostrum (Fig. 30b). Similar species
one pair of lower incisors (Fig. 29b); outer upper Northern Tailless Fruit Bat, M. niphanae, is very

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 similar, distinguished by more pointed muzzle lacking. Muzzle short and thick. One pair of lower
and smaller nostrils; White-collared Fruit Bat, incisors. Rostrum of skull less upturned than in
M. wetmorei, is smaller with more pointed muzzle Sunda Tailless Fruit Bat, M. ecaudatus (Fig. 30b, c).
and (in males) white shoulder tufts; short-nosed Taxonomic notes May be a distinct species
fruit bats, Cynopterus spp., have white or pale from that found in the Philippines, in which case
edges to ears, a short tail and two pairs of lower scientific name of Malaysian species would become
incisors; Dusky Fruit Bat, Penthetor lucasi, has a M. albicollis. Similar species Sunda Tailless Fruit
short tail and darker fur; Grey Fruit Bat, Aethalops Bat, M. ecaudatus, is larger, lacks white neck tufts
alecto, has a more pointed muzzle and a thickly and has a thicker muzzle; Black-capped Fruit Bat,
furred interfemoral membrane. Chironax melanocephalus, has two pairs of lower
Ecology and habitat Has been found mainly in tall incisors and dark head; Grey Fruit Bat, Aethalops
forest, but sometimes also in disturbed forest. alecto, has a more pointed muzzle and thickly furred
Distribution SE Asia: S Thailand and Peninsular interfemoral membrane.
Malaysia, overlapping range of M. niphanae in Ecology and habitat Mainly in mature lowland
Thailand; northern limits of range poorly known forest, where it is most common in canopy.
because of past confusion with M. niphanae. Also Distribution SE Asia: Peninsular Malaysia. Also
Sumatra, Borneo. Sumatra, Borneo and the Philippines.
Status Least Concern. Status Vulnerable, due to loss of lowland rainforest
in all parts of range.
NORTHERN TAILLESS FRUIT BAT
Megaerops niphanae PLATE 5
Measurements FA 53–61, T none. Wt 22–39g. a
Skull: cbl 24.4–26.8
Identification Upperparts brownish with pale grey
base to fur; underparts paler and greyer. Ears with
dark edges. Muzzle sloped in profile with slightly
tubular nostrils (Fig. 30a). One pair of lower incisors.
Interfemoral membrane narrow and thinly haired,
with no visible tail. Similar species Sunda Tailless
Fruit Bat, M. ecaudatus, is very similar, differing b
mainly in the shape of the head, with more upturned
rostrum (Fig. 30b) and protruding nostrils.
Ecology and habitat Found in a wide variety of
forests, including partially disturbed areas.
Distribution SE Asia: throughout much of Laos,
Vietnam, Cambodia and Thailand, except in south.
Most likely in Myanmar, though not yet reported.
Also NE India.
Status Least Concern.
c
WHITE-COLLARED FRUIT BAT
Megaerops wetmorei PLATE 5
Measurements FA 45–51, T 0–4, E 12.
Wt 14–18g. Skull: cbl 21.2–22.0
Identification Body fur pale grey-brown, lower
back slightly darker. Adult males have large white
tufts on sides of neck that extend onto back to form Fig. 30 Skulls of Megaerops niphanae (a),
a broken collar. Interfemoral membrane narrow M. ecaudatus (b) and M. wetmorei (c), showing
and sparsely covered with hairs. Tail very short or differences in rostrum shape.

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 Genus Dyacopterus Similar to Cynopterus, with a Distribution SE Asia: S Thailand and Peninsular
distinct tail, short muzzle and thick jaws (Fig. 26b), Malaysia. Also Sumatra, Java, Borneo and Sulawesi.
but cheek teeth greatly enlarged, with squared Status Least Concern, but declining due to loss
corners, large cusps and ridges (Fig. 28d). Only three of forest.
upper cheek teeth. Dental formula 2/2 1/1 2/3 1/2.
Genus Balionycteris A small, dark fruit bat, easily
DAYAK FRUIT BAT recognised by pale spots on wings. Skull similar
Dyacopterus spadiceus PLATE 7 to Cynopterus, but slightly more elongate with an
Measurements FA 77–81, T 19–24, E 17.5–21. extra, very small upper molar and only one pair of
Wt 75–100g lower incisors. Upper incisors close together and
Identification Upperparts dark grey-brown; angled inwards. Canine lacks a supplementary
underparts paler. Fur short. Some adults have cusp. Dental formula 2/1 1/1 3/3 2/2.
dull orange tufts on each side of the neck. Very
thick jaw, stout muzzle with massive rectangular MALAYAN SPOTTED-WINGED FRUIT BAT
cheek teeth (Fig. 28d). Similar species Rousettes, Balionycteris seimundi PLATE 6
Rousettus spp., have narrow muzzles and smaller Measurements FA 40–45, T none, E 10–12.
teeth; short-nosed fruit bats, Cynopterus spp., differ Wt 10–15g
in colour, and have smaller muzzles and teeth. Identification Smallest fruit bat in region. Upper-
Ecology and habitat Reported roosts include hollow parts dark blackish-brown, darkest on the head;
trees and in the vicinity of limestone caves. Feeds underparts pale grey-brown. Wing membranes dark
mainly in canopy of mature lowland rainforests; has brown, sparsely spotted with buff, especially on joints.
been caught feeding in fig trees. Adult males have Pale spots on edge of ear and in front of eye. One pair
been reported lactating, but it is not known whether of lower incisors. Taxonomic notes Formerly called
they can suckle young. Balionycteris maculata, but that is now considered a
Distribution SE Asia: peninsular Thailand and distinct species restricted to Borneo. Similar species
Malaysia. Also Borneo and the Philippines. Other fruit bats in the region lack spots on wings;
Status Near Threatened, owing to loss of lowland Black-capped Fruit Bat, Chironax melanocephalus,
rainforest. has two pairs of lower incisors.
Ecology and habitat Lowland dipterocarp forest
Genus Chironax Small bat, very similar to where active in understorey. In Peninsular Malaysia
Cynopterus, but lacking a tail; canines lack an inner has been found roosting in small groups in crowns
cusp; premaxillae fused (in contact but not fused in of palms, clumps of epiphytic ferns, hollowed
Cynopterus). Dental formula 2/2 1/1 3/3 1/2. termite nests, rarely in caves.
Distribution SE Asia: peninsular Thailand and
BLACK-CAPPED FRUIT BAT Malaysia.
Chironax melanocephalus PLATE 6 Status Near Threatened, owing to loss of lowland
Measurements FA 43–46, T none, E 13. Wt 15.7g rainforest.
Identification Upperparts dark grey or brown;
head darker, sometimes black; underparts pale Genus Aethalops Small tailless bat similar to
brownish-grey; chin yellowish. Most adults have Balionycteris, but without the second upper molar.
yellow-orange tufts on sides of neck. Two pairs of Rostrum slightly lower and more sloping than that of
lower incisors. Similar species Other small tailless Cynopterus (Fig. 26c, d). Upper incisors about equal
fruit bats have only one pair of lower incisors (as in in length. Interfemoral membrane narrow but thickly
Fig. 29b), and they differ in colour. furred. Dental formula 2/1 1/1 3/3 1/2.
Ecology and habitat Has been netted in the
understorey of tall dipterocarp forest, but more GREY FRUIT BAT
common in canopy. Reported roosts include small Aethalops alecto PLATE 6
groups in tree ferns and shallow caves. Found in Measurements FA 42–46, T none, E 10–13.
lowlands and hills. Wt 14–18g

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 Identification Upperparts dark grey-brown to Distribution SE Asia: N Myanmar, N Thailand and
reddish-brown, underparts slightly paler. Fur thick N Vietnam. Also NE India, E Nepal, Tibet and SW
and long. Interfemoral membrane covered in thick, China.
fluffy fur. Muzzle narrow and pointed. One pair of Status Least Concern.
lower incisors. Similar species Tailless fruit bats,
Megaerops spp., have a thicker, shorter muzzle; Genus Eonycteris Medium to large fruit bat with a
Black-capped Fruit Bat, Chironax melanocephalus, long, narrow muzzle and a very long, sticky tongue.
has two pairs of lower incisors; Spotted-winged Distinguished from all other fruit bats in the region
Fruit Bat, Balionycteris maculata, has distinct pale by the absence of a claw on the second finger. Tail
spots all over the wings; Blanford’s Fruit Bat, well developed. Skull similar to that of Rousettus,
Sphaerius blanfordi, is larger, with white rims to but rostrum longer and lower; braincase more
ears and two pairs of lower incisors. heavily deflected downwards (Fig. 26e, f). Cheek
Ecology and habitat Apparently confined to teeth narrow and elongate with reduced cusps.
montane forest above 1,000m. Lower canines small and simple, heavily curved
Distribution SE Asia: known only from hill forest outwards. Dental formula 2/2 1/1 3/3 2/3.
in Peninsular Malaysia. Also Sumatra, Java, Bali
and Lombok (form in Borneo is now considered a CAVE NECTAR BAT
separate species, A. aequalis). Eonycteris spelaea PLATE 7
Status Least Concern, but some declines due to Measurements FA 62–70, T 15–18, E 17–20.
loss of forest. Wt 45–60g
Identification Upperparts grey-brown; underparts
Genus Sphaerius Medium-small fruit bat with no slightly paler, sometimes tinged with yellow or
tail; interfemoral membrane narrow and thickly orange around the neck. Fur short. Muzzle elongate,
covered with hairs. Long, low rostrum. Dental teeth rather small. Lacks a claw on the second digit.
formula 2/2 1/1 3/3 1/2. Similar species Rousettes, Rousettus spp., are
larger, with a claw on the second digit; nectar bats,
BLANFORD’S FRUIT BAT Macroglossus spp., are much smaller.
Sphaerius blanfordi PLATE 6 Ecology and habitat Roosts in large, noisy colonies
Measurements FA 52–60, T none, E 16–19.5. in caves, often in virtual darkness. Flies far daily
Wt 27–30g in search of flowering trees, to feed on pollen and
Identification Dull grey fur with brown tips, nectar. Important pollinator of many forest trees,
slightly paler below. Interfemoral membrane very including commercially important species such as
narrow, thickly covered with dense fur; no tail. durian, Durio spp., mangroves and Parkia. Forages
Ears dark and hairless, rimmed with white. Wing in canopy of primary forest, as well as in gardens,
bones conspicuously pale, contrasting with darker mangroves and disturbed areas.
membranes. Two pairs of upper and lower incisors; Distribution SE Asia: Myanmar, Thailand, Laos,
incisors relatively well developed, positioned anterior Vietnam, Cambodia and Peninsular Malaysia. Also
to a line joining front of canines. Similar species N India to S China, Sumatra, Java, Borneo, Sulawesi
Short-nosed fruit bats, Cynopterus spp., also have and other Indonesian islands and the Philippines.
white rims to ears, but are brighter in colour Status Least Concern, but has declined in some
(orange or yellow collars in adults), and have broad areas owing to disturbance of cave roosts.
interfemoral membrane without thick fur, a short
tail, larger cheek teeth but smaller incisors; tailless Genus Macroglossus Small bats with long, narrow
fruit bats, Megaerops spp., have light brown fur with muzzle and a very long tongue. Tail minute or
pale grey base, only one pair of lower incisors; Grey lacking. Skull has a very narrow rostrum with
Fruit Bat, Aethalops alecto, is smaller, with only one the braincase strongly deflected downwards (Fig.
pair of lower incisors. 26g). Lower jaw thin and weak. Cheek teeth small
Ecology and habitat Reported from hill and with a large diastema between the first two upper
montane forests at altitudes of 300–2,700m. premolars. Canines long and needle-like, strongly

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 curved outwards on lower jaw. Upper incisors Distribution SE Asia: S Vietnam, Cambodia,
tiny, projecting slightly forwards and separated S Thailand and Peninsular Malaysia, mainly in
from each other and canines by small gaps. coastal areas. Also the Philippines, Java, Borneo
Genetic analyses of mainland populations suggest and Indonesian islands through to New Guinea, the
these species cannot be differentiated based on Solomon Islands and N Australia.
mitochondrial DNA; further analyses are required Status Least Concern.
to determine whether they are actually two different
species. Dental formula 2/2 1/1 3/3 2/3. GREATER LONG-TONGUED NECTAR BAT
Macroglossus sobrinus PLATE 7
LESSER LONG-TONGUED NECTAR BAT Measurements FA 44–50, T none, E 15–17.
Macroglossus minimus PLATE 7 Wt 19–29g
Measurements FA 39–44, T none, E 12–15.5. Identification Upperparts clay-brown with soft,
Wt 11–18g fine fur; underparts paler. Long, narrow muzzle
Identification Upperparts buffy-brown with with angular, slightly jutting chin. Nostrils lacking
pale base; underparts paler and greyer. Wing any distinct grooves between them on the muzzle.
membranes light brown. Long, narrow muzzle with Similar species Lesser Long-tongued Nectar Bat,
very small teeth except for needle-like canines. M. minimus, is smaller with a shorter head, more
Nostrils rounded and facing forwards, with a distinct pointed chin, more forward-pointing nostrils, and
median groove between them running down to distinct median grooves on upper lip.
upper lip, and two smaller grooves under nostrils Ecology and habitat Found primarily in inland
in front. Similar species Greater Long-tongued habitats, including dipterocarp forest and montane
Nectar Bat, M. sobrinus, is slightly larger, with forest up to 2,000m, as well as disturbed areas.
longer head, jutting chin, more lateral-pointing Feeds on nectar and pollen, especially from banana
nostrils and no grooves on upper lip. flowers, both wild and cultivated. Has been found
Ecology and habitat In SE Asia, found principally roosting in young rolled banana leaves.
in coastal habitats, particularly in mangroves, but Distribution SE Asia: Myanmar, Thailand, Laos,
ranges more widely in areas such as Borneo, Vietnam, Cambodia and Peninsular Malaysia. Also
where M. sobrinus is absent. Feeds on nectar and E India, Sumatra, Java and some small adjacent
pollen from many sources, including mangrove and islands.
banana flowers. Often uses disturbed areas. Status Least Concern.

Family RHINOPOMATIDAE MOUSE-TAILED BATS

Medium-sized bats with a very long tail, most of which protrudes beyond the interfemoral membrane. A
thickened band of skin joins the ears across the top of the forehead. There is a thickened pad on the muzzle,
which may have a slight ridge. The ears have a distinct tragus. Three species are currently recognised in
the family, of which one or two have been reported from mainland South-east Asia – there are no recent
records, and there is still some uncertainty about the identity of the specimens that have been recorded
from Myanmar and Thailand. To aid identification of any further specimens, the accounts below include both
species, although only one is illustrated on the plates.

Genus Rhinopoma The only genus in the family. GREATER MOUSE-TAILED BAT
Only one pair of upper incisors and one upper Rhinopoma microphyllum NOT ILLUSTRATED
premolar. Skull has a very short rostrum with Measurements FA 60–75, T 50–77, E 18–22.
swollen nasal cavities. Dental formula 1/2 1/1 Skull: ccl 17.2–22.7
1/2 3/3. Identification Large mouse-tailed bat (FA averages
68). Fur is short and fine, grey-brown above, paler
below, restricted to head and front half of body.
Membranes, lower back and lower belly naked,
with brown skin. Tail long and protruding, usually

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 shorter than forearm. Dermal ridge on muzzle LESSER MOUSE-TAILED BAT
very small and indistinct. Similar species Lesser Rhinopoma hardwickii PLATE 8
Mouse-tailed Bat, R. hardwickii, is smaller, with Measurements FA 53–64, T 56–78, E 17–21.
proportionately longer tail and larger dermal ridge Skull: ccl 15.5–17.5
on muzzle. Identification Medium-sized mouse-tailed bat (FA
Ecology and habitat Found mainly in arid areas, averages 59). Fur is grey-brown above with paler
roosting in small caves, tunnels and buildings, often base, greyer below. Extensive areas of bare skin on
in relatively bright areas. In some parts of India lower back and belly. Tail very long, usually longer
undertakes seasonal migrations of up to 900km. than forearm. Tip of nose forms a thickened pad
When foraging, appears to be a relatively weak with a distinct ridge of skin (dermal ridge) that forms
flier, with rapid wing beats interspersed with steady a rudimentary noseleaf. Similar species Greater
glides. Feeds on insects. Mouse-tailed Bat, R. microphyllum, is larger and has
Distribution SE Asia: individual specimens of a proportionately shorter tail and smaller noseleaf.
Rhinopoma have been recorded from peninsular Ecology and habitat Primarily found in relatively dry
Thailand and Myanmar, but it is uncertain which habitats. Roosts in caves, between large boulders,
species they represent; possibly these were in deserted houses or temples, often using relatively
vagrant, migrating bats. Also Africa, through Middle bright areas. In northern climates builds up fat
East to Afghanistan, Pakistan and India, as well as reserves during periods of insect abundance, then
N Sumatra. moves to more sheltered roosts to live off reserves
Status Least Concern. in winter. Roosts range from a few to several
hundred individuals and are sexually segregated.
Distribution SE Asia: see notes under Greater
Mouse-tailed Bat, R. microphyllum. Also Africa
through Middle East to India.
Status Least Concern.

Family EMBALLONURIDAE SHEATH-TAILED AND TOMB BATS

Small to medium-sized bats with a distinctive short tail that emerges from the middle of the interfemoral
membrane (PLATE 8). When legs are stretched out, membrane largely encloses tail. Muzzle is simple with
no noseleaf. Ears are short to moderate, with a short, rounded tragus. Eyes are relatively large, and these
bats often roost in fairly bright areas, suggesting vision may be important, perhaps for detecting potential
predators. Echolocation usually involves multiple harmonics, and lower frequencies of echolocation calls of
some species are audible to humans. The wings are long and narrow. The skull has a high, full braincase
and a moderately short rostrum (Fig. 31d). The postorbital processes are long and slender, and the
premaxillae bones (where the upper incisors attach) are small and delicate.

Genus Emballonura Small and dark brown. Two membrane. Distinctive posture, supported by
distinct pairs of upper incisors; anterior upper the wrists, and distinct, sharp, audible alarm
premolar very small. Skull with narrow rostrum, call when disturbed enables easy recognition of
very shallow depression between the eyes. Dental this genus at roost. Similar species This is the
formula 2/3 1/1 2/2 3/3. only small, dark, sheath-tailed bat known from
mainland SE Asia; tomb bats, Taphozous spp.,
LESSER SHEATH-TAILED BAT are much larger.
Emballonura monticola PLATE 8 Ecology and habitat Lives in forests, roosting in
Measurements FA 43–45, T 11–14, E 12–13. shallow caves or rock crevices, or under fallen tree
Wt 4.5–5.5g. Skull: ccl 11.8–12.8 trunks.
Identification Body fur uniform dark brown, Distribution SE Asia: peninsular Myanmar,
sometimes with a strong reddish tinge. Short tail Thailand and Malaysia. Also Sumatra, Java, Borneo,
protruding from the middle of the interfemoral Sulawesi and other Indonesian islands.

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 Status Least Concern; widespread, but has declined BLACK-BEARDED TOMB BAT
substantially in many areas due to loss of lowland Taphozous melanopogon PLATE 8
rainforest. Measurements FA 60–63, T 25–28, E 19–22.
Wt 23–26g
Genus Taphozous Medium-large bats, variably Identification Upperparts vary from grey-brown to
coloured. Most species have a flap of skin at the buff-brown; underparts usually paler, sometimes
wrist (metacarpal pouch), forming a pocket in the almost white. Chin covered with fur, lacking a
wing; some species have a glandular pouch on the throat pouch; most adult males have a black
throat. Tomb bats (including Saccolaimus) can be ‘beard’. Fur on upper surface of wing membrane
recognised in flight by long, narrow wings, strong, limited to about one-third length of upper arm
high flight and often audible clicks of echolocation (humerus). Well-developed metacarpal pouch. Tail
calls; species can sometimes be identified by slightly swollen at tip. Wing membrane attaches to
size or colour (such as the very white wings of tibia, above ankle. Wings pale, appearing whitish
Saccolaimus saccolaimus). Only one pair of upper in flight. Similar species Pouched Tomb Bat,
incisors, which are very small and sometimes Saccolaimus saccolaimus, has a reduced wing
fall out. Skull with broad rostrum, deep hollow pouch and a distinct throat pouch; Theobald’s
between eyes (Fig. 31d). Dental formula 1/3 1/1 Tomb Bat, T. theobaldi, is larger and lacks hair on
2/2 3/3. the legs; Long-winged Tomb Bat, T. longimanus,
is slightly smaller, has a naked chin with a throat
THEOBALD’S TOMB BAT pouch in males, longer metacarpals, tail tapering
Taphozous theobaldi PLATE 8 evenly to the tip and wing membrane attached at
Measurements FA 70–76, T 25–30, E 22–28 the ankle.
Identification Upperparts usually dark brown to Ecology and habitat Roosts in caves, large
grey-brown with paler base to hair, underparts crevices in rocks and temples, often in fairly well-lit
paler. Well-developed metacarpal pouch (see areas such as near cave entrance. Colonies range
colour plate). Chin well haired without a throat from a few individuals to several thousand. Like
pouch. Some adult males may have a patch other tomb bats, flies rapidly and high, foraging
of dark hairs forming a beard. Legs, feet and for insects over forests or other habitats, including
interfemoral membrane lack hair. Wings attach to highly disturbed areas.
tibia, above ankle. Similar species Pouched Tomb Distribution SE Asia: Myanmar, Thailand, Laos,
Bat, Saccolaimus saccolaimus, differs in colour Vietnam, Cambodia and Peninsular Malaysia. Also
and has a well-developed throat pouch, but lacks Sri Lanka, India to S China, Sumatra, Java, Borneo
a metacarpal pouch and has shorter ears; Black- and smaller islands.
bearded Tomb Bat, T. melanopogon, is similarly Status Least Concern.
coloured but smaller, with fur on interfemoral
membrane and base of wing membranes. LONG-WINGED TOMB BAT
Ecology and habitat Most known roosts are in Taphozous longimanus PLATE 8
caves, sometimes forming colonies of over 1,000 Measurements FA 54–63, T 20–30, E 16–19
individuals. Feeds on insects flying high over the Identification Colour dark brown to blackish,
forest. occasionally speckled with white. Chin naked;
Distribution SE Asia: S Myanmar, C Thailand, Laos, males have a throat pouch while females have only
Cambodia and Vietnam. Also India, Java, Sumatra, a rudimentary fold. Wings long and narrow, dark
Borneo and Sulawesi. brown and attached at ankle. Longest wing bone
Status Least Concern, but is known from relatively on third finger (metacarpal 3) longer than forearm.
few colonies, and vulnerable to disturbance of cave Moderately developed metacarpal pouch. Tail
roosts and quarrying. tapers narrowly to tip. Short fur on legs, interfemoral
membrane, and upper surface of wing membrane
to about half length of humerus and femur.
Similar species Pouched Tomb Bat, Saccolaimus

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 saccolaimus, is larger, lacks a metacarpal pouch
and has bare legs; Black-bearded Tomb Bat, a
T. melanopogon, has a furred chin in both sexes
with no throat pouch, wing membrane attached to
side of leg above ankle, tail thickened near tip, third
metacarpal shorter, about 90% of forearm length,
less extensive fur on flight membranes.
Ecology and habitat Has been found roosting
under eaves of houses, in hollow trees, under
rocks, in crowns of palm trees and in caves, often b
in relatively bright areas.
Distribution SE Asia: Myanmar, Thailand, Cambodia,
Laos and Peninsular Malaysia. Also Sri Lanka, India,
Sumatra, Java, Borneo, Flores and Bali.
Status Least Concern.
c
NAKED-RUMPED TOMB BAT
Taphozous nudiventris PLATE 8
Measurements FA 71–80, T 22–46, E 18–25
Identification Fur dark brown above with pale
grey base; paler below. Dorsal surfaces of wing
and tail membranes, legs and arms lack fur. d
Lower back and lower abdomen naked. Ears long
and relatively narrow. Wing has a moderately
developed metacarpal pouch, with fur on underside
of wing. Chin and throat naked; males have
a large gular sac and chest gland, which are
lacking in females. Similar species All other
tomb bats usually have fur extending all over body; Fig. 31 Skulls of Nycteris tragata (a – top, b – side),
Pouched Tomb Bat, Saccolaimus saccolaimus, Megaderma spasma (c) and Taphozous melanopogon (d).
lacks metacarpal pouch; Black-bearded Tomb Bat,
T. melanopogon, has a furred chin in both sexes Genus Saccolaimus Medium-large bats, similar
with no throat pouch, wing membrane attached in size, shape and behaviour to Taphozous, but
above ankle, tail thickened near tip. differing in lack of a pouch on the wing, presence
Ecology and habitat Roosts in caves, crevices in of a groove on the lower lip, completely ossified
rocks, temples and other buildings. Has been found auditory bullae and a relatively large anterior upper
roosting under the eaves of houses, in hollow trees premolar. Dental formula 1/3 1/1 2/2 3/3.
and under rocks. Most colonies are small, with only a
few individuals, although a colony of several hundred POUCHED TOMB BAT
individuals has been reported. May develop large Saccolaimus saccolaimus PLATE 8
fat reserves when insects are abundant at end of Measurements FA 71–78, T 33–34, E 19–21.
monsoon period; these are visible under skin on rump. Wt 40–50g
In some areas may undertake seasonal migrations, Identification Upperparts blackish-brown variably
and spends winter in a state of torpor. Forages on marked with white; underparts usually pure white,
insects with rapid, strong flight high in sky. but sometimes all dark brown. No metacarpal
Distribution SE Asia: NW Myanmar. Also Africa, pouch. Has a distinct glandular pouch under the
India and Pakistan. chin in both sexes, although slightly smaller in
Status Least Concern; only a few records from females. Legs and feet hairless. Similar species
region, but common elsewhere. Other species of tomb bat, Taphozous spp., have

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 well-developed metacarpal pouches; Black-bearded crevices. Forages high above the ground with a
and Long-winged Tomb Bats, T. melanopogon and strong, rapid flight.
T. longimanus, are smaller, with fur on the legs; Distribution SE Asia: S Myanmar, S Thailand, S
Theobald’s Tomb Bat, T. theobaldi, lacks pouch Vietnam, Cambodia, and Peninsular Malaysia. Also
under chin and has longer ears. India, Sri Lanka, Sumatra, Java and Borneo through
Ecology and habitat Sometimes found in houses, to New Guinea and Australia.
in colonies varying from a few individuals to a Status Least Concern.
few hundred. Also roosts in hollow trees and rock

Family CRASEONYCTERIDAE HOG-NOSED BATS

This family, with only one known species, was not discovered to science until 1974. Hog-nosed bats
are very small, with no visible tail and no calcar, but a well-developed interfemoral membrane. Muzzle
is thickened around the nostrils and chin, with crescent-shaped nostrils opening to the front of the face,
giving a pig-like appearance. Ears are relatively large, separated on the head, with a well-developed tragus.
Males have a glandular swelling on the throat. Dentition resembles Rhinopoma, but upper incisors are
proportionately larger. Dental formula 1/2 1/1 1/2 3/3.

KITTI’S HOG-NOSED BAT Ecology and habitat Roosts in limestone caves

Craseonycteris thonglongyai PLATE 8 in small colonies. Tolerant of limited disturbance
Measurements FA 22–26, T none, E 9–12. within cave roosts. Leaves the caves just before
Wt 2–3.2g dusk to forage for insects around trees. Can
Identification Fur brown to reddish-brown in manage in areas with extensive agriculture and
adults; juveniles greyer. Distinguished from all other disturbance, provided some areas of trees remain.
bats by very small size, pig-like muzzle with a slight Distribution SE Asia: Found only in a limited area of
ridge on the front and lack of a tail. Thailand and W Thailand and adjacent areas of Myanmar.
Myanmar populations are genetically distinct, and Status Vulnerable, owing to limited range, small
differ in echolocation calls: 71–78kHz (Thailand), population size and evidence of recent declines.
78–83kHz (Myanmar).

Family NYCTERIDAE SLIT-FACED BATS

Medium to large bats with very long ears separated at the base, and a short, rounded tragus. The tail is
long with a T-shaped tip and fully enclosed in the interfemoral membrane. The face has a deep groove in
the middle, bordered by leaf-like flaps of skin. The skull is distinguished by a deep frontal depression with
broad ridges on either side (Fig. 31a, b). The premaxillae are broadly attached to the palate. Dental formula
2/3 1/1 1/2 3/3. The family contains only the genus Nycteris.

MALAYAN SLIT-FACED BAT occurring in Java, N. javanica. Similar species All

Nycteris tragata PLATE 9 other bats in region have very differently shaped
Measurements FA 46–55, T 65–72, E 29–31. noseleaf; false-vampires, Megaderma spp., have
Wt 12–22g ears joined at base, long forked tragus, and no tail.
Identification Fur long and fluffy, greyish-brown Ecology and habitat Roosts in small groups
to pale red-brown. Noseleaf and ears grey-brown in hollow trees or caves, largely in mature
with irregular pale patches. Deep hollow groove rainforest. Feeds on large insects which it gleans
in middle of face, fringed with large flaps which from the surface; may hunt by passive listening
form a type of noseleaf. Ears long and rounded, for prey.
not joined at base. Tragus short and bent. Very Distribution SE Asia: S Myanmar, Thailand and
long tail with T-shaped tip, totally enclosed in Peninsular Malaysia. Also Sumatra and Borneo.
interfemoral membrane. Taxonomic notes Has Status Near Threatened by loss of lowland
sometimes been considered same species as form rainforest.

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 Family MEGADERMATIDAE FALSE-VAMPIRES
Medium to large bats with a large, erect noseleaf and large ears joined across the top of the head. The
tragus is long and forked. The tail is very short, not visible externally, although the interfemoral membrane
is well developed. Despite the name, these bats do not drink blood and are unrelated to South American
vampire bats (which do drink blood). Several genera occur throughout the world. Until recently, only two
species were known from South-east Asia, considered to be in the same genus. Recent genetic analyses
indicate they belong to different genera, and in 2015 a new genus and species, Eudiscoderma thongareeae,
was described from Thailand.

Genus Megaderma Interorbital region of skull Genus Lyroderma Skull similar to Megaderma, but
not especially concave (Fig. 31c). Premaxillae first upper molar with indentation on labial side;
minute with no upper incisors; canines project well upper canine with distinct anterior and posterior
forwards with a distinct secondary posterior cusp; cusps. Dental formula 0/2 1/1 2/2 3/3.
anterior upper premolars minute and displaced
inwards. Dental formula 0/2 1/1 2/2 3/3. GREATER FALSE-VAMPIRE
Lyroderma lyra PLATE 9
LESSER FALSE-VAMPIRE Measurements FA 65–72, T none, E 35–45.
Megaderma spasma PLATE 9 Wt 37–52g. Skull: GL 27.8–31.4
Measurements FA 56–63, T none, E 30–42. Identification Fur long, greyish-brown above,
Wt 18–28g. Skull: GL 24.7–26.6 paler below. Posterior lobe of noseleaf elongate
Identification Fur pale grey to grey-brown. with stiffened central ridge, approximately parallel-
Noseleaf has long dorsal lobe with stiffened central sided flaps on sides and squared off at the top;
ridge and broad convex flaps on the sides. Base of base of lobe (intermediate noseleaf) narrow and
lobe (intermediate noseleaf) heart-shaped, nearly oval, not heart-shaped; anterior noseleaf relatively
as broad as anterior noseleaf. Anterior noseleaf small, not covering muzzle. Muzzle protruding,
also broad, largely covering muzzle, which is well relatively short of hairs. Ears very large, joined
furred. Ears very large, joined at base. Tragus long, at base for 30–50% of length. Similar species
forked into two branches. No visible tail, although Lesser False-vampire, M. spasma, is smaller,
interfemoral membrane is well developed. Similar has shorter posterior noseleaf with convex sides;
species Greater False-vampire, L. lyra, is larger heart-shaped intermediate noseleaf. Thongaree‘s
with posterior noseleaf longer, more parallel-sided False-vampire, E. thongareeae, has disk-shaped
and squared at top, median noseleaf rounded and posterior noseleaf.
not heart-shaped; Thongaree’s False-vampire, E. Ecology and habitat Roosts in caves, temples,
thongareeae, is slightly larger with disk-shaped disused buildings and tunnels. Eats large insects
posterior noseleaf; Malayan Slit-faced Bat, Nycteris and vertebrates, including lizards, small mammals
javanica, has very different-shaped noseleaf, short and birds. Can hunt by listening for prey-generated
tragus and very long tail with a T-shaped tip. sounds, such as rustling in the leaves.
Ecology and habitat Roosts in small groups in Distribution SE Asia: Myanmar, Thailand, Laos,
caves, tunnels and hollow trees. Diet consists Vietnam, Cambodia and Peninsular Malaysia. Also
mainly of large insects, which it can glean from Afghanistan to S China, Pakistan, India and Sri Lanka.
vegetation or catch in the air. Has been known to Status Least Concern.
kill other bats in traps and to eat lizards in captivity,
and hence may sometimes eat small vertebrates Genus Eudiscoderma Similar to Megaderma, but
in the wild. lacks anterior upper premolar, no secondary cusp
Distribution SE Asia: Myanmar, Thailand, Laos, on canine, crown area of first lower premolar larger
Vietnam, Cambodia and Peninsular Malaysia. Also Sri than second, deeper depression in frontal area of
Lanka, India, the Philippines, Sumatra, Java, Borneo, rostrum. Dental formula 0/2 1/1 1/2 3/3.
Sulawesi, Maluku and other Indonesian islands.
Status Least Concern.

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 THONGAREE’S FALSE-VAMPIRE developed. Similar species Lesser False-vampire,
Eudiscoderma thongareeae PLATE 9 M. spasma, is smaller, has posterior noseleaf
Measurements FA 64–66.5, T none, E 32–37.5. somewhat triangular, narrowing to top. Greater False-
Wt 30–36g. Skull: gl 29.2–29.4 vampire, L. lyra, has posterior noseleaf longer, more
Identification Fur pale grey to grey-brown. Noseleaf parallel-sided and squared at top; median noseleaf
has disk-shaped dorsal lobe with stiffened central narrower without lobes on sides; naked muzzle.
ridge and broad flaps on the sides that are the same Ecology and habitat Known only from lowland
width as height. Base of lobe (intermediate noseleaf) rainforest. Has been found roosting in a hollow tree.
heart-shaped, similar width to anterior noseleaf. Prey includes large beetles.
Anterior noseleaf broad, largely covering muzzle Distribution SE Asia: Known only from Halabala
which is covered with fur. Ears very large, joined Wildlife Sanctuary in peninsular Thailand.
at base. Tragus long, forked into two branches. No Status Critically Endangered due to very limited
visible tail, although interfemoral membrane is well known range and threats to remaining habitat.

Family RHINOLOPHIDAE HORSESHOE BATS

Horseshoe bats are small to medium-sized bats with an elaborate noseleaf (Fig. 32). The anterior section
is rounded and roughly horseshoe-shaped. In the middle, behind the nostrils, is a raised portion called the
sella. Behind this is the posterior noseleaf, which usually rises to a long point, called the lancet. The shape
of the sella (Fig. 33) and the connecting process which joins it to the posterior leaf (see colour plates) varies
between species and is a useful diagnostic character. The ears are large, with a prominent fold on the
outside edge, the antitragus. The eyes are small and may be partially hidden from the front by the noseleaf.
The tail is moderately long and almost completely enclosed within the interfemoral membrane.
The family contains only one genus, Rhinolophus. Most of the species can be distinguished by a
combination of size and the shape of the noseleaf, but a few species are very similar in external appearance,
sometimes differing only in skull characters, particularly the shape and size of the nasal chambers. In some
cases, the taxonomy is still not fully understood, and some groups currently recognised as one species may,
after additional research, prove to represent multiple species.
In the field, similar species can often be distinguished by the frequency of the echolocation calls. In the
Rhinolophus genus these calls consist of a relatively long, high-energy, constant-frequency component,
with a short, frequency-modulated sweep at the end. This constant-frequency component is usually
characteristic of the species, at least in a given region, and can be determined using a bat detector (see
page 21 for more information on bat detectors). The echolocation frequencies given in the text refer to this
constant-frequency component. These are given with the location where it was recorded, as they may vary
geographically.
The skull is long with an inflated bump on the rostrum (Fig. 34a) that contains a series of nasal
chambers, which vary in size and shape and are presumably involved in echolocation. Upper incisors are
small and located on prolonged premaxillae, which are attached only to the palate and not to the maxillae.
Anterior upper premolar and middle lower premolar are often very small and displaced inwards. The canines
are heavy but simple, without conspicuous secondary cusps. Dental formula 1/2 1/1 2/3 3/3.
To help with identification of the large number of species, they are divided here into groups with similar
morphological characters – these do not necessarily represent taxonomic groups.

The following five species have well-developed TREFOIL HORSESHOE BAT

lateral lappets at the base of the sella (Fig. 32), with Rhinolophus trifoliatus PLATE 10
long woolly hair. They can be distinguished by colour Measurements FA 45–56, T 27–37, E 23–27.
and size. Wt 10–20g. Skull: ccl 19.5–22.4, mt 8.1–9.4
Identification Fur long and woolly, pale buffy-
brown to brownish-grey. Noseleaf pale yellow; ears
and wing membranes yellowish-brown with yellow

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 Ear

Lancet Antitragus

Posterior noseleaf

Connecting process Eye

Sella Internarial septum

Lateral lappet
Nostril
Anterior noseleaf

Fig. 32 Diagram of Rhinolophus noseleaf showing naming of parts.

elbows and knees. Noseleaf has lateral lappets at (peninsular Thailand and Borneo). Taxonomic
base of sella. Echolocation 50–54kHz (Sabah and notes Newly described in 2015. Form in Thailand
Peninsular Malaysia). Similar species Other woolly is genetically somewhat distinct and considered a
horseshoe bats with lateral lappets have dark brown distinct subspecies, R. f. thailandicus; specimens
fur and a dark noseleaf. from Vietnam also differ genetically. Similar
Ecology and habitat Occurs in understorey of species Trefoil Horseshoe Bat, R. trifoliatus, is
forests from lowlands to hills, including secondary similar in size, but with pale fur and a yellowish
forests. Roosts alone in foliage of forest understorey, noseleaf; Lesser Woolly Horseshoe Bat, R. sedulus,
including under palm and rattan leaves. Hunts by is smaller, while Great Woolly Horseshoe Bat,
hanging from an open branch, echolocating and R. luctus, is much larger.
waiting for insects to fly by. Ecology and habitat Thailand specimen was
Distribution SE Asia: S and W Thailand, S Myanmar caught over a small stream in understorey of
and Peninsular Malaysia. Also NE India, China, evergreen forest; in Borneo found in pristine and
Sumatra, Java, Borneo and some small adjacent evergreen forest at altitudes up to 1,600m.
islands. Distribution SE Asia: Western Thailand, Vietnam.
Status Least Concern. Also Borneo.
Status Not yet assessed.
FRANCIS’S WOOLLY HORSESHOE BAT
Rhinolophus francisi PLATE 10 LESSER WOOLLY HORSESHOE BAT
Measurements FA 52–55, T 30–38, E 24–27. Rhinolophus sedulus PLATE 10
Wt 16–18g. Skull: ccl 21.7–23.9, mt 9.3–9.8 Measurements FA 38–45, T 22–25, E 21–24.
Identification Similar in size and shape to Trefoil Wt 7–11g. Skull: ccl 17.4–18.3, mt 6.6–7.6
Horseshoe Bat, R. trifoliatus, but with dark fur Identification Fur long and fluffy, uniformly dark
and dark noseleaf. Echolocation 49–50kHz brown to dark grey. Noseleaf and ears dark grey.

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 Noseleaf has lateral lappets at base of sella. HILL WOOLLY HORSESHOE BAT
Echolocation 67kHz (Peninsular Malaysia), 62kHz Rhinolophus luctoides NOT ILLUSTRATED
and 76kHz (two individuals in Sabah). Similar Measurements FA 58–64. Wt 22–32g.
species Other woolly horseshoe bats with Skull: ccl 23.6–25.2, mt 10.8–11.6, lower toothrow
lateral lappets are larger; Trefoil Horseshoe Bat, 11.6–12.3
R. trifoliatus, is larger and has pale grey fur and a Identification Very similar in appearance to Great
yellowish noseleaf. Woolly Horseshoe Bat, R. luctus, but differs in
Ecology and habitat Roosts in bushes or hollow genetics and chromosomes; proportionately longer
trees. Forages in the understorey of tall forest up to lower toothrow (ratio of toothrow to mandible >0.60
1,500m. Occurs at relatively low densities. vs. <0.59); longer baculum in males (4.1–4.8mm
Distribution SE Asia: Peninsular Malaysia. Also compared with 3.1–3.2mm). Echolocation 42kHz.
Borneo. Taxonomic notes Newly described in 2015 based
Status Near Threatened owing to loss of lowland on genetic and chromosomal distinctions; a possible
rainforest. hybrid with R. luctus has also been reported.
Ecology and habitat Has been caught in hill and
GREAT WOOLLY HORSESHOE BAT montane forest from 600–1,400m altitude, at higher
Rhinolophus luctus PLATE 10 altitudes than R. luctus in Peninsular Malaysia.
Measurements FA 62–76, T 50–58, E 31–42. Distribution SE Asia: known only from Peninsular
Wt 30–45g. Skull: ccl 24.6–25.5, mt 10.5–10.7, Malaysia, but may occur elsewhere in region.
lower toothrow 11.1–11.3 Status Not yet assessed.
Identification Largest horseshoe bat in the region.
Fur long and woolly. Upperparts brownish-black; The following four species have an elongate sella
underparts slightly greyer. Noseleaf and ears dark (e.g. Fig. 33a), greatly enlarged ears and an
grey-brown. Noseleaf has lateral lappets at base of expanded internarial cup at the base of the sella.
sella. Taxonomic notes Recent analyses suggest Despite the similarities, genetic analyses suggest
that R. luctus may be restricted to Java, and R. morio that the first two are not closely related to the other
is the appropriate name from peninsular Malaysia; two. They can be distinguished from one another by
however, there is considerable variation in size and size and the shape of the sella and the internarial
echolocation frequency throughout the range, and cup at the base of the sella.
further analyses are needed to clarify which names
belong to which populations. Echolocation 40–42kHz BIG-EARED HORSESHOE BAT
(Sabah and Peninsular Malaysia), 32–35kHz (Thailand Rhinolophus macrotis PLATE 11
and Laos). Similar species Hill Woolly Horseshoe Measurements FA 42–47, T 17–25, E 22–28.
Bat, R. luctoides, is very similar, distinguished only Wt 6–9.5g
by tooth and baculum measurements. Francis’s Identification Very large ears with well-developed,
Woolly Horseshoe Bat, R. francisi, is smaller. Trefoil rounded antitragus. Sella broad and tall, broadest
Horseshoe Bat, R. trifoliatus, is smaller with pale fur at base and narrowing gradually to rounded tip;
and a yellowish noseleaf. base expanded into an internarial cup in front of
Ecology and habitat Roosts in small groups in sella. Connecting process tall, parallel to sella at
caves and rock crevices, in hollow trees and among base, rounded at top and forming a distinct notch
roots of trees. Sometimes forages by hanging from where it meets sella, which is angled forwards
a branch, flying out periodically to grab food, then at tip. Lancet well developed, rounded at top.
returning to its branch. Found in both primary and Horseshoe broad, covering muzzle, with well-
disturbed forests. developed secondary noseleaf beneath it. Upperparts
Distribution SE Asia: Myanmar, Thailand, Laos, mid-brown, underparts paler. Echolocation 48kHz
Vietnam, Cambodia and Peninsular Malaysia. Also (Peninsular Malaysia), 51–52kHz (Laos). Similar
N India through S China, Taiwan, Sumatra, Java, species Siamese Horseshoe Bat, R. siamensis, is
Borneo and Bali if all populations are the same species. smaller, with higher-frequency echolocation calls;
Status Least Concern. Marshall’s Horseshoe Bat, R. marshalli, has low,

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 a b

a b c d
c d

e f g h

Fig. 33 Front views of sella of selected species of Fig. 34 Front and side views of typical Rhinolophus
Rhinolophus: R. marshalli (a), R. thailandensis (b), R. (a, b) and Hipposideros (c, d) skulls, showing differences
pearsonii (c), R. robinsoni (d), R. affinis (e), R. stheno (f), in shape of inflated nasal area.
R. malayanus (g) and R. refulgens (h).

arched connecting process, internarial cup continuing Also China.

behind sella, shorter lancet and broader noseleaf. Status Least Concern.
Ecology and habitat Has been found in a variety of
forest types, including primary forest and disturbed KING HORSESHOE BAT
forests, from lowlands to 1,000m. No information Rhinolophus rex PLATE 11
on roosting sites. Measurements FA 53–57, T 21–33, E 32–35.
Distribution SE Asia: scattered records from through- Wt 8–14g
out Thailand, Myanmar, N Laos, N Vietnam and Identification Enormous ears, about 60% of
Peninsular Malaysia. Also Pakistan, N India, Nepal forearm length, with long, narrow antitragus. Sella
and S China. (Note: specimens from the Philippines very broad and tall, with a broad, rounded cup-like
formerly referred to this species are not closely structure at the base that continues behind the sella
related, and represent a distinct species, R. hirsutus.) to join the connecting process. Connecting process
Status Least Concern. low, extending in a long arch to the tip of the sella.
Lancet low and rounded. Horseshoe very broad, and
SIAMESE HORSESHOE BAT completely covering the muzzle with a deep notch in
Rhinolophus siamensis PLATE 11 the front. Fur of upperparts brown with paler base,
Measurements FA 38–42, T 16–22, E 19–22. underparts paler. Taxonomic notes Formerly called
Wt 4.5–5.7g R. paradoxolophus, but recent genetic analyses
Identification Very similar in shape and general indicate this is the same species as R. rex from
appearance to Big-eared Horseshoe Bat, China. Echolocation 24–25kHz (Laos). Similar
R. macrotis, with a similarly shaped noseleaf. species Marshall’s Horseshoe Bat, R. marshalli,
Differs mainly in smaller size and higher has similarly shaped noseleaf but is substantially
echolocation frequency. Taxonomic notes smaller, with proportionately shorter and narrower
Formerly considered a subspecies of R. macrotis, sella, longer lancet and more laterally extended
but both forms occur together, indicating they are internarial cup.
distinct species. Echolocation 68–74kHz (Laos). Ecology and habitat Known roosts all in limestone
Similar species Other small Rhinolophus have caves. Found in a variety of forest types, including
shorter sella and smaller ears, and do not have dry deciduous, pine, moist evergreen and disturbed
expanded internarial cup. riverine forests. Distribution SE Asia: Myanmar,
Ecology and habitat Has been found in disturbed N Thailand, Laos, N Vietnam. Also S China.
forest near limestone caves. Status Least Concern, but probably declining
Distribution SE Asia: N Thailand, Laos and Vietnam. somewhat due to loss of forest.

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 MARSHALL’S HORSESHOE BAT Shamel’s Horseshoe Bat, R. shameli, is very similar,
Rhinolophus marshalli PLATE 11 differing in slightly larger size, broader noseleaf and
Measurements FA 41–48, T 16–26, E 25–30. lower echolocation call frequencies.
Wt 6.5–7.5g Ecology and habitat Has been found roosting
Identification Enormous ears, with long, narrow in limestone caves, in colonies of up to several
antitragus. Sella broad and tall, with a cup-like hundred individuals. Forages in a range of forest
structure at base that continues behind sella to join types, from lowlands to hills.
connecting process (Fig. 33a). Basal cup extends Distribution SE Asia: C Myanmar, Thailand, Laos
laterally to sides. Connecting process low, extending and N Peninsular Malaysia.
in a long, smooth arch to tip of sella. Lancet rounded Status Least Concern.
and relatively short. Horseshoe very broad, covering
muzzle, with a deep anterior notch. Echolocation SHAMEL’S HORSESHOE BAT
40–41kHz (Laos). Similar species King Horseshoe Rhinolophus shameli PLATE 11
Bat, R. rex, is larger, with a proportionately longer Measurements FA 44–49, T 15–23, E 17–21.
sella and shorter lancet; Big-eared Horseshoe Bat, Wt 7.5–12.5g. Skull: gl 19.8–21.6, mt 7.5–8.6
R. macrotis, has a tall, rounded connecting process, Identification Very similar to R. coelophyllus,
and internarial cup not extending behind sella. with same shape noseleaf, but somewhat larger
Ecology and habitat Has been found roosting in and with lower echolocation frequency. Noseleaf
limestone caves, and in a variety of forest types typically broader than muzzle. Echolocation Male
from lowlands to hill forest at 800m. 65–67kHz, female 69–72kHz (Laos). Similar
Distribution SE Asia: known from scattered species Croslet Horseshoe Bat, R. coelophyllus, is
localities in C and N Thailand, N Laos, N Vietnam slightly smaller, with higher echolocation frequency.
and extreme N Peninsular Malaysia. Ecology and habitat Has been found in vicinity
Status Least Concern, although relatively uncommon. of limestone caves. Forages in deciduous and
evergreen forests, including heavily disturbed areas,
The following two species have a distinctive short up to 1,000m.
lancet, thickened at top and sides to enclose base Distribution SE Asia: E. Thailand, Laos, Vietnam
of strongly arched connecting process. Both are and Cambodia.
very similar, distinguished primarily by size and Status Least Concern.
echolocation frequency.
The following species are distinguished by having
CROSLET HORSESHOE BAT a triangular or pointed connecting process. They
Rhinolophus coelophyllus PLATE 11 include some of the smallest horseshoe bats in the
Measurements FA 41–45, T 15–24, E 15–20. region, as well as the medium-sized R. acuminatus.
Wt 6.2–8.6g. Skull: gl 18.2–20.1, mt 6.9–7.8 Within species, the shape of the connecting process
Identification Noseleaf has relatively short lancet, may vary from triangular to horn-shaped, and thus
thickened at sides and top to form lobes that protrude does not appear to be reliable for identification.
forwards and enclose base of connecting process. Some species are known only from dried skins, and
Lancet, including pocket in lobes, covered with long details of their noseleaf shape are not well known.
hairs. Connecting process tall and broadly arched Further research, using genetics and echolocation
from lancet to tip of sella. Sella is roughly parallel- recordings, may help to clarify species limits and
sided with a triangular tip. Noseleaf broad, nearly identification in this group.
covering muzzle, with rudimentary secondary noseleaf
beneath it. Fur typically mid-brown above, buffy- ACUMINATE HORSESHOE BAT
white below, but some individuals are dull orange Rhinolophus acuminatus PLATE 11
or reddish. Shows geographic variation in size and Measurements FA 44–50, T 22–31, E 18–21.
echolocation calls. Echolocation Male 83–84kHz, Wt 11–16g
female 86kHz (Laos), 74–85kHz (Myanmar, Thailand, Identification Noseleaf simple without lateral
with variation among regions). Similar species lappets on sella; connecting process triangular,

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 with blunt or sharp point at tip. Sella approximately MOUNTAIN HORSESHOE BAT
parallel-sided, but in some populations may be Rhinolophus monticolus NOT ILLUSTRATED
narrow in middle. Lancet has concave sides Measurements FA 41–44, T 19–26, E 15.5–19.5.
and narrow pointed tip. Two colour phases. In Wt 6.9–8.6. Skull: gl 17.4–19.7, mt 6.3–6.9, C–C
some, upperparts are dark greyish-brown with 4.2–4.5
pale frosting on tips of fur, underparts only Identification Small horseshoe bat with simple
slightly paler. In others, fur is bright orange all noseleaf and long, triangular, pointed connecting
over, slightly paler below. Noseleaf and ears process. Noseleaf broad, 7–8.5mm wide, covering
dark greyish-brown in both colour phases. Ears muzzle. Lancet tall, bluntly pointed, slightly concave
moderately large. Has a distinctive, strong musky on sides. Sella broad, parallel-sided with squared-
smell. Echolocation Male 86–90kHz, female off tip. Upperpart fur dark grey-brown with pale tips
93– 95kHz (Laos). Similar species Intermediate and white base about 50% of length; underpart
Horseshoe Bat, R. affinis, has rounded connecting hairs brown with pale tips and white bases about
process; other species with pointed connecting 80% of length. Upper canine long, slender, twice
process are much smaller. height of P2. Large median chambers in rostrum.
Ecology and habitat Recorded roosting in caves Taxonomic notes Newly described in 2016.
and hollow trees. Has been found in understorey of Echolocation Male: 85–87kHz, female: 91–93kHz
dry evergreen and dry dipterocarp forest in Laos and (Thailand). Similar species Other small species
lowland rainforest in Malaysia. have higher echlocation calls. Least Horseshoe
Distribution SE Asia: Myanmar, Thailand, Laos, Bat, R. pusillus, is smaller with narrow sella, short
Cambodia, Vietnam and Peninsular Malaysia. Also canine; Blyth’s Horseshoe Bat, R. lepidus, averages
the Philippines, Sumatra, Java, Borneo and many smaller with higher echolocation frequency, narrow
smaller Indonesian islands. sella with rounded tip; Shortridge’s Horseshoe Bat,
Status Least Concern. R. shortridgei, has rounded tip to sella, thicker
canines, smaller median chamber of rostrum.
LEAST HORSESHOE BAT Ecology and habitat Known from evergreen
Rhinolophus pusillus PLATE 11 montane forest at altitudes of 600–1,300m. Has
Measurements FA 33–39, T 13–22, E 11–17. been caught flying over small streams. It likely
Skull: gl 14.9–16.1, mt 5.4–6.1, C–C 3.3–4.1 roosts in hollow trees or rock crevices.
Identification Very small horseshoe bat with Distribution SE Asia: Thailand, Laos.
simple noseleaf and triangular, pointed connecting Status Not yet assessed.
process. Noseleaf small, not covering muzzle.
Lancet broadly pointed, slightly concave on sides. BLYTH’S HORSESHOE BAT
Sella parallel-sided. Upperparts vary from dark Rhinolophus lepidus NOT ILLUSTRATED
brown to smoky-grey or cinnamon-red brown, Measurements FA 38–41, T 15–21, E 13–17.
underparts paler. Taxonomic notes Genetic and Wt 4.5–8.0g. Skull: gl 15.7–17.3, mt 5.7–6.5,
size variation across range suggest it may include C–C 3.7–4.2
more than one species. Echolocation 108–115kHz Identification Small horseshoe bat with connecting
(Laos, Thailand). Similar species Blyth’s Horseshoe process triangular and pointed, sometimes hooked
Bat, R. lepidus, averages larger. slightly forwards. Lancet bluntly pointed, usually
Ecology and habitat Has been found roosting with concave sides. Sella narrow with rounded tip.
in caves, clumps of bamboo and even houses. Noseleaf narrower than muzzle. Three grooves on
Forages in a wide range of forest habitats, including lower lip. Upper canine long and thick, double height
mature forest and disturbed areas. of P4. Taxonomic notes Populations in peninsular
Distribution SE Asia: Myanmar, Thailand, Laos, Thailand and Malaysia now considered a separate
Vietnam, Cambodia and Peninsular Malaysia. Also species, R. refulgens. Echolocation 103kHz
India, S China, Borneo, Java and small adjacent (Thailand). Similar species Least Horseshoe Bat,
islands. R. pusillus, averages smaller, with more sharply
Status Least Concern. pointed connecting process; Convex Horseshoe

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 Bat, R. convexus, has rounded lancet; long, whether they are the same species.
pointed connecting process, Glossy Horseshoe Bat, Distribution SE Asia: known only from Peninsular
R. refulgens, has shorter upper canine. Malaysia and possibly Laos.
Ecology and habitat Found in dry deciduous and Status Data Deficient; uncertain status given
semi-deciduous forests. uncertainty in range.
Distribution SE Asia: Myanmar, N Thailand,
Cambodia, Vietnam. Also Afghanistan, N India, LITTLE NEPALESE HORSESHOE BAT
Pakistan, S China. Rhinolophus subbadius NOT ILLUSTRATED
Status Least Concern. Measurements FA 33–38, T 16–19, E 14–18.
Skull: gl 14.4–15.0, mt 5.2–5.6, C–C 2.7–3.2
GLOSSY HORSESHOE BAT Identification Very small horseshoe bat with pointed
Rhinolophus refulgens PLATE 11 connecting process that is slightly constricted in the
Measurements FA 37–43, T 15–22, E 13–18. middle and angled forwards. Lancet triangular with
Wt 4.0–7.2g. Skull: gl 15.8–17.1, mt 5.8–6.5, concave sides. Upperparts light brown with greyish
C–C 3.9–4.6 base to fur; shoulders and underparts slightly paler.
Identification Small horseshoe bat with connecting Rostrum narrow, as reflected in narrow width across
process triangular and pointed. Lancet bluntly upper canines, and width across upper molars
pointed, usually with concave sides. Noseleaf <5.0mm. Upper canines slender and moderately
narrower than muzzle. Sella with parallel sides long. Echolocation Not recorded. Similar species
and squared-off tip. Three grooves on lower Least Horseshoe Bat, R. pusillus, is very similar in
lip. Fur dark blackish-grey or reddish-brown, in external appearance but is slightly larger, with a
both cases with long pale tips to fur giving a broader rostrum with C–C 3.3–4.1.
glossy, frosted appearance. Upper canine short, Ecology and habitat Recorded from bamboo in
less than 1.5x height of P4. Taxonomic notes dense forest at 1,200m in N Myanmar.
Formerly considered same species as R. lepidus. Distribution SE Asia: N Myanmar. Also Nepal,
Echolocation 98–105kHz. Similar species Least India, Bangladesh, S. China (Yunnan).
Horseshoe Bat, R. pusillus, averages smaller, with Status Least Concern.
more sharply pointed connecting process; Blyth’s
Horseshoe Bat, R. lepidus, has longer canines. SHORTRIDGE’S HORSESHOE BAT
Ecology and habitat Largely restricted to mature Rhinolophus shortridgei NOT ILLUSTRATED
lowland rainforest. Measurements FA 38–42, T 16–25, E 14–19.
Distribution SE Asia: peninsular Thailand and Skull: gl 17.2–18.0, mt 6.5–7.2
Malaysia. Also Sumatra, Java. Identification Small horseshoe bat closely
Status Not assessed. resembling R. lepidus, with triangular connecting
process, but with broader sella and rounded tip,
CONVEX HORSESHOE BAT slightly larger skull, canines thick, twice height
Rhinolophus convexus PLATE 11 of posterior premolar, third lower premolar (P4 )
Measurements FA 42–43, T 18–22, E 15–16. slightly elongate rather than rounded. Upperparts
Wt 7.2–8.2g. Skull: gl 16.9–17.4, mt 6.4–6.6, light brown, hairs with grey base and brown tip;
C–C 4.1–4.2 underparts paler. Echolocation 94.5–100.5kHz.
Identification Small horseshoe bat with tall, Similar species Blyth’s Horseshoe Bat, R. lepidus,
pointed connecting process, low rounded lancet. has broader sella, more rounded third lower
Sella with straight sides that converge towards tip. premolar.
Echolocation 92kHz (Laos). Similar species Other Ecology and habitat Has been caught in pagodas
small Rhinolophus have more pointed lancet. and in caves in lowland, limestone karst near dry
Ecology and habitat Has been found in upper forest.
montane rainforest in Malaysia (1,600m). Similar- Distribution SE Asia: C Myanmar. Also NE India,
looking individuals have been found in hill forest in SW China.
Laos, but genetic studies are needed to determine Status Least Concern.

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 The remaining horseshoe species have an LESSER BROWN HORSESHOE BAT
unmodified noseleaf structure and a rounded Rhinolophus stheno PLATE 13
connecting process. These resemble what is Measurements FA 43–48, T 13–21, E 15–20.
thought to be the ancestral form of Rhinolophus, Wt 7–10.5g. Skull: gl 19.5–21.3, mt 7.2–8.1
without any additional flaps or special elongation Identification Medium-sized horseshoe bat with
of the connecting process. Despite morphological moderately large horseshoe. Connecting process
similarities, these are probably not all closely rounded, joining sella just below tip to form a
related to one another; they are distinguished from slight notch. Sella is narrow, parallel-sided and
one another by a combination of size, shape of the slightly pointed at the top (Fig. 33f). Upperparts
sella (Fig. 33) and shape of the lancet. In some vary from dark brown to reddish-brown; underparts
cases, echolocation calls are the easiest way to contrastingly paler. Noseleaf usually dark at
distinguish species. edges, contrasting paler in middle. Taxonomic
notes Formerly included R. microglobosus as a
INTERMEDIATE HORSESHOE BAT subspecies. Echolocation 85–88kHz (Peninsular
Rhinolophus affinis PLATE 12 Malaysia and Thailand). Similar species Malayan
Measurements FA 48–54, T 20–32, E 19–24. Horseshoe Bat, R. malayanus, is slightly smaller,
Wt 12–20g. Skull: gl 21.1–23.7, mt 8.4–9.7 has broader sella with square top (Fig. 33g), lower
Identification Upperparts dark brown to reddish- frequency echolocation calls; Peninsular Horseshoe
brown; underparts only slightly paler. Ears Bat, R. robinsoni, has dark fur above and below,
moderately large. Noseleaf relatively large, covering with sella abruptly narrower towards tip (Fig. 33d);
muzzle, simple without extra lappets on sella; Intermediate Horseshoe bat, R. affinis, is larger, on
connecting process broadly rounded, originating average, with underparts only slightly paler than
from below tip of sella, forming a slight notch. Lancet upperparts, sella constricted in the middle (Fig.
tall, triangular, with straight sides. Sella is narrow 33e); Thomas’s Horseshoe Bat, R. thomasi, has
with concave sides (Fig. 33e). Second phalanx of darker fur, short lancet with abrupt point in middle;
third digit (longest finger) is long, 66–80% of the Chasen’s Horseshoe Bat, R. chaseni, has more
length of the metacarpal. Echolocation 77–78kHz uniform-coloured fur, darker noseleaf; Indochinese
(Peninsular Malaysia), 73–78kHz (Laos). Similar Brown Horseshoe Bat, R. microglobosus, is smaller
species Lesser Brown Horseshoe Bat, R. stheno, with higher frequency echolocation calls, smaller
is smaller, with a parallel-sided sella, and usually median rostral chamber in skull.
relatively paler underparts; Acuminate Horseshoe Ecology and habitat Roosts in crevices of rock
Bat, R. acuminatus, has a pointed connecting boulders, limestone caves and hollow trees. Forages
process; Pearson’s Horseshoe Bat, R. pearsoni, has mainly in lowland rainforest.
long woolly fur, sella broad at base, constricting Distribution SE Asia: peninsular Thailand and
abruptly in middle; Rufous Horseshoe Bat, R. rouxii, Malaysia, and possibly SC Vietnam.
has shorter lancet with strongly convex sides, and Status Least Concern.
second phalanx of third digit less than 66% of
metacarpal length. INDOCHINESE BROWN HORSESHOE BAT
Ecology and habitat Has been found roosting in Rhinolophus microglobosus NOT ILLUSTRATED
caves. Forages in understorey of forest, including Measurements FA 41–46, T 14–22, E 13–18.
mature lowland rainforest, dry forest and disturbed Wt 5–9g. Skull: gl 18.3–20.1, mt 6.6–7.3
areas. Identification Medium-sized horseshoe bat with
Distribution SE Asia: Myanmar, Thailand, Laos, moderately large horseshoe. Connecting process
Vietnam, Cambodia and Peninsular Malaysia. Also rounded, joining sella just below tip to form a slight
India through China, Sumatra, Java and Borneo. notch. Sella is narrow, parallel-sided and slightly
Status Least Concern; this species is widespread pointed at the top (as in Fig. 33f). Upperparts vary
and common. from dark brown to reddish-brown; underparts
contrastingly paler. Noseleaf usually dark at edges,
contrasting paler in middle. Smaller on average in

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 all dimensions than R. stheno, with much smaller been found in a variety of dry forest types, including
median rostral chamber in skull. Taxonomic notes heavily degraded forest.
Formerly considered a subspecies of R. stheno. Distribution SE Asia: S Myanmar, Thailand, Laos,
Echolocation 94.5–96.5kHz (Laos). Similar N Vietnam and N of Peninsular Malaysia.
species Malayan Horseshoe Bat, R. malayanus, Status Least Concern; locally common in many
is slightly smaller, has broader sella with square areas.
top (Fig. 33g), lower frequency echolocation calls;
Intermediate Horseshoe Bat, R. affinis, is larger, CHASEN’S HORSESHOE BAT
with underparts only slightly paler than upperparts, Rhinolophus chaseni PLATE 12
sella constricted in the middle (Fig. 33e); Thomas’s Measurements FA 43–46, T 20–27, E 17–22.
Horseshoe Bat, R. thomasi, has darker fur, short Wt 8–14.5g
lancet with abrupt point in middle; Chasen’s Identification Medium-sized horseshoe bat with
Horseshoe Bat, R. chaseni, has more uniform- moderate-sized noseleaf that does not completely
coloured fur, darker noseleaf. cover muzzle; connecting process rounded; lancet
Ecology and habitat Roosts in crevices of rock tall and triangular. Sella slightly broader at base,
boulders, limestone caves and hollow trees. narrowing very slightly in middle, broadly rounded at
Forages in rainforest, dry forest and disturbed areas top. Upperparts dark brown, hairs with slightly paler
with some tree cover from lowlands to hills. tips; underparts only slightly paler. Noseleaf dark.
Distribution SE Asia: Myanmar, Thailand, Laos, Taxonomic notes Formerly considered conspecific
Vietnam. with R. borneensis, but genetically and morphologically
Status Least Concern. distinct. Specimens referred to R. borneensis have
been reported from Peninsular Malaysia, but their
MALAYAN HORSESHOE BAT identification has not been confirmed. Echolocation
Rhinolophus malayanus PLATE 13 77.5–78.5kHz (Laos). Similar species Peninsular
Measurements FA 38–44, T 18–27, E 17–20. Horseshoe Bat, R. robinsoni, has broader sella
Wt 5–9g. Skull: gl 17.0–18.2, mt 6.5–7.2 with more marked constriction in middle, lower
Identification Medium-small horseshoe bat with echolocation frequency; Thomas’s Horseshoe Bat,
moderately large horseshoe. Connecting process R. thomasi, has short lancet with abrupt point
rounded, joining sella just below tip to form a slight in middle; Lesser Brown and Malayan Horseshoe
notch. Sella is moderately broad, parallel-sided Bats, R. stheno and R. malayanus, have underparts
and squared off at top (Fig. 33g). Lancet tall and markedly paler than upperparts, paler centre of
triangular. Upperparts usually brown with contrasting noseleaf, different-shaped sella.
pale buff or whitish underparts; sometimes overall Ecology and habitat Has been found in dry
colour is more orange-brown. Noseleaf grey to dipterocarp forest, including in disturbed areas.
dark grey, paler in the middle, but without a Distribution SE Asia: Laos, Cambodia and Vietnam.
sharp contrast in colour. Echolocation 83–91kHz Status Least Concern (as R. borneensis).
(peninsular Thailand), 75–82kHz (Myanmar, Laos,
elsewhere in Thailand, except some populations THOMAS’S HORSESHOE BAT
have 82–86kHz). Similar species Lesser Brown Rhinolophus thomasi PLATE 12
Horseshoe Bat, R. stheno, and Indochinese Brown Measurements FA 42–46, T 18–25, E 17–24.
Horseshoe Bat, R. microglobosus, have narrower Wt 7–11g. Skull: gl 17.9–20.0, mt 6.8–7.7
sella that is slightly pointed at top (Fig. 33f), Identification Fur uniformly medium to dark brown,
and higher frequency echolocation calls; Least only slightly paler below; noseleaf dark grey, slightly
Horseshoe Bat, R. pusillus, has proportionately pinkish in middle. Noseleaf moderately broad, not
much smaller noseleaf with pointed connecting quite covering muzzle; small notch in anterior leaf,
process; Thomas’s Horseshoe Bat, R. thomasi, has sella parallel-sided or slightly broader at base;
darker fur, short lancet with abrupt point in middle. connecting process rounded; lancet short and
Ecology and habitat Roosts in limestone caves, stubby, with a narrow point in the middle. Upper
sometimes in colonies of several hundred; has canines relatively small, only slightly longer than

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 premolars. Echolocation Males 76–77kHz, females finger bone) is 53–67% of length of metacarpal (first
83–85.5kHz (Laos). Similar species Chasen’s bone). Upper canine large, more than 30% longer
Horseshoe Bat, R. chaseni, has tall, well-developed than largest premolar. Echolocation 73–79kHz
lancet; Chinese and Rufous Horseshoe Bats, (Sri Lanka), 84kHz (India). Similar species Chinese
R. sinicus and R. rouxii, very similar, distinguished by Horseshoe Bat, R. sinicus, is similar, but with
larger size (on average), and more massive canines. differently shaped wing; Thomas’s Horseshoe Bat,
Ecology and habitat Roosts in limestone caves. R. thomasi, is smaller, with upper canine only slightly
Has been found in dry forests, including disturbed longer than largest premolar.
forests, around cave areas. Ecology and habitat In India, found mainly in wet
Distribution SE Asia: Myanmar, Thailand, Laos and forested areas, roosting in caves, tunnels, hollow
Vietnam. Also S China (Yunnan). trees, wells and temples. Forages on insects within
Status Least Concern. forest canopy, either on the wing or by hanging from
a perch and sallying out to catch passing prey.
CHINESE HORSESHOE BAT Distribution SE Asia: C Myanmar. Also India and
Rhinolophus sinicus PLATE 12 Sri Lanka.
Measurements FA 44–55, T 19–30, E 16–21. Status Least Concern, though poorly known in
Skull: gl 18.4–21.0, mt 7.0–8.4 region.
Identification Medium to moderately large
horseshoe bat with moderate-size noseleaf that PENINSULAR HORSESHOE BAT
does not cover muzzle. Lancet strongly concave at Rhinolophus robinsoni PLATE 12
sides (hastate), with tip usually short and stubby, Measurements FA 43–46, T 19–25, E 19–23.
but sometimes longer; sella approximately parallel- Wt 7.2–8.9g. Skull: gl 19–20, mt 6.7–7.5
sided, rounded at top; connecting process rounded. Identification Medium-small horseshoe bat with
Wing relatively long: on third digit (longest finger), rounded connecting process. Sella broad at base,
second phalanx (last finger bone) is 66–80% of getting slightly wider towards middle, then narrowing
length of the metacarpal (first bone). Upperparts abruptly just above middle, with squared-off top
dark brown, underside slightly paler; noseleaf dark. (Fig. 33d). Noseleaf moderately large, connecting
Upper canine large, nearly twice as long as largest process rounded, lancet broadly triangular. Well-
premolar. Similar species Rufous Horseshoe developed supplementary leaflet below horseshoe.
Bat, R. rouxii, has more rounded wing; Thomas’s Upperparts brown to reddish-brown, underparts
Horseshoe Bat, R. thomasi, has smaller upper only slightly paler. Noseleaf dark grey, slightly paler
canine, only slightly longer than largest premolar. in middle. Three grooves in lower lip. Taxonomic
Ecology and habitat In India, found mainly in areas notes A separate subspecies, R. robinsoni thaianus,
of higher elevations. has been described from N Thailand, but it is
Distribution SE Asia: N Myanmar and N Vietnam. not clear whether it actually represents the same
Also N India, Nepal and S China. species. Echolocation 64–68kHz (Peninsular
Status Least Concern. Malaysia). Similar species Other similar-sized
Rhinolophus in the region do not have stepwise
RUFOUS HORSESHOE BAT constriction in middle of sella; Pearson’s Horseshoe
Rhinolophus rouxii PLATE 12 Bat, R. pearsoni, is larger with longer noseleaf, long
Measurements FA 44–52, T 20–33, E 15–22. woolly hair, and a single groove in the lower lip.
Skull: gl 20.0–22.8, mt 7.9–9.2 Ecology and habitat Lowland and hill forest,
Identification Medium-sized horseshoe bat mainly in tall undisturbed forest. Has been found
with noseleaf that does not cover muzzle. Two roosting in crevices in rock boulders and in palm
colour phases, which may vary within individuals: leaves in forest understorey.
upperparts dark brown, base slightly paler, underside Distribution SE Asia: Peninsular Thailand and
slightly paler; or uniformly orange-rufous above and Malaysia.
below. Wing is relatively shorter and more rounded: Status Near Threatened; relatively uncommon;
on third digit (longest finger), second phalanx (last declining due to loss of lowland rainforest.

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 PEARSON’S HORSESHOE BAT any notch. Sella tall, broad at base, narrowing
Rhinolophus pearsonii PLATE 13 abruptly in middle (Fig. 33b). A single deep groove
Measurements FA 49–55, T 16–29, E 23–29. on lower lip. Echolocation 49–51kHz (Thailand).
Wt 10–16g. Skull: ccl 18.4–21.8, mt 8.6–9.6 Taxonomic notes Described in 2009 as a species
Identification Medium-large horseshoe bat with distinct from R. yunanensis, which is now thought
long woolly hair; hair is dark brown to greyish- to be restricted to China. DNA barcodes indicate
brown with pale tips, slightly paler on underparts. genetically close to R. chiewkweeae. Similar
Connecting process broadly rounded, joining sella species Pearson’s Horseshoe Bat, R. pearsonii, is
at tip without any notch. Sella tall, broad at similar but smaller, with darker fur, higher frequency
base, narrowing abruptly in middle (Fig. 33c). echolocation calls; Great Woolly Horseshoe Bat,
Noseleaf broad, covering muzzle, dark, with tall R. luctus, has lateral lappets on sella.
triangular lancet. A single deep groove on lower lip. Ecology and habitat Roosts in limestone caves.
Echolocation 65kHz (Thailand), 54–59kHz (Laos, Distribution SE Asia: Myanmar and Thailand.
Vietnam). Similar species Thai Horseshoe Bat, Status Not yet assessed; apparently much less
R. thailandensis, is larger with pale fur; Intermediate common than R. pearsonii.
Horseshoe Bat, R. affinis, has narrow sella, slightly
constricted in middle, shorter fur, and connecting CHIEW KWEE’S HORSESHOE BAT
process joining slightly below tip of sella, forming Rhinolophus chiewkweeae NOT ILLUSTRATED
a notch. Measurements FA 52–57, T 17–21, E 20–27.
Ecology and habitat Roosts in limestone caves. Skull: ccl 21.5–23.5, mt 9.9–10.3
Has been found in a variety of forest types at Identification Medium-large horseshoe bat, closely
altitudes ranging from 160m in Laos to 3,400m resembling Thai Horseshoe Bat, R. thailandensis.
in Nepal. Appears to use torpor regularly, and may Long woolly orange-brown fur, slightly paler below.
hibernate in northern parts of range during cold Noseleaf broad, covering muzzle, with tall triangular
seasons. lancet. Connecting process broadly rounded,
Distribution SE Asia: Myanmar, Thailand, Laos, joining sella at tip without any notch. Sella tall,
Cambodia and N Vietnam. Also N India, Nepal and broad for basal third, narrowing abruptly in middle
S China. and parallel sided to tip (similar to Fig. 33b). A
Status Least Concern. single deep groove on lower lip. Echolocation
53–55kHz. Taxonomic notes Described in
THAI HORSESHOE BAT 2005. Similar species Pearson’s Horseshoe Bat,
Rhinolophus thailandensis PLATE 13 R. pearsonii, averages smaller, while Thai Horseshoe
Measurements FA 56–61, T 18–24, E 26–32. Bat, R. thailandensis, is larger with lower frequency
Wt 20–22g. Skull: ccl 22.9–24.8, mt 10.4–11.4 echolocation calls.
Identification Large horseshoe bat with long woolly Ecology and habitat Has been caught in lowland,
hair; fur either light sandy-brown or dark greyish, hill and submontane rainforest at altitudes 100–
only slightly paler below. Noseleaf broad, covering 1,300m, including on Langkawi islands.
muzzle, with tall triangular lancet. Connecting Distribution SE Asia: peninsular Malaysia.
process broadly rounded, joining sella at tip without Status Not yet assessed.

Family HIPPOSIDERIDAE ROUNDLEAF BATS

The roundleaf bats (sometimes called roundleaf horseshoe bats or leafnose bats) vary in size from very
small species with a forearm of 34mm, weighing only 4g, to moderately large species with a forearm over
100mm, and weights of 60–70g. Like horseshoe bats (Rhinolophidae), they have an elaborate noseleaf
(Fig. 35). The anterior noseleaf is rounded and somewhat horseshoe-shaped (except in Coelops). The
median leaf is a low cushion-like structure expanded laterally, without a sella, while the posterior leaf is low
and rounded, usually divided by vertical septa into several pockets. The internarial septum varies from very
narrow to broadly expanded. Some species have additional lateral leaflets below the anterior noseleaf. The

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 Ear

Antitragus
Frontal sac

Vertical septa
Posterior noseleaf

Intermediate noseleaf
Gland
Median ridge

Internarial septum Lateral

supplementary
Anterior noseleaf leaflets

Fig. 35 Diagram of Hipposideros noseleaf, showing naming of parts.

number of lateral leaflets can be important for species identification. The ears vary from moderately small
to large with a low antitragus. The eyes are small.
Recent genetic and morphological studies indicate that several of the currently recognised species,
including Hipposideros bicolor, H. cf. cineraceus, H. pomona and H. cf. larvatus, are actually a complex of two
or more species. However, the species limits and diagnostic characters have been only partially determined,
and the appropriate names for each form have not yet been worked out, so most of these species complexes
are treated here as single species, although in some cases more than one is illustrated.
Echolocation calls are similar to, but generally much shorter than, those of Rhinolophus bats, with most
of the energy in a characteristic constant-frequency component that can help with species identification;
echolocation frequencies in the species accounts refer to the constant-frequency component.

Genus Hipposideros Typical horseshoe-shaped Medium to small Hipposideros species with a

anterior noseleaf (Fig. 35). Tail short to moderate, relatively small noseleaf and no lateral leaflets or
largely enclosed in interfemoral membrane. Skull one indistinct lateral leaflet.
generally similar to Rhinolophus, but rostrum with
paired swellings on top (Fig. 34c, d), although these GREATER BICOLOURED ROUNDLEAF BAT
are sometimes low. Anterior upper premolar variably Hipposideros bicolor PLATE 16
reduced; upper canines heavy but simple, without Measurements FA 45–49, T 29–34, E 17–20.
prominent supplementary cusps. Dental formula 1/2 Wt 7–10g
1/1 2/2 3/3. To assist with identification, species Identification Upperparts brown to grey-brown,
are grouped based on similar external characters, individual hairs with brown tip, white base;
although these do not necessarily indicate close underparts paler, usually buffy-white. Some
taxonomic relatedness. individuals bright orange. Ears moderately large and

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 rounded. Noseleaf pale pinkish-brown, simple and LARGE-EARED ROUNDLEAF BAT
small, with one indistinct lateral leaflet; internarial Hipposideros pomona PLATE 16
septum straight and only slightly widened at base. Measurements FA 38–45, T 26–36, E 19–26.
Echolocation 127–134kHz (peninsular Malaysia Wt 6–9g
and Thailand). Taxonomic notes Formerly included Identification Fur dark brown with pale base,
H. atrox, which is now considered a distinct species. underparts only slightly paler. Some individuals are
Similar species Lesser Bicoloured Roundleaf Bat, dark orange, with slightly paler orange underparts.
H. atrox, tends to be smaller (mean FA 44), with Noseleaf small and simple. Ears very large and
the noseleaf greyer and more angular, higher rounded. Taxonomic notes Genetic analyses
frequency echolocation call; Ashy Roundleaf Bat, suggest this may be a complex of species, but
H. cf. cineraceus, is smaller with shorter ears, boundaries among species and morphological
internarial septum distinctly swollen in the middle; differences have not yet been determined.
Large-eared Roundleaf Bat, H. pomona, has longer Echolocation 120–126kHz (Laos). Similar species
and broader ears (about 20% larger, on average), Greater Bicoloured Roundleaf Bat, H. bicolor, has
males with long, thin penis. slightly longer forearm, smaller ears; Ashy Roundleaf
Ecology and habitat Roosts mainly in caves or Bat, H. cf. cineraceus, is smaller with small ears and
tunnels, often in large colonies, but also in crevices noseleaf.
between large boulders and probably hollow trees. Ecology and habitat Roosts mainly in caves. Has
Forages in understorey of lowland rainforest. been found in a variety of forest types, including
Distribution SE Asia: peninsular Thailand and disturbed areas.
Malaysia. Also Sumatra, Java, Borneo, the Distribution SE Asia: Myanmar, Thailand, Laos,
Philippines, Sulawesi and smaller islands. Vietnam, Cambodia and N Peninsular Malaysia. Also
Status Least Concern, but has probably declined India and S China.
somewhat due to loss of mature forest. Status Least Concern.

LESSER BICOLOURED ROUNDLEAF BAT MALAY ROUNDLEAF BAT

Hipposideros atrox PLATE 16 Hipposideros nequam NOT ILLUSTRATED
Measurements FA 41–46, T 23–34, E 15–22. Measurements FA 46, E 19. Skull: mt 6.3
Wt 7–10g Identification Medium-small roundleaf bat, known
Identification Upperparts brown to grey-brown, only from a single damaged specimen, so its
individual hairs with brown tip, white base; under- characteristics are not well known. Apparently
parts paler, usually buffy-white. Similar to Greater similar to Greater Bicoloured Roundleaf Bat,
Bicoloured Roundleaf Bat, H. bicolor, but averages H. bicolor, with large rounded ears, but with larger
slightly smaller; has greyer noseleaf with more noseleaf (6 wide × 8 tall). Skull differs in having a
angular sides; more triangular internarial septum; thinner vomer (bone on top of the palate), and in
lacks lateral leaflet. Most easily distinguished by having a small anterior upper premolar inserted
higher frequency calls. Echolocation 138–144kHz. in toothrow (displaced to the side in H. bicolor).
Taxonomic notes Formerly considered part of Taxonomic notes May possibly be an aberrant
H. bicolor, but distinct genetically and morphologically specimen of one of the other small Hipposideros.
with different echolocation calls. Not yet confirmed Ecology and habitat Unknown.
that H. atrox is the appropriate name. Distribution SE Asia: known only from a single
Ecology and habitat Roosts mainly in caves or specimen from Selangor, Peninsular Malaysia.
tunnels, often in large colonies, but also in crevices Status Data deficient; unknown whether it is a valid
between large boulders and probably hollow trees. species.
Forages in understorey of lowland rainforest.
Distribution SE Asia: peninsular Thailand and ASHY ROUNDLEAF BAT
Malaysia. Also Sumatra. Hipposideros cf. cineraceus PLATE 16
Status Least Concern (as H. bicolor), but has Measurements FA 33–42, T 24–30, E 18.5–21.
probably declined due to loss of mature forest. Wt 4–5.5g. Skull: ccl 13.2–13.9

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 Identification Upperparts buffy-brown to greyish- in Thailand, 181–188kHz in Malaysia. Similar
brown; underparts pale brown to buffy-white. Ears species Ashy Roundleaf Bat, H. cf. cineraceus, has
large and rounded. Noseleaf simple, lacking lateral different-shaped internarial septum, shallow notch
leaflets; anterior noseleaf with only a slight notch. on anterior noseleaf.
Taxonomic notes Genetic and morphological Ecology and habitat Roosts in limestone caves.
analyses indicate this name is being used for at Distribution SE Asia: Central and S Thailand, and
least two species that are not closely related. A N Peninsular Malaysia.
larger form (mean FA 37.5), known only from Status Vulnerable due to small and declining
Sabah and Peninsular Malaysia, has internarial population size and fragmented habitat.
septum with a small, slightly raised, flattened oval
disc in the middle; a smaller form (mean FA 35.2) MYANMAR ROUNDLEAF BAT
found throughout the region has internarial septum Hipposideros einnaythu NOT ILLUSTRATED
straight-sided but slightly swollen vertically. It Measurements FA 39.5–40.5, T 24–29, E 16–17.
is not yet known which names are appropriate Skull: ccl 13.5–13.7, mt 5.1–5.2
for each. Echolocation Sabah and Peninsular Identification Noseleaf moderate-sized, largely dark
Malaysian form 137–138kHz (Sabah), 144kHz with slightly paler middle; one pair of narrow lateral
(Peninsular Malaysia); widespread form c.155kHz leaflets; internarial septum swollen in the middle,
(Peninsular Malaysia and Laos). Similar species narrower at the base and tip. Fur of upperparts dark
Thai Roundleaf Bat, H. halophyllus, has kidney- brown with pale base; underside paler brown. Ears
shaped lobes on internarial septum, notched broad and somewhat rounded. Taxonomic notes
anterior noseleaf; Myanmar Roundleaf Bat, Newly described in 2012. Similar species Dayak
H. einnaythu, has larger dark noseleaf with swollen Roundleaf Bat, H. dayacorum, is similar in size, but
internarial septum; Least Roundleaf Bat, H. doriae, has triangular internarial septum, more triangular
lacks supporting septa on posterior noseleaf; ears, lacks lateral leaflets. Ashy Roundleaf Bat,
Bicoloured Roundleaf Bat, H. bicolor, is larger with H. cf. cineraceus, is smaller with pink noseleaf.
straight internarial septum. Ecology and habitat Poorly known; has been found
Ecology and habitat Roosts in caves, usually in roosting in village houses.
small to moderate-size colonies. Has also been Distribution SE Asia: known only from S Myanmar.
found roosting in culverts. Forages in forest of many Status Not yet assessed.
types, where usually found in small numbers.
Distribution SE Asia: Myanmar, Thailand, Laos, DAYAK ROUNDLEAF BAT
Vietnam, Cambodia and Peninsular Malaysia. Also Hipposideros dyacorum PLATE 16
Pakistan, N India, China, Sumatra, Borneo and Measurements FA 38–43, T 19–24, E 15–18.
various small islands, although some may prove to Wt 5–8g
be different species. Identification Fur uniformly dark brown, often
Status Least Concern, but likely to be declining due with slightly paler tips. Noseleaf small and
to loss of forest. simple without lateral leaflets, usually strongly
pigmented dark grey-brown. Internarial septum
THAI ROUNDLEAF BAT slightly swollen at base, narrow in middle. Ears
Hipposideros halophyllus PLATE 17 broadly triangular and pointed. Echolocation
Measurements FA 35–38, T 25–28, E 16. 167–170kHz (Sabah). Similar species Greater
Skull: ccl 12.7–13.1, mt 4.7–4.8 Bicoloured Roundleaf Bat, H. bicolor, tends to
Identification Noseleaf small, simple with no lateral have more pinkish noseleaf, fur with brown tip
leaflets; internarial septum with swollen kidney- and white base, and larger rounded ears; Fawn
shaped lobes on each side; anterior noseleaf with a Roundleaf Bat, H. cervinus, is larger, with a wider
deep notch. Fur of upperparts mid-brown with pale noseleaf and two lateral leaflets.
base; underside whitish. Noseleaf mostly pink, with Ecology and habitat Has been found roosting in
darker lobes on internarial septum and sometimes caves, under rocks and in hollow trees. Forages in
posterior noseleaf. Echolocation 156–188kHz the understorey of tall forest.

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 Distribution SE Asia: peninsular Thailand and pair of lateral leaflets and does not overlap in range.
Peninsular Malaysia. Also Borneo. Ecology and habitat Occurs in lowland dipterocarp
Status Least Concern, although rarely recorded on forest. Roosts in fallen hollow trees; also found in
the mainland. Likely to have declined substantially culverts and drainage pipes.
owing to loss of lowland rainforest. Distribution SE Asia: peninsular Thailand and
Malaysia. Also Borneo.
LEAST ROUNDLEAF BAT Status Vulnerable, owing to loss of lowland
Hipposideros doriae PLATE 17 rainforest.
Measurements FA 34–37. Wt 4–6g.
Skull: ccl 12.9, mt 4.4–5.1 SMALL-DISC ROUNDLEAF BAT
Identification Upperparts dark greyish-brown to Hipposideros orbiculus PLATE 15
brown, underparts paler. Noseleaf very small and Measurements FA 46–49, T 26–34, E 20–23.5.
simple with no lateral leaflets; anterior horseshoe Wt 9.3–10.3g. Skull: ccl 16.1–16.9, mt 6.4–6.6
with small central notch; posterior noseleaf lacks Identification Fur dark chocolate-brown; ears,
supporting septa; internarial septum slightly swollen. noseleaf and wing membranes dark. Ears large,
Ears and noseleaf pigmented grey. Ears broad, rounded. Noseleaf large, covering muzzle; no lateral
rounded, bluntly pointed at top. Echolocation 195– leaflets; internarial septum expanded into a medium-
200kHz (Peninsular Malaysia). Taxonomic notes size disc (1.9–2.2mm across). Echolocation 80kHz
Formerly called H. sabanus. Similar species All (Peninsular Malaysia). Similar species Ridley’s
other small roundleaf bats in region have posterior Roundleaf Bat, H. ridleyi, has larger internarial disc,
noseleaf divided into four compartments by three larger ears, lower frequency echolocation; Thai
supporting septa. Roundleaf Bat, H. halophyllus, is much smaller with
Ecology and habitat Found in relatively undisturbed small noseleaf.
lowland and hill forest up to 1,500m elevation, Ecology and habitat Has been found roosting in
usually at low densities. Roosting sites unknown. drainage pipes. Recorded from a rubber plantation
Distribution SE Asia: Peninsular Malaysia. Also in primary forest (Sumatra), peatswamp forest
Borneo and Sumatra. (Peninsular Malaysia).
Status Near Threatened, due to loss of forest Distribution SE Asia: Peninsular Malaysia. Also
habitat. Sumatra.
Status Vulnerable, owing to limited range, low
Medium-sized Hipposideros species with an densities and loss of forest.
enlarged noseleaf that covers muzzle. Lateral
leaflets lacking or indistinct. ANNAMITE ROUNDLEAF BAT
Hipposideros rotalis PLATE 15
RIDLEY’S ROUNDLEAF BAT Measurements FA 45–49, T 32–39, E 24–27,
Hipposideros ridleyi PLATE 15 Wt 7.5–11g. Skull: ccl 16.8–17.0, mt 6.4–6.6
Measurements FA 47–51, T 25–29, E 25–30. Identification Upperparts brown, individual hairs
Wt 7–12g. Skull: ccl 17.0–17.5, mt 6.6–6.8 with white base, brown middle and pale tip;
Identification Fur uniformly dark brown. Ears, underparts paler. Noseleaf broad, covering muzzle;
noseleaf and wing membranes dark grey. Ears enlarged internarial disc (width 2.3–2.6mm);
very large and rounded. Noseleaf large, covering anterior noseleaf angular with deep indentation in
whole muzzle; no lateral leaflets; internarial septum the middle. Echolocation 68.5–71.5kHz (Laos).
expanded into large flat disc (2.8–3.1 across), which Similar species Ridley’s Roundleaf Bat, H. ridleyi,
largely obscures nostrils. Echolocation 65–67kHz is darker, lacks lateral leaflets and has less indented
(Sabah), 61–62kHz (Peninsular Malaysia). Similar anterior noseleaf; Khaokhouay Roundleaf Bat,
species Small-disc Roundleaf Bat, H. orbiculus, H. khaokhouayensis, has smaller internarial disc.
has slightly smaller noseleaf, shorter ears, smaller Ecology and habitat Has been found near
internarial disc that does not completely cover limestone caves in disturbed evergreen forest, and
nostrils; Annamite Roundleaf Bat, H. rotalis, has one mixed dry dipterocarp and evergreen forest.

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 Distribution SE Asia: known only from C Laos. Distribution SE Asia: Peninsular Malaysia. Also
Status Least Concern, although likely to be declining Sumatra, Java, Borneo, the Philippines and
due to loss of habitat within its limited range. E Indonesia through New Guinea to Australia.
Status Least Concern.
KHAOKHOUAY ROUNDLEAF BAT
Hipposideros khaokhouayensis PLATE 15 CANTOR’S ROUNDLEAF BAT
Measurements FA 46–49, T 35–37.5, E 24–26. Hipposideros galeritus PLATE 15
Wt 7.7–9.6g. Skull: ccl 16.0–17.6, mt 6.4–6.7 Measurements FA 45–50, T 32–42, E 14–17.
Identification Upperparts mid-brown, individual Wt 6.3–10.0g
hairs with white base, brown middle and pale tip; Identification Fur usually dark grey-brown,
underparts paler. Noseleaf broad, covering muzzle; occasionally with a reddish tinge. Noseleaf pinkish-
moderate-sized internarial disc (width 1.5–1.8mm); grey. Ears broad, rounded at base, triangular at tip.
narrow notch in anterior noseleaf; no lateral leaflets. Noseleaf with two lateral leaflets; median noseleaf
Echolocation 87–91kHz (Laos), 94.5 (Vietnam). broader than posterior noseleaf. Echolocation
Similar species Annamite Roundleaf Bat, 109–117kHz (Sabah), 85–93kHz (Peninsular
H. rotalis, has larger internarial disc, well-developed Malaysia), 107–110kHz (Laos). Taxonomic notes
lateral leaflets, lower frequency echolocation calls; Consists of several disjunct populations which may
Small-disc Roundleaf Bat, H. orbiculus, has dark prove to be distinct species. Similar species Fawn
noseleaf with more rounded anterior noseleaf, Roundleaf Bat, H. cervinus, has median noseleaf
shorter tail, lower frequency echolocation. narrower than posterior noseleaf, more triangular
Ecology and habitat Has been found in patches ears, shorter tail.
of intact and disturbed evergreen forest, both near Ecology and habitat Roosts in caves, often in small
limestone caves and far from known caves. groups with the Fawn Roundleaf Bat, H. cervinus,
Distribution SE Asia: C Laos, Cat Ba in Vietnam. but sometimes in colonies of several hundred. Also
Status Vulnerable, due to very limited range and found near large boulders. Forages in a variety of
ongoing loss of habitat. forest types from lowland rainforest to dry disturbed
areas.
Medium to large Hipposideros species with plain Distribution SE Asia: S Laos, S Vietnam, Cambodia,
noseleaf and two or three distinct lateral leaflets. S Thailand and Peninsular Malaysia. Also India, Sri
Lanka, Borneo and Java.
FAWN ROUNDLEAF BAT Status Least Concern.
Hipposideros cervinus PLATE 15
Measurements FA 44–52, T 21–28, E 14–17. INTERMEDIATE ROUNDLEAF BAT
Wt 8–12g Hipposideros cf. larvatus PLATE 14
Identification Fur colour varies from grey-brown Measurements FA 51–67, T 32–35, E 21–25.
or yellowish-brown to bright red-brown or orange; Wt 15–23g
underparts somewhat paler. Noseleaf greyish-pink. Identification Medium-sized roundleaf bat with
Ears broadly triangular. Noseleaf simple with two three lateral leaflets on noseleaf. Upperparts dark
lateral leaflets; median noseleaf narrower than grey-brown to reddish-brown; underparts slightly
posterior noseleaf. Echolocation 118–124kHz paler. Noseleaf, ears and wing membranes brown.
(Sabah), 122–127kHz (Peninsular Malaysia). Similar Taxonomic notes Recent studies indicate that
species Cantor’s Roundleaf Bat, H. galeritus, has this is a complex of several species varying in
median noseleaf broader than posterior noseleaf, size, echolocation frequency and genetics; multiple
ears broader and more rounded, tail longer; Dayak species have been found sympatrically in India,
Roundleaf Bat, H. dyacorum, and other species with Laos and Vietnam, but the appropriate names for
a small noseleaf do not have lateral leaflets. each have not yet been worked out. True H. larvatus
Ecology and habitat Usually roosts in caves, may be restricted to Java. H. grandis has been used
sometimes in very large colonies (up to 300,000 in for some mainland forms, but H. leptophylla may be
Sabah). Feeds in the forest understorey. a senior synonym. Because diagnostic characters

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 and correct names are uncertain, they are maternity colonies of females and young. Forages in
treated together here. Echolocation 97–100kHz various forested habitats, including highly disturbed
(Peninsular Malaysia); large form 93–97kHz, areas.
medium form 86–89kHz, small form 99–102kHz Distribution SE Asia: Myanmar, Thailand, S Laos,
(Laos). Similar species Diadem Roundleaf Bat, S Vietnam, Cambodia and Peninsular Malaysia. Also
H. diadema, is larger, with buff or white shoulder Sumatra, Java, Borneo and the Philippines through
marks; Halong Roundleaf Bat, H. alongensis, and to the Solomon Islands and Australia.
Indian Roundleaf Bat, H. lankadiva, are similar but Status Least Concern.
substantially larger.
Ecology and habitat Roosts in caves, rock BOONSONG’S ROUNDLEAF BAT
crevices, temples and old mines. Cave colonies Hipposideros lekaguli PLATE 15
may be very large. Measurements FA 71–79, T 44–54, E 26–33
Distribution SE Asia: throughout Myanmar, Identification Upperparts pale greyish or brownish-
Thailand, Laos, Vietnam, Cambodia and Peninsular white; underparts and throat dull creamy white. Ears
Malaysia. Also Bangladesh to S China, Sumatra, triangular, large and broad. Anterior noseleaf large,
Java, Borneo, Sumba and adjacent small islands. almost covering muzzle with no notch in middle,
Status Least Concern, although some species in and three well-developed lateral leaflets. Internarial
this complex may be of concern. septum slightly swollen; raised flaps on either
side of nostrils. Intermediate leaf thickened with
Large Hipposideros species generally with more triangular wedge protruding in middle. Posterior leaf
complex noseleaf, often with distinctive characters. formed into three backward-facing pockets, median
pocket smaller than lateral pockets. Echolocation
DIADEM ROUNDLEAF BAT 45–46kHz (Thailand). Similar species No other
Hipposideros diadema PLATE 15 similar-size roundleaf bat has enlarged, backward-
Measurements FA 76–87, T 53, E 27–18.5. facing pockets in posterior leaf, with triangular
Wt 30–47g lappet in intermediate noseleaf.
Identification Fur of upperparts dark brown with Ecology and habitat Roosts in caves. Forages in
pale base, white patches on the shoulders and forested habitats, including very disturbed areas.
sides; underparts greyish-white. Adult females Distribution SE Asia: S Thailand and Peninsular
often with orange or orange-buff replacing the Malaysia. Also reported in the Philippines, although
white. Juveniles dark grey and white. Moulting there is uncertainty whether those are actually the
individuals can be a mixture of brown, grey, orange same species.
and white. Noseleaf with three or four lateral Status Near Threatened, due to limited range and
leaflets; posterior noseleaf large and rounded. population declines as a result of roost disturbance
Echolocation 59–62kHz (Peninsular Malaysia), as well as loss of habitat.
64–66kHz (Sabah). Similar species No other
Asian Hipposideros has pale shoulder patches; INDIAN ROUNDLEAF BAT
Great Roundleaf Bat, H. armiger, and Pendlebury’s Hipposideros lankadiva NOT ILLUSTRATED
Roundleaf Bat, H. pendleburyi, have posterior Measurements FA 89–94, T 44–54, E 24–28.
noseleaf narrower than anterior; Boonsong’s Skull: ccl 29.1–31.2, mt 13.5–14.6
Roundleaf Bat, H. lekaguli, has thickened posterior Identification Very large roundleaf bat, superficially
noseleaf with deep inverted pockets; triangular resembling an enlarged H. cf. larvatus with similar
protuberance on median leaf. noseleaf shape and three well-developed lateral
Ecology and habitat Usually roosts in large colonies leaflets. Posterior noseleaf wider than anterior
in caves, often mixed with other species; has been noseleaf; intermediate noseleaf narrower than both.
recorded in hollows in trees, or roosting alone under Fur fulvous brown above, slightly paler below.
palms. Feeds by hanging from a perch and waiting Echolocation 69–71kHz (Myanmar). Taxonomic
for prey to pass by. May select several perches over notes Subspecies in Myanmar, H. l. gyi, was first
one square kilometre during a night. Forms large described in 2015 and also occurs in NE India; it is

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 much larger than form in peninsular India. Similar 2012. Similar species Pendlebury’s Roundleaf
species Great Roundleaf Bat, H. armiger, is similar Bat, H. pendleburyi, is similar but smaller, and does
size, but with narrow, thickened posterior noseleaf; not overlap in range.
swelling behind noseleaf, especially in males. Other Ecology and habitat Roosts in limestone caves.
species with three lateral leaflets are much smaller. Distribution SE Asia: known only from Vietnam.
Ecology and habitat In Myanmar, a colony of Status Not yet assessed.
1,000 individuals was found in a limestone cave
surrounded by disturbed evergreen forest at about PENDLEBURY’S ROUNDLEAF BAT
250m altitude. Hipposideros pendleburyi PLATE 14
Distribution SE Asia: Myanmar. Also India, Measurements FA 75–81. Wt 24–40g.
Bangladesh, Nepal, Sri Lanka. Skull: ccl 22.3–24.3
Status Least Concern, though known from only one Identification Large roundleaf bat with dark brown
location in region. fur. Anterior noseleaf does not cover muzzle;
four lateral leaflets, the outer one very small;
GREAT ROUNDLEAF BAT posterior noseleaf thickened and narrower than
Hipposideros armiger PLATE 14 median noseleaf with weakly developed swellings
Measurements FA 85–103, T 54–69, E 25–35. on each side behind it. Taxonomic notes
Wt 44–67g (males average larger than females). Formerly considered same species as H. turpis,
Skull: ccl 27.1–29.8, mt 11.7–13.1 which is now considered restricted to Ryukyu
Identification One of the largest roundleaf bats Islands, Japan. Echolocation 85–88kHz. Similar
in SE Asia. Fur generally dark brown all over, species Great Roundleaf Bat, H. armiger, has
sometimes paler beneath. Anterior horseshoe similar-shaped noseleaf, but is larger, with more
relatively small, not covering muzzle, fairly straight developed swellings behind posterior noseleaf;
across; four lateral leaflets, outer ones often very Diadem Roundleaf Bat, H. diadema, has white or
small; median noseleaf swollen; posterior noseleaf buff shoulder patches, posterior noseleaf broadly
thick, but narrower than anterior noseleaf. In males, rounded and wide; Shield-faced Roundleaf Bat,
face behind and around noseleaf may be greatly H. lylei, has posterior noseleaf as wide as anterior,
thickened and swollen. Echolocation 69–71kHz with enlarged lobes behind it.
(Laos), 65–69kHz (Vietnam). Similar species Ecology and habitat Roosts in limestone caves
Pendlebury’s Roundleaf Bat, H. pendleburyi, is in karst areas; forages in forest, both intact and
smaller with less swelling behind noseleaf. Griffin’s disturbed.
Roundleaf Bat, H. griffini, is slightly smaller. Distribution SE Asia: known only from a limited
Ecology and habitat Roosts in caves; may fly low area in peninsular Thailand.
around trees to hunt, but also flies very high in open Status Not yet assessed. Probably at risk due
sky, possibly commuting to foraging sites. to restricted range, small known population and
Distribution SE Asia: Myanmar, Thailand, Vietnam, ongoing loss of forest habitats.
Laos, Cambodia and Peninsular Malaysia. Also
N India, Nepal, S China and Taiwan. HALONG ROUNDLEAF BAT
Status Least Concern. Hipposideros alongensis NOT ILLUSTRATED
Measurements H. a. alongensis FA 66–72.
GRIFFIN’S ROUNDLEAF BAT Wt 20–30g. Skull: ccl 22.4–23.2, mt 9.8–10.1.
Hipposideros griffini NOT ILLUSTRATED H. a. sungi FA 68–76. Wt 22–36g.
Measurements FA 83–90, E 28–30. Skull: ccl 22.8–24.4, mt 9.8–10.5
Skull: ccl 25.5–26.5, mt 11.3–11.5 Identification Medium-large roundleaf bat with
Identification Closely resembles Great Roundleaf dark brown fur, slightly paler below. Superficially
Bat, Hipposideros armiger, but averages smaller, resembles H. cf. larvatus complex, but larger.
with higher frequency echolocation calls and Anterior noseleaf does not cover muzzle; three
genetically distinct. Echolocation 75–79kHz large, well-developed lateral leaflets; posterior
(Vietnam). Taxonomic notes Newly described in noseleaf similar in width to anterior noseleaf and

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 broader than median noseleaf. Mainland subspecies SHIELD-NOSED ROUNDLEAF BAT
(H. a. sungi) is larger than nominate subspecies Hipposideros scutinares PLATE 14
H. a. alongensis, which is found only on Con Measurements FA 78–83, T 50–59, E 28–30.
Son Island. Echolocation 70–75kHz (H. a. sungi ); Wt 38–42g. Skull: ccl 26.5–27.9, mt 11.5–12.3
78.5–80.0kHz (H. a. alongensis). Taxonomic Identification Large roundleaf bat with enlarged
notes Formerly considered same species as lobes behind posterior leaf. Anterior noseleaf broad
H. turpis, but recent research indicates it is not (width 9.5–10.5), rounded, with a conspicuous
closely related. Similar species Intermediate deep notch in the middle and a shallow notch on
Roundleaf Bat, H. cf. larvatus, is smaller; Shield- each side; two lateral leaflets; posterior leaf similar
faced Roundleaf Bat, H. lylei, averages larger, with in width to anterior, and joined on sides to anterior
enlarged lobes behind posterior noseleaf, only two leaf; posterior lobes enlarged in males (7–14
lateral leaflets. tall), smaller in females (3–4), separated by deep
Ecology and habitat Roosts in limestone caves; notch. Echolocation 61–64kHz (Laos). Similar
forages in surrounding forest. species Shield-faced Roundleaf Bat, H. lylei, has
Distribution SE Asia: N. Vietnam. narrower noseleaf with (in males) larger posterior
Status Not yet assessed. Probably at risk due to lobes, smaller skull, higher frequency echolocation;
limited range, forest loss and cave disturbance. Swinhoe’s Roundleaf Bat, H. swinhoei, has longer
forearm, larger skull, narrower anterior noseleaf
SHIELD-FACED ROUNDLEAF BAT with a single shallow central notch, posterior leaf
Hipposideros lylei PLATE 14 not joined to anterior noseleaf.
Measurements FA 73–84, T 48–55, E 30. Ecology and habitat Roosts in limestone caves;
Wt 35–52g. Skull: ccl 24.1–26.5, mt 10.4–11.9 has been found in disturbed evergreen forest near
Identification Large roundleaf bat with prominent limestone.
enlarged lobes behind posterior noseleaf. Anterior Distribution SE Asia: Laos and C Vietnam.
noseleaf moderately broad (width 8.8–9.2), Status Vulnerable, owing to limited range, small
rounded, with a deep notch in the middle and population size and loss of forest.
shallow notches on either side; two lateral leaflets;
posterior noseleaf similar in width to anterior leaf, SWINHOE’S ROUNDLEAF BAT
joined on sides to anterior leaf; posterior lobes of Hipposideros swinhoei PLATE 14
male greatly enlarged, pointed at tip, joined at base Measurements FA 81–89. Skull: ccl 27.8–30.0,
and extending around sides of noseleaf; lobes of mt 12.1–12.8
female small, partly hidden in fur. Ears large and Identification Large roundleaf bat with enlarged
triangular. Fur of upperparts pale grey to light lobes behind posterior leaf. Anterior leaf moderately
brown; underparts similar or paler. Echolocation broad (width 8.3–9.7), rounded, with single shallow
70–75kHz (Thailand). Similar species Shield- notch in front margin; two lateral leaflets; posterior
nosed Roundleaf Bat, H. scutinares, has broader leaf similar in width to anterior leaf, not connected on
noseleaf with less-developed posterior lobes, lower sides; posterior lobes larger in males than females.
frequency echolocation; Swinhoe’s Roundleaf Bat, Echolocation 58–62kHz (China and Vietnam).
H. swinhoei, has larger skull, anterior noseleaf not Taxonomic notes Formerly called H. pratti, but
connected to posterior leaf. recent research shows that H. swinhoei is the
Ecology and habitat Roosts in limestone caves; appropriate name for the species. Similar species
has been found in both understorey and canopy Shield-faced and Shield-nosed Roundleaf Bats,
of lowland primary forest in Malaysia. In Thailand H. lylei and H. scutinares, have shorter forearm and
reported from both disturbed and fragmented forests. smaller skull, posterior noseleaf joined on sides to
Distribution SE Asia: Myanmar, Thailand, anterior leaf, anterior leaf with three notches.
Peninsular Malaysia and N Vietnam; probably Ecology and habitat Roosts in limestone caves.
N Laos. Also S China. Distribution SE Asia: Vietnam. Also S China.
Status Least Concern, but colonies are small and Status Least Concern (as H. pratti), though status is
affected by cave disturbance and hunting. poorly known in region.

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 Genus Coelops Very small bats. Anterior noseleaf represent more than one species. Similar species
divided in middle and expanded to sides. Lateral Malaysian Tailless Roundleaf Bat, C. robinsoni, has
leaflets large and expanded forwards. No visible tail, broad, rounded lobes on lateral leaflets.
although interfemoral membrane is well developed. Ecology and habitat Roosts in caves or hollow
Skull and dentition as Hipposideros, but rostral trees, sometimes in small groups. Forages in forests.
swellings generally low; upper canines have well- Distribution SE Asia: Myanmar, Thailand, Laos,
developed secondary cusp; lower incisors separated Vietnam, Cambodia and Peninsular Malaysia. Also
from canines by a distinct gap. Dental formula 1/2 E India, S China, Taiwan, Sumatra, Java and Bali.
1/1 2/2 3/3. Status Least Concern, though apparently occurs at
very low density throughout its range.
MALAYSIAN TAILLESS ROUNDLEAF BAT
Coelops robinsoni PLATE 17 Genus Paracoelops This genus with the associated
Measurements FA 34–37, T none, E 12–14. species Paracoelops megalotis was included in the
Wt 3.5–5.0g. Skull: ccl 12.7, mt 4.7 first edition of the book, but recent research has
Identification Fur long and soft, brown to dark shown that the genus and species are not valid. The
brown with paler tips; underparts brownish to specimen on which they were based, from Vinh in
slightly grey. Ears rounded, lacking supporting S. Vietnam, is actually a poorly prepared specimen
ridges. Narrow interfemoral membrane with no of Hipposideros closely resembling H. pomona, but
visible tail. Anterior margin of noseleaf deeply with the tail missing and a damaged noseleaf.
notched, forming two separate lobes; one pair of
lateral leaflets expanded forwards to form wide, Genus Aselliscus Posterior noseleaf raised into
rounded lobes under the anterior noseleaf; posterior three points; anterior noseleaf with two lateral
noseleaf low with a small median lobe. Similar leaflets, upper ones joining in the middle; median
species Asian Tailless Roundleaf Bat, C. frithii, has noseleaf contiguous with internarial septum.
lower lobes of noseleaf elongated and narrow, and Tail relatively short, protruding slightly beyond
is usually larger. interfemoral membrane. Skull and dentition similar
Ecology and habitat Known roost sites include one to Hipposideros, but braincase low, rostral swellings
in a cave, and one in hollow buttress of a large tree. relatively low, canines small and weak. Dental
Has been found in understorey of mature lowland formula 1/2 1/1 2/2 3/3.
rainforest, but relatively rarely recorded.
Distribution SE Asia: peninsular Thailand and COMMON TRIDENT BAT
Malaysia. Also Borneo. Aselliscus stoliczkanus PLATE 17
Status Vulnerable, due to population declines as a Measurements FA 39–45, T 33–44, E 10–13.
result of loss of lowland rainforest. Wt 4.5–6.5g. Skull: gl 14.5–15.2, mt 4.9–5.3,
upper canine 1.6–1.8.
ASIAN TAILLESS ROUNDLEAF BAT Identification Small roundleaf bat with posterior
Coelops frithii PLATE 17 noseleaf raised into three points; median noseleaf
Measurements FA 34–44, T none, E 14–16. merges with internarial septum to form nearly
Wt 3.0–7.0g. Skull: mt 5.7 triangular pad in middle of nose; two lateral leaflets,
Identification Fur long and soft, brown to blackish upper ones joined in front of face. Ears short,
with paler tips; underparts slightly greyer. Ears broad with point at top. Upperparts brown, fur with
rounded and lacking supporting ridges. Narrow white base, underparts whitish. Adult males may
interfemoral membrane with no visible tail. Anterior have tuft of longer yellowish fur on either side of
margin of noseleaf deeply notched, forming two chest. Echolocation 124–130kHz (Laos, Vietnam,
lobes; one pair of lateral leaflets extending from Thailand, Myanmar). Taxonomic notes A population
underneath them to form narrow, elongated lobes; in N Vietnam was recently differentiated as a distinct
posterior noseleaf low with a small median lobe. species, A. dongbacana (see page 248). Genetic
Taxonomic notes Specimens referred to this studies suggest there may be additional species not
species exhibit a very large size range, and may yet recognised. Similar species Similar-sized small

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 Hipposideros lack points on posterior noseleaf, have Skull: gl 14.9–15.5, mt 5.3–5.5, upper canine
larger ears. Dong Bac Trident Bat, A. dongbacana, 1.9–2.1.
currently known only from N Vietnam, has relatively Identification Very similar to Common Trident
longer canines. Bat, A. stoliczkanus, but with longer canines.
Ecology and habitat Roosts in limestone caves. Echolocation 125–130kHz (Vietnam). Taxonomic
Forages in forest, including disturbed areas. notes Described in 2015 based on genetic and
Distribution Myanmar, Thailand, Laos, Vietnam morphological differences from A. stoliczkanus.
and N Malaysia. Also S China. Ecology and habitat Recorded around limestone
Status Least Concern, although susceptible to caves, including disturbed areas.
disturbance of caves. Distribution Known only from N Vietnam.
Status Not yet assessed. Possibly at risk due to
DONG BAC TRIDENT BAT disturbance, damage to caves and loss of habitat
Aselliscus dongbacana NOT ILLUSTRATED within limited range.
Measurements FA 41–44, T 39, E 12. Wt 4.5g.

Family VESPERTILIONIDAE COMMON BATS

The largest, most diverse and most widespread family of bats, occurring on every continent except
Antarctica. The nose is simple, without any noseleaf, at least in all South-east Asian species. The ears
vary from relatively small to very large, but always with a well-developed tragus. The shape of the ear
and tragus can be very helpful for identification, especially for distinguishing the different genera. The
tail is long, and completely or nearly completely enclosed in the interfemoral membrane, although the
terminal vertebra sometimes protrudes. The skull and dentition vary among genera and species, and are
often important for identification (Figs 36–38). The ancestral condition for these bats is believed to have
had two upper and three lower incisors, and three upper and lower premolars, a condition still found in
Kerivoula and most Myotis, although the middle premolar is often reduced and sometimes missing
in Myotis. In most other genera, there has been a trend towards shortening of the jaws and reduction in
the number of teeth.
South-east Asian species are grouped into four subfamilies containing 22 genera. The bent-winged bats
were formerly considered part of this family, but are now considered to be their own family, Miniopteridae.
The Myotinae are now considered a distinct subfamily separate from the Vespertilioninae.
When trying to identify vespertilionid bats, it is very helpful first to determine to which subfamily and
genus the species belongs, by comparing it with the colour plates as well as the genus descriptions. In
most cases, the genus can be determined by a combination of external features, such as ear and wing
shape, as well as dental characters. With care, it is possible to examine the teeth in a live bat by gently
opening the mouth with a toothpick or the edge of a glove, though a magnifying glass is often necessary
to see the teeth clearly. Overall skull shape can usually also be determined in a live bat, because it is
reflected in the head shape. When handling bats, it is often possible to smooth the fur with a finger to
determine the shape of the head.
Once the genus has been determined, species can be identified by a combination of size, fur colour,
dental characters and other characters as described in each account. In a few groups, especially the
Pipistrellus, reliable external identification characters are poorly known, and it may not be possible to identify
some species except through detailed examination of museum specimens. In these cases, for the benefit
of museum workers, the text also gives some information on selected skull and baculum identification
characters for these bats.

Subfamily MYOTINAE
Although formerly considered part of the Vespertilioninae, this is now considered a distinct subfamily.
Most species in this family are currently placed in the genus Myotis. Recent genetic analyses suggest that
Eudiscopus should also be in this subfamily.

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 a b

c d

e f

g h

i j

0 5

Fig. 36 Skull profiles of selected vespertilionid bats and Miniopterus: Myotis annamiticus (a), Myotis rosseti (b),
Eudiscopus denticulus (c), Tylonycteris fulvida (d), Thainycteris aureocollaris (e), Hesperoptenus blanfordi (f),
Miniopterus magnater (g), Glischropus bucephalus (h), Murina cyclotis (i) and Kerivoula picta (j).

Ears moderately long and triangular at the tip; tragus tapered, bluntly pointed and bent slightly forwards
(see colour plates). Skull relatively unspecialised with variably elevated braincase, narrow rostrum (Fig.
36a, b). Usually three upper and three lower premolars; middle premolar small and displaced inwards in
many species, lacking in three species (Fig. 37); anterior premolar always well developed and not displaced
inwards, unlike Pipistrellus. Dental formula 2/3 1/1 2–3/2–3 3/3.

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 The genus has traditionally been divided into several subgenera, but recent genetic analyses indicate
these do not represent evolutionary groupings. Nevertheless, the characters separating them can be useful
for identification, so they are considered here as species groups. Recent genetic analyses indicate that
several additional species may occur in some groups. Some of these have been recently described, but in
other cases species limits and diagnostic characters have not yet been determined.

‘Myotis’ group Moderately large Myotis with feet have smaller feet, shorter ears, middle upper
slightly enlarged; ears relatively long; braincase premolar greatly reduced and intruded in toothrow;
low, skull profile gradually sloping from rostrum to Hairy-faced Myotis, M. annectans, lacks middle
back of skull; middle upper and lower premolars upper and lower premolars.
somewhat reduced, but usually still within toothrow; Ecology and habitat Only specimen from region
anterior two pairs of lower incisors with four small was found in a limestone cave near a stream.
lobes. Distribution SE Asia: N Thailand. Also S China.
Status Least Concern.
CHINESE MYOTIS
Myotis chinensis PLATE 19 ‘Chrysopteron’ group Moderately large Myotis
Measurements FA 65–69, T 53–58, HF 16–18, with distinctive woolly yellow to red-orange fur
E 20–23. Wt 25–30g. Skull: ccl 20.5–22.2, with black and orange pattern on wings. A
mt 9.5–9.9 recent study showed there are several more
Identification Largest Myotis in region. Upperparts species in this group than previously recognised,
dark greyish-brown, hairs with blackish base; of which three occur in South-east Asia. Ears
underparts paler, with grey tip to hairs. Ears long moderate-sized, typical Myotis shape, broadened
and slender, tragus tapered and angled slightly in middle and tapering bluntly to top with moderate
forwards. Middle premolars slightly more than half antitragal fold extending halfway up ear; tragus
size of anterior premolars and in toothrow. Similar relatively long, bluntly pointed and straight, not
species All other Myotis in region are smaller; other bent forwards; feet not especially enlarged; wing
large vespertilionids have different-shaped ears and membrane attached at base of toes. Anterior two
fewer teeth. pairs of lower incisors with four small lobes.
Ecology and habitat Has been recorded in or near
limestone caves with rivers nearby. Altitudes from BLACK-AND-ORANGE WOOLLY MYOTIS
50–1,000m. Moderately common in some areas. Myotis rufoniger PLATE 18
Distribution SE Asia: Myanmar, Thailand and Measurements FA 45–56, T 52, HF 11, E 19.
Vietnam. Also China. Wt 12g. Skull: ccl 15.3–17.7, mt 6.9–8.1
Status Least Concern. Identification Moderately large Myotis with orange
to orange-red body and head. Hairs of upperparts
SZECHUAN MYOTIS with dark base, pale middle, darkening to orange
Myotis altarium PLATE 19 or orange-red at the end, with small black tip;
Measurements FA 43–46, HF 11–12, E 22–24. underside paler base with orange to orange-red
Skull: gl 15.5–16.0 tip, lacking black. Black rim to ears, black thumb
Identification Upperparts light brown with long and underside of feet. Tail membrane orange with
hairs, underparts paler, individual hairs with black edge; wing membrane black in between
dark base and whitish tip. Ears long and narrow, fingers, but orange along fingers and near body.
extending 5mm beyond muzzle when folded Taxonomic notes Formerly called M. formosus,
forwards; tragus long, thin and bluntly pointed. which is now known to be a distinct species. Similar
Hind foot moderately long, over 60% of tibia length; species Herman’s Myotis, M. hermani, has similar
wing membrane attached on side of foot at base of coloration, but is larger; Hodgson’s Woolly Myotis,
toes; calcar weakly lobed at midpoint. Middle upper M. formosus, has paler, yellowish fur, without black
and lower premolars small. Similar species Large tips to hairs, little black on ears, brown thumb;
Brown Myotis, M. montivagus, and related species Painted Bat, Kerivoula picta, is smaller with more

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 funnel-shaped ears, more pointed tragus, no black darker red fur with black tips; black rims to ears;
on head, different dentition. Painted Bat, Kerivoula picta, is smaller with more
Ecology and habitat Has been caught over a river funnel-shaped ears, more pointed tragus, different
in evergreen forest. dentition.
Distribution SE Asia: Laos, Vietnam. Also China, Ecology and habitat No information from SE Asia.
Korea, Taiwan, Japan. Distribution SE Asia: Vietnam. Also Afghanistan,
Status Not yet assessed separately from India, Nepal, China, Taiwan.
M. formosus. Status Least Concern, although only one record
from region.
HERMAN’S WOOLLY MYOTIS
Myotis hermani NOT ILLUSTRATED ‘Selysius’ group Small to medium-sized Myotis
Measurements FA 56–60, T 58, HF 12, E 20. with relatively small feet, wing membrane attached
Wt 16g. Skull: cbl 19.5–20.0, mt 8.6 at base of toes. Ears not especially long. Tragus
Identification Similar in colour and pattern to narrow and tapered, typically angled slightly
Black-and-Orange Woolly Myotis, M. rufoniger, forwards. Middle lower incisors with only three
but larger. Body fur orange-red to orange-brown, lobes.
individual dorsal hairs with dark base, pale middle,
reddish-orange end and black tip; underside paler ASIAN WHISKERED MYOTIS
orange. Black rim to ears, black thumb, black feet; Myotis muricola PLATE 18
black and orange pattern on wing membranes. Measurements FA 33–37, T 35–41, E 12.5–14.5,
Similar species Black-and-Orange Woolly Myotis, HF 6–7. Wt 4–7g. Skull: ccl 11.5–11.8, mt 4.8–5.4
M. rufoniger, is smaller; Painted Bat, Kerivoula Identification Upperparts brown to grey with dark
picta, is much smaller with more funnel-shaped base; underparts with dark base, light brown
ears, more pointed tragus, no black on head; tip. Ears moderately long, tragus slender, bent
Hodgson’s Woolly Myotis, M. formosus, has paler forwards and bluntly pointed. Feet small with wing
yellow fur without black tips; little black on ears; membrane attached at base of toes. Middle upper
brown thumb and hind foot. premolar small and often slightly intruded from the
Ecology and habitat Has been caught over rivers toothrow (as in Fig. 37c). Upper canine much longer
in lowland rainforest. than posterior upper premolar. Taxonomic notes
Distribution SE Asia: peninsular Thailand and Genetic studies indicate that bats currently referred
Malaysia. Also Sumatra. to as M. muricola represent a complex of species,
Status Data Deficient: apparently rare and if but diagnostic characters and appropriate names
dependent on mature forest, probably declining. have not yet been worked out. Similar species
Eurasian Whiskered Myotis, M. mystacinus, is paler
HODGSON’S WOOLLY MYOTIS with whiter underparts; Peters’s Myotis, M. ater,
Myotis formosus PLATE 18 is larger, with a larger skull and smaller second
Measurements FA 45–53. Skull: ccl 15.8–17.3, premolars; Small-toothed Myotis, M. siligorensis,
mt 6.9–7.9 is smaller with a domed braincase, lower canines
Identification Moderately large Myotis with long small; Anna Tess’s Myotis, M. annatessae, has
woolly fur, generally pale yellow to orange-yellow. smaller canines; Ridley’s Myotis, M. ridleyi, has
Dorsally, hairs predominantly pale yellow, with a shorter forearm, darker fur and lacks the small
small dark base (10%) and sometimes with browner second premolar; Horsfield’s Myotis, M. horsfieldii,
tip; underside hairs usually uniformly pale yellow. has larger feet with wing membrane attached
Ears with no or only slight dark edges; thumb and at side of the foot; pipistrelles, Pipistrellus spp.,
base of hind foot brown. Wing membranes orange have different-shaped tragus and only two upper
to orange-brown with large dark patches between premolars, the anterior displaced inwards; woolly
the fingers. Taxonomic notes Recently shown bats, Kerivoula spp., have more funnel-shaped
to be distinct from M. rufoniger. Similar species ears with long, straight, pointed tragus and longer,
Black-and-Orange Woolly Myotis, M. rufoniger, has fluffier hair.

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 Ecology and habitat Forages in relatively open Ecology and habitat Has been found near a cave
areas, including over streams and rivers near in disturbed habitats at 1,400m.
forests, and agricultural areas and fields. Roost Distribution Myanmar. Also through much of
sites include furled central leaves of banana plants Palaearctic, from Europe through Nepal and India.
and vegetated areas near cave entrances. Poorly known in region, with only one confirmed
Distribution SE Asia: throughout region. Also record; may have been overlooked owing to
E India, S China, Indonesia from Sumatra through to confusion with M. muricola.
New Guinea and the Philippines. Status Least Concern.
Status Least Concern.
PETERS’S MYOTIS
ANNA TESS’S MYOTIS Myotis ater PLATE 18
Myotis annatessae NOT ILLUSTRATED Measurements FA 34–39, T 43–48, E 13.5,
Measurements FA 32.5–35.5, T 37–39, E 12–13, HF 6.5–7.5. Wt 5.0–7.0g. Skull: cbl 14.4–14.5,
HF 6–8. Wt 2.9–4.8g. Skull: ccl 11–12, mt 4.6–5.0 mt 5.8–5.9
Identification Upperparts grey-brown, hairs Identification Upperparts dark brown, hairs with
blackish at base, brown at tips. Underpart hairs with slightly paler brown tips; underparts with long,
blackish base, brown middle, silvery tip. Hind feet dark brown base and buffy-brown tip; second
moderately small, calcar without well-defined keel. upper and lower premolars usually very small
Rostrum slender, braincase slightly more domed and displaced inwards, so that the first and third
than M. muricola. Upper canine small, about 1.2 × premolars are touching, or nearly so. Taxonomic
height of posterior molar. Lower canine only slightly notes It is unclear whether bats referred to this
taller than posterior molar. Lower molars myotodont species from mainland South-east Asia are actually
(Fig. 39). Taxonomic notes Newly described in the same species as those in Maluku, which is the
2013. Similar species Asian Whiskered Myotis, type locality of the name. Similar species Asian
M. muricola, averages slightly larger and has Whiskered Myotis, M. muricola, is very similar,
longer upper canine about 1.5 × height of posterior but has smaller skull and teeth, tends to have
premolar. Small-toothed Myotis, M. siligorensis, paler fur, usually shorter forearm; Large Brown
and relatives have canine shorter than premolar, Myotis, M. montivagus, has slightly longer forearm,
nyctalodont-like lower molars, larger hind feet. substantially larger skull.
Ecology and habitat Has been caught over small Ecology and habitat Has been found over streams
streams. in lowland rainforest in Malaysia. Probably uses a
Distribution SE Asia: Vietnam, Laos. variety of habitats.
Status Not yet assessed. Distribution SE Asia: Vietnam, peninsular
Thailand and Malaysia. Also Sumatra, Borneo,
EURASIAN WHISKERED MYOTIS the Philippines, Sulawesi and Indonesian islands
Myotis mystacinus PLATE 18 through to New Guinea.
Measurements FA 34–37, T 32–40, E 12–14, Status Least Concern.
HF 7–8. Wt 4–5.5g. Skull: ccl 12.3, mt 5.4
Identification Upperparts greyish buff-brown, hairs LARGE BROWN MYOTIS
with dark brown base; underparts with broad white Myotis montivagus NOT ILLUSTRATED
to creamy white tip with dark base. Ears moderately Measurements FA 39–41. Skull: ccl 14.0–14.1,
long, tragus slender, bent forwards and bluntly mt 6.1–6.2
pointed. Feet small with wing membrane attached Identification Upperpart fur with dark brown base,
at base of toes. Middle upper premolar small and mid-brown tip; underparts similar but fur with paler tip.
slightly intruded from toothrow. Upper canine much Feet relatively small with wing membrane attached at
longer than posterior upper premolar. Similar base of toes. Upper middle premolar small and only
species Asian Whiskered Myotis, M. muricola, is partially intruded; lower middle premolar small (as
darker with buffy-brown underparts, slightly shorter in Fig. 37e) but within toothrow. Taxonomic notes
toothrow. Several populations formerly considered part of

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 a

0 5

Fig. 37 Upper (on left) and lower (on right) toothrows of selected Myotis, showing variation in number, size and position of
premolars: M. rosseti (a), M. silgorensis (b), M. horsfieldii (c), M. hasseltii (d), M. indochinensis (e) and M. annectans (f).

M. montivagus are now considered separate species, Status Least Concern, although this should be
including M. indochinensis, M. federatus and reviewed, as assessment included several
M. borneoensis. Similar species Hairy-faced Myotis, populations now considered separate species.
M. annectans, is similar in size, but lacks middle
upper and lower premolars (Fig. 37f), has longer, LARGE INDOCHINESE MYOTIS
paler fur with whitish underparts; Szechuan Myotis, Myotis indochinensis PLATE 18
M. altarium, has relatively large middle premolars Measurements FA 43–46, T 44–51, E 15–16,
within toothrow, larger feet. Large Indochinese Myotis, HF 8.5–9.5. Wt 12–18g. Skull: ccl 15.6–16.4,
M. indochinensis, has longer forearm, larger skull. mt 7.0–7.5
Ecology and habitat Poorly known. Recorded at Identification Upperparts with medium-length
altitudes up to 1,500m. uniformly brown fur; underparts similar but paler,
Distribution SE Asia: N Myanmar. Also S China, hairs with dark base and distal third pale. Feet
NE India. Possibly also Vietnam, but identity of relatively small with wing membrane attached at
specimens not yet confirmed. base of toes. Middle upper premolar very small

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 and completely intruded in toothrow; middle lower underparts with dark brown base and silvery-white
premolar small and partially intruded. Taxonomic tip, except middle of belly, where tips of hairs are
notes Newly described in 2013 as distinct from orange-brown. Middle upper and lower premolars
Myotis montivagus. Similar species Hairy-faced usually lacking (Fig. 37f); sometimes present
Myotis, M. annectans, is similar in size, but lacks asymmetrically, but very small. Ears moderately
middle upper and lower premolars (Fig. 37f), has large; tragus long and slender, bent forwards and
longer fur with pale tips, whitish underparts and rounded at tip; wing membrane inserts on side
often an orange patch on the belly; Szechuan of foot just above base of toe; calcar with little or
Myotis, M. altarium, has relatively large middle no keel. Similar species Large Brown Myotis,
premolars within toothrow, larger feet; Hasselt’s M. montivagus, and Large Indochinese Myotis,
Large-footed Myotis, M. hasseltii, has large feet M. indochinensis, have shorter, darker fur, middle
with wing membrane attached at ankle. upper and lower premolars small but present;
Ecology and habitat Recorded from understorey Szechuan Myotis, M. altarium, is darker, with well-
of mature evergreen and secondary forests, at developed middle premolars.
altitudes of 300–1,000m. Ecology and habitat Recorded from hill forest
Distribution SE Asia: Laos and Vietnam. Also in Laos.
China. Distribution SE Asia: N Thailand, Laos, Cambodia
Status Not yet assessed. and Vietnam. Also NE India.
Status Least Concern.
LARGE MALAYAN MYOTIS
Myotis federatus NOT ILLUSTRATED THICK-THUMBED MYOTIS
Measurements FA 39.5–42.5. Wt 12.5g. Myotis rosseti PLATE 18
Skull: ccl 15.0–15.1, mt 6.4–6.8 Measurements FA 29–31, T 35, E 13–14, HF 7.
Identification Fur uniformly very dark brown. Wt 4.5–6.0g
Ears moderately large and narrow, tragus bent Identification Small Myotis with relatively short
forward, reaching half height of ear. Feet relatively fur, greyish-brown above, paler underneath, the
small with wing membrane attached at base individual hairs with dark brown base and pale
of toes. Middle upper premolar very small and tip. Enlarged, thickened, rounded pad on base of
completely intruded in toothrow; lower middle thumb, slightly pinker than rest of wing, which is
premolar small and intruded so not visible from very dark grey; small triangular pad on foot. Ears
the side. Taxonomic notes Formerly considered relatively long, extend 2–3mm past muzzle when
a subspecies of M. montivagus. Similar species bent forwards. Tragus thickened at base, angled
Large Indochinese Myotis, M. indochinensis, is forwards and tapering to tip. Feet relatively small
larger, fur is not as dark, middle lower premolar with wing membrane attached at side of base of
relatively larger. toes. Only two upper and lower premolars (Fig.
Ecology and habitat Recorded from understorey 37a). Similar species Ridley’s Myotis, M. ridleyi,
of mature and disturbed forests, at altitudes from is more blackish and lacks thumb pad; Thick-
lowlands up to 900m. thumbed Pipistrelle, Glischropus tylopus, has
Distribution SE Asia: Peninsular Malaysia. Also shorter, broader ears, smaller and more rounded
Sumatra. tragus, smaller and pinker thumb and footpads,
Status Not yet assessed. distinctive upper incisors.
Ecology and habitat Poorly known. Known to
HAIRY-FACED MYOTIS forage over disturbed areas including fields; has
Myotis annectans PLATE 18 been found over a small stream. Roost sites include
Measurements FA 45–49, T 44–49, E 15.5–17.5, houses, hollow trees and bamboo.
HF 8.5–10.5. Wt 8.5–13g. Skull: ccl 14.8–15.7, Distribution SE Asia: Thailand, Laos, Vietnam and
mt 6.6–6.9 Cambodia.
Identification Upperparts with long fur, base dark Status Least Concern, although known only from
brown with short, shiny pale tip, giving frosted effect; small number of records.

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 RIDLEY’S MYOTIS HORSFIELD’S MYOTIS
Myotis ridleyi PLATE 18 Myotis horsfieldii PLATE 19
Measurements FA 27–32, T 30–36, E 10–12, Measurements FA 35–38, T 37–39, HF 10–11,
HF 6.0–7.5. Wt 4–6g E 13.5–17. Wt 5–7.5g. Skull: cbl 14.3–14.7,
Identification A small, dark bat with short wings mt 5.8–6.0
but a relatively heavy body and a typical Myotis ear Identification Upperparts grey-brown; underparts
and tragus shape. Upperparts dark grey-brown; greyer. Ears large. Feet moderately large with wing
underparts paler and greyer. Feet small with wing membrane attached to side of foot 1–2mm from
membrane attached to side of foot. Has only two base of toes. Second upper and lower premolars
upper and lower premolars (as in Fig. 37a). Similar only slightly displaced inwards (Fig. 37c). Similar
species Most other Myotis have three upper species Hasselt’s Myotis, M. hasseltii, has wing
and lower premolars, although middle one can membrane attached at ankle, middle premolar tiny
be small and hard to see; Small-toothed Myotis, and protruded inwards (Fig. 37d); Asian Whiskered
M. siligorensis, is paler, with a lighter body, short Myotis, M. muricola, and its relatives have smaller
canines and a conspicuous middle premolar; Thick- feet with wings attached at base of toes.
thumbed Myotis, M. rosseti, has thickened thumb Ecology and habitat Roosts in crevices or bell-
pad, longer ears; pipistrelles also have only two holes in caves, usually not far from large streams or
premolars, but anterior upper premolar is small and rivers. Feeds low over open surfaces of water such
usually displaced inwards, ears and tragus are a as wide streams.
different shape. Distribution SE Asia: Laos, Vietnam, Cambodia,
Ecology and habitat Known only from lowland Thailand and Peninsular Malaysia. Also known from
rainforest. Forages in understorey, especially over India, S China, Borneo, Java, Bali, Sulawesi and the
small streams. Roost sites include caves, fallen logs Philippines.
and under rocks. Status Least Concern.
Distribution SE Asia: peninsular Thailand and
Malaysia. Also Borneo. HASSELT’S MYOTIS
Status Near Threatened. Population has declined Myotis hasseltii PLATE 19
due to extensive forest loss in its range. Measurements FA 38–43, HF 10–11. Wt 7.5–
12.5g. Skull: cbl 14.1–14.6, mt 5.6–5.8
Singapore Myotis, Myotis oreias, was listed in Identification Upperparts dark brown to dark grey
the first edition of this book, but recent study has with pale grey tips to hairs; underparts greyish-
shown it is not a valid species. The type specimen white with dark base. Fur short and velvety. Flight
is a skin which is actually a Kerivoula, resembling membranes pale grey-brown. Fur appears silvery
K. papillosa, but the species is uncertain. The when spot-lit in flight at night. Feet large, with
skin was mismatched with a damaged skull wing membrane attached at ankle. Second upper
of a Myotis which had confused subsequent premolar very small and displaced inwards so that
researchers. The collection locality was recorded first and third premolars are in contact or nearly
only as ‘the Indian continent’ which at the time of so (Fig. 37d). Similar species Horsfield’s Myotis,
description (1840) could have included SE Asia. It M. horsfieldii, is smaller, with wing membrane
is unclear whether this could be a valid name for attached at side of foot; Rickett’s Myotis,
any currently recognised Kerivoula, but it is not a M. ricketti, is much larger, with very large
valid Myotis. hind feet; Australasian Large-footed Myotis,
M. adversus, has no confirmed records in region,
‘Leuconoe’ group Small to medium Myotis with but occurs nearby and may yet be found. It is
enlarged hind feet, with the wing attached at the similar, but distinguished by denser, woolly fur,
ankle or side of the foot. These large feet are used relatively larger middle upper and lower premolars
for scooping insects off water and, in some species, that are within toothrow or only slightly intruded.
for catching small fish. Middle lower incisors with Ecology and habitat Forages for food low over
only three lobes. larger areas of open water, such as large rivers,

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 lakes and sea, including coastal areas near and light-bodied. Feet medium-sized with wing
mangroves. Catches food by dipping its large feet membrane attached on side of foot about 1mm
into the water. Has been found roosting in caves. from base of toes. Ears reach just past tip of nose
Distribution SE Asia: Myanmar, Thailand, Vietnam, when folded forwards. Second upper and lower
Cambodia and Peninsular Malaysia, especially in premolars only slightly smaller than corresponding
coastal areas and along large rivers and lakes. Also first premolar and in line in toothrow. Skull has
Sumatra, Java, Borneo and smaller intervening distinctly domed braincase (as in Fig. 36a), which is
islands. reflected in live animal by forehead rising abruptly
Status Least Concern. from rostrum. Canines small, upper canine similar
in height to posterior upper premolar, lower canine
RICKETT’S MYOTIS about same height as anterior lower premolar.
Myotis pilosus PLATE 19 Lower molars nyctalodont-like (Fig. 39). Taxonomic
Measurements FA 53–56, T 51–55, HF 15.5– notes SE Asian form M. s. alticraniatus may
17.5, E 18–20. Wt 14–22g be a separate species from M. s. siligorensis in
Identification Large Myotis with very large feet, Nepal. Similar species Asian Whiskered Myotis,
about 80% of tibia length, with long, curved M. muricola, is larger, with longer canines and a
claws; wing membrane attached at ankles. flatter skull; Ridley’s Myotis, M. ridleyi, has a shorter
Upperparts buffy grey-brown, hairs with darker forearm but is otherwise darker and heavier, with
base; underparts greyish-white with darker base. only two premolars; Indochinese Water Myotis,
Taxonomic notes Formerly called M. ricketti. M. laniger, is similar but slightly larger, with longer
Similar species Hasselt’s Myotis, M. hasseltii, is ears and slightly larger canines.
smaller, and has smaller feet. Ecology and habitat Has been found roosting in
Ecology and habitat Roosting sites not reported, rock fissures in caves. Occurs in lowlands up to
but has been found along rocky streams near 1,600m in both primary forest and disturbed areas.
limestone outcrops and likely roosts in caves. Usually associated with water, including small
Extremely large feet used for catching prey off streams.
water, including small fish. Distribution SE Asia: Myanmar, Thailand, Laos,
Distribution SE Asia: Laos and N Vietnam. Also Vietnam, Cambodia and Peninsular Malaysia. Also
China. found from Nepal through S China and in Borneo.
Status Near Threatened due to loss of suitable Status Least Concern.
clean streams for foraging.
PHAN LUONG’S MYOTIS
‘Siligorensis’ group These Myotis are character- Myotis phanluongi NOT ILLUSTRATED
ised by small size, short weak dentition, and only Measurements FA 33–36, T 35, E 12–13, HF 7.5.
slightly enlarged hind feet, with the wing attached Skull: ccl 11.2–11.9, mt 4.9–5.3
at the side of the foot near the toes. They are often Identification Similar to Small-toothed Myotis, M.
caught near water, and may scoop insects off siligorensis, but generally slightly larger. Canines
water. Many species are very similar to each other shorter than upper molars. Wing membrane
and hard to identify. attached to side of foot, just above base of toe.
Baculum very small and narrow. Rostrum narrow,
SMALL-TOOTHED MYOTIS braincase conspicuously domed. Lower molars
Myotis siligorensis PLATE 19 nyctalodont (Fig. 39). Taxonomic notes Newly
Measurements FA 30–35, T 28–38, E 11–12.5, described in 2008. Similar species Small-toothed
HF 6.5–7.0. Wt 2.3–4.0g. Skull: ccl 10.2–10.8, Myotis, M. siligorensis, is smaller; Annamite Myotis,
mt 4.2–4.6 M. annamiticus, has larger foot, wing membrane
Identification Upperparts light reddish-brown inserted on side of foot.
to sandy-brown, the fur with dark brown base; Ecology and habitat Has been caught over small
underparts paler with buffy or white tip to fur. streams in montane evergreen forest at altitudes of
Relatively long forearm, but otherwise very small 1,200–1,500m.

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 Distribution SE Asia: Vietnam. species Small-toothed Myotis, M. siligorensis, is
Status Not yet assessed. smaller, with shorter canines, shorter ears, slightly
smaller feet; Annamite Myotis, M. annamiticus,
ANNAMITE MYOTIS is slightly smaller with smaller upper canines,
Myotis annamiticus NOT ILLUSTRATED nyctalodont-like lower molars; Horsfield’s Myotis,
Measurements FA 30.5–34.5, T 33–36, E 13–15, M. horsfieldii, has larger canines, longer than the
HF 7.0–8.2. Wt 3.3–5.5g. Skull: ccl 11.3–11.6, premolars, and larger, more flattened skull.
mt 4.8–5.1 Ecology and habitat Forages low over water,
Identification Upperparts brown, fur with dark especially small streams. Probably roosts in caves.
base, underparts greyish-brown. Feet somewhat Distribution SE Asia: Laos and Vietnam. Also S China.
enlarged; wing membrane inserted on side of foot Status Least Concern.
1–2mm from base of toes. Ears long and narrow.
Skull with domed braincase that rises abruptly from Genus Eudiscopus Ear shape somewhat similar
rostrum (Fig. 36a), reflected in steep forehead in to Myotis, with narrow tragus bent forwards, but
live bat. Canines small, upper canine similar in characterised by large pads on feet and very
height to posterior premolar; lower canine shorter flattened skull (Fig. 36c). Only two upper premolars,
than posterior premolar. Lower molars nyctalodont- both within toothrow; three lower premolars, middle
like (Fig. 39), Taxonomic notes First described ones very small and completely intruded within
in 2000. Similar species Small-toothed Myotis, toothrow. Dental formula 2/3 1/1 2/3 3/3.
M. siligorensis, has smaller canines, shorter ears;
Indochinese Water Myotis, M. laniger, is slightly DISC-FOOTED BAT
larger, especially in skull measurements, with larger Eudiscopus denticulus PLATE 22
upper canine and myotodont lower molars. Measurements FA 35–38, T 36–40, E 13–14.
Ecology and habitat Forages over water surface Wt 5.0g. Skull: ccl 12.8–13.4
on small streams in forested and disturbed areas. Identification Upperparts medium sandy-brown;
Probably roosts in rock crevices and limestone underparts paler buffy-brown. Ears shaped
caves. somewhat like Myotis, with narrow tragus that
Distribution SE Asia: Laos and Vietnam. Also bends forwards in middle. Sole of foot modified into
S China. an enlarged pink pad like a suction cup, with short
Status Data Deficient; only known from a few toes; wing membrane inserted at end of pad at
specimens, but region has not been well surveyed. base of outer toe; calcar extends from base of pad
to support tail membrane with well-developed keel.
INDOCHINESE WATER MYOTIS Thumb short, thickened at base, but not forming a
Myotis laniger PLATE 19 pad. Skull very flat. Similar species Bamboo bats,
Measurements FA 31–36, T 33–36, E 13–15, Tylonycteris spp., also have enlarged footpads,
HF 7.2–8.2. Wt 3.3–4.0g. Skull: ccl 11.4–13.0, but are much smaller, wing membrane inserts at
mt 4.7–5.7 ankle opposite calcar, head is much flatter, ear is
Identification Upperparts brown, fur with dark a different shape; Myotis spp. have similar-shaped
base, underparts greyish-brown. Feet somewhat ears, but braincase is more elevated, feet do not
enlarged; wing membrane inserted on side of foot have enlarged pads.
1–2mm from base of toes. Ears very long and Ecology and habitat Roosts inside bamboo
narrow, extending 4–5mm beyond nostrils when internodes; footpad provides enough suction to
folded forwards. Skull with domed braincase that hang from a smooth surface. Apparently forages
rises abruptly from rostrum (as in Fig. 36a), reflected near thickets, bamboo and forests, with slow,
in steep forehead in live bat. Canines small, upper manoeuvrable flight, including hovering.
canine similar in height to posterior premolar; Distribution SE Asia: Myanmar, Thailand, Laos
lower canine shorter than posterior premolar. and Vietnam.
Lower molars myotodont (Fig. 39). Taxonomic Status Least Concern, but relatively rare compared
notes Formerly included in M. daubentoni. Similar with other bamboo specialists.

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 a b c

d e f

g h i

0 10

Fig. 38 Upper left toothrows of selected pipistrelle-like bats: Pipistrellus abramus (a), Glischropus tylopus (b),
Philetor brachypterus (c), Arielulus circumdatus (d), Hesperoptenus tomesi (e), Scotomanes ornatus (f),
Thainycteris aureocollaris (g), Scotophilus heathii (h) and Eptesicus pachyomus (i).

Subfamily VESPERTILIONINAE
This subfamily includes several morphologically distinct genera, which can be distinguished by a
combination of dental characters, ear and tragus shape. Recent genetic work has shown that some
traditionally recognised genera should actually be considered separate.

Genus Barbastella Broad, forward-facing ears brown. Muzzle broad, with prominent swellings.
joined across forehead. Skull with enlarged Taxonomic notes Appropriate name has not yet
braincase, sloping steeply to narrow front of been determined. Formerly called B. leucomelas,
shortened rostrum. Anterior upper premolar tiny, but recent analyses show it is not closely related;
completely displaced inwards. Dental formula 2/3 morphologically Vietnam specimens resemble
1/1 2/2 3/3. B. darjelingensis from Nepal, but genetically
they are quite distinct. Similar species No other
FAR-EASTERN BARBASTELLE vespertilionids in region have ears joined on
Barbastella sp. PLATE 23 forehead.
Measurements FA 38–43, T 40–47, E 15–17. Ecology and habitat Vietnamese specimens
Skull: ccl 13.4–14.2 recorded from montane forest at altitude of nearly
Identification Upperparts black, slightly glossy on 2,000m. Other members of genus roost singly or in
the lower back; underparts dark blackish-brown. small groups, under bark and in tree hollows.
Ears relatively broad, facing forwards, joined at base Distribution SE Asia: Laos, N. Vietnam. Also China.
across forehead. Membranes and ears blackish- Status Not yet assessed. Apparently rare in region.

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 Genus Pipistrellus This genus was formerly Some species appear extremely similar to one
considered to comprise a broad range of species another, and appropriate methods for distinguishing
characterised by having only two upper and lower them in the field have not yet been worked out.
premolars (Fig. 38). More recent analyses indicate In particular, for a few species, the main known
that many of these species are not closely related identification characters are the size and shape
and should be separated into several different of the penis and the baculum (the bone inside the
genera, including, within SE Asia, Hypsugo, penis). These differences in reproductive organs
Arielulus and Falsistrellus, as well as Pipistrellus indicate that the species are reproductively isolated
itself. All of them are distinguished from Myotis by from one another and hence valid species, but are
shape of ears and tragus (Fig. 40). only sometimes perceptible in the field and not
In Pipistrellus the anterior premolars, although helpful in identifying females. Nevertheless, they
much smaller than the posterior premolar, are are mentioned here for the benefit of museum
usually reasonably well developed, the upper ones workers, and because close examination of these
similar in size to the second incisor and only partially characters may help with discovering additional
intruded within the toothrow. Ears simple, usually differences that are more useful for distinguishing
somewhat rounded. Tragus usually somewhat these species in life.
club-shaped, rounded at the tip. Calcar has well-
developed, rounded lobe on the outside. Incisors JAVAN PIPISTRELLE
relatively small. Males usually have relatively long Pipistrellus javanicus PLATE 20
penis, with elongate baculum, sometimes extremely Measurements FA 30–36, T 34–40, E 10.5–11.5.
long. Skull variable, but with braincase generally Wt 4.0–7.0g. Skull: ccl 11.9–13.1, mt 4.6–5.2,
low; rostrum only slightly shortened. Lower molars M–M 5.6–6.7
typically with nyctalodont cusp pattern (Fig. 39). Identification Upperparts dark brown with dark
Dental formula 2/3 1/1 2/2 3/3. base; underparts slightly paler, tip mid-brown,
Although some species are very common, base dark brown. Ear moderate length, narrow,
others are known from only a few specimens, with distinct fold at posterior edge; tragus narrow,
possibly because their behaviour makes them hard relatively straight, with rounded tip (Fig. 40a).
to catch, rather than because they are rare. Recent Anterior upper premolar slightly displaced inwards
surveys and genetic studies suggest a number (as in Fig. 38a). Braincase somewhat elevated,
of additional species occur in the region, but the giving a distinct rostral depression. Upper canine
identification characters and their names have not usually has secondary cusp. Male has moderately
yet been sorted out. long penis (more than 6mm); with medium-length

Trigonid Trigonid

Entoconid Hypoconid

Hypoconid

Postcritid
Postcritid Entoconid
Hypoconulid Hypoconulid

Fig. 39 First lower left molars showing cusp patterns of a nyctalodont (left: Pipistrellus javanicus) and myotodont (right:
Hypsugo pulveratus) tooth. In the nyctalodont, the postcristid ridge connects the hypoconid to the hypoconulid, while in
the myotodont it is further forwards and connects to the entoconid.

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 tragus narrow, rounded, angled slightly forwards (as
in Fig. 40a). Upper canine with a small secondary
cusp on posterior cutting edge. Penis long, baculum
c.9mm with strong double curve. Similar species
Javan Pipistrelle, P. javanicus, very similar except
for smaller penis in male, slightly larger skull,
generally darker brown fur; Mount Popa Pipistrelle,
a b c d P. paterculus, lacks secondary cusp on canine,
male has longer penis, long, straight baculum.
Ecology and habitat In N Vietnam, common
around human settlements and heavily disturbed
areas.
Distribution SE Asia: Myanmar, and Vietnam. Also
E Russia, China, Korea and Japan.
Status Least Concern.
e f g
MOUNT POPA PIPISTRELLE
Fig. 40 Right tragus of selected pipistrelles: Pipistrellus Pipistrellus paterculus PLATE 21
javanicus (a), P. tenuis (b), P. ceylonicus (c), Arielulus Measurements FA 27–34, T 29–34, E 7.5–12.0.
societatis (d), Pipistrellus stenopterus (e), Falsistrellus Wt 4.5–6.0g. Skull: ccl 10.6–11.6, mt 4.1–4.8,
affinis (f) and Hypsugo macrotis (g). M–M 5.3–5.9
Identification Fur colour uniformly dark brown
baculum. Similar species Japanese Pipistrelle, above, base essentially same colour as tip;
P. abramus, and Mount Popa Pipistrelle, underparts with sandy-brown tip, dark brown base.
P. paterculus, have slightly different fur colours, Upper canine usually without a secondary cusp. Ear
longer penis and baculum; Least Pipistrelle, subtriangular with rounded tip, short, blunt tragus
P. tenuis, is smaller with shorter, more sloping skull, angled slightly forwards. Male has extremely long
shorter baculum; Myotis spp. have distinctively penis, which is bent at right angles, about 4.5mm
shaped ear with tapered tragus, usually three upper from body to bend, then 12.5mm on straight
and lower premolars. part to tip; narrow, straight baculum, 9–12mm.
Ecology and habitat Known from a wide variety of Similar species Japanese Pipistrelle, P. abramus,
habitats, from primary hill forest up to 2,000m to also has long penis, but baculum has distinctive
heavily disturbed areas such as rubber plantations curve; females are difficult to distinguish, although
and towns. Has been found roosting in treeferns, fur colour may be helpful, at least within local
fallen logs, caves and urban areas. populations.
Distribution SE Asia: throughout the region. Also Ecology and habitat Has been found in disturbed
Afghanistan, Pakistan, India, China (Tibet), Sumatra, habitats and around villages. Roost sites include
Java, Borneo and elsewhere in Indonesia and the holes in tree stumps, thatched roof of a hut.
Philippines. Distribution SE Asia: Myanmar, Laos and Vietnam.
Status Least Concern. Also India and SW China.
Status Least Concern; relatively common.
JAPANESE PIPISTRELLE
Pipistrellus abramus PLATE 20 COROMANDEL PIPISTRELLE
Measurements FA 29–33, T 27–31, E 9.5–10.5. Pipistrellus coromandra PLATE 20
Wt 3.8–5.8g. Skull: ccl 10.9–11.1, mt 4.2–4.4, Measurements FA 26–34. T 28–35, E 9. Wt 3.4g.
M–M 5.2–5.4 Skull: ccl 10.6–11.9, mt 3.9–4.6, M–M 5.0–6.0
Identification Fur of upperparts with greyish-brown Identification Upperparts generally dark brown,
tip, darker base; underparts with pale brown tip and sometimes with reddish-brown cast; underparts
contrasting dark brown base. Ear with rounded tip, noticeably paler with pale brown tip, dark base to

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 fur. Skull similar to P. javanicus but slightly smaller, Distribution SE Asia: Myanmar, Thailand, Laos,
braincase somewhat flattened with relatively Vietnam, Cambodia and Peninsular Malaysia. Also
straight rostral profile. Upper canine has distinct Afghanistan through India, S China, Java, Sumatra,
secondary cusp. Penis moderate size, baculum Borneo, the Philippines and Sulawesi.
thin and straight, 3–4mm long. Similar species Status Least Concern.
Javan Pipistrelle, P. javanicus, averages slightly
larger with more domed braincase, moderately KELAART’S PIPISTRELLE
long penis; Japanese and Mount Popa Pipistrelles, Pipistrellus ceylonicus PLATE 20
P. abramus and P. paterculus, have much longer Measurements FA 33–42, T 30–35, E 13.
penis, latter tends to have more domed skull; Wt 7–8g. Skull: ccl 13.1–14.3, mt 5.2–5.9,
Least Pipistrelle, P. tenuis, has smaller skull that M–M 6.2–7.2
usually does not overlap in size in areas where Identification Relatively large Pipistrellus; fur of
both species occur. upperparts varies from grey-brown to reddish-
Ecology and habitat A wide variety of habitats from brown with dark base; underparts paler, hairs with
forest to agricultural fields and urban areas. Roost dark base and pale grey tip. Tragus broadly rounded
sites include under bark or among tree leaves and (Fig. 40c). Skull large and broad. Anterior upper
in buildings. premolar relatively large, similar in size to anterior
Distribution SE Asia: Myanmar, Thailand, Laos, incisor, displaced inwards, but canine and second
Vietnam and Cambodia. Also Afghanistan, India, premolar do not touch; lower anterior premolar
Nepal, Pakistan and S China. about 80% crown area of second. Canine has
Status Least Concern. secondary cusp in Indian examples, but not in only
known specimen from Myanmar. Baculum slender,
LEAST PIPISTRELLE about 4mm long. Similar species Narrow-winged
Pipistrellus tenuis PLATE 20 Pipistrelle, P. stenopterus, has short fifth finger;
Measurements FA 25–31, T 29–35, E 9–11.5. other Pipistrellus spp. are smaller; Hypsugo spp.
Wt 3–4g. Skull: ccl 9.3–10.7, mt 3.5–4.1, have larger, broader ears, with no keel on calcar,
M–M 4.5–5.2 very small anterior upper premolar; Doria’s False-
Identification Upperparts mid- to dark brown; serotine, Hesperoptenus doriae, lacks small anterior
underparts similar or slightly paler. Ear relatively premolar and has large inner incisors.
short, oval, with weak folds on posterior edge; Ecology and habitat Poorly known in region.
tragus short and rounded, angled slightly forwards Distribution SE Asia: N Myanmar, S Laos and
(Fig. 40b). Second upper premolar slightly displaced N Vietnam. Also Sri Lanka, Pakistan, India, S China
inwards. Upper canine with distinct secondary and Borneo.
cusp. Rostrum of skull short, narrow, with evenly Status Least Concern, although poorly known in
sloping profile from nose to top of skull. Penis of region.
moderate size; baculum thin and straight, c.4mm
long. Taxonomic notes Probably includes more NARROW-WINGED PIPISTRELLE
than one species across its very wide geographic Pipistrellus stenopterus PLATE 20
range. Similar species Coromandel Pipistrelle, Measurements FA 37–42, T 55–57, E 14–16.
P. coromandra, averages larger, especially in skull Wt 14–22g. Skull: ccl 15.1–15.5, mt 5.7–5.8,
dimensions, but can be hard to distinguish in life; M–M 7.6–7.9
other similar pipistrelles are larger, with much longer Identification Fur short; upperparts uniform
penis; Ridley’s Myotis, Myotis ridleyi, has a typical reddish-brown to dark brown; underparts slightly
Myotis ear shape, first upper premolar in line in paler and greyer. Muzzle broad and heavy. Ears
toothrow and not displaced inwards (as in Fig. 37a). moderate, broadly rounded. Tragus broad, hatchet
Ecology and habitat Found in a wide range shaped, strongly angled forwards (Fig. 40e). Wing
of habitats from forest to highly disturbed open narrow – fifth finger short, not much longer than
areas. Roosts include hollow branches, among dead metacarpal of fourth finger. Anterior upper premolar
leaves and in buildings. well developed, similar in size to outer upper incisor,

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 displaced inwards; anterior lower premolar slightly Scotophilus spp., have elongate narrow tragus, only
larger in crown areas than posterior premolar. Skull one upper incisor and one upper premolar; Asian
large and robust, forehead sloping or slightly domed Serotine, Eptesicus pachyomus, has narrower ear
in profile. Similar species Kelaart’s Pipistrelle, and tragus with narrow outer ear edge, broader
P. ceylonicus, is smaller, with longer fifth finger; wing, only one upper premolar.
Doria’s False-serotine, Hesperoptenus doriae, has Ecology and habitat Feeds high over canopy.
enlarged conical inner upper incisors, only one Unknown whether species is resident in S Vietnam
upper premolar; Hypsugo spp. have greatly reduced or migrates from farther north.
anterior upper premolars, longer fifth finger. Distribution SE Asia: N Myanmar, S Vietnam. May
Ecology and habitat Forages in open areas, also occur in Laos. Also in China and Himalayas.
over fields and rivers, and most likely over Status Not yet assessed separately from N. noctula.
houses. Roosts in hollow trees in forests and Only a few records from region.
rubber plantations, as well as in roofs of houses,
sometimes with Scotophilus kuhlii. Genus Glischropus Similar to Pipistrellus, but
Distribution SE Asia: S peninsular Thailand and sole of foot and base of thumb with thickened,
Peninsular Malaysia. Also Sumatra, Borneo and the unpigmented (pink) pads, smaller than dark pads
Philippines. of bamboo bats, Tylonycteris; second (outer) upper
Status Least Concern. incisor displaced outwards from toothrow (Fig. 38b).
Dental formula 2/3 1/1 2/2 3/3.
Genus Nyctalus Medium-sized to large
vespertilionid bats with relatively narrow, elongate SUNDA THICK-THUMBED PIPISTRELLE
wings; fifth finger short, only slightly longer than Glischropus tylopus NOT ILLUSTRATED
fourth metacarpal. Wing membrane attached at side Measurements FA 28–31, T 32–36, E 9–11.
of foot. Ears relatively short and broad, triangular; Wt 3.7–4.8g. Skull: ccl 11.6–12.4, mt 4.6–4.9
tragus short, broadened at tip, angled forwards and Identification Upperparts dark brown; underparts
bluntly pointed. Anterior upper premolar very small, paler buffy-brown. Fur rather long and shaggy.
usually completely intruded within toothrow, hidden Short, broad face with rounded head. Base of thumb
in lateral view by canine. Outer upper incisor similar and sole of foot have thickened, unpigmented,
in crown area to inner incisor, but about half its whitish or pink pads; wing membrane inserted
height. The taxonomic status of Asian specimens is at base of toes; calcar with well-developed keel.
uncertain. Dental formula 2/3 1/1 2/2 3/3. Second upper incisor displaced outwards; anterior
upper premolar small but in line in toothrow
ASIAN NOCTULE (Fig. 38b). Taxonomic notes Previous records of
Nyctalus cf. labiata PLATE 23 Glischropus from Indochina are now recognised
Measurements FA 49–58. Wt 23g. as a different species, G. bucephalus. Similar
Skull: ccl 17.1–18.6, mt 7.1–7.7, M–M 8.5–9.5 species Indochinese Thick-thumbed Pipistrelle,
Identification Fur of upperparts uniformly coloured G. bucephalus, is larger with more elevated
from base to tip, varying among individuals from braincase, no overlap in range; Least False-
dark brown to mid-brown; underparts similar but serotine, Hesperoptenus blanfordi, has dark thumb
slightly paler. Ear broadly triangular, posterior edge pads, short smooth fur, and the second upper
broadened from halfway down ear around below incisor displaced inwards (as in Fig. 38e); bamboo
tragus to near side of mouth; tragus short and bats, Tylonycteris spp., have very flat skull and
rounded, broader at tip than base. Wing with fifth large, rounded, dark thumb pads; other pipistrelles
finger substantially shorter than fourth finger, third have second upper incisor in line in toothrow, lack
finger elongate. Long calcar with well-developed pads on feet.
lobe. Taxonomic notes Formerly considered same Ecology and habitat Roosts in dead or damaged
species as N. noctula. Recent studies indicate bamboo stems, as well as rock crevices or new
it is distinct, but uncertain whether N. labiata is banana leaves.
appropriate name. Similar species House bats, Distribution SE Asia: peninsular Thailand and

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 Malaysia. Also Sumatra, Borneo and the Philippines. somewhat flattened, ears broadly triangular,
Status Least Concern. rounded at tip and prominent. Wings narrow
with fifth digit shortened, tip reaching only to
INDOCHINESE THICK-THUMBED halfway along first phalanx of fourth finger. Only
PIPISTRELLE one, shortened, upper premolar (Fig. 38c). First
Glischropus bucephalus PLATE 22 upper incisor long and narrow with two cusps,
Measurements FA 32–36, T 28–39, E 10–11, second incisor small and conical. Canine with
HF 6.1–7.5. Wt 4.3–5.2g. Skull: ccl 11.6–12.4, large secondary cusp. Similar species Narrow-
mt 4.6–4.9 winged Pipistrelle, Pipistrellus stenopterus, also
Identification Upperparts with relatively long dark has a narrow wing, but is larger and has an extra
brown fur, slightly paler at tip; underside fur with upper premolar; Surat Helmeted Bat, Cassistrellus
dark brown base and distal third pale yellow-brown. dimissus, has short, broad upper incisors, fifth digit
Base of thumb and sole of foot with thickened pink not shortened and less elaborate genitalia.
pads. Second upper incisor displaced outwards Ecology and habitat Roosts include hollow trees.
such that incisors are almost in a straight row; Has been found in both pristine forest and disturbed
anterior upper premolar displaced inwards and areas, but usually near intact forest.
largely hidden from side. Skull with distinctly Distribution SE Asia: Peninsular Malaysia. Also
elevated braincase, thickened rostrum (Fig. 36h). Sumatra, Java, Borneo, the Philippines, Sulawesi
Taxonomic notes Newly described as distinct from and New Guinea.
G. tylopus in 2011. Similar species Sunda Thick- Status Least Concern. Population is likely to have
thumbed Pipistrelle, G. tylopus, is smaller with a declined somewhat due to loss of forest.
sloping skull profile, anterior upper premolar not
displaced inwards; does not overlap in range. Genus Hypsugo Until recently, included within
Ecology and habitat Has been found in dry Pipistrellus. Genetic analyses suggest that some
dipterocarp forest and disturbed areas near small species now placed in Hypsugo are not closely
streams and rivers. May roost inside bamboo. related to each other and may not belong in this
Distribution SE Asia: Thailand (north of peninsula), genus. Ear similar to Pipistrellus, but often broader
Laos, Cambodia, Vietnam. Possibly Myanmar. and wider; tragus short, broad, curved slightly
Status Not yet assessed, but likely to be Least forwards (e.g. Fig. 40g). Anterior upper premolar
Concern, as widespread and able to use disturbed usually small, completely intruded inside toothrow,
areas. so that canine and posterior premolar are touching;
sometimes missing. Rostrum short, skull usually
Genus Philetor Externally similar to Pipistrellus, sloping evenly in profile from front to back. Outer
but with short fifth finger (as in P. stenopterus). upper incisor usually only slightly smaller than inner
External genitalia quite elaborate: male has elongate one, with very small supplementary cusps; inner
penis with bristly pad near end of main shaft, upper incisor usually with a small secondary cusp;
followed by a narrow shaft supporting a broadened canine may or may not have a small second cusp,
triangular structure; female has five separate pads depending on the species. Lower molars typically
around vulva. Skull slopes evenly in profile; rostrum with myotodont pattern (Fig. 39). Calcar with only
thickened. Only one upper premolar; inner upper very narrow keel. Baculum short, relatively broad.
incisors long and narrow with two cusps (Fig. 38c). Dental formula 2/3 1/1 2/2 3/3.
Dental formula 2/3 1/1 1/2 3/3.
CHINESE PIPISTRELLE
NARROW-WINGED BROWN BAT Hypsugo pulveratus PLATE 21
Philetor brachypterus PLATE 20 Measurements FA 32–37, T 32–38, E 10.2–13.8.
Measurements FA 30–36, T 30–38, E 13–16. Wt 4.1–6.5g. Skull: ccl 12.4–12.9, mt 4.7–5.3,
Wt 8–13g. Skull: cbl 14.5, mt 4.7 M–M 5.6–6.2
Identification Upperparts dark brown; underparts Identification Upperparts with long, thick fur,
paler and greyer. Fur short and dense. Head dark brownish-black for much of the length, with

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 narrow golden-brown tip; underparts similar, but tragus and shorter canine, less than 1.5× height of
more broadly tipped with paler buff-brown. Ears large premolar.
and wings dark blackish-grey, ears sometimes with Ecology and habitat Has been found in lowland
paler edge; ears broadly rounded; tragus short, evergreen forest in Laos and Vietnam; individuals in
narrow, rounded at tip, curved slightly forwards. Laos were caught near a large river.
Wings moderately broad, fifth metacarpal similar Distribution SE Asia: Laos, Vietnam.
in length to fourth. Both upper incisors similar in Status Not yet assessed.
crown area; anterior upper premolar not especially
reduced, and only slightly smaller than crown BIG-EARED PIPISTRELLE
area of incisors, but completely intruded; upper Hypsugo macrotis PLATE 21
canine without a secondary cusp; anterior lower Measurements FA 33–34. Skull: ccl 11.7–12.1,
premolar about half height of posterior premolar. mt 4.0–4.3, M–M 5.3–5.4
Skull has moderately long, narrow rostrum with Identification Fur of upperparts reddish-brown
distinctly elevated braincase and rostral depression with very dark brown base; underparts similar but
in profile; zygomata robust, with slight dorsal paler. Ears large, broad and triangular, extending
process. Baculum short (2.4mm). Similar species well above head; tragus short, broad and hatchet-
Similar-sized Pipistrellus have narrower ears, shaped (Fig. 40g). Wing membranes translucent
different fur colour, longer penis and baculum; white with a brown tinge. Rostrum short, skull with
Cadorna’s Pipistrelle, H. cadornae, has greatly evenly sloping profile. Anterior upper premolar
reduced anterior upper premolar; Long-toothed tiny, completely displaced inwards, so canine and
Pipistrelle, H. dolichodon, has short broad tragus, second premolar are touching. Similar species
long upper canine. No other pipistrelle in its range has similarly large
Ecology and habitat Has been found roosting in ears and white wings; White-winged Pipistrelle,
caves in limestone areas; forages within relatively Pipistrellus vordermanni, known from Borneo and
open dry dipterocarp forest, as well as highly Billiton Island off SE Sumatra, also has white wings
disturbed areas of mixed scrub and open areas. and large ears, but is smaller (FA c.30, ccl 10.8).
Distribution SE Asia: Myanmar, Thailand, Laos and Ecology and habitat Has been seen feeding low
Vietnam. Also China. over coastal lagoons near mangroves.
Status Least Concern. Distribution SE Asia: Peninsular Malaysia. Also
Sumatra, Bali and adjacent islands.
LONG-TOOTHED PIPISTRELLE Status Data Deficient; very few records; uncertain
Hypsugo dolichodon NOT ILLUSTRATED whether rare or simply not well surveyed.
Measurements FA 35–38, T 35–39, E 13.5–14.0,
HF 5.9–7.0. Wt 6.1–7.3g. Skull: ccl 12.4–13.3, MYANMAR PIPISTRELLE
mt 4.8–5.2 Hypsugo lophurus NOT ILLUSTRATED
Identification Upperparts with long, thick fur, Measurements FA 35, T 39. Skull: ccl 13.1,
blackish-brown, with paler tip; underparts pale dirty mt 4.8, M–M 6.2
white, with basal half of hairs dark brown. Ears and Identification A medium-sized pipistrelle, known
wings dark blackish-grey. Ears relatively short and only from one male specimen. Upperparts with
broadly rounded. Tragus broad and short, slightly long, silky hair, dark brown with blackish base;
curved forwards. Skull evenly sloping in profile. underparts paler also with black base. Large tuft of
Upper canine long, narrow, twice length of second gland-like hairs at base of tail on upper surface of
(large) upper premolar, with no secondary cusps; interfemoral membrane, about 12mm in diameter,
lower canine longer than second lower premolar. with hairs 5–6mm in length (not known whether this
Anterior upper premolar displaced inwards, but is also present in female). Upper incisors both well
relatively large. Calcar on heel relatively long with developed with secondary cusps; anterior upper
narrow keel. Taxonomic notes Newly described premolar moderately reduced, about two-thirds
in 2014. Similar species Chinese Pipistrelle, crown area of inner incisor; anterior lower premolar
H. pulveratus, is similar in size, but has narrower half to two-thirds crown area of second. Skull

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 has moderately shortened rostrum, profile almost tip; tragus short and angled forwards. Dorsal surface
straight but with slight rostral depression; zygomatic of penis has bristly pad near tip. Baculum tiny or
arches have well-developed post-orbital process. rudimentary. Skull has shortened rostrum, broad,
Similar species No other pipistrelle is known to rounded braincase; zygomatic arches with slight
have glandular tuft at base of tail. descending process near third molar; incisors small
Ecology and habitat Only known specimen was and weak, outer incisor about half crown area of
found in a cleared area near a village. inner; upper canine with distinct posterior secondary
Distribution SE Asia: S Myanmar. cusp; anterior upper premolar tiny, completely
Status Data Deficient; only known from one displaced inwards; anterior lower premolar similar in
specimen. height and crown area to posterior lower premolar.
Taxonomic notes Includes H. anthonyi, which was
CADORNA’S PIPISTRELLE previously considered a separate species. Similar
Hypsugo cadornae PLATE 21 species Among other species with shortened
Measurements FA 33–37, T 32–34, E 12.5–14. fifth finger, Narrow-winged Brown Bat, Philetor
Wt. 4.2–7.0g. Skull: ccl 12.7–13.1, mt 4.6–4.7, brachypterus, is similar in shape and dentition,
M–M 5.8–5.9 but lacks anterior upper premolar, is generally
Identification Upperparts reddish-brown with smaller and has different genitalia; Eurasian Noctule,
slightly darker base; underparts similar but tip much Nyctalus noctula, is substantially larger; Narrow-
paler. Ears moderately large, broadly rounded; tragus winged Pipistrelle, Pipistrellus stenopterus, has well-
relatively short, broad, angled slightly forwards. developed anterior premolar.
Wings not especially narrow, fifth metacarpal about Ecology and habitat No information.
equal in length to fourth or third. Both upper Distribution SE Asia: known from Kachin Hills in
incisors similar in crown areas; first upper premolar N Myanmar and N Vietnam. Also India and Nepal.
greatly reduced, a quarter to a half the crown area Status Data Deficient; recent records indicate
of incisors. Skull with straight profile from side, species is more widespread than previously
zygoma robust with dorsal projection in middle. recognised.
Penis relatively small. Similar species Similar-
sized Pipistrellus spp. and Chinese Pipistrelle, Anthony’s Pipistrelle, Hypsugo anthonyi, was
H. pulveratus, have larger anterior upper premolar. included in the first edition of this book, but recent
Ecology and habitat Has been found over rivers research suggests the colour differences are just
and pools in dry dipterocarp forest, as well as individual variation, and it is the same species
disturbed areas and secondary forest. as Hypsugo joffrei.
Distribution SE Asia: N Myanmar, N Thailand, Laos
and N Vietnam. Also NE India. Genus Falsistrellus This genus was, until recently,
Status Least Concern; probably more widespread included within Pipistrellus. Fur is long and woolly.
than current known distribution. Skull has elongate braincase; relatively long, narrow
rostrum; distinct rostral depression in profile.
JOFFRE’S PIPISTRELLE Baculum broad, proximally widened, with deep
Hypsugo joffrei NOT ILLUSTRATED groove on ventral surface, not swollen at tip. Dental
Measurements FA 36–41, T 37–41, E 14. formula 2/3 1/1 2/2 3/3.
Skull: ccl 13.9–14.3, mt 5.1–5.3, M–M 7.0–7.2
Identification A medium-sized pipistrelle with a very CHOCOLATE PIPISTRELLE
short fifth finger, tip reaching only halfway along Falsistrellus affinis PLATE 21
first phalanx of fourth digit. Fur short and velvety, Measurements FA 36–38, T 31–34, E 12.0–13.5.
upperparts glossy dark brown to mid-brown, hairs Skull: ccl 12.9–13.4, mt 5.3, M–M 6.1–6.3
uniformly coloured to base. Underparts paler brown, Identification Fur long and soft, dark blackish-
hairs with darker base, sharply demarcated from brown with narrow pale grey-brown tip, giving a
upperparts on sides. Sides of muzzle swollen and frosted appearance; underparts with greyish-white
thickened. Ears broad and triangular with rounded base, dark tip, sometimes all white near base of tail.

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 Ears broad and rounded; tragus broad, pointed at Pipistrelle, A. societatis, is similar but smaller, with
anterior end (Fig. 40f). Muzzle moderately long and reduced posterior upper molar.
narrow. First upper premolar similar in crown area to Ecology and habitat Most records in region are
incisors, only slightly displaced inwards; first lower from hill forest, 1,300–2,000m.
premolar in toothrow about half size of posterior Distribution SE Asia: Myanmar, Thailand, Vietnam
premolar. Taxonomic notes Description is based and Peninsular Malaysia. Also India, Nepal, S China
on specimens from C Myanmar. Specimens referred and Java.
to this species from India have darker underparts, Status Least Concern.
are larger (FA 38.5–41.5, ccl 13.7–14.5), and have
a slightly different-shaped baculum; the type locality BENOM GILDED PIPISTRELLE
is in N Myanmar in between these two locations; Arielulus societatis PLATE 23
it is not clear whether all are the same species Measurements FA 35–38, T 33–37, E 8.5–9.0.
or, if not, which one is actually F. affinis. Similar Wt 4.3–5.5g. Skull: ccl 14.1, mt 5.2, M–M 6.7
species Other similar-sized pipistrelles generally Identification Very similar to Black Gilded
have shorter fur, shorter, broader muzzle, with dark Pipistrelle, A. circumdatus, with similar fur and head
base to fur on underparts. colour, but shorter forearm, smaller skull and teeth.
Ecology and habitat In Myanmar, has been found Skull has shorter, narrower rostrum, more inflated
roosting in a cave. braincase; teeth less massive, with third upper
Distribution SE Asia: Myanmar. Also India, Nepal, molar reduced, posterior cusp virtually absent.
Sri Lanka, China (Yunnan). Ecology and habitat All records from lowland
Status Least Concern, although poorly known in rainforest. Has been found roosting in a hole in a tree.
region. Distribution SE Asia: Peninsular Malaysia.
Status Vulnerable, owing to limited range and
Genus Arielulus Bats with a distinctive pale edge ongoing loss of its lowland forest habitat.
to ears, black fur with reddish tip on upperparts and
yellowish tip on belly. Ears moderate, oval-shaped Genus Thainycteris Similar to Arielulus and
with rounded top, distinct lobe on side of face sometimes included in that genus. Differs in larger
below tragus; tragus relatively short, angled sharply size, broader, heavier skull and distinctive fur
forwards with blunt tip (Fig. 40d). Outer upper incisor coloration. Skull is heavy and very broad; inner
very small, short and about one-quarter crown area upper incisor large, about half height of canine;
of inner incisor; anterior premolar tiny, completely second upper incisor tiny and hard to see; canine
displaced inwards, sometimes missing (Fig. 38d). without posterior cusp on cutting edge; anterior
Anterior lower premolar about one-third to one- upper premolar very small and completely displaced
quarter crown area of second in toothrow. Baculum inwards, sometimes missing (Fig. 38g); anterior
Y-shaped. Dental formula 2/3 1/1 2–1/2 3/3. lower premolar about half height and two-thirds
surface area of posterior lower premolar. Dental
BLACK GILDED PIPISTRELLE formula 2/3 1/1 2–1/2 3/3.
Arielulus circumdatus PLATE 23
Measurements FA 40–44, T 36. Skull: ccl 14.7– GOLDEN-COLLARED BAT
15.4, mt 6.0–7.2, M–M 7.1–7.5 Thainycteris aureocollaris PLATE 23
Identification Fur of upperparts long and thick with Measurements FA 47–51, T 43–58, E 15–18.
black base and red-orange tip; fur of underparts Wt 12.5–17g
with dark brown base and pale yellow or greyish- Identification Medium-large bat with very distinctive
white tip; pale patch on side of neck behind ear. coloration: upperparts with long fur (10mm), base
Ear moderately sized with distinct lobe at base near black, tip buff to coppery-gold, giving a frosted
eye; ears and tragus dark brown with buff rim. appearance; underparts similar with white to gold
Anterior upper premolar very tiny and displaced tip; broad band of bright buff to orange under chin
inwards (sometimes lacking); canine and second and across throat, forming a collar; irregular streak
premolar touching. Similar species Benom Gilded of buff hair across top of head. Ears, nose and wing

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 membranes dark grey. Ears fleshy, broadly triangular tip. Ear narrow, triangular at top with rounded
with rounded tip; tragus moderately short, broadened tip, thickened rims; tragus narrow, angled slightly
at top half and angled forwards. Similar species forwards, rounded tip; front part of ear and posterior
Gilded pipistrelles, Arielulus spp., are smaller, without lobes covered with short, orangish hair. Ears, muzzle
buff collar and with pale rim to ears; Harlequin Bat, and wing membranes dark brownish-grey. Muzzle
Scotomanes ornatus, has white streak down reddish broad, forehead sloping evenly. Posterior upper molar
back, white patches on shoulder and head. narrow with only one V-shaped cusp. Taxonomic
Ecology and habitat Known from forested areas notes Formerly considered part of E. serotinus from
at 200–1,500m; has been found at forest edges Europe, but Asian populations are now considered a
and over ponds and streams, but regular foraging distinct species. The name E. andersoni has been
areas not known. used for SE Asian forms, but E. pachyomus from India
Distribution SE Asia: Thailand, Laos and Vietnam. appears to be an older name for the same species.
Status Least Concern. Similar species Other Eptesicus are smaller; Asian
Noctule, Nyctalus cf. labiata, has broader ear, short
Genus Eptesicus Similar to Pipistrellus, but with mushroom-shaped tragus, shortened fifth finger;
only one upper premolar (Figs 38i, 41). Outer upper Tomes’s False-serotine, Hesperoptenus tomesi, has
incisor usually small, inner incisor large; upper more rounded ears, broad, rounded facial profile,
canine usually without extra cusps. Ear broadly second upper incisor displaced inwards.
triangular with rounded top; tragus narrow, relatively Ecology and habitat Has been found in wet forest
uniform in width, angled forwards with rounded tip. and dry dipterocarp forest.
Skull robust, forehead sloping to tip. Calcar with Distribution SE Asia: N Myanmar, N Thailand, N Laos
keel, but sometimes fairly narrow. Baculum very and N Vietnam. Also China, India through to Iran.
short, less than 2mm, usually triangular or slightly Status Probably Least Concern, though not yet
Y-shaped. Dental formula 2/3 1/1 1/2 3/3. assessed separately from E. serotinus.

ASIAN SEROTINE THICK-EARED SEROTINE

Eptesicus pachyomus PLATE 23 Eptesicus pachyotis NOT ILLUSTRATED
Measurements FA 53–57, T 53–60, E 18–20. Measurements FA 38–39, T 40–41, E 13–14,
Wt 20–24g. Skull: ccl 18.5–19.5 HF 8–9. Skull: gl 21.2
Identification A large vespertilionid bat; upperparts Identification Upperparts dark brown, underparts
dark brown with pale tip, giving a frosted appearance; slightly paler. Ears triangular, posterior edge of ear
underparts similar but with contrasting whitish hair thick and fleshy from just below middle down to

0 5

Fig. 41 Left upper toothrows from side and below of Eptesicus pachyotis (a) and Cassistrellus dismissus (b).

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 lobes at base of ear. Tragus short, expanded at top, of head loose, sometimes forming fold of skin
curved inwards. Wing membrane inserts at base between ears. Wing membrane attached on side
of toes. Head flat, muzzle short and broad. Second of foot near ankle; calcar with well-developed
upper incisor small, about half height of bicuspid keel. Incisors relatively short, outer upper incisor
first incisor; upper premolar large, about two-thirds about three-quarters height of first incisor; canine
height of canine; anterior lower premolar about half with small cusp on posterior edge. Posterior upper
size of posterior premolar and compressed into molar with relatively well-developed posterior
toothrow; posterior upper molar moderately narrow cusp about half size of anterior cusp (Fig. 41b).
with only trace of second cusp (Fig. 41a). Similar Taxonomic notes Formerly included in the genus
species Similar-sized pipistrelles have an extra Eptesicus, but recognised as a distinct genus in
upper premolar (although it can be small and hard 2017. Similar species Thick-eared Serotine,
to see, especially in Hypsugo spp.), and a broader, E. pachyotis, and similar-sized pipistrelles have
unthickened ear; Surat Helmeted Bat, Cassistrellus wing membranes inserted at base of toes, different
dimissus, has wing membrane inserted on side of dentition.
foot, different dentition. Ecology and habitat Known records are from hilly
Ecology and habitat Unknown. terrain in mixed forest at altitudes of 200–700m.
Distribution SE Asia: N Thailand and possibly N Distribution SE Asia: peninsular Thailand, Laos.
Myanmar. Also NE India, Bangladesh and China. Also Nepal.
Status Least Concern. Although poorly known, Status Data Deficient. Only one specimen each
species is widespread. known from Thailand and Laos, but several from
Nepal; uncertain whether rare or simply rarely caught.
Genus Cassistrellus Previously included in Eptesicus
owing to a similar dental formula, this genus was YOK DON HELMETED BAT
newly described in 2017. Both known species Cassistrellus yokdonensis NOT ILLUSTRATED
have short chestnut-brown fur, paler underneath. Measurements FA 48, T 52, E 16, HF 12. Wt 15g.
Wings narrow with short pointed tip. Muzzle broad Skull: ccl 16.5, mt 6.6, C–C 6.2
with swollen glands on sides. Calcar extending Identification Upperparts with sparse, short fur,
halfway to tail, with narrow keel. Skull is robust clove-brown, not glossy; underparts beige, creamy
and angular with an almost straight rostral profile. on throat. Wings blackish-brown, membranes
Well-developed crests on top of skull look like a attach to distal end of metatarsus near base
helmet and provide the name. Deep, well-defined of toes. Calcar extends half way to tail with no
basisphenoid pits on underside, between auditory conspicuous lobe. Wings relatively narrow, short
bullae. Upper canine with distinct secondary cusp on and pointed. Canines long, with distinct secondary
rear edge. Posterior upper and lower molars relatively cusp. Taxonomic notes Newly described in 2017.
larger than Eptesicus (Fig. 41b). Baculum very small, Similar species Surat Helmeted Bat, C. dimissus,
triangular. Dental formula 2/3 1/1 1/2 3/3. is genetically distinct, with shorter forearm and
toothrow, wing membrane attached on side of foot.
SURAT HELMETED BAT Serotines (Eptesicus spp.) have smaller posterior
Cassistrellus dimissus NOT ILLUSTRATED molar, broader wings, longer fur.
Measurements FA 38–42, T 36–41, E 14–16. Ecology and habitat Only known specimens were
Wt 15g. Skull: ccl 14.6–16.5, mt 5.8–6.4, caught in open dry dipterocarp forest.
C–C 5.6–6.4 Distribution SE Asia: known only from Vietnam.
Identification Medium-sized bat, upperparts with Status Not yet assessed. Only two specimens
short reddish-brown fur, individual hairs with pale known.
base and brown tip; underparts with hairs uniform
pale brown in colour; fur extends in a triangle Genus Ia Very large vespertilionid bat with only
onto base of tail. Ear relatively short and rounded; one recognised species; sometimes included in
tragus short and broad, angled slightly forwards, Pipistrellus. Outer incisor tiny, barely reaching
slightly thickened. Muzzle very broad, skin on top height of cingulum of enlarged inner incisor;

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 anterior premolar very small, completely intruded separating first incisor from canine. Similar species
into toothrow so that canine and posterior premolar Tomes’s False-serotine, H. tomesi, is substantially
are touching; anterior lower premolar about half larger, with second upper incisor displaced inwards
height and crown area of posterior premolar. Calcar (Fig. 38e); Narrow-winged Pipistrelle, Pipistrellus
with little or no lobe. stenopterus, has small upper incisors and an extra
upper premolar.
GREAT EVENING BAT Ecology and habitat Recorded from lowland forest.
Ia io PLATE 22 Has been found roosting in a palm tree.
Measurements FA 71–79, T 68–70, E 27–28. Distribution SE Asia: Peninsular Malaysia. Also
Wt 48–54g. Skull: ccl 25–26, mt 10.5–11.0, Borneo.
M–M 11.0–11.6 Status Data Deficient; known from only a few
Identification Largest vespertilionid bat in region. specimens. May be at risk from loss of lowland
Upperparts dark grey-brown with pale tip, producing rainforest.
a slightly glossy effect; underparts paler dull greyish-
brown. Ears broadly triangular, rounded at top, with LEAST FALSE-SEROTINE
thickened posterior edge; tragus short, broadened Hesperoptenus blanfordi PLATE 22
at top, bluntly pointed to front; muzzle relatively bare Measurements FA 24–29, T 27–32, E 10–12.
of fur. Similar species All other vespertilionid bats Wt 5.0–8.5g
in region are substantially smaller. Identification Very small, with short wings but
Ecology and habitat Known roosts all in limestone relatively heavy body. Upperparts dark brown,
caves. Recorded from a variety of mature forest sometimes with a reddish tone; underparts similar;
types, including wet rainforest, mixed forest and fur sleek and slightly glossy. Base of thumb and
deciduous forest. sole of foot have small, thickened dark brown pads;
Distribution SE Asia: N Myanmar, N Thailand, Laos thumb pad slightly triangular. Second upper incisor
and N Vietnam. Also NE India, Nepal and China. small and displaced inwards (as in Fig. 38e). Head
Status Least Concern. slightly pointed with smoothly sloping profile (Fig.
36f). Similar species Bamboo bats, Tylonycteris
Genus Hesperoptenus Ears fairly short and spp., have a much flatter head with larger round
rounded, tragus slightly hatchet-shaped, pointed thumb pads, inner incisors small and separated
forwards. One species has a small, thickened from canine by second incisor; Thick-thumbed
thumb pad. Only one upper premolar; distinctive Pipistrelle, Glischropus tylopus, has longer fur,
upper incisors: first (inner) incisor very large and more rounded head, white or pink thumb pads and
conical, about half height of canine; second incisor second upper incisor displaced outwards (Fig. 38b);
small, in subgenus Milithronycteris it is displaced Thick-thumbed Myotis, Myotis rosseti, has typical
inwards so that the canine and first incisor are in Myotis-shaped ears, fluffier fur.
contact (Fig. 38e); in subgenus Hesperoptenus it Ecology and habitat Has been found roosting in
separates canine and first incisor. Dental formula entrances of caves in small groups; also known
2/3 1/1 1/2 3/3. from forests away from caves. Forages in open
areas above forests, in gaps and above streams and
DORIA’S FALSE-SEROTINE rivers within forests.
Hesperoptenus doriae PLATE 22 Distribution SE Asia: S Myanmar, Laos, Cambodia,
Measurements FA 38–41. Skull: cbl 13.3–13.6, S Vietnam, Thailand and Peninsular Malaysia. Also
mt 4.9–5.0 Borneo.
Identification Fur uniformly blackish-brown, Status Least Concern.
slightly glossy and fluffy. Head profile rounded. Ear
moderately broad, tragus short, bluntly pointed, TOMES’S FALSE-SEROTINE
angled nearly horizontally forwards. Only one upper Hesperoptenus tomesi PLATE 22
premolar; first upper incisor large and conical; Measurements FA 50–53, T 49–53, E 17–18.
second incisor slightly displaced inwards, but still Wt 30–32g. Skull: cbl 20.4, mt 8.5

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 Identification Upperparts uniform dark blackish- Genus Scotomanes Moderately large vespertilionid
brown; underparts similar. Head rounded, with fur bat, similar to Eptesicus but with only one pair
smoothly curved from forehead to tip of nose; ears of upper incisors, and with distinctive coloration.
broad, somewhat rounded; tragus relatively short, Dental formula 1/3 1/1 1/2 3/3.
angled horizontally forwards and bluntly pointed.
Teeth large with well-developed cusps; first upper HARLEQUIN BAT
incisor large and conical, touching canine; second Scotomanes ornatus PLATE 23
incisor small and displaced inwards (Fig. 38e); Measurements FA 54–61, T 50–60, E 20–22.
only one upper premolar. Similar species Doria’s Wt 20–30g. Skull: ccl 19.4–19.9
False-serotine, H. doriae, is smaller, with second Identification Distinctively coloured large bat;
upper incisor not displaced inwards; house bats, upperparts orange-brown with white patches on top
Scotophilus spp., have paler brown fur with only one of head, shoulders and sides, and white line down
pair of upper incisors. middle of back; underside mixed pattern of black,
Ecology and habitat Has been found over streams orange and white, with black-and-white collar.
in primary lowland rainforest. Wing bones orange-brown, wing membranes dark
Distribution SE Asia: S Thailand and Peninsular grey-brown, tail membrane brown. Ear moderate,
Malaysia. Also Borneo. bluntly triangular; tragus relatively short, narrow,
Status Vulnerable; has probably declined curved forwards with rounded tip. Similar species
considerably in recent years, owing to loss of Golden-collared Bat, Thainycteris aureocollaris, has
lowland rainforest. black upperparts frosted with pale tip, without white
patches, second upper incisor small but usually
TICKELL’S FALSE-SEROTINE present.
Hesperoptenus tickelli PLATE 22 Ecology and habitat Has been found roosting
Measurements FA 49–56, T 47–57, E 16–18. on tree branches, among leaves, but also occurs
Wt 15–21g around limestone caves. Flies in open areas well
Identification Upperparts light yellowish-brown above ground.
to orange; underparts slightly paler; ears, face Distribution SE Asia: N Myanmar, N Thailand, Laos
and wing bones light orange-brown; wing and N Vietnam. Also NE India and S China.
membranes contrasting dark grey to black; base Status Least Concern.
of tail membrane and wing membrane near
body covered with short orange hairs. Ears oval, Genus Scotophilus Ears moderate; tragus very
moderate in length, broadly rounded at top; tragus long, tapering slightly and curved forwards. Skull
short, pointed horizontally forwards. First upper thick and heavy with only one pair of upper incisors,
incisor large and conical, touching canine; second which are large and well developed (Fig. 38h).
incisor small and displaced inwards (as in Fig. Two species in region differ mainly in size. Dental
38e); only one upper premolar. Similar species formula 1/3 1/1 1/2 3/3.
Tomes’s False-serotine, H. tomesi, has thick, dark
brown fur; house bats, Scotophilus spp., have LESSER ASIAN HOUSE BAT
much longer, pointed tragus, dark wing bones, Scotophilus kuhlii PLATE 23
only one pair of upper incisors. Measurements FA 45–52, T 39–53, E 13–15.
Ecology and habitat Occurs in and around Wt 17–26g. Skull: ccl 18.0–18.7
deciduous and semi-deciduous forest. Forages Identification Upperparts brown; underparts paler
high above forest canopy, coming low to drink over yellowish-brown; sometimes with a strong orange
streams and ponds. Probably roosts in hollow trees. tinge. Fur soft and short. Ear with long, narrow tragus
Distribution SE Asia: Myanmar, Thailand, Laos, bent forwards. Only one pair of upper premolars and
Vietnam and Cambodia. Also India, Sri Lanka, Nepal one pair of large conical upper incisors (as in Fig.
and Bhutan. 38h). Similar species Greater Asian House Bat,
Status Least Concern. S. heathii, has forearm longer than 55mm; Harlequin
Bat, Scotomanes ornatus, also has only one pair of

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 upper incisors, but has distinctive pattern of white MAINLAND GREATER BAMBOO BAT
patches on back and neck; Tickell’s False-serotine, Tylonycteris malayana PLATE 22
Hesperoptenus tickelli, is similar in colour, but has Measurements FA 26–30, T 29–36, E 11.
short, broad, hatchet-shaped tragus, second upper Wt 6.5–8.5g. Skull: ccl 12.3–12.7, mt 4.3–4.5
incisor displaced inwards. Identification Upperparts dark brown, hairs with
Ecology and habitat Roosts under house roofs, in pale tip giving glossy effect – looks pale in some
palm tree leaves and in hollow trees. Forages for light; underparts paler greyish-brown to sandy-
aerial insects in open areas, including around towns brown. Fur very smooth and sleek. Flattened disc-
and over forests. like pads at base of thumb and on sole of foot,
Distribution SE Asia: throughout region. Also usually grey, but sometimes fairly pinkish. Lower
Pakistan, India, Sri Lanka, Bangladesh, S China, part of posterior margin of ear thickened. Head
Taiwan, Sumatra, Borneo, Java, Bali and the (skull) and body extremely flattened – can squeeze
Philippines. through a slot less than 5mm wide. Taxonomic
Status Least Concern. notes Formerly called T. robustula, but that species
is now considered restricted to Borneo and Sumatra.
GREATER ASIAN HOUSE BAT Similar species Mainland Lesser Bamboo Bat,
Scotophilus heathii PLATE 23 T. fulvida, overlaps in forearm length but has
Measurements FA 57–69, T 57–72, E 15–19. smaller body and skull; more reddish, fluffy fur,
Wt 29–53g. Skull: ccl 19.3–20.9 thinner ear; Thick-thumbed Pipistrelle, Glischropus
Identification Upperparts dark orange to reddish- tylopus, has more rounded head; smaller pink
brown, hairs with yellowish base; fur smooth and thumb and footpads, longer toes, wing membrane
shiny; underparts paler yellow-brown to orange. inserted at base of toes; Disc-footed Bat, Eudiscopus
Ears and wing membranes contrasting dark brown, denticulus, has large sucker-like discs on feet, but
hairless. Ear medium-large with well-developed only small thumb pads, longer forearm, ears similar
lobe at base; tragus long, narrow, bent forwards. to Myotis; Least False-serotine, Hesperoptenus
One upper premolar. Only one pair of large, conical blanfordi, is less flattened, has thumb pad much
upper incisors (Fig. 38h). Taxonomic notes Shows smaller and triangular, distinctive upper incisors.
considerable size variation across range, and may Ecology and habitat Roosts in the internodes of
represent more than one species. Similar species bamboos, entering through narrow slits created by
Lesser Asian House Bat, S. kuhlii, is smaller, beetles.
forearm less than 55mm; Tickell’s False-serotine, Distribution SE Asia: Myanmar, Thailand, Laos,
Hesperoptenus tickelli, has short, broad, hatchet- Vietnam, Cambodia and Peninsular Malaysia. Also
shaped tragus and two upper incisors. S China, Sumatra, Java, Borneo, Sulawesi and
Ecology and habitat Often roosts under roofs of smaller islands, and the Philippines.
houses. Forages for aerial insects around towns, in Status Least Concern.
open areas and over forests.
Distribution SE Asia: Myanmar, Thailand, Laos, MAINLAND LESSER BAMBOO BAT
Vietnam and Cambodia. Also Sri Lanka, Pakistan, Tylonycteris fulvida PLATE 22
India, S China and Taiwan. Measurements FA 24–28, T 27–29, E 8–9.
Status Least Concern. Wt 3.5–5g. Skull: ccl 10.4–10.6, mt 3.1–3.5
Identification Upperparts brown to reddish-brown;
Genus Tylonycteris Body and skull extremely underparts slightly paler, usually strongly tinged
flattened (Fig. 36d); large brown disc-shaped pad orange. Fur rather short and fluffy. Body and skull
on foot, toes short, wing membrane inserted on extremely flattened with enlarged disc-shaped pads
side of foot; smaller round pad on base of thumb; on thumb and feet. Taxonomic notes Formerly
ears short, bluntly triangular, thickened and fleshy called T. pachypus, but that name is now considered
on front and rear edge; tragus short, narrow at restricted to Sumatra and Borneo. Recent
base, broader and rounded at top. Dental formula genetic studies suggest Chinese and Indochinese
2/3 1/1 1/2 3/3. populations may be a distinct species from those

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 in peninsular Malaysia, but their appearance is very TONKIN GREATER BAMBOO BAT
similar. Similar species Mainland Greater Bamboo Tylonycteris tonkinensis NOT ILLUSTRATED
Bat, T. malayana, overlaps in forearm length, but is Measurements FA 25.1–27.8. Skull: ccl 10.8–
otherwise much larger, with sleeker fur, thickened 11.6, mt 3.8–4.1
lower third of posterior margin of ear. Identification Upperparts with sleek fur, dark brown
Ecology and habitat Roosts in small groups in the with paler reddish-brown base; underside paler.
internodes of bamboos, entering through small slits Looks essentially the same as Mainland Greater
created by beetles. Frequently chooses small live Bamboo Bat, T. malayana, with no described
bamboo stems, while Mainland Greater Bamboo morphological differences, but genetically quite
Bat, T. malayana, prefers larger, often dead stems, distinct in both mitochondrial and nuclear markers.
although both species may sometimes use the Taxonomic notes Newly described in 2017.
same roost holes. Similar species Mainland Greater Bamboo Bat,
Distribution SE Asia: Myanmar, Thailand, Laos, T. malayana, can only be distinguished genetically;
Vietnam, Cambodia and Peninsular Malaysia. Also probably does not overlap in range.
NE India, S China, Sumatra, Java, Borneo and Ecology and habitat Has been found in disturbed
smaller Indonesian islands, and the Philippines. areas with large bamboos.
Status Least Concern. Distribution SE Asia: N Laos, N Vietnam. Probably
adjacent areas of China.
Status Not yet assessed.

Subfamily MURININAE
All the members of this subfamily have the nostrils expanded into short tubes, which protrude on either
side of the muzzle. The ears are rounded, not funnel-shaped; tragus long and pointed. Most species are
placed in the genus Murina. The genus Harpiola, with only one species in the region, looks similar externally,
but differs in having the canines and premolars all the same size. The genus Harpiocephalus is currently
considered to have only one species which is much larger, and has reduced cusps on the molars.
In most species, females average larger than males. When comparing two similar species that differ in
size, large females of the smaller species may overlap with smaller males of the larger species, but can be
separated if the sex is recorded as well as measurements.

Genus Murina Skull has a long palate and large is important in differentiating some species (Figs 42,
rostrum. Two upper and lower premolars, both 43). Dental formula 2/3 1/1 2/2 3/3.
large, well developed and in line in the toothrow.
At the time of the first edition of this book, only The following species have both upper premolars
nine species were recognised in the genus from similar in size (‘cyclotis’ type), with canines generally
the region. Since then, an additional 11 species of much longer than premolars.
Murina have been newly recognised or described
in the region. ROUND-EARED TUBE-NOSED BAT
Most species can be identified by a combination Murina cyclotis PLATE 24
of dental characters, size and fur colour, although Measurements FA 29–34, T 32–44, E 13–17.
a few species can be difficult to distinguish from Wt 4–8g. Skull: ccl 14–16.1, mt 5.2–5.9,
each other except by range or genetics. Among C–C 3.7–4.5, M–M 5.1–6.3
dental characters, the relative size of the premolars Identification Upperparts pale orange mixed with
and canines is a useful character. In some species grey or grey-brown; underparts paler and greyer.
(‘cyclotis’-type dentition), both upper premolars are Tail membrane thinly covered in long reddish hairs.
nearly the same size (e.g. Fig. 42d, e, f), while in other Ear rounded, tragus long, pointed and white. Nostrils
species (‘suilla’ type) the anterior upper premolar is large and tubular. Both upper premolars large. First
about half the height of the posterior premolar (e.g. two lower molars have greatly reduced posterior
Fig. 42a, b, c). The pattern of cusps on the molars section (talonid), less than one-third surface area of

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 a

0 5

Fig. 42 Left upper toothrows from side and below of: Murina eleryi (a), M. feae (b), M. leucogaster (c),
M. rozendaali, (d), M. cyclotis, (e), M. huttoni (f) Harpiola isodon (g) and Harpiocephalus harpia (h).

anterior section (trigonid) (Fig. 43a); first two upper Distribution SE Asia: Myanmar, Thailand north of
molars with V-shaped notch on outer edge (Fig. the peninsula, Laos, Vietnam and Cambodia. Also
42e). Taxonomic notes Recent genetic studies Sri Lanka, India, S China.
show that the larger form from peninsular Malaysia Status Least Concern.
is a separate species, M. peninsularis. Similar
species Hutton’s Tube-nosed Bat, M. huttoni, has PENINSULAR TUBE-NOSED BAT
larger median outer cusp on upper molars (Fig. 42f), Murina peninsularis PLATE 24
relatively larger talonid on lower molars (Fig. 43b); Measurements FA 34–41, T 32–50, E 14–17.
Hairy-winged Bat, Harpiocephalus harpia, is larger, Wt 6–12g. Skull: ccl 14.9–16.9, mt 5.5–6.4,
with heavier teeth and greatly reduced third upper C–C 4.5–5.3, M–M 5.4–6.2
molar (Fig. 42h). Identification Upperparts orange, base of hairs
Ecology and habitat Found in a wide variety of buff or grey; underparts paler buff-orange with pale
forest types, including wet evergreen forest and or grey base to fur. Tail membrane, legs and feet
semi-deciduous forest. covered in long reddish hairs. Ears rounded, tragus

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 on chin or much of underside. Tail membrane,
a legs and feet sparsely covered with orange-brown
hairs. Ears rounded, tragus tapered and white.
Anterior upper premolar similar to or slightly shorter
than posterior premolar; upper canine substantially
heavier and longer than posterior premolar (as
b in Fig. 42e). Taxonomic notes Newly described
in 2013 based on large genetic differences from
M. cyclotis. Similar species Round-eared Tube-
nosed Bat, M. cyclotis, is similar but slightly smaller,
and does not overlap in range. Peninsular Tube-
Fig. 43 Lower right toothrows of Murina cyclotis (a) nosed Bat, M. peninsularis, averages larger within
and M. huttoni (b), showing difference in relative size a sex, with larger skull, more massive teeth, tends
of talonid on molars. to be brighter orange above, underparts more buff
or orange.
long, pointed and white. Nostrils large and tubular. Ecology and habitat Understorey of secondary
Both upper premolars similar in size; upper canine forest and undisturbed primary evergreen forest.
much larger and taller than posterior premolar; first Distribution SE Asia: peninsular Thailand. Also
two lower molars have greatly reduced posterior Nicobar Islands.
section (talonid), less than one-third surface area Status Not yet assessed.
of anterior section (trigonid) (as in Fig. 43a);
first two upper molars with V-shaped notch on FIONA’S TUBE-NOSED BAT
outer edge (as in Fig. 42e). Taxonomic notes Murina fionae PLATE 25
Formerly considered part of M. cyclotis. Similar Measurements FA 34–38.5, T 37–44, E 14–16.5.
species Round-eared Tube-nosed Bat, M. cyclotis, Wt 7.5–12.5g. Skull: ccl 15.9–16.6, mt 6.1–6.5,
is smaller; Guillén’s Tube-nosed Bat, M. guilleni, is C–C 4.5–5, M–M 6.1–6.5
smaller with greyer underparts; Fiona’s Tube-nosed Identification Upperparts pale orange, long fur
Bat, M. fionae, is similar size and shape, but has with pale buff base and orange-brown tip; long pale
pale base to fur and does not overlap in range; guard hairs create a slightly frosted appearance.
Hutton’s Tube-nosed Bat, M. huttoni, is similar but Underparts with uniformly coloured pale hairs, light
with larger median outer cusp on upper molars buff-orange in centre and more whitish near chin.
(Fig. 42f), relatively larger talonid on lower molars Tail membrane, legs, feet and tail covered with
(Fig. 43b); Hairy-winged Bat, Harpiocephalus long orange-brown hairs. Wing membrane inserted
harpia, is larger, with heavier teeth and greatly on side of toe, near base of claw. Ears rounded
reduced third upper molar (Fig. 42h). with moderately long, tapered tragus. Molars with
Ecology and habitat Reported only from reduced talonid on lower molars (as in Fig. 43a),
understorey of tall rainforest. and notch on outer edge of upper molars (as in
Distribution SE Asia: peninsular Thailand and Fig. 42e). Taxonomic notes Newly described in
Malaysia. Also Borneo. 2012. Similar species Round-eared Tube-nosed
Status Not yet assessed. Bat, M. cyclotis, is smaller with darker base to fur;
Peninsular Tube-nosed Bat, M. peninsularis, usually
GUILLÉN’S TUBE-NOSED BAT has some grey bases to fur, posterior upper molar
Murina guilleni NOT ILLUSTRATED narrower than second upper molar (similar width
Measurements FA 32–36, T 28–42, E 11.5–15. in M. fionae) does not overlap in range; Harrison’s
Wt 3–8g. Skull: ccl 14.5–15.8, mt 5.4–5.9, Tube-nosed Bat, M. harrisoni, also has pale base to
C–C 4.1–4.4, M–M 5.4–5.9 fur, but wing membrane inserts at side of foot, half-
Identification Upperparts orange-brown, the hairs way between base and claw, and has molars more
with dark grey base; underparts usually grey or like Hutton’s Tube-nosed Bat, M. huttoni.
greyish-white; sometimes with strong orange cast Ecology and habitat Has been caught in wet

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 evergreen hill forest at of altitude 830–1,150m in hairs. Wing membrane attaches to foot at side of
Vietnam and Laos, and in semi-deciduous forest at toe, halfway between base and tip. Anterior upper
250m in Cambodia. premolar similar in height to posterior molar, canine
Distribution SE Asia: Laos, Vietnam, Cambodia. about twice height of premolars; second incisor
Status Not yet assessed. about half height of anterior incisor; lower molars
with relatively well-developed talonid (similar to
HUTTON’S TUBE-NOSED BAT but not quite as large as Fig. 43b). Females with
Murina huttoni PLATE 24 larger skull and more robust rostrum than males.
Measurements FA 32–37, T 31–42, E 15–17.5. Taxonomic notes Includes M. tiensa, which is
Wt 5–11g. Skull: ccl 15.0–16.5, mt 5.6–6.3, now considered to be the same species. Similar
C–C 4.2–5.1, M–M 5.6–5.9 species Hutton’s Tube-nosed Bat, M. huttoni, has
Identification Upperparts greyish-brown to reddish- dark base to fur; Fiona’s Tube-nosed Bat, M. fionae,
brown, hairs with dark grey-brown base, pale middle has reduced talonids on lower molars and deep
and brown to reddish-brown tip; underparts similar, notch on side of upper molars, wing membrane
but tips paler. Ears long, reaching 3–4mm past inserted closer to base of claw; Greater Tube-nosed
nostrils if folded forwards; tragus long and white. Bat, M. leucogaster, has relatively smaller anterior
Wing membrane attached to side of toe near claw. upper premolar, shorter canine.
Both upper premolars similar in size, about twice Ecology and habitat Has been found flying over a
as tall as molars, but about half height of canine; stream in disturbed semi-deciduous lowland forest.
anterior lower premolar about 20% smaller than Distribution SE Asia: Myanmar, Thailand,
posterior premolar. Posterior section of lower molars Cambodia, Laos, Vietnam. Also China.
(talonid) nearly as large as anterior section (trigonid) Status Least Concern, although only known from a
(Fig. 43b); first two upper molars have moderately limited number of specimens.
well-developed outer median cusp, so there is only
a shallow notch on outside edge (Fig. 42f). Similar BRONZED TUBE-NOSED BAT
species Round-eared Tube-nosed Bat, M. cyclotis, Murina aenea PLATE 24
Fiona’s Tube-nosed Bat, M. fionae, and Peninsular Measurements FA 34–38, T 35–45, E 13.5–15.5.
Tube-nosed Bat, M. peninsularis, have shorter Wt 6–8.5g. Skull: cbl 15.3–16.1, mt 5.7–6.0,
ears and different-shaped molars (Figs 42, 43); C–C 4.7–4.8.
Harrison’s Tube-nosed Bat, M. harrisoni, has wing Identification Fur of upperparts brown mixed
membrane inserted on side of foot, pale base to fur, with shiny gold-orange; short hairs with bands
lower molars with slightly smaller talonids. of buff and dark brown, long hairs dark brown
Ecology and habitat Most records from hill forest with shiny golden or bronze tip; underparts buffy,
at altitudes of 400–1,500m. individual hairs with dark brown base and pale
Distribution SE Asia: N Myanmar, N Thailand, Laos, buff-brown or orange-brown tip. Nostrils long and
Vietnam and Peninsular Malaysia. Also Pakistan, tubular. Lower molars with reduced talonid, similar
N India, Tibet and S China. to M. cyclotis (as in Fig. 43a), but less extreme.
Status Least Concern. Poorly known in Peninsular Similar species Rozendaal’s Tube-nosed Bat,
Malaysia, as only one record exists. M. rozendaali, is smaller and paler, back fur
with yellow tip, underparts white with pale base;
HARRISON’S TUBE-NOSED BAT Golden Tube-nosed Bat, M. aurata, is smaller
Murina harrisoni PLATE 25 with whitish underparts, smaller teeth, especially
Measurements FA 34–40, T 36–45, E 13.5–18.5. anterior premolars; Round-eared Tube-nosed Bat,
Wt 8.5–12g. Skull: ccl 15.9–17, mt 5.8–7.1, M. cyclotis, has similar dentition, but fur is reddish-
C–C 4.7–5.3, M–M 5.6–6.5 brown without gilded tips; groove-toothed bats,
Identification Upperparts uniform reddish-brown, Phoniscus spp., have similar colour but less tubular
hairs only slightly darker at tip; underparts white, nostrils, very different dentition.
individual hairs pale to base. Upper surface of tail Ecology and habitat Has been found in lowland
membrane uniformly covered in reddish-brown dipterocarp forest and hill forest.

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 Distribution SE Asia: Peninsular Malaysia and Bat, M. cyclotis, has shorter dark base to fur, greatly
adjacent S Thailand. Also Borneo. reduced mesostyle on upper molars and talonids on
Status Vulnerable due to loss of lowland rainforest. lower molars; Hutton’s Tube-nosed Bat, M. huttoni,
is substantially larger.
ROZENDAAL’S TUBE-NOSED BAT Ecology and habitat Has been caught in recently
Murina rozendaali PLATE 24 logged evergreen forest, and patches of mixed
Measurements FA 28–32, T 30–35, E 13–14. evergreen and deciduous forest at altitudes of
Wt 3.8–4.8g. Skull: cbl 13.7–13.8, mt 5.2–5.3, 500–1,100m.
C–C 3.8–4.1 Distribution SE Asia: Laos, Vietnam.
Identification Fur of upperparts with dark brown Status Not yet assessed.
base and shining yellow or golden-brown tip;
underparts white to the base with a buffy or The following species have the anterior upper
yellowish tinge. Nostrils tubular. Both upper premolar distinctly smaller than the posterior molar,
premolars similar in size; canine long, about twice and the canine similar in size to posterior molar
height of premolars (Fig. 42d); lower molars with (‘suilla-type’ dentition).
well-developed talonids (as in Fig. 43b). Similar
species Bronzed Tube-nosed Bat, M. aenea, is BROWN TUBE-NOSED BAT
larger and darker, with dark base to belly fur; Bala Murina suilla PLATE 24
Tube-nosed Bat, M. balaensis, has dark base to fur Measurements FA 27–32, T 26–35, E 12.5–13.5.
of underparts, anterior upper premolar small, half Wt 3–5g. Skull: cbl 12.6–13.3, mt 4.7–4.9,
size of posterior premolar, shorter canine (as in Fig. C–C 3.6–3.7
42a); Brown Tube-nosed Bat, M. suilla, has small Identification Upperparts brown, sometimes with
anterior upper premolar and short canine (as in Fig. orange or greyish tone; individual hairs with grey
42b) and lacks golden tips to fur. base, paler sub-terminal band, then brown tip;
Ecology and habitat Found flying low over streams underparts greyish-white, the hairs with grey base,
in disturbed lowland dipterocarp forest. especially on flanks. Tubular nostrils. Anterior upper
Distribution SE Asia: peninsular Thailand and premolar half height of posterior premolar (as in
Malaysia. Also Borneo. Fig. 42b). Similar species Rozendaal’s Tube-
Status Vulnerable, owing to low densities and loss nosed Bat, M. rozendaali, has pale yellowish tips
of lowland rainforest. to back fur, is slightly larger and has relatively
large anterior premolar and canine (Fig. 42d); Bala
ANNAMITE TUBE-NOSED BAT Tube-nosed Bat, M. balaensis, has guard hairs with
Murina annamitica NOT ILLUSTRATED long golden tip, whiter underparts, cusp on base of
Measurements FA 29–34, T 32–40, E 12–15. upper canine; Whitehead’s Woolly Bat, Kerivoula
Wt 4.5–8g. Skull: ccl 13.6–15.1, mt 5.2–5.7, whiteheadi, has small nostrils, funnel-shaped ears;
C–C 3.7–4.3, M–M 5.2–5.8 very different dentition (Fig. 45).
Identification Upperparts with fluffy orange-brown Ecology and habitat Lowland rainforest, apparently
fur overall; individual hairs with dark grey base tolerant of moderate disturbance.
(40%), buffy middle and orange-brown to brown tip; Distribution SE Asia: peninsular Thailand and
underparts greyish-buff, individual hairs with dark Malaysia. Also Sumatra, Java and Borneo.
grey base (60–70%) and buffy tip. Tail membrane, Status Least Concern, but has probably declined
legs and feet covered with long orange-brown hairs. due to loss of lowland rainforest.
Ears rounded, tragus long and tapered. Anterior
upper premolar similar height to posterior molar, FEA’S TUBE-NOSED BAT
and about half height of canine (as in Fig. 42f). Murina feae PLATE 24
Upper molars with well-developd mesostyle (middle Measurements FA 27–34, T 30–40, E 12–15.
cusp); lower molars with well-developed talonids (as Wt 3.0–5.5g. Skull: ccl 12.8–14.3, mt 4.8–5.4,
in Fig 43b). Taxonomic notes Newly described in C–C 3.3–4.0, M–M 5.0–5.7
2012. Similar species Round-eared Tube-nosed Identification Upperparts appear dark brown

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 mottled with buff; individual hairs with dark grey- secondary forest and mature montane forest, at
brown base (about 60–70% of length), then buffy- elevations of 400–1,600m.
grey, then dark grey-brown; long guard hairs have Distribution SE Asia: Vietnam.
dark base and white tip, giving an appearance of Status Not yet assessed.
four bands of colour with frosted tips; underparts
greyish-white, fur with long blackish-brown base WALSTON’S TUBE-NOSED BAT
and white tip. Interfemoral membrane sparsely Murina walstoni PLATE 25
haired with grey-brown hairs. Anterior premolar Measurements FA 30–35, T 27–32, E 12.5–14.
small, about half height of posterior premolar, Wt 4.5–7.5g. Skull: ccl 13.3–13.7, mt 5.1–5.7,
canine only slightly longer than posterior molar C–C 3.8–4.3, M–M 5.3–5.9
(Fig. 42b). Posterior lobes of lower molars Identification Upperparts with long, woolly brown
relatively small. Taxonomic notes Formerly called to orange-brown fur; individual hairs white for
M. tubinaris, which is now considered a separate about 70% of length with brown to orange-brown
species not found in the region. Has also been tip; scattered long guard hairs pale to tip. Underpart
called M. cineracea, but M. feae is an older name fur all white, sometimes with greyish or buffy tinge.
for the same species. Similar species Brown Upper surface of wing and tail membrane largely
Tube-nosed Bat, M. suilla, is browner with less naked except along base of tail; forearm with
distinct bands of colour; Beelzebub Tube-nosed Bat, only short, sparse hairs. Ear rounded with a small
M. beelzebub, is similar but has blacker fur without notch (emargination) on posterior edge. Anterior
brown tinge, averages larger. upper premolar small, about half height and basal
Ecology and habitat Most records from evergreen area of posterior premolar; upper canine narrow,
hill forest. slightly longer than posterior premolar (as in Fig.
Distribution SE Asia: Myanmar, N Thailand, Laos 42b). Taxonomic notes Newly described in 2011.
and Vietnam. Similar species Other small Murina with ‘suilla’-
Status Not yet assessed separately from M. type dentition have dark base to fur, and much more
tubinaris. Apparently widespread and relatively extensive fur on tail and wing membranes.
common, but has probably declined owing to loss Ecology and habitat Has been caught in relatively
of evergreen forest. open seasonally dry forest and disturbed areas,
unlike most other Murina, which primarily occur in
BEELZEBUB TUBE-NOSED BAT mature forest.
Murina beelzebub PLATE 25 Distribution SE Asia: N Thailand, Laos, Vietnam.
Measurements FA 34–38, T 33–45, E 13–14. Status Not yet assessed.
Wt 3–5.5g. Skull: ccl 14.5–15.0, mt 5.3–5.7,
C–C 3.8–4.0, M–M 5.2–5.8 GILDED TUBE-NOSED BAT
Identification Upperparts overall very dark; fur with Murina eleryi PLATE 24
blackish base (about 80% of length), then a light Measurements FA 27–31, T 26–32, E 11.5–13.5.
grey band with a dark blackish tip; scattered long Wt 3–5.5g. Skull: ccl 12.2–13.2, mt 4.5–4.9,
guard hairs are all pale grey; underparts hairs with C–C 3.2–3.6, M–M 4.6–5.2
dark brownish-black base (about 70% of length) Identification Upperparts yellowish-brown,
and white tip. Upper surface of tail membrane, legs individual hairs banded with blackish base, then
and feet covered in dark brown hairs; underside of buff-white middle, then orange-brown to dark
tail membrane covered in whitish hairs. Dentition brown. Long guard hairs have pale golden tip;
of ‘suilla’ type, with small anterior upper premolar, underparts extensive black base with white tip. Tail
upper canine similar height to posterior premolar membrane, feet and legs with scattered yellow-
(as in Fig. 42b). Taxonomic notes Newly described brown hairs. Ears small and rounded, tragus long,
in 2011. Similar species Fea’s Tube-nosed Bat, straight, pointed white. Braincase very tall and
M. feae, is paler and browner overall, with brownish rounded, anterior upper premolar small, half height
tips to fur, smaller skull. of posterior premolar; upper canine slightly longer
Ecology and habitat Has been caught in disturbed than posterior premolar; lower canine shorter than

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 posterior premolar (Fig. 42a). Taxonomic notes DA LAT TUBE-NOSED BAT
Newly described in 2011 as a species distinct from Murina harpioloides PLATE 25
M. aurata, which does not occur in SE Asia. Similar Measurements FA 29–30, T 27–30, E 12–13.
species Fea’s Tube-nosed Bat, M. feae, is greyer Wt 4–4.5g. Skull: ccl 12.3, mt 4.7, C–C 3.4,
with less domed braincase; Bala Tube-nosed Bat, M–M 4.9
M. balaensis, is very similar, but has shorter upper Identification Very small bat. Upperparts appear
canine with slightly larger cusp, greyer underparts, bicoloured dark brown and golden; underfur
does not overlap in range; Golden-haired Tube- uniformly dark brown with slightly paler tip, guard
nosed Bat, M. chrysochaetes, has yellow-tipped hairs dark brown at base (50%) with buff band then
guard hairs on belly, shorter, more slender upper brown band then long orange-gold tip; underparts
canine, more domed braincase. dark brown at base, tipped with pale greyish-white.
Ecology and habitat Evergreen hill forest. Entire tail membrane and wing membrane near body
Distribution SE Asia: N Myanmar, N Thailand, Laos covered with long gold-tipped hairs; forearm and
and Vietnam. Also Nepal, NE India and S China. legs covered with short gold hairs. Ears wide and
Status Least Concern. rounded with small notch (emargination) on posterior
edge; muzzle and ears darkly pigmented. Anterior
BALA TUBE-NOSED BAT upper premolar very small, less than half height
Murina balaensis NOT ILLUSTRATED and about one-quarter crown area of posterior
Measurements FA 28–30, T 30–34, E 12–13.5. premolar; upper canine small and slender, shorter
Wt 3–4g. Skull: ccl 12.3–13, mt 4.7–4.9, than posterior premolar. Lower canine shorter than
C–C 3.4–3.6, M–M 4.9–5.0 posterior premolar, with well-developed cusp on
Identification Upperparts golden-brown overall; anterior edge. Taxonomic notes Newly described in
fur with grey base (40%), then pale buff gradually 2008. Similar species Golden-haired Tube-nosed
darkening to orange-brown or dark brown at tip; Bat, M. chrysochaetes, is similar, but short hairs
long guard hairs on back and head shiny golden; on dorsal fur are banded, underside has gold not
underparts white, hairs with blackish-grey base whitish tips, ear does not have notch; Even-toothed
(40%) and white tip. Tail membrane, forearm and Tube-nosed Bat, Harpiola isodon, is similar colour
legs sparsely covered with golden hairs; underside but has very large anterior premolars.
of tail with white hairs. Ears rounded, light brown; Ecology and habitat Known only from Da Lat
tragus tapered and white. Anterior upper premolar plateau, in mixed broadleaf and coniferous forest at
small, half height of posterior premolar; upper 1450–1,800m altitude.
canine similar length to poster premolar (as in Fig. Distribution SE Asia: Vietnam.
42a); canine with distinct cusp on base. Taxonomic Status Not yet assessed.
notes Newly described in 2013. Similar species
Gilded Tube-nosed Bat, M. eleryi, is very similar but GOLDEN-HAIRED TUBE-NOSED BAT
has smaller cusp on canine, higher braincase, does Murina chrysochaetes PLATE 25
not overlap in range; Brown Tube-nosed Bat, M. Measurements FA 28–30, T 24–26, E 12–12.5.
suilla, is browner, without golden guard hairs, and Wt 4–5g. Skull: ccl 12.8, mt 4.7–5.4, C–C 3.6,
no cusp on canine; Rozendaal’s Tube-nosed Bat, M–M 4.9
M. rozendaali, has white base to fur on belly and Identification Upperparts with long gold and black
‘cyclotis’-type dentition. fur; individual hairs with black base (65%), then a
Ecology and habitat All records are from lowland yellow band then a dark brown tip; interspersed with
rainforest. long guard hairs with black base (60%) and long
Distribution SE Asia: S peninsular Thailand and gold tip; underpart hairs largely black, interspersed
Malaysia. with long gold-tipped guard hairs except around
Status Assessed as Critically Endangered, but chin and base of tail where guard hairs have white
additional records from Malaysia suggest may be tip. Black hairs around edge of face. Upperside of
more widely distributed than previously thought. tail membrane, legs and feet sparsely covered with
gold-tipped hairs. Ears relatively short and rounded

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 with no notch, tragus tapered and white. Anterior dark grey base (30%), rest pale brownish-grey.
upper premolar small, half height of posterior Blackish hairs around eye and muzzle, white
premolar; upper canine slender and slightly shorter collar under chin. Upper surface of tail membrane,
than posterior premolar; lower canine similar height legs and feet covered with long yellow-brown
to posterior premolar with distinct small anterior hairs. Ears relatively long, with slight notch on
cusp. Mesostyle (middle cusp) on upper molars posterior edge. Anterior premolars small, half or
reduced. Taxonomic notes Newly described in less height of corresponding posterior premolar;
2011. Similar species Gilded Tube-nosed Bat, upper canine substantially longer than posterior
M. eleryi, has white tips to belly fur, upper canine premolar. Taxonomic notes Newly described in
longer than premolar, less reduced mesostyles 2015. Similar species Other tube-nosed bats
on upper molars; Da Lat Tube-nosed Bat, with ‘suilla’-type dentition differ in colour (are not
M. harpioloides, has slightly different fur colour, brownish-grey, or lack golden guard hairs), have
small notch on posterior edge of ear. shorter ears and usually shorter canines not much
Ecology and habitat All records are from montane longer than posterior premolars.
broadleaved forest at altitudes of 950–1,950m. Ecology and habitat Only known specimen was
Distribution SE Asia: N Vietnam. Also S. China. caught along a stream in relatively undisturbed
Status Not yet assessed. montane forest at an altitude of 1,800m.
Distribution SE Asia: Vietnam.
GREATER TUBE-NOSED BAT Status Not yet assessed. Known from only one
Murina leucogaster PLATE 24 specimen.
Measurements FA 42, T 40, E 14. Wt 9g.
Skull: ccl 16.5, mt 6.2, M–M 6.5, C–C 4.5 LORELIE’S TUBE-NOSED BAT
Identification Largest Murina in region; upperparts Murina lorelieae PLATE 25
reddish-brown, hairs with dark base; underparts Measurements FA 33–35.5, T 31–41, E 14–16.
pale yellowish-white, hairs with dark base only on Wt 4–6.5g. Skull: ccl 13.5–14.1, mt 5.2–5.6,
sides. Anterior upper and lower premolars small, C–C 3.7–3.9, M–M 5.3–5.6
about half height and surface area of corresponding Identification Upperparts long, woolly, reddish-
posterior premolars; canine relatively low, no more brown overall; individual hairs with dark grey base
than 30% taller than posterior premolar (Fig. 42c). (40%), a buff band and reddish-brown tip, mixed
Similar species Other large Murina have anterior with uniformly red long guard hairs; underparts with
premolar nearly same size as posterior premolar, dark grey base (70%) and greyish-white tip. Short
less contrasting colours of upper and underparts. dark hairs around mouth and eyes. Basal half of tail
Ecology and habitat Has been found over a stream membrane, legs and feet covered in long reddish
in disturbed secondary forest. hairs. Ears long, rounded, with no notch. Anterior
Distribution SE Asia: N Vietnam. Also NE India, upper premolar larger than most ‘suilla’-type
Nepal and S China. bats, 60–70% height of posterior premolar; upper
Status Least Concern, but poorly known in region canine slightly longer than posterior premolar; lower
with only one confirmed record – previously canine similar height to posterior premolar. Middle
published records from Thailand were based on cusp of upper molars well developed. Taxonomic
misidentified specimens. notes Newly described in 2011. Similar species
Annamite Tube-nosed Bat, Murina annamitica, is
KONTUM TUBE-NOSED BAT similar in external appearance, but has ‘cyclotis’-
Murina kontumensis PLATE 25 type dentition, with much longer upper and lower
Measurements FA 32, T 38, E 18.5. Wt 5g. canines, both premolars more similar in size.
Skull: ccl 13.3, mt 5.6, C–C 3.7, M–M 5.3 Ecology and habitat Has been found in the
Identification Upperparts brownish-grey overall, understorey of wet montane evergreen forest at
hairs with dark brown base, light brown middle, elevations of 1,100–1,700m in Vietnam.
brownish-grey tip; guard hairs with yellow tip, Distribution SE Asia: Vietnam. Also China.
denser on head, sparse on back. Underpart hairs Status Not yet assessed.

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 Genus Harpiola Similar to Murina, with tubular Genus Harpiocephalus Similar to Murina, with
nostrils and the same dental formula, but canine tubular nostrils and the same dental formula, but
and premolars in both upper and lower jaws similar posterior upper molar very small, sometimes lacking
in size, incisors relatively large (Fig. 42g). Dental (Fig. 42h); anterior upper molars with reduced
formula 2/3 1/1 2/2 3/3. cusps except the central one, thus resembling
the premolars; canines low and massive; rostrum
EVEN-TOOTHED TUBE-NOSED BAT slightly shortened. Dental formula 2/3 1/1 2/2 3/3.
Harpiola isodon PLATE 25
Measurements FA 31–36, T 25–35, E 12–13. HAIRY-WINGED BAT
Skull: ccl 13.7, mt 5–5.6, C–C 3.6–4, Harpiocephalus harpia PLATE 26
M–M 4.9–5.5 Measurements FA 45–51, T 45–55, E 16–19.
Identification Upperparts with long woolly fur, a mix Wt 12–23g. Skull: cbl 18.5–21.1, mt 6.6–7.4,
of dark brown and gold overall; individual hairs of C–C 6.0–7.8, M–M 6.9–7.7
underfur with dark base, yellow band in middle and Identification Colour of upperparts varies from
brown tip; long guard hairs with dark brown base and bright orange with dark base (individuals from
shiny gold tip; underpart fur shorter, dark brown at Malaysian rainforests) to dull orange-brown with
base and light brown at tip. Tail membrane, above and grey base. Underparts similar but slightly greyer.
below, legs, tail and forearm covered with fur, golden Nostrils elongate and tubular. Ears rounded with long,
above, greyer below. Ears rounded, moderately long, pointed tragus. Forearm, much of wing membrane
with medium-length tragus reaching to notch in ear. and interfemoral membrane covered in short orange
In both upper and lower toothrows, anterior premolar hairs. Two upper premolars similar in size; third
is larger than second premolar and similar in size to upper molar tiny, sometimes lacking (Fig. 42h).
canine (Fig. 42g); upper incisors large, similar to or Taxonomic notes This was formerly treated as two
exceeding height of posterior premolar. Lower canine species, with the form H. mordax differing in larger
distinctly bifid, with well-developed anterior cusp. skull and more massive teeth, but recent genetic
Taxonomic notes Newly described in 2006. Similar analyses suggest there is only one species in the
species All other similar tube-nosed bats (Murina region, with females much larger than males. Similar
spp.) have anterior premolar smaller than posterior species Tube-nosed bats, Murina spp., are smaller,
premolar, and small upper incisors. with a relatively larger third upper molar (Fig. 42a–f).
Ecology and habitat In Vietnam, has been caught Ecology and habitat Understorey of lowland
over a small stream in montane moss forest at dipterocarp forest, as well as disturbed areas.
an altitude of 2,250m. In Taiwan, was reported Distribution SE Asia: Myanmar, Thailand, Peninsular
from montane forest (altitude 1,000–2,400m) in Malaysia, Laos and Vietnam. Also India, Nepal,
coniferous and broadleaf forest. S China, Taiwan, Sumatra, Java, Borneo and Maluku.
Distribution SE Asia: N Vietnam. Also Taiwan. Status Least Concern, but has probably declined
Status Data Deficient. Only known from one due to loss of lowland forest.
specimen in Vietnam and several from Taiwan.

Subfamily KERIVOULINAE
This subfamily includes two genera that differ in fur colour, the colour of the tragus and the presence or
absence of a large groove in the upper canines.

Genus Kerivoula Ears funnel-like, with a fold at in any of the cheek teeth (Figs 44, 45). Dental
the back and a large flap on the outside; tragus formula 2/3 1/1 3/3 3/3. Myotis spp., have same
long, narrow and pointed. Fur long and woolly, dental formula, but different-shaped ears with a
often covering much of the face. Nostrils small. shorter, bluntly pointed tragus, shorter rostrum
Skull usually has a high domed braincase (with and reduced second premolar (Fig. 37). Recent
some exceptions) and a long rostrum (Figs 36j, 44). genetic analyses suggest that some taxa, such as
Teeth relatively unspecialised, with no reduction Kerivoula hardwickii and the Kerivoula papillosa/

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 a a

b b

Fig. 44 Skull profiles of Kerivoula papillosa (a) Fig. 45 Upper left toothrows of Kerivoula minuta (a) and
and Kerivoula kachinensis (b). Kerivoula whiteheadi (b).

lenis complex, include more species than are pointed tragus. Braincase high and domed (as in
currently recognised. Fig. 44a). Similar species Papillose Woolly Bat,
K. papillosa, has larger skull, longer toothrow;
PAPILLOSE WOOLLY BAT Titania’s Woolly Bat, K. titania, is smaller,
Kerivoula papillosa PLATE 27 with a shorter, flatter skull; Kachin Woolly Bat,
Measurements FA 39–49, T 49–56, E 14–17, K. kachinensis, has flatter skull.
Wt 6–13g. Skull: ccl 15.4–17.1, mt 7.1–7.8, Ecology and habitat Understorey of forest,
M–M 6.7–7.1 including both wet and dry forest.
Identification Fur long and woolly. Upperparts brown Distribution SE Asia: Peninsular Malaysia. Also
to buffy-brown; underparts paler. Ears funnel-shaped India and Borneo.
with a long, pointed tragus. Three upper and lower Status Least Concern.
premolars, all of which are well developed. Braincase
high, with domed forehead (Fig. 44a). Similar KACHIN WOOLLY BAT
species Indian Woolly Bat, K. lenis, overlaps in Kerivoula kachinensis NOT ILLUSTRATED
forearm length, but has shorter skull and toothrow; Measurements FA 41–43, T 56–61, E 14–16.
Kachin Woolly Bat, K. kachinensis, has flatter skull. Wt 6.5–9.1g. Skull: ccl 15.5, mt 6.8, M–M 6.4
Ecology and habitat Forages in understorey of Identification Fur long and woolly; upperparts grey-
lowland rainforest. Roosts in small holes within live brown with dark grey base to fur; underparts slightly
trees. Has also been found inside bamboo stems. paler. Ears funnel-shaped with a long, pointed
Distribution SE Asia: Thailand, Laos, N Vietnam, tragus. Three upper and lower premolars, all well
Cambodia and Peninsular Malaysia. Also Sumatra, developed. Braincase relatively flat in profile (Fig.
Java, Borneo and Sulawesi. 44b). Similar species Other large Kerivoula spp.
Status Least Concern. have high domed braincase (Fig. 44a), producing
an abruptly rising forehead, which can be felt in the
INDIAN WOOLLY BAT live bat if the fur is gently pressed down.
Kerivoula lenis NOT ILLUSTRATED Ecology and habitat Has been found in evergreen
Measurements FA 37–41. Skull: ccl 14.5–15.1, and mixed deciduous forest.
mt 6.6–6.8, M–M 6.0–6.6 Distribution SE Asia: Myanmar, Thailand, Laos and
Identification Upperparts brown to buffy-brown, N Vietnam.
underparts paler. Ears funnel-shaped with a long, Status Least Concern.

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 HARDWICKE’S WOOLLY BAT Distribution SE Asia: Myanmar, Thailand, Laos,
Kerivoula hardwickii PLATE 27 Vietnam.
Measurements FA 29–34, T 40–50, E 12–14. Status Not yet assessed.
Wt 3.5–6.0g. Skull: ccl 12.4–13.6, mt 5.2–5.9
Identification Fur long and woolly; upperparts INDOCHINESE WOOLLY BAT
grey-brown with dark grey base to fur (about Kerivoula dongduongana NOT ILLUSTRATED
50%) followed by a pale band then a browner Measurements FA 30–33. Wt. 4.5g.
tip; underparts paler and greyer. Ears moderately Skull: ccl 11.7–12.8, mt 5.1–5.5
large. Braincase elevated (similar profile to Fig. Identification Fur long and woolly; upperparts dark
44a). Taxonomic notes Recent morphological grey; underparts paler grey. Ears pale with dark
and genetic analyses identified Kerivoula depressa, edge. Braincase relatively flat (similar in profile
K. furva and K. dongduongana as distinct species to Fig. 44b). Taxonomic notes Newly described
from K. hardickii. Similar species Titania’s Woolly in 2018. Similar species Miller’s Woolly Bat,
Bat, K. titania, is greyer and slightly larger; Miller’s K. depressa, and Dark Woolly Bat, K. furva, also
Woolly Bat, K. depressa, Indochinese Woolly Bat, have similarly flat braincases and are only reliably
K. dongduongana, and Dark Woolly Bat, K. furva, distinguished by genetic analysis.
all have a flat braincase; Clear-winged Woolly Bat, Ecology and habitat Found in a range of forested
K. pellucida, has pale orange fur with pale base, habitats in the area around the Annamite mountains.
long, narrow ears; Small Woolly Bat, K. intermedia, Distribution SE Asia: Laos, Cambodia, Vietnam.
has orange-brown fur with short ears. Status Not yet assessed.
Ecology and habitat Understorey of various forest
types, including evergreen and deciduous. Has been DARK WOOLLY BAT
found roosting in hollow trees and among clumps Kerivoula furva NOT ILLUSTRATED
of dead leaves. Measurements FA 30.5–36.5, Tibia 15.8–18.6.
Distribution SE Asia: Myanmar, Thailand, Laos, Skull: ccl 12.2–13.8, mt 5.3–6.1
Vietnam, Cambodia and Peninsular Malaysia. Also Identification Fur long and woolly; upperparts dark
Sri Lanka, India, S China, Sumatra, Java, Sulawesi, grey-brown to blackish, individual hairs usually
the Philippines and smaller Indonesian islands. uniformly dark from base to tip; underparts paler
Status Least Concern. grey or grey-brown, fur with dark grey base, pale
grey tip (sides) or brownish tip (middle). Ears pale
MILLER’S WOOLLY BAT with dark edge. Braincase relatively flat (similar in
Kerivoula depressa NOT ILLUSTRATED profile to Fig. 44b). Ratio of tibia/forearm length
Measurements FA 30.5–34, Tibia 15.5–16. 0.48–0.55. Taxonomic notes Newly described in
Skull: ccl 11.8–12.8, mt 4.9–5.6 2017. Similar species Hardwicke’s Woolly Bat,
Identification Upperparts grey-brown to buffy- K. hardwickii, has more domed braincase; Titania’s
brown, with dark-grey base to fur; underparts paler. Woolly Bat, K. titania, has proportionately longer
Braincase relatively flattened, similar in profile to Fig. tibia; Miller’s Woolly Bat, K. depressa, usually has
44b. Taxonomic notes Recently recognised as a fur with contrasting dark base, paler middle and tip;
distinct species from K. hardwickii. Similar species Indochinese Woolly Bat, K. dongduongana, is only
Kachin Woolly Bat, K. kachinensis, and Titania’s reliably distinguished by genetics.
Woolly Bat, K. titania, also have flattened skull, but Ecology and habitat In Myanmar, has been found
are larger; Hardwicke’s Woolly Bat, K. hardwickii, in degraded lowland forest, at a foothill near
has more domed braincase; Indochinese Woolly Bat, evergreen forest; in Taiwan, found in a range of
K. dongduongana, and Dark Woolly Bat, K. furva, are forested habitats.
only reliably distinguished by genetics, but latter tends Distribution SE Asia: Kachin in Myanmar,
to have more uniformly dark fur without pale tips. N Vietnam. Also India, S China, Taiwan.
Ecology and habitat Has been caught in evergreen Status Not yet assessed.
and semi-deciduous forests, including in disturbed
areas.

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 TITANIA’S WOOLLY BAT orange; wing membranes between fingers with
Kerivoula titania NOT ILLUSTRATED large patches of contrasting black. Similar species
Measurements FA 34–36, T 48–53, Tibia 18.4– No other Kerivoula has contrasting orange and black
19.7, E 14–15. Wt 4.4–4.8g. Skull: ccl 14.0–14.4, wings; Black-and-Orange Woolly Myotis, Myotis
mt 6.4 rufoniger, has similar colour pattern on wings but is
Identification Upperparts grey, individual hairs considerably larger, with different-shaped ears and
with blackish base, paler middle, dark grey tip; teeth, black edges to ears and feet.
underparts paler. Skull moderately flattened, but not Ecology and habitat Recorded from dry forest and
as much as in K. kachinensis. Ratio of tibia/forearm scrubby areas; roosts among dead leaves in trees
length 0.55–0.57. Taxonomic notes Formerly and bananas. Flight slow and manoeuvrable.
confused with K. flora, which does not occur in Distribution SE Asia: Myanmar, Thailand, Laos,
region. Similar species Hardwicke’s Woolly Bat, Vietnam, Cambodia and Peninsular Malaysia (Penang
K. hardwickii, has shorter forearm, smaller skull; Island). Also India, Sri Lanka, Bangladesh, S China,
Kachin Woolly Bat, K. kachinensis, is larger, with Sumatra, Java and smaller Indonesian islands.
relatively flatter skull; Dark Woolly Bat, K. furva, has Status Least Concern, although rarely encountered
uniformly dark dorsal fur, relatively shorter tibia. in most parts of range.
Ecology and habitat Found mainly in evergreen
forest, often near limestone or rivers. CLEAR-WINGED WOOLLY BAT
Distribution SE Asia: Myanmar, Laos, Vietnam, Kerivoula pellucida PLATE 27
Thailand and Cambodia. Measurements FA 29–32, T 40–50, E 14.5–17.
Status Least Concern. Wt 3.5–5.0g. Skull: ccl 12.0–12.5, mt 5.3–5.6
Identification Upperparts pale orange-brown,
KRAU WOOLLY BAT individual hairs with paler greyish-white base;
Kerivoula krauensis PLATE 27 underparts greyish-white. Wing membranes and
Measurements FA 29–33, T 35–37, E 11–12. ears pale orange-brown, almost translucent. Ears
Wt 2.5–3.5g. Skull: ccl 11.3–11.5, mt 4.9–5.0 long and narrow. Similar species Hardwicke’s
Identification Hair of upperparts largely dark brown Woolly Bat, K. hardwickii, has grey-brown fur
to black (90% of length) with shiny golden tips; with dark base, shorter ears; Painted Woolly Bat,
underparts blackish base with broad (20%) buff K. picta, has black patches on wing membranes;
to white tip. Similar species Hardwicke’s Woolly Small Woolly Bat, K. intermedia, has darker orange
Bat, K. hardwickii, is similar in size but fur is grey- fur with dark base, small ears.
brown with only base of hair dark; Small Woolly Bat, Ecology and habitat Roosts in clumps of dried
K. intermedia, is generally orange-brown in colour with leaves, including dead, curled banana leaves.
shorter ears; Whitehead’s Woolly Bat, K. whiteheadi, Forages in understorey of tall forest.
has pale underparts, elongated premolars. Distribution SE Asia: peninsular Thailand and
Ecology and habitat Known records from Malaysia. Also Sumatra, Java, Borneo and the
understorey of mature lowland dipterocarp forest. Philippines.
Distribution SE Asia: peninsular Thailand and Status Near Threatened, owing to loss of lowland
Malaysia. Also Borneo, Sumatra. rainforest.
Status Data Deficient, but may be at risk due to loss
of lowland rainforest. SMALL WOOLLY BAT
Kerivoula intermedia PLATE 27
PAINTED WOOLLY BAT Measurements FA 26–31, T 37–41, E 9–11.5.
Kerivoula picta PLATE 27 Wt 2.9–4.2g. Skull: cbl 11.1–11.8, mt 4.6–5.0
Measurements FA 32–39, T 37–47, E 13–15. Identification Upperparts orange-brown with
Wt 4.5–5.5g. Skull: ccl 12.2–13.3, mt 5.5–5.8 dark base; underparts paler. Ears relatively small.
Identification Upperparts with soft, fluffy light Premolars small and rounded (as in Fig. 45a).
orange fur; underparts similar but slightly paler. Similar species Least Woolly Bat, K. minuta, is
Ears, wing bones, tail membrane and feet bright very similar and overlaps in forearm length, but

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 has smaller skull, lighter body weight; Whitehead’s incisors tall, about two-thirds length of canine, with
Woolly Bat, K. whiteheadi, has different coloration; well-developed secondary cusps. Similar species
pale brown above, white below, longer ears and very Other woolly bats, Kerivoula spp., have premolars
narrow premolars (Fig. 45b); Hardwicke’s Woolly Bat, with nearly circular cross-section (Fig. 45a), and
K. hardwickii, is slightly larger, with larger ears and less contrasting colour of underparts.
greyer fur with dark grey base; Clear-winged Woolly Ecology and habitat In Sabah, has been caught
Bat, K. pellucida, has pale fur with whitish base and in kerangas forest. A roost of 20–30 individuals
very long ears; Krau Woolly Bat, K. krauensis, has was found in a hanging cluster of large dead leaves
gold-tipped dark fur. by a river. In the Philippines, has been found in
Ecology and habitat Understorey of tall lowland secondary forest, scrub and open grasslands.
forest. Roost sites unknown. Distribution SE Asia: peninsular Thailand. Also
Distribution SE Asia: Peninsular Malaysia. Also Borneo and the Philippines.
Borneo. Status Least Concern globally, but status in region
Status Near Threatened, owing to loss of lowland uncertain, as only one record from early 1900s
rainforest, although still relatively common in some exists and the species has not been found since.
forest reserves.
Genus Phoniscus Similar to Kerivoula, with long
LEAST WOOLLY BAT woolly fur, but tragus white with distinct notch in
Kerivoula minuta PLATE 27 posterior margin near base; fur banded with four
Measurements FA 25–29.5, E 8–10. Wt 1.9– bands of colour including pale tip; upper canines
2.3g. Skull: cbl 10.0–11.1, mt 4.1–4.6 large with longitudinal grooves on outer face;
Identification Very small, light bat. Upperparts outer upper incisor very small, about one-quarter
orange-brown with brown base to fur; underparts height of anterior incisor; anterior lower premolars
slightly paler. Ear relatively small. Premolars small elongate, more than 50% longer than wide. Dental
and rounded. Similar species Small Woolly Bat, formula 2/3 1/1 3/3 3/3.
K. intermedia, is very similar in appearance but
is generally heavier with larger skull; Whitehead’s GREATER GROOVE-TOOTHED BAT
Woolly Bat, K. whiteheadi, has whitish belly, brown Phoniscus jagorii PLATE 27
upperparts, narrow premolars (Fig. 45b). Measurements FA 35–42, T 38–45, E 14–17.
Ecology and habitat Understorey of lowland Wt 7–11g. Skull: cbl 15.3–15.9, mt 6.7–7.1
rainforest, including some disturbed areas. Flight is Identification Upperparts golden-brown and black
slow and fluttering. overall, hairs banded with four colours: dark grey-
Distribution SE Asia: peninsular Thailand and brown base, then a buff band, then dark brown, then
Malaysia. Also Borneo. golden tip; underparts paler with slightly greyer tips;
Status Near Threatened, due to lowland rainforest short shiny yellow hairs along forearm and fingers.
loss. Tragus white with deep notch on posterior edge.
Canines long and grooved. Middle upper premolar
WHITEHEAD’S WOOLLY BAT elongate, projecting beside anterior premolar on inside.
Kerivoula whiteheadi PLATE 27 Similar species Lesser Groove-toothed Bat, P. atrox,
Measurements FA 28–29. Skull: cbl 11.8–11.9, is smaller, with more rounded second upper premolar;
mt 5.0 tube-nosed bats, Murina spp. have enlarged nostrils,
Identification Upperparts brown, fur with dark grey only two upper and lower premolars (Fig. 42).
base; underparts greyish-white. Ears moderately Ecology and habitat Lowland rainforest, as well as
long. Wing membranes apparently with white tip dry dipterocarp and semi-evergreen forest.
in mainland form, K. w. bicolor, but type and only Distribution SE Asia: Thailand, Laos, Vietnam and
known specimen has faded and it is no longer Peninsular Malaysia. Also S China, Java, Borneo,
possible to check the colours. Anterior two upper Bali, Sulawesi and the Philippines.
and lower premolars elongate and oval in cross- Status Least Concern, although there are relatively
section with knife-like cusps (Fig. 45b); inner upper few records from most areas.

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 LESSER GROOVE-TOOTHED BAT premolar; woolly bats, Kerivoula spp., have shorter
Phoniscus atrox PLATE 27 canines without any groove, no more than two
Measurements FA 31–35, T 35–40, E 12.5–14. bands of colour on hairs; tube-nosed bats, Murina
Wt 4.0–5.5g. Skull: cbl 12.9–14.1, mt 5.6–6.2 spp., have long, tubular nostrils and only two upper
Identification Upperparts golden-brown and premolars (Fig 42).
black overall, hairs banded with grey and brown Ecology and habitat Found in understorey of
base, black middle and orange-brown or buff tip; lowland dipterocarp forest and disturbed areas
underparts paler and greyer. Tragus white with near primary forest. Has been found roosting in old
deep notch in posterior edge. Upper canines long hanging birds’ nests.
with groove on outside; middle upper premolar Distribution SE Asia: S Thailand and Peninsular
rounded, similar in shape to anterior premolar. Malaysia. Also Sumatra and Borneo.
Similar species Greater Groove-toothed Bat, Status Near Threatened, owing to loss of lowland
P. jagorii, is larger with more elongate upper rainforest.

Family MINIOPTERIDAE BENT-WINGED BATS

The bent-winged bats were formerly considered part of the family Vespertilionidae, but are now
recognised to be evolutionarily very distinct and belong in their own family. All members of this family
are placed in the same genus, Miniopterus. They have a distinctive wing shape: third finger has a very
short first phalanx and very long terminal phalanx (see colour plates). Ear is short, slightly rounded,
with a moderate posterior fold and a short, blunt tragus curved slightly forwards. Fur is generally black
or dark brown, occasionally with reddish patches or even completely reddish. All species are very
similar in appearance, and difficult to distinguish except by size. Forearm length and weight separate
most individuals, but since there is some overlap, skull measurements are sometimes necessary for
confirmation. Owing to similarities in appearance, the taxonomy is somewhat uncertain – it is difficult
to determine which forms in different areas belong to the same species. Genetic studies may result in
some changes to the currently recognised species boundaries. Skull with large high braincase, slender
rostrum (Fig. 36g). Dental formula 2/3 1/1 2/2 3/3.
The conservation status of some Miniopterus may need to be reviewed after further taxonomic work, as
some species currently thought to be widespread may turn out to be multiple species with more restricted
ranges. Cave colonies are vulnerable to disturbance or exploitation, and some large colonies have been lost.

LARGE BENT-WINGED BAT EASTERN BENT-WINGED BAT

Miniopterus magnater PLATE 26 Miniopterus fuliginosus PLATE 26
Measurements FA 47–53. Wt 13–20g. Measurements FA 42–46. Wt 10–12.5g.
Skull: cbl 15.8–16.8, mt 6.4–7.3, M–M 7.4–8.0 Skull: cbl 14.5–16.0, mt 5.8–6.7, M–M 6.3–7.3
Identification See general description of bent- Identification See general description of bent-
winged bats. Largest of the Miniopterus, with winged bats. Forearm length overlaps range of
an especially wide palate, as measured by width Large Bent-winged Bat, M. magnater, but averages
across outer molars. shorter; body and skull are smaller and narrower.
Ecology and habitat Roosts in caves, but has also Taxonomic notes Formerly considered part of
been found away from known caves. Flies above M. schreibersii from Europe.
canopy catching insects; can be seen flying low over Ecology and habitat Roosts in caves, sometimes
streams and small bodies of water. mixed with other species of bent-winged bat.
Distribution SE Asia: Myanmar, Thailand, Laos, Distribution SE Asia: Myanmar, Thailand, Laos,
Vietnam, Cambodia and Peninsular Malaysia. Also Vietnam, Cambodia and Peninsular Malaysia.
NE India, SE China, Sumatra, Java, Borneo and However, only records from Vietnam have been
other Indonesian islands through to New Guinea. confirmed by genetic analyses as this species. Also
Status Least Concern. Pakistan, Nepal, China, Russia, Korea, Taiwan.
Status Least Concern.

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 MEDIUM BENT-WINGED BAT SMALL BENT-WINGED BAT
Miniopterus medius PLATE 26 Miniopterus pusillus PLATE 26
Measurements FA 38–44. Wt. 8.0–10.0g. Measurements FA 40–43. Wt 7.0–10.0g.
Skull: cbl 13.8–14.5, mt 5.5–6.0, M–M 5.8–6.5 Skull: cbl 12.7–13.4, mt 5.0–5.4, M–M 5.4–5.7
Identification See general description of bent- Identification See general description of bent-
winged bats. Intermediate in size between Eastern winged bats. Similar to Medium Bent-winged Bat,
and Small Bent-winged Bats, M. fuliginosus and M. medius, but slightly shorter and narrower skull
M. pusillus. Overlap in forearm length, but skull and dentition.
averages larger than that of Small Bent-winged Bat, Ecology and habitat Roosts in limestone caves
M. pusillus. and possibly other habitats as well. Forages in
Ecology and habitat Roosts in caves. Forages over open areas, including over small streams and
low rivers in forest. rivers.
Distribution SE Asia: S Thailand and Peninsular Distribution SE Asia: Myanmar, Thailand, Laos,
Malaysia. Also Java, Borneo, Sulawesi, the Philippines Vietnam and Cambodia. Also S India, Nicobar
and New Guinea. Islands and Hong Kong.
Status Least Concern. Status Least Concern.

Family MOLOSSIDAE FREE-TAILED BATS

Medium-small to large bats distinguished by their thick, relatively short tail, which protrudes for well over
half its length from the interfemoral membrane. The muzzle lacks a noseleaf and projects well beyond the
lower jaw. Lips often have a series of folds in the skin, appearing wrinkled. Ears thick, sometimes joined
over the top of the head by a band of skin. Wings long and narrow. In skull, premaxillae well developed and
attached to the palate and the maxillae.

Genus Tadarida Fur thick and short. Upper incisors records. Potentially at risk in region, as only known
long and well developed, evenly spaced between colony in Laos was being harvested for food;
the canines. Skull somewhat flattened without a tendency to roost in large cave colonies makes it
sagittal crest. Premaxillae in skull not fused. Dental susceptible to over-harvesting.
formula 1/2–3 1/1 2/2 3/3.
Genus Chaerephon Formerly included in Tadarida.
LA TOUCHE’S FREE-TAILED BAT Short, thick fur. Premaxillae in skull fused, forming
Tadarida latouchei PLATE 28 small foramina. Braincase slightly inflated with low
Measurements FA 53–55, T 47–49, E 26–27. sagittal crest. Dental formula 1/2 1/1 2/2 3/3.
Wt 18–26g
Identification Relatively large, with long, narrow, ASIAN WRINKLE-LIPPED BAT
naked wings. Relatively large ears, separated Chaerephon plicatus PLATE 28
except for a narrow skin fold at base. Fur short, Measurements FA 40–50, T 30–40, E 21–24,
dense and velvety; upperparts uniformly dark brown HF 8–9. Wt 17–31g
or greyish-brown; underparts greyer. Taxonomic Identification Fur short, dense and soft;
notes Formerly included in Tadarida teniotis. upperparts dark brown; underparts paler with grey
Similar species Wroughton’s Free-tailed Bat, tip to hairs. Upper lip heavily wrinkled, nostrils
Otomops wroughtoni, has ears joined for much of protruding slightly in front; ears moderate, thick
length, pale patches on shoulders; other free-tailed and rounded, joined across front of head by flap
bats (Mops and Chaerephon) are smaller. of skin. Two upper premolars, the anterior quite
Ecology and habitat Known to roost in caves. small; posterior upper molar well developed,
Recorded in dry evergreen hill forest. about half the area of the second molar (Fig.
Distribution SE Asia: Laos and Thailand. Also China 46a). Similar species Johore Wrinkle-lipped Bat,
and Japan. C. johorensis, has enlarged ‘pocket’ in membrane
Status Data Deficient, as known from only a few joining ears; Sunda Free-tailed Bat, M. mops,

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 has darker fur, more massive skull, with only one
upper premolar and reduced posterior upper molar
(Fig. 46b). a
Ecology and habitat Roosts in caves in large
densely packed colonies, sometimes containing
hundreds of thousands or millions of individuals.
Often flies out before darkness in dense flocks to
forage high above ground.
Distribution SE Asia: Myanmar, Thailand, Laos,
Vietnam, Cambodia and Peninsular Malaysia. Also
Sri Lanka, India, S China, Hainan, Sumatra, Java,
Borneo, Lesser Sunda Islands and the Philippines.
Status Least Concern. Although still abundant b
in many areas, with some large colonies well
protected, its tendency to roost in large colonies
makes it vulnerable to hunting or disturbance; all
large colonies have disappeared in the Philippines,
and some large colonies have been lost in Cambodia
and Laos.

JOHORE WRINKLE-LIPPED BAT

Chaerephon johorensis PLATE 28 Fig. 46 Upper left toothrows of Chaerephon plicatus (a)
Measurements FA 44–49, T 36–43. Wt 15–25g and Mops mops (b).
Identification Upperparts covered in short, dark
brown fur; underparts paler with grey tips to SUNDA FREE-TAILED BAT
fur. Upper lip heavily wrinkled; ears moderate, Mops mops PLATE 28
joined across top of head by flap of skin that Measurements FA 41–46, T 35–42, E 18–22.
is raised at centre and extended backwards to Wt 23–35g. Skull: cbl 19.1–19.4, mt 7.0–7.4,
form a ‘pocket’ between ears; under pocket, long C–C 5.2–5.7
tuft of hairs that can be everted and erected. Identification Fur thick; upperparts and underparts
Small anterior upper premolars (similar to Fig. uniform dark brown to reddish-brown; crown of
46a). Similar species Asian Wrinkle-lipped Bat, head nearly naked. Ears large and rounded, joined
C. plicatus, has narrower band of skin joining ears, at front across top of head by narrow flap of skin.
without ‘pocket’; Sunda Free-tailed Bat, M. mops, Upper lip heavily wrinkled. Only one upper premolar;
also lacks pocket, has darker fur, more massive posterior molar reduced (Fig. 46b). Similar species
skull, with only one upper premolar and reduced Asian Wrinkle-lipped Bat, C. plicatus, has shorter
posterior upper molar (Fig. 46b). fur, paler underparts, smaller skull with extra
Ecology and habitat Mainly associated with forest, small anterior upper premolar and well-developed
foraging high over canopy, but also flies lower in posterior molar (Fig. 46a).
large gaps and over rivers. Ecology and habitat Forages high above forest
Distribution SE Asia: Peninsular Malaysia. Also or large clearings, and sometimes flies low over
Sumatra. rivers. Has been found roosting in hollows in
Status Vulnerable, owing to loss of intact forest. trees, sometimes in association with Naked Bat,
Cheiromeles torquatus.
Genus Mops Fur thick and short. Upper incisors Distribution SE Asia: peninsular Thailand and
long and well developed, evenly spaced between Malaysia. Also Sumatra and Borneo.
the canines. Only one upper premolar. Dental Status Near Threatened, owing to loss of intact
formula 1/2 1/1 1/2 3/3. forest.

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 Genus Otomops Ears long, pointing forwards in Genus Cheiromeles Large and naked with only
front of head, and joined for much of length by fold scattered hairs on the skin. Ears separate. Skull very
of skin; virtually no tragus or anti-tragus; anterior robust with a prolonged sagittal crest. Anterior lower
upper premolar well developed, within toothrow. premolar minute. Dental formula 1/2 1/1 1/2 3/3.
Dental formula 1/2 1/1 2/2 3/3.
NAKED BAT
WROUGHTON’S FREE-TAILED BAT Cheiromeles torquatus PLATE 28
Otomops wroughtoni PLATE 28 Measurements FA 74–86, T 60–75, E 25–32.
Measurements FA 63–67, T 36, HF 10–14, E 35. Wt 150–185g
Skull: gl 24–25, mt 8.9–9.3 Identification Heaviest insectivorous bat in the
Identification Upperparts dark brown apart from world. Body almost completely naked except for
contrasting pale greyish-white collar across back scattered short hairs on the feet and a cluster of
and around under neck and throat; lower belly long hairs around a scent gland in the neck. Bare
greyish-brown; small tuft of white hairs at base skin of body dark grey. Feet large with long, sharp
of ears and on skin joining ears; thin line of claws. Ears well developed, separate. In flight
white hairs down side beside wings. Ears long, can be recognised by large size, strong flight and
pointing forwards over head, joined near front by audible clicking of echolocation call. Exposed tail
band of skin. Similar species La Touche’s Free- and large separate ears can sometimes be seen in
tailed Bat, Tadarida latouchei, is slightly smaller, flight as well.
with less elongate, separated ears, uniform dark Ecology and habitat Roosts in large caves or hollow
upperparts. trees, sometimes associated with Sunda Free-tailed
Ecology and habitat One individual was found in Bat, Mops mops. Feeds in open areas over streams
open semi-deciduous dipterocarp forest. Audible or clearings, or high over forest canopy.
component to echolocation call. In India, only known Distribution SE Asia: peninsular Thailand and
colony roosts in a cave. Malaysia. Also Sumatra, Java, Borneo, Palawan and
Distribution SE Asia: Cambodia. Also S India. some nearby small islands.
Status Data Deficient; appears to be very rare, but Status Least Concern. However, many populations
too little information is available to assess threats. have declined due to loss of forest habitat as well
as trapping for food; some formerly large colonies
in caves have been virtually wiped out by hunting.

Order PRIMATES
Lorises, monkeys and gibbons
Primates are represented in mainland South-east Asia by the lorises (family Lorisidae), the true monkeys,
including macaques and leaf monkeys (family Cercopithecidae), and the gibbons (family Hylobatidae), as
well as by the Hominidae, the family that includes humans. All primates have hands and feet that can grasp,
digits generally with nails rather than claws, and both eyes on the front of the face. The Lorisidae are strictly
nocturnal, while the other species are mainly active during the day. Although all species are protected to
varying degrees in the region, many primates are threatened due to clearing and conversion of forest, as
well as illegal hunting and trapping both for food and for the pet trade. In some cases, accurate survey
information is not available, but population declines are inferred from extensive loss of suitable habitat. A
few species are now Critically Endangered, with very small remaining populations and a high probability of
extinction in the near future.

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 Family LORISIDAE LORISES
Lorises are small, stocky primates with a very short tail, and they are largely nocturnal and arboreal. Unlike
most other primates they climb slowly and deliberately, almost never jumping and rarely moving quickly
except when threatened or when striking at prey. All the digits have nails, with the exception of the second
digit of the foot, which has a short claw. The thumb and big toe are both opposing, allowing the animal to
grip branches very strongly.

SUNDA SLOW LORIS species Sunda Slow Loris, N. coucang, averages

Nycticebus coucang PLATE 29 smaller and has a dark stripe continuing to top of
Measurements HB 260–300, T 15–25, HF 48–63. head and forking towards eyes and ears; hybrids
Wt 375–900g. Skull: gl 55–62 may occur in area of overlap in peninsular Thailand;
Identification General coloration varies from pale Pygmy Loris, N. pygmaeus, is smaller, with larger
grey-brown to reddish-brown, with a broad dark ears.
brown to black stripe from lower back to top of head, Ecology and habitat Nocturnal and arboreal.
where it branches into four lines connecting to the Occurs in both evergreen and deciduous forests,
eyes and ears. Usually has a dark-coloured ring as well as degraded areas such as bamboo groves.
around each eye. Fur soft and dense. Eyes reflect Feeds mainly on fruits, tree gums and nectar, but
light clearly at night with a reddish colour. Similar also on some animal matter, including large insects,
species Asian Slow Loris, N. bengalensis, averages nestling birds and lizards.
larger, with paler head, narrower dorsal stripe that Distribution SE Asia: Myanmar, Thailand, Laos,
usually ends on neck with, at most, indistinct forks Vietnam and Cambodia. Also NE India, Bangladesh
to eyes and ears; hybrids may occur in area of and S China.
overlap in peninsular Thailand. Status Vulnerable. Populations declining due to loss
Ecology and habitat Nocturnal and usually of forest habitat as well as hunting and trapping for
arboreal, mostly in small to medium-sized trees. food and pet trade.
Feeds mainly on fruits, flower nectar, gum and sap,
but also eats some small animals, mostly insects. PYGMY LORIS
Occurs in tall and secondary forests, as well as Nycticebus pygmaeus PLATE 29
gardens and cocoa plantations. Mainly solitary. Measurements HB 210–290. Skull: gl 48–55
Distribution SE Asia: peninsular Thailand and Identification Small loris with finely textured
Malaysia south of the Isthmus of Kra. Also Sumatra. reddish-buff fur, though some animals may be
Populations in Java, Borneo and the Philippines are dull brown. Dorsal stripe varies from faint or
now considered separate species. absent to broad and dark, with indistinct forks
Status Vulnerable. Populations throughout the area on head. Ears long and relatively conspicuous.
have suffered from severe loss of habitat due to Some researchers suggest that this may comprise
clearing of forests, as well as illegal hunting and a mixture of two species, while others believe
trapping for the pet trade. differences are mainly age-related, young animals
being more orange-coloured and lacking dorsal
ASIAN SLOW LORIS stripe. Genetic studies may be required to resolve
Nycticebus bengalensis PLATE 29 this uncertainty. Similar species Asian Slow Loris,
Measurements HB 300–380, T 10–20, E 20–25, N. bengalensis, is larger with less conspicuous
HF 55–70. Wt 1,000–2,000g. Skull: gl 62–67 ears and a thin dorsal stripe that does not continue
Identification Overall colour orange-buff to light onto top of head.
brown, with extensive greyish frosting on neck, head Ecology and habitat Nocturnal and arboreal.
and forelimbs, making head and neck distinctly Occurs in a wide variety of forest habitats, including
paler than back; thin, brown dorsal stripe ends on semi-evergreen and evergreen forest as well
neck with, at most, indistinct fork on head; dark as degraded forests and bamboo, mainly below
rings around eyes. Taxonomic notes Formerly 500m altitude. Feeds heavily on animal material,
considered a subspecies of N. coucang. Similar especially insects, as well as plant sap and flowers.

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 Also takes some small vertebrates such as lizards, Status Vulnerable. Declining due to hunting and
eggs and nestlings of birds. trapping for the food and pet trade.
Distribution SE Asia: Laos, E Cambodia and
Vietnam. Also extreme S China.

Family CERCOPITHECIDAE MONKEYS

Monkeys in South-east Asia are represented by two distinct groups: the langurs or leaf monkeys (subfamily
Colobinae) and the macaques (subfamily Cercopithecinae). The colobines are mainly arboreal, have a long
tail, lack cheek pouches and have a large sacculated stomach with many chambers, which permits the
breakdown of leafy material into digestible substances and helps detoxify poisonous leaves. The macaques
are partly terrestrial, with a short or long tail, cheek pouches for temporary storage of food, and a simple
stomach capable of breaking down only a limited amount of leafy material. The skull has a large, domed
braincase with forward-pointing orbits for the eyes (Fig. 47).
Footprints can be recognised by the five distinct fingers/toes, with an opposable first digit separated
from the rest (Fig. 48). Only the macaques regularly forage on the ground, but footprints for other species
may be found near water, where they come down to drink or to search for minerals. Tracks of langurs have
a relatively longer, narrower foot and shorter first digit compared with those of macaques, but the species of
langurs probably cannot be readily distinguished from tracks.

BANDED LANGUR Ecology and habitat Diurnal and arboreal. Travels

Presbytis femoralis PLATE 30 in groups, typically of three to six individuals. Occurs
Measurements HB 460–590, T 695–765, in dipterocarp forest. Feeds on leaves and fruits.
HF 170–185 Distribution SE Asia: P. f. femoralis: S Peninsular
Identification Dorsally dark brown or blackish, Malaysia (Johore) and Singapore. Reduced to only
underside medium to light grey, with conspicuous a few groups in Singapore, but status on adjacent
pale patches, especially on inside of thighs. Head mainland is unknown. P. f. robinsoni: peninsular
with a distinct central crest, pale grey bare skin Myanmar and peninsular Thailand from about 13°N,
around eyes, but rings much darker and less south to N Peninsular Malaysia (Perak). Also occurs
conspicuous than in Dusky Langur, Trachypithecus in Sumatra.
obscurus. Newborn infants have ‘cruciger’ pattern Status Near Threatened. Declining due to loss and
– white or beige, with dark lines down the back conversion of forest habitat.
and across the shoulders. Separate populations
in N and S Peninsular Malaysia/Thailand differ PALE-THIGHED LANGUR
somewhat in appearance: P. f. femoralis: blackish Presbytis siamensis PLATE 30
above, sometimes with red-brown tone; sharply Measurements HB 430–690, T 680–840
demarcated mid-grey below with white mid- (130–160% of HB)
ventral line and pale patches on inside of legs; Identification Upperparts, including top of head
P. f. robinsoni: upperparts dark brown to black, with and top of arms, brown to greyish-brown; hands,
tail and feet black; underside mid-grey with slightly feet and distal half of tail black; underside,
paler areas on inside of thighs. Some individuals including underside of arms and legs, as well as a
are all blonde or beige. Voice Adult male loud call large patch on outside of legs, pale grey to whitish.
is a staccato ‘ke-ke-ke’. Similar species Sundaic Bare face skin dark grey to nearly black, but
Silvered Langur, Trachypithecus cristatus, has sometimes skin around eye may be paler, forming
more uniform fur colour, with no white markings indistinct rings. Similar species Banded Langur,
(although parts of the body may appear whitish in P. femoralis, is generally darker above, without
bright sunlight) and orange-coloured infants; Dusky contrasting pale outer thighs. Dusky Langur,
Langur, Trachypithecus obscurus, has much more Trachypithecus obscurus, has conspicuous pale
conspicuous white rings around eyes, and a very rings around the eyes.
different voice. Ecology and habitat Found in a wide variety

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 of forest types from lowlands to hills, including
disturbed forests, orchards and plantations.
Distribution SE Asia: Peninsular Malaysia, in
between range of P. f. femoralis and P. f. robinsoni.
Also in Sumatra.
Status Near Threatened. Declining due to loss of
forest.

SUNDAIC SILVERED LANGUR

Trachypithecus cristatus PLATE 30
Measurements HB 415–540, T 600–760,
HF 145–174. Wt 4.0–6.5kg
Identification Entirely dark grey, the individual
hairs frosted with pale grey tip. Face dark grey.
May appear totally black in poor light. Infants are
bright orange. Voice Relatively quiet; calls include
rattling grunt, as well as shrill squeals and screams,
0 20
mainly from young. Taxonomic notes Peninsular
Malaysian form sometimes considered a separate
species, T. selangorensis, from the Sumatran Fig. 47 Skull of typical leaf monkey, Presbytis sp.
and Bornean forms. Similar species Banded
Langur, Presbytis femoralis, has contrastingly paler Distribution SE Asia: W coast of Peninsular
underparts; infants are not orange; Dusky Langur, Malaysia. Also Sumatra, Borneo and some adjacent
T. obscurus, has large pale rings around eyes, pale islands.
lips and less of a crest on the head. Status Near Threatened. Declining due to extensive
Ecology and habitat Diurnal and generally loss of coastal forests including mangroves.
arboreal. Found in coastal, riverine and swamp
forests, as well as some adjacent plantations, INDOCHINESE SILVERED LANGUR
rarely more than 40km inland. Diet includes Trachypithecus germaini PLATE 30
leaves, shoots and fruits, including many mangrove Measurements HB 490–570, T 720–840
species. Usually found in moderate to large groups Identification Dark black upperparts; slightly paler
of 10 to 50 individuals. underparts with paler grey patches on arms and

a c
b

F
H

H 0 100 H

Fig. 48 Left hind (H) and front (F) footprints of Macaca fasciculata (a) and M. nemestrina (b), and a langur,
Presbytis sp. (c). Within each species, print size varies considerably with age and sex.

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 legs, although hands and feet are dark. Tail dark crest on top of head contrasting paler grey; face
grey. Long, straight whitish hairs on head around skin dark grey with strongly contrasting incomplete
face, including crest. Infants orange. Similar white rings around eye, which are narrow on
species Annamese Silvered Langur, T. margarita, top, broad on the outside, but usually incomplete
has much paler upperparts with contrasting dark and dark grey near the nose; contrasting bare
hands and feet, pink ring around each eye; Phayre’s pink patches on upper and lower lips. Newborn
Langur, T. phayrei, has pale marks around eye, lacks young pale orange. Several subspecies have been
contrasting pale hairs around face. described from some offshore islands, differing in
Ecology and habitat Occurs in evergreen, semi- colour. Some are much darker than the mainland
evergreen and mixed forest, especially in riverine form, but all have the distinctive, conspicuous
areas, mainly at low elevations. eye-rings. Voice Loud double honking call ‘ceng-
Distribution SE Asia: SE Thailand, Cambodia and kong’. Similar species Pale-thighed Langur,
S Vietnam W of Mekong River. Presbytis siamensis, lacks pale eye-rings, has
Status Endangered. Much of forest habitat has contrasting dark tail and feet, different voice;
been destroyed; occurs at very low densities in most Tenasserim Langur, T. barbei, has dark body
remaining areas, probably due to excessive hunting without contrasting paler areas; Phayre’s Langur,
and trapping. T. phayrei, is generally more uniformly coloured
and paler.
ANNAMESE SILVERED LANGUR Ecology and habitat Mainly arboreal, living in a
Trachypithecus margarita PLATE 30 variety of forest types from lowlands to hills; lives
Measurements HB 500–600, T 700–800 in groups of 5 to 20, with one or more adult males,
Identification Upperparts medium to dark grey occupying a home range of 5–20 hectares. Feeds
with paler frosting; underparts paler, almost whitish mainly on leaves and shoots but also eats some
with very dark, nearly black, lower arms and feet. fruits, especially unripe ones.
Head with distinct crest, long pale grey ‘whiskers’ Distribution SE Asia: peninsular Myanmar, Thailand
on side of face. Face dark grey except for well- and Malaysia. Also some small offshore islands.
marked pale pink ring around each eye. Infants Status Near Threatened. Declining due to loss of
orange. Taxonomic notes Sometimes considered forest and hunting.
the same species as T. germaini. Similar species
Indochinese Silvered Langur, T. germaini, is more TENASSERIM LANGUR
uniformly dark grey except for pale hairs around Trachypithecus barbei PLATE 31
face, lacks pink rings around eye. Phayre’s Langur, Measurements HB 500–700, T 700–800
T. phayrei, has generally paler body colour, pale Identification Body dark greyish-black without any
whitish patch around mouth. frosting, only slightly paler underneath; long, dark
Ecology and habitat Found in tall primary forest grey tail, slightly paler than body with a pale patch at
and coastal forest. the base; face grey with large white rings around eye
Distribution SE Asia: Cambodia, Laos and C and and whitish area of bare skin around mouth. Similar
S Vietnam, largely or exclusively E of Mekong River. species Dusky Langur, T. obscurus, has legs and
Status Endangered (assessed with T. germaini). crown paler than body; Phayre’s Langur, T. phayrei,
Much of the coastal forest has been lost, and the has browner body colour with paler underparts;
species has further declined in some parts of its Indochinese Silvered Langur, T. germaini, has dark
range due to hunting and trapping. face with contrasting pale hairs around head.
Ecology and habitat Forest.
DUSKY LANGUR Distribution SE Asia: known only from a small area
Trachypithecus obscurus PLATE 31 in SE Myanmar and SW Thailand.
Measurements HB 500–700, T 700–800. Status Data Deficient. No recent observations;
Wt 6.5–7.5kg likely threatened due to limited range, loss of forest
Identification Upperparts greyish-brown to dark habitat and apparently low densities.
grey; underparts, outside of hind legs, tail and

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 INDOCHINESE GREY LANGUR much darker, nearly black, on the back. Indochinese
Trachypithecus crepusculus PLATE 31 Grey Langur, T. crepusculus, has more uniform grey
Measurements HB 520–620, T 600–850, upperparts, contrasting dark hands and feet.
HF 170–180. Wt 6–9kg Ecology and habitat Primary or secondary
Identification Overall colour of back, legs and head evergreen and semi-evergreen forest, but also in
mid- to light grey; underparts whitish. Hands and some more disturbed habitats, including bamboo-
feet dark grey, blending into lighter grey of arms dominated areas. Also in forest on limestone
and legs. Bare facial skin dark grey except for outcrops.
pale greyish-white patch around mouth and large Distribution SE Asia: Myanmar. Also E India,
conspicuous pale rings around eyes. Taxonomic Bangladesh and S China.
notes Formerly considered a subspecies of Phayre’s Status Endangered. In many parts of the range
Langur, T. phayrei. Similar species Dusky Langur, populations have declined severely, owing to loss
T. obscurus, has thighs paler than back; Tenasserim of habitat and to hunting, including for medicinal
Langur, T. barbei, is much darker, nearly black, on values, especially for ‘bezoar stones’ that form inside
the back; Phayre’s Langer, P. phayrei, is browner the animals when they drink from mineral springs.
with hands and feet same colour as legs. Populations in Myanmar poorly known, but likely
Ecology and habitat Primary or secondary evergreen declining severely; in India and Bangladesh, largely
and semi-evergreen forest, but also in some more restricted to small fragmented protected areas.
disturbed habitats, including bamboo-dominated
areas. Also in forest on limestone outcrops. FRANCOIS’S LANGUR
Distribution SE Asia: SE Myanmar, Thailand, Laos Trachypithecus francoisi PLATE 32
and N Vietnam. Also S China. Measurements HB 470–630, T 740–960
Status Endangered (as part of T. phayrei). In many Identification Fur nearly all black, except for a band
parts of range has declined severely, owing to loss of elongated white hairs from corner of mouth to
of habitat and to hunting, including for medicinal above ears like an extended moustache. Tall, pointed
values, especially for ‘bezoar stones’ – mineral crest on top of head. Newborn babies are largely
deposits that form inside the animals when they orange, turning black with a pale orange head.
drink from springs rich in minerals, and that are Similar species Hatinh Langur, T. hatinhensis, has
thought to have magical powers. Still found in white facial stripes continuing around to back of
moderate numbers in some protected areas. head.
Ecology and habitat Found only in sub-tropical and
PHAYRE’S LANGUR tropical rainforests and monsoon forest on limestone
Trachypithecus phayrei PLATE 31 karst outcrops. Often shelters in limestone caves.
Measurements HB 520–620, T 600–850, Feeds mainly on leaves as well as some shoots,
HF 170–180. Wt 6–9kg flowers and fruit.
Identification Fur colour of back and head of Distribution SE Asia: N Vietnam. Also extreme
nominate subspecies (T. p. phayrei in W Myanmar S China.
and India) varies from grey-brown to reddish-brown Status Endangered. Total population in Vietnam
with sharply contrasting white underparts. Fur probably fewer than 500 animals, fragmented into
around face dark. Legs, feet and tail similar colour many relatively small sub-populations due to loss
to back. Bare facial skin dark grey except for pale of intervening forests. Major threat is from hunting,
greyish-white patch around mouth and large pale especially for traditional medicine.
rings around eyes. Subspecies found in N and E
Myanmar (T. p. shanicus) is light brownish-grey CAT BA LANGUR
overall, with underparts paler but not contrasting Trachypithecus poliocephalus PLATE 32
strongly; rings around eyes incomplete and Measurements HB 490–590, T 800–900
noticeable only on the inner side of the eye. Similar Identification Body dark chocolate-brown; head
species Dusky Langur, T. obscurus, has thighs and neck vary from pale yellowish to dark orange;
paler than back; Tenasserim Langur, T. barbei, is patch of paler grey on rump and upper thigh. Tail

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 long and black. Similar species No other langur Distribution SE Asia: known only from C Laos.
with pale head occurs within its range. Status Vulnerable. Declining due to loss of habitat,
Ecology and habitat Occurs in forests on limestone excessive hunting and trapping.
karst hills. Arboreal and terrestrial. Shelters in caves.
Feeds mainly on leaves, as well as some shoots, DELACOUR’S LANGUR
fruit, flowers and bark. Trachypithecus delacouri PLATE 32
Distribution SE Asia: found only on Cat Ba Island, Measurements HB 570–730, T 730–970
N Vietnam. Identification Most of body and head glossy black
Status Critically Endangered. Fewer than 100 above and below; large white patch on rump and
individuals remain, fragmented into several outside of thighs; cheeks with long pale grey hairs,
populations. Ongoing threats include disturbance of topped by ‘moustache’ from sides of mouth to above
remaining habitat, illegal hunting and trapping. ears. Tail very long, glossy black. Similar species
No other monkey has such contrasting white rump
HATINH LANGUR and legs.
Trachypithecus hatinhensis PLATE 32 Ecology and habitat Found only in forest on
Measurements HB 500–660, T 810–870 limestone karst hills, although can tolerate degraded
Identification Body largely glossy black overall forest. Shelters in limestone caves. Feeds mainly on
except for white moustache that extends from sides leaves, but also some shoots, flowers, fruit and bark.
of mouth over behind ears to nearly touch on back Distribution SE Asia: found only in a limited area of
of neck. Forehead has white band in young animals, Vietnam S of the Red River.
but is glossy black in adults. Distinct black crest. Status Critically Endangered. Fewer than 250
Similar species Francois’s Langur, T. francoisi, has individuals remain, fragmented into many small
moustache that does not continue behind ears; Lao sub-populations owing to loss of habitat. Most
Langur, T. laotum, has white forehead even in adults. populations too small for long-term viability –
Ecology and habitat Forested habitat in limestone several have been lost in recent years. Major current
karst and rocky outcrops in mountainous areas. threat is illegal hunting.
Arboreal and terrestrial. Eats mainly leaves.
Distribution SE Asia: E Laos and NC Vietnam. INDOCHINESE BLACK LANGUR
Status Endangered. Populations have declined Trachypithecus ebenus PLATE 32
owing to loss of habitat, though moderate Measurements HB 600–700, T 800–900
populations remain in some protected areas. Major Identification Glossy black all over, including head,
threats are illegal hunting and trapping. except for a few scattered pale hairs on chin, hands
and feet. Hairs on head raised in a distinct crest.
LAO LANGUR Taxonomic notes Sometimes considered a colour
Trachypithecus laotum PLATE 32 morph of T. hatinhensis. Similar species No other
Measurements HB 460–530, T 800–900 langurs are completely glossy black without white
Identification Overall colour of body glossy black on the head or body.
above and below; head with long white whiskers that Ecology and habitat Lives in forests on limestone
extend from sides of mouth, above ears and around karst, as well as other rock outcrops in steep hills
back of head nearly to nape; broad white band on or mountainous areas. Feeds mainly on leaves.
forehead; black crest joined by narrow black stripe Arboreal and terrestrial.
down back of neck to black colour on back. Similar Distribution SE Asia: C Laos and Vietnam.
species Francois’s Langur, T. francoisi, lacks white Status Endangered (as T. hatinhensis). It occurs in
forehead band; Hatinh Langur, T. hatinhensis, has a limited range, where it is threatened by hunting
pale forehead only in juveniles; Indochinese Black and trapping.
Langur, T. ebenus, has all-black head.
Ecology and habitat Found only in forest on CAPPED LANGUR
limestone karst. Arboreal and terrestrial. Feeds Trachypithecus pileatus PLATE 31
mainly on leaves. Measurements HB 500–700, T 800–1,000

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 Identification Grey to pale brown on back, chestnut. Face skin yellow-brown except around
contrasting with paler buff to yellowish-orange on mouth and chin, which is white; broad black frontal
face, sides of body and underparts; crown of head band above face; long white cheek whiskers.
with distinct black cap, hairs short, sticking straight Tail very long, white, with thick tassel at tip. Eyes
up; tail coloured as back, with black tip. Colour of slanting, forming an angle of >20° from horizontal.
underside varies from whitish to dark orange, sides Similar species Black-shanked Douc, P. nigripes,
of face usually more orange than rest. Similar has black lower arms and legs, bluish-grey facial
species No other species has such a distinct dark skin with black streak on sides of face.
cap contrasting with yellowish face. Ecology and habitat Largely arboreal, living in
Ecology and habitat Found in evergreen, semi- tall evergreen and semi-evergreen forests, from
evergreen and moist deciduous forests, as well lowlands up to about 2,000m, including on some
as bamboo forests and relatively open woodlands. limestone outcrops. Feeds mainly on leaves and
Mainly arboreal, feeding on leaves, but sometimes buds, as well as some seeds, flowers and fruit.
descends to ground. Moves noisily through branches when relaxed, but
Distribution SE Asia: NW Myanmar, W of Chindwin disappears quietly when disturbed. Usually found in
River. Also NE India and Bangladesh. small groups of three or more individuals.
Status Vulnerable. Populations in Myanmar may Distribution SE Asia: N and C Vietnam and Laos
be at higher risk due to loss of habitat, hunting and between about 14°N and 20°N.
trapping for food and the pet trade. Status Endangered. Has suffered extensive habitat
loss, especially in Vietnam; biggest current threat is
SHORTRIDGE’S LANGUR hunting and trapping, both for food and for the pet
Trachypithecus shortridgei PLATE 31 trade.
Measurements HB 600–750, T 900–1,050
Identification Uniformly silver-grey on head and BLACK-SHANKED DOUC
body, including underside; slightly paler on legs Pygathrix nigripes PLATE 34
except for nearly black hands and feet and distal Measurements HB 610–760, T 560–760
half of tail. Hairs on top of head stick up vertically. Identification Upperparts and crown dark
Bare face skin dark grey. Eyes pale orange-yellow. speckled grey, underparts paler grey; chin and
Similar species Phayre’s Langur, T. phayrei, has throat white; arms and legs largely black with
underparts paler than upperparts, white patches on some paler frosting on arms. Bare skin on face
lips and around eyes. blue-grey except for yellow-orange rings around
Ecology and habitat Evergreen and semi-evergreen eyes; eyes relatively straight, forming an angle of
forest. Largely arboreal, feeding on leaves, but <10° from horizontal; white whiskers on cheeks
sometimes comes down to the ground. relatively short; black frontal band above face
Distribution SE Asia: NE Myanmar. Also adjacent extends down side of face to connect with black
NE India and S China. of shoulders, and also forms a wedge on side of
Status Endangered. Believed to have declined face like a sideburn. Tail very long, white, with thin
at least 50% in recent years due to habitat loss tassel. Sometimes hybridises with Red-shanked
and hunting, although only limited information is Douc, P. nemaeus, in zone of overlap (c.13°N
available on current status. to 14°N). Similar species Red-shanked Douc,
P. nemaeus, has white lower arms, red lower legs,
RED-SHANKED DOUC orange facial skin, white on sides of face; Grey-
Pygathrix nemaeus PLATE 34 shanked Douc, P. cinerea, has pale grey legs,
Measurements HB 610–760, T 560–760 yellow-brown facial skin.
Identification A strikingly coloured primate with Ecology and habitat Largely arboreal, but will travel
back, belly and tops of upper arms speckled grey; on ground. Found in evergreen, semi-evergreen and
shoulders, inside of upper arms, upper legs and mixed deciduous primary and disturbed forest.
rump, hands and feet black; lower arms and scrotal Feeds mainly on leaves and seeds, but also eats
region white; lower legs (shanks) dark reddish- buds, flowers and fruits.

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 Distribution SE Asia: S Vietnam and adjacent head and body, largely black, frosted with pale hairs
Cambodia from about 10°30’N to 14°30’N. along top and a pale tip. Similar species No other
Status Endangered owing to habitat loss and primate in region has similar face pattern; Cat Ba
hunting. Langur, Trachypitheus poliocephalus, has a pale
head, but dark brown body, different head shape,
GREY-SHANKED DOUC does not overlap in range.
Pygathrix cinerea PLATE 34 Ecology and habitat Occurs only in forests on
Measurements HB 610–760, T 560–760 limestone karst mountains. Largely arboreal, rarely
Identification Body, crown, most of arms speckled descending to the ground. Feeds on leaves and
light grey, paler on underside and lower arms; fruits. Sleeps on lower branches of trees.
shoulders, upper legs and part of rump, feet and Distribution SE Asia: known only from a limited
hands black; lower legs dark speckled grey. Bare area in N Vietnam.
skin on face yellow-brown, except around mouth Status Critically Endangered. Remaining world
and chin, which is white; long white whiskers on population estimated at fewer than 300 individuals,
sides of face; eye slant intermediate, forming an restricted to a few reserves; threatened by illegal
angle c.15° from horizontal; narrow black band on logging and hunting.
forehead not connected to shoulders; throat white
with a broad orange collar bordered by a black line MYANMAR SNUB-NOSED MONKEY
that joins black patches on shoulders and inner Rhinopithecus strykeri PLATE 33
surfaces of upper arms. Tail long and white with Measurements HB 555, T 680
thin tassel; white patch on rump at base of tail. Identification Fur largely black on most of head,
Similar species Other doucs have darker bodies; body, arms, legs and tail; fur on sides of face and
Red-shanked Douc, P. namaeus, has red lower legs, underparts may be dark brownish-black. Face
contrasting white forearms; Black-shanked Douc, flattened with very short nose and enlarged pinkish
P. nigripes, has uniformly blackish arms and legs, lips. Bare pink or pinkish-grey skin on face and
dark sideburns, blue facial skin; hybrids between around eyes. White beard (and sometimes other
P. namaeus and P. nigripes may be found in same facial hairs), especially in older males, white tufts of
area as P. cinerea, but do not have grey shanks hair on ears, white around genital area. Tail relatively
or white underparts, instead resembling one or long, about 40% longer than head and body in adult
other parent. male. Infants are grey to dark grey, with light grey
Ecology and habitat Evergreen and semi-evergreen faces. Similar species Tonkin Sub-nosed Monkey,
primary rainforest, although also in some degraded R. avunculus, has pale underparts, more white on
forests. Largely arboreal, feeding mainly on leaves, head, and does not overlap in range. Taxonomic
but also some buds, fruits and flowers. notes Newly described in 2011.
Distribution SE Asia: recorded only from C Vietnam, Ecology and habitat Occurs only in forests
between 14°N and 14°46’N. (broadleaf and coniferous) in mountains of
Status Critically Endangered. Rarest of the doucs, N Myanmar and adjacent China, primarily at
threatened by forest clearing, hunting and trapping. altitudes of 2,400–3,300m, but sometimes lower,
especially in winter. Largely arboreal, but also travels
TONKIN SNUB-NOSED MONKEY on the ground. Home range for a troop estimated
Rhinopithecus avunculus PLATE 33 up to 23km2.
Measurements HB 510–620, T 660–920 Distribution SE Asia: N Myanmar. Also nearby
Identification Back, including back of neck, tops areas in S China.
of arms and legs, largely black; top and front of Status Critically Endangered. World population
head white to brownish-white; underparts including estimated at less than 1,000 individuals; ongoing
insides of arms and legs white; bare skin on face pale threats of hunting and forest loss.
bluish to bluish-grey, with white rings around eyes;
face flattened with very short nose and enlarged
pinkish lips. Tail very long, about 50% longer than

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 Genus Macaca (Macaques) Six species of NORTHERN PIG-TAILED MACAQUE
macaque are currently recognised in the region. Macaca leonina PLATE 35
The most distinctive is the Stump-tailed Macaque, Measurements HB 470–585, T 140–230,
M. arctoides, which has a short, stump-like tail and HF 140–170. Wt adult males 7–9kg, adult females
bare skin on its face. The Southern and Northern 4–6kg
Pig-tailed Macaques, M. nemestrina and M. leonina, Identification Similar to Southern Pig-tailed
are easily distinguished from other macaques by Macaque, M. nemestrina, but lighter build, shorter
their short tail and distinctive facial pattern, but muzzle; tail slightly hairier, especially at tip, tends to
differences from each other are more subtle. The be carried over back arched towards rear, with tip
remaining three species have a well-developed tail pointing down. Body fur generally brown to golden-
and differ in shape, average tail length, fur colour, brown; whiter on face. Less black on back than
range and habitat. Footprints (Fig. 48) are readily M. nemestrina, with distinctive red streaks on face
distinguished from those of other animals by the pointing diagonally upwards from eyes. Possible
opposable first digit, but difficult to distinguish from hybrids with N. nemestrina have been reported
one another. The more terrestrial Pig-tailed and from peninsular Thailand around the Isthmus of
Stump-tailed Macaques tend to have larger, broader Kra (around 7°N to 8°N). Identification of animals
feet than the other more arboreal species. in some areas is complicated by feral animals that
have escaped or been released from captivity.
SOUTHERN PIG-TAILED MACAQUE Ecology and habitat Diurnal. Forages mainly on the
Macaca nemestrina PLATE 35 ground but also climbs trees; inclined to escape into
Measurements HB 470–585, T 140–230, trees if disturbed. Rarely found around temples or
HF 140–170. Wt adult males 7–9kg, adult females gardens, unlike some other macaques. Found in a
4–6kg wide variety of forest habitats, including disturbed
Identification Upperparts vary from greyish- forests.
brown to reddish-brown, with paler, often whitish Distribution SE Asia: Myanmar, Thailand (S to about
underparts. Top of head, neck and middle of back 8°N), Laos, Vietnam and Cambodia. Also N India,
distinctly dark brown or blackish. Tail is short and Bangladesh and S China (Yunnan).
sparsely haired; often held curled above the back, Status Vulnerable. Declining due to degradation of
especially in adult males. Similar species Northern habitat, hunting and trapping for pets or for use in
Pig-tailed Macaque, M. leonina, which has minimal climbing coconut trees.
overlap in range, is on average slightly smaller with
shorter face and shorter limbs; distinct diagonal STUMP-TAILED MACAQUE
stripes from corner of eye; crown not as dark; back Macaca arctoides PLATE 35
without distinct black colour; tail with slight tuft of Measurements HB 485–635, T 37–78, HF 145–
hair near tip. 177. Wt 8–12kg
Ecology and habitat Diurnal. Group size usually Identification Stout, heavily built macaque with
15–40 animals, but solitary males are also tail reduced to a short, barely visible stump about
encountered. Found in lowland and hill forests. 10% of head and body length that is usually
Natural diet includes fruits and small vertebrate kept down except when the animal is alarmed.
and invertebrate animals, but sometimes enters Upperparts vary from dark brown to reddish-brown
plantations and gardens, where it can cause or blackish, paler underneath. Juveniles tend to be
damage to grain and fruit crops. If disturbed, tends much paler. Hairs on back and sides of head can
to run off on the ground to escape. be very long and shaggy, and may form a beard
Distribution SE Asia: peninsular Thailand (S of about on adult males. Prominent eyebrows. Bald forehead
7°30’N) and Peninsular Malaysia. Also Sumatra, and largely bare skin on face. Facial skin pink to
Bangka Island and Borneo. reddish, becoming darker with age, but also turning
Status Vulnerable. Declining owing to habitat loss bright red when animal is excited. Similar species
and degradation, as well as hunting. Other macaques have a more prominent tail and a
different head shape.

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 Ecology and habitat Found mainly in upland areas, LONG-TAILED MACAQUE
in primary or secondary forest as high as 2,000m. Macaca fascicularis PLATE 35
Feeds mainly on the ground, on seeds, fruits, buds, Measurements HB 450–550, T 440–540
insects and small animals. Typically in relatively (90–120% of HB), HF 120–145. Wt 3.5–6.5kg
large groups, sometimes of up to 50 individuals. If Identification Grey-brown to reddish-brown, paler
disturbed, usually runs off through undergrowth, but on the underparts. Cheek whiskers prominent. Infants
may climb trees. blackish. Groups often detected by their calls, the
Distribution SE Asia: Myanmar, Thailand, Vietnam, most common being ‘krra!’. Individuals tend to be
Laos, Cambodia and extreme N Peninsular less noisy than langurs when travelling through the
Malaysia. Also NE India (Assam) and S China tree canopy, but groups are noisier. Tail typically about
(Yunnan, Guangxi, Guandong). 10–20% longer than head and body in southern
Status Vulnerable. Declining owing to loss or populations, slightly shorter than head and body
degradation of habitat, and hunting and trapping for in populations from Bangladesh through Myanmar
food, medicinal purposes and pet trade. to W Thailand. Similar species Other macaques
have shorter tail; Rhesus Macaque, M. mulatta, has
ASSAMESE MACAQUE reddish hindquarters; Silvered Leaf Monkey, Presbytis
Macaca assamensis PLATE 35 cristatus (often in same habitat), is entirely dark
Measurements HB 510–735, T 150–300 blackish-grey with frosted fur (infants orange), and
(30–40% of HB), HF 155–175. Wt 5–10kg has distinct crest of hairs on top of head.
Identification Heavy-set macaque with well-haired, Ecology and habitat Active periodically from dawn
relatively short tail. Fur varies from yellowish-grey to until dusk. Often travels in groups of 20 to 30
dark brown; shoulders, head and arms tend to be individuals, sometimes more, with 2 to 4 adult males,
paler than hindquarters; hair on underparts relatively 6 to 11 adult females and the remainder immatures.
pale and sparse, often with bluish skin showing Usually only part of the group can be seen at one
through. Hindquarters greyish and haired to edge of time. Males sometimes solitary or in small groups.
callosities (thickened bare skin on rump). Head has A group occupies an area of up to several tens of
fringe of dark hair on cheeks directed backwards hectares, travelling from 150 to 1,500m daily. Unlike
to ear; hair on crown typically parted in middle; other monkeys, spends a large proportion of the
facial skin dark brownish to purplish. Calls include time active in low trees and thick scrub. Common
a musical ‘pio’. Subspecies in NE India has a longer in coastal forests including mangrove and beach,
tail, about 50–60% of head and body. Similar and along rivers. Also around gardens, villages and
species Rhesus Macaque, M. mulatta, is lighter plantations. Diet mostly ripe fruits and a wide array
in build, with reddish hindquarters, hairs on crown of animal material including insects, frogs’ eggs,
directed backwards and not parted in middle; Long- crabs and other coastal invertebrates. Sometimes a
tailed Macaque, M. fascicularis, and leaf monkeys pest in commercial crops.
have a much longer tail. Distribution SE Asia: Myanmar, S Thailand,
Ecology and habitat Found mainly in upland areas, Peninsular Malaysia, S Laos, Cambodia and
as high as 3,500m in the hills, generally in forested S Vietnam. Also S Bangladesh, Sumatra, Java, Bali,
areas. Omnivorous, feeding on fruit, insects and Borneo and many smaller offshore islands.
small animals. Spends much of time in trees, but Status Least Concern.
will descend to the ground to run from predators,
and sometimes raids crops. RHESUS MACAQUE
Distribution SE Asia: N Myanmar, Thailand, Laos Macaca mulatta PLATE 35
and N Vietnam. Also NE India, Nepal, Pakistan and Measurements HB 470–585, T 205–280
S China. (40–70% of HB), HF 140–170. Wt 3–6kg
Status Near Threatened. Declining in most parts of Identification Upperparts brownish, with reddish-
range owing to habitat loss and illegal hunting. brown hindquarters, greyer-brown foreparts;
paler underneath. Tail is well haired, about half
length of head and body. Face is relatively hairy,

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 with bare areas pinkish. Hairs on crown directed often in close proximity to human settlements. May
backwards. Voice Similar to that of Long-tailed become a pest in agricultural areas. Reported to
Macaque, M. fascicularis, but more squealing and be declining in some areas as a result of loss of
less harsh. Similar species Assamese Macaque, habitat and persecution, including trapping for use
M. assamensis, has a similar-length tail, but in medical and other research.
heavier build and greyish hindquarters; Long-tailed Distribution SE Asia: N and C Myanmar, Thailand,
Macaque, M. fascicularis, has relatively longer tail, Laos, Cambodia and Vietnam. Also Nepal, Pakistan,
hindquarters same colour as back. India and China. Minimal overlap in range with
Ecology and habitat Typically gregarious, forming M. fascicularis, but some hybridisation occurs where
large troops that vary from 10 to over 50 individuals. ranges meet.
Frequents secondary forests and disturbed areas, Status Least Concern.

Family HYLOBATIDAE GIBBONS

Gibbons are slender, totally arboreal primates that rarely descend to the ground. They are distinguished from
other primates by their long, slender arms and lack of a tail. They can travel very rapidly in the forest canopy,
swinging by their arms. Many authors place these all in the same genus, Hylobates, but recent research
suggests they should be grouped into four separate genera, all of which have representatives in the region,
with a total of 10 different species.
Most species are separated from each other geographically, often by major rivers, with the exception of
the Siamang, Symphalangus syndactylus, which overlaps the range of H. lar and H. agilis. In areas where
two species meet, there is sometimes limited hybridisation. Species can usually be distinguished by colour
patterns as well as the calls, especially the great calls of the females.
The distributional and species limits of crested gibbons, Nomascus spp., in Indochina are not well
understood. Many areas have, until recently, been difficult to access owing to war. Recent taxonomic and
genetic studies have been largely based on captive animals, for which the precise origin is not always
known. Although increased field work is now underway, many areas of the original habitat have been heavily
disturbed by logging or hunting, with loss of animals from parts of their former range. Furthermore, in areas
where they remain, gibbons become very shy and difficult to observe. Analysis of recordings of their songs
appears to be a useful tool, but many areas have not yet been thoroughly surveyed.
All gibbons are protected under Appendix I of CITES (Convention on International Trade in Endangered
Species), meaning that no international commercial trade in them is permitted. Many species have
experienced severe population declines, and some populations are now threatened with extinction. The
major threats are habitat loss (since gibbons are largely dependent on tall forest), as well as illegal hunting
for food, medicine and the pet trade.

SIAMANG have pale brows, hands or cheeks, and are smaller

Symphalangus syndactylus PLATE 36 and more slender.
Measurements HB 750–900. Wt adult males Ecology and habitat Diurnal and fully arboreal,
10.4–15kg, females 8–11kg dependent on tall forest. Can survive in partially
Identification Largest of the gibbons, all black with logged forests only if sufficient large trees remain.
long, shaggy hairs, especially on legs and arms, Generally in small family groups of a pair with up
giving thickset appearance. Both sexes have a throat to three immature offspring; unmated adults are
pouch that enlarges during calls. Voice Calls consist solitary. Usually start calling later in the morning
of a loud boom, followed by series of loud whoops. than other gibbons, often more than two hours after
Female utters some whoops, as well as a series of sunrise. Diet consists of fruits, flowers, leaves and
short barks that gradually accelerate, at the climax shoots, with some animal food such as insects or
of which male gives an extremely loud yell. Calls can birds’ eggs. Moves less and eats proportionately
be heard several kilometres away. Similar species more leaves than smaller gibbons.
Dark-phase animals of other gibbons within range Distribution SE Asia: mountains and hills of extreme

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 S peninsular Thailand and Malaysia. Also Barisan Identification Adults have two colour phases,
Mountains of Sumatra and Indonesia. which are not related to sex. Dark phase is black
Status Endangered. Populations declining severely or very dark brown, sometimes with a reddish hue;
due to loss of forest, selective logging, illegal hunting pale phase is buff to blonde. Both sexes have white
and trapping for the pet trade. brow band (which may be separated); males also
have pale cheeks and sometimes a whitish ‘beard’,
WHITE-HANDED GIBBON usually lacking in females. Hands and feet same
Hylobates lar PLATE 36 colour as rest of body or slightly darker. Dark phase
Measurements HB 450–600, HF 133–155. most common in mainland SE Asia, but in Sumatra
Wt 4–7kg about half of animals are pale. Voice Loud call of
Identification Body colour varies from dark brown female a series of high-pitched rising and falling
or nearly black to buff or cream. Both pale and dark notes, similar to that of White-handed Gibbon, H. lar,
forms may occur in same family, but colour is not but shorter in duration. Both sexes give a distinctive
related to sex. Tops of hands and feet, brow band and ‘whoo-aa’ call, quite distinct from ‘hoo’ calls given
sides of face contrastingly pale or white. Voice Loud by H. lar. Similar species White-handed Gibbon,
call of female a series of slow, high-pitched rising H. lar, has pale hands and feet, complete white
and falling whoops; typically about 7–10 notes in a facial ring. Limited hybridisation with H. lar has been
series over 15–25 seconds. Both sexes give single reported from S Thailand, where the ranges meet for
‘hoo’ calls. Similar species Agile Gibbon, H. agilis, the two species.
has hands and feet same colour as body, less white Ecology and habitat Arboreal. Usually found in
on face; Pileated Gibbon, H. pileatus, females are pale small groups of one adult male, one adult female and
brown, with black patches on underparts and crown; one to three young. Each group defends a territory
males are similar to dark-phase H. lar, but have of 20–30 hectares. Diet consists of ripe fleshy fruits,
incomplete white facial ring and pale patch extending young leaves and small insects. Normally found only
above ears. Limited hybridisation with H. pileatus and in tall dipterocarp forests. Peak calling activity is
H. agilis has been reported in areas of overlap. soon after dawn.
Ecology and habitat Arboreal, rarely descending to Distribution SE Asia: extreme S Thailand and
the ground. Normally in a family group with adult N Peninsular Malaysia; species range occurs in
male and female and up to three offspring. Sleeps between range of White-handed Gibbon, H. lar, in
in tops of tall trees, either crouched in fork of tree, Peninsular Malaysia and Thailand. Also S Sumatra.
or with arms outspread grasping smaller branches. Status Endangered. Substantial habitat has been
Peak calling activity one to two hours after dawn. lost recently, especially in Sumatra. Status of
Diet mostly fruits, leaves, new shoots and flowers, remaining populations uncertain.
with some animal food including insects; may travel
several kilometres a day around territory to visit PILEATED GIBBON
scattered fruiting trees. Hylobates pileatus PLATE 36
Distribution SE Asia: Myanmar, Thailand, NW Laos Measurements HB 470–600. Wt 4–7kg
and Peninsular Malaysia. Also China (S Yunnan) and Identification Sexually dimorphic in adults. Adult
N Sumatra. males (age 3–4 years or older) are all black except
Status Endangered. Populations have declined white feet and hands; thick white eyebrow band
severely owing to loss of forest habitat and hunting. that continues in a partial ring around the eyes;
Although significant numbers remain in some crown hair flattened towards back of head, with
protected areas, major threat is now illegal hunting whitish hairs over ears; white genital area. Females
for food and trapping for the pet trade. are grey or greyish-buff, with dark patch on belly
and dark crown, white eyebrows and partial rings
AGILE GIBBON around eyes. Young are greyish-white, acquiring
Hylobates agilis PLATE 36 black patches on chest and top of head as they
Measurements HB 450–650. Wt adult males mature. Voice Loud call involves series of ‘hoo-ha’
4.9–7.3kg, females 4.5–6.8kg notes from male, and series of rising hoots from

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 female that accelerates into a long bubbling trill; three to six individuals. Feeds mainly on fruits as
female great call averages 60–80 notes over 15–25 well as leaves and buds, with a very small amount of
seconds. Similar species White-handed Gibbon, animal matter such as insects. Male song may begin
H. lar, dark phase has complete white facial ring and before dawn, with male-female duets occurring
lacks pale tufts over ears; pale phase does not have mainly one to three hours after dawn.
dark patches on underparts and head; great call of Distribution SE Asia: N Myanmar. Also NE India
female lacks accelerating trill. Limited hybridisation (Assam) and SW China (SW Yunnan).
between H. lar and H. pileatus has been reported in Status Endangered. Major population declines due
areas of overlap in SE Thailand. to ongoing loss of habitat, especially in India, as well
Ecology and habitat Arboreal, found principally in as hunting.
tall forests. Occurs in small family groups. Calling
typically starts soon after daybreak, but may Genus Nomascus These gibbons have tufts of hair
continue to late morning. Diet consists mainly of on the top of their head, forming a crest, and patches
fruits, especially figs, and also young leaves and of bushy hair on their cheeks that are contrastingly
shoots, with some insects. pale in some species. Juveniles of most species go
Distribution SE Asia: SE Thailand, SW Laos and through a range of colour changes, starting all buffy
Cambodia W of Mekong. with a dark face, then progressing to dark body with
Status Endangered. Although population appears to pale head that eventually turns black. Although some
be stable in Thailand, range is very restricted and species can be difficult to tell apart, there is limited
population is declining in Cambodia. overlap in range among the species.

HOOLOCK GIBBON WESTERN BLACK-CRESTED GIBBON

Hoolock hoolock PLATE 37 Nomascus concolor PLATE 37
Measurements HB 450–650. Wt 6–9kg Measurements HB c.500. Wt 6–9kg
Identification Adult male largely black with bushy Identification Male completely black, with small
white eyebrows, sometimes with a tuft of white on throat sac visible only during loud vocalisations.
chin; adult female buffy-brown to brown, middle of Female greyish-brown with extensive black on
underparts and cheeks darker, often blackish; white underside, as well as dark streak on crown and
face-ring and white streak from cheeks up under eyes inside of limbs. Juvenile completely black. Both
and across top of nose. Newborn young are white, sexes have hairs on crown of head raised in a
but turn all black by end of first year. In populations tuft. Voice Males produce a low boom (during
W of Chindwin River in Myanmar (H. h. hoolock) inflation of throat sac), short, staccato single notes,
males have eyebrows joined together and black and ‘whoops’ that rise, or fall then rise rapidly in
underparts; E of Chindwin River (H. h. leuconedys, frequency; females produce only the last of these
sometimes considered a separate species) eyebrows in their great call. Similar species Hainan Crested
are separated, male tends to be pale brownish on Gibbon, N. hainanus, male is very similar, but female
underparts; intermediates occur in headwaters of lacks black on underparts; other Nomascus gibbons
Chindwin. Small laryngeal sac is present in both have pale cheeks in males; Hoolock Gibbon, Hoolock
sexes. Voice Male and female vocalisations similar, hoolock, male has white eyebrows.
consisting of mixture of two-part notes (‘ow-wa’), Ecology and habitat Arboreal and diurnal. Largely
simple hoots and growls; these are interspersed in an restricted to tall forests in sub-tropical and montane
accelerating duet during the great call. Taxonomic forests. Feeds on fruits, leafbuds and shoots.
notes Genus formerly called Bunopithecus. Similar Distribution SE Asia: restricted areas in parts of
species White-handed Gibbon, Hylobates lar, has N Laos and N Vietnam, N of about 20°N. Also China
white hands, call lacks ‘ow-wa’. (S Yunnan).
Ecology and habitat Arboreal and diurnal, being Status Critically Endangered. Formerly widespread
most active from dawn until middle of afternoon. in much of S China and N Indochina, but global
Found mainly in primary evergreen and some semi- population now estimated at fewer than 2,000
evergreen forests. Occurs in small family groups of animals, restricted to a few isolated areas with

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 no more than a few hundred in each of Laos orange or buff with no dark hairs on underparts;
and Vietnam. Several known populations have dark brown elongate patch on crown, white hairs
disappeared recently. Threats include loss of habitat, around face, forming a distinct face-ring. Voice
illegal hunting and trapping. Similar to that of Black-crested Gibbon, N. concolor:
the male has a mixture of a low boom, short hoots
BUFF-CHEEKED GIBBON and strongly frequency-modulated whoops, while
Nomascus gabriellae PLATE 37 the female has a series of rising whoops that
Measurements HB c.500. Wt 6–10kg increase in pitch and frequency during the great call,
Identification Male black with pale reddish or typically with more notes than N. concolor. Similar
reddish-yellow patches on cheeks rounded at top, species Buff-cheeked Gibbon, N. gabriellae, male
reaching less than halfway up ears; hairs on top of has more reddish cheeks with hairs that stick out to
head raised into a crest; chest brownish, contrasting side, brownish patch on chest; female has indistinct
with rest of body. Female orange-brown, rarely with buff face-ring.
any white on face, dark triangular patch on crown, Ecology and habitat Arboreal and diurnal. Largely
no dark on belly. Hairs on cheeks appear brushed restricted to tall forests.
sideways in both sexes. Voice Male produces short Distribution SE Asia: Laos and Vietnam, E to Black
hoots as well as descending long hoots that each River, N of about 19°N. Also adjacent extreme
end in a sharp upwards note; female produces long, S China (SW Yunnan), although Chinese population
upwardly modulated calls that increase in pitch and may now be extinct.
frequency during the great call. Taxonomic notes Status Critically Endangered. Remaining populations
The more northern populations in the Annamite highly fragmented and greatly reduced due to loss of
mountains were proposed in 2010 as a separate habitat, illegal hunting and trapping.
species, N. annamensis, based on small genetic
differences and differences in the voice, but they are SOUTHERN WHITE-CHEEKED GIBBON
indistinguishable in appearance and could also be Nomascus siki PLATE 37
considered subspecies. Similar species Southern Measurements HB c.500. Wt 6–10kg
White-cheeked Gibbon, N. siki, has paler cheeks Identification Males and juveniles black except
and darker chest in males; more distinct white face- for white cheek whiskers; cheek patches reach
ring in females. only halfway up ears with a pointed upper end,
Ecology and habitat Arboreal and diurnal. Found but extend at bottom to margin of upper lips and
in tall evergreen and semi-evergreen forest. Feeds onto sides of chin. Female creamy orange with dark
mainly on fruits. Group size typically three to five triangular patch on crown, white ring around face.
individuals. Voice Similar to that of Northern White-cheeked
Distribution SE Asia: E Cambodia, S Laos, Gibbon, N. leuocogenys, but typically with fewer
S Vietnam from Mekong Delta N to about 15°30’N. notes in great call. Similar species Northern White-
Status Endangered. Although more common cheeked Gibbon, N. leucogenys, is very similar, but
than other crested gibbons (Nomascus spp.), it with longer hair, and male has white cheek patches
has disappeared from many suitable areas, and higher on face; Buff-cheeked Gibbon, N. gabriellae,
habitat has been greatly reduced in parts of range, male has orange or yellow cheeks with pale patch
especially in Vietnam. Threats include loss of habitat, on chest, female lacks white ring around face.
illegal hunting and trapping for food and pet trade. Ecology and habitat Arboreal and diurnal. Largely
restricted to tall forests.
NORTHERN WHITE-CHEEKED GIBBON Distribution SE Asia: C Vietnam and Laos between
Nomascus leucogenys PLATE 37 15°45’N and 20°N.
Measurements HB c.500. Wt 6–10kg Status Endangered. Although moderate populations
Identification Male largely black with some silvery continue to persist, many families occur in forest
hairs, except for white or very pale yellow cheek-tufts patches too small to ensure long-term persistence,
that are pointed at top, reaching level of top of ears and species is still threatened by ongoing loss of
but not touching corners of mouth; female creamy forest, illegal hunting and trapping.

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 EASTERN BLACK-CRESTED GIBBON to Hainan. Similar species Western Black-crested
Nomascus nasutus NOT ILLUSTRATED Gibbon, N. concolor, is very similar but female
Measurements HB c.500. Wt 6–10kg has dark patches on underparts; other Nomascus
Identification Adult male all black with relatively gibbons have pale cheeks on males.
long hair; adult female light brownish-grey to Ecology and habitat Arboreal and diurnal. Largely
brownish-yellow, with no black on underparts, restricted to tall tropical valley and montane
indistinct elongate oval dark patch on crown. Voice rainforests.
Male produces short, staccato notes, and sharply Distribution SE Asia: Cao Bang province in Vietnam,
rising whoops; female great call is strongly vibrato, NE of Red River.
like a trill, increasing in pitch and accelerating to Status Critically Endangered. Fewer than 250
end. Taxonomic notes Formerly considered a form remaining wild adults.
of N. hainanus, which is now considered restricted

Order CARNIVORA
Dogs, bears, martens, weasels, otters, civets, mongooses, cats and seals
The carnivores are a diverse group of mammals that have largely evolved, as their name implies, with
adaptations for hunting and killing vertebrate prey, ranging from fish in the case of otters, to large mammals
in the case of tigers and leopards. However, many species also eat plant food, especially fruits, and some
of the civets are largely frugivorous. There is considerable variation among families in the skull and form of
dentition, as might be expected given this variety of diets.

Family CANIDAE DOGS

The skull of a dog has a relatively elongate muzzle (Fig. 49), with dentition adapted for eating meat. The
dental formula is 3/3 1/1 4/4 2/2. The canine is relatively large and pointed; there are usually three
upper and lower incisors, with a gap (diastema) between the incisors and the upper canine, into which
the lower canine fits when the mouth is closed. The last upper premolar and first lower molar are formed
into carnassials – elongated sharpened teeth used for tearing flesh. There are only two upper molars, the
last relatively small.
Apart from the Domestic Dog, Canis familiaris, which occurs throughout the region, there are four species
of wild dog currently known to occur in South-east Asia. Each of these has a distinctive shape and coloration,
and is fairly easily separated from others if seen well. However, care must be taken not to confuse them with
free-ranging Domestic Dogs, which can show a wide variety of shapes and colour patterns.
The footprints of dogs have a small pad on the sole and four large pads on the toes, with broad, blunt
nails (Fig. 50); those of wild dogs are difficult to distinguish from those of Domestic Dogs.

GOLDEN JACKAL Domestic Dogs, Canis familiaris, rarely have black

Canis aureus PLATE 38 saddle pattern.
Measurements HB 600–800, T 200–250. Ecology and habitat Largely found in drier, open
Wt 7–10kg areas. Forms large groups in India, but in Thailand
Identification Generally greyish-brown to golden- usually seen singly or in pairs. Groups may call at
brown, with black-tipped hairs on shoulders and night, especially just after dusk and before dawn.
back tending to form a saddle-like pattern. Ears Distribution SE Asia: Myanmar, Thailand, Laos,
broadly pointed, tail moderately bushy and dark at Vietnam and Cambodia. Also from N Africa, S Europe
tip. Voice A series of ascending whines, followed by through to India.
several quick yelps. Similar species Dhole, Cuon Status Least Concern. Generally tolerant of human
alpinus, is larger and heavier, more reddish-brown disturbance, but has declined in some areas owing
without back pattern, all dark tail; free-ranging to excessive trapping.

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 a

F H

0 50 0 100

Fig. 49 Skulls of two large carnivores: Dhole, Cuon Fig. 50 Footprints of Dhole, Cuon alpinus, in soft soil
alpinus (a), and Leopard, Panthera pardus (b). (top), and on firm soil (bottom).

RED FOX RACCOON DOG

Vulpes vulpes PLATE 38 Nyctereutes procyonoides PLATE 38
Measurements HB 550–700, T 350–450. Measurements HB 500–680, T 130–250.
Wt 3.5–5.5kg Wt 4–6kg
Identification Relatively slender canid with long tail, Identification Generally yellowish-brown, with black
pointed face and pointed ears. Usually upperparts tip on shoulders, back and tail. Black ‘mask’ on face
uniformly reddish-brown; paler underneath; that goes around eyes and under chin. Similar
bushy, reddish tail sometimes with white tip, species Some civets also have dark patterns
though usually all dark in Asian populations; black on face, but are a different shape with a much
behind ears. Some animals are all dark brown. longer tail.
Voice A high-pitched bark and various other calls. Ecology and habitat Mainly in forested areas.
Similar species Dhole, Cuon alpinus, is considerably Largely solitary, most active at night. In northern
heavier, with a thicker muzzle, dark tail and rounded parts of range, species hibernates after increasing
ears with white centre; Golden Jackal, Canis aureus, weight by nearly 50%, but presumably not farther
is greyish-brown and has a shorter tail. south, although little is known of the species in
Ecology and habitat No information from region, SE Asia. Feeds on small animals, including rodents,
but elsewhere found in a wide variety of habitats frogs, fish, crabs and insects, as well as fruits.
from forests to open areas that have been heavily Distribution SE Asia: N Vietnam. Also S Siberia
disturbed by humans. Feeds on small mammals, through E China and Japan, and widely introduced
birds, frogs, insects and sometimes fruits. in W Russia and E Europe.
Distribution SE Asia: N Vietnam. Also throughout Status Least Concern, but has become very rare
much of northern hemisphere, including Europe in region, probably owing to loss of forest and
through Asia to Himalayas, N India and parts of excessive trapping.
China, as well as North America.
Status Least Concern, although status in region poorly
known; may only be an irregular visitor from China.

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 DHOLE Status Endangered. Has declined dramatically owing
Cuon alpinus PLATE 38 to loss of habitat, loss of prey base and hunting.
Measurements HB 800–1,050, T 300–450,
SH 420–550. Wt male 15–21kg, female 10–17kg DOMESTIC DOG
Identification Largest and heaviest wild dog in Canis familiaris NOT ILLUSTRATED
region. Body reddish-brown, often paler underneath. Measurements HB usually 800–1,000, T usually
Tail long and bushy, largely black or very dark. Ears 300–350
rounded with conspicuous long white hairs inside. Identification Domestic Dogs have been bred into
Voice Relatively quiet, but occasional yaps or a tremendous diversity of shapes and sizes, ranging
whistling sounds. Similar species Domestic Dogs, from miniature ‘toy’ forms to extremely large forms.
Canis familiaris, are sometimes a similar reddish Free-ranging dogs may be encountered in wild areas
colour, but if so they lack the long, bushy black tail in almost any part of the region. Some of these are
and rounded ears with white centre. Golden Jackal, similar in shape to some of the wild dogs; many are
Canis aureus, is smaller, more greyish-brown, with dull orange-brown, although black or pied forms are
dark saddle-like pattern on back. often encountered. Nevertheless, they rarely have
Ecology and habitat Found in a wide variety of the exact coloration and shape of the wild forms,
forested areas, including evergreen forest, montane and are usually less shy.
forests and semi-open forest. Avoids contact with Ecology and habitat Usually associated with humans.
humans. Hunts in packs, usually at night. Preys on a Feral (free-ranging) dogs occasionally encountered far
variety of animals, including pigs and deer. from human settlements, often in poor condition.
Distribution SE Asia: Myanmar, Thailand, Laos, Distribution SE Asia: may be found throughout
Vietnam, Cambodia and Peninsular Malaysia. Also region, especially in areas where there are human
S Siberia to India, Sumatra and Java. settlements.

Family URSIDAE BEARS

Bears are large, powerfully built omnivorous mammals with a short tail and good sense of smell, but poor
sight and hearing. Two species of bear occur in the region, both of them mainly black apart from pale
markings on the chest. The species can be distinguished by fur texture and length, head and ear shapes,
and size. Despite mainly feeding on fruit and plant material, insects or smaller animals, bears are potentially
dangerous to humans, owing to the unpredictability of their behaviour. The Red Panda, Ailurus fulgens, has
sometimes been placed in the bear family, but is here considered to be in its own family, the Ailuridae.
Bear footprints have an elongate sole, slightly triangular with five toe prints all similar in size, and very long
claw marks, longest on the front feet (Fig. 51). The prints of the Asian Black Bear, Ursus thibetanus, are larger
than those of the Sun Bear, Helarctos malayanus.

SUN BEAR trees. The claw marks are much more conspicuous
Helarctos malayanus PLATE 39 than the faint marks of other carnivores or monitor
Measurements HB 1.1–1.4m, T 30–70. lizards. Voice Occasionally utters hoarse grunts
Wt 27–63kg or loud roars; rarely short barks like that of a
Identification Body entirely black, except for grey muntjac, Muntiacus spp. Similar species Asian
muzzle and a white or yellowish V- or C-shaped Black Bear, Ursus thibetanus, is much heavier, with
mark on the upper chest. Chest mark normally long shaggy fur, large ears; Binturong, Arctictis
prominent but occasionally very faint. Ears short binturong, is much smaller, with a prominent long
and rounded. Fur short and smooth. Has distinct bushy tail.
posture when walking, with head held very low. Ecology and habitat Active periodically during day
Signs are more often seen than the bear itself: and night, on the ground and in tall trees. Builds
prominent claw marks gouged into a tree trunk nests of small branches in tall trees for sleeping.
where it has climbed the tree, or remains of bee Diet includes bees’ nests, termites, small animals,
or termite nests ripped open in standing or fallen fruits and the hearts of coconut palms. Occurs in

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 extensive areas of forest, but sometimes enters
gardens or oil palm plantations.
Distribution SE Asia: Myanmar, Thailand, Laos,
Cambodia, Vietnam and Peninsular Malaysia. Also
Sumatra and Borneo. F
Status Vulnerable due to loss of forest as well as
illegal hunting and trapping.

ASIAN BLACK BEAR

Ursus thibetanus PLATE 39
Measurements HB 1.2–1.5m, T 65–100, HF 175–
195, E 120–180. Wt up to 100kg or more
Identification A large bear, mainly black apart from a
white ‘V’ on the chest and a pale muzzle. Pale marks
on muzzle rarely extend as far back as eyes. Has large
erect ears and long shaggy hair. Signs include deep
claw marks on trees. Occasionally, brown or blonde H
individuals have been reported. Similar species Sun
Bear, Helarctos malayanus, is about half the body
weight, has short fur, short rounded ears that barely
extend above head in profile, more stooped posture
and more extensive pale area on muzzle.
Ecology and habitat Generally solitary, except in
breeding season. Mainly in forested areas, both 0 100
evergreen and deciduous. Sometimes enters cultivated
areas near forest. In southern part of range, found Fig. 51 Prints of left front (F) and hind (H) feet of
only in hill forest. Usually rests during day in hollow Sun Bear, Helarctos malayanus.
trees, caves or rock crevices, emerging in the evening
to feed, but sometimes active during the day. Feeds Cambodia and Vietnam. Also Pakistan, India, Nepal,
mainly on fruits, berries, beehives, invertebrates and China, Siberia and Japan.
small vertebrates such as amphibians and rodents, but Status Vulnerable; declining due to illegal hunting
sometimes hunts larger mammals as well. and trapping, especially for alleged medicinal
Distribution SE Asia: Myanmar, Thailand, Laos, values, as well as loss of forest habitat.

Family AILURIDAE RED PANDAS

Red Pandas have sometimes been placed in the same family as bears, along with the Giant Panda. Other
researchers have considered them part of the Procyonidae or Raccoon family. They are currently considered to
belong in their own family. They are readily distinguished by colour and shape from all other animals in the region.

RED PANDA long bushy tail and black marks under the eyes.
Ailurus fulgens PLATE 39 Ecology and habitat Found mainly in montane
Measurements HB 510–630, T 280–480. Wt 3–6kg forests at 2,500–4,000m, often in bamboo thickets,
Identification Body fur long and soft, upperparts though sometimes at lower altitude. Mainly nocturnal,
reddish, darkest in middle of back; underparts sleeping curled on a branch during the day. Climbs
paler. Tail long and bushy, inconspicuously ringed. well, but also feeds on the ground on bamboo shoots,
Head pale, with white rim to ears, dark marks under grasses, roots, fruits, acorns and sometimes insects,
eyes. Voice Includes short whistles and squeaking eggs and small vertebrates. In winter is largely
notes. Similar species No other mammal in region dependent on bamboo leaves for food. May be
has this combination of colour and shape, with solitary, in pairs or in small family groups.

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 Distribution SE Asia: N Myanmar. Also Nepal, India, habitat, fragmentation of remaining habitat leading
Bhutan and S China. to isolation of populations, and illegal hunting and
Status Endangered. Major threats include loss of trapping.

Family MUSTELIDAE MARTENS, WEASELS, BADGERS AND OTTERS

The mustelids are a rather diverse group and are usually divided into subfamilies. The subfamily Mustelinae
includes martens and weasels, both slender, agile mammals adapted to hunting small vertebrate animals,
as well as invertebrates. Martens tend to be more arboreal than weasels. The subfamily Melinae (which
is sometimes combined with the Mustelinae) includes the ferret-badgers and Greater Hog Badger. The
ferret-badgers are much smaller than the Greater Hog Badger with distinctive patterns as a genus,
but the species are hard to tell apart unless the animal is captured so that the teeth can be examined.
The subfamily Lutrinae includes the otters, which can be distinguished by their semi-aquatic existence,
webbed or partially webbed feet, and broad muzzle. The different species of large otter are difficult to
distinguish in the field, unless seen well.
Mustelids appear to be closely related to the Viverridae, or civet family, but can usually be distinguished
by having 34 to 38 teeth, whereas all viverrids have a total of 40 teeth. Mustelids also tend to have stockier
limbs and broader feet, and are generally more strictly carnivorous.
Footprints vary considerably within the family, though all species show five toe prints, unlike dogs or cats,
which show only four. The sole pad has a relatively complex pattern of lobes (Fig. 52). Greater Hog Badger
prints have distinctive long claw marks (Fig. 52d), while otter prints usually show distinct webbing between
the toes (Figs 52f, g, h) and sometimes do not show claw marks.

STONE MARTEN Back colour varies from medium brown to pale

Martes foina PLATE 40 yellowish-brown; lower back, hind legs, lower half
Measurements HB 400–540, T 220–300 (about of front legs and tail contrastingly darker, varying
55–60% of HB). Wt 1.1–2.3kg from dark brown to black; conspicuous black stripe
Identification Colour mostly brown, frequently with on side of neck behind ear; top of head varies from
reddish, yellowish or grey tinge; underside slightly black, continuous with neck stripe, to dark brown.
paler than upperparts. Throat white or yellowish with Chin, throat and chest yellowish, whitish or buff,
variable brown patches, sometimes largely brown. contrasting with slightly darker lower belly. In the
During moult, fur can have pale creamy white field, the long, slender body, long tail and lithe,
patches. Similar species Yellow-throated Marten, bounding motion are distinctive. Tail may be held
M. flavigula, has longer tail, yellow throat with dark down, up with the tip drooping, or almost horizontal
head and/or sides of throat; weasels, Mustela spp., (if moving quickly). Voice A series of soft, rapid
are much smaller with a more slender body. ‘chuk’s. Similar species Stone Marten, M. foina,
Ecology and habitat In region, usually at high lacks dark border to throat and has shorter tail;
altitudes, 1,300m up to over 3,000m. Inhabits weasels, Mustela spp., with white throats have
conifer forests or open rocky areas. shorter tail and lack the contrasting black or dark
Distribution SE Asia: N Myanmar. Also Europe brown pattern on the neck, lower legs and tail;
through to China, Himalayas. mongooses have a more pointed muzzle, tend to
Status Least Concern. Only marginally occurs in hold head and tail down while walking, and walk
region where status is uncertain, but populations quickly but rarely bound; civets are normally active
apparently stable elsewhere. only at night and have different colour patterns.
Ecology and habitat Active mainly during the day,
YELLOW-THROATED MARTEN but sometimes at night. Agile, moving fairly quickly
Martes flavigula PLATE 40 on the ground or in the tree canopy. Diet includes a
Measurements HB 450–650, T 370–450, wide range of small vertebrates and invertebrates,
HF 90–110, E 28–40. Wt 1.3–3.0kg bees’ nests and nectar. Rests in tree holes and
Identification Colour pattern varies geographically. on large branches. Usually alone or in pairs, but

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 sometimes in family groups. Occurs in a wide variety same colour as back. Similar species Yellow-bellied
of forest types, both natural and disturbed. Often Weasel, M. kathiah, is pale yellow underneath and
enters plantations and gardens in search of food. lacks mask; Malay Weasel, M. nudipes, has white
Occurs from sea level up to 4,500m altitude. head extending to behind ears.
Distribution SE Asia: Myanmar, Thailand, Laos, Ecology and habitat Found in a variety of habitats
Vietnam, Cambodia and Peninsular Malaysia. Also from dense forest to dry areas and villages, mainly
Pakistan, Nepal, India, China, Russia, Sumatra, Java at 1,500–5,000m, but can occur at lower altitudes
and Borneo. outside region. Eats small animals including rats,
Status Least Concern, although some populations and has been known to attack domestic fowl.
may have declined due to loss of forest or hunting. Distribution SE Asia: N Myanmar, N Thailand, Laos.
May be expected in N Vietnam. Also Himalayas,
MALAY WEASEL China, Siberia, Taiwan, Korea and Japan.
Mustela nudipes PLATE 41 Status Least Concern, but has declined recently in
Measurements HB 300–360, T 220–260, parts of China.
HF 49–55, E 23–25. Wt 1kg
Identification Body colour typically orange to TONKIN WEASEL
golden-brown, but sometimes more greyish- Mustela tonkinensis PLATE 41
brown. Basal half of tail same colour as back, but Measurements HB 200, T 90 (45% of HB)
distal half is typically all white, though sometimes Identification Smallest weasel in region, with
only the extreme tip. Head white to behind ears. brown upperparts, white underparts, relatively
Tail bushy. Similar species Siberian Weasel, short tail. Taxonomic notes Formerly considered
M. sibirica, has white limited to front of muzzle, a subspecies of the widespread Mustela nivalis,
usually has a dark mask around eyes, and does not which is no longer considered to occur in the region.
overlap in range; some large squirrels are similar Similar species Other weasels in region are larger,
in size and proportions, but have differently shaped with longer tail.
heads, and none has same colour pattern of pale Ecology and habitat Unknown. Only recorded
head and tail tip, with orange or reddish body. specimen likely from a montane area. The related
Ecology and habitat Mainly terrestrial. Apparently Mustela nivalis is found elsewhere in various
active both day and night. Sleeps in holes in the habitats, including forested, open and disturbed
ground. Diet includes small animals. Recorded in areas.
lowland and hill rainforests up to 1,700m altitude. Distribution SE Asia: known from only one specimen
Also uses disturbed areas, and appears to be from Chapa, Tonkin, Vietnam.
tolerant of a range of habitats. Status Data Deficient. Unknown whether the
Distribution SE Asia: peninsular Thailand and species is very rare or simply overlooked.
Malaysia. Also Sumatra and Borneo.
Status Least Concern, although it is rarely YELLOW-BELLIED WEASEL
encountered, and there are likely some declines due Mustela kathiah PLATE 41
to clearing and conversion of forests. Measurements HB 200–290, T 130–180
(60–70% of HB)
SIBERIAN WEASEL Identification Upperparts dark chocolate-brown,
Mustela sibirica PLATE 41 sometimes with reddish tinge; underparts yellow or
Measurements Male: HB 280–390, T 150–210, light orange-brown with whitish on chin and upper
HF 40–50, Wt 0.65–0.82kg. Female: HB 250– lip; tail moderately long and bushy, same colour
305, T 130–165, HF 35–45. Wt 0.35–0.45kg as upperparts. Similar species Siberian Weasel,
Identification Colour of body varies from golden- M. sibirica, has dark underparts nearly same colour
brown and dark chocolate to reddish, only slightly as upperparts, except for pale throat and muzzle.
paler underneath. Pale patch on front of muzzle and Ecology and habitat Has been reported from
often upper throat, usually with dark mask around forested areas 1,000m or higher, including above
and in front of eyes. Tail medium-long and bushy, tree line, though sometimes also in lowlands. May

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 a
f
e
F H

F F
H H

F H g

F H F H

d h

F
F H H
0 100

Fig. 52 Prints of left front (F) and hind (H) feet of selected Mustelidae and Herpestidae: Mustela nudipes (a), Martes
flavigula (b), Melogale sp. (c), Arctonyx collaris (d), Urva javanica (e), Amblonyx cinerea (f), Lutra lutra (g) and Lutrogale
perspicillata (h).

be tolerant of disturbed habitats. Feeds on birds, throat, extending into a buff or cream stripe down
rodents and other small mammals. the centre of the belly. Tail medium-long and bushy.
Distribution SE Asia: N. Myanmar, Laos, Thailand, Similar species No other species of similar size
Cambodia and Vietnam. Also Nepal, N India (Assam) and shape has a single pale stripe down the middle
and S China. of the back; Javan Mongoose, Urva javanica, lacks
Status Least Concern. stripe and differs in shape.
Ecology and habitat Primarily recorded from
STRIPE-BACKED WEASEL evergreen forest in hills and mountains up to
Mustela strigidorsa PLATE 41 2,500m, but also some records in disturbed areas
Measurements HB 275–325, T 145–205 (c.60% and scrub. Has been observed foraging on the
of HB), HF 47–54, E 20–23 ground, where it presumably catches a range of
Identification Dark chocolate-brown to reddish- small animals.
brown above and below, including tail, except for a Distribution SE Asia: Myanmar, Thailand, Laos and
thin white or buff stripe down the back, from the top Vietnam. Also Nepal, NE India (Assam) and S China.
of the head to the base of the tail, and a yellowish- Status Least Concern. Recent surveys suggest
white patch on the lower cheeks, chin and upper more common than previously thought.

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 LARGE-TOOTHED FERRET-BADGER rostrum; further taxonomic studies are required to
Melogale personata PLATE 40 confirm the taxonomic names and to determine
Measurements HB 330–390, T 145–210, distinguishing features of live animals. Similar
HF 55–70. Wt 1–3kg species Large-toothed Ferret-badger, M. personata,
Identification Head has distinct pattern of black has longer white stripe on back, more white on tail,
(or dark brown) and white; pattern varies among large teeth; Masked Palm Civet, Paguma larvata,
individuals. Body brownish to greyish, paler also has black-and-white face pattern, but is much
underneath, sometimes with orange tinge. Sides of larger, more slender, with much longer tail.
body heavily frosted white, contrasting slightly with Ecology and habitat Nocturnal and largely
darker back. White stripe on back of neck extending terrestrial, although can climb trees. Ecological
at least to middle of back, and often as far as rump differences from M. personata not well understood.
or tail; bushy tail pale, usually white on distal half. Distribution SE Asia: N Myanmar, Laos and Vietnam.
Large, massive teeth, especially fourth premolar, Also E India (Assam), C and SE China, Hainan and
which is over one-third length of all cheek teeth (Fig. Taiwan.
53b). Similar species Small-toothed Ferret-badger, Status Least Concern.
M. moschata, tends to have shorter, interrupted
white back streak, white on tail restricted to tip, GREATER HOG BADGER
more black on face; reliably identified only by much Arctonyx collaris PLATE 40
smaller teeth (Fig. 53a). Measurements HB 550–700, T 120–170,
Ecology and habitat Nocturnal and largely HF 115–135. Wt 7–14kg
terrestrial, digging burrows with its strong claws, Identification A large badger with a long, pig-like
but also sometimes climbs trees. Diet mainly snout, short tail, dark stripes on head. Overall body
invertebrate prey such as large insects, snails, colour varies from greyish to brown. Stripes on head
earthworms. Found in forested areas as well as vary from dark brown to black, and vary in extent,
grasslands and rice fields. with lower dark stripe sometimes reduced. Similar
Distribution SE Asia: Myanmar, Thailand, Laos, species Ferret-badgers, Melogale spp., are much
Cambodia and Vietnam. Also Nepal and India smaller, with longer tail.
(Assam). Ecology and habitat Nocturnal and terrestrial,
Status Least Concern, but some uncertainty sleeping during the day in burrows. A powerful digger,
in status as many field records have not been feeding on tubers and roots as well as invertebrates
distinguished from M. moschata.

SMALL-TOOTHED FERRET-BADGER a b
Melogale moschata PLATE 40
Measurements HB 330–380, T 140–160
(40–45% of HB)
Identification Distinct black-and-white head
pattern, generally with more black than M. personata
but quite variable. Body colour greyish to light brown,
with flanks similar colour to back, belly whitish. White
stripe on top of head narrow and incomplete, rarely
extending much past shoulders. Tail proportionately
shorter than that of Large-toothed Ferret-badger,
M. personata, darker with only tip white. Teeth
relatively small with distinct gaps between premolars
(Fig. 53a). Taxonomic notes The species Melogale
cucphuongensis was described in 2008 from Fig. 53 Skulls of Melogale moschata (a) and
Vietnam, based on a single specimen. It apparently M. personata (b), showing difference in relative and
has similar teeth to M. moschata, but a narrower absolute size of teeth.

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 such as worms, insects and other small animals. externally. Skull flatter than that of Smooth Otter,
Usually in forests as well as plantations near forests. Lutrogale perspicillata (Fig. 54c). Voice Contact call
Mainly in hills, but also in lowlands in Cambodia. between otters a single-syllabic chirp; adult females
Distribution SE Asia: Myanmar, Thailand, Laos, call to cubs with a staccato chatter. Similar species
Vietnam and Cambodia. Also NE India. Note that Eurasian Otter, L. lutra, has long, pale guard hairs
populations in China and those in Sumatra are now on a dark coat, underparts paler than upperparts;
each recognised as separate species. Smooth Otter, Lutrogale perspicillata, can be hard
Status Vulnerable. Populations declining due to to distinguish in field, but has smoother coat, paler
hunting and loss of habitat. underparts and hairless rhinarium.
Ecology and habitat Occurs in coastal areas and
EURASIAN OTTER on larger inland rivers, solitary or in groups of up to
Lutra lutra PLATE 42 four. Diet includes fish. Pairing of male and female
Measurements HB 550–720, T 375–480, may be limited to the breeding period.
HF 100–130
Identification Upperparts rather dark brown, with
paler chin and upper throat. Underparts markedly a
paler. Fur dense, consisting of short hairs and longer,
paler guard hairs, producing a somewhat grizzled
appearance. Rhinarium (tip of nose) hairless; edge
of fur and nose forms ‘W’ shape. End of tail circular
in cross-section. SE Asian form, L. lutra barang, is
smaller than races found in Europe. Similar species
Hairy-nosed Otter, L. sumatrana, and Smooth Otter,
Lutrogale perspicillata, generally paler, without
frosted appearance, and live primarily in coastal b
and flatland habitats; Oriental Small-clawed Otter,
Amblonyx cinereus, is substantially smaller.
Ecology and habitat Diet includes fish, other small
vertebrates and crustaceans. Most SE Asian records
are from hill or mountainous areas in streams and
lakes.
Distribution SE Asia: Myanmar, Thailand, Laos
and Vietnam. Also Europe, N Africa, through much
of N Asia, India, Sri Lanka, China, Taiwan, Japan,
Sumatra and possibly Borneo.
Status Near Threatened globally. However, status
c
in region very poorly known; likely to be declining
owing to loss of habitat, including damming of rivers,
as well as hunting.

HAIRY-NOSED OTTER
Lutra sumatrana PLATE 42
Measurements HB 500–800, T 370–500,
HF 105–130
Identification Entirely brown, except lips, chin and
upper throat, which are whitish. Fur rather rough but 0 20
short. Rhinarium covered in hair. Tail flattened, oval in
cross-section. Feet fully webbed between the digits. Fig. 54 Skulls of Amblonyx cinerea (a), Lutragale
Claws prominent. Penis of adult male not visible perspicillata (b) and Lutra sumatrana (c).

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 Distribution SE Asia: Myanmar, S Thailand, Status Vulnerable. Population declining owing to
Cambodia, S Vietnam and Peninsular Malaysia. Also loss and degradation of lowland wetland habitats,
Sumatra and Borneo. as well as hunting.
Status Endangered. Relatively few confirmed
records. Species is threatened by loss of lowland ORIENTAL SMALL-CLAWED OTTER
wetland habitats and hunting for fur and meat. Amblonyx cinereus PLATE 42
Measurements HB 360–550, T 225–350,
SMOOTH OTTER HF 85–110
Lutrogale perspicillata PLATE 42 Identification Upperparts dark brown or greyish-
Measurements HB 650–750, T 400–450, brown; underparts slightly paler. Chin, throat,
HF 100–140 cheeks and sides of neck buff. Digits only partially
Identification Upperparts mid-brown, underparts webbed. Claws short, not extending beyond the end
distinctly paler, buffy. Throat and sides of neck cream- of the digits. Voice Contact call is ‘wiuk’; various
coloured. Fur short, smooth and sleek. Rhinarium other calls are also made. Taxonomic notes Has
hairless; edge of fur and nose forms straight line. Tail been placed in genus Aonyx, but recent genetic
flattened on underside, rounded on top. Feet webbed research shows this is more closely related to
only up to last joint on digits. Claws prominent. Penis Lutrogale than the African Aonyx. Similar species
of adult male protrudes beyond body wall. Dorsal Other otters in region are substantially larger and
edge of skull slightly rounded in profile (Fig. 54b). have prominent claws.
Voice Contact call ‘wiuk’. Taxonomic notes Recent Ecology and habitat Diurnal. Diet includes crabs,
genetic research suggests this is closely related to other crustaceans and molluscs, as well as some
Amblonyx cinereus and should potentially be placed fish. Occurs in many habitats where there is
in the same genus. Similar species Hairy-nosed permanent water and some tree cover, from the
Otter, Lutra sumatrana, has belly same colour as coast to inland hill forest, including seacoast, large
back, rhinarium covered with hair, more fully webbed rivers, small hill streams, ponds and lakes. Solitary
feet; Eurasian Otter, Lutra lutra, has long, pale guard individuals sometimes encountered, but often in
hairs on a dark coat, tail rounded in cross-section. large groups.
Ecology and habitat Lives on coast or inland Distribution SE Asia: Myanmar, Thailand, Laos,
in extensive flatlands, including lakes, rivers and Vietnam, Cambodia and Peninsular Malaysia. Also
streams. Several individuals may cooperate in India, S China, Sumatra, Java, Borneo and Palawan.
fishing. Sometimes forages on dry land. Status Vulnerable. Formerly common and
Distribution SE Asia: Myanmar, Thailand, Laos, widespread, but has declined dramatically in much
Vietnam, Cambodia and Peninsular Malaysia. Also of region due largely to hunting and trapping for fur,
Afghanistan, Bangladesh, India, S China, Sumatra, meat and traditional medicine.
Java and Borneo.

Family VIVERRIDAE CIVETS

Civets are a diverse group of carnivores, mostly active at night. Some species are largely terrestrial, while
others spend nearly their whole lives in trees. Most species have a relatively pointed muzzle and long
tail (except the Otter Civet, Cynogale bennetti, which has a broad muzzle and short tail); and a total of
40 teeth. Most civets (except male Small-toothed Palm Civet, Arctogalidia trivirgata) have perineal scent
glands. Linsangs were formerly considered part of this family but are now considered to belong in their own
family, Prionodontidae.
Viverrids share many features with the mustelids, but the latter have fewer teeth and lack perineal scent
glands. Most species can be fairly readily identified by colour patterns.
Footprints of most civets have five distinct toe impressions, except those of the Viverra and Viverricula,
which have only four toes visible and closely resemble the prints of dogs (Fig. 55).

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 LARGE INDIAN CIVET Large Indian Civet, V. zibetha, has proportionately
Viverra zibetha PLATE 43 slightly smaller head, (usually) wavy bands rather
Measurements HB 750–850, T 380–460, than spots on flanks, tail with complete white
HF 110–140. Wt 8–9kg bands; skin sheath on toenails and hairy soles of
Identification Large civet with bold black-and- feet (visible only if animal has been captured); Malay
white throat pattern consisting of three black collars Civet, V. tangalunga, is smaller, with many more
separated by white collars. General colour grey or bands on tail.
tawny-brown with dark pattern on flanks consisting Ecology and habitat Poorly known; recent records
of wavy lines, often with some spotting on legs; are largely from evergreen forest, but it is also found
pattern is sometimes very indistinct. Erectile crest of in disturbed forests. Forages largely at night and
hairs from back of neck to base of tail. Tail banded on ground, hunting for prey in leaf litter. Appears to
to tip, with five or six broad black bands separated be located mainly in lowland forests below 300m,
by narrow but complete white rings. Bottoms of feet although there are some records from higher
heavily haired between pads, with skin sheath over plateaux. Less tolerant of habitat fragmentation than
nail on third and fourth toes. Taxonomic notes other Viverra civets.
Taynguyen Civet, Viverra tainguensis, was described Distribution SE Asia: Myanmar, Cambodia,
from Vietnam, but reanalyses of specimens suggest Thailand, Vietnam, Laos and N Peninsular Malaysia.
it is the same species as V. zibetha. Similar species Also S China.
Malay Civet, V. tangalunga, is smaller, with many Status Endangered. Declining due to forest loss,
more black bands on tail, and black dorsal crest hunting and trapping.
continues along top of tail. Large-spotted Civet,
V. megaspila, has larger head with longer, more MALAY CIVET
swollen muzzle; body with distinct dark spots or Viverra tangalunga PLATE 43
bands on flanks; black crest continuing along top Measurements HB 610–670, T 285–355,
of tail so that white bands are broken; distal half of HF 94–105. Wt 4–5kg
tail usually dark. Identification Upperparts greyish with numerous
Ecology and habitat Nocturnal and largely black spots; a black stripe along the midline extends
terrestrial, although can also climb trees. Found to tip of tail; underparts white with bold black
mainly in forested habitats, but also uses secondary markings on the throat; legs blackish; tail with about
forests and some plantations. Eats any animal it can 15 black bands. Similar species Large Indian and
catch, including lizards, small mammals, birds and Large-spotted Civets, V. indica and V. megaspila, are
fish, as well as fruits and other plant materials. larger, with fewer bands on tail.
Distribution SE Asia: Myanmar, Thailand, Laos, Ecology and habitat Nocturnal and usually
Vietnam, Cambodia and Peninsular Malaysia. Also terrestrial, but occasionally climbs into trees. Diet
Nepal, NE India and S China. includes a wide variety of invertebrates and small
Status Least Concern, although likely to be declining vertebrates taken mainly from the forest floor. May
in some areas due to hunting. visit forest camps to feed on food scraps. Occurs
in forests and cultivated land adjacent to forest.
LARGE-SPOTTED CIVET Tolerant of moderate forest disturbance.
Viverra megaspila PLATE 43 Distribution SE Asia: Peninsular Malaysia. Also
Measurements HB 720–850, T 300–370, Sumatra and adjacent small islands, Borneo,
HF 130–138. Wt 8–9kg Sulawesi and the Philippines. Apparently introduced
Identification Large civet with bold black-and- on several islands.
white throat pattern; body colour varies from greyish Status Least Concern.
to brown, with pattern of dark brown or black spots
on flanks. Pattern very variable, but usually spots are SMALL INDIAN CIVET
distinct, or fused into vertical or horizontal bars. Tail Viverricula indica PLATE 43
with black crest along top, usually dark on distal half, Measurements HB 530–640, T 300–430,
incomplete white bands at base. Similar species HF 85–100. Wt 2–4kg

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 a b
c

F H
F H
F H f

e
d

F H

F H
F H
i
h

F H
F H F H
0 100

Fig. 55 Left footprints of front (F) and hind (H) feet of civets: Cynogale bennetti (a), Arctogalidia trivirgata (b), Prionodon
linsang (c), Viverricula indica (d), Paradoxurus hermaphroditus (e), Hemigalus derbyanus (f), Viverra tangalunga (g),
Paguma larvata (h) and Arctictis binturong (i).

Identification General body colour grey, light COMMON PALM CIVET

grey or buff, with numerous dark spots that form Paradoxurus hermaphroditus PLATE 44
longitudinal rows along back and sides. Throat white Measurements HB 420–500, T 330–420 (usually
with dark lines. Tail with six to nine black-and-white 70–90% of HB), HF 70–76. Wt 2–3kg
rings and long pale tip. Similar species Viverra Identification Upperparts vary from olive-brown
civets are larger, with longer legs, larger, broader or occasionally reddish-brown to dark grey-brown;
muzzle, a dark crest down back, less distinct rows underparts paler. Limbs and tail dark brownish or
of spots, bolder black-and-white markings on throat, black. Cheeks and front of face black, forming a
generally dark tip to tail. dark mask; often with pale spots on sides of head,
Ecology and habitat Primarily nocturnal and which sometimes extend over forehead in a large
terrestrial, but sometimes active during the day. white area; pattern varies among individuals. Three
Frequents relatively open areas, including long grass indistinct, broken dark lines along midline of back,
and scrub; may enter villages or rural buildings. Diet with irregular rows of spots on flanks, sometimes
includes birds, small mammals, frogs, reptiles, forming extra lines. In northern part of range, spot
insects and fruits. pattern can be indistinct and obscured in thick
Distribution SE Asia: Myanmar, Thailand, Laos, winter coat. Adult females have three pairs of
Vietnam, Cambodia and Peninsular Malaysia. mammae. Similar species Small-toothed Palm
Also Pakistan, India, Nepal, Bangladesh, S China, Civet, Arctogalidia trivirgata, has tail longer than
N Sumatra, Java and Bali. head and body; three stripes on back but without
Status Least Concern. additional spots on sides of body; dark head usually

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 with narrow white stripe on forehead; larger skull
with relatively straight toothrow, narrow molars
and widely spaced teeth (Fig. 56b); two pairs of a b
mammae in females. Masked Palm Civet, Paguma
larvata, is larger, lacks spots or stripes on back or
sides, has pale stripe on forehead.
Ecology and habitat Nocturnal; sleeps during day
in trees or sometimes in buildings. Arboreal and
terrestrial, but more often active on the ground
than Small-toothed Palm Civet. Diet includes fruits,
leaves, arthropods, worms and molluscs. Occurs in
tall and secondary forests, plantations and gardens.
Often seen near human settlements.
Distribution SE Asia: Myanmar, Thailand, Laos,
Vietnam, Cambodia and Peninsular Malaysia.
Also Sri Lanka, India, Nepal, Bangladesh, S China,
Sumatra, Java, Borneo, Sulawesi and smaller Fig. 56 Right upper toothrow of Paradoxurus
Indonesian islands and the Philippines. hermaphroditus (a) and Arctogalidia trivirgata (b).
Status Least Concern.
Distribution SE Asia: Myanmar, Thailand, Laos,
MASKED PALM CIVET Vietnam, Cambodia and Peninsular Malaysia. Also
Paguma larvata PLATE 44 Himalayas, including Nepal, N India, China, Taiwan,
Measurements HB 510–760, T 510–640, Hainan, Sumatra and Borneo.
HF 95–104. Wt 3–5kg Status Least Concern, but populations have
Identification Body colour highly variable, from declined in some parts of Vietnam and Laos owing
very light brown, almost blonde, to dark brown to excessive hunting.
or reddish, darker on legs and top of head. Tail
may be dark throughout or only on distal half, BINTURONG
sometimes with white tip. Dark ‘mask’ around Arctictis binturong PLATE 45
eyes and on muzzle, with white on cheeks or Measurements HB 650–950, T 500–800,
sometimes most of head. White stripe on top of HF 120–180. Wt 6–9kg
head from nose to back of neck, shorter and less Identification Long, coarse black fur frosted with
conspicuous in Peninsular Malaysia. Northern whitish or reddish grizzling. Grizzling heaviest
forms (near Himalayas) may have head largely dark around head, making head paler than body. Ears
with only narrow white stripe on forehead. Similar round, edged in white, with long tufts of hair at ends.
species Common Palm Civet, Paradoxurus Tail long, thickly haired, especially near base, and
hermaphroditus, has dark face mask, but lacks prehensile. Similar species Sun Bear, Helarctos
white stripe on forehead and has dark stripes malayanus, is larger, has a short tail and no ear tufts,
and spots on back and sides; Small-toothed and very different shape; melanistic (dark) wild cats
Palm Civet, Arctogalidia trivirgata, has only small have shorter hair, a slender, non-prehensile tail and
white stripe on forehead, no conspicuous ‘mask’, no ear tufts; other civets have shorter fur and are
smaller, more slender build, three stripes on back, rarely all dark.
relatively longer tail. Ecology and habitat Mainly arboreal, but also seen
Ecology and habitat Mainly nocturnal, generally on the ground. Mainly active at night, but sometimes
arboreal, but also travels and hunts on the ground. also during the day. Moves slowly in trees, using tail
Sleeps in tree holes or forks in large trees. Diet for balance and to cling to branches while feeding.
includes fruits and small animals. Occurs in tall Diet includes ripe fruits, especially figs, and small
and secondary forests; may enter plantations and animals. Occurs in tall and secondary forests;
gardens to feed. sometimes in cultivated areas near forest.

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 Distribution SE Asia: Myanmar, Thailand, Laos, those of cats. Similar species Owston’s Civet,
Vietnam, Cambodia and Peninsular Malaysia. Also C. owstoni, has similar pattern, but has dark spots
NE India, S China (Yunnan), Sumatra, Java, Borneo on sides and legs, more elongate muzzle and does
and Palawan. not overlap in range; Banded Linsang, Prionodon
Status Vulnerable. Appears to be relatively rare in linsang, is smaller with shorter legs and an entirely
much of range and declining due to loss of forest as banded tail.
well as hunting and trapping for the pet trade. Ecology and habitat Nocturnal. Travels and feeds
mainly on the ground, but sleeps in holes either in
SMALL-TOOTHED PALM CIVET the ground or in trees. Diet includes earthworms,
Arctogalidia trivirgata PLATE 44 insects and other small animals, both invertebrate
Measurements HB 440–530, T 480–660 (about and vertebrate. Largely restricted to lowland forest,
110–120% of HB), HF 75–95. Wt 2–2.5kg but also found in some secondary forests.
Identification Fur colour varies from olive-brown Distribution SE Asia: peninsular Myamar, Thailand
to greyish, rarely reddish-brown. Underfur reddish- and Malaysia. Also Sumatra and Borneo.
brown. Face, ears, feet and much of tail blackish. Status Near Threatened. Populations declining due
Usually has three fine, dark stripes or series of to loss of lowland forest and some hunting.
close, dark spots extending along midline from
neck to base of tail. Usually has a narrow pale OWSTON’S CIVET
stripe from forehead to tip of nose. Females have Chrotogale owstoni PLATE 45
two pairs of mammae. Similar species Common Measurements HB 510–630, T 380–480
Palm Civet, Paradoxurus hermaphroditus, has a Identification Body colour varies from greyish-
shorter tail, distinct ‘mask’ on face without white white to buffy-brown, paler underneath; distinct
on forehead, additional rows of spots on flanks, broad black bands across back, and stripes on
smaller skull but with relatively large, broad teeth, neck and face; dark spots on sides of neck and
curved toothrow (Fig. 56a); Masked Palm Civet, legs. Pattern of spots and bands individually
Paguma larvata, is larger and heavier, with usually variable. Tail banded at base, dark on distal two-
broader white stripe on forehead, pale cheeks, no thirds. Underparts strongly washed with bright
black stripes on back. orange-red in adult males, yellowish in adult
Ecology and habitat Usually nocturnal and females. Skull has elongate rostrum with relatively
arboreal, rarely descending to the ground. Very narrow, widely spaced teeth. Similar species
agile. Diet includes fruits and small animals. Occurs Banded Civet, Hemigalus derbyanus, lacks spots
in tall and secondary forests. on sides of legs and neck, has larger teeth, does
Distribution SE Asia: Myanmar, Thailand, Laos, not overlap in range.
Vietnam, Cambodia and Peninsular Malaysia. Also Ecology and habitat Nocturnal, largely terrestrial.
NE India, S China (Yunnan), Sumatra, Java, Bali, Feeds on invertebrates such as earthworms. Found
Borneo and some smaller islands. largely in wet evergreen broadleaf forest often above
Status Least Concern. 1,000m, but sometimes in disturbed areas.
Distribution SE Asia: N and C Laos, and N Vietnam.
BANDED CIVET Also S China (Yunnan).
Hemigalus derbyanus PLATE 45 Status Endangered. Low population densities in
Measurements HB 450–560, T 250–360, limited range, threatened by illegal hunting and
HF 72–88. Wt 1–3kg trapping.
Identification Body colour usually pale buff to
golden-brown, paler underneath, with distinct dark OTTER CIVET
brown or black bars across back, dark longitudinal Cynogale bennettii PLATE 45
stripes on neck and face. Some individuals are Measurements HB 575–680, T 120–205,
light grey, while others can be very reddish. Tail HF 102–110. Wt 3.5–5.6kg
mostly dark brown, banded only at base. Feet Identification Entirely dark brown, with faint grey
have strongly curved claws that are retractile, like grizzling and pale underfur; lips prominent, broad

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 and white, with very long white whiskers; a faint
pale spot above each eye. Ears small. Feet with
partial webbing between digits. Tail short. Similar
species Otters have longer, tapering tail, pale chin
and throat, and no white on muzzle or above eyes;
Flat-headed Cat, Prionailurus planiceps, is paler with
white on chin and chest, and short, slender muzzle.
Ecology and habitat Little known. Terrestrial and
aquatic. Diet includes aquatic animals. Largely
restricted to lowland forest near rivers, but has been
recorded at altitudes of up to 1,000m in Borneo. Has 0 20
limited tolerance of disturbed forest.
Distribution SE Asia: S Thailand and Peninsular Fig. 57 Skull of Otter Civet, Cynogale bennetti.
Malaysia. Also Sumatra and Borneo.
Status Endangered. Very few recent records in declined dramatically due to loss of forest, trapping
mainland SE Asia. Population is likely to have and degradation of wetland areas.

Family HERPESTIDAE MONGOOSES

Mongooses can be distinguished in the field from civets by their sharply pointed muzzle, tapering tail
with hair longer near the base than at the tip, and rather slender legs with long claws. They are largely
active during the day, unlike civets. Distinctive posture while walking, with hindquarters arched above the
level of the forequarters and tail. The footprints have relatively narrow toe pads (Fig. 52e). Taxonomic
notes All Asian mongooses were formerly assigned to the genus Herpestes, but recent genetic research
has shown they are not closely related to African species in that genus and should be assigned instead
to the genus Urva.

SHORT-TAILED MONGOOSE JAVAN MONGOOSE

Urva brachyura PLATE 46 Urva javanica PLATE 46
Measurements HB 380–445, T 205–250 (less Measurements Males: HB 360–420, T 275–315
than 55% of HB), HF 75–90 (70–75% of HB), HF 60–70. Wt 0.88–1.8kg.
Identification Entirely blackish-brown with fine Skull: cbl 76–82. Females: HB 320–360, T 250–
sandy-brown or orange speckling (speckling visible 265 (70–75% of HB), HF 55–65. Wt 0.53–0.84kg.
only from close range); chin and throat pale brown. Skull: cbl 64–79
Head and tail somewhat paler than body. Tail Identification Individual hairs of fur finely
relatively short. Similar species Other mongooses banded dark brown and pale buff with strong
are paler in colour, with longer tail, and usually found reddish-orange tinge, especially around head. Tail
in more open habitats. moderately long. Similar species Small Indian
Ecology and habitat Mainly diurnal and terrestrial. Mongoose, U. auropunctata, averages smaller
Diet includes arthropods and small vertebrate and lacks orange tinge; Short-tailed Mongoose,
animals. Mainly restricted to lowland forest, but U. brachyura, is much darker, has shorter tail and is
sometimes enters secondary forest, plantations and largely restricted to forest; Crab-eating Mongoose,
gardens. U. urva, is substantially larger, pale grey with pale
Distribution SE Asia: Peninsular Malaysia. Also stripe on neck.
Sumatra, Borneo. Ecology and habitat Active both day and night.
Status Near Threatened. Declining due to loss of Primarily found in grasslands and scrubby areas;
lowland forest. avoids dense evergreen forest. Feeds on rodents
and many other small vertebrates including birds,
reptiles and frogs, as well as crabs and large
insects.

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 Distribution SE Asia: E Myanmar, Thailand, Laos, Mongoose, U. javanica, is smaller with yellow or
Vietnam, Cambodia and Peninsular Malaysia. Also orange tinge; Crab-eating Mongoose, U. urva, has
S China, Java. long white patch on each side of neck.
Status Least Concern. Ecology and habitat Active by day or night. In
India occurs largely in open areas, including scrub
SMALL INDIAN MONGOOSE and cultivated lands. In some areas enters villages,
Urva auropunctata NOT ILLUSTRATED where often seen during the day. Feeds on rodents
Measurements HB 250–370, T 190–290. and a wide variety of vertebrate and invertebrate
Skull: cbl 56–71 (males), 55–67 (females) prey, sometimes including domestic chickens.
Identification Individual hairs of fur finely banded Distribution SE Asia: Formerly occurred between
dark brown and pale buff, giving overall impression Penang and Malacca in Peninsular Malaysia, where
of grizzled yellowish to olive-brown, lacking any it was believed to have been introduced, but now
reddish tinge. Tail moderately long. Taxonomic thought to be extirpated. Natural range includes Iran,
notes U. auropunctata has sometimes been Arabia, Pakistan, India and Sri Lanka.
considered the same species as U. javanica, Status Least Concern elsewhere in range.
but recent morphological and genetic analyses
confirm they are distinct. Similar species Javan CRAB-EATING MONGOOSE
Mongoose, U. javanica, is more reddish-orange Urva urva PLATE 46
and distinctly larger within respective sexes; Short- Measurements HB 440–480, T 260–310
tailed Mongoose, U. brachyura, is much darker, has (c.60–65% of HB), HF 90–110. Wt 3–4kg
shorter tail and is largely restricted to forest; Crab- Identification Large mongoose with comparatively
eating Mongoose, U. urva, is substantially larger, short tail. Overall colour brown to grey, with individual
pale grey with pale stripe on neck. guard hairs banded black and white, giving grizzled
Ecology and habitat Active both day and night. appearance. Conspicuous white stripe on side
Found in grasslands, around fields and scrubby of neck, and white chin. Rest of underparts dark.
areas. Preys on a range of small animals, including Long guard hairs make animal appear very shaggy.
rats, mice, snakes, lizards, frogs, scorpions, Similar species Javan Mongoose, U. javanica, is
centipedes and large insects. smaller, lacks white neck stripe and has shorter coat
Distribution SE Asia: W Myanmar. Also Pakistan, hairs; Indian Grey Mongoose, U. edwardsii, lacks
N India, Nepal. Introduced to many islands in shoulder stripe and has longer tail.
Caribbean and elsewhere. Ecology and habitat Active both day and night.
Status Least Concern. Found in a variety of forest and scrub habitats, often
near streams. Feeds on aquatic animals such as
INDIAN GREY MONGOOSE crabs, frogs, fish and molluscs, but probably takes
Urva edwardsii PLATE 46 other prey as well. Can squirt a strong-smelling
Measurements Males: HB 400–480, T 370–420 fluid from anal glands near base of tail, presumably
(80–95% of HB). Wt 1.4–2.0kg. Females: HB 370– for defence.
410, T 330–390 (80–95% of HB). Wt 0.9–1.3kg Distribution SE Asia: Myanmar, Thailand, Laos,
Identification Individual hairs banded black and Vietnam, Cambodia and Peninsular Malaysia. Also
white, giving grizzled grey appearance overall; worn Nepal, Bangladesh, NE India and S and E China.
fur acquires a brownish tinge. No pale shoulder Status Least Concern, although likely to be declining
stripe. Tail long with pale tip. Similar species Javan due to loss of forest and hunting.

Family PRIONODONTIDAE LINSANGS

Linsangs were formerly considered part of the civet family, Viverridae, but recent genetic studies have shown
they are not closely related to civets and should be placed in their own family. They have a long, slender body
with a long tail, and are believed to be largely carnivorous.
The two species of linsangs are readily separated by the pattern of spots/bands on their backs. The
footprints of both species have five distinct toe impressions (Fig. 55c).

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 BANDED LINSANG SPOTTED LINSANG
Prionodon linsang PLATE 44 Prionodon pardicolor PLATE 44
Measurements HB 350–450, T 300–420, Measurements HB 350–370, T 313–340,
HF 50–70. Wt 600–800g HF 60–70. Wt 600g
Identification Whitish to golden or buff, with a Identification Whitish to light brown to orange-buff,
pattern of large, dark brown spots that merge into with many irregular black spots on back and sides
about five dark transverse bands on upperparts, of body. Spots tend to remain separate, not blending
and longitudinal bands on sides of neck. Tail has into blotches, except for dark lines on either side of
black and pale bands along full length, typically neck. Tail with black and buff bands, typically about
about seven dark bands. Small, slender and cat- eight or nine complete dark bands. Similar species
like. Claws retractile, like a cat’s. Similar species Banded Linsang, P. linsang, has larger splotches
Spotted Linsang, P. pardicolor, has smaller spots that tend to blend into dark bands on back, fewer
on body that do not blend into large splotches bands on tail; Owston’s Civet, Chrotogale owstoni,
or bars, tail with more, narrower bands; Banded has distinct dark bands on back, and largely dark
Civet, Hemigalus derbyanus, has broad bars tail; Small Indian Civet, Viverricula indica, has many
on upperparts, but no spots, and a mainly black more spots that form narrow rows with lines on
tail; Leopard Cat, Prionailurus bengalensis, has a throat, and different posture.
differently shaped head, lacks broad barring along Ecology and habitat Largely nocturnal and
length of tail. arboreal, but also hunts on the ground. Most
Ecology and habitat Nocturnal. Largely arboreal records are from mountain and hill forest to at least
but also active on the ground. Sleeps during the day 2,000m altitude, but also occurs in secondary
in a nest made in a hole, either in the ground or in forest. Believed to prey largely on small mammals
a tree. Diet includes small mammals, birds, reptiles and birds, but probably eats arthropods and other
and arthropods. Occurs in tall and secondary forests, animals as well.
plantations and gardens. Distribution SE Asia: N Myanmar, N Thailand, Laos,
Distribution SE Asia: peninsular Myanmar, Thailand Cambodia and N Vietnam. Also E India, Nepal and
and Malaysia. Also Sumatra and adjacent islands, SE China.
Java and Borneo. Status Least Concern, although not often
Status Least Concern, though infrequently encountered and likely to be declining due to
encountered and likely to be declining somewhat habitat loss.
due to loss of forest.

Family FELIDAE CATS

Members of the cat family range in size from very large species, such as the Tiger and the Leopard, which
prey upon large mammals such as deer or pigs, down to several smaller species, similar in size to a domestic
cat, which feed mainly on small animals such as rodents or fish. All species have basically similar structure
and body form. The skull has a relatively short rostrum, with teeth highly specialised for carnivory. The dental
formula is 3/3 1/1 2–3/2 1/1 (Fig. 49b). The canines are very long, and the posterior premolar is greatly
enlarged in the form of a carnassial for tearing meat. The anterior small premolar is sometimes missing, and
the upper molar is very small.
The footprints show only four toes on both the front and hind feet (Fig. 58); the front feet have a fifth
toe, but it is small and does not touch the ground. The claws are retractile, and do not usually appear in
footprints. The main variation among species in footprints is in size, although there is some variation in the
shape of the footpads (Fig. 58).
There is considerable uncertainty in the generic relationships of cats, owing to the limited number of
differentiating characters. Some authorities place most species in the genus Felis, while others split them
into several genera, the approach followed here.

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 TIGER hair, ear tufts, no coat pattern, prehensile tail and
Panthera tigris PLATE 47 very different shape.
Measurements HB 1,700–2,300, T 950–1,150. Ecology and habitat Found in a wide range of
Wt 180–245kg habitats, though mainly in wooded areas.
Identification Easily recognised by very large Distribution SE Asia: Myanmar, Thailand, Laos,
size and pattern of vertical black and pale orange Vietnam, Cambodia and Peninsular Malaysia. Also
stripes along the body. Pattern of striping varies throughout much of Africa, Asia as far north as
considerably, and can be used to recognise N China and Java.
individuals in photographs. Tail is proportionately Status Vulnerable. Populations have declined owing
shorter than that of other large cats. Footprints to loss of habitat, declines in prey base and illegal
can be distinguished by their large size and by the hunting.
absence of claw marks (front feet 90mm or longer
– Fig. 58g, h). MAINLAND CLOUDED LEOPARD
Ecology and habitat Tigers are found in a wide Neofelis nebulosa PLATE 47
range of forested and partially open habitats, provided Measurements HB 650–950, T 550–800.
suitable prey is available. Main prey items are deer Wt 15–23kg
and pigs, though many other animals may be taken, Identification Usually pale sandy-brown to orange,
ranging from fish to rodents and birds. Occasionally, with ‘cloud’ patterns on sides and neck consisting
tigers may hunt humans or take domestic animals, of darker brown splotches with incomplete dark
but most remaining tigers in the region are very shy borders. Rarely, base colour is very dark and
of humans, after years of persecution. animals appear almost black. Footprints shorter
Distribution SE Asia: Myanmar, Thailand, Laos, with narrower pads than those of Leopard (Fig.
Peninsular Malaysia, possibly still in Vietnam, Cambodia. 58f). Upper canines very large relative to skull,
Also India, Nepal, China, Siberia and Sumatra. with clear diastema between them and cheek
Status: Endangered. Until recently was found in all teeth. Taxonomic notes Populations in Sumatra
countries in the region, but has disappeared from and Borneo now considered a separate species,
much of range and may now be extirpated from N. diardi. Similar species Marbled Cat, Pardofelis
Vietnam and Cambodia. Major threats are illegal marmorata, is smaller, with more irregular pattern
hunting and trapping, but also suffering from loss of lines and dark areas that do not form discrete
of habitat and prey base. Presumed Extinct on Java clouds, more extensive small spots on legs; Leopard,
and Bali. Panthera pardus, has rosettes of spots, not clouds,
longer legs with more erect posture, thinner tail.
LEOPARD Ecology and habitat Mainly nocturnal and arboreal,
Panthera pardus PLATE 47 but sometimes active during the day. In logged
Measurements HB 1,050–1,300, T 800–1,000. forest, often travels on the ground. Diet includes
Wt 45–65kg pigs, deer, primates and smaller mammals. Occurs
Identification Large cat with two colour phases. mainly in tall forests, but also secondary forests and
Spotted form has pale yellowish or yellowish-brown other disturbed habitats.
base colour, with black spots clustered in rosettes Distribution SE Asia: Myanmar, Thailand, Laos,
on back and singly on sides. Dark form also has Vietnam, Cambodia and Peninsular Malaysia. Also
rosettes, but base colour is also black, so rosettes Nepal, NE India, S China, Taiwan.
can be seen only in good light. Dark phase is most Status Vulnerable. Threatened by excessive hunting
common in S Thailand and Peninsular Malaysia. and loss of habitat.
Footprints (Fig. 58e) are similar to Tiger, P. tigris, but
generally shorter (length less than 80mm). Similar MARBLED CAT
species Mainland Clouded Leopard, Neofelis Pardofelis marmorata PLATE 48
nebulosa, is smaller, with irregular pattern of ‘clouds’ Measurements HB 450–530, T 470–550,
on coat; Binturong, Arctictis binturong, is sometimes HF 115–120, E 35–40. Wt 2–4kg
mistaken for a black Leopard, but has long shaggy Identification Sides and back brownish with dark

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 a b c

F H
d
F H
F H
F H
e f

F H
F H

g
h

H
F 0 100

Fig. 58 Left footprints of front (F) and hind (H) feet of various cats: Felis temminckii (a), Prionailurus viverrinus (b),
P. planiceps (c), P. bengalensis (d), Panthera pardus (e), Neofelis nebulosa (f) and Panthera tigris, showing a
relatively large (g) and relatively small (h) individual.

splotches and black lines, reminiscent of Mainland Vietnam, Cambodia and Peninsular Malaysia. Also
Clouded Leopard, Neofelis nebulosa, but black- Nepal, NE India, S China (Yunnan), Sumatra and
edged blotches on sides of body less distinct, tending Borneo.
to blend together in a marbled pattern, rather than Status Near Threatened. Overall rare, and declining
forming discrete ‘clouds’; black spots on legs smaller due to habitat loss and hunting. However, recent
and more numerous. Fur soft and thick. Similar camera trap surveys show it is more common than
species Mainland Clouded Leopard, N. nebulosa, is previously suspected.
larger, with patterns forming discrete ‘clouds’.
Ecology and habitat Mainly nocturnal and arboreal, ASIAN GOLDEN CAT
though also forages on ground. Diet probably Catopuma temminckii PLATE 48
includes a broad range of small animals, including Measurements HB 760–840, T 430–500 (about
rodents. Occurs mainly in tall forests, including 60% of HB). Wt 12–15kg
regenerating forest. Identification A large cat with relatively plain coat,
Distribution SE Asia: Myanmar, Thailand, Laos, usually lacking distinct markings except for black and

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 white stripes on sides and front of head. Coat colour Status Endangered. Relatively rare, population
quite variable, ranging from golden-brown to light declining due to loss of lowland forest and illegal
tawny-brown to greyish. Some individuals are very hunting and trapping.
dark brown or reddish. Generally darker along spine
and paler underneath. Tail moderately long, darker LEOPARD CAT
above, pale below, often with dark tip. Has indistinct Prionailurus bengalensis PLATE 48
spots or blotches on belly, as well as sometimes Measurements HB 400–550, T 230–290 (more
on legs and shoulder. Race C. t. tristis, which may than 50% of HB), HF 110–125, E 40–45. Wt 3–5kg
occur in N Myanmar, has extensive dark spots, Identification Reddish-orange or yellowish-buff,
stripes and splotches on upperparts (somewhat with black spots over entire upperparts including
similar to Marbled Cat, Pardofelis marmorata), but tail; underparts contrasting white with black spots.
can be recognised by distinctive face pattern and Pattern and size of spots varies considerably among
white underside to tail. Rarely, some individuals are individuals, ranging from large, round black spots to
all dark brown. Similar species Jungle Cat, Felis numerous small spots to large irregular splotches
chaus, has taller, pointed ears, bands on tail, lacks with browner centres. Spots may be rounded or
white stripes on face; Flat-headed Cat, Prionailurus elongate in shape; often form into lines on back; top
planiceps, is smaller with a very short tail. of head and back of neck with several distinct black
Ecology and habitat Reported from both dry stripes. Similar species Fishing Cat, P. viverrinus, is
forests and tropical rainforests. Occurs in disturbed larger and greyer, with shorter tail, buffy underparts,
areas but only near forest. Mainly terrestrial but can black spots usually fairly small, forming distinct
climb trees. Diet includes various medium and large rows; Marbled Cat, Pardofelis marmorata, has more
animals including hares, small deer and large birds. complex pattern of lines, dark and pale areas, not
Distribution SE Asia: Myanmar, Thailand, Laos, just spots; Banded Linsang, Prionodon linsang, is
Vietnam, Cambodia and Peninsular Malaysia. Also smaller, with different shape, relatively thick tail with
Nepal, NE India, S China and Sumatra. bands, not spots, along entire length.
Status Near Threatened. Still occurs in much of Ecology and habitat Most widespread and common
region, but declining due to loss of habitat and illegal small cat in the region, occupying a wide range of
hunting. habitats, including forests, plantations and gardens.
Usually nocturnal, mainly terrestrial but can climb
FLAT-HEADED CAT trees; swims well. Diet includes small mammals,
Prionailurus planiceps PLATE 48 lizards, amphibians and large insects.
Measurements HB 445–505, T 130–170 (27– Distribution SE Asia: Myanmar, Thailand, Laos,
34% of HB), HF 95–107, E 36–39. Wt 1.5–2.2kg Vietnam, Cambodia and Peninsular Malaysia. Also
Identification Appears brownish at a distance, Pakistan, India, Nepal, China, Siberia, Taiwan,
but hair of upperparts brown with fine grey Sumatra, Java, Bali, Borneo and the Philippines.
and pale buff speckling. Chin and chest white. Status Least Concern.
Indistinct dark and pale stripes on sides of face
and forehead. Ears small, top of head long and FISHING CAT
flattened. Similar species Asian Golden Cat, Prionailurus viverrinus PLATE 48
Catopuma temminckii, is substantially larger with Measurements HB 720–780, T 250–290 (c.40%
longer tail; Otter Civet, Cynogale bennetti, is darker of HB), E 44–50, HF 150–170. Wt 7–11kg
with very broad muzzle. Identification Medium-large, greyish or olive-brown
Ecology and habitat Nocturnal and terrestrial. in base colour with black stripes on top of head and
Probably feeds mostly on fish. Found mainly in tall rows of small black spots on back and sides. Spots
lowland forests near streams or wetlands. May be vary from rounded to elongate. Underside paler,
tolerant of some forest disturbance, but very rarely buffy to greyish, but not white. Tail relatively short.
detected on camera traps. Similar species Leopard Cat, P. bengalensis, is
Distribution SE Asia: peninsular Thailand and substantially smaller, more yellowish or orange
Malaysia. Also Sumatra and Borneo. with contrasting white underside, black spots often

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 larger, tail longer (more than 50% of head and body). climbing trees. Mostly nocturnal, though sometimes
Ecology and habitat Mainly found in drier, scrubby active during the day. Feeds on rodents, lizards, birds
habitat, usually near water, including swamps and and other animals, as well as carrion.
marshes. Feeds on fish, crabs, rodents, birds, Distribution SE Asia: Myanmar, Thailand, Laos,
molluscs and other animals. Vietnam and Cambodia. Also Middle East,
Distribution SE Asia: Myanmar, Thailand, Vietnam Afghanistan, Pakistan, India and S China (Yunnan).
and Cambodia; not confirmed in Laos. Also Pakistan, Status Least Concern globally, but populations in SE
Sri Lanka, S China (Yunnan), India, Nepal, Java and Asia have declined dramatically, with very few recent
Sumatra. records. Threats include loss of natural habitat and
Status Vulnerable. Rare and declining in many illegal trapping.
parts of the region due to loss of wetland habitats,
persecution and hunting. DOMESTIC CAT
Felis catus NOT ILLUSTRATED
JUNGLE CAT Measurements HB about 450–550, T variable,
Felis chaus PLATE 48 often short and bent
Measurements HB 500–650, T 260–310 (about Identification Coloration variable, often white with
50% of HB), HF 145–154, E 70–80. Wt 4–6kg irregular orange patches. Unlike wild cats, free-
Identification Fur light ashy-grey to yellowish- ranging Domestic Cats rarely flee from people and
brown, paler underneath; no stripes or spots on are often seen during the day. Similar species Wild
body. Tail with four or five dark rings on distal half. cats usually have a more symmetrical coat pattern
Ears tall and pointed, with black tuft on tip; legs and are very shy of people.
relatively long. Similar species Asian Golden Cat, Ecology and habitat Always associated with
Catopuma temminckii, has shorter, more rounded humans. Domestic Cats abandoned in disused
ears, stripes on forehead and cheeks, bicoloured tail temporary settlements fare much better than
without rings. Domestic Dogs, Canis familiaris, in the same
Ecology and habitat Found in relatively open, drier situation, and tend to stay near the old buildings,
habitats with suitable natural cover, such as tall grass, feeding on a variety of small animals.
open deciduous forest or scrub, usually near wetlands, Distribution Found worldwide, near most human
including streams or rivers. Mainly terrestrial, rarely settlements.

Family OTARIIDAE FUR SEALS AND SEALIONS

Most species of fur seals and sealions live in cold temperate or arctic waters in the northern or southern
oceans. One species has been recorded in South-east Asian waters, with recent confirmed records off the
coast of Vietnam. Members of this family have enlarged front flippers and flexible hind flippers, allowing
them to walk on land. They generally have an enlarged forehead with a distinct snout and small but distinct
external ears. The dental formula of Callorhinus is 3/2 1/1 4/4 2/1. The canines are enlarged, while the
molars and premolars are cone-shaped with single roots, and well-developed cusps.

NORTHERN FUR SEAL Ecology and habitat Breeds on islands mainly

Callorhinus ursinus PLATE 49 around Bering Sea, off Japan and California.
Measurements Male: HB up to 2.1m. Wt up to Remains at breeding colonies for only 1–2 months,
270kg. Female: HB up to 1.5m. Wt up to 60kg spending rest of year at sea.
Identification Males much larger than females. Fur Distribution SE Asia: Vietnam. Main distribution is
largely dark brown or blackish, paler in females. North Pacific, Bering Sea. Only occasionally found in
Relatively small head; short snout; blunt nose. Very warmer waters off South-east Asia.
large flippers and hind feet. Similar species Spotted Status Vulnerable; historically hunted heavily for
Seal, Phoca largha, has different head shape, much fur. Although commercial hunting has stopped,
smaller flippers. No other fur seals or sealions are populations continue to decline.
likely to be found in South-east Asian waters.

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 Family PHOCIDAE SEALS
The true seals are sometimes called earless seals because they do not have visible external ear flaps,
although they do have hidden ears and good hearing. Most species live predominantly in cold temperate or
arctic waters in the northern or southern oceans, although one species occurs in the Mediterranean (Monk
Seal, Monachus monachus) and two species occur in large inland water bodies (the Caspian Seal, Pusa
caspica, and the Baikal Seal, Pusa sibirica). Only one species has been recorded in South-east Asian waters,
with recent confirmed records off the coast of Vietnam. Members of this family have relatively small front
flippers. The hind flippers extend backwards and are used for swimming, but cannot be turned forwards for
walking. The dental formula varies among species. In the only South-east Asian species it is 3/2 1/1 4/4 1/1.

SPOTTED SEAL moult into adult colour after a few weeks. Similar
Phoca largha PLATE 49 species Northern Fur Seal, Callorhinus ursinus, has
Measurements HB up to 1.7m. Wt up to 130kg. different-shaped head, much larger flippers.
Identification Fur colour light grey to silver when Ecology and habitat Breeds on sea ice. Spends
fresh, often darkening to brown several months most of the year at sea.
after moult; darker above than below. Covered Distribution SE Asia: Vietnam. Occurs from North
with numerous conspicuous black spots above and Pacific to Arctic Ocean, off coast of Japan to Siberia
below. Front flippers relatively small. Males only and Alaska. Only occasionally comes to SE Asia.
slightly larger than females. Pups pure white, but Status Least Concern.

Order CETACEA
Whales, dolphins and porpoises
The cetaceans include the largest animals that have ever lived in the world – the largest Blue Whales are
larger than the largest-known dinosaurs. Because they live completely in the water they are often thought
to be fish, but they are true warm-blooded mammals that have lost their legs and most of their hair, and
acquired fins and other modifications for living in the sea. They still breathe air, and they give birth to live
young and feed them on milk.
If not seen well, some dolphins and small whales could be mistaken for sharks, as the dorsal fin
sometimes appears similar. However, among other differences, dolphins have a blowhole on top of the head,
visible when the animal breathes, and horizontal tail flukes instead of a vertical tail. They also swim quite
differently, rising up and down, and sometimes even jumping out of the water.
Whales and dolphins are often difficult to identify, as they are usually hard to see well in the water. Even
experienced observers familiar with the behaviour of the animals cannot identify every cetacean seen, and it
can be especially difficult for a beginner. However, some species are quite tame, allowing good views, while
others show themselves several times, eventually allowing enough to be seen for identification.
Until recently, relatively little was known about cetaceans in South-east Asia. Most confirmed records
related to specimens that became stranded, or that had died and were then washed up on a beach. Since
the 1990s there has been substantially increased boat-based survey work in the region, prompted by
concern about declining populations. This has provided increased information on the status of many of
the smaller porpoises and dolphins that are regularly found in the offshore waters, including the Irrawaddy
Dolphin that regular occurs in some of the largest freshwater rivers. Several other species, including
species of large whales, are likely to pass through South-east Asian waters at least occasionally. Because
so little is known about them, all of the species known to occur in the tropical Pacific or Indian Oceans are
included here, even if they have not definitely been recorded from the region, in the expectation that they
are likely to show up.
Unfortunately, many cetaceans are now of significant conservation concern. Most species breed slowly,
not starting to reproduce until they are many years old, and then producing only one young every few

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 years. As a result, most populations cannot sustain hunting of more than 1 or 2% of the total population
per year without starting to decline. Historically, large whales were hunted very heavily, with the result that
nearly all species declined dramatically, some almost to extinction. Although commercial whaling has been
largely stopped, most populations have not yet recovered, and are still threatened by entanglement in nets,
collisions with ships and pollution. Smaller cetaceans such as dolphins were rarely hunted commercially on
a large scale, but in many areas, including South-east Asia, they are threatened by accidental entanglement
in fishing nets, which leads to drowning, injury from illegal dynamite fishing, collisions with boats and some
targeted hunting by local fishermen despite legal protection in most areas. More than half the species that
occur in South-east Asia are thought to have experienced major population declines and to be locally at
risk. Cetacean numbers are exceptionally low off the coast of Vietnam, probably due to excessive mortality
from fishing and other operations.

Family DELPHINIDAE OCEANIC DOLPHINS

This family includes all of the well-known dolphins, as well as the larger Orca or Killer Whale. All species have
rows of peg-like teeth in both jaws, well-developed dorsal fins and often a distinct beak.

MELON-HEADED WHALE Dorsal fin tall and curved back in centre of back;
Peponocephala electra PLATE 50 flippers rounded at the tip. Upper jaw has 8–11
Measurements TL 2.2–2.7m teeth on each side, lower jaw has 11–13. Similar
Identification No prominent beak; forehead species Melon-headed Whale, Peponocephala
smoothly curved downwards in profile; head conical electra, can be hard to separate but head shape
or triangular from above. Dorsal fin tall and curved. is more triangular (from above), flippers are more
Flippers long, slim and pointed. Colour generally pointed; False Killer Whale, Pseudorca crassidens,
black, sometimes paler on the belly. Teeth 20–25 is longer and more slender, more uniformly dark,
in each row. Similar species False Killer Whale, with hump on flippers and proportionately smaller
Pseudorca crassidens, is substantially larger, has dorsal fin.
a slightly more rounded profile and bent flippers; Ecology and habitat Often forms large groups, but
Pygmy Killer Whale, Feresa attenuata, has rounded less active than many dolphins. May sometimes
flippers and a dark cape contrasting with paler sides, attack other cetaceans.
a pattern rarely present in Melon-headed Whale. Distribution Worldwide, in tropical and sub-tropical
Ecology and habitat Usually occurs in large herds seas. SE Asia: occurs off east coast of India,
of up to several hundred animals, which can travel Indonesia and in South China Sea; records from
very quickly. Primarily in deeper, offshore waters. Thailand and Vietnam.
Feeds on squid and small fish. Status Data Deficient. Occurs in low numbers and
Distribution Found in tropical and sub-tropical poorly known; possibly declining substantially due to
oceans, mainly in deep water. SE Asia: frequently fisheries bycatch, plastic pollution and other human-
reported around the Philippines, with records in related threats.
Peninsular Malaysia, the Gulf of Thailand and
Vietnam; also occurs near India and thus likely to FALSE KILLER WHALE
occur throughout Bay of Bengal. Pseudorca crassidens PLATE 50
Status Least Concern. Measurements TL 3.2–4.0m; males up to 6.1m;
females up to 4.9m
PYGMY KILLER WHALE Identification Mainly black, sometimes slightly
Feresa attenuata PLATE 50 paler around the head and on the chest. Long,
Measurements TL 2.2–2.6m slender body with relatively small, rounded head.
Identification General colour dark grey-brown to Conspicuous dorsal fin, curved backwards; flippers
black with paler sides, giving an indistinct caped with hump on leading edge, unlike other black
look; there are irregular patches of white on the whales. Large conspicuous teeth, 8–11 in each row.
belly, chin and lips. Head rounded with no beak. Similar species Killer Whale, Orcinus orca, is much

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 larger, with a tall, triangular fin, broad, rounded head with regional records from Thailand and Vietnam.
and large white patches; pilot whales, Globicephala Probably only migrates through local waters.
spp., have a more rounded, bulbous head, broader Status Data Deficient; not assessed owing to
lower dorsal fin and long, pointed flippers; Pygmy uncertainty about number of taxonomic units
Killer and Melon-headed Whales, Feresa attenuata involved.
and Peponocephala electra, are smaller and have
paler markings, often with white on the lips. SHORT-FINNED PILOT WHALE
Ecology and habitat Gregarious, usually in groups, Globicephala macrorhynchus PLATE 50
often forming large herds; very active, frequently Measurements TL males 4.2–5.4m; females
playing near boats. Eats mainly squid and larger fish, 3–4m
but has been reported attacking other cetaceans. Identification Generally black with greyish-white
Distribution Throughout tropical seas and into marks under the throat and usually large pale patch
warmer temperate areas. SE Asia: recorded from behind dorsal fin. Dorsal fin long and low. Thick,
coastal waters off Pakistan and India, through bulbous forehead with no visible beak in adults.
Sumatra, Borneo and E Indonesia. In region, Flippers sickle-shaped. Has 7–9 teeth in each row.
recorded from Singapore, Cambodia, Thailand and Similar species False Killer Whale, Pseudorca
Vietnam. crassidens, has a more tapered head and a more
Status Data Deficient; relatively rare, and possibility slender, erect dorsal fin; Killer Whale, Orcinus orca,
of substantial declines due to fisheries bycatch and is larger, with tall, erect dorsal fin; Long-finned Pilot
other threats. Whale, Globicephala melas, is extremely similar but
not known to occur in the tropics: it is slightly larger,
KILLER WHALE has 8–12 teeth in each row and longer flippers, but
Orcinus orca PLATE 50 can rarely be distinguished at sea.
Measurements TL 6–9.5m; adult males up to Ecology and habitat Found in small groups, often
9.5m; females up to 7m, but usually smaller in the company of dolphins. Feeds mainly in deep
Identification Tall, triangular dorsal fin, lower and waters, but often comes close to the shore. Believed
more curved in females. Black with large, well- to feed mainly on squid.
marked white areas behind the eye, on the throat Distribution Worldwide in tropical and sub-tropical
and belly, and behind the dorsal fin – pattern varies waters. SE Asia: has been recorded off coasts of
with individuals. Eye patch often visible when Borneo and Java, and in Bay of Bengal; regional
swimming. Flippers broad and rounded. Adult records from Peninsular Malaysia, Cambodia and
males easily recognised; females smaller, with a Thailand.
more dolphin-like dorsal fin, but often in company Status Data Deficient; not assessed owing to
of adult male. Similar species False Killer Whale, uncertainty about number of taxa involved.
Pseudorca crassidens, is more slender, with no
white markings and more rounded head; Risso’s RISSO’S DOLPHIN
Dolphin, Grampus griseus, is much smaller, usually Grampus griseus PLATE 50
paler and greyer, with rounded, beakless head; pilot Measurements TL 3.6–4.0m
whales, Globicephala spp., have bulbous forehead Identification High, domed forehead, bisected
and low, broad-based dorsal fin. in the middle by a deep, longitudinal crease; no
Ecology and habitat Diet varies among populations, conspicuous beak. Body generally dark grey when
some eating mainly fish and others specialising young, turning pale grey to whitish with age,
in marine mammals such as seals and other especially around the head; extensively covered
cetaceans. Very social, with group size varying from with long scars; tall, erect dorsal fin, slightly curved
a few individuals to 30 or more; larger groups may back. No upper teeth and usually fewer than seven
represent several pods coming together. pairs of peg-like lower teeth. Similar species
Distribution Worldwide, but less often seen in False Killer Whale, Pseudorca crassidens, is darker
tropical waters. SE Asia: recorded from Bay of Bengal and generally unscarred, forehead not grooved;
and around Indonesian islands, including Borneo, Common Bottlenose and Indo-Pacific Humpbacked

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 Dolphins, Tursiops truncatus and Sousa chinensis, a white belly. Fin, flippers and flukes dark. Similar
have a long beak; beaked whales, Mesoplodon spp., species Striped Dolphin, Stenella coeruleoalba,
have a smaller dorsal fin and tapered head, and has a narrower dark side stripe, distinct pattern of
grow to be much larger. white stripes on sides, longer beak and larger fin;
Ecology and habitat Feeds mainly on squid, but Common Dolphin, Delphinus delphis, has larger
also some fish; usually in groups, but sometimes fin, V-shaped dark dorsal saddle and distinct pale
singly or in pairs. Prefers deep water. patches on sides.
Distribution Worldwide, in tropical to sub-tropical Ecology and habitat Eats fish, shrimp and squid,
seas, primarily in deep waters. SE Asia: recorded often at depths of 250–500m. Usually in large
from South China Sea, with records from Sarawak groups, sometimes mixed with other species. Mainly
and off coast of Vietnam; also known from Sri Lanka offshore in deep waters.
and likely to occur in Bay of Bengal. Distribution Tropical seas in most parts of the
Status Least Concern, but some mortality from world. SE Asia: occurs off Sri Lanka, Borneo (type
fishing. specimen was found stranded in Sarawak), Taiwan
and the Philippines, with regional records in Thailand
LONG-BEAKED COMMON DOLPHIN and Vietnam.
Delphinus capensis PLATE 51 Status Least Concern, but populations may be
Measurements TL 1.8–2.5m, but usually less than affected by excessive hunting in some areas.
2.3m
Identification Back brownish-black to black, belly PANTROPICAL SPOTTED DOLPHIN
white; dark V-shaped saddle below dorsal fin, with Stenella attenuata PLATE 51
large buff or white patch in front, and large grey Measurements TL 2–2.5m
patch behind, which appear to form criss-crossing Identification Dark grey to brownish-grey cape that
stripes. Dorsal fin tall and triangular, all dark, or is very broad below dorsal fin, narrow on forehead;
with pale patch in middle. Beak long and narrow. underside whitish except dark flippers and a thin
Taxonomic notes Form found around Asia may stripe from beak to flippers; pale grey stripe from
represent a distinct species, D. tropicalis. Similar side to lower back; can be heavily spotted with pale
species Striped Dolphin, Stenella coeruleoalba, spots above, dark spots below, but some animals
has thin dark stripes from the eye to the anus and are relatively unspotted. Beak long and narrow, often
from the eye to the flippers, and has a different side with white tip and lips. Similar species Bottlenose
pattern. dolphins, Tursiops spp., similar to unspotted
Ecology and habitat Feeds mainly on small individuals, but more robust with thicker, shorter
schooling fish and squid. Very active during the beak, plainer coloration; Indo-Pacific Humpbacked
day, often jumping and ‘playing’ in the water. Most Dolphin, Sousa chinensis, sometimes heavily spotted,
frequently seen near shore. but usually has low dorsal fin with thickened base
Distribution Tropical and sub-tropical seas. SE and more uniform mantle pattern; Spinner, Striped
Asia: reported from Bay of Bengal, Borneo, Straits and Common Dolphins differ in shape of mantle.
of Malacca off Malaysia, Gulf of Thailand, Cambodia Ecology and habitat Feeds on fish and squid; often
and Vietnam. congregates in large herds, sometimes mixed with
Status Data Deficient; uncertainty whether other species; found in both coastal and offshore
population is declining fast enough to be at risk. waters. Very active, often leaping high or ‘playing’ in
the water.
FRASER’S DOLPHIN Distribution Throughout tropical waters and
Lagenodelphis hosei PLATE 51 sometimes in warm temperate seas. SE Asia: likely
Measurements TL 2.3–2.7m in waters throughout region including Borneo, the
Identification Short beak; relatively small, pointed Philippines and Taiwan, with confirmed records from
flippers and triangular dorsal fin. Generally dark Thailand, Cambodia and Vietnam.
blue-grey above, with paler sides, distinct black Status Least Concern. Regionally affected by
stripe on each side from the eye to the anus, and mortality in fishing nets.

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 STRIPED DOLPHIN Ecology and habitat Forms very large, active
Stenella coeruleoalba PLATE 51 groups, often jumping and twisting (from which
Measurements TL 1.8–2.7m they get their name); sometimes mixed with other
Identification Long, narrow black stripes from species. Dwarf form occurs in relatively shallow
eye to flipper and eye to anus; broad, pale grey waters and feeds mainly on bottom-dwelling and
streak above eye to below dorsal fin. Back dark coral reef organisms; larger form in deep offshore
grey to bluish-grey; sides light grey; belly white. Fin waters feeds mainly on fish and squid.
moderate and slightly curved; beak long and distinct. Distribution Tropical and sub-tropical regions
Similar species Spinner Dolphin, S. longirostris, of the world’s oceans. SE Asia: dwarf form,
has a more even grey stripe on the side without S. l. roseiventris, which may prove to be a distinct
black stripes, and less curved back fin; Common species, recorded from Myanmar, Peninsular
Dolphin, Delphinus delphis, has a different pattern of Malaysia, Gulf of Thailand, Cambodia, Vietnam and
stripes; bottlenose and Pantropical Spotted dolphins, probably in coastal waters throughout the region;
Tursiops spp. and Stenella attenuata, sometimes typical larger form, S. l. longirostris, regularly seen
have an indistinct pale blaze on the shoulder, but around the Philippines and probably occurs in
lack the black stripes on the sides; Fraser’s Dolphin, deeper waters throughout South China Sea.
Lagenodelphis hosei, has broader black side stripes, Status Data Deficient; uncertain extent to which
shorter snout and small dorsal fin. global populations are declining; populations in SE
Ecology and habitat Eats mainly small fish, shrimp Asia threatened by excessive mortality in fishing
and squid. Usually in large groups well offshore, but operations.
sometimes enters near-shore waters. Very active,
often jumping clear of the water. ROUGH-TOOTHED DOLPHIN
Distribution Worldwide in warm-temperate to Steno bredanensis PLATE 52
tropical seas. SE Asia: occurs around Indonesia and Measurements TL 1.8–2.8m
the Philippines, with confirmed records off Thailand Identification Forehead and sides of head slope
and Vietnam; potentially occurs anywhere in region. evenly into long, slender beak. Prominent, tall
Status Least Concern, but SE Asian population now dorsal fin and large flippers. Generally dark grey to
very small, affected not only by fishing mortality in blackish, with pale lips and underparts; pale streaks
the region, but also by over-hunting in Japan, where along sides set off dark back in an indistinct narrow
they are believed to migrate seasonally. saddle; often has pale splotches on sides. Similar
species Other beaked dolphins have a distinct
SPINNER DOLPHIN crease where the bulbous forehead meets the beak;
Stenella longirostris PLATE 51 bottlenose dolphins, Tursiops spp., have shorter
Measurements TL 1.3–2.3m beaks and more uniform grey coloration; Indo-
Identification Long, narrow beak with relatively low Pacific Hump-backed Dolphin, Sousa chinensis, can
forehead; dorsal fin nearly triangular and sometimes sometimes be similar in colour, and may lack hump,
even curved forwards in adult males; tail base often but has smaller fin and differently shaped head;
with a strong keel or ridge. Upperparts dark grey, Pantropical Spotted Dolphin, Stenella attenuata, and
paler grey stripe along sides forming a relatively Spinner Dolphin, Stenella longirostris, have more
straight border from dark mantle; underparts white distinct colour patterns.
with a dark line from eye to flipper. Dwarf form, S. l. Ecology and habitat Eats mainly fish and squid.
roseiventris, which is most frequently seen in region, Typically in groups of 10 to 20 individuals, mainly in
is substantially smaller than the typical deepwater deeper offshore waters.
forms. Similar species Bottlenose dolphins, Distribution Widely distributed in tropical and
Tursiops spp., are larger, with a more curved fin and warm-temperate seas. SE Asia: recorded from Bay
a shorter beak with a pale tip and lips; Common of Bengal through Java and Borneo, with reports
Dolphin, Delphinus delphis, has a V-shaped dark from Thailand and Vietnam.
saddle on back; Striped Dolphin, S. coeruleoalba, Status Least Concern.
has a distinct black stripe on sides.

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 INDO-PACIFIC HUMPBACKED DOLPHIN Risso’s Dolphin, Grampus griseus, has no beak;
Sousa chinensis PLATE 52 adults are paler with white around the head and no
Measurements TL 2.4–2.8m distinct cape; Pantropical Spotted Dolphin, Stenella
Identification Adults usually have a long, low hump attenuata, has a more complex pattern, usually with
on the back with a small triangular fin on top, but spots, and a longer, slimmer beak.
some forms lack the hump and have a larger fin. Ecology and habitat Found mainly in shallow
Quite variable in coloration, from uniform dull coastal waters, usually in small groups.
grey to pure white or pink, with some speckled Distribution Coastal waters around Indian Ocean
individuals. Colour variation is partly related to age, and South Pacific to N Australia. SE Asia: probably
with younger animals dark grey, becoming mottled occurs in coastal waters throughout region,
grey then whitish or pink, but there may be genetic with records off Myanmar, Peninsular Malaysia,
variation in colour as well. Taxonomic notes Recent Cambodia, Thailand and Vietnam.
genetic studies show that the populations in India (S. Status Data Deficient; possibly substantial declines,
plumbea) and Australia (S. sahulensis) are different due to entanglement in fishing nets and deliberate
species. Similar species Bottlenose dolphins, hunting.
Tursiops spp., can be very similar to forms without a
hump, but tend to be more uniform grey with shorter COMMON BOTTLENOSE DOLPHIN
beak and taller dorsal fin, and usually form larger, Tursiops truncatus NOT ILLUSTRATED
more active groups. Measurements TL 2.4–3.8m; males average
Ecology and habitat Eats mainly fish. Usually in slightly larger than females
sheltered coastal waters; sometimes in mangrove Identification Very similar to Indo-Pacific Bottlenose
swamps and estuaries. Dolphin, T. aduncus, and difficult to distinguish in
Distribution Shallow coastal waters from the field, but tends to be rather larger, with more
Bangladesh around SE Asia to the Philippines and robust body, stouter beak, more convex melon and
China; related species occur in India and Australia. proportionately smaller dorsal fin, and is generally
SE Asia: coastal waters throughout region, including slightly darker and lacking speckling.
Myanmar, Peninsular Malaysia, Thailand, Cambodia Ecology and habitat Eats a wide variety of food,
and Vietnam. including fish and invertebrates. Found in shallow to
Status Vulnerable; population declining due mainly oceanic waters, usually in small groups. Frequently
to entanglement in fishing gear. Additional threats quite tame, approaching ships or swimmers.
include chemical contamination from pollution and Distribution Worldwide except in very cold waters.
boat collisions. SE Asia: poorly known, as most records were not
distinguished from T. aduncus. Most frequently seen
INDO-PACIFIC BOTTLENOSE DOLPHIN near shore, but probably regular in deeper waters,
Tursiops aduncus PLATE 52 with confirmed records off Borneo and Vietnam.
Measurements TL 2.0–2.6m; males slightly larger Status Least Concern; potentially significant
than females mortality from fishing activities in region.
Identification Fairly uniform grey body, slightly
darker on back and paler on sides, forming an IRRAWADDY DOLPHIN
indistinct cape; whitish on the belly, often with Orcaella brevirostris PLATE 52
dark spots. Beak moderately short and thick; large Measurements TL 2.0–2.5m
flippers; dorsal fin tall and curved back. Similar Identification Varies from mid-grey to dark slate-
species Common Bottlenose Dolphin, T. truncatus, blue, paler underneath, with no distinctive pattern.
is very similar but averages slightly larger; Indo- Small dorsal fin behind middle of back, rounded
Pacific Humpbacked Dolphin, Sousa chinensis, if or slightly falcate. Forehead high and rounded; no
it lacks hump, is usually whiter and has smaller, beak. Broad, rounded flippers. Taxonomic notes
more triangular dorsal fin; Rough-toothed Dolphin, Populations around Australia are morphologically
Steno bredanensis, has a more conical head with no distinct and they may represent a different species.
crease separating the beak, and a very narrow cape; Similar species Indo-Pacific Finless Porpoise,

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 Neophocaena phocaenoides, is smaller and has no and S Vietnam. Seen most often in estuaries and
back fin; Indo-Pacific Humpbacked Dolphin, Sousa bays; also in inland rivers, including Irrawaddy, up
chinensis, is larger, has a longer beak and larger to 1,400km inland in Myanmar, and Mekong inland
dorsal fin, and is generally more active. to S Laos.
Ecology and habitat Prefers inshore waters and Status Endangered; large declines due to
estuaries, including very muddy and murky waters; accidental mortality during fishing, degradation of
also occurs quite far inland on large rivers. Quiet and water in rivers and deliberate hunting. Several local
inconspicuous, rarely leaping or jumping. populations, including those on large rivers, have
Distribution India through Indonesia to Australia. SE experienced dramatic declines and are considered
Asia: coastal waters from Myanmar, around Malay Critically Endangered, at risk of imminent extinction.
Peninsula, through Gulf of Thailand to Cambodia

Family PHOCOENIDAE PORPOISES

Similar to dolphins, but with no beak. The only species found in the region has no dorsal fin.

INDO-PACIFIC FINLESS PORPOISE Ecology and habitat Eats small squid, prawns and
Neophocaena phocaenoides PLATE 52 small fish. Found in shallow coastal waters and river
Measurements TL 1.5–2.0m estuaries, in small groups or singles.
Identification Uniform grey, sometimes very pale, Distribution Coastal waters from India through to
but usually slate. Rounded head; no dorsal fin. China. SE Asia: confirmed records include coastal
Taxonomic notes Northern Asian population in waters and estuaries off Thailand, Peninsular
China and Japan is now considered a separate Malaysia, Cambodia and Vietnam; may also occur
species, N. asiaeorientalis. Similar species off Myanmar.
Irrawaddy Dolphin, Orcaella brevirostris, occurs Status Vulnerable; many local populations have
in similar areas and also has a rounded head, but experienced dramatic declines, due largely to
is larger, with a small but distinct dorsal fin; other deliberate and accidental catch by fisheries, and
dolphins are larger with large dorsal fins. species is now relatively rare.

Family PHYSETERIDAE SPERM WHALES

This family includes one of the largest whales as well as some of the smallest, but they appear to be related.
They all have a huge, bulbous forehead containing a large cavity full of oil and spermaceti, a wax-like
substance highly valued by whalers. The lower jaw is relatively small and underslung, with a row of well-
developed teeth that fit into sockets in the upper jaw.

GREAT SPERM WHALE colour; beaked whales, Mesoplodon spp., have a high
Physeter macrocephalus PLATE 54 forehead, but long beak and prominent dorsal fin;
Measurements TL males 10–12m, maximum blowhole well back on head.
18m; females 8–9m, maximum 12m Ecology and habitat Feeds primarily on large
Identification Huge head, with large square squid. Usually in deep water; sometimes dives to
forehead. Overall colour dark brownish-grey to over 1,000m. Remains near surface for 10 minutes
brown; ‘wrinkled’ skin on back. Distinct dorsal hump or more to breathe after a deep dive.
with a series of low bumps leading to the tail – clearly Distribution Worldwide. SE Asia: occurs in both
visible when tail is thrown high before a deep dive. Indian Ocean and South China Sea, but only a few
Tail flukes straight across the end, all dark below. confirmed records from SE Asian waters.
Teeth on lower jaw only. Distinctive blow, angled Status Vulnerable; populations declined dramatically
forwards and to the left. Taxonomic notes Formerly due to historic over-hunting; unclear whether
called P. catadon. Similar species Humpback Whale, population has started to recover after protection
Megaptera novaeangliae, could appear similar when from hunting; currently still some mortality from
diving at a distance, but tail flukes differ in shape and entanglement in fishing nets and collisions with boats.

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 PYGMY SPERM WHALE DWARF SPERM WHALE
Kogia breviceps PLATE 53 Kogia sima PLATE 53
Measurements TL 2.7–3.4m (males and females Measurements TL 2.1–2.7m
similar) Identification Coloration and shape very similar to
Identification Head appears shark-like, with that of Pygmy Sperm Whale, K. breviceps – dark
underslung lower jaw. General colour dark bluish- grey above, whitish below – but slightly smaller,
grey fading to whitish on belly, with a gill-shaped dorsal fin larger and further forwards (as in Common
light and dark mark at the side of the head. Dorsal Bottlenose Dolphin, Tursiops truncatus), sometimes
fin low, behind centre of back. Teeth thin, curved a few creases or grooves on throat, and fewer,
inwards and sharp; 12 to 16 pairs in lower jaw, smaller teeth (7 to 12 pairs on lower jaw – Pygmy
none in upper; up to 30mm long, 4.5mm in usually has 12 to 16). Similar species Dolphins
diameter. Usually swims slowly and sluggishly with similar fins have beaked heads; Pygmy Killer
when on the surface, sometimes resting. Similar Whale, Feresa attenuata, and Melon-headed Whale,
species Dwarf Sperm Whale, K. sima, is smaller, Peponocephala electra, have different head shapes,
has a taller, dolphin-like fin and fewer, smaller teeth; but could appear similar at a distance; both are
some beaked whales, Mesoplodon spp., may appear much more active and usually in groups.
similar at a distance, but have a quite different head Ecology and habitat Feeds mainly on squid, but
shape. also fish and crustaceans. Can dive up to 300m, but
Ecology and habitat Feeds mainly on squid and often in inshore waters. Sometimes lies motionless
octopus, but also some fish and crabs. Usually alone on the surface of the water.
or in small groups. Often in shallow water, frequently Distribution Worldwide, in tropical to temperate
stranded. seas, but until recently confused with Pygmy Sperm
Distribution Most parts of the world, in temperate to Whale, K. breviceps, and many records have not
tropical waters. SE Asia: records from all around the been separated. SE Asia: confirmed records from
region (east of India, South China Sea and Sarawak) all around the region (E India, Lesser Sundas and
but no confirmed records within region. Taiwan), but none from within the region.
Status Data Deficient; relatively rare; possibly Status Data Deficient; relatively rare; uncertain
declining due to threats such as entanglement in whether declining significantly; threats include
fishing nets and ingestion of plastic. entanglement in fishing nets and ingestion of plastic.

Family ZIPHIIDAE BEAKED WHALES

A poorly known group of medium to small whales, with an indistinct beak. The lower jaw has a few large
teeth in males, only vestigial teeth in females. Bodies often covered with long, pale scars.

CUVIER’S BEAKED WHALE Ecology and habitat Feeds mainly on squid and
Ziphius cavirostris PLATE 53 deepwater fish. Can probably dive very deeply;
Measurements TL 5.5–7.5m; males typically about submerges for up to 30 minutes. Often raises flukes
0.5m shorter than females before diving. Apparently wary of boats and hard to
Identification Sloping forehead; poorly defined observe.
beak, lower jaw longer than upper. Colour extremely Distribution Worldwide except in Arctic waters,
variable, from dark rust-brown to slate-grey or though less common in the tropics. SE Asia:
fawn; belly usually paler; head sometimes whitish recorded from Indian Ocean, from Java to Maluku
in older adults; back and sides usually covered with in Indonesia, off Sabah and off coast of Thailand.
linear scars. Males have two conical teeth at front Status Least Concern; apparently one of the more
of jaw. Similar species Blainville’s Beaked Whale, common beaked whales; some threats related to
Mesoplodon densirostris, has a longer beak, teeth at entanglement in fishing nets, especially drift nets.
side of jaw; Longman’s Beaked Whale, Indopacetus
pacificus, is of a similar size but has a more bulbous
forehead and elongated beak like a dolphin.

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 BLAINVILLE’S BEAKED WHALE beak, a more sloped forehead in profile and an
Mesoplodon densirostris PLATE 53 unarched jaw; male Blainville’s Beaked Whale,
Measurements TL 4.4–4.7m M. densirostris, often has more scarring, and has
Identification Generally black or dark grey, paler a more strongly arched jaw with a conspicuous
below, often with extensive pale blotches and white single large tooth.
scars (presumably from fighting). Forehead low with Ecology and habitat Little known; presumed to
a distinct beak; lower jaw highly arched, especially in feed mainly on squid.
males, which have a single large tooth in the middle Distribution Indian Ocean and tropical to temperate
of the arch. Similar species Cuvier’s Beaked Whale, N Pacific. SE Asia: not yet recorded, but there are
Ziphius cavirostris, has a less prominent beak, less nearby confirmed records from Sri Lanka, E Sumatra
arched jaw and often a pale forehead. Male Ginkgo- and Taiwan.
toothed Beaked Whale, M. ginkgodens, has less Status Data Deficient; appears to be relatively
arched jaw with smaller, bilobed tooth. Females and common, but trends uncertain; potentially threatened
young probably indistinguishable in the field from by plastic pollution and loud underwater noise.
other Mesoplodon.
Ecology and habitat Seen most often in slope LONGMAN’S BEAKED WHALE
waters 500–1,000m deep. Feeds on small fish and Indopacetus pacificus NOT ILLUSTRATED
squid. Measurements TL 6–7m, possibly longer
Distribution Reported from tropical and sub- Identification Generally black or dark grey, with
tropical waters in most parts of the world. SE Asia: well-developed dorsal fin that curves back at tip.
recorded from Nicobar Islands in Indian Ocean and Has a distinct beak with a domed forehead (like
the Philippines, China and Taiwan, with one record a dolphin) and two small teeth protruding in front
from Thailand. of lower jaw. Has a small pale patch behind eye
Status Data Deficient; appears to be relatively and may have linear scars from fighting as well
common, but trends uncertain; potentially threatened as small round scars from cookie-cutter shark
by plastic pollution and loud underwater noise. bites. Juveniles have distinctive contrasting pale
underparts and side of face. Similar species Other
GINKGO-TOOTHED BEAKED WHALE beaked whales have less domed forehead; dolphins
Mesoplodon ginkgodens PLATE 53 are much smaller.
Measurements TL 4.7–5.2m Ecology and habitat Presumed to feed mainly on
Identification Generally black or dark grey, paler squid. Has been reported travelling in groups of
below. Tends not to have linear scars, unlike 20–100 individuals.
other Mesoplodon. Forehead low with a distinct Distribution Indian Ocean and tropical Pacific. SE
beak; lower jaw moderately arched, especially in Asia: likely occurs throughout region; a stranded
males, which have a bilobed tooth in the middle specimen has been reported from Myanmar.
of the arch. Similar species Cuvier’s Beaked Status Data Deficient; field identification characters
Whale, Ziphius cavirostris, has a less prominent only determined in early 2000s.

Family BALAENOPTERIDAE RORQUALS OR BALEEN WHALES

The rorquals, or baleen whales, include some of the largest whales, and can be recognised by their huge size
and small dorsal fins near the back of the body. They feed by filtering food and water through huge baleen
plates in their mouths. At a long distance they can often be distinguished from the Sperm Whale by the shape
of the spout when they breathe – tall and conical or low and bushy, as opposed to the Sperm Whale’s spout,
which is angled forwards and to the left. This, however, is less conspicuous in warm, tropical air.
Several of the rorquals can be separated only when seen well. Key features include size, the shape and
position of the dorsal fin, the shape of the head and the coloration. The size and colour of the baleen can be
helpful for distinguishing dead whales found stranded on beaches.

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 BLUE WHALE B. musculus, averages larger, and is mottled blue-
Balaenoptera musculus PLATE 54 grey with U-shaped head and tiny dorsal fin; Bryde’s
Measurements TL 22–24m, formerly up to 30.5m, Whale, B. edeni, and Sei Whale, B. borealis, can be
but excessive hunting has led to the extermination of difficult to distinguish, but lower jaw and baleen are
most of the larger, older individuals. Females slightly grey on both sides, dorsal fin is steeper (over 45°
larger than males. angle) and visible at same time as blow.
Identification Largest of the whales. General colour Ecology and habitat Eats a wide variety of food,
light bluish-grey mottled with greyish-white; belly including crustaceans, squid and small fish. Can
sometimes yellowish. Head flat from the side; broad dive up to 200m, but also feeds near the surface.
and U-shaped from above. Dorsal fin tiny, in last Winters in tropical areas, and migrates to colder,
quarter of back – rarely visible until whale dives. richer feeding areas in summer.
Flukes lifted only slightly out of the water or not at Distribution Worldwide. SE Asia: no confirmed
all when diving. Baleen relatively short, stiff and all records from the region, but individuals of northern
black. Similar species Fin Whale, B. physalus, is population reach the Philippines and are likely to
closest to this species in size but has a plain grey occur in South China Sea.
back and asymmetrical white on the underparts, a Status Endangered due to historic declines
more V-shaped head and a larger dorsal fin. associated with whaling; uncertain whether
Ecology and habitat Feeds mainly on krill (small recovering; ongoing threats include boat strikes.
crustaceans) within 100m of the surface. Migrates
long distances between tropical seas and cold SEI WHALE
waters. Balaenoptera borealis PLATE 54
Distribution Worldwide, with seasonal migrations Measurements TL 12–14m, maximum 17–21m;
between polar and tropical areas. SE Asia: southern females larger than males
animals migrate regularly to the Indian Ocean, and Identification Body uniform dark grey with ovoid
are likely to occur occasionally in the Andaman Sea, white scars; belly whitish. Conspicuous dorsal fin
while northern animals migrate regularly as far about one-third of way forwards from tail; angle of
south as Taiwan and may be expected in the South front of fin usually more than 40°. Single prominent
China Sea. A specimen in a museum in Myanmar ridge from blowhole to front of rostrum (rostral ridge);
was apparently stranded on the coast there. tip of snout slightly down-turned in profile. Dorsal fin
Status Endangered due to very large historic usually visible at same time as blow. Baleen uniform
declines associated with excessive whaling; grey-black with white fringes; sometimes a few half-
population recovery potentially threatened by white plates near front of mouth. Similar species
climate-change impacts on Antarctic krill. Fin Whale, B. physalus, has shallower dorsal fin,
visible just after the blow, and asymmetric white
FIN WHALE lower lips; Bryde’s Whale, B. edeni, is very similar,
Balaenoptera physalus PLATE 54 distinguished reliably only by having three ridges on
Measurements TL 18–20m, maximum 24–27m; top of the head.
males about 2m shorter than females Ecology and habitat Rarely dives very deeply,
Identification Dark grey to brownish-black above often feeding near the surface, breathing smoothly
with little or no mottling; whitish below. Often paler at regular intervals. Eats krill, squid and small fish.
patch behind blowhole. Asymmetrical head colour – Often in small groups. Migrates annually from high-
lower lip and palate white on right side, dark on left. latitude feeding grounds to low-latitude wintering
Right front baleen white, remainder striped dark grey grounds.
and whitish; up to 720mm long and 300mm wide. Distribution Worldwide, mainly in temperate and
Dorsal fin somewhat variable in shape, shallowly sub-tropical waters. SE Asia: uncommon in region,
angled; usually conspicuous shortly after blow. but confirmed from Gulf of Thailand, where an
Head broadly V-shaped from above, with a central individual was found on a beach in the peninsula.
ridge. Tail flukes rarely raised before diving. Tall, Status Endangered due to historic declines
cone-shaped blow. Similar species Blue Whale, associated with whaling.

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 BRYDE’S WHALE coloured baleen – white at the front of the mouth,
Balaenoptera edeni PLATE 54 dark grey at the back, with more white on the right
Measurements TL 10–12m in SE Asia (can be up side than the left side.
to 14m elsewhere) Ecology and habitat Feeds mainly on krill in the
Identification Dark grey above; whitish below. Three south, but also on small shoaling fish in the north.
prominent ridges on top of head. Baleen slate-grey, Populations in colder seas migrate seasonally to
up to 420mm long × 240mm wide. Rolls sharply to warmer waters.
expose fin and base of tail before diving. Taxonomic Distribution Found from Arctic waters to tropical
notes Sittang Whale, B. e. edeni, is substantially waters, including Indian Ocean, though largely north
smaller and genetically distinct from populations of the equator. SE Asia: potentially in any of the
elsewhere (B. e. brydei), and the two are sometimes coastal waters; recorded off coasts of Malacca and
considered different species. Similar species Other Penang in Peninsular Malaysia as well as peninsular
large whales have only one ridge on the head; Fin Thailand.
Whale, B. physalus, is larger, with white right lower Status Least Concern globally, but some populations
lip; Sei Whale, B. borealis, is larger than typical Asian are threatened by fisheries bycatch and hunting.
forms, with different feeding patterns.
Ecology and habitat Primarily feeds on schooling HUMPBACK WHALE
fish; best-studied populations usually dive deeply, Megaptera novaeangliae PLATE 54
returning irregularly to surface to breathe; not Measurements TL 11–13m, males maximum
known whether SE Asian form has similar habits. 15m; females maximum 16m
Distribution Throughout tropical and sub-tropical Identification Generally black or dark grey, with
seas. SE Asia: Sittang Whale, B. e. edeni, occurs in varying amount of white on belly. Dorsal fin usually
shallow waters in eastern Indian Ocean and west relatively small, sometimes similar to Blue or Fin
Pacific; confirmed records from Myanmar, Malaysia Whales, Balaenoptera musculus or B. physalus, but
and Thailand, as well as around the Philippines and often stepped. Very long flippers, white below and
South China Sea. usually partially white above. Tail raised high out
Status Data Deficient, due to uncertainty about of the water on deep dives, with distinctive deep
number of species involved. notch in middle, and individually variable amounts
of white and dark grey on the underside. Baleen
MINKE WHALE generally all black, 700mm long × 300mm wide.
Balaenoptera acutorostrata PLATE 54 Blow bushy and rounded. Similar species Other
Measurements TL 7–8m, males maximum 9.8m; rorquals have ridges instead of knobs on the head
females maximum 10.7m and smaller flippers, and do not raise their tail very
Identification Smallest of the rorquals, with narrow, high or at all before diving; Sperm Whale, Physeter
pointed V-shaped rostrum and a single ridge on top macrocephalus, can also appear hump-backed
of the head. Black to dark grey above, white below, when diving, but its tail is all dark, without a deep
usually with pale grey areas on the sides or back; notch, the head shape is quite different when seen
usually has conspicuous white bands on flippers. well, and the blow is forward.
Baleen about 200mm long × 120mm wide, mainly Ecology and habitat Feeds on krill and schooling
cream-coloured or white. Similar species Fin, Sei fish. Migratory.
and Bryde’s Whales, B. physalus, B. borealis and Distribution Worldwide, migrating between
B. edeni, are much larger with more rounded heads temperate and polar feeding areas and tropical
and more conspicuous blows; beaked whales have breeding areas. SE Asia: only a few records in the
similar dorsal fins, but distinctive head shapes. The region, though could potentially occur anywhere;
very similar Antarctic Minke Whale, B. bonaerensis, nearest known breeding areas to the west of India,
is found primarily in the southern hemisphere, but and north of Taiwan.
may occasionally cross the equator and could show Status Least Concern; experienced large population
up in SE Asia. It is distinguished by pale flippers that declines due to historic over-hunting, but populations
usually lack a banding pattern, and asymmetrically are now recovering in many areas.

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 Order SIRENIA
Family DUGONGIDAE DUGONG
The order Sirenia contains two families: the Trichechidae, found in the Americas, and the Dugongidae, found
in Africa, Asia and Australasia. Like the cetaceans, the sirenians are adapted to a life completely in the water,
but they appear to have evolved separately.
Sirenians live in shallow coastal seas, feeding only on plant material. The forelimbs are flippers and there
is a tail flipper, or fluke, but no dorsal fin. In adult Dugongs there is one pair of tusk-like incisors in the upper
jaw and none in the lower jaw. There are no canines or premolars, but a total of 10–12 molars, only some of
which are visible at one time; the deeper ones replace the top ones as they wear out.

DUGONG vegetation in shallow waters at night. Usually found

Dugong dugon PLATE 52 in small groups, which are believed to travel long
Measurements TL 2.5–3.5m. Wt 150–250kg distances between feeding areas.
Identification Broad, rounded head with two nostrils Distribution Seas off E Africa through India, Sri
at front of face and mouth on the underside; thick, Lanka, Indonesia, the Philippines, New Guinea
paddle-like flippers; fluked tail. Colour generally and N Australia. SE Asia: still occurs off coasts of
dull brownish-grey. Has a few sparsely scattered peninsular Myanmar, Malaysia, Thailand, Cambodia
hairs on body and stiff bristles on upper lip. Similar and Vietnam.
species Distinguished from small cetaceans by very Status Vulnerable globally, but locally Endangered;
different head shape, nostrils in front of face (not on one of the most threatened marine mammals in
top of head) and no dorsal fin. Less active than most the region; numbers have declined dramatically in
small cetaceans. recent years due to hunting, entanglement in fishing
Ecology and habitat Usually rests during the day gear and changes in coastal habitats resulting from
in deep water, and feeds on sea grasses and other the clearing of mangroves and from trawling.

Order PROBOSCIDEA
Family ELEPHANTIDAE ELEPHANTS
Elephants are the largest land animals still alive today, and include two genera, Loxodonta and Elephas. The
former includes the African elephants, which differ from the Asian Elephant in several respects, of which
the most conspicuous is their much larger ears. The Asian Elephant has two sorts of teeth: tusks and cheek
teeth. It possesses only four cheek teeth at a time (one on each jaw), which are shed periodically when they
are worn and no longer functional; a sixth and final set emerges at an age of about 40 years.
Elephants have been domesticated for a long time in Asia, being used particularly for moving heavy
loads, including hauling logs in forestry operations, as well as for transport and even in warfare. In many
areas where there has been extensive loss of forest, wild elephants may come into conflict with people by
feeding on agricultural crops, particularly palms. They are also subject to poaching, especially for their ivory.
Elephants can be dangerous to humans, especially if disturbed accidentally.

ASIAN ELEPHANT bristly hair. Adult males may have tusks up to 2m

Elephas maximus PLATE 55 long, although only about half is usually visible from
Measurements SH 1.5–3.0m, HB 3.0– 6.0m, outside; tusks are usually not visible externally in
T 1.0–1.5m. Wt 3,000– 5,000kg females and younger males. Footprints very large
Identification Very large mammal with distinctive (about 450mm across in adult), oval, with three toe
long trunk. Generally grey-brown, but paler when prints at front and one at side, all within the oval
dry and blackish when wet. Infants covered in black (Fig. 59a). Voice Has a broad vocal range including

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 very low frequency booms for long-distance larger groups have sometimes been reported.
communication; loud trumpeting when alarmed; Groups split and merge, but normally contain one
softer snorts within herd, often silent. or more adult females with young of both sexes and
Ecology and habitat Mostly active from about various ages. Adult males occur loosely attached to
two hours before dusk, through the night until two herds, or as solitary individuals or in groups of two
hours after dawn. Diet consists mainly of tender or three. They can swim across rivers.
parts of various plants, including soft grasses, the Distribution SE Asia: Myanmar, Thailand, Laos,
growing parts of palms and banana stems. Fruits Vietnam, Cambodia and Peninsular Malaysia. Also
and the bark of latex-producing trees are also India, Sri Lanka, Sumatra and Borneo.
eaten, but only in relatively small quantities. Water Status Endangered. Populations have declined
and minerals are important to elephants, and may dramatically in much of range, owing to illegal
influence their distribution. Usually found in groups hunting as well as conflicts between humans and
of typically between 3 and 40 individuals, although elephants in agricultural areas.

a b

H
c

F
H
H

0 200

Fig. 59 Prints of left feet of Elephas maximus (a), Rhinoceros sondaicus (b), Dicerorhinus sumatraensis (c) and Tapirus
indicus (d). H = hind foot, F = front foot. Print of fourth front toe on tapir may not be visible on firm ground.

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 Order PERISSODACTYLA
Odd-toed ungulates: tapirs and rhinoceroses
Members of this order are large herbivorous mammals with one or three toes on each foot (except tapirs,
which have four toes on each front foot). They include horses, rhinoceroses and tapirs. The stomach is rather
small and simple, while the caecum (large intestine) is large. Until recently, three species occurred in the
region, but the two rhinoceroses are now extinct in mainland South-east Asia.

Family TAPIRIDAE TAPIRS

Four species of tapir are currently recognised, all in the same genus. Three of these occur in South America,
and only one species occurs in Asia. All are very similar in shape, with an elongate nose, curved back and
short tail.

ASIAN TAPIR with different colour pattern; pigs, Sus spp., are
Tapirus indicus PLATE 56 smaller, with longer, more pointed head, thinner
Measurements SH 900–1,050, HB 2,000–2,400, legs, different footprints.
T 50–100. Wt 250–350kg Ecology and habitat Found mainly in intact
Identification Adult has front part of body black, rainforest, from lowlands up to about 2,000m in
including head and front legs, as well as lower part northern part of range where most lowland forest
of hind legs, the remainder being white. Newborn has been lost. Most active at night. Often found near
young dark grey, completely covered with horizontal streams, but does not wallow in mud baths.
white stripes and rows of spots; gradually starts to Distribution SE Asia: S Myanmar, S Thailand and
acquire adult pattern at about four months of age. Peninsular Malaysia. Also Sumatra.
Tracks similar to those of rhinoceros, but smaller Status Endangered. Still found in many protected
(150–170mm across), front foot with fourth toe, areas in Malaysia, but overall population has declined
though this is usually visible only on soft ground dramatically because of loss and fragmentation of
(Fig. 59d). Similar species Asian Two-horned habitat and hunting.
Rhinoceros, Dicerorhinus sumatrensis, is larger,

Family RHINOCEROTIDAE RHINOCEROSES

Rhinoceroses are characterised by a large, stocky body; short, stout legs with three toes on each foot; and
either one or two horns, made of compacted hairs on top of the muzzle. There are five living species of
rhinoceros (two in Africa and three in Asia) of which two occur in this region. However, rhinoceroses have a
long fossil history, and millions of years ago there were also forms occurring in Europe and the Americas. All
of the rhinos are legally protected throughout their range, but are still illegally persecuted for their horns and
other parts of the body, which are falsely believed to have medicinal properties.
Two species were still found in the region when the first edition of this book was published, but both
are now believed to be extinct in mainland South-east Asia. Small populations of both species remain in
Indonesia, but they are severely threatened and at risk of global extinction.

ASIAN TWO-HORNED RHINOCEROS to see in field. Only one fold of skin crosses back,
Dicerorhinus sumatrensis PLATE 55 behind shoulder; folds of skin on neck and lower
Measurements SH 1.2–1.3m, HB 2.4–2.6m, belly do not cross over back; no folds of skin on hind
T 650. Wt 900–1,000kg legs. Two horns, front one relatively narrow, typically
Identification General coloration usually dark 250–300mm in males and smaller in females; rear
brown, but appearance may vary after bathing horn short, rarely more than 100mm, and often not
in water or mud. Skin thick with extensive sparse visible in field. More often detected by tracks than
hairs, especially when young, but hairs may be hard seen: footprints of adults on firm soil 185–235mm

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 at widest point, showing three clear toe marks Status Critically Endangered. Formerly widespread
(Fig. 59c). Mud wallows can be distinguished throughout mainland SE Asia, but illegal hunting
from those of pigs by clear, deep horn marks in and loss of habitat has reduced numbers to fewer
sides of wallow. Dung consists of balls of coarsely than 100 individuals in isolated areas in Sumatra
chopped woody material about 90mm in diameter, and Borneo.
usually found in small piles, sometimes in mounds
frequently used over long periods. When disturbed LESSER ONE-HORNED RHINOCEROS
unexpectedly, this rhinoceros usually flees rapidly, Rhinoceros sondaicus PLATE 55
sometimes giving a series of short, hoarse barks. Measurements SH 1.6–1.8m, HB 3.0–3.2m,
Similar species Lesser One-horned Rhinoceros, T 700. Wt 1,500–2,000kg
Rhinoceros sondaicus, is larger, with three folds of Identification Large, distinctly shaped heavy
skin crossing back; footprints of similar shape but animal; thick, dark grey skin; three folds of skin
larger; Asian Tapir, Tapirus indicus, has similar but across back, one on neck, one behind shoulders
smaller footprints, conspicuous black-and-white and one over lower back; an additional horizontal
body colour; large pigs, Sus spp. have much lighter fold on rump just below tail, and on hind leg. There
bodies with slender legs, relatively large head and is a single relatively thick horn on the tip of the
two-hoofed footprints. nose, rarely longer than 150mm in male (maximum
Ecology and habitat Mainly active from late c.250mm); smaller and inconspicuous in female.
afternoon to mid-morning. Usually rests during hot Footprints of adults large, about 280–300mm
hours of the day in a mud wallow, shaded spot or across, with three conspicuous toe prints (Fig. 59b).
ridge top. Feeds on mature leaves and twigs from Similar species Asian Two-horned Rhinoceros,
a wide range of woody plants, including saplings, Dicerorhinus sumatrensis, is smaller, with only one
lianas and small trees, which may be pushed over fold of skin across back; often has a second horn.
to obtain the leaves. Occasionally eats fallen fruits. Ecology and habitat Currently restricted to
Usually found within 10–15km of natural mineral dense lowland rainforest with extensive wet areas,
sources. Formerly occurred in a variety of forest including streams and mud wallows. Formerly
types, but now largely in lowland rainforest, although occurred in a greater variety of habitats, including
sometimes in fairly hilly areas. Uses regenerating mixed forest and grasslands.
forests, but rarely enters open areas. Distribution SE Asia: formerly occurred throughout
Distribution SE Asia: formerly occurred throughout region. Last remaining small population occurred
much of region, but last remaining population in in Vietnam, but last individual was illegally killed in
Peninsular Malaysia was declared extinct in 2015. 2010. Now known only from W Java.
Still occurs in very small numbers in Sumatra and Status Critically Endangered; only about 60
Borneo. individuals left in Java.

Order ARTIODACTYLA
Even-toed ungulates: pigs, mousedeer, deer, cattle and sheep
The even-toed ungulates are terrestrial mammals with two functional hoofed toes and two small dew toes on
each foot. The dew toes are situated well above the main toes and, except in pigs, rarely appear in footprints
except in very soft soil or mud. Most species, other than pigs, feed mainly on plant material. The stomach is
rather large and complex, with two or three chambers in pigs, three in mousedeer and four in other species.
The caecum is small. For all species other than pigs, food is partially digested in the stomach and then
brought back up into the mouth one or more times for further chewing. This increases the amount of nutrition
that can be obtained from grasses and leaves, which contain a high percentage of cellulose that is difficult to
digest. It also permits ruminants to ingest large amounts of food quickly before moving to sheltered places,
better protected against predators, to continue their chewing.

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 Family SUIDAE PIGS
Unlike other artiodactyls, pigs have incisors in the upper jaw (Fig. 61) and an omnivorous diet, including
animal material. Pigs can be distinguished by their elongate snout with a flattened tip and nostrils in the
middle. Canines of males are greatly enlarged, forming tusks that protrude sideways.

EURASIAN WILD PIG Distribution SE Asia: Myanmar, Thailand, Laos,

Sus scrofa PLATE 56 Vietnam, Cambodia and Peninsular Malaysia. Also
Measurements SH 600–800, HB 1,350–1,500, much of Europe, N Africa and mainland Asia, with
T 200–300. Wt of large male 75–200kg introduced populations elsewhere.
Identification Colour varies from blackish to reddish; Status Least Concern.
muzzle elongate without a beard; long black hairs
on upper back and neck, forming a mane. Young BEARDED PIG
are born dark brown with elongate white stripes Sus barbatus PLATE 56
along body. Footprints more rounded and symmetric Measurements SH 700–900, HB 1,200–1,500,
than those of deer, with imprints of the dew toes T 170–250. Wt 50–120kg (sometimes considerably
apparent except on very hard ground (Fig. 60). Voice heavier when storing fat)
Pigs in herds are relatively noisy, making many Identification Colour varies from blackish (in young
grunting and squealing noises; also often sounds animals) to reddish-brown or yellow-grey; may
of breaking vegetation. Taxonomic notes ‘Banded also be affected by colour of mud wallow. Head
Pig’, S. scrofa vittatus, from Peninsular Malaysia and long; fleshy protuberance (‘wart’) above each side
Indonesia, is distinctly smaller with sparser hair, and of mouth with upwards-pointing bristles, larger in
sometimes considered a separate species. Similar males than females; large beard of long wiry hairs
species Bearded Pig, S. barbatus, averages taller, on snout. Lower canines of males relatively straight,
has longer legs, longer muzzle with beard and bristly protruding to sides. Body size varies greatly with
warts; domestic pigs, which are descended from food supply. Females have five pairs of mammae.
S. scrofa, are sometimes very similar, but generally Mud wallows are a distinctive feature of pig activity.
tamer, and usually lack a black mane. Footprints similar to those of other wild pigs (Fig.
Ecology and habitat Found in a wide variety of 60). Voice Various squeals and grunting, as well as
habitats, including mature forests, disturbed areas, loud snorts or crunching sounds of pigs feeding on
secondary forest, gardens and plantations. Feeds on hard plant material. Similar species Eurasian Wild
roots, seeds and a wide variety of other vegetable and Pig, S. scrofa, is smaller, with shorter legs, lacks
animal matter, such as invertebrates, worms and small facial protuberances and beard.
vertebrates including birds’ eggs and nestlings, if Ecology and habitat Active day and night; tends
encountered on the ground. In gardens and plantations to be more nocturnal where heavily disturbed. Diet
can cause extensive damage to growing crops. includes fallen fruits and seeds, roots, herbs and
other plant material, earthworms and other small
animals which it finds by grubbing on the ground.
Can cause damage to plantations, feeding on the
growing parts of young palms and on cocoa fruits.
In regions of extensive forest, periodically forms very
large herds which travel great distances in search
of food, swimming across rivers and climbing into
mountain ranges. Adult females make nests of
saplings and shrubs, which are bitten or torn off
and piled up on the ground, where they give birth
0 100 to young (from 3–11 piglets at one time). Nests are
made on dry ground not liable to flooding.
Fig. 60 Left hind footprints of Sus scrofa on hard ground Distribution SE Asia: Peninsular Malaysia. Also
(left) and soft ground (right). Sumatra, Borneo and some adjacent islands.

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 Status Vulnerable. Populations declining throughout
range due to over-hunting and loss of large areas of
continuous forest. In Peninsular Malaysia population
is at imminent risk of extinction; it has disappeared
even from large protected areas, possibly due to
disruption of traditional migration routes or hunting.

VIETNAMESE WARTY PIG

Sus bucculentus NOT ILLUSTRATED 0 100
Measurements Skull: gl 350–390
Identification Uncertain whether this represents a Fig. 61 Skull of Sus barbatus.
legitimate species. Known only from two skulls from
Vietnam, and an additional skull of a juvenile from and relatively larger premolars. However, recent
Laos. Supposedly differs from Eurasian Wild Pig, genetic analyses of the Lao specimen indicated that
S. scrofa, in having a more elongate skull, straighter it fitted within genetic sequences of typical S. scrofa,
dorsal margin of the braincase, higher occipital crest suggesting it may not be a distinct species.

Family TRAGULIDAE MOUSEDEER

Mousedeer (also called chevrotains) are small forest-dwelling ungulates with slender legs and no antlers.
The back is strongly arched. The lower canines are long, and may protrude out of the mouth from the upper
jaw of males. Three or four species are currently recognised from the region.

LESSER MOUSEDEER coloration on upperparts and different throat markings;

Tragulus kanchil PLATE 57 Silver-backed Mousedeer, T. versicolor, has orange-
Measurements SH 200–230, HB 400–550, brown on sides of neck and shoulders, contrasting with
T 60–90. Wt 1.4–2.5kg finely speckled silver lower back and rump.
Identification Upperparts with relatively fine fur, Ecology and habitat Active periodically both night
reddish-brown mixed with fine black, appearing and day. Diet includes fallen fruits, leaf shoots and
fairly uniform coloured overall; centre of nape usually fungi. Usually solitary. Both adults and young rest in
blacker than rest of back, often forming a distinct any sheltered spot under forest cover. Occurs in tall
stripe. Underparts white, with variable brown stripes up and secondary forests, sometimes entering gardens
middle or along sides of belly; distinctive dark brown and mixed scrub.
and white markings on throat and upper chest, typically Distribution SE Asia: Myanmar, Thailand, Laos,
with triangular white stripe up middle, bordered by dark Vietnam, Cambodia and Peninsular Malaysia. Also
brown triangle, then one diagonal stripe on each side, SW China, Sumatra, Borneo and many adjacent
which usually join on chin; in profile white appears as a islands.
single line from chin to side of chest. Some geographic Status Least Concern, but populations in many
variation in colour, with populations in peninsular areas are declining as a result of forest loss and
Thailand and Malaysia darker and more reddish; those excessive hunting.
on rest of mainland duller, often lacking nape stripe;
additional colour variations occur on islands. Footprints GREATER MOUSEDEER
similar in shape to other deer, but much smaller (Fig. Tragulus napu PLATE 57
62). Taxonomic notes Formerly considered part of Measurements SH 300–350, HB 520–572,
species T. javanicus, which is now considered to be T 60–100, HF 140–157. Wt 3.5–4.5kg
restricted to Java; population in N Thailand is larger Identification Upperparts appear coarsely mottled
and sometimes considered a separate species, orange-buff, grey-buff and blackish; hairs on
T. williamsoni, but only the type specimen is known, upperparts grey-buff to orange-buff with blackish
and its distinctiveness is uncertain. Similar species tip; darker in midline and paler on sides of body;
Greater Mousedeer, T. napu, is larger, with more flecked often with a darker nape patch. Intensity of

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 a

0 100

Fig. 62 Left hind footprints on hard ground (left) and soft ground (right) of Muntiacus muntjak (a), Cervus unicolor (b),
Tragulus napu (c) and T. kanchil (d).

coloration varies between individuals. Underparts declined throughout range, owing to loss of forest
white, usually without brown stripes on belly; pattern habitat and excessive hunting.
of brown and white markings on upper chest and
underside of neck, typically with triangular white SILVER-BACKED MOUSEDEER
stripe in centre, bordered by dark brown stripes, Tragulus versicolor PLATE 57
then two separate diagonal white stripes on each Measurements SH 250, HB 400–450. Wt 1.7kg
side, one originating near front of chin, and one in Identification Head, sides of neck and shoulders
middle of throat, though sometimes these join; in reddish-orange mottled with black, especially on
profile, usually appears as two separate white bars back of neck, which typically shows a distinct
on side of neck. Footprints similar in shape to those darker stripe; rest of back, flanks and hindquarters
of Lesser Mousedeer, but distinctly larger (Fig. 62c). silver-grey finely speckled with dark grey-brown.
Similar species Lesser Mousedeer, T. kanchil, is Underparts white in middle; upper chest and throat
substantially smaller, with more finely patterned with pattern of reddish-brown and white stripes,
reddish fur, usually only a single white stripe on side usually with one continuous white stripe on side
of throat; Silver-backed Mousedeer, T. versicolor, is of neck. Similar species Lesser Mousedeer,
smaller, with silver-grey speckling on back. T. kanchil, is slightly smaller, with upperparts all
Ecology and habitat Occurs in tall and secondary mottled reddish-brown and black.
forests, sometimes entering gardens. Mainly Ecology and habitat Believed to inhabit understorey
nocturnal, but sometimes also active during the of tall lowland forest.
day. Diet includes fallen fruits, leaf shoots and other Distribution SE Asia: S Vietnam.
vegetation. Status Data Deficient. Very few historic records,
Distribution SE Asia: S Myanmar, S Thailand and with no recent information on whether the species
Peninsular Malaysia. Also Sumatra, Borneo and still persists; likely to be threatened by loss of
adjacent islands, and Palawan (the Philippines). lowland forest and hunting, and may be Endangered
Status Least Concern, but populations have or even Extinct.

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 Family MOSCHIDAE MUSK-DEER
Medium-sized ungulates closely related to deer with relatively small head and enlarged ears. Both sexes
lack antlers, but males have long, narrow upper canines that protrude from upper jaw as backwards-curving
tusks. Hind legs are longer than forelegs, so that shoulder is slightly lower than rump; this allows them to
jump very strongly. All four toes on feet relatively well developed, and can be used to support animal when
climbing in lower branches of trees. Several species of musk-deer occur in China, the Himalayas and Siberia,
with two species in the region, one in N Myanmar and one in N Vietnam. Adult males have a gland on the
underside of the belly which produces a strong-smelling musk that is highly sought-after for perfumes;
illegal hunting to obtain this musk has caused declines in all species, and some are now Endangered. A few
species are now farmed in China.

FOREST MUSK-DEER Status Endangered. In Vietnam, population has

Moschus berezovskii PLATE 58 declined from several hundred individuals to only
Measurements SH 500–600, HB 700–800, T 40. a few, largely due to illegal hunting. In China,
Wt 6–9kg still moderate numbers remaining, but declining
Identification Upperparts dark brown, finely dramatically due to loss of habitat and hunting.
speckled with olive-brown or orange-brown spots,
darker on lower back and rump as well as top of BLACK MUSK-DEER
head; underparts including legs reddish-brown to Moschus fuscus PLATE 58
yellowish-orange; throat and neck with long, pale Measurements SH c.600, HB c.700. Wt 8kg
orange or white stripes on underside, one in the Identification Upperparts dark brown to blackish;
middle and one on each side, sometimes broken head, neck, ears black; underparts dark, similar to
into separate spots; ears orange-brown at base, back; throat dark, often with two incomplete yellowish
black at tip, white inside. Newborn young blackish- collars. Juveniles dark brown with buff spots. Similar
brown above with rows of buff spots on back. species Forest Musk-deer, M. berezovskii, has pale
Similar species Black Musk-deer, M. fuscus, is underparts, long yellow stripes on neck.
darker with dark underparts, black ears, reduced Ecology and habitat Forest, at higher altitudes near
markings on neck. tree line, often with pines and rhododendrons.
Ecology and habitat Largely forested areas on Distribution SE Asia: N Myanmar. Also S China
limestone karst outcrops in Vietnam. In China, (Yunnan, Tibet), NE India (Assam) and Nepal.
occurs in a variety of forested habitats, mostly at Status Endangered. Population believed to have
higher elevations. declined dramatically due to loss of habitat and
Distribution SE Asia: N Vietnam. Also S and C China. excessive hunting, especially for illegal trade in musk.

Family CERVIDAE DEER

Males have distinctive antlers, which are usually shed and regrown at regular intervals (probably annually),
unlike the horns of Bovids, which are permanent. Antlers are a form of bone and grow from permanent
projections, called pedicels, on the skull. There has been some confusion about the appropriate generic
relations for large deer. The most recent genetic analyses indicate that the three remaining large SE Asian
species are all closely related and should be placed in the same genus, Cervus. They tend to walk with the
head held high and the back fairly straight. Muntjacs, Muntiacus spp., walk with the head carried low, the
back slightly arched and the hindquarters high, lifting the feet high off the ground at every step. Adult male
muntjacs have long upper canines which protrude beyond the lip and are loose in their sockets. A rattling
noise, probably produced by these loose teeth, is sometimes heard from muntjacs observed at close range.

SAMBAR antlers along greatest curve typically 300–550

Cervus unicolor PLATE 59 Identification Large deer with dark brown skin and
Measurements SH 1,400–1,600, HB 1,500– hair; upperparts generally grey-brown, sometimes
2,000, T 210–280. Wt 180–260kg. Length of slightly reddish, usually darker along midline;

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 underparts same colour as back or darker. Tail on each antler. Footprints similar to those of other
bushy, mainly blackish except for white on base of large deer. Voice During mating season, male gives
underside. Middle of rump sometimes paler than rest loud bugling call. Taxonomic notes Mainland forms
of back. Adult males with long, coarse hair on neck. may be a different species from those in Japan, in
Young usually without spots. Adult males typically which case name may change. Similar species
have three tines to each antler, one in front and two Sambar, C. unicolor, is larger, without spots in
at the tip of the main beam; inner branch of terminal adults, antlers usually have fewer tines; Eld’s Deer,
fork of antler normally somewhat smaller than the C. eldii, has brow tine of antler forming continuous
outer, which appears to be a continuation of the curve with main branch.
main beam of the antler. Two-year-old males have Ecology and habitat Poorly known in region.
antlers with only one point; three-year-old males Probably largely nocturnal and solitary.
have antlers with two points. Footprints relatively Distribution SE Asia: Vietnam. Also E China, Russia,
large, pointed at front (Fig. 62b). Taxonomic notes Japan and Taiwan.
Formerly placed in the genus Rusa. Voice Call is a Status Least Concern globally due to healthy
distinctive yelp or shrill bark. Similar species Eld’s populations in Japan and Taiwan, but elsewhere
Deer, C. eldii, is more reddish, underside pale, tail populations greatly reduced. Vietnam population
not black, brow tine of antler forming continuous believed extinct in wild, but persists in captivity, as
curve with main branch. well as semi-captive populations in some reserves.
Ecology and habitat Active mainly at night, also
early morning and late afternoon. Diet includes ELD’S DEER
grasses, herbs, shrubs, young leaves of woody plants Cervus eldii PLATE 59
and fallen fruits. Often visits natural mineral sources. Measurements SH 1,200–1,300, HB 1,500–
Usually solitary, but groups of two are sometimes 1,700, T 220–250. Wt 95–150kg
seen, consisting of adult male and female, female Identification Upperparts reddish-brown, males
and young, or adult females. Nocturnal activity and darker than females; underparts whitish. Adult
solitary nature possibly a consequence of heavy males have thick long hair on neck, forming a mane.
hunting pressure. Most common in secondary Brow tine of antler angled strongly forwards, forming
forests on gently sloping terrain, but also in tall continuous curve with main branch, in a bow shape;
dipterocarp forests on steep terrain and in swamp many small tines at tip of antler. Fawns have many
forests. Enters gardens and plantations to feed. pale spots, which sometimes persist in adults.
Distribution SE Asia: Myanmar, Thailand, Laos, Footprints similar to those of Sambar, C. unicolor
Vietnam, Cambodia and Peninsular Malaysia. Also (Fig. 62b). Taxonomic notes Formerly placed in
Sri Lanka, India, S China, Sumatra, Borneo and the genus Rucervus. Similar species Sambar,
many larger adjacent islands and the Philippines. C. unicolor, is more greyish-brown with a black tail,
Status Vulnerable: has declined dramatically in antlers with front tine angled backwards.
many parts of range, especially Indochina, as result Ecology and habitat Lowland swampy areas, as
of excessive hunting. well as open, dry dipterocarp forest and grasslands.
Feeds mainly on grasses; sometimes enters rice
SIKA fields.
Cervus nippon PLATE 59 Distribution SE Asia: Myanmar, Laos, Cambodia.
Measurements SH 650–1,100, HB 950–1,800, Formerly occurred in Thailand and Vietnam, but
T 75–150. Wt 80kg likely now extirpated. Also India, China.
Identification Moderately large deer with dark Status Endangered. Populations declining rapidly in
brown to reddish-brown upperparts, usually with most areas, owing to hunting and habitat loss.
rows of large white spots; underparts paler. In
winter, coat is darker and longer, with fewer white SCHOMBURGK’S DEER
spots. Rump has small white patch around base Rucervus schomburgki PLATE 59
of tail. Tail small. Young with extensive spots. Ears Measurements SH 1,050
relatively small. Adult males typically have five tines Identification Upperparts uniform brown, slightly

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 darker on top of tail, white under tail, forehead and TUFTED DEER
legs reddish; underparts pale. Antlers with short Elaphodus cephalophus PLATE 58
main beam and numerous branches with a total of Measurements SH 500–700, HB 1,000–1,200,
10–30 tines on both antlers. Fawns with pale spots. T 70–150. Wt 17–30kg
Similar species Other deer have many fewer tines Identification Upperparts dark chocolate-brown,
on antlers. blacker on legs, slightly greyer on head and neck;
Ecology and habitat Formerly found mainly in underparts paler; hair coarse and slightly shaggy.
swampy plains and river floodplains, in areas of long Top of head pointed in profile with distinct tuft of
grasses, canes and shrubs. black hair; pedicel very short with small antlers,
Distribution SE Asia: Formerly C Thailand. often hidden by tuft of hair. Ears with conspicuous
Status Extinct; last reliable reports of live animals in white marks; tail dark with white underside. Upper
1930s, although some antlers still found occasionally canines elongate, visible outside mouth in males.
in trade. Species was lost due to conversion of Voice A sharp bark. Similar species Muntjacs,
habitat to rice farms and excessive hunting. Muntiacus spp., tend to be somewhat smaller,
without tuft on top of head and with longer antlers.
HOG DEER Ecology and habitat Upper montane areas in damp
Axis porcinus PLATE 58 forest near snowline.
Measurements SH 650–720, HB 1,400–1,500, Distribution SE Asia: formerly N Myanmar. Also
T 170–210. Wt 70–110kg China.
Identification Upperparts brown to dark brown Status Near Threatened. No recent records from
in winter, greyer in summer; underparts paler. Tail Myanmar and possibly extirpated; still persists in
coloured as back, with white underside; centre of China, but declining due to excessive hunting.
back darker; fawns with rows of small white spots
that may persist in adults. Antlers of males slender, Genus Muntiacus Muntjacs can be distinguished
forked at tip of main beam, with small third prong at from other deer by their relatively small size, and
base pointing upwards. Footprints similar in shape by their distinctive posture with curved back and
to those of Sambar (Fig. 62b), but smaller. Similar head held low. Males have large bony pedicels,
species Sambar, Cervus unicolor, is much larger, and generally small antlers. Several species are
with longer legs, larger antlers, dark tail; Large- currently recognised from the region, some of which
antlered Muntjac, Muntiacus vuquangensis, is have been supported by genetic studies. There is
similar in size, but has much larger pedicel, antlers considerable uncertainty about how to identify them,
not forked at tip, short triangular tail, not spotted, however, as some have been reported only from
different habitat. trophies found in villages, and the field characters
Ecology and habitat Natural habitat mainly wet or are not described. In particular, there is uncertainty
moist tall grasslands in lowland floodplains, feeding over the number of species of small dark muntjacs
mainly on grasses. found in Indochina, and no information is available
Distribution SE Asia: Myanmar and Cambodia, on how to tell them apart in the field.
formerly Thailand, Laos and Vietnam. Also
Pakistan, N India, Nepal and Bangladesh, with RED MUNTJAC
some introduced populations elsewhere. Muntiacus muntjak PLATE 57
Status Endangered. Populations declining dramati- Measurements SH 500–550, HB 900–1,100,
cally owing to hunting and conversion of wet T 170–190. Wt 20–28kg. Antler length 70–130,
grasslands to agriculture. Probably extirpated from pedicel length 70–150
Thailand (though subsequently reintroduced in some Identification Small reddish deer, males with
areas), Laos and Vietnam, with small remaining very large pedicels and relatively small antlers.
populations in Myanmar and Cambodia. Upperparts vary from deep reddish-brown or rufous
to paler reddish-yellow, generally darker along
midline; underparts pale whitish or greyish; tail
coloured as back above, white below. Antlers rough

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 with small spike near base, sharp curve near tip.
Pedicel thick and straight, with a burr where antler
joins; continues as bony ridge down forehead,
usually with conspicuous black lines. Female has a
tuft of stiff hair in place of pedicel or antlers. Young
normally with white spots. Footprints with relatively
elongate toes, pointed at front (Fig. 62a), probably
indistinguishable from those of other muntjacs
except by size. Taxonomic notes Populations north
of peninsular Thailand are sometimes considered a
distinct species, Northern Red Muntjac, M. vaginalis,
based in part on chromosomal differences, but
there has been no comprehensive analysis of genes
and morphology to support this. Voice Adults of 0 100
both sexes give short, loud, barking calls. Similar
species Fea’s Muntjac, M. feae, is browner, with Fig. 63 Skull of male (top) and female (bottom)
dark legs, pale top of head; Gongshan Muntjac, Muntiacus muntjak.
M. gongshanensis, has black belly and legs; Leaf
Muntjac, M. putaoensis, is smaller, with short similar, but may be darker with shorter antlers; field
pedicels; Large-antlered Muntjac, M. vuquangensis, identification characters poorly known because
is substantially larger, with long antlers and relatively species descriptions are based on incomplete skulls
short pedicels. and trophies.
Ecology and habitat Found in a wide variety Ecology and habitat Apparently found mainly in
of forest habitats, from tropical rainforest to dry evergreen wet hill forest.
dipterocarp forest, lowlands to hills. Browses on Distribution SE Asia: N Laos, possibly N Myanmar
leaves and twigs more often than grazing on grass; and probably N Vietnam.
also eats fallen fruits. Mainly nocturnal, although Status Data Deficient. Uncertain owing to confusion
also active during the day in areas where not hunted. with M. truongsonensis; however, all small muntjacs
Largely solitary except during breeding season. in the M. rooseveltorum complex are heavily hunted
Distribution SE Asia: Myanmar, Thailand, Laos, and likely to be declining.
Vietnam, Cambodia and Peninsular Malaysia. Also
Sri Lanka, India, S China, Taiwan, Sumatra, Java, ANNAMITE MUNTJAC
Borneo and some adjacent islands. Muntiacus truongsonensis PLATE 57
Status Least Concern, though declining owing to Measurements Estimated SH c.400. Wt c.15kg
excessive hunting in some areas. This is the most Identification Small blackish muntjac, with short
widespread muntjac in the region, and the only pedicels (about 40mm) and very short antlers, about
species in many areas. 20mm long. Canines relatively large in both males
and females. A muntjac described in life from Laos,
ROOSEVELT’S MUNTJAC which may have been this species, had dark brown
Muntiacus rooseveltorum NOT ILLUSTRATED to black fur on most of the body and legs, with
Measurements Estimated SH c.400 orange-brown crown and light brownish neck, very
Identification Small muntjac with dark brown coat, short antlers. Taxonomic notes The relationships of
white throat, short brown tail; large glands on chin, this species with M. puhoatensis, M. putaoensis and
dark legs; moderate antlers. Taxonomic notes The M. rooseveltorum are uncertain. Similar species
number of species of small muntjacs in the region Roosevelt’s Muntjac, M. rooseveltorum, may have
is uncertain. This is the oldest name among them, longer antlers, but field characters to identify species
and hence would be a valid name if several prove uncertain; Gongshan Muntjac, M. gongshanensis,
to be the same species. Similar species Annamite is similar in colour but substantially larger, with a
Muntjac, M. truongsonensis is apparently very heavier build and proportionately larger head.

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 Ecology and habitat Evergreen wet hill forest GONGSHAN MUNTJAC
above 1,000m in Annamite Mountains. Muntiacus gongshanensis PLATE 57
Distribution SE Asia: mountains of Vietnam, Laos Measurements SH 500–550, HB 900–1,100,
and possibly Myanmar. T 200. Wt 21–28kg
Status Data Deficient. Camera trap and hunter Identification Body mid- to dark brown, with belly,
surveys suggest small dark muntjacs are still outsides of legs and top of tail black; insides of legs,
regularly encountered in hill forest above 1,000m underside of tail contrasting white. Head pale brown
altitude, but species identification is uncertain. with dark lines on pedicels. Antlers relatively short.
Likely to be threatened by excessive hunting. Taxonomic notes Some studies have suggested
this is a form of M. crinifrons from E China, but
PUHOAT MUNTJAC further analyses are required. Similar species
Muntiacus puhoatensis NOT ILLUSTRATED Red Muntjac, M. muntjak, is uniformly reddish,
Identification Small dark muntjac that apparently with larger pedicels and antlers; Leaf Muntjac,
differs from Annamite Muntjac, M. truongsonensis, M. putaoensis, is smaller; Fea’s Muntjac, M. feae,
in genetic characters, but differences in appearance has underparts same colour as back, rather than
of live animals not presently known. May prove black, with brown rather than black upper legs.
to be the same species as M. rooseveltorum or Ecology and habitat Sub-tropical and temperate
M. truongsonensis. forest in montane areas, up to alpine scrub.
Ecology and habitat Evergreen wet hill forest Distribution SE Asia: N Myanmar. Also S China
above 1,000m in Annamite Mountains. (Yunnan). May also occur in India and Vietnam.
Distribution SE Asia: Vietnam. Status Data Deficient. Uncertain range and status
Status Data Deficient. Uncertain how or whether this owing to limited records with confirmed identify.
differs from M. truongsonensis and M. rooseveltorum. Probably threatened by excessive hunting and
trapping.
LEAF MUNTJAC
Muntiacus putaoensis PLATE 57 FEA’S MUNTJAC
Measurements Estimated SH c.400. Wt 15kg Muntiacus feae PLATE 57
Identification Small muntjac similar in size to Measurements SH 500–550, HB 880, T 100.
Annamite Muntjac, M. truongsonensis, also with Wt 22kg
short pedicels (less than 40mm in length) and short, Identification Upperparts dark brown, finely
unbranched antlers 10–35mm long, but differing speckled with yellow; underparts brown, sometimes
genetically and in colour. Fur generally reddish- with pale areas; legs darken to black near hoof,
brown with slightly darker legs and face; some white stripe on front of hind leg; face dark brown
individuals may be darker. Taxonomic notes The on sides, yellowish-brown on crown, pedicel and
relationships with M. rooseveltorum, M. puhoatensis, base of ears, except for black stripe from just
and M. trongsonensis remain uncertain. Similar above eyes on inside of pedicel; tail short with
species Annamite Muntjac, M. truongsonensis, is black upperside, white underside. Pedicels well
believed to be darker on body, paler on head; Red developed, antlers relatively short. Well-developed
Muntjac, M. muntjak, is much larger, with longer lachrymal gland (in front of eye). Similar species
pedicels and larger antlers. Red Muntjac, M. muntjak, is reddish-orange in
Ecology and habitat Temperate and sub-tropical colour without contrasting paler top of head;
hill forest. Regularly detected at altitudes of 700– Gongshan Muntjac, M. gongshanensis, has black
1,200m in Myanmar. belly, legs all black.
Distribution SE Asia: N Myanmar. Also N India and Ecology and habitat Evergreen forest in hilly or
S China. mountainous areas.
Status Data Deficient. Status uncertain, but camera Distribution SE Asia: S Myanmar and peninsular
trapping suggests it may be reasonably common Thailand.
in some areas. Probably threatened by excessive Status Data Deficient. Distribution and status
hunting and trapping. uncertain as limited confirmed records; photographs

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 can be difficult to identify with confidence. Could be of male; tail short and triangular, coloured as back
threatened owing to apparently restricted range, above, white below. Similar species Red Muntjac,
habitat loss and some hunting. M. muntjak, is smaller, with reddish coat, longer tail,
shorter antlers and longer pedicel; Hog Deer, Axis
LARGE-ANTLERED MUNTJAC porcinus, has darker, greyer fur, short pedicel, with
Muntiacus vuquangensis PLATE 58 thinner, more forked antlers.
Measurements SH 650–700. Wt 34kg Ecology and habitat Main habitat is mature
Identification Relatively large muntjac, with evergreen forest, mostly in hills, but somewhat
distinctive large antlers in male; antlers up to tolerant of disturbed forest.
280mm long on moderately long, stout pedicels; Distribution SE Asia: Annamite Mountains of Laos
fork at base well developed, up to 100mm long. and Vietnam.
Fur colour yellowish-brown or tan to grey-brown, Status Critically Endangered. Has very limited range,
with grizzled effect caused by banding on individual is rarely detected on camera traps and appears to
hairs; black lines on forehead; white spot on knee have declined dramatically due to excessive hunting.

Family BOVIDAE CATTLE, BUFFALO, ANTELOPES, GOATS AND SHEEP

All of the species in this family have horns that are never shed and that continue to grow as the animal gets
older. Both males and females have horns in all species in the region, though males are usually larger, with
larger horns.
Most species in this group have suffered substantial population declines resulting from loss of habitat and
hunting. At least one form, the Kouprey, Bos sauveli, is probably extinct, although there is some controversy
as to whether it was a distinct species or a hybrid between Banteng, Bos javanicus, and domestic cattle.
An additional species, Spiral-horned Ox, Pseudonovibos spiralis, was once described based on horns from
Vietnam. However, genetic analysis and closer study of the horns indicate that this was based on artificially
altered horns of domestic cattle, and no such animal ever existed. A number of strains of domestic cattle
may resemble some of the wild cattle, in some cases having been interbred with Banteng, but can usually be
distinguished by their relatively tame behaviour, and by being near human settlements.

BANTENG Banteng, except by less muscular backs in males

Bos javanicus PLATE 60 and tamer behaviour.
Measurements SH 1,500–1,700, HB 1,900– Ecology and habitat Mainly nocturnal, probably
2,250, T 650–700. Wt 600–800kg partly as a result of hunting pressure. Found in a
Identification Adult males dark chestnut-brown variety of forest types, including dry semi-open
in most parts of mainland; adult females and forest, lowland forest and swamp forest. Often along
young males pale brown to bright rufous-brown. In banks of major rivers. Feeds mainly in clearings
S Thailand (as well as Java and Borneo), males are and open areas, where diet is mainly grasses, with
darker, often jet-black, both sexes with distinctive some herbaceous and low woody vegetation. Visits
white band around muzzle, white buttocks and natural mineral sources. Now found mainly in fairly
white ‘stockings’ on lower part of the legs. Adult small groups, but formerly herds were much larger.
females smaller and more slender than adult males, Domestic forms are kept in some areas.
with smaller horns. Horns of male narrow, curving Distribution SE Asia: Myanmar, Thailand, Laos,
outwards and forwards, then inwards at tip; female Vietnam and Cambodia. Formerly Peninsular
horns closer together and more upright. Horny patch Malaysia. Also Java, Borneo. May have formerly
of thick skin between horns. Footprints relatively occurred in Yunnan, China.
large, with curved hoof (Fig. 64b). Similar species Status Endangered. Wild populations have declined
Gaur, B. gaurus, is larger and more muscular, without throughout range, due to loss of habitat and hunting.
white on rump or muzzle. Most domestic cattle lack Also some risk from hybridisation with and disease
white on the buttocks, but some can be very hard to transmission from domestic cattle. Uncertain
distinguish, in particular those descended from wild whether still persists in Laos and Vietnam.

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 GAUR rest of back; females and young greyish, with paler
Bos gaurus PLATE 60 underparts. Male with very large flap of skin (dewlap)
Measurements SH 1,700–1,850, HB 2,500– under neck, sometimes nearly reaching the ground.
3,000, T 700–1,050. Wt 650–900kg Horns of male curve outwards and forwards, with
Identification Large and muscular, adult male with a tendency for tip to split; horns of female smaller,
high ridge of muscles on back; relatively short neck; lyre-shaped, with a corkscrew-like twist. Similar
variably developed flap of skin under neck (dewlap). species Banteng, B. javanicus, has white rump,
Body largely black; legs, from knees down, greyish- differently shaped horns; Gaur, B. gaurus, has pale-
white or golden-yellow. Newborn young brown with coloured hollow forehead shield, smaller dewlap,
black stripe on back, turns to black by 4–5 months. larger back muscles, horns curved backwards and
In Indian populations, and possibly some in region, upwards.
young are golden-brown, turning to reddish-brown Ecology and habitat Was found in mixed savanna,
in young males and adult females. Forehead of open grasslands and patches of dense forest.
male raised up in pale yellowish-brown ridge that Foraged mainly in grasslands, but sheltered in
is concave in profile. Horns relatively short, rounded forests.
in cross-section, relatively far apart on head, curved Distribution SE Asia: formerly found in SE Thailand,
outwards and upwards. Domestic descendents of S Laos, Cambodia and S Vietnam.
Gaur, called Mithan or Gayal, are sometimes kept Status Critically Endangered, possibly Extinct. No
in region, but are generally much less muscular. confirmed records of the species since c.1969.
Footprints similar to those of Banteng, B. javanicus Apparently hunted to extinction.
(Fig. 64b), but average somewhat larger. Taxonomic
notes Sometimes called Bos frontalis, which is now WATER BUFFALO
considered just the name of the domestic form. Bubalus arnee PLATE 60
Similar species Banteng, B. javanicus, has white Measurements SH 1,600–1,900, HB 2,400–
rump patch, usually much browner, especially in 2,800, T 600–850. Wt 800–1,200kg
females, smaller horns, less muscular. Identification Entirely greyish, except for a pale
Ecology and habitat Found principally in forested V-shape usually present on the underside of the
areas; may enter grasslands to feed, but returns neck. Horns long, swept out to sides and curved;
to forest for shelter. Feeds on various grasses, somewhat triangular in cross-section. Footprints
including bamboo, as well as leaves. Mainly very large and broad, often rather splayed in soft
nocturnal in most of region, but also active during ground (Fig. 64a). Taxonomic notes Sometimes
the day in areas where protected from disturbance called Bubalus bubalis, which is the name of the
by humans. Regularly visits natural mineral sources domestic form. Similar species Domestic Water
and salt licks. Buffalo looks similar but is generally smaller, less
Distribution SE Asia: Myanmar, Thailand, Laos, muscular, relatively tame. Gaur, Bos gaurus, has
Vietnam, Cambodia and Peninsular Malaysia. Also white lower legs, horns shorter and rounded in
India, Nepal, Bangladesh and extreme S China. cross-section.
Status Vulnerable, but populations in SE Asia Ecology and habitat Wild Water Buffalo is
declining much faster than those in India, due to associated with grassland on alluvial soil.
loss of habitat and excessive hunting. Also at risk Distribution SE Asia: N Myanmar, W Thailand, E
from diseases of domestic cattle. Cambodia. Probably no wild animals still occur in Laos
or Vietnam. Also NE India, Nepal and Bangladesh.
KOUPREY Formerly much more widespread in region. Domestic
Bos sauveli PLATE 60 animals found in many parts of the world, with
Measurements SH 1,700–1,900, HB 2,100– escaped wild populations in several areas.
2,200, T 1,000–1,100. Wt 700–900kg Status Endangered. Wild populations severely
Identification Adult males largely dark brown to reduced by conversion of habitat to rice farming and
black; lower legs white or greyish; area on forehead by illegal hunting. Also at risk from hybridisation with
between horns black, rump dark, same colour as domestic forms, and diseases from domestic cattle.

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 a b

Female above
Male below

0 100

Fig. 64 Footprints of Bubalus arnee (a), Bos javanicus (b), and Capricornis sumatraensis (c).For all species, the toes
are more spread out in soft ground, and the hind toes may be visible.

TAKIN reach salt licks. Often in medium-sized herds of

Budorcas taxicolor PLATE 61 20–30 animals, but sometimes seen in herds of up
Measurements SH 700–1,400, HB 1,000–2,400, to 300 animals. Old bulls may be solitary.
T 70–120. Wt 150–400kg Distribution SE Asia: N Myanmar. Also NE India
Identification Thick-bodied animal with short, thick (Assam), S China including Tibet.
legs, arched back, thick neck, face with distinctive Status Vulnerable. Rare in Myanmar, where
outward-curved profile. Horns rise near middle threatened by illegal hunting and trapping.
of forehead, turn outwards, then curve abruptly
backwards. Females and young have straighter horns SAOLA
than males. Colour varies from dark greyish-black Pseudoryx nghetinhensis PLATE 61
with white patches through dark brown to golden- Measurements SH 800–900, HB 1,500.
yellow brown. Females and young usually greyer than Wt 85–100kg
males, and often with dark line down back. Hair long Identification Slender, antelope-like animal with
and shaggy. Similar species Gorals, Naemorhedus long, smooth horns slightly curved backwards. Fur
spp., and serows, Capricornis spp., have thinner legs, dark grey-brown to reddish-brown. Distinctive face
different-shaped head, straighter horns. pattern, with white stripe above eye and several white
Ecology and habitat Found in hill forests and spots and stripes on chin and muzzle. Legs generally
thickets in montane areas at altitudes of 1,000– darker than body, with white marks on ankle. Horns of
4,000m. In summer may be near tree line, but young animals short and goat-like. Similar species
comes lower in winter. Feeds by browsing on bushes Serows, Capricornis spp., have shorter, more curved
and grazing on grass. May travel long distances to horns and lack white markings on head.

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 Ecology and habitat Wet evergreen hill forests, to fur on flanks, with pale lower legs; Chinese Goral,
apparently restricted to large contiguous blocks of Naemorhedus griseus, is much smaller, with smaller
forest. head, thinner neck, paler body colour.
Distribution SE Asia: Laos and Vietnam. Ecology and habitat Thick forests, usually on fairly
Status Critically Endangered. Population has been steep terrain, including limestone outcrops and hill
declining dramatically; likely to be fewer than 250 forest. Usually solitary, except female with young.
mature animals left, in limited range in Annamite Distribution SE Asia: S Thailand and Peninsular
Mountains. Main threats are illegal hunting and Malaysia. Also Sumatra.
trapping, as well as forest loss. Status Vulnerable. Main threats are loss of forest
habitat, but also illegal hunting and trapping.
BLUE SHEEP
Pseudois nayaur PLATE 61 INDOCHINESE SEROW
Measurements SH 800–900, HB 1,200–1,650, Capricornis milneedwardsi PLATE 62
T 100–200. Wt 55–70kg Measurements SH 850–940, HB 1,400–1,550,
Identification Adult male light greyish-brown to T 110–160kg
blue-grey, with black line along flanks and along Identification Similar to Southern Serow, differing
front edge of both back and front legs; black chest; mainly in colour. Hairs on most of body black with
white belly; white on back sides of all legs with white extensive white base showing through, giving overall
marks on ankles; white rump and tail. Males have greyish colour; black stripe on back; head browner;
long, smooth horns. Females similar but without underside often paler; white or pale brown hairs on
black markings and with shorter horns. Similar throat, often forming pale patch; legs black on upper
species Serows, Capricornis spp. have shorter, half, contrasting reddish or whitish on lower half; long
narrower horns, longer legs, thick mane, occur in mane with long white hairs, sometimes extensively
more forested habitats. mixed with black. Similar species Southern Serow,
Ecology and habitat Montane habitats, often above C. sumatraensis, is blacker, without white hair base
tree line, in bare rocky areas and open grasslands at on most of body, black legs; could possibly hybridise
2,500–6,500m altitude. in areas of overlap in peninsular Thailand.
Distribution SE Asia: extreme N Myanmar. Also Ecology and habitat Occurs mainly in hill forest
China, Himalayas (N India, Nepal, Bhutan and N and rugged terrain such as limestone karst, though
Pakistan). sometimes in lowland forest. Tolerant of some forest
Status Least Concern globally, but very rare and degradation, but not clearing for agriculture.
probably threatened by hunting in Myanmar. Distribution SE Asia: Myanmar, Thailand, Laos,
Vietnam and Cambodia. Also S China.
SOUTHERN SEROW Status Near Threatened. Populations fragmented in
Capricornis sumatraensis PLATE 62 many areas, owing to loss of habitat and intervening
Measurements SH 850–940, HB 1,400–1,550, forest clearance, with excessive hunting in many
T 110–160. Wt 85–140kg areas.
Identification Long-legged, relatively short-bodied,
goat-like animal with short, nearly straight horns RED SEROW
with marked striations (circular ridges). Colour Capricornis rubidus PLATE 62
generally black, greyer in younger animals, with Measurements SH 850–950, HB 1,400–1,550,
black legs. Black crest of hairs on back, adults T 110–160kg
with well-developed mane of long hairs on back of Identification Shape and size similar to other
neck, varying from dirty-white to buff or reddish, serows, but overall fur colour red-brown, hairs with
sometimes black mixed with a few white hairs. Has black base; usually with dark stripe down back;
large, open gland in front of eye. Footprints more mane relatively short, dark reddish, blending with
rounded than those of deer, but narrower than those back. Often with white patch on throat and white
of pigs (Fig. 64c). Similar species Indochinese belly. Some individuals are darker, black with
Serow, C. milneedwardsi, has extensive white base reddish tone, and may sometimes hybridise with

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 Chinese Serow, C. milneedwardsi. Similar species at 1,000–4,000m altitude, often near cliffs. Feeds
Red Goral, Naemorhedus baileyi, is smaller, without on leaves, grasses and twigs. Usually in small
distinct mane, fur smoother and shorter, shorter groups of 5–10 animals.
legs, horns smaller and more slender. Distribution SE Asia: N Myanmar and N Thailand.
Ecology and habitat Tropical and sub-tropical Also S China.
forest, mainly in hills. Status Vulnerable. Populations have declined
Distribution SE Asia: N Myanmar. Also NE India. substantially due to hunting and habitat loss.
Status Near Threatened. Population declining
throughout range due to loss of habitat and RED GORAL
excessive hunting. Naemorhedus baileyi PLATE 62
Measurements SH 550–600, HB 900–1,050,
CHINESE GORAL T 100–150. Wt 20–30kg
Naemorhedus griseus PLATE 62 Identification Upperparts fairly uniform reddish-
Measurements SH 500–700, HB 800–1,200, brown, without white or black flecking; narrow dark
T 80–200. Wt 20–30kg stripe down middle of back; legs coloured as back;
Identification Relatively small, goat-like animal with underparts paler, chin and throat with pale patch.
short fur and short, slender horns. Fur light greyish- Similar species Red Serow, Capricornis rubidus,
brown to brown, with darker brown stripe down is substantially larger with longer legs, bigger head
middle of back and darker brown forehead and ridge and neck, longer, coarser fur and distinct mane;
of nose; legs paler, golden-brown with dark wedge Chinese Goral, N. griseus, is greyer.
on front; underside pale greyish-brown; throat white Ecology and habitat Mainly in coniferous hill
to orange-white; tail mostly dark brown. Similar forests with rocky outcrops.
species Serows, Capricornis spp., are much larger, Distribution SE Asia: N. Myanmar. Also S China and
with darker fur with long coarse hairs, thicker neck NE India (Assam).
and larger head. Status Vulnerable. Population is declining, owing to
Ecology and habitat Generally in steep, hilly terrain illegal hunting.

Order RODENTIA
Squirrels, rats, mice, voles, kha-nyou and porcupines
Rodents can be recognised by their dentition, with large, curved, chiselling incisors in both the upper and
lower jaws; no canines and a wide diastema (toothless gap) before the cheek teeth (Fig. 65). All rodents in
South-east Asia have five well-developed toes on the hind foot, but unlike insectivores and treeshrews, only
four long, clawed digits on each front foot. The thumb is short, with a nail instead of a claw, and does not
show in the footprint. Footprints of porcupines can be distinguished by size; those of squirrels differ from
rats in having the front toes closer together and relatively larger footpads (Fig. 67). Although many species
of rat have distinctive patterns of footpads, these can rarely be distinguished clearly enough to separate
species based on prints.

Family SCIURIDAE SQUIRRELS

The squirrel family includes both regular squirrels and flying or gliding squirrels. Formerly, the flying squirrels
were thought to belong in their own family because of their distinctive shape and morphology, but recent
genetic research indicates that they evolved as an offshoot of one branch of tree squirrels. The remaining
squirrels in the region are allocated to two separate subfamilies. Ratufinae includes the two species of giant
squirrel, while Callosciurinae includes several genera that are mainly arboreal, although a few species spend
much of their time on the ground. Squirrel skulls can be distinguished from those of murid rodents by being
generally broader, with well-developed supraorbital processes (Fig. 66), and four or five cheek teeth (Fig. 65).

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 a

Fig. 65 (above) Lateral view of skull of Plantain Squirrel,

Callosciurus notatus.

Fig. 66 (right) Dorsal views of skulls of a squirrel, Belomys

pearsoni (a) and a rat, Rattus andamanensis (b). 0 20

a b

F H

c F
H

d e

F H
F H F H

0 50

Fig. 67 Footprints of selected rodents: Trichys fasciculata (a), Hystrix brachyura (b), Ratufa affinis (c),
Lariscus insignis (d) and a typical rat, Rattus sp. (e). F = front foot, H = hind foot.

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 Subfamilies RATUFINAE, CALLOSCIURINAE GIANT, TREE AND GROUND SQUIRRELS
Squirrels are among the small mammals most frequently seen by people, as they are active during the
day; some species regularly occur in gardens, orchards and other disturbed areas, although others are
restricted to more natural forests. Most squirrels have distinctive coloration, but identification is complicated
by extensive variation in colour within some species, particularly in Callosciurus. Much of this variation is
geographical, such that populations of the same species in different areas can be very different colours. In
addition, some species, particularly the Variable Squirrel, Callosciurus finlaysonii, may have different colour
morphs occurring together in the same area. In some cases these are thought to represent seasonal changes
in colour pattern of the same individuals, while others may be due to small genetic differences within
interbreeding populations – further research is needed to determine which type of variation is occurring in
each area. This variation complicates understanding of speciation in these squirrels, and in several groups
the currently recognised species limits are still very provisional. Genetic analyses of squirrels from a wide
geographic area are required to resolve these uncertainties.
Identifying squirrels in the field is usually easiest if they can be observed with binoculars, so that the
colours and shape can be seen more clearly. Some of the key patterns to look for include the presence or
absence of stripes on the belly, sides or back, and the colour of the belly and back. Conditions are rarely
ideal, however, and many ground squirrels are glimpsed only briefly in dim light, while tree squirrels are often
obscured by leaves or silhouetted against the sky. Some species can be trapped fairly easily in cage traps for
closer study, though traps may need to be placed in trees to catch the more arboreal species.
Treeshrews, Tupaia spp., could be mistaken for squirrels but they generally have more pointed muzzles
(although some squirrels also have pointed muzzles), a pale mark on their shoulder and very different
dentition, without the enlarged yellowish incisors typical of rodents (Fig. 23, page 210).

Genus Ratufa These are among the largest have similar colour patterns, but are substantially
squirrels in the world, with a large body and a long, smaller and usually have more extensive white on
bushy tail. They are largely confined to tall forest, head, tail or feet.
where they remain mainly in the canopy. They have Ecology and habitat Primarily in canopy of tall
a distinctive posture when sitting or feeding, the tail forest, rarely descending to the ground. Feeds
often hanging down vertically. Two species occur in mainly on fruits and nuts, but also some insects and
the region, one of them widespread, the other mostly animal matter.
confined to peninsular Thailand and Malaysia. Distribution SE Asia: Myanmar, Thailand, Laos,
Vietnam, Cambodia and Peninsular Malaysia. Also
BLACK GIANT SQUIRREL E Nepal, Assam, S China, Sumatra, Java and Bali.
Ratufa bicolor PLATE 63 Status Near Threatened; although widespread,
Measurements HB 370–405, T 425–500, declining rapidly in many areas due to hunting as
HF 84–91, E 30–38 well as loss of tall forest.
Identification Largest tree squirrel in the region.
Upperparts typically black in fresh pelage, but CREAM-COLOURED GIANT SQUIRREL
some individuals may be dark brown or reddish- Ratufa affinis PLATE 63
tinged black; underparts vary from pale buff Measurements HB 310–380, T 370–440,
to orange. Cheeks white with dark mark like a HF 66–86. Wt 875–1,500g
moustache. Some geographic variation in extent Identification A very large tree squirrel with
of white on forelegs and presence or absence of orange-brown to pale buff upperparts, buffy to white
ear tufts. Similar species Cream-coloured Giant underparts. Animals in the northern peninsula are
Squirrel, R. affinis, is much paler, with little contrast generally darker orange, while those in the southern
between upper and underparts; Prevost’s Squirrel, peninsula are paler. Outside the region, in Borneo,
Callosciurus prevostii, is smaller, with distinct white individuals may be much darker, closely resembling
stripe on flank separating black back from orange the Black Giant Squirrel, R. bicolor. When moving
belly. Some Variable Squirrels, C. finalysonii, may quickly through the tree canopy, the tail is held out

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 horizontally, but when sitting on a branch it hangs stripe; no other Callosciurus in its range has similar
vertically. Voice Loud call is a short, harsh chatter, colour patterns.
often audible for several hundred metres. It also Ecology and habitat Diurnal, most active in the
has a distinctive soft call, a series of ‘hgip ... hgip early morning and late afternoon. Usually arboreal,
…’, audible only when close to the animal. Similar descending to the ground only to cross gaps in the
species Pale forms of Variable Squirrel, Callosciurus tree canopy. Diet includes fruits, especially those
finlaysonii, are smaller and not found in the same with a sweet or oily flesh, and insects, notably ants,
geographic areas. termites and beetle larvae, which are gnawed out of
Ecology and habitat Diurnal, usually emerging dying wood. Occurs in tall and secondary forests.
from the nest well after dawn and retiring for the Enters gardens and plantations from adjacent forest
night before dusk. Nest is a neat, globular array to feed on fruits.
of twigs, usually built in the crown of a tall tree. Distribution SE Asia: peninsular Thailand and
Mostly active in tall trees, descending to the Malaysia. Also Sumatra, Borneo, Sulawesi and
ground only to cross gaps in the tree canopy. Diet smaller Indonesian islands.
mainly seeds with some leaves and bark. Occurs Status Least Concern, but likely to be declining due
in lowland and hill forests with tall trees, including to loss of forest.
some selectively logged forests, but rarely enters
plantations. PLANTAIN SQUIRREL
Distribution SE Asia: peninsular Thailand, Malaysia Callosciurus notatus PLATE 63
and Singapore. Also Sumatra, Borneo and some Measurements HB 175–225, T 160–210,
smaller Indonesian islands. HF 42–52. Wt 150–280g
Status Near Threatened due to destruction of tall Identification Upperparts brownish-agouti. Under-
forests, as well as hunting. parts reddish-orange, varying from very pale to fairly
dark, sometimes with a greyish wash. Distinctive
Genus Callosciurus Moderately large tree stripes on flanks, with black stripe next to belly, buff
squirrels; many species are very colourful with stripe above it. Tail tip usually orange, though orange
distinctive patterns. There is extensive geographic restricted to a few hairs in some individuals. Voice
variation in the colour patterns of many species, A shrill, scolding chatter as well as sharp, bird-
with some exhibiting very different patterns across like chirp, repeated continuously for long periods.
their range. The taxonomic limits for some of these Similar species Sunda Black-banded Squirrel,
species are not well known and may not be fully C. nigrovittatus, also has black and buff stripes on
resolved until genetic studies have been done. sides, but has grey underparts and no reddish on tip
From the perspective of field identification it can be of tail; Pallas’s Squirrel, C. erythraeus, lacks black
very helpful to check the species distribution maps, and buff stripes on sides.
because there is only limited overlap among the Ecology and habitat Diurnal, most active early
more variable species. morning and late afternoon. Travels and feeds
mainly in small trees. Diet includes a wide variety of
PREVOST’S SQUIRREL fruits and insects, mostly ants. Commonly found in
Callosciurus prevostii PLATE 63 gardens, plantations and secondary forests, and can
Measurements HB 200–270, T 200–270, live and breed entirely in monoculture plantations.
HF 45–60. Wt 250–500g Less frequently found in the interior of undisturbed
Identification Upperparts largely black with grey on tall forests, but usually common in coastal and
side of face; belly dark reddish-orange; broad white swamp forests.
stripe on flanks. Outside region, for example in Distribution SE Asia: peninsular Thailand and
Borneo, colour varies considerably, with white stripe Malaysia, and Singapore. Also Sumatra, Java,
thin or lacking in some forms, and grey or brown Borneo and many smaller offshore islands around
upperparts in others. Voice A loud, harsh chatter. them.
Similar species Black Giant Squirrel, Ratufa bicolor, Status Least Concern. One of the most common
is larger, with paler underparts and no distinct flank squirrels of disturbed areas in its range.

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 SUNDA BLACK-BANDED SQUIRREL larger, lacks pale patch on hips, has dark reddish
Callosciurus nigrovittatus PLATE 63 underparts with (usually) agouti stripe in middle of
Measurements HB 170–240, T 145–230, belly; Grey-bellied Squirrel, C. caniceps, is similar to
HF 40–50 C. p. stevensi but lacks pale hip patch and does not
Identification Upperparts speckled brownish- overlap in range.
agouti; buff stripe above black stripe on sides of Ecology and habitat Diurnal and arboreal.
body; underparts uniform grey. Tail as back colour, Broadleaved and coniferous evergreen forest,
without reddish tip. Sides of head and upper chin including disturbed areas and secondary growth.
reddish-brown. Voice A rattling chatter or chuckle; Distribution SE Asia: NW and C Myanmar. Also
also a sharp, repeated bird-like chirp. Similar China (Xijang), Nepal, Bangladesh and E India.
species Plantain Squirrel, C. notatus, has sides of Status Least Concern.
head same colour as back, reddish belly and reddish
tip to tail. PHAYRE’S SQUIRREL
Ecology and habitat Diurnal and arboreal, mainly Callosciurus phayrei PLATE 63
in canopy and middle storey of both primary and Measurements HB 210–250, T 230–270
secondary lowland forest. Identification Upperparts mottled grey and brown
Distribution SE Asia: peninsular Thailand and agouti; underparts pale buff to orange-brown,
Malaysia. Also Sumatra, Java, Borneo and many usually separated from back colour by a distinct,
smaller offshore islands around them. wide black stripe. However, stripe may be indistinct
Status Near Threatened; population declining due or missing in some individuals. Tail agouti with
to loss of natural forest. distinct black tip. Similar species Some colour
forms of Pallas’s Squirrel, C. erythraeus, may look
IRRAWADDY SQUIRREL similar, but lack a black stripe on the flank (or have
Callosciurus pygerythrus PLATE 63 at most a very thin stripe).
Measurements HB 170–200, T 170–200, Ecology and habitat Diurnal and arboreal. Primary
HF 41–45 and secondary broadleaved forest.
Identification Upperparts agouti, with overall colour Distribution SE Asia: Myanmar, between lower
varying from slightly reddish to olive-brown to grey. Sittang and Salween Rivers. Also adjacent parts of
Underparts vary geographically from pale buff to China (SW Yunnan).
grey to orange. Within populations, fur colour varies Status Least Concern.
seasonally, being brighter in wet season than dry
season. In wet season, has pale patches on hips, PALLAS’S SQUIRREL
but these are often lacking in dry season. Major Callosciurus erythraeus PLATE 64
racial variation within Myanmar is as follows: C. p. Measurements HB 200–260, T 190–250,
pygerythrus (W of Sittang, E of lower Irrawaddy): HF 44–55, E 18–23
underparts reddish, more buff near front, hip patch Identification Considerable geographic variation in
buff or lacking; C. p. janetta (W of Irrawaddy, E of colour pattern; may include more than one species,
lower Chindwin): underparts light buff, hip patch but appropriate species limits are uncertain. In most
white; C. p. owensi (W of Irrawaddy, E of upper forms, upperparts are speckled olive-brown (agouti),
Chindwin): more reddish upperparts and underparts, sometimes with a black stripe down the back, and
hip patch buff; C. p. mearsi (W of lower Chindwin): underparts are reddish, though this varies from
underparts buff, hip patch white, buff eye-ring; dark maroon to red-brown. In many populations
C. p. stevensi (W of upper Chindwin): underparts there is a stripe of agouti down the middle of the
grey, upperparts reddish, hip-patch buff. Similar belly, separating the reddish belly into two bands.
species Inornate Squirrel, C. inornatus, never The major variations within the region are as
has pale patches on hips and does not overlap in follows: C. e. erythraeus group (Peninsular Malaysia,
range; Phayre’s Squirrel, C. phayrei, has black stripe W Thailand, S and NW Myanmar, NE India, N Vietnam
on flank and usually darker orange underparts; and N Laos): upperparts agouti, feet and tail as back,
Pallas’s Squirrel, C. erythraeus, tends to be slightly underparts reddish, with an agouti stripe up the

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 middle – in some parts of S Myanmar underparts individuals through the year, is required to clarify
may be all agouti without any red; in W Thailand these patterns. Plate shows some of the major colour
and S Myanmar, lower back has a distinct broad variations, but many other combinations occur. The
black stripe (sometimes considered a separate major population variations are described here as
subspecies, C. e. atrodorsalis). C. e. flavimanus subspecies, but there is intergradation among these,
group (S Vietnam, S Laos and Cambodia): upperparts especially where ranges overlap, and genetic studies
agouti, feet pale, often yellowish-orange, underparts have not yet been done to determine how distinct
vary from cream to rich reddish-brown. C. e. sladeni these populations really are. The mainland variations
group (Myanmar between Chindwin and Irrawaddy are: C. f. sinistralis (NW Thailand): upperparts agouti,
Rivers): a highly variable form with some populations often with strong reddish cast, underparts reddish-
similar to typical C. erythraeus, but others may be orange, tail deep red-brown with white ring around
greyish-white above, pale orange below and on base. C. f. menamicus (E part of N Thailand and N
the feet; medium grey mixed with agouti above, Laos): varies from largely reddish-brown with agouti
cream below and on the feet; or even completely flanks to reddish-brown upperparts with white belly,
dark rufous-red all over and only slightly paler on feet and head, and many intermediates in between;
the underparts. Taxomonic notes C. flavimanus is tail varies from red to grey with a buff tip or white with
sometimes considered a separate species. Similar grey on top. C. f. bocourti (E C Thailand, from S to N):
species Variable Squirrel, C. finlaysonii, never has varies from completely creamy white to having black
the combination of agouti upperparts and tail with back, with white head, underparts, legs and tail; tail
reddish underparts; however, some populations sometimes black above, white below. C. f. boonsongi
can resemble some of the variations of the C. e. (NE Thailand): usually very dark grey to black all
sladeni group, though typically not those in the same over, sometimes with white or red edges to ears;
geographic areas; Phayre’s Squirrel, C. phayrei, some individuals have extensive white on underside,
has a black band on the sides; Plaintain Squirrel, feet and face. C. f. annellatus (SE Thailand, C and
C. notatus, has a black and buff band on the flanks. E Cambodia, and S Laos W of Mekong): uniform
Ecology and habitat Diurnal and arboreal. In deep reddish-brown except for pale ring around
southern part of range mainly confined to hill base of tail. C. f. williamsoni (E part of Mekong plain
forests, but elsewhere occurs in a variety of habitats, in Laos and N Cambodia): red to orange above,
including primary and secondary forests as well as clearly separated from dark chestnut below; red tail.
disturbed areas and plantations. C. f. cinnamomeus (SE Thailand and SW Cambodia):
Distribution SE Asia: Myanmar, Thailand, Laos, all red, or largely red with some agouti on head and
Vietnam, Cambodia and Peninsular Malaysia (where limbs. C. f. floweri (SC Thailand, near Bangkok):
confined to central mountain range above 1,000m olive-brown upperparts including head with white
elevation). Also Bangladesh, NE India, and S and marks around eye, sometimes forming a complete
E China. ring; white underparts. C. f. nox (SE Thailand): all
Status Least Concern. black. C. f. ferrugineus (E Myanmar): typically all
reddish-brown. In addition, there are distinctive
VARIABLE SQUIRREL populations on several of the small islands off the
Callosciurus finlaysonii PLATE 65 coasts of Thailand and S Vietnam. C. f. finlaysonii
Measurements HB 210–220, T 220–240, (Si Chang island, Thailand): all yellowish-white.
HF 45–50, E 19–23 C. f. folletti (Phai island, Thailand): greyish-white
Identification A highly variable squirrel with no to beige upperparts; creamy underparts often with
single combination of features that can diagnose an orange wash; light, buffy tail. C. f. trotteri (Lan
the species. There is extensive variation both island, Thailand): mid-grey upperparts with lower
among geographic areas and within populations. back darker; pale grey belly; white tail. C. f. frandseni
Some of this variation appears to be seasonal, but (Chang island, Thailand): dark reddish-brown above
this has been little studied; it appears that some mixed with brownish-olive (agouti) on head, neck,
squirrels may change from maroon-red to white at flanks and upper parts of legs; underparts buffy-
different times of year. Further field study, following orange. C. f. albivexilli (Kut island, Thailand): all black

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 except for a white tip to the tail. C. f. harmandi (Phu also a bird-like cheep repeated for long periods.
Quoc island, Vietnam): upperparts dark reddish- Similar species Sunda Black-banded Squirrel,
brown to grey-brown, underparts orange-red; tail C. nigrovittatus, has black and buff bands on flanks
greyish-white. C. f. germaini (Con Dao Is, Vietnam): without orange on upperparts; Pallas’s Squirrel,
all black. Similar species Some populations of C. erythraeus, has underparts all reddish, or reddish
Pallas’s Squirrel, C. erythraeus, may be similar to with agouti in the middle, lacks golden-orange tinge
some populations of C. finlaysonii, but can usually be on back, and has a pale tail tip; Inornate Squirrel,
separated by details of the colour pattern, especially C. inornatus, is very similar but lacks black tail tip
an agouti banded tail. and does not overlap in range.
Ecology and habitat Diurnal and arboreal, found Ecology and habitat Diurnal and primarily arboreal.
in a variety of habitats including dense forest, open Found in a wide range of forest types, including
woods and coconut plantations. plantations, cultivated areas, second growth and
Distribution SE Asia: S Vietnam, Thailand, gardens, as well as intact forest. May range up to
Cambodia, Laos and part of E Myanmar. Introduced 1,500m in hills, but mainly in lowlands.
in Singapore. Distribution SE Asia: N Laos W of Mekong River,
Status Least Concern; still common in many areas. SE Myanmar, C and W Thailand, and Peninsular
Malaysia.
STRIPE-BELLIED SQUIRREL Status Least Concern.
Callosciurus quinquestriatus PLATE 64
Measurements HB 210–240, T 160, HF 40 INORNATE SQUIRREL
Identification Upperparts speckled brown (agouti), Callosciurus inornatus PLATE 64
often with a strong reddish tinge. Underparts white, Measurements HB 220–230, T 200–240
with a black stripe on each flank, and a black stripe Identification Upperparts mottled brown (agouti),
up the middle of the belly. Tail lightly banded with a sometimes with a reddish tinge, underparts uniform
black tip. Similar species No other species has a grey. Similar species Grey-bellied Squirrel,
white belly with black stripes on each flank and in C. caniceps, has black tail tip, and does not overlap
middle of belly. in range, occurring only to west of Mekong River.
Ecology and habitat Diurnal and arboreal. Ecology and habitat Diurnal and arboreal.
Evergreen hill and montane forest. Evergreen forests including conifer hill forest, as well
Distribution SE Asia: NE Myanmar, E of upper as degraded forest and scrub. Found from lowlands
Irrawaddy River. Also adjacent parts of China up to 2,000m altitude.
(Yunnan). Distribution SE Asia: N and C Laos, and N Vietnam,
Status Near Threatened due to limited range and E of Mekong River. Also S China (Yunnan).
loss of forest. Status Least Concern.

GREY-BELLIED SQUIRREL Genus Tamiops A group of small, striped squirrels,

Callosciurus caniceps PLATE 64 distinguished from most other squirrels by their
Measurements HB 210–240, T 185–245, arboreal habits and pattern of dark and pale stripes
HF 49–54, E 19–23 on the back. Currently, four species are recognised,
Identification Upperparts vary from grey-agouti differing in size, in the brightness and width of
to golden-brown; underparts usually silvery-grey, the stripes and in belly colour. However, there is
but may have reddish wash on part of underparts; also moderate variation within each species, and
tip of tail black in most parts of Thailand, but grey further study, preferably involving genetic analysis,
in Peninsular Malaysia. In northern parts of range, is required to determine whether these are actually
back is a very bright orange in the dry season, but appropriate species limits. Field identification is
greyish-agouti in the wet season. In Peninsular aided by the fact that there appears to be relatively
Malaysia, back is mottled orange-brown year little overlap in distribution among the species,
round. Some populations have patches of rufous although parts of Vietnam and Laos may have more
on underparts. Voice A loud, harsh chuckle; than one species.

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 WESTERN STRIPED SQUIRREL orange. Similar species Western Striped Squirrel,
Tamiops mcclellandii PLATE 66 T. mcclellandii, has outer pale stripes wider and
Measurements HB 110–125, T 110–140, more conspicuous than inner pair; Eastern Striped
HF 27–32, E 15–22 Squirrel, T. maritimus, is larger, has greyish-white
Identification Upperparts mottled grey and brown, underparts, outer stripes brighter than inner and not
with five dark stripes and four pale stripes down connected to cheek stripe.
middle of back. Central back stripe usually blackest, Ecology and habitat Diurnal and arboreal. Found in
with others varying from nearly black to only slightly a variety of habitats with trees, from forest to scrub,
darker than rest of body. Pale stripes vary from white plantations and gardens.
to buff with outer pair typically much wider, paler and Distribution SE Asia: E Thailand, S Laos, Cambodia
brighter than inner pair. Outer stripe contiguous with and S Vietnam.
white stripe on cheeks. Underparts usually buff to Status Least Concern; common in many areas.
dull orange, with grey base to fur. Conspicuous white
tufts on ears. There is some geographic variation, SWINHOE’S STRIPED SQUIRREL
with pale stripes narrower and less conspicuous Tamiops swinhoei PLATE 66
in some northern populations. Voice A bird-like Measurements HB 120–155, T 100–120,
chirp repeated for long periods at intervals of about HF 30–38
one second; also a descending trill of shrill chirps. Identification Upperparts brownish, with five dark
Similar species Cambodian Striped Squirrel, stripes separated by four pale stripes on back. Dark
T. rodolphii, has central stripes similar in width and stripes very black in fresh pelage; outer stripes
brightness to outer stripes, underparts often more become browner in winter. Inner pale stripes relatively
orange; other Tamiops usually have cheek stripes dull, only slightly paler than nape and head; outer
separate from side stripe. pale stripes much brighter, varying from orange-buff
Ecology and habitat Diurnal and arboreal. Uses to white. Outer pale stripes do not connect with cheek
a variety of forest types, from primary hill forest stripes. Pelage relatively thick and bushy. Similar
to gardens and fruit trees. Diet consists of fruits, species Eastern Striped Squirrel, T. maritimus, is
vegetable matter and some insects. Uses holes very similar, but dark stripes away from centre tend
in trees for shelter. Primarily found above 700m to be browner and less distinct, pelage is thinner; has
altitude in Thailand and Malaysia, but also in only limited overlap in distribution.
lowlands farther north. Ecology and habitat Diurnal and arboreal. Forest,
Distribution SE Asia: N Myanmar, N and including disturbed areas with trees, primarily
W Thailand, N Laos (W of Mekong River) and south in montane and hill areas at elevations of 1000–
through Peninsular Malaysia. 3900m.
Status Least Concern. Distribution SE Asia: NE Myanmar and NW Vietnam.
Also SW China.
CAMBODIAN STRIPED SQUIRREL Status Least Concern.
Tamiops rodolphii PLATE 66
Measurements HB 105–125, T 115–130, EASTERN STRIPED SQUIRREL
HF 25–30 Tamiops maritimus PLATE 66
Identification Upperparts mottled grey-brown, Measurements HB 100–145, T 95–120, HF 30–35
with five dark stripes and four pale stripes down Identification Upperparts brownish with five dark
middle of back. Pale stripes all similar in width and stripes and four pale stripes in middle of back.
brightness, but inner stripes tend to be more buff- Median dark stripe black, but outer dark stripes tend
orange than outer ones. Central black stripe usually to be mixed with reddish-brown. Outer pale stripes
has a thin, indistinct brown stripe in middle. Back brighter and more conspicuous than inner stripes,
of neck and top of head with strong reddish tinge but not connecting with cheek stripes. Pelage
in parts of Thailand, dull brown further south in relatively thin. Underparts greyish-white, hairs with
Cambodia and Vietnam. Outer pale stripe continuous grey base and white tip. Similar species Swinhoe’s
with pale stripe on side of head. Underparts dull Striped Squirrel, T. swinhoei, is very similar and

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 most reliably distinguished by range; Cambodian brown and underparts buffy-white, sometimes with a
Striped Squirrel, T. rodolphii, has all four pale stripes reddish tinge. Indistinct pale reddish-buff ring around
relatively bright, underparts mixed orange and grey. each eye. Tail short and bushy. Voice Most commonly
Ecology and habitat Diurnal and arboreal, but heard call consists of a series of bird-like ‘chik’s.
may be found in low shrubs and sometimes on the Similar species Slender Squirrel, S. tenuis, has a
ground. Occurs in a wide variety of forest types, longer, thinner tail and greyer underparts; Shrew-
including disturbed areas. faced Ground Squirrel, Rhinosciurus laticaudatus, is
Distribution SE Asia: Laos, Vietnam. Also SE China, larger, with more pointed head, more reddish tinge
Hainan and Taiwan. to fur.
Status Least Concern. Ecology and habitat Diurnal. Most active in the
early morning and late afternoon. Diet includes
Genus Sundasciurus A group of small to medium- fruits, insects and fungi. Travels and feeds in small
sized squirrels with many species in the Sunda standing trees, fallen trees and on the ground.
Islands and the Philippines, but only three species Occurs in tall and secondary forests, including
in the region, restricted to S peninsular Thailand scrub near forests, primarily lowland forest below
and Malaysia. Most species are smaller and plainer- 900m.
coloured than Callosciurus, with the exception of Distribution SE Asia: peninsular Thailand and
S. hippurus, which is relatively large and bright. They Malaysia. Also Sumatra, Borneo and some smaller
are both arboreal and terrestrial, spending more Indonesian islands.
time on the ground than most Callosciurus. Status Least Concern.

HORSE-TAILED SQUIRREL SLENDER SQUIRREL

Sundasciurus hippurus PLATE 66 Sundasciurus tenuis PLATE 66
Measurements HB 210–290, T 240–290, Measurements HB 115–155, T 125–132
HF 54–64 (85–95% of HB), HF 30–36
Identification Upperparts reddish-brown with Identification Upperparts speckled olive-brown;
contrasting grey head, shoulders and thighs; underparts grey with white or buffy-brown tip. Pale
underparts dark reddish-orange; tail thick and marks around eyes and above facial whiskers.
bushy, largely black. Voice Most commonly heard Facial profile slightly turned up. Tail rather long
call is ‘CHEK! CHEK! chekchekchekchek ...’ Similar and slender. Similar species Low’s Squirrel,
species No other squirrel has the contrasting S. lowii, has a shorter bushy tail and white or buffy
greyish and reddish pattern on the upperparts, with underparts without grey base; Grey-bellied Squirrel,
rufous underparts. Callosciurus caniceps, is larger with reddish or
Ecology and habitat Diurnal. Most often seen in orange tinge to back and bushier tail that often has
small trees, shrubs or on the ground, but sometimes black tip; Lesser Treeshrew, Tupaia minor, has more
climbs high into trees. Diet includes seeds, fruits pointed muzzle and relatively longer tail.
and insects. Occurs in primary and secondary Ecology and habitat Diurnal and primarily arboreal,
forests, mainly in lowlands and lower hill forest up to ranging from bushes near the ground to canopy. Diet
1,000m, but most common in intact forest. includes inner bark and insects from tree trunks,
Distribution SE Asia: peninsular Thailand and fruits and seeds. Occurs in tall and secondary
Malaysia. Also Sumatra and Borneo. forests as well as scrub and gardens.
Status Near Threatened; declining due to loss of Distribution SE Asia: extreme S peninsular Thailand
forests. and Malaysia, Singapore. Also Sumatra, Borneo and
some smaller Indonesian islands.
LOW’S SQUIRREL Status Least Concern; common in many areas.
Sundasciurus lowii PLATE 66
Measurements HB 130–150, T 80–100 (about Genus Dremomys Medium-sized squirrels
60–70% of HB), HF 30–35 with relatively long noses, found mainly in hill
Identification Upperparts speckled brown to reddish- or montane forest, where they are active mostly

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 in relatively low vegetation or on the ground. All than sympatric populations of Red-cheeked Squirrel,
species are largely brown, distinguished from one D. rufigenis, up to 3,000m.
another by the colour of the underparts and the Distribution SE Asia: N Vietnam. Also adjacent
pattern of red (or lack thereof) around the head and areas of Yunnan in S China.
tail. There is only limited overlap in the distribution Status Least Concern, although relatively poorly
of species in the region, except in N Vietnam and known.
N Myanmar.
RED-HIPPED SQUIRREL
RED-CHEEKED SQUIRREL Dremomys pyrrhomerus PLATE 67
Dremomys rufigenis PLATE 67 Measurements HB 215, T 120, HF 54
Measurements HB 170–210, T 143–147, Identification Upperparts, including top of tail,
HF 50–51, E 22–24 speckled olive-brown mixed with grey; cheeks,
Identification Upperparts, including top of tail, sides of face, hips, thighs and underside of tail
speckled olive-brown mixed with grey; cheeks, contrasting reddish-chestnut; underside white with
side of face and underside of tail contrasting light grey underfur, sharply demarcated from olive-
reddish-chestnut; underside white with light grey brown upperparts. Similar species Red-cheeked
underfur, sharply demarcated from olive-brown Squirrel, D. rufigenis, has slightly shorter rostrum,
upperparts. Small white or buff patch behind ear. more conspicuous red cheeks, with hips and outer
Similar species Red-throated Squirrel, D. gularis, thighs grey.
has darker underparts with red throat; Red-hipped Ecology and habitat Diurnal.
Squirrel, D. pyrrhomerus, has less conspicuous red Distribution SE Asia: extreme N Vietnam. Also
cheeks, large red patch on thigh; no other squirrels S and E China, Hainan.
have such contrasting red cheeks. Status Least Concern, although poorly known in
Ecology and habitat Diurnal. Found both in trees region.
and on the ground. Occurs in primary and secondary
forests, mainly in hills. Eats both insects and plant ORANGE-BELLIED SQUIRREL
material. Animals in northern part of range tend to Dremomys lokriah PLATE 67
be more arboreal than those in Peninsular Malaysia. Measurements HB 170–200, T 120–140,
Occurs from lowlands to 2,200m altitude in some HF 20–23
areas. Identification Upperparts finely speckled olive-
Distribution SE Asia: widespread throughout hillier brown and grey. Some individuals have dark line
areas of region, from N Myanmar through Thailand, down middle of back. Underparts mixture of orange
Laos (also in lowlands), Vietnam and Cambodia to or buff and grey, brightest on throat, but with
hills of Peninsular Malaysia. Also adjacent areas of considerable individual variation. Buff spot behind
S China. each ear. Similar species Callosciurus squirrels
Status Least Concern. have longer, bushier tail, tend to be more arboreal
and have different colour patterns (either greyer or
RED-THROATED SQUIRREL redder underparts).
Dremomys gularis PLATE 67 Ecology and habitat Diurnal. Found mainly near
Measurements HB 220, T 168, HF 52 ground level in both primary and secondary forest.
Identification Upperparts, including top of tail, Distribution SE Asia: W and N Myanmar. Also from
speckled olive-brown mixed with grey. Dull red on Assam west to C Nepal and nearby parts of China.
sides of face, throat, underside of legs and under Status Least Concern.
tail. Rest of underparts grey with white tip to fur
in middle. Similar species Red-cheeked Squirrel, PERNY’S LONG-NOSED SQUIRREL
D. rufigenis, has whiter underparts, with no red on Dremomys pernyi PLATE 67
throat. Measurements HB 180–210, T 130–165,
Ecology and habitat Diurnal. Found mainly near HF 42–28
ground level in forest. Occurs at higher elevations Identification Upperparts finely speckled olive-

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 brown and grey (agouti), as other species in genus. INDOCHINESE GROUND SQUIRREL
Underparts greyish-white. Orange spots behind Menetes berdmorei PLATE 66
ears. Often with some reddish around base of tail. Measurements HB 180–210, T 140–175,
Muzzle relatively elongate. Similar species Orange- HF 40–46
bellied Squirrel, D. lokriah, has less elongate muzzle, Identification Overall colour speckled brown,
orange throat and breast. Other Dremomys have sometimes slightly redder on the back, with two pale
orange cheeks. stripes on each side, separated by a darker stripe. Dark
Ecology and habitat Diurnal. Active mainly near stripe also present above top pale stripe in some areas,
ground level in shrubs and lower parts of trees. but inconspicuous or absent in others. Brightness
Occurs in primary and secondary forests, including of stripes varies geographically and seasonally; pale
highly disturbed areas. stripes vary from buff to rufous. Similar species
Distribution SE Asia: N Myanmar and Vietnam. Also Striped squirrels, Tamiops spp., are smaller, more
S China. arboreal, and have stripes in the middle of the back;
Status Least Concern. Three-striped Ground Squirrel, Lariscus insignis, has
three dark stripes on back, but no pale stripes.
Genus Lariscus Medium-sized squirrels, largely Ecology and habitat Diurnal and mainly terrestrial,
terrestrial. Only one species occurs in the region, though also forages in low bushes, bamboo and
but additional species occur in Sumatra, Borneo and scrub. Found mainly in drier, more open forests and
the Mentawai Islands. savanna.
Distribution SE Asia: Myanmar, Thailand, Laos,
THREE-STRIPED GROUND SQUIRREL Vietnam and Cambodia. Also China (SW Yunnan).
Lariscus insignis PLATE 67 Status Least Concern.
Measurements HB 170–230, T 100–135,
HF 33–46, E 14–18 Genus Rhinosciurus Medium-sized ground
Identification Upperparts brown with three black squirrel with elongate nasal bones, making the head
stripes along back; underparts white or buff (turning profile relatively pointed and superficially resembling
dark yellow in old skins), sometimes with a faint Tupaia treeshrews. Only one species currently
red tinge. Similar species Other species of striped recognised in genus.
squirrel (Menetes and Tamiops) have buff as well as
black stripes on back; Shrew-faced Ground Squirrel, SHREW-FACED GROUND SQUIRREL
Rhinosciurus laticaudatus, is also terrestrial and Rhinosciurus laticaudatus PLATE 67
similar in colour, but lacks stripes and has a more Measurements HB 195–233, T 131–170 (usually
pointed muzzle. less than 70% of HB), HF 40–46
Ecology and habitat Diurnal and mainly terrestrial. Identification Upperparts dark brown without stripes;
Diet includes fruits and insects. Occurs in tall and underparts white or buff, turning to yellow in old skins.
secondary forests, sometimes entering disturbed Tail short and bushy, often held upwards with fluffed-
areas. Mainly at altitudes of 300–1,500m. out hairs when the squirrel is active. Muzzle pointed.
Distribution SE Asia: peninsular Thailand and Similar species Treeshrews, Tupaia spp., have a
Malaysia. Also Sumatra, Java, Borneo and some relatively long tail, even more pointed muzzle, darker
intervening islands. underparts and very different dentition.
Status Least Concern, but declining due to loss of Ecology and habitat Diurnal and terrestrial.
forest. Diet includes insects taken from fallen trees and
branches. Occurs in lowland rainforest, including
Genus Menetes A medium-sized ground squirrel mature forests and selectively logged forests.
with many similarities to Lariscus, but can be Distribution SE Asia: Peninsular Malaysia, extreme
distinguished by pattern of dark and pale stripes on S Thailand and Singapore. Also Sumatra, Borneo
the sides instead of black stripes on the back. Only and adjacent small islands.
one species currently recognised in genus. Status Near Threatened; populations declining due
to loss of forest.

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 Subfamily SCIURINAE, Tribe PTEROMYINI FLYING SQUIRRELS
Flying squirrels, although incapable of true flight, unlike bats, have membranes stretched between the fore
and hind legs, which enable them to glide long distances between trees. Unlike in the Colugo, which also
glides, the tail is not enclosed by a membrane.
Flying squirrels are mainly nocturnal and most active high up in trees, so they are difficult to see,
especially the small species. The larger flying squirrels are often active only around dusk. If the light is
adequate, or if a powerful headlamp or spotlight is available, the larger species can often be distinguished
by colour patterns, especially with binoculars. Several of the small to medium-sized flying squirrels are quite
similar in appearance, and may be difficult to identify unless seen at close range. Pygmy flying squirrels have
a diagnostic white tip to the tail.
The skulls of flying squirrels have relatively large, well-developed auditory bullae with complex
patterns of septa within them. Differences in shape and size of the auditory bullae can be helpful for
distinguishing genera of museum specimens, especially some superficially similar groups such as
Hylopetes and Petinomys (Fig. 68).
Small flying squirrels can occasionally be captured by field workers for study in mist nets at night, or in
cage traps set in trees, well above the ground, but are more often caught by searching nest holes in tree
trunks. Many flying squirrels caught in this way are immature, which can make them more difficult to identify.

Genus Petaurista Very large flying squirrels with melanotus) has entire body dark reddish except
long, rounded tail. Cheek teeth have complex, for black on nose, chin, eye-ring, behind ears,
folded cusp patterns. Auditory bullae moderately feet and tail tip. In S Myanmar and W Thailand
large, with four or fewer dividing septa internally. (P. p. taylori), colour is similar but with light speckling
Currently, five species recognised from the region, on head and back. In most of rest of Thailand and
but with considerable geographic variation in all Myanmar (P. p. candidula), overall colour is pale
species; taxonomic limits, both within and among orange-brown with white speckling, especially on
species, are not well understood. For example, some head; tail brownish or greyish with black tip; throat
populations currently allocated to P. petaurista were white; rest of underparts off-white, pale line over
formerly considered part of P. alborufus, a species each shoulder. Similar species Lesser Giant Flying
that otherwise occurs only in China. Conversely, Squirrel, P. elegans, has conspicuous white spots on
some populations now allocated to P. philippensis back; Indian Giant Flying Squirrel, P. philippensis,
were formerly included within P. petaurista. Recent has either all-dark or all-grey tail, depending on
molecular genetic studies from China indicate that the population.
P. yunanensis, formerly considered a subspecies of Ecology and habitat Mostly nocturnal, but
P. philippensis, is actually a distinct species. Further becomes active shortly before dusk and may be
genetic studies may lead to additional changes seen climbing up large trees or gliding between
in species limits within South-east Asia. The text trees just before dark. Occasionally active during the
describes characteristics of different populations, day or seen resting on exposed parts of tall trees
and the major different types are illustrated, but in until mid-morning. Can make continuous glides of
some areas there may be intermediates between up to 100m. Nests in holes in large trees, usually at
these forms. least 10m above the ground. Found alone or in small
groups. Diet includes leaves and seeds. Occurs in
RED GIANT FLYING SQUIRREL forests, including highly disturbed areas with some
Petaurista petaurista PLATE 68 tall trees.
Measurements HB 400–520, T 400–600, Distribution SE Asia: Myanmar, Thailand, S Laos,
HF 70–100. Wt 1,600–2,900g (larger in north Cambodia and Peninsular Malaysia. Also Nepal,
than south) N India, S China, Java, Sumatra and Borneo.
Identification Considerable geographic variation Status Least Concern, but likely to be declining
in colour pattern; possibly more than one species due to loss of forests. Relatively rarely recorded in
is involved. Peninsular Malaysian form (P. p. SE Asia outside of Malaysia.

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 INDIAN GIANT FLYING SQUIRREL Ecology and habitat Nocturnal. Occurs in dipterocarp
Petaurista philippensis PLATE 68 hill and lower montane forests.
Measurements HB 400–490, T 400–550, Distribution SE Asia: Myanmar, Thailand, Laos,
HF 65–90 Cambodia and Vietnam. Also Sri Lanka, India,
Identification Moderate geographic variation in S China, Taiwan and Hainan.
colour pattern, and may include more than one Status Least Concern, but experiencing some
species. W Myanmar forms (P. p. cineraceus): declines due to loss of forest and hunting.
upperparts, including tail and membranes, medium
grey; back heavily frosted with long white tip to hairs; YUNNAN GIANT FLYING SQUIRREL
underparts greyish-white. E Myanmar and Thailand Petaurista yunanensis PLATE 68
(P. p. lylei): upperparts very dark grey, heavily frosted Measurements HB 400–500, T 400–550,
with white; dorsal surface of membrane, feet and tail HF 65–90
black; underparts buff. Vietnam (P. p. annamensis): Identification Upperparts dark grey, with reddish
upperparts dark brown or dark red, heavily frosted base to fur, and white frosting on many hairs; gliding
with white, underparts pale orange-brown, tail membranes dull red; feet black; tail dark grey.
dark. Similar species Red Giant Flying Squirrel, P. Taxonomic notes Formerly considered a form of
petaurista, in areas of overlap, is more rufous with P. philippensis, but recent genetic studies indicate it
paler head, tail with a discrete black tip. is distinct and more closely related to P. petaurista.

a b

Fig. 68 Underside of posterior half of skull of selected small flying squirrels, showing variation in shape of
auditory bullae: Petinomys genibarbis (a), Petinomys setosus (b), Petinomys vordermanni (c), Hylopetes spadiceus (d)
and Petaurillus kinlochii (e).

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 Similar species Red Giant Flying Squirrel, tip; Indian Giant Flying Squirrel, P. philippensis, is
P. petaurista, is more rufous with paler head, tail greyer without large white spots.
with a discrete black tip; Indian Giant Flying Squirrel, Ecology and habitat Nocturnal. Occurs in dipterocarp
P. philippensis, does not have combination of grey hill and lower montane forests. In areas of sympatry
back and red gliding membranes. with other species of Petaurista, more often occurs
Ecology and habitat Nocturnal. Occurs in hill at higher elevations, though may extend to lowlands.
forests. Occurs sympatrically with P. philippensis Distribution SE Asia: Myanmar, Thailand, Laos,
in China, but differences in ecology have not been Vietnam and Peninsular Malaysia. Also S China
studied as only recently recognised as distinct. (Yunnan), Java, Sumatra and Borneo.
Distribution SE Asia: Myanmar, N Laos and Status Least Concern. Still relatively common in
N Vietnam. Also SW Yunnan in China. Precise some areas of hill forest.
distribution limits not yet determined, as only
recently recognised as distinct. Genus Aeromys This genus contains two species
Status Not yet assessed. of large flying squirrels, of which one occurs in
the region. Superficially similar to Petaurista, with
CHINDWIN GIANT FLYING SQUIRREL long, rounded tail, they differ in a number of skull
Petaurista sybilla PLATE 68 characters, including simpler patterns of raised,
Measurements HB 350–400, T 350–370, HF 60 rounded cusps on the molars and pale yellow
Identification Similar in size and shape to instead of orange incisors. The one species in the
P. elegans, but without any white spots. Overall, region is readily identified by colour.
largely reddish-brown with slightly greyer-brown
on back, without distinctive characters. Taxonomic BLACK FLYING SQUIRREL
notes Sometimes considered a form of P. elegans. Aeromys tephromelas PLATE 69
Similar species Red Giant Flying Squirrel, Measurements HB 355–426, T 410–470,
P. petaurista, is larger with a black tail tip; Lesser Giant HF 67–78. Wt c.900g
Flying Squirrel, P. elegans, has white spots on back. Identification Upperparts, tail and cheeks dark
Ecology and habitat Nocturnal. Hill forest. grey-brown, almost black, with fine, pale speckling
Distribution SE Asia: mountains of N Myanmar. Also on the back; underparts slightly paler, with sparse,
adjacent areas of Yunnan in China. fluffy hair. Similar species Other large flying
Status Not assessed separately from P. elegans. squirrels, Petaurista spp., are coloured differently,
although they can look black in poor light at night;
LESSER GIANT FLYING SQUIRREL Smoky Flying Squirrel, Pteromyscus pulverulentus,
Petaurista elegans PLATE 69 is much smaller and has creamy underparts.
Measurements HB 340–365, T 340–365, Ecology and habitat Nocturnal. Occurs in tall and
HF 60–66. Wt 840–1,240g secondary forests, including some tree plantations
Identification In Peninsular Malaysia and extreme and gardens.
S Thailand, upperparts, including membranes, Distribution SE Asia: Peninsular Malaysia and
dark rufous mixed with black, with extensive S Thailand. Also Sumatra and Borneo.
large white ‘spots’ of heavy white flecking; tail Status Data Deficient; relatively rarely recorded, but
black; underparts pale rufous. Elsewhere in likely to be declining due to loss of lowland forest.
region: upperparts brown, with fewer white spots
mainly on head and centre of back; feet and tail Genus Biswamoyopterus This genus was first
reddish-brown. Outside region, in Java, may lack described in 1981 from India, with a second
white spots. May be some intermediate forms in S species described from Laos in 2015. Very large
Thailand. Similar species Chindwin Giant Flying squirrels, superficially similar to Petaurista, with
Squirrel, P. sybilla, lacks spots; Red Giant Flying a long, rounded tail. They differ in a number of
Squirrel, P. petaurista, is either uniform reddish or skull characters, including pale yellow instead
frosted white, without discrete clumps of white, of orange incisors; cheek teeth with simplified
and tail is reddish or brown with discrete black cusp patterns; and very large auditory bullae with

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 many internal septae (>12) forming a complex creamy white. Tail bushy, rounded above, flattened
honeycomb pattern. The one species in the region below; reddish-brown with a paler grey base. Feet
can be distinguished from other large flying squirrels have long, sensory hairs. Tufts of long hairs behind
by colour. ears. Similar species Particoloured Flying Squirrel,
Hylopetes alboniger, is more uniformly coloured
LAO GIANT FLYING SQUIRREL without a contrasting reddish-brown tail and lacking
Biswamoyopterus laoensis PLATE 69 tuft of hairs behind ears; Smoky Flying Squirrel,
Measurements HB 455, T 620, HF 74.5, E 52. Pteromyscus pulverulentus, is larger, greyer and
Wt 1,800g. Skull: gl 74 does not overlap in range.
Identification Upperparts, including head, Ecology and habitat Nocturnal. Nests in holes in
dark reddish brown and black, mixed with long trees. Occurs in broadleaved hill forest.
whitish hairs on back and head, giving a grizzled Distribution SE Asia: N Myanmar, Thailand, Laos
appearance. Tail very long, rounded in cross-section and Vietnam. Also E Nepal, S China, Hainan and
and mostly black except at base. Underside mostly Taiwan.
pale orange, partially mixed with grey in the middle. Status Data Deficient; likely to be declining in region
Gliding membrane thickly haired, dark reddish- due to loss of forest and hunting.
brown and black without white hairs above; greyish
with extensive fine black lines and thin orange hairs Genus Pteromyscus Cheek teeth have relatively
below. Ears dark grey without long hairs, but with complex pattern of ridges, but differ in pattern from
long tufts of hair near their base. Taxonomic notes those of Belomys. Hairs on tail longer on sides than
Newly described in 2015. Similar species None of on top or bottom, producing a slightly flattened
the other large flying squirrels in the region have effect. Auditory bullae relatively large, divided into
the combination of frosted dark brown upperparts discrete sections by five septa.
and pale orange underparts; also Petaurista spp.
lack black lines on gliding membrane and have dark SMOKY FLYING SQUIRREL
orange incisors. Pteromyscus pulverulentus PLATE 70
Ecology and habitat Unknown. Only known Measurements HB 220–290, T 215–235,
specimens were found for sale in markets; HF 41–44. Wt 232–305g
presumably had been caught in nearby forests. Identification Upperparts very dark grey-brown
Distribution SE Asia: known only from a limited with fine, pale greyish speckling; underparts creamy
area in C Laos. with some grey. Tail dark grey-brown with grey-
Status Not yet assessed; limited range and hunting buff at base. Cheeks greyish. Tail slightly flattened
pressures suggest likely to be at risk. above, flat below. Similar species Black Flying
Squirrel, Aeromys tephromelas, is larger with darker
Genus Belomys Cheek teeth have an exceptionally upperparts, greyish fluffy hair on the underparts.
complex pattern of ridges, similar to Trogopterus, Ecology and habitat Nocturnal. Nests in tree holes.
with which it is sometimes combined. The auditory Occurs in tall lowland rainforest.
bullae are relatively small but slightly bulbous, with a Distribution SE Asia: Peninsular Malaysia. Also
honeycomb pattern of septa that may be hard to see Sumatra and Borneo.
owing to thick bone. Status Endangered; relatively rare, and is likely to
have experienced large declines due to clearing
HAIRY-FOOTED FLYING SQUIRREL of primary forest in its range and apparently low
Belomys pearsonii PLATE 70 tolerance of disturbed areas.
Measurements HB 190–200, T 150–160, HF 34
Identification Upperparts blackish, with reddish- Genus Iomys Only one species in the region, with
brown or white tip, giving an overall speckled an additional species in Indonesia. This species
reddish or speckled grey appearance; gliding can be recognised by its medium-large size and
membrane darker, almost black, with a pale edge; generally dull orange colour, especially on the tail.
underparts, including throat, breast and belly, The genus is characterised by relatively square

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 cheek teeth with a distinctive cusp in each corner. on the cheek behind the eye. Auditory bullae with
The auditory bullae are relatively small but bulbous, honeycomb pattern. Similar species No other flying
with one or two internal septa. squirrels have a pinkish rump or prominent tuft of
whiskers behind the eye.
HORSFIELD’S FLYING SQUIRREL Ecology and habitat Nocturnal. Occurs mainly
Iomys horsfieldii PLATE 70 in primary lowland rainforest, but also in some
Measurements HB 165–230, T 160–207, secondary forests.
HF 33–40. Wt 135–215g Distribution SE Asia: Peninsular Malaysia. Also
Identification Upperparts brown to dark grey, hairs Sumatra, Java and Borneo.
with buff or dull orange tip; underparts orange- Status Vulnerable; has declined due to loss of
buff or white with orange around the edges. lowland rainforest.
Gliding membrane fringed with rusty-brown. Tail
relatively round and rusty-brown. Cheeks buffy or TEMMINCK’S FLYING SQUIRREL
rusty, contrasting with darker top of head. Similar Petinomys setosus PLATE 71
species Vordermann’s Flying Squirrel, Petinomys Measurements. HB 105–125, T 93–115,
vordermanni, is similar colour but much smaller; HF 21–25. Skull: cbl 27.6–29.6
Hylopetes flying squirrels have a white margin to the Identification Upperparts dark brown or black
gliding membranes and a flatter tail. with pale buff tip; underparts white with pale
Ecology and habitat Nocturnal. Occurs in forests, grey underfur. Fur soft and silky. Margin of gliding
plantations and gardens with tall trees. Usually membrane not distinctly pale. Tail dark brownish-
sleeps in tree holes, but may sometimes use leafy grey, with pale patch of whitish hairs at base.
nests. Cheeks greyish. Dark ring around eye that usually
Distribution SE Asia: Peninsular Malaysia and connects with a black streak to the nose. Form in
Singapore. Also Sumatra, Java and Borneo. N Myanmar and NW Thailand is slightly larger, with
Status Least Concern; relatively common in many a white tip to the tail. Similar species Phayre’s
areas. Flying Squirrel, Hylopetes phayrei, is substantially
larger, without a white tip to the tail and with a
Genus Petinomys Several species in the region, different skull structure; other small flying squirrels
superficially similar to Hylopetes, mostly with have at least some orange on the upperparts, base
distichous (feather-like) tail. Auditory bullae broad of tail or cheeks.
and relatively flat with multiple septa. In some Ecology and habitat Nocturnal. One record of a
species septa are both transverse and horizontal, nest hole, 19mm wide, in a tree trunk 0.5m above
forming a complex honeycomb pattern (Fig. 68a, c); the ground. Occurs in lowland rainforest in southern
in others septa are in a radial pattern forming part of range, but deciduous hill forest in north.
discrete chambers (Fig. 68b), but with bullae still Distribution SE Asia: N Myanmar, NW Thailand,
relatively flat. Most species can be distinguished by extreme S Thailand and Peninsular Malaysia. Also
colour patterns. Sumatra and Borneo.
Status Vulnerable; declining due to loss of lowland
WHISKERED FLYING SQUIRREL rainforest.
Petinomys genibarbis PLATE 71
Measurements HB 160–180, T 155–188, VORDERMANN’S FLYING SQUIRREL
HF 31–32 Petinomys vordermanni PLATE 71
Identification Upperparts reddish-brown with Measurements HB 95–120, T 89–115, HF 21–23.
grey underfur, speckled with grey anteriorly, and Skull: cbl 28.2–30.0
with reddish posteriorly, giving an unusual pinkish Identification Hair of upperparts blackish with
tinge; underparts cream or dull orange-buff. Gliding rusty-coloured tip; underparts buffy-white. Top
membrane with a white margin. Tail rusty-coloured of head and tail brown to reddish-brown. Gliding
with darker brown streaking. Whitish hairs at the membrane with a buff (not white) margin. Tail brown
base of each ear and a distinct tuft of long whiskers to reddish-brown with paler buffy hairs at base;

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 slightly rounded above, but very flat below. Cheeks Singapore. Also Sumatra, Java and Borneo.
orange. Black ring around each eye. Tuft of bristle- Status Least Concern.
like hairs at front of base of ear. Auditory bullae of
skull with honeycomb pattern. Similar species GREY-CHEEKED FLYING SQUIRREL
Hylopetes spp. have white margins on the gliding Hylopetes platyurus PLATE 71
membranes and a different face pattern; Horsfield’s Measurements HB 115–135, T 118–120,
Flying Squirrel, Iomys horsfieldii, is larger. HF 29–30. Skull: gl 29–38
Ecology and habitat Nocturnal. Nest holes Identification Upperparts blackish or dark grey-
recorded at 0.3–6.0m above the ground. Mainly brown with rust-coloured markings, especially in the
reported from lowland rainforest. Occurs in tall and midline; underparts white or buffy-white with grey
secondary forests. underfur. Gliding membrane with a narrow white
Distribution SE Asia: S Myanmar, S Thailand and margin. Tail narrow at base, then broad, then tapers
Peninsular Malaysia. Also Borneo and small islands to tip, dark brownish-grey to black. Cheeks and
off E Sumatra. patch on each side of base of tail pale grey, often
Status Vulnerable; substantial population declines tinged with yellow but never distinctly orange. In
due to loss of lowland rainforest. skull, auditory bullae are relatively small. Taxomonic
notes Formerly considered part of H. lepidus from
Genus Hylopetes A genus of about ten medium Java and Borneo. Similar species Red-cheeked
to small species, of which four occur in the region. Flying Squirrel, H. spadiceus, has orange cheeks
Tail is usually distichous (feather-like). Superficially and base of tail; Vordermann’s Flying Squirrel,
similar to many species of Petinomys, but skull is Petinomys vordermanni, has buff margin to gliding
very different, with auditory bullae that are bulbous membrane, yellowish-buff cheeks, reddish tail.
with only two or three septa (Fig. 68d). Ecology and habitat Nocturnal. Occurs mainly
in lowland rainforests, but apparently tolerant of
RED-CHEEKED FLYING SQUIRREL moderate habitat disturbance.
Hylopetes spadiceus PLATE 71 Distribution SE Asia: extreme S Thailand and
Measurements HB 157–184, T 152–166, Peninsular Malaysia. Also Sumatra.
HF 29–35. Wt 80–157g. Skull: gl 33–39 Status Data Deficient; likely to be declining from
Identification Upperparts blackish or dark grey- loss of habitat, but uncertain how well it adapts to
brown with rust-coloured markings, especially along disturbed forest.
midline; underparts white with orange tinge and
grey base. Gliding membrane blackish with a thin PARTICOLOURED FLYING SQUIRREL
white margin. Tail dark, slightly orange-brown with Hylopetes alboniger PLATE 70
buffy underfur; distinctly orange at the base. Cheeks Measurements HB 175–225, T 180–230,
orange-brown on grey. In skull, auditory bullae are HF 36–45. Skull: gl 46–51
relatively large. Similar species Grey-cheeked Flying Identification Upperparts with dark greyish base to
Squirrel, H. platyurus, is somewhat smaller and has fur, paler brown or greyish tip. Underparts white. Tail
principally greyish-white cheeks and base of tail, with greyish to greyish-brown, paler at base, darker at
only traces of a yellowish tinge; Vordermann’s Flying tip, flattened below, but somewhat rounded above.
Squirrel, Petinomys vordermanni, has buff margin Similar species Phayre’s Flying Squirrel, H. phayrei,
to gliding membrane, distinct face pattern and is is similar but usually smaller, with proportionately
smaller; Phayre’s Flying Squirrel, H. phayrei, has no larger auditory bullae in skull.
orange, and has tail rounded above. Ecology and habitat Nocturnal. Occurs mainly in
Ecology and habitat Nocturnal. Nest holes about hill and montane forest, up to 3,000m in Thailand,
32mm wide recorded at 0.3–3.3m above the especially in oak forest.
ground. Occurs in a variety of forest types; generally Distribution SE Asia: N Myanmar, Thailand, Laos,
more abundant in tall forest than disturbed areas. Vietnam and Cambodia. Also Nepal through Assam
Distribution SE Asia: Myanmar, Thailand, S Laos, to S China, Hainan.
S Vietnam, Cambodia, Peninsular Malaysia and Status Least Concern, although likely to be declining.

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 PHAYRE’S FLYING SQUIRREL Genus Petaurillus Among the smallest flying
Hylopetes phayrei PLATE 70 squirrels in the world, with two species, only one of
Measurements HB 145–195, HF 30–36. which occurs in the region. Tail distichous (feather-
Skull: gl 37–43 like). In skull, auditory bullae relatively swollen and
Identification Overall colour of upperparts greyish- bulbous with two septa, similar to Hylopetes but
brown to mottled brown and black; individual hairs smaller (Fig. 68e).
with dark slaty to blackish base, and very extensive
greyish-buff to brown tip. White mark behind ear. MALAYAN PYGMY FLYING SQUIRREL
Underparts pale buff. Tail narrow at base, then Petaurillus kinlochii PLATE 71
broad, then tapering gradually to tip; rounded Measurements HB 80–90, T 80–98, HF 19–20.
above, flattened below; greyish-brown to slightly Skull: cbl 25.2
cinnamon-brown, with darker tip. Similar species Identification Upperparts very dark grey with pale
Particoloured Flying Squirrel, H. alboniger, is very buff streaks, especially in the midline; underparts
similar but larger, with proportionately smaller off-white with grey underfur. Tail buffy at base,
auditory bullae; Red-cheeked Flying Squirrel, becoming blacker near the end, with a white tip.
H. spadiceus, has tail flattened above and below, Cheeks buffy-white with a strong grey tinge beneath
reddish cheeks and base to tail. the eye. A whitish spot behind each ear. Taxonomic
Ecology and habitat Nocturnal. Occurs in tall and notes Sometimes considered a form of P. hosei from
secondary forests, including disturbed areas and Borneo. Similar species Other small flying squirrels
cultivated orchards. Feeds on fruits. in its range are larger and do not have a white tail tip.
Distribution SE Asia: Myanmar, Thailand, Vietnam. Ecology and habitat Nocturnal. Roosts and nests in
Probably also Laos. Also Nepal through Assam to S holes in trees. Reported from both natural forest and
China, Hainan. adjacent rubber plantations.
Status Least Concern. Distribution SE Asia: Peninsular Malaysia.
Status Data Deficient; few records so distribution
and habitat requirements poorly known.

Superfamily MUROIDEA RATS AND MICE

Rodents of the superfamily Muroidea, commonly called rats and mice, can be found nearly everywhere
on Earth, whether native or introduced by humans. Worldwide, this group contains over 1,500 recognised
species, of which 70 are presently recognised in mainland South-east Asia.
The selection of English names for this group is particularly challenging. Even the choice of the name ‘rat’
or ‘mouse’ is often rather arbitrary. In Europe, the name ‘rat’ is applied to larger species, of which the best
known are in the genus Rattus, while the name ‘mouse’ is applied to smaller species, especially in the genus
Mus. However, these names have since been applied to genera all over the world, largely on the basis of size,
with little regard for actual relationships. Many species called ‘mouse’ are much more distantly related to
Mus than Mus is to Rattus. Recently, there has been a tendency to develop new English group names based
on the scientific names, to reflect relationships more accurately. For some genera, such as Maxomys and
Niviventer, these new names are used here in the interests of consistency and to avoid long, complicated
names. However, whenever practical, the familiar names of ‘mouse’ or ‘rat’ have been retained as part of the
English name to make them easier to remember and recognise.
Some rats and mice are confined to human settlements or to vegetation that has been disturbed or
modified by humans. Others are strictly forest dwellers, either terrestrial or partially or mainly arboreal. All
South-east Asian rats and mice are mainly active at night.
It is normally necessary to catch rats and mice for positive identification. In some cases, it may be
necessary to prepare the animal as a museum specimen and examine the skull to confirm the identity. For
the benefit of people who may be working in a museum, or who may find a dead animal, the text sections
feature some information on skull characters, particularly those that help to diagnose the genus, including
the position and relative length of the incisive foramina and the bony palate (e.g. Fig. 69).

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 a c

Fig. 69 Undersides of skulls of selected rats, showing

differences in size and position of incisive foramina
Incisive foramina and posterior edge of palate relative to teeth:
Chiromyscus langbianis (a), Rattus argentiventer (b)
Posterior edge of palate 0 10
and Leopoldamys edwardsi (c).

The size and shape of the teeth, especially the

pattern of cusps on the molars, are also important a b c d
for identification, especially for separating genera;
some genera have many separate cusps on the
molars, while in others the cusps are fused to form
parallel ridges (Figs 70, 71). In a live animal, teeth
can usually be examined only if it has first been
anaesthetised. Some care is required in examining
cusp patterns, because the teeth become worn
with age, and cusps that are separate in a young
animal may appear joined in a very old animal.
In most cases there are also external characters
that help with identification, and with practice it
should be possible to identify most species in life.
The presence and relative density of spines in
the fur are important for identifying many rats. The
0 3
spines are often hard to see but are easily detected
by feel – they are stiff, but not sharp, and can be
found by running a finger backwards along the fur.
Also important are the relative length and colour Fig. 70 Upper right toothrows, with anterior at top,
pattern of the tail (particularly whether it is all dark of selected mice and voles: Mus musculus (a),
or bicoloured). Apodemus draco (b), Micromys erythrotis (c) and
The sizes and shapes of footpads are also useful Eothenomys melanogaster (d).
identification characters (Figs 72, 73). All rats have
pads on the bottom of their feet, which vary among species in size, shape and the amount of fine striations
(like fingerprints) on the bottom. Species that climb well tend to have shorter feet with larger, better developed
footpads and more conspicuous striations.
The arrangement of the mammary glands (mammae) is a useful identification character for females, and
can help to distinguish genera and species (Fig. 74). The number of pairs of mammae is given as the number
near the front legs (axillary – typically one pair just in front and one or two pairs just behind the base of the
leg) and the number near the back legs (abdominal and inguinal) using a formula such as mammae: 3 + 3.
The number of pairs varies among most species from two (0 + 2) to six (3 + 3), but one species (Bandicota
bengalensis) sometimes has up to 10 or more pairs.

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 a b c d e

i
f g h

Fig. 71 Upper right toothrows, with

anterior at top, of selected medium
and large murid rodents:
Rattus rattus (a), Niviventer
confucianus (b), Lenothrix canus (c),
Pithecheir parvus (d), Hapalomys
longicaudatus (e), Leopoldamys
edwardsi (f), Sundamys muelleri (g),
Bandicota indica (h) and Dacnomys
millardi (i).
0 10

Immature rats are frequently found and can be

difficult to identify as they often differ in appearance
from adults. Not only are the measurements smaller,
but the colour and texture of the fur is different –
usually softer and greyer. Any rat or mouse that
differs from all of the illustrated species and is
distinctly smaller than the most similar species
described may be an immature. Immatures can be
recognised by disproportionately large heads and
feet (which grow faster than the rest of the body)
and incompletely developed sexual organs. Adult
females have several pairs of distinct, readily visible a b
nipples, while adult males have a large scrotum
and testes; both are small and inconspicuous
in immatures. Immatures moulting into adult
pelage can sometimes be recognised by having
a patchwork of immature and adult fur colour. In
addition, young rats have unworn teeth that may
not be fully erupted from the gums; and their skull
bones are not solidly fused, tending to fall apart if Fig. 72 Undersides of left hind feet of Rattus argentiventer
the animal is prepared as a museum specimen. (a) and R. rattus (b). In addition to having larger pads, R.
Even among sexually mature rats, measurements rattus has much deeper and more conspicuous striations
must be considered somewhat cautiously because (like fingerprints) on the pads (not shown in sketch).

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 a b c d e
3
2

4
1
omt
imt

Fig. 73 Undersides of left hind feet of Chiromyscus chiropus (a), Chiromyscus langbianis (b), Niviventer fulvescens (c),
Saxatilomys paulinae (d) and Tonkinomys daovantieni (e), showing differences in size and shape of the four
interdigital (1–4), inner metatarsal (imt; also called thenar) and outer metatarsal (omt; also called hypothenar) pads.

a b c d

Axillary

Inguinal

Fig. 74 Undersides of adult female rats, showing some of the different patterns of axillary and inguinal mammae
(teats): 3 + 3 (a); 2 + 3 (b); 3 + 2 (c); 2 + 2 (d), as described in the species accounts. The pair of mammae in front of
the arms are sometimes called pectoral, and those immediately behind them post-axillary.

most murids continue to grow as adults. It can be useful to examine the teeth – an animal with very worn
teeth is likely to be an older individual that may be larger, for a particular species, than a younger adult with
less worn teeth.
A rat or mouse that does not seem to fit any description and yet appears to be mature could be an
undescribed species. Several new species and even new genera have been described from the region in
recent years, and additional species are likely. An animal that is entirely new to specialists working in the
field is sometimes collected and preserved as a museum specimen, either as a skin and skull or preserved

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 in formalin or alcohol as described in the introduction. The colours are described as accurately as possible,
as they tend to change with time: red sometimes fades, and fur can turn yellowish. Colour photographs help,
ideally taken with a colour reference strip in the photo. The specimen is labelled with the date, location and
habitat of capture, and sent to a reputable museum for identification.
The order in which families, subfamilies and genera are presented varies considerably among published
books. Here, these are arranged based generally on size and morphological similarities, starting with the
most widespread family (Muridae) and the genus Rattus, followed by other genera of larger species (rats) in
this family, followed by the various genera of smaller species (mice).

Family MURIDAE RATS AND MICE

This is one of the largest families of rodents, with more than 560 species currently recognised worldwide in
126 genera. All of the South-east Asian forms are in the subfamily Murinae. These can be distinguished from
other families of rodents in the region by having only three cheek teeth, with patterns of cusps on the teeth
that vary from many separate cusps to parallel ridges (Figs 70a, b, c, 71). The tail is usually moderately long
to very long, covered with scales and only sparsely haired.

Genus Rattus Small to large rats. Tail is usually upper surface of foot usually brown with white hairs
entirely dark except in some Norway Rats; spines on toes, sometimes all white. Front foot has dark fur
numerous on upperparts, scarce on underparts, but over wrist with white hairs on toes. Indian/European
stiffness varies somewhat among species; generally forms of R. rattus, which may occur in some towns
soft, like stiff hairs. In skull, incisive foramina elongate, (‘Black Rat’) are all dark blackish-grey including
extending behind first upper molar, posterior edge of underparts. Mammae usually 2 + 3, sometimes 3 +
bony palate extending beyond third upper molar (Fig. 3, in which case both post-axillary pairs are generally
69b). Most cusps on teeth fused into curved ridges very close together. Taxonomic notes This is actually
(Fig. 71a). Differences among species subtle and a complex of species, with South-east Asian forms
somewhat variable; it is important to examine several often called Rattus tanezumi, differing in chromosome
features when identifying species. Small Rattus, count from true Rattus rattus, which is apparently
including juveniles, can be distinguished from Mus native to India and introduced in much of the rest of
by elongate instead of rounded inner metatarsal pad the world. However, recent genetic studies suggest
on hind foot (Fig. 73). Mammae vary among species that there are more than two similar species, and
from 3 + 3 to as few as 2 + 2 (Fig. 74). Experience is the most appropriate names for each are uncertain,
required to identify specimens accurately, especially so they are treated together here. Similar species
with dry skins where fur colour may have altered. Malaysian Wood Rat, R. tiomanicus, always has
whitish underparts and sleeker upperparts with less
HOUSE RAT prominent guard hairs; Ricefield Rat, R. argentiventer,
Rattus rattus PLATE 72 has upperparts more speckled with black, often with
Measurements HB 105–215, T 120–230 (95– a small tuft of orange fur in front of ear; underparts
120% of HB; usually just over 100%), HF 26–40, grey to silvery; feet darker with less developed
E 17–25. Wt 100–200g. Skull: cbl 33–43, footpads; White-footed Indochinese Rat, R. nitidus,
mt 6.2–7.0 has narrower hind feet, pure white front and hind feet,
Identification Medium-sized rat. Upperparts finely darker woollier dorsal fur; Indochinese Forest Rat,
grizzled olive-brown, sometimes with greyish or R. andamanensis, has larger pads on hind feet, longer
reddish tone; underparts usually buffy-brown with tail, more prominent guard hairs along entire length of
grey base but sometimes creamy white. Dorsal fur back; Pacific Rat, R. exulans, is usually much smaller.
moderately spiny, especially on flanks, with long black Ecology and habitat Nocturnal and sometimes
guard hairs, especially on lower back. Ears large and diurnal. Diet includes a wide range of plant and
thinly furred. Tail similar length to body, dark above animal matter. In most areas, largely confined
and below, very rarely with a white tip. Hind foot broad to human settlements, plantations and gardens,
with prominent pads and clear striations; hairs on where it is a major pest in stored crops. In areas

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 where other species, such as R. argentiventer, are RICEFIELD RAT
absent, including parts of Laos, can be abundant Rattus argentiventer PLATE 72
in agricultural fields. Climbs well, including inside Measurements HB 140–210, T 130–205
buildings, but also spends much time on the ground. (80–125% of HB, usually just under 100%),
Distribution SE Asia: Myanmar, Thailand, Laos, HF 30–40, E 16–24. Wt 85–240g.
Vietnam, Cambodia and Peninsular Malaysia. R. Skull: cbl 35–45, mt 6.7–8.2
‘tanezumi’ form also in Nepal, N India, Bangladesh, S Identification Upperparts pale brown to orange-
and C China, Korea, Japan, Sumatra, Java, Borneo, the brown with black speckling, giving a mottled
Philippines, Sulawesi and many other Indonesian and appearance; underparts wholly silvery-grey, often
Philippine Islands (probably introduced in most island with a dark streak along the middle. Fur moderately
areas, including Sunda Shelf). Other forms of Rattus spiny. Ears large, lightly furred. Tail entirely dark
rattus group occur in much of the rest of the world. brown. Feet rather slender with relatively small
Status Least Concern. Considered a pest in most footpads (Fig. 72a), with inconspicuous patterns of
areas. ridges. Often has an orange-brown tuft of fur in front
of each ear, but this may fade in older individuals
MALAYSIAN WOOD RAT or dry skins. Mammae 3 + 3. Taxonomic notes
Rattus tiomanicus PLATE 72 Includes the recently described R. hoxaensis from
Measurements HB 140–190, T 150–200 Vietnam, which is now considered a synonym of
(95–120% of HB), HF 28–35. Wt 55–150g. this species. Similar species No other Rattus has
Skull: cbl 34–45, mt 6.0–8.0 a tuft of orange fur in front of ear, but this may
Identification Upperparts finely grizzled olive-brown; be missing in older animals. House Rat, R. rattus,
underparts usually pure white or slightly off-white. has long guard hairs on lower back, underparts
Fur smooth and sleek with short, stiff spines; black either pure white (not grey) or with grey base and
guard hairs short to moderate length distributed buff tip; Malaysian Wood Rat, R. tiomanicus, has
evenly through the pelage. Tail entirely dark brownish. sleeker, more uniform-coloured upperparts with less
Feet quite broad with well-developed pattern of fine prominent guard hairs, creamy white underparts,
ridges on footpads for climbing (similar to R. rattus, broader feet with well-developed ridges; Lesser
Fig. 72b). Ears large and thinly furred. Mammae 2 + Ricefield Rat, R. losea, averages slightly smaller with
3. Similar species House Rat, R. rattus, has longer smaller furry ears, upperparts softer orange-brown,
dark guard hairs on lower back, giving it a more underparts with grey base but white or cream tip.
shaggy appearance, and usually has darker base to Ecology and habitat Nocturnal. Largely associated
fur on underside; Ricefield Rat, R. argentiventer, has with open areas including rice fields, grassland and
grey-tinged underparts, orange tuft in front of ear, plantations. Mainly in lowlands but may extend to
dark speckling on back, more slender feet with less higher altitudes in areas of extensive cultivation.
developed pads; Annandale’s Rat, R. annandalei, has Active principally on the ground, but can climb some
softer, shaggy fur, without stiff spines, fewer mammae. trees. Burrows extensively and nests in holes in the
Ecology and habitat Nocturnal. Occurs in secondary ground. Diet includes many types of plant material,
and coastal forests, plantations, gardens, scrub and such as rice plants, grain, flowers and fruit of oil
grassland, but rarely in houses or tall dipterocarp palm, as well as insects. Can breed rapidly, producing
forests. Climbs well and spends much time in trees, several large litters per year when food is abundant.
but also often found on the ground. Diet includes Distribution SE Asia: Thailand, Laos, Vietnam,
a wide range of plant and animal matter, including Cambodia and Peninsular Malaysia. Also Sumatra,
oil palm fruits. Shelters in piles of cut palm fronds, Java, Borneo, Sulawesi, the Philippines, New Guinea
stumps, fallen logs and crowns of palms. and many intervening islands.
Distribution SE Asia: peninsular Thailand (S of Isthmus Status Least Concern. Widespread and abundant;
of Kra) and Peninsular Malaysia. Also Sumatra, Java, one of the major agricultural pests in the region.
Borneo and many smaller adjacent islands. Probably native to Indochina but introduced into
Status Least Concern. Very abundant in some areas many other areas with the spread of agriculture,
and considered a pest in oil palm plantations. especially rice cultivation.

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 PACIFIC RAT brown; underparts paler, grey to grey-brown, without
Rattus exulans PLATE 72 a sharp contrast; sometimes with a pale chest patch.
Measurements HB 90–140, T 105–160 (about Tail dark above, usually slightly paler below, but
110% of HB), HF 21–26, E 15–18. Wt 25–60g. sometimes mottled with pale patches; thinly covered
Skull: cbl 27–30, mt 4.5–6.0 with sparse hairs. Eyes and ears relatively small.
Identification Smallest Rattus. Upperparts usually Front and hind feet largely white. Footpads poorly
greyish-brown on mainland, but may be reddish- developed with low inconspicuous ridges. All-black
brown elsewhere; coarse spiny fur, spines white with melanistic forms occur in some areas. Mammae
dark brown tip. Underparts with cream or white tip 3 + 3. Similar species House Rat, R. rattus, is
and grey base. Facial whiskers very long, reaching smaller, with more mottled upperparts, flattened
beyond ears when folded back. Tail brown. Hind spines, longer and more uniformly dark tail, less
foot with elongate inner metatarsal pads with well- evenly coloured upperparts, larger footpads with fine
developed ridges (similar to Fig. 72b). Mammae 2 + ridges, larger ears; bandicoot rats, Bandicota spp.,
2. Similar species House Mouse, Mus castaneus, have larger ears, dark feet, broader incisors.
and relatives are smaller, with inner metatarsal pad Ecology and habitat Mostly nocturnal. Largely
short, rounded and near front of foot; House Rat, terrestrial. Builds large and complex burrow systems
R. rattus, is much larger when full-grown, with in some areas. Diet includes plant and animal
longer guard hairs, more mammae. material. Lives largely in towns and cities, especially
Ecology and habitat Diet includes plant and ports, where it feeds extensively on garbage and
animal material. In region, mainly found in gardens, waste and is considered an urban pest. Sometimes
secondary areas around houses, and inside houses; occurs in agricultural fields, but mostly near cities.
raids grain stores. Also sometimes enters rice fields, Distribution SE Asia: Myanmar, Thailand, Vietnam,
especially near buildings. In Pacific islands that lack Peninsular Malaysia, Singapore, possibly Laos and
some of the other rat species, occurs in many more Cambodia. Also most of the rest of the world, mainly
habitats and can be a major pest of many crops. in large cities. Probably native to Japan, Siberia and
Tends to grow larger on smaller islands. Climbs N China, and spread elsewhere by humans.
well, often ascending large grasses, small trees and Status Least Concern. Non-native in region and
houses. Makes nest of grasses in fields of tall grass considered a pest.
or thatched roofs of village houses.
Distribution SE Asia: C and S Myanmar, Thailand, INDOCHINESE FOREST RAT
Laos, Vietnam, Cambodia, Peninsular Malaysia and Rattus andamanensis PLATE 73
Singapore. Also Bangladesh, Taiwan, Sumatra, Java, Measurements HB 155–200, T 185–240 (115–
Borneo, Sulawesi and many smaller Indonesian 120% of HB), HF 32–37, E 23–25. Wt 100–150g.
Islands, the Philippines, New Guinea, New Zealand, Skull: gl 40–43, mt 6.5–7.5
Hawaii and other Pacific islands. Probably native Identification Upperparts orange-brown with
to mainland South-east Asia and spread to islands conspicuous long black guard hairs all along back,
either with humans or through natural rafting on giving a shaggy appearance. Underparts white or
drifting vegetation. cream, sometimes with red-brown patch on chest.
Status Least Concern. Widespread and common; Ears longest of any Rattus in region. Face whiskers
considered a pest in some areas. (vibrissae) long and thick, extending well behind
ears when folded back. Hind foot thickly haired with
NORWAY RAT mix of white and dark hairs on top; large, prominent
Rattus norvegicus PLATE 73 pads on soles of feet. Tail dark, sometimes with
Measurements HB 160–265, T 170–250 short white tip. In skull, auditory bullae relatively
(80–97% of HB), HF 35–50. Wt 200–500g. small. Mammae 3 + 3. Taxonomic notes Formerly
Skull: cbl 41–56, mt 6.8–8.8 R. sikkimensis, R. remotus or R. koratensis, but
Identification Large, mainly terrestrial rat. Fur recent research suggests all are the same, and
without flattened spines, but with moderately long R. andamanensis is the oldest name for the
guard hairs, giving shaggy apprearance. Upperparts species. Similar species House Rat, R rattus, has

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 proportionately shorter tail, smaller ears, guard hairs spines, medium-length black guard hairs protruding
mainly restricted to lower back, smaller footpads. no more than 10mm from fur. Upperparts brown to
Ecology and habitat Mostly nocturnal. A highly reddish-brown; underparts with grey base and white,
arboreal species found in a variety of forest and cream or buff tip. Tail shorter than body, dark above
forest edge habitats including native forest and and below. Some individuals in peninsular Thailand
disturbed forest, as well as clumps of large bamboo. are very dark brown. Feet narrow, footpads relatively
Has been found in gardens near forest. small (even smaller than in Fig. 72a), smooth, lacking
Distribution SE Asia: Myanmar, Thailand (N of striations; fur on feet varies from whitish-grey to
Isthmus of Kra except for some offshore islands), grey-brown. Mammae 2 + 3. Taxonomic notes
Laos, Vietnam and Cambodia, including some Recent research has shown this is a distinct species
offshore islands. Also Nepal, NE India, S China, from R. losea, which occurs alongside it in some
Andaman and Nicobar Islands. parts of Vietnam and Cambodia. Similar species
Status Least Concern. Most other Rattus are larger, with tail longer than
body; most species other than larger Ricefield Rat,
WHITE-FOOTED INDOCHINESE RAT R. argentiventer, have striated pads on soles of feet
Rattus nitidus PLATE 73 for climbing; Pacific Rat, R. exulans, is smaller with
Measurements HB 160–180, T 165–190 (100– long tail, fewer mammae; Osgood’s Rat, R. osgoodi,
105% of HB), HF 34–40, E 20–24. Wt 110–140g. is smaller with darker fur, shorter tail, shorter
Skull: gl 34–44, mt 6.0–7.2 toothrow, and is found higher in mountains; Losea
Identification Medium-sized with soft, woolly fur and Rat, R. losea, tends to have more grey-brown fur.
long, broad snout. Upperparts brown, belly creamy Ecology and habitat Mostly nocturnal. Grassy and
white with grey base to hairs. Hind and front feet, scrub habitats, mainly terrestrial in lowlands up to
including lower part of front legs, white. Narrow feet, about 850m altitude. Has been found in natural
but with large, well-developed, ridged footpads for grasslands in pine hill forest, and in disturbed areas.
climbing. Tail dark, slightly paler below. Mammae 3 Can be a significant pest in rice fields, where it
+ 3. Similar species House Rat, R. rattus, usually burrows and feeds on plants. Most common where
has dark tops to feet, paler coarser upperparts with R. argentiventer is uncommon or absent.
stiffer spines, broader feet; Indochinese Forest Rat, Distribution SE Asia: Thailand (N of peninsular),
R. andamanensis, has white-based fur on underparts, Laos, Vietnam and Cambodia.
longer tail, long guard hairs, dark tops to feet. Status Least Concern. Common in many areas and
Ecology and habitat Mostly nocturnal. In region, sometimes considered a pest.
mainly restricted to hills. In Thailand, found mainly
on hills, but also in fields. In S China, a major LOSEA RAT
agricultural pest where found in many fields. Rattus losea NOT ILLUSTRATED
Distribution SE Asia: Myanmar, N Thailand, Measurements HB 120–185, T 125–175,
Laos and Vietnam. Also Nepal, Bhutan, N India, HF 24–32, E 18–21.
Bangladesh and much of S China including Hainan, Identification Medium-small rat, generally similar
where probably native; the Philippines, Sulawesi and in appearance and size to Lesser Ricefield Rat, R.
New Guinea, where probably introduced. sakeratensis, with which it was formerly confused,
Status Least Concern. Considered a pest in some but differing genetically. Fur dull grey-brown mixed
parts of China. with pale and dark hairs; paler on sides blending
into buffy underparts with grey base, tail dark above,
LESSER RICEFIELD RAT slightly paler below. Taxonomic notes Recent
Rattus sakeratensis PLATE 72 research has shown R. sakeratensis is a distinct
Measurements HB 120–180, T 110–165 (typically species from R. losea, but further analyses are
c.90% of HB), HF 24–32, E 15–20. Wt 40–90g. needed to determine distinguishing characters in
Skull: gl 33–38, mt 6.3–7.1 areas where both species occur.
Identification Medium-small rat. Fur long, slightly Ecology and habitat Found in grassy areas and
shaggy but sometimes sleek, with soft transparent scrub, including cultivated fields.

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 Distribution SE Asia: Vietnam, Cambodia. Also Status Least Concern, although may be adversely
China. affected by conversion of forest cover to agriculture.
Status Least Concern.
Genus Bandicota Medium to large rats with
OSGOOD’S RAT long, shaggy fur with very long guard hairs, most
Rattus osgoodi PLATE 72 conspicuous on lower back. Distinguished externally
Measurements HB 125–170, T 100–140 (85% of from Rattus by broad upper incisors (combined
HB), HF 26–37. Skull: gl 31–36, mt 5.3–6.0 width more than 3.5mm), relatively small pads
Identification Fur dense, long and soft. Upperparts on feet and hence poor climbing ability. Cusps on
rich dark brown, sides slightly paler buffy-brown. molars largely fused to form nearly straight parallel
Underparts dark grey, with brownish wash in middle. ridges (Fig. 71h). Skull has large auditory bullae,
Ears and upper sides of feet dark brown. Tail dark long incisive foramina that extend back to level
brown. Mammae 2 + 3. Similar species Lesser with the front or middle of the first molar, long
Ricefield Rat, R. losea, has thinner, shaggier fur and bony palate that extends posterior to last molar.
paler underparts, and is larger with slightly longer tail, Mammae 3 + 3, but B. bengalensis varies from
longer toothrow; Pacific Rat, R. exulans, is smaller with 2 + 3 up to 3 + 7. Genus contains three species. An
greyer upperparts, spiny fur, pale grey underparts, additional species, B. bangchakensis, was described
long tail, fewer mammae; other Rattus are larger with in 1989 from Thailand, but is now considered to be a
longer tail, coarser fur, paler upperparts. synonym of B. savilei.
Ecology and habitat Mostly nocturnal. Found in
montane areas at 900–2,000m. Believed to be GREATER BANDICOOT RAT
mainly terrestrial in grasslands and low shrubs. Bandicota indica PLATE 73
Distribution SE Asia: S Vietnam, where known only Measurements HB 190–330, T 190–280,
from the Langbian plateau. HF 46–60, E 25–33. Wt 400–900g.
Status Least Concern. Has a limited range, but not Skull: gl 49–64, mt 10.7–12.4
currently considered at risk; surveys are required Identification Very large, dark rat with long, shaggy
to determine whether it occurs in other hill areas in fur. Upperparts dark blackish-brown, somewhat
the region. greyer along sides, with very long black guard
hairs. Underparts dark brownish-grey. Tail slightly
ANNANDALE’S RAT shorter than head and body, very dark brown with
Rattus annandalei PLATE 73 relatively large scales. Feet very large and broad,
Measurements HB 145–220, T 160–270 dark brown to blackish. Upper incisors very broad
(100–150% of HB), HF 35–40, E 18–23 (more than 4mm across the pair), angled straight
Identification Upperparts greyish-brown, under- down or slightly curved backwards. On skull, nasal
parts white or pale yellow. Fur soft and shaggy bones long, hiding nostrils and incisors from above.
without prominent spines. On skull, incisive foramina Mammae 3 + 3. Similar species Other Bandicota
short, anterior to first molar, auditory bullae relatively are substantially smaller and paler in colour, and can
large and inflated. Mammae 2 + 2. Similar species be distinguished from juvenile B. indica by shorter
Malaysian Wood Rat, R. tiomanicus, has smooth toothrows and narrower feet; Rattus are generally
fur with short, stiff spines, females have more smaller, with shorter feet; Müller’s Rat, Sundamys
mammae, skull with long incisive foramina; Müller’s muelleri, is browner with sharply demarcated pale
Rat, Sundamys muelleri, has even longer, shaggier underparts, tail longer than head and body, fewer
fur, averages larger. mammae in females, relatively narrow incisors.
Ecology and habitat Nocturnal. Secondary forest, Ecology and habitat Mostly nocturnal. Naturally
scrub and rubber plantations. Rarely found in primary occurs in wet swampy areas, but now exploits wet
forest. Found mainly low in trees, but sometimes rice fields in both upland and lowland areas. Usually
also on ground. Not known to be a significant pest. near human activity. Swims well. Builds extensive
Distribution SE Asia: Peninsular Malaysia and tunnel systems, sometimes in colonies with many
Singapore. Also Sumatra. adults and young. Feeds on plant materials as

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 well as invertebrates; food selection may vary with Upperparts with mixture of buff and dark brown hairs
population density. and moderately long black guard hairs, giving overall
Distribution SE Asia: Myanmar, Thailand, Laos, colour dark brownish-grey grizzled with black;
Vietnam and Cambodia. Introduced in parts of underparts grey sometimes tipped with buff. Tail all
Peninsular Malaysia. Also India, Nepal, Sri Lanka, dark brown; feet with dark brown or brownish-grey
Bangladesh, S China and Taiwan, and introduced in hairs, long narrow claws. Upper incisors broad with
Java. pale orange to creamy enamel; protruding forwards.
Status Least Concern; very common and considered On skull, nasal bones short, not hiding nostrils from
a pest in some areas; hunted for food. above. Mammae vary from 2 + 3 up to 3 + 7 or even
more in India; most commonly 7–9 pairs. Similar
SAVILE’S BANDICOOT RAT species Savile’s Bandicoot Rat, B. savilei, has more
Bandicota savilei PLATE 73 pointed snout, incisors angled backwards, longer,
Measurements HB 150–240, T 125–230, narrower feet.
HF 33–44, E 20–30. Wt 150–320g. Skull: 42–52, Ecology and habitat Mostly nocturnal. Found in
mt 8.6–10.0 villages and towns in rural areas, and associated
Identification Medium-sized. Moderately shaggy crop lands. A good swimmer, often found in wet
fur with long guard hairs; pelage softer than other rice fields. Constructs elaborate burrow systems.
Bandicota. Upperparts with mixture of black and Feeds on plants and grains as well as animal matter,
buff hairs, overall tone varying from brownish-grey including crustaceans and molluscs.
to reddish-brown; underparts greyish-buff, the hairs Distribution SE Asia: Myanmar. Introduced in
with grey base and buff tip. Feet grey. Tail slightly Penang in Peninsular Malaysia, and Phuket Island
shorter than head and body, uniformly grey-brown, in Thailand. Also native to Pakistan, India, Sri Lanka,
sometimes with a short white tip. Upper incisors Nepal and Bangladesh, and introduced in Sumatra,
straight down or slightly curved backwards. On skull, Java and Saudi Arabia.
nasal bones long, hiding nostrils and incisors from Status Least Concern. Considered a pest in many
above. Mammae 3 + 3. Similar species Greater areas.
Bandicoot Rat, B. indica, larger with broader feet,
larger teeth, even in juveniles; Lesser Bandicoot Genus Berylmys Medium to large rats with
Rat, B. bengalensis, has more upturned snout, paler smooth, stiff fur; upperparts steely-grey turning
forwards-projecting incisors, shorter, broader feet, slightly browner when worn; underparts pure
usually more mammae in females; Norway Rat, white, sharply demarcated from upperparts.
Rattus norvegicus, has white feet, narrower incisors. Incisors relatively wide and stout, usually with
Ecology and habitat Mostly nocturnal. Occurs in very pale enamel, white or cream to pale orange,
open degraded habitats including rice fields and though sometimes darker orange in B. bowersi or
corn fields, as well as edges of natural forest. Mainly B. mackenziei; in contrast, most other murid rodents
in lowland areas. have dark orange enamel on outside of incisors.
Distribution SE Asia: C Myanmar, Thailand, S Laos, Upper incisors either stick straight down or protrude
Vietnam and Cambodia. slightly forwards, not curved backwards. Molars
Status Least Concern; locally abundant and have some discrete cusps, but cusps tend to appear
considered a pest in some areas; also hunted for fused and are hard to differentiate when teeth are
food. worn; posterior molars small relative to others. Skull
with incisive foramina ending anterior to or level with
LESSER BANDICOOT RAT front molars; palate relatively short, ending anterior
Bandicota bengalensis NOT ILLUSTRATED to posterior molars. Braincase broadly triangular
Measurements HB 160–210, T 110–160, when viewed from above, rather than the more oval
HF 27–38, E 20–24, Wt 200–400g. shape of Rattus. Long, narrow hind feet with six
Skull: gl 36–44, mt 6.6–8.3 medium-sized pads lacking ridges. Mammae 2 + 2
Identification Medium-sized with moderately in B. bowersi, 3 + 2 in other species.
shaggy, harsh fur, blunt, slightly upturned snout.

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 BERDMORE’S RAT elsewhere; further study is needed to determine
Berylmys berdmorei PLATE 74 whether these populations are specifically distinct.
Measurements HB 190–260, T 150–190, Similar species Mackenzie’s Rat, B. mackenziei,
HF 37–46, E 22–29, Wt 180–300g. averages smaller and greyer, has more white on
Skull: gl 38–50, mt 6.6–8.0 tail in areas of overlap in Myanmar; females have
Identification Upperparts with smooth, stiff fur, steel- additional pair of post-axillary mammae; other
grey; underparts pure white, sharply demarcated Berylmys are considerably smaller.
from underparts. Tail dark brown on top and sides; Ecology and habitat Mostly nocturnal. Terrestrial,
dark brown or mottled pale greyish-white underneath; digging extensive burrow systems. Most specimens
lacking a white tip; considerably shorter than head and from 1,000–1,500m altitude, but has been
body. Front and hind feet white or grey above. Upper recorded from as low as 200 to as high as 1,600m.
incisors angled slightly forwards; auditory bullae In Malaysia, most common in primary forest, but
relatively larger than those of other Berylmys. Upper also found in disturbed forest and secondary scrub.
incisors 2.8–3.5 across. Similar species Other Feeds mainly on plant matter, including fruits; more
Berylmys have tail equal to or longer than head and rarely on insects and snails.
body, often with white tip; Bowers’s Rat, B. bowersi, is Distribution SE Asia: Myanmar, Thailand, Laos,
considerably larger, especially feet and skull. Vietnam and Peninsular Malaysia. Also NE India,
Ecology and habitat Mostly nocturnal. Mainly S China and NE Sumatra.
terrestrial. Digs extensive burrows. Found in forest Status Least Concern.
and scrub, but sometimes enters agricultural areas
near forest; reported from swampy forest and MACKENZIE’S RAT
marshy areas in Thailand. Found from sea level to Berylmys mackenziei NOT ILLUSTRATED
1,400m. In most areas infrequently found, but can Measurements HB 155–240, T 150–250,
be locally common. HF 44–51, E 25–35. Skull: gl 40–54, mt 8.1–9.8.
Distribution SE Asia: S Myanmar, Thailand, Laos, Specimens from S Vietnam average about 15%
S Vietnam, Cambodia. larger than those from Myanmar.
Status Least Concern; may occasionally be a pest in Identification Moderately large. Upperparts with
some agricultural areas. smooth, relatively thick fur, dark grey or steel-grey;
underparts pure white, sharply demarcated from
BOWERS’S RAT upperparts. Tail colour varies geographically: all dark
Berylmys bowersi PLATE 74 brown, sometimes with a short white tip in Vietnam;
Measurements HB 235–300, T 240–310, dark brown base with distal one-third to two-thirds
HF 52–61, E 27–37. Wt 270–650g. white in Myanmar. Feet white on sides and toes,
Skull: gl 51–63, mt 9.0–10.6 dark brown on top. Mammae 3 + 2. Upper incisors
Identification Upperparts with smooth, stiff fur, 3.0–3.9 across. Taxonomic notes Further study
colour varying from dark grey to bright brownish- is required to ascertain whether these apparently
grey with tan on sides; upperparts pure white, separate populations are really the same species.
sharply demarcated from underparts. Tail colour Similar species Bowers’s Rat, B. bowersi, averages
varies geographically: uniformly dark brown, larger, females with fewer mammae, less white on
sometimes slightly paler below in Laos and tail in Myanmar population.
Vietnam; dark brown at base with distal one-half Ecology and habitat Mostly nocturnal. Apparently
to one-third white in Malay Peninsula; dark brown occurs mainly at higher altitudes, 1,000–3,000m,
with short white tip in N Myanmar. Feet white on in wet evergreen or semi-deciduous montane forest.
sides and toes, dark brown on top. Mammae 2 Distribution SE Asia: C and S Myanmar, and
+ 2. Upper incisors 3.5–4.5 across. Taxonomic S Vietnam. Also NE India and S China (Sichuan).
notes Populations in peninsular Thailand, Malaysia Previously published records from Thailand have
and Sumatra are geographically isolated and differ been reidentified as B. bowersi.
in some ways from those elsewhere; specimens Status Data Deficient; further surveys are required
from N Myanmar and Laos are smaller than those to determine full extent of range.

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 MANIPUR RAT long, grey; lips unpigmented pinkish. Tail relatively
Berylmys manipulus PLATE 74 short with stubby tip, dark grey on top and sides
Measurements HB 135–185, T 140–185, for 60–75% of length, unpigmented pinkish-white
HF 33–40, E 23–25. Skull: gl 36–41, mt 6.2–6.7 on underside and all round at tip. Similar species
Identification Smallest of the Berylmys. Upperparts Lao Limestone Rat, Saxatilomys paulinae, is smaller
with smooth, stiff fur, steel-grey; underparts pure with longer tail, dark feet; Berylmys spp. have grey
white, sharply demarcated from upperparts. Tail upperparts, sharply contrasting white underparts,
dark brown at base, sometimes paler underneath, much smaller footpads; Leopoldamys spp. have
with distal one-half to two-thirds white. Front and contrasting white underparts, tail longer than head
hind feet white above. Upper incisors 2.3–2.9 and body; all species of Niviventer are smaller with
across, protrude slightly forwards. Similar species different fur colour, longer tail.
Berdmore’s Rat, B. berdmorei, is not known to Ecology and habitat Mostly nocturnal. Known only
overlap in range, has shorter tail without white tip; from limestone outcrops where it has been found
other Berylmys are larger. among vine-covered limestone boulders and inside
Ecology and habitat Mostly nocturnal. Strictly caves.
terrestrial, digging extensive burrows. Found in Distribution SE Asia: N Vietnam.
wooded and scrub habitats, as well as moist Status Data Deficient. Only known records are from
rainforest; not reported near human settlements. Huu Lien Nature Reserve in Vietnam, but probably
Feeds on plants, including leaves and seeds, as well occurs in other limestone areas in the region.
as on insects and other invertebrates. Altitudinal
range 100–1,500m. Genus Saxatilomys A recently discovered genus
Distribution SE Asia: N and C Myanmar. Also India with only one known species, first described in
and S China (Yunnan). 2005. Similar in many respects to Niviventer, it
Status Data Deficient, but has probably declined differs in colour pattern, narrow incisive foramina
following extensive loss of forest cover. that end anterior to the upper molars, long bony
palate that extends beyond molars, toothrows
Genus Tonkinomys This genus, with only one diverging posteriorly and other details of skull. Feet
known species, was first described in 2006. Feet with with large bulbous pads, lacking fine striations,
large bulbous pads lacking fine striations, apparently probably adapted for climbing on rocks (Fig. 73d);
adapted for climbing on limestone rocks (Fig. 73e); all toes with pointed claws. Incisors smooth on
all toes with pointed claws. Incisive foramina short surface, dark orange enamel on uppers, pale orange
and broad, bony palate long, extending far behind on lowers. Mammae 2 + 2.
molars. Incisors smooth with pale orange enamel.
Mammae 2 + 2. LAO LIMESTONE RAT
Saxatilomys paulinae PLATE 74
TONKIN LIMESTONE RAT Measurements HB 145–164, T 165–200 (110–
Tonkinomys daovantieni PLATE 74 130% of HB), HF 29–32, E 24–25. Wt 110–130g.
Measurements HB 185–215, T 155–185 (85% of Skull: gl 40–44, mt 7.3–7.9
HB), HF 37–41, E 29–31. Wt 140–205g. Identification Upperparts shiny greyish-black with
Skull: gl 47–53, mt 8.0–8.8 soft dark grey underfur, moderate-length flexible
Identification Upperparts slightly shaggy, shiny dark spines with pale grey base and dark tip, and long
greyish-black, often with a white blaze on forehead; guard hairs with pale base and black tip extending
fur a mixture of soft dark grey underfur, moderate- 5–10mm beyond underfur. Underparts lightly frosted
length flexible spines pale grey with dark tip, and dark grey, hairs uniformly grey with unpigmented tip,
long guard hairs with pale base and dark brownish- giving a slight frosting effect; blends into colour of
black tip. Underparts uniformly grey except for white upperparts. Feet covered with dark brown hairs.
patch on chest, colour blending into darker colour Tail long with slender tip, dark brown above and on
of upperparts. Feet whitish-pink except for brown sides, pale underneath, but not contrasting sharply.
patch in middle on tops of hind feet. Ears moderately Ears moderately long, grey; lips unpigmented

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 pinkish. Similar species Tonkin Limestone Rat, Ecology and habitat Mostly nocturnal and
Tonkinomys daovantieni, has similar colour and terrestrial, but can also climb. Often found near
footpads but is much larger, has relatively short tail streams. Feeds on plant and animal matter. Occurs
with white tip, usually has white patches on chest in lowland tropical forests, but also in some areas of
or forehead; Rattus spp. have smaller footpads with secondary forest near primary forest.
fine striations in arboreal species; Niviventer have Distribution SE Asia: peninsular Myanmar, Thailand
brown or brown-grey upperparts, contrasting white and Malaysia. Also Sumatra, Borneo, Palawan and
underparts. adjacent islands.
Ecology and habitat Mostly nocturnal. Known only Status Least Concern. However, populations have
from limestone rocky outcrops and surrounding likely declined in many areas due to loss of primary
forests. lowland rainforest.
Distribution SE Asia: C Laos, Vietnam.
Status Data Deficient; known only from a few Genus Niviventer Tail usually at least 125% of
individuals found by villagers or in owl pellets around head and body length (except in N. hinpoon), usually
Khammouane Limestone, and a few trapped at one with longer hairs at tip forming a slight tuft, often
site in Quang Binh, Vietnam. bicoloured. Fur varies in texture from soft to semi-
spiny, sometimes in the same species, and varies
Genus Sundamys Very large rats with long dark in length from short to moderately long; guard
tail. Shaggy fur with prominent long black guard hairs hairs moderately long and conspicuous in some
on upperparts. Skull with moderate-length, narrow species, inconspicuous in others. In most species,
incisive foramina ending near anterior molars; bony upperparts some shade of brown or reddish-brown,
palate ending slightly posterior to molars; relatively underparts pure white, but in preserved animals
small auditory bullae; large, stocky molars with white may discolour to bright yellow, especially
relatively large posterior molar, cusps similar in in formalin. Hind feet long and slender in most
shape to Rattus (Fig. 71g). Feet with relatively small species (Fig. 73c), but relatively broad in the most
pads. Several species in Greater Sundas, but only arboreal species, N. cremoriventer and C. langbianis
one in region. Mammae 2 + 2. (Fig. 73b). Hind feet with six well-developed pads,
somewhat larger in arboreal species. Incisors
MÜLLER’S RAT smooth with dark orange enamel, uppers angled
Sundamys muelleri PLATE 73 straight downwards or slightly backwards. Molars
Measurements HB 210–280, T 250–370 ( 110– relatively small, anterior molars narrow, third molar
120% of HB), HF 47–55, E 20–27. Wt 200–470g. much smaller than others (Fig. 71b). Skull long and
Skull: gl 53–62, mt 9.4–11.5 slender with long, narrow incisive foramina ending
Identification Very large with long dark tail. just in front of molars; bony palate ending roughly
Upperparts dark tawny-brown, darkest on top and level with back of molars (as in Fig. 69a). Auditory
paler on sides; fur harsh and slightly shaggy but bullae relatively small. Mammae usually 2 + 2,
without stiff spines, prominent long black guard sometimes 1 + 2. Taxonomic notes Taxonomy of
hairs 10–15mm beyond overfur; underparts group still quite uncertain, as many species can
with short dense fur, sharply demarcated from be difficult to distinguish. A recent genetic and
upperparts, usually white or pale grey, sometimes morphological analysis in Vietnam recognised six
darker grey with buff tinge. Tail longer than head species (excluding those now considered part of
and body, entirely dark brown. Ears round and Chiromyscus). However, the appropriate names for
dark brown. Similar species Most other rats some of these species have not been confirmed by
are smaller with shorter, sleeker fur; bandicoot comparison with specimens from the type localities
rats, Bandicota spp., have even longer, shaggier or elsewhere in SE Asia. Pending confirmation of
fur, underparts darker, not sharply demarcated these names, these additional species are mentioned
from upperparts, tail shorter than head and body, under ‘Taxonomic notes’ within the accounts for the
females with more mammae. traditionally recognised species.

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 DARK-TAILED NIVIVENTER Identification Upperparts bright reddish-brown,
Niviventer cremoriventer PLATE 75 fur with many flexible spines and long black guard
Measurements HB 130–165, T 150–200 (usually hairs, as in N. cremoriventer. Underparts white. Tail
about 125% of HB), HF 20–29, E 15–21. very long, dark brown above and all around at base,
Wt 53–100g. Skull: gl 33–38, mt 5.5–6.4 whitish on remainder of underside and all around
Identification Upperparts reddish-brown to orange- at tip. Mammae 2 + 2. Taxonomic notes Formerly
brown, darker on back and rump, sometimes with called N. rapit, but that species is now considered to
scattered patches of white hairs. Fur a mixture of four occur only in Borneo. Similar species Dark-tailed
types of hairs: soft grey underfur with dull orange- Niviventer, N. cremoriventer, has smaller hind feet,
brown tip; longer overfur with grey base and bright all-dark tail; Indomalayan Niviventer, N. fulvescens,
orange-brown tip; similar-length stiff, flattened is smaller with yellowish-brown upperparts.
spines, translucent at base and tipped with black Ecology and habitat Mostly nocturnal. Occurs in
on back and rump, orange-brown on sides; and forests above 1,000m where generally found on the
long black-tipped guard hairs extending 15–20mm ground.
beyond overfur. Underparts whitish or creamy, also Distribution SE Asia: known only from Cameron
with a mixture of flexible spines and softer hairs, Highlands, Peninsular Malaysia.
all hair types white to base; sometimes turns Status Vulnerable; has very limited range, and
yellowish, especially in older skins. Short face with population likely to be declining due to conversion
relatively large ears and long facial whiskers. Feet of montane forest to agriculture and tea plantations.
dark brown above, relatively short and broad with
large footpads (as in Fig. 73b). Tail usually entirely CONFUCIAN NIVIVENTER
dark brown, but sometimes paler below (especially Niviventer confucianus PLATE 75
in Borneo); covered in short hairs with longer, more Measurements HB 125–170, T 150–220 (125–
prominent hairs on the distal 20mm forming a slight 135% of HB), HF 28–32, E 20–23. Skull: gl 34–41,
tuft extending 5–8mm beyond tip. Mammae 2+2. mt 5.6–5.9
Similar species Cameron Highlands Niviventer, Identification Upperparts brown, sometimes
N. cameroni, is larger (especially hind foot), tail with yellowish or greyish tinge; fur long and soft,
pale underneath without tuft of hairs; Indomalayan usually without stiff spines or conspicuous guard
Niviventer, N. fulvescens, has bicoloured tail and, in hairs. Underparts white, sharply demarcated from
areas of overlap in range, is more yellowish-brown. upperparts. Tail moderately long, dark brown
Ecology and habitat Nocturnal. A good climber, above, pale below, sometimes pale all around at
active in small trees, lianas and shrubs to at least tip; short tuft of hairs (c.5mm) at tip. Mammae
5m above ground; also active on ground. Occurs in 2 + 2. Taxonomic notes Formerly considered
tall and secondary forests, forest edge and lightly part of N. niviventer from Nepal. Recent genetic
wooded areas from near sea level to 1,900m. Feeds analyses suggest that N. confucianus and
on plant matter including fruits and seeds, as well N. tenaster are both part of a group of at least four
as insects. closely related species in Vietnam that may include
Distribution SE Asia: peninsular Thailand and N. niviventer. These four species differ in size (two
Peninsular Malaysia. Also Sumatra, Java, Borneo are much smaller than others), relative tail length
and adjacent smaller islands. and colour (reddish vs. more yellowish). However,
Status Least Concern. However, population has more information from the type localities and
probably declined substantially owing to loss of elsewhere in the range is needed to determine the
lowland forest. appropriate names for each species, and whether
size or colour vary geographically. Similar species
CAMERON HIGHLANDS NIVIVENTER Other Niviventer in its range have coarser fur with
Niviventer cameroni PLATE 75 stiff spines and conspicuous guard hairs, but this
Measurements HB 130–170, T 205–270 (130– is somewhat variable; can be difficult to distinguish
180% of HB), HF 30–38, E 18–25. Wt 58–129g. from Indomalayan Niviventer, N. fulvescens.
Skull: 40–43, mt 6.3–7.2 Ecology and habitat Mostly nocturnal. In region,

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 found at high elevations in moss forest, but in China analyses indicate N. fulvescens is one of two closely
occurs in a variety of habitats including disturbed related but distinct species in Vietnam. One is more
and secondary forest. reddish with a longer tail (140–150% of HB) that
Distribution SE Asia: N Myanmar, NW Thailand and sometimes has an all-dark or all-pale tip and is
NW Vietnam. Also China. primarily montane. The other is more yellowish-
Status Least Concern, but only a few records from brown, with a shorter tail (120–130% of HB) that
Thailand and Vietnam. is bicoloured to the tip and widely distributed in
the lowlands. The first has been tentatively called
INDOCHINESE MOUNTAIN NIVIVENTER N. fulvescens (originally described from Nepal)
Niviventer tenaster NOT ILLUSTRATED and the latter N. huang (originally described from
Measurements HB 120–190, T 175–235 (125– China); however, comparison with type material is
140% of HB), HF 32–35, E 23–26. Wt 50–140g. needed to confirm these names. Populations from
Skull: gl 37–42, mt 5.9–6.5 S Thailand, Peninsular Malaysia and Sunda shelf are
Identification Upperparts yellowish-brown with often separated as N. bukit, but genetic analyses
conspicuous dark brown guard hairs. Underparts of Malaysian specimens are not yet available to
white, sharply demarcated from upperparts. Tail determine their relationships. Similar species
dark brown above, paler below, often with a pale Dark-tailed Niviventer, N. cremoriventer, has all-
tip. Ears relatively long. Mammae 2 + 2. Taxonomic dark tail; Confucian Niviventer, N. confucianus,
notes See comments under N. confucianus. Similar tends to have less spiny fur, but this is somewhat
species Indochinese Tree Rat, C. langbianis, has all- variable and can be difficult to distinguish;
dark tail; Indomalayan Niviventer, N. fulvescens, in Indochinese Mountain Niviventer, N. tenaster, is
areas of overlap in range, is more reddish-brown. more yellowish-brown with longer ears.
Ecology and habitat Mostly nocturnal. Hill forest, Ecology and habitat Mostly nocturnal. Found in a
from 1,000–2,200m. Mainly active on the ground, wide variety of forested habitats as well as gardens
low vines and fallen logs. and disturbed areas with vegetation. Active both on
Distribution SE Asia: C and S Myanmar, NW ground and in trees, frequently climbing large lianas
Thailand, Laos, Vietnam and SW Cambodia. Also and vines.
China (Yunnan and Hainan). Distribution SE Asia: Myanmar, Thailand, Laos,
Status Least Concern. Vietnam, Cambodia and N Peninsular Malaysia.
Also Nepal, N India, Bangladesh, S China (including
INDOMALAYAN NIVIVENTER Hainan, Hong Kong), Sumatra, Java and Bali.
Niviventer fulvescens PLATE 75 Status Least Concern; relatively common in many
Measurements HB 130–170, T 155–220 (115– areas.
150% of HB), HF 25–35, E 18–21. Wt 60–135g.
Skull: gl 31–42, mt 5.4–6.4 LIMESTONE NIVIVENTER
Identification Upperparts vary from yellowish- Niviventer hinpoon PLATE 75
brown to reddish-brown, hairs with a mixture of Measurements HB 120–160, T 120–160 (c.100%
yellowish or reddish overfur, dark brown-tipped of HB), HF 25–28, E 17–21. Wt 50–70g.
spines and conspicuous black guard hairs on Skull: gl 34–37
back. Underparts pure white, sometimes slightly Identification Upperparts dull buffy-grey, hairs
yellow, sharply demarcated on sides; also with mixed with extensive spines; underparts dull buff,
extensive short, stiff white spines. Tail very long, hairs with grey base and buff tip. Tail similar in
dark brown above, white below, sometimes with length to head and body, sharply bicoloured dark
all-pale or all-dark tip, no tuft of hairs at tip. Top above, pale below for entire length. Mammae 2 + 2.
of hind foot silvery grey to pale brown, sometimes Similar species Other Niviventer in its range have
with dark brown patch in middle. Colour varies white underparts; Rattus spp. have a dark tail.
geographically, with Malaysian individuals more Ecology and habitat Mostly nocturnal. Forested
yellow-brown. Feet relatively long (Fig. 73c). limestone outcrops, in scrubby forest at base of
Mammae 2 + 2. Taxonomic notes Recent genetic cliffs, as well as inside caves.

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 Distribution SE Asia: known only from limestone larger and darker, lacks ear tuft, has longer tuft of
karst in C Thailand. hair on tail; other Niviventer in range have white
Status Endangered; limited range and loss of habitat bellies with stiff spines in pelage.
due to forest clearing and limestone quarrying. Ecology and habitat Mostly nocturnal. Mainly
terrestrial. Recorded from scrub and wet montane
BRAHMAN NIVIVENTER temperate forests around areas of boulders at
Niviventer brahma PLATE 75 2,400–3,300m altitude.
Measurements HB 135–145, T 200–225 (150% Distribution SE Asia: N Myanmar. Also Nepal, India
of HB), HF 28–33, 20–25. Skull: gl 34–36, and S China.
mt 6.4–7.0 Status Least Concern, though poorly known in region.
Identification Medium-sized with very long tail. Fur
relatively long (15mm above, 10mm on underparts), Genus Chiromyscus Similar to Niviventer but
dense and soft without stiff spines. Upperparts distinguished by relatively short feet with large
bright orange-brown to yellow-brown; dark brown pads, and shortened first hind toe (hallux), with
ring c.2mm around each eye, connected across top either rounded nail or very small claw (Fig.
of muzzle by narrow dark brown band. Underparts 73a, b). Taxonomic notes Recent genetic and
greyish-white, individual hairs with grey base and morphological analyses in Vietnam have shown that
distal half white. Ears dark brown with short hairs. this genus includes at least three species, rather
Tail very long, brownish-black above, brownish-grey than just one. The holotype of C. chiropus (from
below, well haired, hairs longer towards tip forming Myanmar) was found not to be the same as the
distinct brush 6–8mm long. Hind feet silver-grey on species with a conspicuous black eye-ring, that was
sides, grey-brown on top. Mammae 1 + 2. Similar traditionally associated with the name; that species
species Smoke-bellied Niviventer, N. eha, is paler has now been given the name C. thomasi. The name
brown, eye-rings do not connect over muzzle, has C. chiropus is now being used for a different species
tuft of hairs at base of ear; other Niviventer in its from Vietnam, based on similarities in appearance
range have all-white belly, stiff spines in fur. with the holotype. Further analyses of additional
Ecology and habitat Mostly nocturnal. Reported specimens, preferably with genetic data, are needed
from wet montane broadleaf forest at 2,500m to confirm that the Myanmar and Vietnamese
altitude. Found on the ground and fallen logs. populations are actually the same species. In
Distribution SE Asia: N Myanmar. Also NE India and addition, the species formerly called Niviventer
China (NW Yunnan). langbianus has been shown to be closely related,
Status Least Concern. and also belongs in Chiropus. Mammae 2 + 2.

SMOKE-BELLIED NIVIVENTER FEA’S TREE RAT

Niviventer eha PLATE 75 Chiromyscus chiropus NOT ILLUSTRATED
Measurements HB 110–130, T 165–195 (150% Measurements HB 122–148, T 200–225,
of HB), HF 27–31, E 17–20. Skull: gl 30–33, (128–148% HB). Skull: gl 36–41, mt 5.6–7.5
mt 5.1–5.6 Identification Upperparts orange-brown with
Identification Fur thick and soft. Upperparts conspicuous black guard hairs along back; cheeks
pale brownish-orange; dark rings around eye but and parts of chest light orange; rest of underparts
without a dark mask-like mark joining to other eye. white, separated by a narrow band of light orange
Underparts light grey, individual hairs with grey base from upperparts. Feet and toes largely unpigmented,
and distal one-third white. Ears dark brown, densely with orange or white hairs on top. Tail all dark.
covered with long hairs, and a long brownish-black First hind toe shortened with rounded nail instead
tuft of hairs at base of each ear. Hind feet silver-grey of pointed claw. Taxonomic notes The species
on sides, grey-brown on top. Tail very long, brownish- previously associated with this name is now called
black above, grey below, with moderate-length hairs C. thomasi – see notes under genus name for
forming short tuft at tip. Mammae 1 + 2. Similar more details. Similar species Thomas’s Tree Rat,
species Brahman Niviventer, N. brahma, averages C. thomasi, averages larger, has brighter orange

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 around face and rump and conspicuous black eye- INDOCHINESE TREE RAT
ring; Indochinese Tree Rat, C. langbianus, also has Chiromyscus langbianis NOT ILLUSTRATED
reduced first toe with very small claw, but claw is Measurements HB 125–150, T 150–200 (c.145%
distinctly pointed (Fig. 73b), tail is all dark, back of HB), HF 27–32, E 20–25. Skull: gl 33–40,
has extensive long black guard hairs, face is not as mt 6.0–6.7
orange. Identification Upperparts brownish-grey mixed with
Ecology and habitat Nocturnal. Arboreal with feet pale yellowish-orange with distinctive olive-grey
well adapted for climbing; found in a variety of forest tone; darker on back and lower rump owing to more
types including moist deciduous and evergreen. May long black hairs; paler on sides, with yellow-orange
use degraded forest, but not open areas. at lower edge of flanks. Underparts contrasting white
Distribution SE Asia: E Myanmar, Vietnam. Probably or cream. Fur a mixture of softer hairs, spines and
also Thailand, Laos. long guard hairs, as in N. cremoriventer, but often
Status Not yet assessed under current species somewhat softer with fewer spines. Cheeks, sides of
definition. neck and upper arms often brighter yellow-orange,
contrasting with back. Tail very long, uniformly dark
THOMAS’S TREE RAT brown with tuft of hairs at tip. Ears relatively large,
Chiromyscus thomasi PLATE 75 dark brown. Feet relatively short and broad with large
Measurements HB 145–180, T 200–230 (128– footpads (Fig. 73b). Skull has long incisive foramina
132% HB), HF 27–29, E 18–20. Skull: gl 41–43, and relatively large auditory bullae. Mammae 2 + 2.
mt 7.0–8.0 Taxonomic notes Formerly considered in the genus
Identification Upperparts orange-brown, with light Niviventer. Similar species Dark-tailed Niviventer,
brown guard hairs; sides of fore and hind legs, N. cremoriventer, is similar in shape and size but
rump and face up to behind ears relatively bright more reddish-brown with smaller ears, does not
orange with conspicuous blackish ring around overlap in range; Thomas’s Tree Rat, C. thomasi, is
eye; underparts white, sharply demarcated from similar in size and shape but has brighter orange on
upperparts. Feet and toes largely unpigmented, but face with dark eye-ring, rounded nail on first toe,
with orange hairs on top. Tail bicoloured, dark above bicoloured tail.
and pinkish-grey below. First hind toe shortened with Ecology and habitat Mostly nocturnal. Tropical
rounded nail instead of pointed claw (as in Fig. 73a). evergreen forest as well as mixed forest, at altitudes
Taxonomic notes Newly named in 2014; species from near sea level to 2,800m. Spends much time
had previously been considered part of C. chiropus, in trees and vines, but also sometimes found on the
which is now recognised as the appropriate name ground.
for a different species. Similar species Fea’s Tree Distribution SE Asia: Myanmar, Thailand (N of
Rat, C. chiropus, is smaller, with black guard hairs, Isthmus of Kra), Laos, Vietnam and SW Cambodia.
all-dark tail; Indochinese Tree Rat, C. langbianus, Also NE India.
also has reduced first toe with very small claw, but Status Least Concern, but probably declining owing
claw is distinctly pointed (Fig. 73b), tail is all dark, to loss of forest.
back has extensive long black guard hairs and is
not as orange. Genus Dacnomys Large rats, somewhat similar
Ecology and habitat Nocturnal. Arboreal with feet to Niviventer or Leopoldamys in skull morphology.
well adapted for climbing; found in a variety of forest Auditory bullae relatively very small; incisive
types including moist deciduous and evergreen. May foramina long, extending past anterior face of
use degraded forest, but not open areas. anterior molar; teeth wide and chunky with very high
Distribution SE Asia: N Laos, Vietnam. Possibly crowns; toothrow proportionately and absolutely
Thailand and S China. very long (>20% of skull length); cusps on teeth
Status Not yet assessed; newly described as angular and partly fused into V-shaped ridges (Fig.
distinct from C. chiropus. 71i). Upper incisors orange. Mammae 2 + 2.

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 MILLARD’S GIANT RAT Identification Large, long-tailed rat. Fur sleek and
Dacnomys millardi PLATE 73 smooth. Upperparts buffy-brown to orange-brown,
Measurements HB 230–290, T 260–360 (115– darker brown in middle of back, brighter orange-brown
140% of HB), HF 50–56. Skull: gl 53–61, on sides; underparts creamy white. Tail dark all around
mt 11–12.5 at base and on top for most of length, pale underneath
Identification Very large. Fur thick, short, lacking and at tip; pattern of dark and pale areas irregular and
stiff spines. Upperparts brown flecked with buff; somewhat variable. Juveniles plain light brown above.
underparts whitish, varying from creamy to light Taxonomic notes Populations elsewhere in Thailand
grey with buff tinge. Feet large and brown. Long and Indochina now considered a distinct species,
tail all dark brown. Similar species Most other rats L. herberti. Sometimes called L. vociferans, with
are smaller, with much smaller toothrows; Edward’s the name L. sabanus restricted to Borneo. Similar
Giant Rat, Leopoldamys edwardsi, has more uniform species Other Leopoldamys have dark brown or grey-
brown upperparts without pale flecks, extensive brown upperparts, without contrasting orange flanks.
short spines in fur, smaller teeth. Ecology and habitat Nocturnal. Mainly terrestrial and
Ecology and habitat Mostly nocturnal. Recorded in lower parts of trees up to at least 3m above the
only in upper montane rainforest. ground. Occurs in tall and secondary forests, mainly
Distribution SE Asia: N Laos and NW Vietnam. in lowlands up to 1,200m in Peninsular Malaysia.
Also NE India, E Nepal and S China. Might also be Distribution SE Asia: Peninsular Thailand (north
expected in N Myanmar. to Kanchanaburi) and Malaysia. Also Sumatra and
Status Data Deficient; known from widely scattered Borneo (although those found here are sometimes
localities but with very few records, so status and considered distinct species).
requirements poorly understood. Status Least Concern; very common in some
forested areas.
Genus Leopoldamys Large rats with a very long
tail; sleek, smooth fur, guard hairs only slightly HERBERT’S GIANT RAT
longer than rest of fur, numerous spines that are soft Leopoldamys herberti NOT ILLUSTRATED
and hair-like; upperparts dark, colour varying with Measurements HB 200–260, T 300–370 (at least
species, sharply demarcated from pale underparts; 135% of HB), HF 42–48. Skull: gl 50–58,
tail mixed dark and pale. In skull, incisive foramina mt 8.9–10.2
relatively short, ending in front of molars; auditory Identification Large, long-tailed rat. Upperparts
bullae relatively small; posterior edge of bony buffy-brown to orange-brown, with extensive black
palate level with end of toothrow (Fig. 69c). Large, streaking, especially on back; lighter brown on
strong incisors with bright orange enamel. Cusps sides; underparts creamy white. White on upper
on molars partly fused into V-shaped ridges (Fig. lip reaches to vibrissae or eye-ring. Tail dark above
71f). Taxonomic notes Recent genetic studies have and about halfway along, pale underneath and at
shown this genus is more diverse than previously distal half; pattern of dark and pale areas irregular
thought. Currently six species are recognised in and somewhat variable. Taxonomic notes Formerly
the region, with several changes to previously considered part of L. sabanus, but recent genetic
recognised names. However, populations in many analyses have shown it is quite distinct from both
areas have not yet been sampled genetically, so L. sabanus and L. ciliatus. Similar species Long-
further changes may be expected. Some species are tailed Giant Rat, L. sabanus, is more orange and
difficult to distinguish, but generally only one or two does not overlap in range. Neill’s Rat, L. neilli,
species occur in any given area. Mammae 2 + 2. averages smaller, is darker brown with relatively
shorter tail, brown sides of face.
LONG-TAILED GIANT RAT Ecology and habitat Nocturnal. Mainly terrestrial
Leopoldamys sabanus PLATE 76 and in lower parts of trees up to at least 3m above
Measurements HB 200–275, T 270–415 (at least the ground.
135% of HB), HF 42–52. Wt 250–500g. Distribution SE Asia: Thailand (excluding peninsula),
Skull: cbl 51–60, mt 8.7–10.3 Laos, Vietnam, Cambodia.

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 Status Least Concern (as part of L. sabanus). Not Measurements HB 225–280, T 260–340
yet assessed separately. (110–120% HB). Wt 350–500g. Skull: gl 56–62,
mt 9.0–10.2
SUNDAIC MOUNTAIN RAT Identification Similar to L. edwardsi, but with very
Leopoldamys ciliatus NOT ILLUSTRATED dark grey-brown upperparts. Tail generally all dark,
Measurements HB 215–255, T 300–390 (120– relatively short; skull has relatively large auditory
165% of HB), HF 45–55, E 25–32. Wt 300–425g. bullae. Taxonomic notes Recent genetic research has
Skull: cbl 54–63, mt 9.5–10.8 confirmed this is a distinct species. Similar species
Identification Large, long-tailed rat. Upperparts and Other Leopoldamys have paler brown upperparts.
flanks dull dark brown, underparts white, often with Ecology and habitat Mostly nocturnal. Hill forest
dark patch on chest between front legs. Tail usually above 1,500m.
all dark. Taxonomic notes Formerly considered part Distribution SE Asia: known only from Langbian
of L. edwardsi. Similar species Long-tailed Giant highlands in S Vietnam.
Rat, L. sabanus, has contrasting orange flanks, lacks Status Least Concern as no apparent threats,
dark patch on chest, usually found at lower altitude; but further research is needed to understand
other Leopoldamys do not overlap in range. distribution limits.
Ecology and habitat Mostly nocturnal. Montane
forest, usually above 1,000m. Probably mainly NEILL’S RAT
terrestrial and low in trees and shrubs. Leopoldamys neilli PLATE 76
Distribution SE Asia: Peninsular Malaysia. Also Measurements HB 200–235, T 240–300,
Sumatra. HF 39–45, E 25–29. Wt 220g. Skull: gl 50–62,
Status Least Concern, though may be declining mt 8.8–10.3
somewhat due to loss of hill forest, especially in Identification Upperparts mixed black and buffy-
Sumatra. brown; underparts plain white. Sides of face brown.
Tail dark basally and along top for two-thirds of
EDWARD’S GIANT RAT length, white at tip and underneath. Feet pale with
Leopoldamys edwardsi PLATE 76 dusky line on top. Juvenile plain grey above. Similar
Measurements HB 210–280, T 290–360, HF 46– species Edward’s Rat, L. edwardsi, is substantially
54, E 29–33. Wt 340g. Skull: gl 51–59, mt 8.6–9.7 larger; Herbert’s Rat, L. herberti, is more orange
Identification Upperparts uniform, plain brown, with more white on face; Tonkin Limestone Rat,
slightly darker in front of eye; underparts sharply Tonkinomys daovantieni, has short tail, dark grey
demarcated white. Tail mainly dark, but with some underparts.
white hairs on underside near tip. Taxonomic notes Ecology and habitat Mostly nocturnal. Most records
Recent genetic studies suggest species is restricted are from limestone outcrops, where it climbs among
to northern part of region. Similar species Herbert’s rocks and caves; also reported from some areas of
Long-tailed Giant Rat, L. herberti, is darker in middle lowland bamboo scrub near limestone.
of back with contrasting orange sides, more white Distribution SE Asia: N and SW Thailand, Laos,
on upper lip; Neill’s Rat, L. neilli, is distinctly smaller Vietnam.
with mottled upperparts; Sundaic Mountain Rat, L. Status Least Concern; relatively common in some
ciliatus, does not overlap in range. localities.
Ecology and habitat Mostly nocturnal, primarily
montane, separating altitudinally from L. sabanus. Genus Diomys This genus has only one recognised
Distribution SE Asia: N Myanmar, N Thailand (only species. Thumb small, reduced to a small pad. Soles of
in Phu Kadeung plateau), N Laos and N Vietnam. front feet unpigmented, with three pads between toes
Also NW India, and S and C China. and two behind them. Hind feet with greatly reduced
Status Least Concern. pads; four moderately small pads between base of
toes and only one small metatarsal pad in middle of
MILLET’S GIANT RAT foot; lacking second elongate inner metatarsal pad
Leopoldamys milleti NOT ILLUSTRATED found in most other rats (e.g. Fig. 72 or 73). Fifth toe

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 relatively short, similar in length to first toe. Upper Identification Fur soft but relatively short.
incisors relatively pale, slightly protruding forwards, Upperparts light brown to greyish; underparts white
often faintly grooved. In skull, incisive foramina long or pale greyish-white. Hands and feet white. Tail
and thin, extending to level of middle of anterior distinctly bicoloured, dark above, whitish below,
molars; bony palate extends 0.5–1.0mm posterior of sometimes with white tip, distinctly shorter than
molars. Possibly closely related to Millardia. head and body. Upper incisors narrow, yellow and
plain; auditory bullae relatively large, about 20%
CRUMP’S SOFT-FURRED RAT length of skull. Only four pads on soles of feet.
Diomys crumpi PLATE 74 Similar species Crump’s Soft-furred Rat, Diomys
Measurements HB 100–135, T 110–135 (100– crumpi, is slightly smaller with darker upperparts,
105% of HB), HF 24–28, E 20–26. Skull: gl 29–33, longer fur, relatively longer tail; most other rats
mt 5.2–5.6 with bicoloured tail have stiff spines in fur, better
Identification Relatively small. Fur soft, silky developed plantar footpads.
and short (10mm long on rump). Upperparts dark Ecology and habitat Mostly nocturnal. Terrestrial.
blackish-grey to brownish-grey; underparts greyish- Found in dry areas of C Myanmar, where it has
white, with grey base and white tip. Tail sharply been recorded in scrubby areas with sandy or stony
bicoloured, dark brown to blackish above, greyish- ground. Presumably burrows in ground.
white below. Front and hind feet white, contrasting Distribution SE Asia: known only from C Myanmar.
with dark body. Ears relatively large, dark brown. Status Least Concern; apparently locally abundant
Incisor breadth 2mm. Similar species White- in some areas.
toothed rats, Berylmys spp., are generally larger
with stiff, spiny fur, tail all dark or with pale tip. Popa Genus Maxomys Tail bicoloured, similar in length
Soft-furred Rat, Millardia kathleenae, has paler to head and body. Fur short and dense, without
upperparts, averages slightly larger, with tail usually conspicuous long guard hairs. Most species in the
shorter than head and body. region have extensive stiff, prominent spines on
Ecology and habitat Mostly nocturnal. Terrestrial. upperparts, but one species has soft fur. Hind feet
Burrows in the ground. Has been reported from long and narrow, with relatively small footpads,
areas of tall grass in N Myanmar. smooth and not well adapted for climbing; only five
Distribution SE Asia: N Myanmar. Also NE India and pads, lacking outer metatarsal pad in most species;
SW Nepal. if pad is present, it is small and rounded. Upper
Status Data Deficient; only one record from the incisors orange, not protruding forwards; molars
region, and poorly known elsewhere. relatively wide. Skull has relatively broad braincase
with elongate rostrum; short, wide incisive foramina
Genus Millardia A genus of primarily Indian rats, that end well in front of toothrow, posterior margin
of which one species is found in the region. These of palate slightly in front of back edge of posterior
rats are mainly terrestrial, found in dry areas. Fur is molars. Mammae 2 + 2 (but some species outside
generally soft. Hind feet have greatly reduced fifth region have 1 + 2).
toe, similar to Dacnomys. Most species have only
four or five small plantar footpads. In skull, incisive RAJAH MAXOMYS
foramina long and thin, extending to level of middle Maxomys rajah PLATE 76
of anterior molars; bony palate relatively long. Measurements HB 165–225, T 160–210
Mammae 0 + 2 in only species in the region (but 1 (90–95% of HB), HF 35–43. Wt 95–220g.
+ 2 or 2 + 2 in other recognised species elsewhere). Skull: gl 41–49, mt 6.9–8.1
Identification Upperparts brown, sometimes tinged
POPA SOFT-FURRED RAT reddish or orange, darker in the midline and with
Millardia kathleenae PLATE 74 numerous stiff grey-brown spines; underparts white,
Measurements HB 130–165, T 120–155 (90– with many short, soft white spines, usually with dark
95% of HB), HF 25–30, E 20–23. Skull: gl 34–36, brown streak along middle in adults. White on inner
mt 5.5–6.0 sides of thighs normally extends unbroken to feet.

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 Tail sharply bicoloured, dark brown above, pale WHITEHEAD’S MAXOMYS
below, thinly haired. Similar species Red Spiny Maxomys whiteheadi PLATE 76
Maxomys, M. surifer, is brighter reddish, lacks dark Measurements HB 105–150, T 90–125 (85–90%
streak on underparts, usually has orange collar of HB), HF 22–30. Wt 35–80g. Skull: gl 29–34,
under neck and orange band around leg; immatures mt 5.1–6.2
of both species are very difficult to distinguish from Identification Small and spiny; upperparts reddish-
each other. brown to dark brown with grey underfur, brown-
Ecology and habitat Nocturnal and predominantly tipped overfur and numerous stiff pale grey spines
terrestrial. Occurs in tall and secondary lowland with black tip; underparts grey to orange-grey, the
forests, possibly in drier areas than M. surifer; the hairs with grey base and orange-buff tip mixed with
two species have rarely been found in the same numerous soft, pale spines; tail dark above, pale
locations. below. Hind feet with only five pads. Similar species
Distribution SE Asia: peninsular Thailand and Malayan Mountain Maxomys, M. inas, is bigger, with
Malaysia. Also Sumatra, Borneo and adjacent longer tail; Niviventer spp. are larger with tail longer
smaller islands. than head and body, six large footpads, white belly;
Status Vulnerable; Believed to have declined Pacific Rat, Rattus exulans, has entirely dark tail
considerably owing to extensive loss of lowland significantly longer than head and body.
forest, although still common in remaining areas. Ecology and habitat Nocturnal. Mainly terrestrial.
Diet includes ants and other insects as well as plant
RED SPINY MAXOMYS matter. Occurs in tall and secondary forests, mainly
Maxomys surifer PLATE 76 in lowlands, including plantations and disturbed
Measurements HB 160–210, T 155–210 (95– areas, but only if adjacent to forest.
100% of HB), HF 38–47, E 22–28. Wt 125–285g. Distribution SE Asia: peninsular Thailand and
Skull: gl 40–48, mt 6.0–7.5 Malaysia. Also Sumatra, Borneo and adjacent
Identification Upperparts distinctly orange- or islands.
reddish-brown, slightly darker along midline, with Status Vulnerable. Populations have declined
numerous short, stiff, dark spines; underparts considerably owing to loss of lowland forest in many
white with soft white spines. Orange-brown usually areas.
extends around part or all of underside of neck,
forming a collar, and around inner side of front leg MALAYAN MOUNTAIN MAXOMYS
above ankle. Tail bicoloured. Taxonomic notes Maxomys inas PLATE 76
Considerable geographic variation in size and Measurements HB 125–160, T 135–165 (95–
genetics, and may represent more than one species. 105% of HB), HF 31–33, E 19–22. Skull: gl 36–40,
Similar species Rajah Maxomys, M. rajah, is less mt 6.2–6.7
brightly coloured, usually has a dark brown streak in Identification Upperparts reddish-brown with
middle of underparts, lacks orange-brown collar and extensive stiff spines in fur; underparts grey with
band around leg; juveniles of both species are grey chestnut tip to fur. Tail usually slightly longer than
and difficult to distinguish. head and body, sharply bicoloured dark above and
Ecology and habitat Nocturnal and mainly white below. Hind feet with only five pads. Similar
terrestrial. Occurs in primary and secondary forests species Whitehead’s Maxomys, M. whiteheadi, is
and adjacent gardens, but not heavily disturbed smaller with shorter tail.
areas. Ecology and habitat Mostly nocturnal. Montane
Distribution SE Asia: Myanmar, Thailand, Laos, forests, usually above 900m.
Vietnam, Cambodia and Peninsular Malaysia. Also S Distribution SE Asia: known only from Peninsular
China (Yunnan), Sumatra, Java, Borneo and adjacent Malaysia, though might also be expected in extreme
islands. S Thailand.
Status Least Concern, but populations in some Status Least Concern.
areas have probably declined substantially as a
result of deforestation.

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 INDOCHINESE MAXOMYS Distribution SE Asia: Peninsular Malaysia. Also
Maxomys moi PLATE 76 Tuangku Island (W of Sumatra), Borneo.
Measurements HB 140–215, T 155–200 (95– Status Least Concern. Reportedly locally common
105% of HB), HF 37–44, E 22–27. Skull: gl 40–46, in some areas.
mt 5.8–6.6
Identification Upperparts very bright orange, fur Genus Pithecheir Medium-sized rats with naked,
dense and soft without spines. Underparts white, prehensile tail; feet adapted for climbing, first toe
sharply demarcated from dorsum. Hind feet usually widely separated from others with enlarged pads at
with only five pads. Similar species Red Spiny tip; claws instead of nails on all toes. Long, soft fur.
Maxomys, M. surifer, in Indochina has duller- Molars have cusps distinctly separated (Fig. 71d),
coloured fur with extensive spines, averages larger instead of fused into ridges like many other rats. Two
and usually has six footpads; Niviventer spp. have tail known species, one from Java and the other from
substantially longer than head and body, usually less Peninsular Malaysia.
bright orange with extensive spines, larger footpads.
Ecology and habitat Mostly nocturnal. Found in MALAYAN WOOLLY TREE RAT
primary and secondary hill forest up to 1,500m, but Pithecheir parvus PLATE 74
some records as low as 200m. Not found in heavily Measurements HB 150–180, T 170–215,
disturbed areas. HF 26–30, E 15–19. Wt 80–135g. Skull: gl 39–44,
Distribution SE Asia: known only from highlands of mt 7.5–8.2
S Laos (Bolaven plateau) and adjacent S Vietnam. Identification Fur very long and soft without spines,
Status Least Concern, but suffering from some extending about 18mm along base of tail. Upperparts
habitat loss owing to deforestation; still relatively brownish-red, with slate-grey basal half of hairs;
common in some areas. underparts creamy white with buff tinge on flanks
and chin. Ears short and translucent. Face relatively
Genus Lenothrix Dense, woolly hair without spines; short. Feet pink, hairs on upper side white. Tail
tail dark at the base, white at the distal end. Skull hairless beyond base, with smooth tip. Taxonomic
has relatively small auditory bullae; short incisive notes Formerly included with P. melanurus, which is
foramina ending well in front of molars; posterior now considered restricted to Java. Similar species
edge of bony palate ends between molars; teeth Marmoset Rats, Hapalomys spp., have nails on first
with well-developed, discrete cusps not fused into toe instead of claw, different fur colour, distinctive
ridges (Fig. 71c). Mammae 3 + 2. teeth; other arboreal rats such as Rattus, Niviventer
and Chiromyscus have shorter hair, usually mixed
GREY TREE RAT with soft spines, sparse hairs along tail.
Lenothrix canus PLATE 74 Ecology and habitat Nocturnal. Largely arboreal.
Measurements HB 165–220, T 190–270, HF Found mainly in primary lowland rainforest up
30–37. Wt 80–220g. Skull: gl 43–48, mt 8.2–9.3 to 1,000m altitude, but has been reported from
Identification Fur soft and woolly. Upperparts grey disturbed forest.
to greyish-brown; underparts white to pale buff. Tail Distribution SE Asia: known only from Peninsular
dark near body, white on distal third. Short, broad Malaysia.
hind feet adapted for climbing but with claws on all Status Data Deficient; populations have likely
toes. Taxonomic notes Formerly called L. malaisia, declined owing to loss of primary lowland forest, but
but now considered the same species as L. canus tolerance of disturbance is poorly known.
from Sumatra. Similar species Bowers’s Rat,
Berylmys bowersi, also has white tip to tail, but is Genus Hapalomys Marmoset-rats are medium-
substantially larger with coarser fur. sized, rat-like rodents, well adapted for climbing
Ecology and habitat Nocturnal. Largely arboreal, in bamboo and small branches. Feet adapted for
in trees and bushes. Recorded from primary forest, gripping; broad and wide with long, well-spaced
disturbed forest and tree plantations, mainly in digits; underside of digits with extensive ridges;
lowlands. ridged pad on sole of foot, and broadened pad

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 at tip of toe. Hind toe is opposable, with flattened described for genus. Tail relatively long, sparsely
nail instead of pointed claw. Probably most closely haired to tip. Individuals from N Laos have smaller
related to Chiropodomys, which is smaller. Teeth skulls and longer hairs on the tail, compared with
with distinctive U-shaped cusps (Fig. 71e). Two Vietnamese individuals. Similar species Greater
recognised species, both occurring in the region. Marmoset-Rat, H. longicaudatus, is larger with
Mammae 2 + 2. grey-brown upperparts including face, contrasting
orange-brown on flanks; Indomalayan Pencil-tailed
GREATER MARMOSET-RAT Tree-mouse, Chiropodomys gliroides, is smaller with
Hapalomys longicaudatus PLATE 78 grey fur, distinct tuft of hair at tip of tail.
Measurements HB 150–170, T 170–200, Ecology and habitat Nocturnal. Known individuals
HF 26–32, E 12–15. Skull: gl 39.5–41.5, in the region found in branches and trees in
mt 7.5–8.5 deciduous and evergreen hill forests at 1,200–
Identification Medium-sized marmoset-rat with 1,500m. Little else known about habitat, but might
short rostrum, soft, dense woolly fur. Upperparts be expected in bamboo and branches of low trees.
greyish-brown including upper lips and cheeks; Has not been found in disturbed areas.
underparts white with strip of orange-brown on Distribution SE Asia: Laos and Vietnam. Also S
flanks separating upperparts from underparts; China and Hainan.
slight orange-brown tinge elsewhere on underparts. Status Vulnerable. Population presumed to have
Dark brown ears fringed with long, black whisker- declined as much of hill forest in known range has
like hairs originating inside ear. Feet as described been damaged or converted to agriculture; rarely
for genus. Tail all dark, sparsely haired at base detected, and distribution may be more widespread
with longer hairs at tip. Similar species Lesser than currently recognised.
Marmoset-Rat, H. delacouri, is smaller, with dark
reddish-brown upperparts, no orange band on Genus Mus A widespread genus of small mice, with
flanks, sparsely haired tip of tail; Fea’s Tree Rat, several species closely associated with humans,
Chiromyscus chiropus, has nail on hind toe, but foot sometimes as pests. Distinguished from other
is more elongate with relatively shorter toes that lack mice by large first molar, more than half length of
swollen pads at tip, head is elongate like Niviventer, toothrow, with only two cusps in back row of cusps;
upperparts are orange-brown with flat spines in fur. third molar very small (Fig. 70a). Hind feet with six
Ecology and habitat Nocturnal. In Malaysia, mainly pads on sole of foot, all small and rounded, and
in hills above 400m, but possibly in lowlands further towards front of foot. Mammae 3 + 2 in all species
north. Associated with limited range of bamboo in the region.
species on which it feeds. Nests inside internodes
of bamboo, making holes 35mm in diameter. Hole ASIAN HOUSE MOUSE
surrounded by 10mm band where outer layer is Mus musculus PLATE 77
stripped. Lines nest with bamboo leaves. Measurements HB 65–90, T 67–92 (c.105% of
Distribution SE Asia: Peninsular Malaysia; possibly HB), HF 14.5–18.5. Wt 9–17g. Skull: gl 18.5–22,
SE Myanmar and S Thailand, but no recent records. mt 2.9–3.5
Status Endangered; very restricted range, and Identification Upperparts dark brown and grey,
much of suitable habitat has been destroyed. often with reddish-brown patches, underfur grey;
underparts only slightly paler greyish-brown. Fur
LESSER MARMOSET-RAT soft. Tail entirely dark brownish. Feet same colour
Hapalomys delacouri PLATE 78 as back except for pale toes. Taxonomic notes SE
Measurements HB 100–135, T 135–170, Asian form is often considered a separate species,
HF 22–24, E 13–15. Skull: gl 29–34, mt 5.8–6.7 Mus castaneus. Similar species Other Mus in the
Identification Fur thick and silky. Upperparts region have pale underparts; Pacific Rat, Rattus
dark orange-brown; underparts white, extending exulans, is larger with stiff spines in fur, elongate
to upper lips and edges of cheeks. Relatively long inner metatarsal pad on sole of foot, similar to
brown ears with long brown hairs inside. Feet as R. rattus (Fig. 72b).

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 Ecology and habitat Nocturnal. Diet includes wide Identification Upperparts brownish-grey; under-
range of plant and animal material. Restricted to parts pale, varying from almost white to light grey.
buildings in towns. Runs very fast. Tail mid-grey above, paler below, shorter than
Distribution SE Asia: Myanmar, Thailand, Laos, body. Upper incisors with pale buff enamel, curved
Vietnam, Cambodia and Peninsular Malaysia. SE slightly forwards, then angled straight down. In
Asian form, M. m. castaneus, also found in India skull, nasal bones long, hiding incisors from above,
and China, and introduced into many Pacific islands. incisive foramina long, ending level with middle
Other forms native to Europe and now introduced of first molars. Taxonomic notes In Thailand and
throughout much of the world. Myanmar, forms found in natural forest are larger
Status Least Concern. than those in rice fields and have been considered a
distinct subspecies, M. c. popaeus, but they appear
RICEFIELD MOUSE to interbreed where they meet. Similar species
Mus caroli PLATE 77 Ricefield Mouse, M. caroli, has darker tail as long as
Measurements HB 65–85, T 65–95 (c.100% of head and body, dark orange incisors; Fragile-tailed
HB), HF 16.5–19.2. Wt 8–14g. Skull: gl 19–22, Mouse, M. fragilicauda, is very similar in appearance,
mt 3.1–3.5 but has slightly longer tail with loose skin.
Identification Upperparts vary from rich brown Ecology and habitat Mostly nocturnal. Occurs in
to brownish-grey; underparts white or white with rice fields as well as natural grasslands in hill forest
grey base to fur. Tail very dark grey above, sharply savanna. Feeds on seeds and insects.
contrasting whitish below. Feet vary from dark grey Distribution SE Asia: Myanmar, Thailand, Laos,
to white. Upper incisors with dark orange enamel, Vietnam and Cambodia. Also N India, S China,
curved slightly forwards then angled straight down. Sumatra and Java; probably inadvertently introduced
In skull, nasal bones short so that incisors are visible to Indonesian localities.
from above, incisive foramina short, ending level Status Least Concern.
with front of first molars. Similar species Fawn-
coloured Mouse, M. cervicolor, has shorter tail, FRAGILE-TAILED MOUSE
paler grey above, pale upper incisors; Asian House Mus fragilicauda NOT ILLUSTRATED
Mouse, M. musculus, has grey-brown underparts Measurements HB 66–88, T 55–67 (c.80–85%
and entirely dark tail. of HB), HF 14–16. Wt 9–15g. Skull: gl 9.3–10.1,
Ecology and habitat Occurs in rice fields and mt 3.5–4.0
grassland, often in the same fields as M. cervicolor. Identification Upperparts brown with a mixture of
Also reported from grassy areas in open pine pale brown and dark brown hairs; underparts paler
savanna. Digs small holes in mud banks for brownish-grey. Feet white. Tail relatively short, dark
nests, which can be recognised by round pellets brown above, somewhat paler and pinkish below;
of excavated mud at entrance. Diet includes plant tail skin tears and comes off easily when handled;
material and animals such as insects. this may help the animal to escape from predators.
Distribution SE Asia: S Myanmar, Thailand, Laos, Taxonomic notes Only described as a distinct
Vietnam, Cambodia and Peninsular Malaysia (where species in 2003. Similar species Fawn-coloured
possibly introduced). Also S China, Taiwan, Ryukyu Mouse, M. cervicolor, is very similar in appearance
Islands, Sumatra, Java and some smaller islands; but differs genetically and tail skin is not especially
possibly inadvertently introduced in Indonesian prone to tear; Shortridge’s Mouse, M. shortridgei,
localities. also sheds tail skin readily but is much larger with
Status Least Concern. spiny fur.
Ecology and habitat Mostly nocturnal. Has been
FAWN-COLOURED MOUSE found in dry rice fields and patches of scrub.
Mus cervicolor PLATE 77 Distribution SE Asia: SC Thailand. S Laos.
Measurements HB 70–95, T 50–70 (c.75% of Status Least Concern; possibly more widespread
HB), HF 15–18. Wt 10–25g. Skull: gl 19.5–23.5, than currently recognised, as it is difficult to identify
mt 3.2–3.9 specimens.

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 COOK’S MOUSE INDOCHINESE SHREWLIKE MOUSE
Mus cookii PLATE 77 Mus pahari PLATE 77
Measurements HB 80–105, T 71–95 (90–100% Measurements HB 75–105, T 70–100 (90–100%
of HB), HF 17–21. Wt 18–30g. Skull: gl 23–26, of HB), HF 19–22. Wt 20–25g. Skull: gl 23–26,
mt 4.0–4.4 mt 3.4–4.0
Identification Relatively large, with large teeth. Identification Relatively long, shrew-like nose, ears
Upperparts brown to greyish-brown with quite small, scarcely projecting above nape, eyes small.
stiff fur; underparts pale grey with dark base. Feet Fur with extensive spines. Upperparts dark bluish-
grey or brown. Tail dark grey above, paler below, grey to brownish-grey; underparts white with grey
only slightly shorter in length than head and body. base to fur. Feet white. Tail grey, moderately long.
Upper incisors curved slightly backwards, with pale Taxonomic notes May include multiple species
enamel. Taxonomic notes Populations from NW across its range. Similar species Shortridge’s
Myanmar and India are smaller (about 15% shorter Mouse, M. shortridgei, has shorter tail, pale brown
head and body and tail), and sometimes considered upperparts, large ears and eyes; other Mus spp.
a separate species, M. nagarum. Similar species in region have fur without spines, larger eyes, less
Ricefield Mouse, M. caroli, is smaller with shorter pointed head.
toothrow, dark forwards-curved incisors; Fawn- Ecology and habitat Mostly nocturnal. Found
coloured Mouse, M. cervicolor, has shorter tail, in grassy areas within montane forest; probably
smaller toothrow and forwards-curved incisors. tolerant of disturbed forest.
Ecology and habitat Mostly nocturnal. Mainly in Distribution SE Asia: N Myanmar, Thailand, Laos,
hilly areas, including grassy fields in pine savanna, Vietnam and Cambodia. Also Bhutan, India, China.
open dipterocarp forest, and hill rice fields. Also Status Least Concern, although not common
some records in lowlands in riverine grassland. anywhere.
Distribution SE Asia: Myanmar, Thailand, Laos and
Vietnam. Also Nepal, India and S China. SHORTRIDGE’S MOUSE
Status Least Concern. Mus shortridgei PLATE 77
Measurements HB 95–120, T 60–90 (70% of HB),
LITTLE INDIAN FIELD MOUSE HF 19–22. Skull: gl 26–30, mt 4.4–5.2
Mus booduga PLATE 77 Identification Fur with extensive stiff, flattened
Measurements HB 60–75, T 45–55 (80–90% of spines; ears relatively large; eyes not especially
HB), HF 14.5–17. Wt 9–15g. Skull: gl 19–21.5, small. Upperparts greyish-brown; underparts
mt 3.0–3.6 greyish-white, fur with grey base and white tip, ears
Identification Very small mouse with narrow rostrum, relatively large. Tail dark grey, sometimes weakly
relatively short tail. Upperparts light brown; underparts bicoloured pink underneath. Skin on body and tail
pure white or white with grey base, sometimes with loose; tail skin sheds easily if animal is picked up
brown spot on chest. Tail bicoloured, dark above and by the tail. Similar species Other Mus spp. average
paler below. Upper incisors curved backwards. Broad smaller and lack spines; Pacific Rat, Rattus exulans,
first upper molar. Taxonomic notes Populations has relatively longer tail, elongate pads on sole of
in C Myanmar may represent a distinct species, foot, darker underparts.
M. lepidoides. Similar species Other Mus are usually Ecology and habitat Mostly nocturnal. Found in
larger, many species with incisors angled forwards; dry grass and clumps of small bamboo in open dry
Asian Harvest Mouse, Micromys erythrotis, has dipterocarp and pine forest.
relatively longer, thinner tail, small ears. Distribution SE Asia: Myanmar, Thailand, Laos,
Ecology and habitat Mostly nocturnal. Dry scrubby Vietnam and Cambodia.
areas and agricultural fields. In India, occurs in wet Status Least Concern; distribution fragmented,
rice fields. but can be locally abundant and tolerant of some
Distribution SE Asia: C Myanmar. Also Pakistan, disturbance.
Nepal, India, Sri Lanka and Bangladesh.
Status Least Concern; may be a pest in some crops.

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 Genus Micromys Includes some of the smallest INDOMALAYAN PENCIL-TAILED
species of rodents. Bony palate does not extend TREE-MOUSE
much behind teeth. Teeth small with multiple Chiropodomys gliroides PLATE 78
separate cusps (Fig. 70c). First digit (big toe) of hind Measurements HB 70–105, T 95–145 (c.135% of
foot has pointed claw. HB), HF 15–22, E 13–19. Wt 15–35g.
Skull: gl 24–27, mt 3.3–4.2
ASIAN HARVEST MOUSE Identification Fur short, thick and soft. Upperparts
Micromys erythrotis PLATE 78 bright, varying from pale red-brown to grey-brown
Measurements HB 46–68, T 54–88 (115–130% colour; underparts white. Sometimes narrow orange-
HB), HF 13–16, E 8–10. Wt 5–7g. Skull: gl 17.9– red band on flanks separating white underparts from
19.1, mt 2.7–3.1 upperparts. Tail brown, hairy, with brush of hairs
Identification Very small with short, rounded head, 4–5mm long at tip. Long whiskers on muzzle much
small, rounded ears, well haired. Upperparts greyish- longer than half of head and body length. Foot with
brown, underparts silvery-grey. Tail longer than head nail on first toe. Similar species Marmoset-rats,
and body, thin and naked, partly prehensile; dark Hapalomys spp., have nail on first toe, but toe is
pinkish-brown above, pink below. Feet relatively relatively longer and animals are larger.
broad for gripping on grass stems. Taxonomic Ecology and habitat Nocturnal. Occurs in a wide
notes Formerly considered the same species as variety of forest types, including heavily disturbed
M. minutus from Europe and N Asia (which is more areas, usually associated with extensive bamboo.
orange with a whiter belly and shorter tail), but Nests in tree holes, internodes of bamboo and
recent genetic studies confirm it is distinct. Similar similar places; in bamboo makes neat circular hole
species Majority of other mice in the region are 25mm in diameter.
larger with more conspicuous ears, thicker tail; Distribution SE Asia: Myanmar, Thailand, Laos,
Long-tailed Climbing Mouse, Vandeleuria oleracea, Vietnam, Cambodia and Peninsular Malaysia. Also
has nails instead of claws on outer toes. NE India, S China, Sumatra, Java and Bali.
Ecology and habitat Mostly nocturnal. Habits and Status Least Concern.
behaviour of Asian Harvest Mouse, M. erythrotis, not
described. Elsewhere, Micromys feeds on seeds, Genus Vandeleuria Small mice with flattened nails
fruits, buds and insects; found in grassy fields, instead of pointed claws on first and fifth digits of
including grain fields, low shrubs and bracken; well hands and feet. Mammae 2 + 2. Three species
adapted for climbing on herbaceous vegetation; currently recognised, of which only one occurs in
builds a globe-shaped nest of grass 60–130mm in region.
diameter, attached to stems of large grasses.
Distribution SE Asia: N Myanmar and NW Vietnam. ASIAN LONG-TAILED CLIMBING MOUSE
Also China (Sichuan). Vandeleuria oleracea PLATE 77
Status Not yet assessed. Measurements HB 55–85, T 90–130 (150–180%
of HB), HF 16–18, E 12–14. Wt 10–20g
Genus Chiropodomys Medium-small arboreal mice Identification Small with soft fur, very long tail.
with a short head, small body, large eyes, long Upperparts orange-brown to pinkish-brown,
whiskers on muzzle and above eye, long tail with a underparts white, sometimes tinged light brown, not
brush-like tip of long hairs and specialised feet for sharply demarcated from upperparts. Tail uniformly
climbing; feet are short and broad, first digit short brown with short hairs along length, no tuft at end.
with a broad pad and a short flat nail instead of a Outer toes on hind feet have nails instead of claws
sharp claw; remaining digits with enlarged pad at and are opposable, allowing them to grip well around
tip, with short, sharp claws. Only one species in the grass stalks and other vegetation. Taxonomic
region, but additional species occur in Borneo, the notes Geographic variation in chromosomes and
Mentawei Islands and Palawan. Mammae 0 + 2. other features suggests may represent more
than one species. Similar species Asian Harvest
Mouse, Micromys erythrotis, has short, rounded

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 ears, claws instead of nails on all toes; Vernay’s long. Front feet with four well-developed toes, all
Climbing Mouse, Vernaya fulva, has pointed claws with claws; hind feet with five well-developed toes,
on all toes; Indomalayan Pencil-tailed Tree-mouse, all with claws. Incisors narrow with yellow enamel.
Chiropodomys gliroides, averages larger, with broad Incisive foramina moderate, ending in front of or
pad at tip of toes, flat nail only on first digit with claw level with front of first molar; bony palate extending
on fifth digit, tuft of hair at tip of tail. posterior to last molar. Mammae vary between the
Ecology and habitat Mostly nocturnal. Dense species. Widely distributed throughout Europe and
tangles of vines, tall cane and brush. Arboreal, Asia, with at least 20 species. Two species have
climbing on small branches and tall grasses. Builds been reported in the literature from Myanmar, but
globe-shaped nest of grasses in cane, 1–2m above there is some uncertainty as to whether both actually
ground. occur there; published reports of A. latronum from
Distribution SE Asia: Myanmar, Thailand (N of Myanmar have toothrow measurements that fit
Isthmus of Kra), Vietnam and SW Cambodia. within the range for A. draco. Both species are
Probably Laos. Also India, Sri Lanka, S Nepal and described here, but further research is required to
SE China. confirm the number of species in Myanmar.
Status Least Concern.
SOUTH CHINA WOOD MOUSE
Genus Vernaya Small mice, superficially similar to Apodemus draco PLATE 78
Vandeleuria but all digits have pointed claws instead Measurements HB 80–105, T 90–130 (100–125%
of nails, except vestigial first finger (thumb) on of HB), HF 20–25. Skull: gl 24–28, mt 3.7–4.4
front feet. Mammae 1 + 2. Poorly known, with few Identification Upperparts yellowish-brown to dark
reported specimens. orange-brown, the hairs with grey base; underparts
sharply contrasting greyish-white, the individual
VERNAY’S CLIMBING MOUSE hairs with grey base and silvery tip. Fur short,
Vernaya fulva PLATE 78 soft and velvety. Ears relatively large, dark. Tail
Measurements HB 60–80, T 100–140 (170– moderately long, dark above, usually somewhat
200% of HB), HF 16–18. Skull: gl 22, mt 3.3 paler below, sparsely covered with short whitish
Identification Small with very long tail. Upperparts hairs. Mammae 2 + 2. Taxonomic notes Includes
brown with yellowish tinge, especially on flanks A. orestes, which has sometimes been considered
and cheeks; underparts grey with buff tip to hairs. a separate species. Similar species Large-eared
Fur without spines. All toes on hind feet have claws Wood Mouse, A. latronum, averages larger with a
instead of nails. Tail all dark. Similar species longer toothrow.
Asian Harvest Mouse, Micromys erythrotis, is Ecology and habitat Forest heath and scrubby
smaller with shorter tail; Long-tailed Climbing areas in mountains at 1,300–3,800m.
Mouse, Vandeleuria oleracea, has flattened nails Distribution SE Asia: N and EC Myanmar. Also NE
instead of claws on outer toes; Mus spp. have India and China.
shorter, thicker tail. Status Least Concern; common in some areas.
Ecology and habitat Mostly nocturnal. Known from
mountains above 2,100m; Myanmar specimens LARGE-EARED WOOD MOUSE
found in areas of low shrubs and thick bracken. Apodemus latronum NOT ILLUSTRATED
Distribution SE Asia: N Myanmar. Also S China. Measurements HB 95–110, T 100–120
Status Least Concern, but relatively poorly known, (100–120% of HB), HF 24–27. Skull: gl 28–30,
with few records. mt 4.5–5.1
Identification Relatively large with long, soft silky
Genus Apodemus Medium to small soft-furred fur. Upperparts orange-brown to greyish-brown, the
mice, generally similar to typical mice (Mus) but hairs with dark grey base; underparts contrasting
first and second molars with three well-developed greyish-white, individual hairs with grey base and
posterior cusps (Fig. 70b), rather than only two as in silvery-white tip. Tail generally somewhat longer
Mus. Ears relatively large, tail generally moderately than head and body, varying from bicoloured (dark

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 above, whitish below) to all dark; sparsely haired Ecology and habitat Hill and lower montane areas
with short hairs. Ears large, blackish, noticeably in a variety of habitats, including scrub and bracken.
darker than back. Mammae 1 + 2. Similar species Distribution SE Asia: N Myanmar. Also S China.
South China Wood Mouse, A. draco, has smaller Status Least Concern; believed to be common in
skull and toothrow. some areas.

Family PLATACANTHOMYIDAE PYGMY-DORMICE

This family has some resemblance to dormice of the family Gliridae (which do not occur in South-east Asia)
and has sometimes been included with them, but differs in having only three cheek teeth on each side of
each jaw and small auditory bullae without septa.
The family includes two genera, Platacanthomys with spiny fur, which is now restricted to India although
fossil species have occurred more widely, and Typhlomys from South-east Asia and China.

Genus Typhlomys Only one species is currently on distal half forming a distinct brush, often with a
recognised in the genus, though the only population white tip. Hands white, hind feet dusky. Taxonomic
known from the region, in N Vietnam, has sometimes notes Form in N Vietnam has sometimes been
been considered a separate species from that found considered a separate species, T. capensis, which
in China. Cusps on molars form folded ridges that differs in slightly larger size and darker underparts
are angled diagonally inwards, quite unlike the from typical Chinese cinereus, but recent research
separate cusps or horizontal ridges of mice in the suggests it is within the variation of the species,
family Muridea (e.g. Fig. 70a, b, c) or the zigzag and should at most be considered a subspecies,
patterns of Cricetid voles (Fig. 70d) T. c. capensis. Similar species Other rodents
have different patterns of cusps on the molars;
SOFT-FURRED PYGMY-DORMOUSE long-tailed mice, Vandeleuria and Vernaya spp. are
Typhlomys cinereus PLATE 78 smaller, with even longer tail, broader feet adapted
Measurements HB 70–100, T 95–135 for climbing.
(125–135% of HB), HF 20–23. Skull: gl 25–26 Ecology and habitat High montane forest, at
Identification Small and mouse-like with prominent elevations of 1,200–2,100m. Has been recorded
ears, soft, dense fur, small eyes and narrow hind from moss forest with undergrowth of bamboo, but
feet. Upperparts uniformly dark brownish-grey, poorly known.
sometimes almost black. Underparts with grey base Distribution SE Asia: NW Vietnam. Also mountains
and buffy tip in Vietnamese form (greyish-white tip of S China.
in Chinese populations). Tail long and thin, scaly Status Least Concern, but very little known about
with sparse hairs near base, longer, denser hairs form in Vietnam.

Family CRICETIDAE, Subfamily ARVICOLINAE VOLES

The family Cricetidae is currently considered to include several subfamilies, of which only one is represented
in the region. This subfamily is widely distributed through the Old and New Worlds, primarily in the northern
hemisphere.
Most voles in the region have a short, blunt head with short, rounded ears, rounded body and relatively
short tail. The molars have a complex zigzag pattern that is quite unlike those of other rodents in the region,
and lack roots (e.g. Fig. 70d).

Genus Eothenomys Similar to Microtus, but only two are known in the region. Two additional
posterior margin of palate in skull straight across. species have been reported from SW Yunnan, near
Mammae 0 + 2. Cusps of first lower molar matched the border with Myanmar, and might be expected
up on opposite sides, forming loops (Fig. 70d). Tail to occur in Myanmar. These are E. olitor, which is
moderately covered with short hairs that partly hide similar to E. melanogaster but smaller in size, with
scales. Eight species currently recognised, of which dark greyish-brown upperparts with pale frosted tip;

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 and E. miletus, which is similar in size and colour CLARKE’S VOLE
to E. cachinus but with a relatively shorter tail and Microtus clarkei PLATE 79
higher skull profile. Measurements HB 105–120, T 60–65 (50–60%
of HB), HF 19–20, E 14–16. Skull: gl 27–28.5,
KACHIN VOLE mt 6.3–6.6
Eothenomys cachinus PLATE 79 Identification Relatively large with moderately
Measurements HB 110–125, T 43–60 (40–50% long hair. Upperparts dark brown; underparts slate-
of HB), HF 18–21, E 12–15. Skull: gl 27–28, grey with silver tip to fur. Medium-long tail covered
mt 6.2–7.0 with short hairs, thicker at tip forming a slight tuft,
Identification Upperparts bright tawny-brown, not completely covering scales. Taxonomic notes
underparts grey with buff tips. Medium-length tail Sometimes included in genus Volemys, but recent
(for a vole). Taxonomic notes Formerly sometimes morphological analyses suggest it is closer to
considered a subspecies of E. miletus from China. Microtus. Similar species Other voles have tail
Similar species Père David’s Vole, E. melanogaster, <50% of head and body; Eothenomys spp. have inner
is smaller with darker, less bright underparts, shorter and outer cusps of lower molars opposite each other.
tail; Clarke’s Vole, Microtus clarkei, has cusps of Ecology and habitat Coniferous forest and alpine
lower molar more alternating, darker fur, longer tail. meadows in high mountains at 3,300–4,300m.
Ecology and habitat Montane forest at 2,300– Distribution SE Asia: N Myanmar. Also S China
3,200m. (SE Tibet, Yunnan and Xizang).
Distribution SE Asia: NE Myanmar, west of Salween Status Least Concern.
River valley. Also adjacent NW Yunnan in China.
Status Least Concern. Genus Neodon Small voles, similar to Microtus
but with cusps of first lower molar forming only
PÈRE DAVID’S VOLE three alternating closed triangles. Taxonomic
Eothenomys melanogaster PLATE 79 relationships are still somewhat uncertain, and
Measurements HB 90–100, T 21–42 (20–40% of the genus is sometimes considered a subgenus of
HB), HF 15–17, E 11–14. Skull: gl 23–26, Microtus, Pitymys or Phaiomys.
mt 5.5–6.6
Identification Upperparts dark brown to blackish, FORREST’S MOUNTAIN VOLE
underparts slate-grey, sometimes tinted buffy or Neodon forresti PLATE 79
brown. Medium-short tail. Similar species Kachin Measurements HB 95–115, T 30–45 (40% of HB),
Vole, E. cachinus, averages larger, with a longer tail, HF 17–19, E 14–16. Skull: 25–27, mt 5.5–6.5
brighter fur. Identification Upperparts deep dark brown with
Ecology and habitat In Myanmar, reported from blackish-grey base; underparts pale brown or greyish.
relatively open habitats, including meadows and Tail bicoloured, dark above, paler below. Upper incisors
edges of cultivated fields, at altitudes of 1,200– relatively wide with pale to dark orange enamel,
2,750m. Reported from mossy rhododendron sometimes with faint grooves on outer surface.
forest in N Thailand. Makes burrows in earth at Taxonomic notes Formerly considered a subspecies
sides of banks. of N. irene, but distinctly larger and with longer, darker
Distribution SE Asia: N Myanmar, N Thailand and fur. Similar species Clarke’s Vole, Microtus clarkei,
N Vietnam. Also S China. has slightly larger skull and relatively longer tail, fewer
Status Least Concern. closed triangles in first lower molar; Kachin Vole,
Eothenomys cachinus, has paler upperparts.
Genus Microtus Similar to Eothenomys, but in Ecology and habitat High mountains at 3,300–
skull, central septum of bone extends backwards 3,600m.
from posterior edge of palate. First lower molar with Distribution SE Asia: N Myanmar. Also China (NW
alternating cusps forming five closed triangles. Upper Yunnan).
incisors broad with reddish enamel, sometimes with Status Data Deficient; no recent information on
a slight groove in Asian species. Mammae 0 + 2. population status or threats.

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 Family SPALACIDAE, Subfamily RHIZOMYINAE BAMBOO RATS
The bamboo rats are sometimes placed in their own family, but more recently have been considered a
subfamily of the family Spalacidae, which includes several different groups from Africa and Asia. Bamboo
rats are characterised by a short face and thick, rounded head; small eyes; thick, tube-like body with no
obvious neck; short, smooth, sparsely haired tail without scales; short legs and stout claws. All of these
are adaptations for burrowing. The zygomatic arches are broad, the skull somewhat flat with a triangular
braincase. The incisors are thick and relatively blunt, and used both for digging and eating; like those of most
rodents, they continue growing throughout the life of the animal.

Genus Cannomys Smaller than other bamboo rats, molar larger than second. Ears short, visible through
with prominent incisors that protrude forwards, fur. Three recognised species in genus, all occuring
large gap between incisors and molars (diastema) in the region.
exceeding 40% of condylobasal length, first upper
molar larger than second, and footpads that are HOARY BAMBOO RAT
smooth, without granular ridges. Currently only one Rhizomys pruinosus PLATE 80
species recognised in genus, but some authorities Measurements HB 260–350, T 100–120
suggest this may actually be a complex of species. (35–40% of HB), E 18–25, HF 45–60. Wt 1–3kg.
Skull: cbl 56–67, mt 13.5–15.5
LESSER BAMBOO RAT Identification Upperparts dark greyish-brown to
Cannomys badius PLATE 80 chocolate-brown with grey base to hairs; many
Measurements HB 150–265, T 45–75, E 7–11, long guard hairs with long white tip, giving frosted
HF 30–35. Wt 0.5–0.8kg. Skull: cbl 40–45, appearance; underparts slightly paler. Some
mt 8–12 individuals, especially in south, are smaller and
Identification Small, stout rat with short, broad paler, making guard hairs less conspicuous. Fur is
head, short legs, strong feet. Fur soft and dense, dense and soft, especially in north, but sometimes
upperparts dark reddish-brown with dark slate- more sparse and harsh in south. Feet brown with
grey underfur that shows through as fur becomes granular pads on soles; two separate posterior pads
worn; underparts slightly paler with thinner hair. on hind foot. Similar species Chinese Bamboo
Sometimes has white patch on forehead. Ears Rat, R. sinensis, averages larger, with pale throat,
short and largely hidden in fur. Tail short, with soft more uniform colour without white tip to hairs;
wrinkled skin, sparse short hairs. Similar species Indomalayan Bamboo Rat, R. sumatrensis, is larger,
Other bamboo rats, Rhizomys spp., are much larger, with coarse hair, contrasting reddish face, only one
with granular pads on soles of feet, different fur posterior pad on hind foot.
colour and visible ears. Ecology and habitat Scrub and hill forest, often
Ecology and habitat Bamboo thickets in hilly with grassy areas and extensive stands of bamboo at
or mountainous areas. Spends much of the time altitudes of 1,000–4,000m. Digs extensive burrows
underground in fairly conspicuous burrows that it where it stays during day, emerging at night to feed,
digs into side of a bank, under bamboo or elsewhere. though also feeds on roots underground.
Emerges at night to feed on bamboo roots and Distribution SE Asia: E Myanmar, Thailand, Laos,
shoots, grass seeds and fallen fruits. Vietnam, Cambodia and N Peninsular Malaysia. Also
Distribution SE Asia: Myanmar, Thailand, Laos, NW NE India and S China.
Vietnam and N Cambodia. Also Nepal, NE India, Status Least Concern, but exploited for food and
Bhutan, Bangladesh and S China. does not breed quickly, so may be declining in
Status Least Concern; relatively common in many some areas.
areas, though heavily trapped for food.
CHINESE BAMBOO RAT
Genus Rhizomys Large bamboo rats with gap Rhizomys sinensis PLATE 80
between incisors and molars (diastema) less than Measurements HB 230–450, T 50–90 (15–25%
40% of condylobasal length of skull, first upper of HB), HF 35–60. Skull: cbl 69–80, mt 14–19

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 Identification Medium to large, with very short tail; fur Skull: cbl 71–78, mt 14–15
thick and soft; overall colour buffy-brown to reddish- Identification Large bamboo rat with sparse
brown, mixed with grey; fur has extensive pale to coarse hair, relatively long tail. Upperparts pale
mid-grey base. Two separate posterior granular pads brownish-grey, darker on top of head. Cheeks often
on hind foot. Similar species Hoary Bamboo Rat, contrasting reddish-brown, especially in young
R. pruinosus, averages smaller, with darker fur and animals. Posterior granular footpads on hind foot
white tip to many hairs, proportionately longer tail. joined together to form a single large pad. Similar
Ecology and habitat Mainly in hill and lower species Hoary Bamboo Rat, R. pruinosus, is smaller,
montane forest, especially in areas with extensive with dense hair, different colour, two separate
bamboo. Digs burrows like other members of genus. posterior footpads.
Distribution SE Asia: N Myanmar, N Vietnam and Ecology and habitat Secondary forest with
might be expected in Laos. Also S China. extensive bamboo in lowlands and lower hills. Feeds
Status Least Concern; relatively common in some on bamboo roots and other plant material, often
areas, although heavily exploited for food and may coming above ground to forage at night.
be declining. Distribution SE Asia: Myanmar, Thailand, Laos,
Vietnam, Cambodia and Peninsular Malaysia. Also
INDOMALAYAN BAMBOO RAT S China (Yunnan), Sumatra.
Rhizomys sumatrensis PLATE 80 Status Least Concern. Still common in some areas,
Measurements HB 280–480, T 100–200 (35– but breeds slowly and has declined dramatically in
50% of HB), E 20–36, HF 46–67. Wt up to 2kg. areas where heavily hunted for food.

Family DIATOMYIDAE KHA-NYOU

The sole known living member of this family, the Kha-nyou, Laonastes aenigmamus, was first described to
science in 2005. At the time, it was thought to represent a new family of mammal, but subsequent research
showed that it belonged to a family previously known only from fossils more than 11 million years old. The
relationship of this family to other rodents is uncertain – it shows features that are somewhat intermediate
between squirrels and rats, but it may be closer to the group that includes porcupines. This species has four
well-developed cheek teeth on each side, all of similar size (one premolar and three molars; most squirrels
have an extra upper small premolar; rats lack premolars), each with two broad ridge-like cusps. The feet
have large, well-developed pads on the base, lacking fine striations; presumably these help the animals to
grip on sharp limestone rocks.

KHA-NYOU longer tail, shorter face, more slender body, more

Laonastes aenigmamus PLATE 80 bounding gait, moving both feet at the same time;
Measurements HB 210–290, T 120–160 (55– rats and mice have a thin hairless tail.
60% of HB), HF 37–44, E 21–26. Wt 330–420g. Ecology and habitat Apparently mainly nocturnal.
Skull: gl 61–71, mt 13.2–14.8 Known only from limestone hills and outcrops in
Identification Pointed, rat-like head, but with a C Laos and Vietnam. Diet is mainly plant material,
bushy, squirrel-like tail. Upperparts dark greyish- including leaves and seeds, with some insects.
black with a mixture of soft grey hairs with pale tip Distribution SE Asia: Laos, Vietnam.
and stiff black guard hairs; amount of grey frosting Status Least Concern. Known only from limestone
varies among individuals, with some nearly black areas in C Laos and Vietnam, but currently
and others extensively grey; underparts silvery-grey, protected. Evidence of high genetic isolation among
individual hairs with grey base and white tip; extent local populations, and some genetically distinct
of pale tip varies among individuals. Tail coloured populations are possibly at risk from destruction of
as back, very bushy. Large hind legs; walks with a limestone or hunting.
waddle-like gait. Similar species Squirrels have a

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 Family HYSTRICIDAE PORCUPINES
Porcupines are larger and more heavily built than squirrels or rats, with characteristic hard spines or quills
over most of the upperparts. The incisors and molars are large and powerful. All three species found in
South-east Asia are primarily terrestrial and usually nocturnal, though both the smaller species can climb
trees. They sleep during the daytime in underground holes or burrows.

LONG-TAILED PORCUPINE Ecology and habitat Mainly terrestrial, digging

Trichys fasciculata PLATE 81 extensive burrows under forest floor. Feeds on fallen
Measurements HB 375–435, T 150–240, fruits including oil palm, as well as tree bark, roots
HF 61–67. Wt 1.5–2.0kg and tubers. Occurs in forests and cultivated areas.
Identification Upperparts brown; underparts Distribution SE Asia: Myanmar, Thailand, Laos,
whitish. Spines short and flattened, dark brown Vietnam, Cambodia and Peninsular Malaysia. Also
towards ends, whitish towards base. Tail brown India, Nepal, S China, Sumatra and Borneo.
and scaly, with brush of flattened bristles at tip. Status Least Concern.
Like a large rat with coarse fur. Part or all of tail
sometimes missing. Cannot bristle or rattle quills. BRUSH-TAILED PORCUPINE
Similar species Brush-tailed Porcupine, Atherurus Atherurus macrourus PLATE 81
macrourus, is larger, has longer spines, and a Measurements HB 380–520, T 140–230,
shorter, thicker tail with a bigger brush; large rats HF 64–75
(Muridae) have much smaller, softer spines. Identification Upperparts greyish-brown to
Ecology and habitat Often seen feeding on the brown; underparts whitish. Body largely covered
ground, where it has been observed feeding on large with flattened spines, longest in the middle of the
tree seeds as well as bamboo shoots, but can also back. Tail scaly with tuft of whitish quills at tip,
climb well. Occurs in forests and cultivated areas. quills shaped like a string of beads that rattle when
Distribution SE Asia: Peninsular Malaysia. Also shaken. Tail sometimes broken off. Similar species
Sumatra and Borneo. Malayan Porcupine, Hystrix brachyura, is larger,
Status Least Concern. black and white in colour, with long, conspicuous
quills and a short tail; Long-tailed Porcupine, Trichys
MALAYAN PORCUPINE fasciculata, has smaller, inconspicuous quills, small
Hystrix brachyura PLATE 81 brush on tail with flattened spines that do not rattle.
Measurements HB 590–720, T 60–110, HF 80–95 Ecology and habitat Mainly terrestrial, but can also
Identification Generally black; long spines or quills climb well. Nocturnal, living in burrows during the
on lower back white, with black band in middle; short day. Eats roots, tubers, fruits, some cultivated crops
spines on front parts of body mostly blackish, some and tree bark. Occurs in forests and plantations,
with paler base and tip. Spines along back of neck rarely cleared areas.
may form a crest. Tail has hollow, goblet-shaped Distribution SE Asia: Myanmar, Laos, Thailand,
quills that rattle when shaken. Similar species Vietnam, Cambodia and Peninsular Malaysia. Also
Brush-tailed Porcupine, Atherurus macrourus, is E India, S China and Sumatra.
smaller, with shorter brown quills, longer scaly tail Status Least Concern, though likely declining due to
with brush on tip, beaded tail quills. habitat loss and hunting.

Order LAGOMORPHA
Pikas, hares and rabbits
Lagomorphs differ from rodents in having two pairs of upper incisors, although the second pair is located
directly behind the first pair, and does not have a sharp cutting edge. Both pairs of incisors are completely
covered in enamel, which is generally white, not orange as in most rodents. The order includes two families,
both of which are very distinctive in shape, and both with representatives in the region.

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 Family OCHOTONIDAE PIKAS
There are about 30 species of pika currently recognised worldwide, more than two-thirds of which have
been reported from China, although there is still considerable uncertainty about exact species limits. All are
included in the same genus, Ochotona, and are generally similar in shape with compact body, short face,
small, rounded ears that protrude above the head, relatively short legs and no visible tail.

MOUPIN PIKA FORREST’S PIKA

Ochotona thibetana PLATE 81 Ochotona forresti NOT ILLUSTRATED
Measurements HB 140–180, T none, HF 24–32, Measurements HB 155–185, T none, HF 27–30,
E 17–23. Wt 75–135g. Skull: gl 36–42 E 18–23. Wt 110–150g. Skull: gl 37–41
Identification Small, with rounded ears and no tail. Identification Small, with rounded ears and no
Fur colour varies seasonally. In summer, upperparts tail. In summer, upperparts blackish-brown to dark
brownish, varying from sandy-brown to reddish- reddish-brown, underparts similar; in winter, dark
brown to dark brown, sometimes with light speckling; grey-brown, underparts only slightly paler. Ears
underparts paler varying from light grey to greyish- light chestnut with white rim. Patch of dark grey fur
yellow, except for buffy-brown collar on throat. In behind ears that may form collar on nape. Feet dull
winter, upperparts become paler, buffy-brown to dull white. Similar species Moupin Pika, O. thibetana,
brown. Ears are rounded, dark brown on outside with has paler underparts, buffy patches behind ears
white rim. Pale buffy area behind each ear. Similar that do not meet around nape, shorter front claws,
species Forrest’s Pika, O. forresti, has darker narrower skull, especially across zygomatic arches.
underparts, with dark grey patch behind ears. Ecology and habitat Forested areas on mountain
Ecology and habitat Found at altitudes of 1,800– slopes at elevations of 2,600–4,400m, especially
4,000m in bamboo and rhododendron forests and on south-facing slopes. Poorly known; may dig
subalpine forests, where it digs burrows for shelter. burrows, but may also occur around rock piles.
May be seen in openings in forest, near shrubs in Distribution SE Asia: N Myanmar. Also N India,
more open areas, or around scrub piles in logged Bhutan and China (NW Yunnan and SE Tibet).
areas. Status Least Concern, though status and distribution
Distribution SE Asia: N Myanmar. Also N India and in Myanmar very poorly known; may be declining
S China. due to loss of montane forest.
Status Least Concern, although status in region
poorly known.

Family LEPORIDAE HARES AND RABBITS

This family includes many species worldwide, of which three are found in South-east Asia. The two species
of hare are relatively large, with long ears and long, powerful hind legs, readily distinguished from any other
mammals in the region, but similar to each other. The Annamite Striped Rabbit, Nesolagus timminsi, which
was first described in 2000, is very similar to the only other member of the genus, Nesolagus netscheri, from
Sumatra, but very different from all other mammals in the region.

BURMESE HARE greyish sides, white below. Upper incisors with

Lepus peguensis PLATE 81 Y-shaped groove, usually filled with a cement-
Measurements HB 360–500, T 65–80, like substance. Similar species Chinese Hare,
HF 90–105, E 65–85. Skull: gl 80–95 L. sinensis, has brown on top of tail, V-shaped
Identification Typical hare shape with large hind groove on incisors without cement.
feet, moderately long ears. Upperparts mottled Ecology and habitat Found in dry dipterocarp
brown, buff and pale grey with reddish-orange forest and grasslands, mainly in lowlands, often in
patch on nape; underparts white, mixed with disturbed areas. Primarily active at night, resting
patches of brown including on throat. Ears long during the day in a sheltered spot under a bush or
with small black tip. Tail with narrow black top, in thick grass.

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 Distribution SE Asia: Myanmar, Thailand (N of Status Least Concern, but very rare in Vietnam,
Malay Peninsula), Cambodia, Laos and S Vietnam. where there are no confirmed records since the
Status Least Concern. 1990s.

CHINESE HARE ANNAMITE STRIPED RABBIT

Lepus sinensis PLATE 81 Nesolagus timminsi PLATE 81
Measurements HB 350–450, T 40–60, HF 80–110, Measurements HB 350–400, HF 60–70, E 30–40.
E 65–85. Wt 1–2kg. Skull: gl 67–93 Skull: gl 70–80
Identification Similar shape and size to Burmese Identification Small with elongate but relatively
Hare. Upperparts overall mottled mixture of dark short ears, short tail, distinctive back pattern.
brown, buff and pale grey, tending to be paler and Upperparts pale brownish to brownish-grey with
more yellowish in winter; orange-brown patch on broad black or very dark brown stripes along sides
nape; underparts white with areas of pale brown. of upper back and across lower back. Orange-
Ears moderately long, usually with black tip. Tail dark brown patch on rump. Underparts paler greyish-
brown on top, buffy on sides, whitish below. Upper white. Similar species Hares, Lepus spp., are
incisors with V-shaped groove lacking cement. larger, with long ears, lack black stripes; pikas,
Similar species Burmese Hare, L. peguensis, Ochotona spp., are much smaller, with round ears,
has black on top of tail, Y-shaped groove on upper no stripes.
incisors. Ecology and habitat Understorey of wet hill forest
Ecology and habitat Open grassy areas in hills up in Annamites; altitudes as low as 200m.
to 4,000m altitude. Mainly nocturnal, but sometimes Distribution SE Asia: Annamite Mountains of
active during the day. Feeds on vegetation. Nests in C Vietnam and C Laos.
burrows dug by other animals. Status Data Deficient, but likely to be declining due
Distribution SE Asia: NE Vietnam. Also SE China to habitat loss and excessive hunting and trapping.
and Taiwan.

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 GLOSSARY

adult sexually mature. digit finger or toe.

agouti a speckled colour pattern found in many dipterocarp forest forest dominated by large trees
rodents, formed from alternating bands of pale in the family Dipterocarpaceae.
and dark colours on the hairs. distal towards the end (e.g. away from the body).
anterior towards the front or head end of an animal. distichous flattened and feather-like in shape
antitragus flap of skin on back of ear of (referring to a tail).
horseshoe bats, which often lack a true tragus diurnal active during daylight.
(see Figs 32, 35). dorsal relating to the back or upper surface.
arboreal living in trees. feral domestic animals that have escaped from
arthropod invertebrate animals with hard, jointed captivity and breed in the wild state.
exoskeletons, including insects, spiders and flukes broadened tail of whales and dolphins.
crustaceans. foramen (plural foramina) hole in bone, especially
auditory bulla (plural bullae) thin-walled bony the skull, where nerves or blood vessels enter.
structure on the bottom of the skull that forearm (FA) part of the arm between the elbow
encloses the inner and middle ear. and the wrist on a bat (see Fig. 3).
baculum bone that supports the penis in many fossorial adapted to live underground, usually with
mammals. strong feet for digging.
baleen large fibrous plates found in the mouth of greatest length (gl) the greatest length from the
some whales, used for filtering food. back to the front of a skull excluding the teeth
bifid with two forks or lobes. (see Figs 5, 7).
buff or buffy pale yellowish-brown. grizzled dark coloration with white or pale
calcar (in bats) long bone protruding from heel speckling at tips of hairs.
that supports posterior edge of interfemoral guard hairs hairs that are longer than the main
membrane. coat of hairs.
canine width (C–C) width across the outside of gular sac small pocket of skin under the chin.
the base of the upper canines (see Fig. 7). incisors front teeth (see Fig. 7).
canines pointed, usually long, teeth behind the infant baby animal entirely dependent on its
incisors (see Fig. 7). mother.
cheek teeth the chewing teeth inside the cheeks, interfemoral membrane (in bats) membrane
consisting of both molars and premolars. between the hind legs, often enclosing the tail.
cingulum ridge around the base of a tooth. internarial in between the nostrils.
condylobasal length (cbl) length of the skull from juvenile young animal, not yet fully grown, but
the back of the occipital condyles to the front of partially or totally independent of its mother.
the premaxillae (see Figs 5, 7). karst landscape dominated by limestone
condylocanine length (ccl) length of the skull formations, often with many caves.
from the back of the occipital condyles to the lancet tall, often pointed, part of posterior noseleaf
front of the canines (see Fig. 7). of a horseshoe bat (see Fig. 32).
cusp raised point on a tooth. lateral lappets flaps at the base of the sella in
decurved curved downwards. some horseshoe bats (see Fig. 32).
dental formula shorthand notation for indicating the liana large woody vine.
arrangement and number of teeth (see page 14). mammae mammary glands or nipples; many
dew toes hind two toes on pigs, deer and other mammals have several pairs, allowing suckling
ungulates that only touch the ground on soft soil. of several young at the same time.
dewlap loose flap of skin under the neck maxilla (plural maxillae) bone in the skull that
(especially in cattle). supports all of the upper teeth except the
diastema natural gap between adjacent teeth. incisors.

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 maxillary toothrow (mt) length of the upper premaxilla (plural premaxillae) the small bones at
toothrow from the back of the molars to the the front of the skull that support the incisors;
front of the canines (see Figs 5, 7). these vary greatly in size and shape, especially
melon large, rounded expanded area on the within the bats.
forehead of some whales and dolphins. premolars teeth between the molars and canines
metacarpal bone between the wrist and the finger (see Fig. 7) that are preceded by ‘baby’ teeth,
bones (see Fig. 25). which are shed.
metacarpal pouch (in some bats) extra flap of rhinarium moist area at the tip of the nose or
skin between the forearm and metacarpal at muzzle.
the wrist that forms a pouch. rostrum narrower part of the skull in front of the
molar width (M–M) width across the outside of position of the eyes.
the upper molars at the bases. sagittal crest ridge of bone in the middle of the
molars relatively large posterior cheek teeth, skull of many mammals.
distinguished from the premolars by not having sella elevated part of the centre of the noseleaf of
deciduous precursors or ‘baby’ teeth (see a horseshoe bat (see Fig. 32).
Fig. 7). septum (plural septa) thin wall inside a chamber,
muzzle part of an animal’s face in front of its which divides it into compartments.
eyes, including the nose. sympatric occurring together in the same
myotodont pattern of ridges and cusps on the geographic area as another species.
molars of most Myotis and Hypsugo bats (see talonid posterior section of lower molars of bats
Fig. 39). (see Fig. 43).
nape back of neck. terrestrial active on the ground.
nocturnal active only or mainly at night. tines prongs on the antlers of deer.
noseleaf (in bats) flaps of skin around the nose torpor state of reduced activity and awareness, in
that are thought to be related to echolocation. which the body temperature drops to conserve
nyctalodont pattern of ridges and cusps on the energy.
molars of Nyctalus, Pipistrellus and some tragus in the ear of most bats, flap of skin
Myotis bats (see Fig. 39). supported by cartilage, located immediately in
olive a greenish-brown colour. front of the ear canal.
orbit area in the skull where the eyes are located. trigonid anterior section of lower molars of bats.
palate upper part (roof) of the mouth. underparts throat, chest, belly and insides of the
pedicel bony projection on the skull of deer from legs.
which the antlers grow. unicuspids (in shrews) small, single cusped teeth
pelage coat of fur or hair of a mammal. between the incisors and the multi-cusped
perineal around the anus. molars.
phalanx (plural phalanges) finger bones (see upperparts the back or ‘top’ of a mammal,
Fig. 25). including the outsides of the legs.
pinna (plural pinnae) externally visible part of ear. zygomatic arch a curved arch of bone below the
posterior towards the back. eye socket, found in most mammals (e.g. Fig.
prehensile referring to a tail with a tip that can 5), but not shrews (see Fig. 6).
curl and grasp branches.

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 SELECTED BIBLIOGRAPHY

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 FIGURE CREDITS

The line drawings in the text were derived from a Sergei Kruskop, from Borissenko and Kruskop
variety of sources. Some were prepared by digitally (2003): 9, 10, 11, 30a, 31 (c, d), 36, 39; from
tracing over photographs of museum material. Kruskop (2013): 42 (g).
Others were reproduced from published materials
or from original drawings prepared by other Alex Borisenko: 7, 37.
artists, with permission of the original publisher or Mammal Society of Japan, by Patricia J. Wynne
artist as listed below. For figures previously used from Lunde et al. (2003): 17, 18, 19; by Patricia
in published sources, the original publication is J. Wynne from Lunde et al. (2004): 22.
listed in the form of author and year, and refers
to entries in the Selected Bibliography. Note that Bloomsbury Publishing: 1, 4.
in many cases these drawings were rearranged
R. I. Pocock, from Pocock (1939–41): 53.
or resized, and in a few cases edited slightly from
their original form. The Sabah Society and Karen Phillipps, from
Payne, Francis and Phillipps (1985): 2, 3, 5, 12,
American Museum of Natural History, by Jennifer 20, 23, 24, 25, 26 (a, b, c, e), 28, 29, 31 (a, b),
Perrott Emry from Musser (1973): 72; by Patricia 32, 34, 35, 47, 54, 56, 57, 61, 65.
J. Wynne from Musser et al. (2006): 73.
Nico Van Strien, from Strien (1983): 13, 15, 48,
Charles M. Francis: 6, 8 (with R. Budden), 14, 16, 50, 51, 52, 55, 58, 59, 60, 62, 64, 67.
21, 26 (d, f, g), 27, 30 (b, c), 33, 38, 40, 41, 42
(a–f, h), 43, 44, 45, 46, 49, 63, 66, 68, 69, 70, Copyright remains with the original sources as
71, 74. credited.

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 INDEX

Numbers in bold refer to plate numbers.

Numbers in roman are page numbers.

Aeromys tephromelas 69, 364 Bat, Beelzebub Tube-nosed 25, 277 Bat, Greater Tube-nosed 24, 279
Aethalops aequalis see A. alecto Bat, Big-eared Horseshoe 11, 230 Bat, Grey Fruit 6, 220
Aethalops alecto 6, 220 Bat, Black-bearded Tomb 8, 224 Bat, Griffin’s Roundleaf 245
Ailurus fulgens 39, 306 Bat, Black-capped Fruit 6, 220 Bat, Guillén’s Tube-nosed 274
Amblonyx cinereus 42, 312 Bat, Blanford’s Fruit 6, 221 Bat, Hairy-winged 26, 280
Anourosorex squamipes 2, 201 Bat, Blyth’s Horseshoe 233 Bat, Halong Roundleaf 245
Aonyx sp. see Amblonyx cinereus Bat, Boonsong’s Roundleaf 15, 244 Bat, Hardwicke’s Woolly 27, 282
Apodemus draco 78, 394 Bat, Bronzed Tube-nosed 24, 275 Bat, Harlequin 23, 270
Apodemus latronum 394 Bat, Brown Tube-nosed 24, 276 Bat, Harrison’s Tube-nosed 25, 275
Apodemus orestes see A. draco Bat, Cantor’s Roundleaf 15, 243 Bat, Hill Woolly Horseshoe 230
Arctictis binturong 45, 315 Bat, Cave Nectar 7, 221 Bat, Horsfield’s Fruit 6, 218
Arctogalidia trivirgata 44, 316 Bat, Chasen’s Horseshoe 12, 236 Bat, Hutton’s Tube-nosed 24, 275
Arctonyx collaris 40, 310 Bat, Chiew Kwee’s Horseshoe 238 Bat, Indian Roundleaf 244
Arielulus circumdatus 23, 266 Bat, Chinese Horseshoe 12, 237 Bat, Indian Woolly 281
Arielulus societatis 23, 266 Bat, Clear-winged Woolly 27, 283 Bat, Indochinese Brown Horseshoe
Aselliscus dongbacana 248 Bat, Common Trident 17, 247 235
Aselliscus stoliczkanus 17, 247 Bat, Convex Horseshoe 11, 234 Bat, Indochinese Woolly 282
Atherurus macrourus 81, 399 Bat, Croslet Horseshoe 11, 232 Bat, Intermediate Horseshoe 12,
Axis porcinus 58, 344 Bat, Da Lat Tube-nosed 25, 278 235
Bat, Dark Woolly 282 Bat, Intermediate Roundleaf 14, 243
Badger, Greater Hog 40, 310 Bat, Dayak Fruit 7, 220 Bat, Johore Wrinkle-lipped 28, 287
Balaenoptera acutorostrata 54, 334 Bat, Dayak Roundleaf 16, 241 Bat, Kachin Woolly 281
Balaenoptera bonaerensis 334 Bat, Diadem Roundleaf 15, 244 Bat, Khaokhouay Roundleaf 15, 243
Balaenoptera borealis 54, 333 Bat, Disc-footed 22, 257 Bat, King Horseshoe 11, 231
Balaenoptera edeni 54, 334 Bat, Dong Bac Trident 248 Bat, Kitti’s Hog-nosed 8, 226
Balaenoptera edeni edeni see Bat, Dusky Fruit 6, 218 Bat, Kontum Tube-nosed 25, 279
B. edeni Bat, Eastern Bent-winged 26, 285 Bat, Krau Woolly 27, 283
Balaenoptera musculus 54, 333 Bat, Even-toothed Tube-nosed 25, Bat, La Touche’s Free-tailed 28,
Balaenoptera physalus 54, 333 280 286
Balionycteris maculata 220 Bat, Fawn Roundleaf 15, 243 Bat, Large Bent-winged 26, 285
Balionycteris seimundi 6, 220 Bat, Fea’s Tube-nosed 24, 276 Bat, Large-eared Roundleaf 16, 240
Bamboo Rat, Chinese 80, 397 Bat, Fiona’s Tube-nosed 25, 274 Bat, Least Horseshoe 11, 233
Bamboo Rat, Hoary 80, 397 Bat, Forest Short-nosed Fruit 6, 217 Bat, Least Roundleaf 17, 242
Bamboo Rat, Indomalayan 80, 398 Bat, Francis’s Woolly Horseshoe Bat, Least Woolly 27, 284
Bamboo Rat, Lesser 80, 397 10, 229 Bat, Lesser Asian House 23, 270
Bandicota bengalensis 377 Bat, Gilded Tube-nosed 24, 277 Bat, Lesser Bicoloured Roundleaf
Bandicota indica 73, 376 Bat, Glossy Horseshoe 11, 234 16, 240
Bandicota savilei 73, 377 Bat, Golden-collared 23, 266 Bat, Lesser Brown Horseshoe 13,
Banteng 60, 347 Bat, Golden-haired Tube-nosed 25, 235
Barbastella sp. 23, 258 278 Bat, Lesser Groove-toothed 27, 285
Barbastella darjelingensis see Bat, Great Evening 22, 269 Bat, Lesser Long-tongued Nectar
Barbastella sp. Bat, Great Roundleaf 14, 245 7, 222
Barbastella leucomelas see Bat, Great Woolly Horseshoe 10, Bat, Lesser Mouse-tailed 8, 223
Barbastella sp. 230 Bat, Lesser Sheath-tailed 8, 223
Barbastelle, Far-eastern 23, 258 Bat, Greater Asian House 23, 271 Bat, Lesser Woolly Horseshoe 10,
Bat, Acuminate Horseshoe 11, 232 Bat, Greater Bicoloured Roundleaf 229
Bat, Annamite Roundleaf 15, 242 16, 239 Bat, Little Nepalese Horseshoe 234
Bat, Annamite Tube-nosed 276 Bat, Greater Groove-toothed 27, 284 Bat, Long-winged Tomb 8, 224
Bat, Ashy Roundleaf 16, 240 Bat, Greater Long-tongued Nectar Bat, Lorelie’s Tube-nosed 25, 279
Bat, Asian Tailless Roundleaf 17, 7, 222 Bat, Mainland Greater Bamboo 22,
247 Bat, Greater Mouse-tailed 222 271
Bat, Asian Wrinkle-lipped 28, 286 Bat, Greater Short-Nosed Fruit 6, Bat, Mainland Lesser Bamboo 22,
Bat, Bala Tube-nosed 278 217 271

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 Bat, Malay Roundleaf 240 Berylmys bowersi 74, 378 Chiromyscus thomasi 75, 384
Bat, Malayan Horseshoe 13, 236 Berylmys mackenziei 74, 378 Chironax melanocephalus 6, 220
Bat, Malayan Slit-faced 9, 226 Berylmys manipulus 74, 379 Chiropodomys gliroides 78, 393
Bat, Malayan Spotted-winged 6, 220 Binturong 45, 315 Chodsigoa caovansunga 2, 201
Bat, Malaysian Tailless Roundleaf Biswamoyopterus laoensis 69, 365 Chodsigoa parca 2, 200
17, 247 Blarinella griselda 2, 198 Chrotogale owstoni 45, 316
Bat, Marshall’s Horseshoe 11, 232 Blarinella quadraticauda 199 Civet, Banded 45, 316
Bat, Medium Bent-winged 26, 286 Blarinella wardi 199 Civet, Common Palm 44, 314
Bat, Miller’s Woolly 282 Bos frontalis see Bos gaurus Civet, Large Indian 43, 313
Bat, Mountain Horseshoe 233 Bos gaurus 60, 348 Civet, Large-spotted 43, 313
Bat, Myanmar Roundleaf 241 Bos javanicus 60, 347 Civet, Malay 43, 313
Bat, Naked 28, 288 Bos sauveli 60, 348 Civet, Masked Palm 44, 315
Bat, Naked-rumped Tomb 8, 225 Bubalus arnee 60, 348 Civet, Otter 45, 316
Bat, Narrow-winged Brown 20, 263 Bubalus bubalis see Bubalus arnee Civet, Owston’s 45, 316
Bat, Northern Tailless Fruit 5, 219 Budorcas taxicolor 61, 349 Civet, Small Indian 43, 313
Bat, Painted Woolly 27, 283 Buffalo, Water 60, 348 Civet, Small-toothed Palm 44, 316
Bat, Papillose Woolly 27, 281 Bunopithecus sp. see Hoolock Civet, Taynguyen see Civet, Large
Bat, Pearson’s Horseshoe 13, 238 hoolock Indian
Bat, Pendlebury’s Roundleaf 14, 245 Coelops frithii 17, 247
Bat, Peninsular Horseshoe 12, 237 Callorhinus ursinus 49, 323 Coelops robinsoni 17, 247
Bat, Peninsular Tube-nosed 24, 273 Callosciurus caniceps 64, 357 Colugo, Sunda 29, 211
Bat, Pouched Tomb 8, 225 Callosciurus erythraeus 64, 355 Craseonycteris thonglongyai 8, 226
Bat, Ridley’s Roundleaf 15, 242 Callosciurus finlaysonii 65, 356 Crocidura annamitensis 208
Bat, Round-eared Tube-nosed 24, Callosciurus flavimanus see Crocidura attenuata 205
272 C. erythraeus Crocidura cranbrooki 208
Bat, Rozendaal’s Tube-nosed 24, Callosciurus inornatus 64, 357 Crocidura fuliginosa 3, 204
276 Callosciurus nigrovittatus 63, 355 Crocidura guy 209
Bat, Rufous Horseshoe 12, 237 Callosciurus notatus 63, 354 Crocidura hilliana 207
Bat, Shamel’s Horseshoe 11, 232 Callosciurus phayrei 63, 355 Crocidura horsfieldi see C.
Bat, Shield-faced Roundleaf 14, 246 Callosciurus prevostii 63, 354 indochinensis and C. wuchihensis
Bat, Shield-nosed Roundleaf 14, Callosciurus pygerythrus 63, 355 Crocidura indochinensis 3, 206
246 Callosciurus quinquestriatus 64, 357 Crocidura kegoensis 3, 206
Bat, Shortridge’s Horseshoe 234 Canis aureus 38, 303 Crocidura malayana 204
Bat, Siamese Horseshoe 11, 231 Canis familiaris 305 Crocidura cf. monticola 3, 205
Bat, Small Bent-winged 26, 286 Cannomys badius 80, 397 Crocidura negligens 204
Bat, Small-disc Roundleaf 15, 242 Capricornis milneedwardsi 62, 350 Crocidura phanluongi 207
Bat, Small Woolly 27, 283 Capricornis rubidus 62, 350 Crocidura phuquocensis 208
Bat, Spotted-winged Fruit 221 Capricornis sumatraensis 62, 350 Crocidura rapax 207
Bat, Sunda Free-tailed 28, 287 Cassistrellus dimissus 268 Crocidura sapaensis 209
Bat, Sunda Short-Nosed Fruit 6, 217 Cassistrellus yokdonensis 268 Crocidura sokolovi 208
Bat, Sunda Tailless Fruit 5, 218 Cat, Asian Golden 48, 321 Crocidura tanakae 205
Bat, Surat Helmeted 268 Cat, Domestic 323 Crocidura vorax 3, 207
Bat, Swinhoe’s Roundleaf 14, 246 Cat, Fishing 48, 322 Crocidura wuchihensis 206
Bat, Thai Horseshoe 13, 238 Cat, Flat-headed 48, 322 Crocidura zaitsevi 208
Bat, Thai Roundleaf 17, 241 Cat, Jungle 48, 323 Cuon alpinus 38, 305
Bat, Theobald’s Tomb 8, 224 Cat, Leopard 48, 322 Cynocephalus sp. see Galeopterus
Bat, Thomas’s Horseshoe 12, 236 Cat, Marbled 48, 320 variegatus
Bat, Titania’s Woolly 283 Catopuma temminckii 48, 321 Cynogale bennettii 45, 316
Bat, Tonkin Greater Bamboo 272 Cervus eldii 59, 343 Cynopterus cf. brachyotis ‘Forest’
Bat, Trefoil Horseshoe 10, 228 Cervus nippon 59, 343 6, 217
Bat, Walston’s Tube-nosed 25, 277 Cervus unicolor 59, 342 Cynopterus cf. brachyotis ‘Sunda’
Bat, White-collared Fruit 5, 219 Chaerephon johorensis 28, 287 6, 217
Bat, Whitehead’s Woolly 27, 284 Chaerephon plicatus 28, 286 Cynopterus horsfieldi 6, 218
Bat, Wroughton’s Free-tailed 28, Cheiromeles torquatus 28, 288 Cynopterus sphinx 6, 217
288 Chimarrogale hantu 202
Bat, Yok Don Helmeted 268 Chimarrogale himalayica 3, 202 Dacnomys millardi 73, 385
Bear, Asian Black 39, 306 Chimarrogale phaeura see C. hantu Deer, Eld’s 59, 343
Bear, Sun 39, 305 Chimarrogale styani 3, 202 Deer, Hog 58, 344
Belomys pearsonii 70, 365 Chiromyscus chiropus 383 Deer, Schomburgk’s 59, 343
Berylmys berdmorei 74, 378 Chiromyscus langbianis 384 Deer, Tufted 58, 344

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 Delphinus capensis 51, 327 Euroscaptor longirostris see E. orlovi Gibbon, Buff-cheeked 37, 302
Delphinus tropicalis see D. capensis Euroscaptor malayana 196 Gibbon, Eastern Black-crested 303
Dendrogale murina 4, 211 Euroscaptor micrura 196 Gibbon, Hainan Crested 301
Dhole 38, 305 Euroscaptor orlovi 197 Gibbon, Hoolock 37, 301
Dicerorhinus sumatrensis 55, 337 Euroscaptor parvidens 197 Gibbon, Northern White-cheeked
Diomys crumpi 74, 387 Euroscaptor parvidens ngoclinhensis 37, 302
Dog, Domestic 305 197 Gibbon, Pileated 36, 300
Dog, Raccoon 38, 304 Euroscaptor subanura 197 Gibbon, Southern White-cheeked
Dolphin, Common Bottlenose 329 37, 302
Dolphin, Fraser’s 51, 327 False-serotine, Doria’s 22, 269 Gibbon, Western Black-crested 37,
Dolphin, Indo-Pacific Bottlenose 52, False-serotine, Least 22, 269 301
329 False-serotine, Tickell’s 22, 270 Gibbon, White-handed 36, 300
Dolphin, Indo-Pacific Humpbacked False-serotine, Tomes’s 22, 269 Glischropus bucephalus 22, 263
52, 329 False-vampire, Greater 9, 227 Glischropus tylopus 262
Dolphin, Irrawaddy 52, 329 False-vampire, Lesser 9, 227 Globicephala macrorhynchus 50,
Dolphin, Long-beaked Common 51, False-vampire, Thongaree’s 9, 228 326
327 Falsistrellus affinis 21, 265 Globicephala melas 326
Dolphin, Pantropical Spotted 51, Felis catus 323 Goral, Chinese 62, 351
327 Felis chaus 48, 323 Goral, Red 62, 351
Dolphin, Risso’s 50, 326 Feresa attenuata 50, 325 Grampus griseus 50, 326
Dolphin, Rough-toothed 52, 328 Ferret-badger, Large-toothed 40, Gymnure, Chinese 1, 194
Dolphin, Spinner 51, 328 310 Gymnure, Large-eared 195
Dolphin, Striped 51, 328 Ferret-badger, Small-toothed 40, Gymnure, Short-tailed 1, 194
Douc, Black-shanked 34, 295 310
Douc, Grey-shanked 34, 296 Flying-fox, Indian 5, 216 Hapalomys delacouri 78, 390
Douc, Red-Shanked 34, 295 Flying-fox, Intermediate see Flying- Hapalomys longicaudatus 78, 390
Dremomys gularis 67, 360 fox, Indian Hare, Burmese 81, 400
Dremomys lokriah 67, 360 Flying-fox, Island 5, 216 Hare, Chinese 81, 401
Dremomys pernyi 67, 360 Flying-fox, Large 5, 215 Harpiocephalus harpia 26, 280
Dremomys pyrrhomerus 67, 360 Flying-fox, Lyle’s 5, 216 Harpiocephalus mordax see N. harpia
Dremomys rufigenis 67, 360 Flying Squirrel, Black 69, 364 Harpiola isodon 25, 280
Dugong 52, 335 Flying Squirrel, Chindwin Giant 68, Helarctos malayanus 39, 305
Dugong dugon 52, 335 364 Hemigalus derbyanus 45, 316
Dyacopterus spadiceus 7, 220 Flying Squirrel, Grey-cheeked 71, Herpestes sp. see Urva sp.
367 Hesperoptenus blanfordi 22, 269
Echinosorex gymnurus 1, 194 Flying Squirrel, Hairy-footed 70, 365 Hesperoptenus doriae 22, 269
Elaphodus cephalophus 58, 344 Flying Squirrel, Horsfield’s 70, 366 Hesperoptenus tickelli 22, 270
Elephant, Asian 55, 335 Flying Squirrel, Indian Giant 68, 363 Hesperoptenus tomesi 22, 269
Elephas maximus 55, 335 Flying Squirrel, Lao Giant 69, 365 Hipposideros alongensis 245
Emballonura monticola 8, 223 Flying Squirrel, Lesser Giant 69, 364 Hipposideros armiger 14, 245
Eonycteris spelaea 7, 221 Flying Squirrel, Malaysian Pygmy Hipposideros atrox 16, 240
Eothenomys cachinus 79, 396 71, 368 Hipposideros bicolor 16, 239
Eothenomys melanogaster 79, 396 Flying Squirrel, Particoloured 70, Hipposideros cervinus 15, 243
Eothenomys miletus see E. cachinus 367 Hipposideros cf. cineraceus 16, 240
Episoriculus baileyi 200 Flying Squirrel, Phayre’s 70, 368 Hipposideros diadema 15, 244
Episoriculus caudatus 2, 200 Flying Squirrel, Red-cheeked 71, Hipposideros doriae 17, 242
Episoriculus leucops see E. baileyi 367 Hipposideros dyacorum 16, 241
Episoriculus macrurus 2, 200 Flying Squirrel, Red Giant 68, 362 Hipposideros einnaythu 241
Eptesicus andersoni 267 Flying Squirrel, Smoky 70, 365 Hipposideros galeritus 15, 243
Eptesicus dimissus see Cassistrellus Flying Squirrel, Temminck’s 71, 366 Hipposideros grandis see H. cf.
dimissus Flying Squirrel, Vordermann’s 71, larvatus
Eptesicus pachyomus 23, 267 366 Hipposideros griffini 245
Eptesicus pachyotis 267 Flying Squirrel, Whiskered 71, 366 Hipposideros halophyllus 17, 241
Eptesicus serotinus see Flying Squirrel, Yunnan Giant 68, Hipposideros khaokhouayensis 15,
E. pachyomus 363 243
Eudiscoderma thongareeae 9, 228 Fox, Red 38, 304 Hipposideros lankadiva 244
Eudiscopus denticulus 22, 257 Hipposideros lankadiva gyi see
Euroscaptor grandis 1, 196 Galeopterus variegatus 29, 211 H. lankadiva
Euroscaptor klossi 1, 196 Gaur 60, 348 Hipposideros cf. larvatus 14, 243
Euroscaptor kuznetsovi 197 Gibbon, Agile 36, 300 Hipposideros lekaguli 15, 244

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 Hipposideros leptophylla see H. cf. Lagenodelphis hosei 51, 327 Manis pentadactyla 29, 192
larvatus Langur, Annamese Silvered 30, 292 Marmoset-rat, Greater 78, 390
Hipposideros lylei 14, 246 Langur, Banded 30, 290 Marmoset-rat, Lesser 78, 390
Hipposideros nequam 16, 240 Langur, Capped 31, 294 Marten, Stone 40, 307
Hipposideros orbiculus 15, 242 Langur, Cat Ba 32, 293 Marten, Yellow-throated 40, 307
Hipposideros pendleburyi 14, 245 Langur, Delacour’s 32, 294 Martes flavigula 40, 307
Hipposideros pomona 16, 240 Langur, Dusky 31, 292 Martes foina 40, 307
Hipposideros ridleyi 15, 242 Langur, Francois’s 32, 293 Maxomys, Indochinese 76, 389
Hipposideros rotalis 15, 242 Langur, Hatinh 32, 294 Maxomys, Malayan Mountain 76,
Hipposideros sabanus see H. doriae Langur, Indochinese Black 32, 294 388
Hipposideros scutinares 14, 246 Langur, Indochinese Grey 31, 293 Maxomys, Rajah 76, 387
Hipposideros swinhoei 14, 246 Langur, Indochinese Silvered 30, Maxomys, Red Spiny 76, 388
Hipposideros turpis see H. alongensis 291 Maxomys, Whitehead’s 76, 388
and H. pendleburyi Langur, Lao 32, 294 Maxomys inas 76, 388
Hoolock hoolock 37, 301 Langur, Pale-thighed 30, 290 Maxomys moi 76, 389
Hylobates agilis 36, 300 Langur, Phayre’s 31, 293 Maxomys rajah 76, 387
Hylobates lar 36, 300 Langur, Shortridge’s 31, 295 Maxomys surifer 76, 388
Hylobates pileatus 36, 300 Langur, Sundaic Silvered 30, 291 Maxomys whiteheadi 76, 388
Hylomys megalotis 195 Langur, Tenasserim 31, 292 Megaderma spasma 9, 227
Hylomys sinensis 1, 194 Laonastes aenigmamus 80, 398 Megaerops albicollis see M. wetmorei
Hylomys suillus 1, 194 Lariscus insignis 67, 361 Megaerops ecaudatus 5, 218
Hylopetes alboniger 70, 367 Lenothrix canus 74, 389 Megaerops niphanae 5, 219
Hylopetes lepidus see H. platyurus Lenothrix malaisia see L. canus Megaerops wetmorei 5, 219
Hylopetes phayrei 70, 368 Leopard 47, 320 Megaptera novaeangliae 54, 334
Hylopetes platyurus 71, 367 Leopard, Mainland Clouded 47, 320 Melogale cucphuongensis see
Hylopetes spadiceus 71, 367 Leopoldamys ciliatus 386 M. moschata
Hypsugo anthonyi 265 Leopoldamys edwardsi 76, 386 Melogale moschata 40, 310
Hypsugo cadornae 21, 265 Leopoldamys herberti 385 Melogale personata 40, 310
Hypsugo dolichodon 264 Leopoldamys milleti 386 Menetes berdmorei 66, 361
Hypsugo joffrei 265 Leopoldamys neilli 76, 386 Mesoplodon densirostris 53, 332
Hypsugo lophurus 264 Leopoldamys sabanus 76, 385 Mesoplodon ginkgodens 53, 332
Hypsugo macrotis 21, 264 Lepus peguensis 81, 400 Micromys erythrotis 78, 393
Hypsugo pulveratus 21, 263 Lepus sinensis 81, 401 Micromys minutus see M. erythrotis
Hystrix brachyura 81, 399 Linsang, Banded 44, 319 Microtus clarkei 79, 396
Linsang, Spotted 44, 319 Millardia kathleenae 74, 387
Ia io 22, 269 Loris, Asian Slow 29, 289 Miniopterus fuliginosus 26, 285
Indopacetus pacificus 332 Loris, Pygmy 29, 289 Miniopterus magnater 26, 285
Iomys horsfieldi 70, 366 Loris, Sunda Slow 29, 289 Miniopterus medius 26, 286
Lutra lutra 42, 311 Miniopterus pusillus 26, 286
Jackal, Golden 38, 303 Lutra sumatrana 42, 311 Miniopterus schreibersii see
Lutrogale perspicillata 42, 312 M. fuliginosus
Kerivoula depressa 282 Lyroderma lyra 9, 227 Mogera insularis see M. latouchei
Kerivoula dongduongana 282 Mogera latouchei 198
Kerivoula flora see K. titania Macaca arctoides 35, 297 Mole, Blyth’s 197
Kerivoula furva 282 Macaca assamensis 35, 298 Mole, Himalayan 196
Kerivoula hardwickii 27, 282 Macaca fascicularis 35, 298 Mole, Kloss’s 1, 196
Kerivoula intermedia 27, 283 Macaca leonina 35, 297 Mole, Kuznetsov’s 197
Kerivoula kachinensis 281 Macaca mulatta 35, 298 Mole, La Touche’s 198
Kerivoula krauensis 27, 283 Macaca nemestrina 35, 297 Mole, Large Chinese 1, 196
Kerivoula lenis 281 Macaque, Assamese 35, 298 Mole, Long-tailed 1, 198
Kerivoula minuta 27, 284 Macaque, Long-tailed 35, 298 Mole, Malayan 196
Kerivoula papillosa 27, 281 Macaque, Northern Pig-tailed 35, Mole, Orlov’s 197
Kerivoula pellucida 27, 283 297 Mole, Short-tailed 197
Kerivoula picta 27, 283 Macaque, Rhesus 35, 298 Mole, Small-toothed 197
Kerivoula titania 283 Macaque, Southern Pig-tailed 35, Mole-shrew 2, 201
Kerivoula whiteheadi 27, 284 297 Mongoose, Crab-eating 46, 318
Kha-nyou 80, 398 Macaque, Stump-tailed 35, 297 Mongoose, Indian Grey 46, 318
Kogia breviceps 53, 331 Macroglossus minimus 7, 222 Mongoose, Javan 46, 317
Kogia sima 53, 331 Macroglossus sobrinus 7, 222 Mongoose, Short-tailed 46, 317
Kouprey 60, 348 Manis javanica 29, 192 Mongoose, Small Indian 318

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 Monkey, Myanmar Snub-nosed 33, Murina harrisoni 25, 275 Myotis annatessae 252
296 Murina huttoni 24, 275 Myotis annectans 18, 254
Monkey, Tonkin Snub-nosed 33, Murina kontumensis 25, 279 Myotis ater 18, 252
296 Murina leucogaster 24, 279 Myotis borneoensis 253
Moonrat 1, 194 Murina lorelieae 25, 279 Myotis chinensis 19, 250
Mops mops 28, 287 Murina peninsularis 24, 273 Myotis daubentoni see M. laniger
Moschus berezovskii 58, 342 Murina rozendaali 24, 276 Myotis federatus 254
Moschus fuscus 58, 342 Murina suilla 24, 276 Myotis formosus 18, 251
Mouse, Asian Harvest 78, 393 Murina tiensa see M. harrisoni Myotis hasseltii 19, 255
Mouse, Asian House 77, 390 Murina tubinaris see M. feae Myotis hermani 251
Mouse, Asian Long-tailed Climbing Murina walsoni 25, 277 Myotis horsfieldii 19, 255
77, 393 Mus booduga 77, 392 Myotis indochinensis 18, 253
Mouse, Cook’s 77, 392 Mus caroli 77, 391 Myotis laniger 19, 257
Mouse, Fawn-coloured 77, 391 Mus castaneus see M. musculus Myotis montivagus 252
Mouse, Fragile-tailed 391 Mus cervicolor 77, 391 Myotis muricola 18, 251
Mouse, Indochinese Shrewlike 77, Mus cervicolor popaeus see Myotis mystacinus 18, 252
392 M. cervicolor Myotis oreias 255
Mouse, Large-eared Wood 394 Mus cookii 77, 392 Myotis phanluongi 256
Mouse, Little Indian Field 77, 392 Mus fragilicauda 391 Myotis pillosus 19, 256
Mouse, Ricefield 77, 391 Mus musculus 77, 390 Myotis ricketti see M. pillosus
Mouse, Shortridge’s 77, 392 Mus nagarum see M. cookii Myotis ridleyi 18, 255
Mouse, South China Wood 78, 394 Mus pahari 77, 392 Myotis rosseti 18, 254
Mouse, Vernay’s Climbing 78, 394 Mus shortridgei 77, 392 Myotis rufoniger 18, 250
Mousedeer, Greater 57, 340 Musk-deer, Black 58, 342 Myotis siligorensis 19, 256
Mousedeer, Lesser 57, 340 Musk-deer, Forest 58, 342 Myotis siligorensis alticraniatus see
Mousedeer, Silver-backed 57, 341 Mustela kathiah 41, 308 M. siligorensis
Muntiacus crinifrons see Mustela nivalis see M. tonkinensis
M. gongshanensis Mustela nudipes 41, 308 Naemorhedus baileyi 62, 351
Muntiacus feae 57, 346 Mustela sibirica 41, 308 Naemorhedus griseus 62, 351
Muntiacus gongshanensis 57, 346 Mustela strigidorsa 41, 309 Nectogale elegans 3, 202
Muntiacus muntjak 57, 344 Mustela tonkinensis 41, 308 Neodon forresti 79, 396
Muntiacus puhoatensis 346 Myotis, Anna Tess’s 252 Neodon irene see N. forresti
Muntiacus putaoensis 57, 346 Myotis, Annamite 257 Neofelis diardi see N. nebulosa
Muntiacus rooseveltorum 345 Myotis, Asian Whiskered 18, 251 Neofelis nebulosa 47, 320
Muntiacus truongsonensis 57, 345 Myotis, Australasian Large-footed Neophocaena asiaeorientalis see
Muntiacus vaginalis see M. muntjak 255 N. phocaenoides
Muntiacus vuquangensis 58, 347 Myotis, Black-and-orange Woolly Neophocaena phocaenoides 52, 330
Muntjac, Annamite 57, 345 18, 250 Neotetracus sp. see Hylomys sinensis
Muntjac, Fea’s 57, 346 Myotis, Chinese 19, 250 Nesolagus timminsi 81, 401
Muntjac, Gongshan 57, 346 Myotis, Eurasian Whiskered 18, 252 Niviventer, Brahman 75, 383
Muntjac, Large-antlered 58, 347 Myotis, Hairy-faced 18, 254 Niviventer, Cameron Highlands 75,
Muntjac, Leaf 57, 346 Myotis, Hasselt’s 19, 255 381
Muntjac, Northern Red see Muntjac, Myotis, Herman’s Woolly 251 Niviventer, Confucian 75, 381
Red Myotis, Hodgson’s Woolly 18, 251 Niviventer, Dark-tailed 75, 381
Muntjac, Puhoat 346 Myotis, Horsfield’s 19, 255 Niviventer, Indochinese Mountain
Muntjac, Red 57, 344 Myotis, Indochinese Water 19, 257 382
Muntjac, Roosevelt’s 345 Myotis, Large Brown 252 Niviventer, Indomalayan 75, 382
Murina aenea 24, 275 Myotis, Large Indochinese 18, 253 Niviventer, Limestone 75, 382
Murina annamitica 276 Myotis, Large Malayan 254 Niviventer, Smoke-bellied 75, 383
Murina aurata see M. eleryi Myotis, Peters’s 18, 252 Niviventer brahma 75, 383
Murina balaensis 278 Myotis, Phan Luong’s 256 Niviventer bukit see N. fulvescens
Murina beelzebub 25, 277 Myotis, Rickett’s 19, 256 Niviventer cameroni 75, 381
Murina chrysochaetes 25, 278 Myotis, Ridley’s 18, 255 Niviventer confucianus 75, 381
Murina cineracea see M. feae Myotis, Singapore 255 Niviventer cremoriventer 75, 381
Murina cyclotis 24, 272 Myotis, Small-toothed 19, 256 Niviventer eha 75, 383
Murina eleryi 24, 277 Myotis, Szechuan 19, 250 Niviventer fulvescens 75, 382
Murina feae 24, 276 Myotis, Thick-thumbed 18, 254 Niviventer hinpoon 75, 382
Murina fionae 25, 274 Myotis adversus 255 Niviventer huang see N. fulvescens
Murina guilleni 274 Myotis altarium 19, 250 Niviventer langbianis see
Murina harpioloides 25, 278 Myotis annamiticus 257 Chiromyscus langbianis

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 Niviventer niviventer see Physeter macrocephalus 54, 330 Pygathrix nemaeus 34, 295
N. confucianus Pig, Banded see Pig, Eurasian Wild Pygathrix nigripes 34, 295
Niviventer rapit see N. cameroni Pig, Bearded 56, 339 Pygmy-dormouse, Soft-furred 78,
Niviventer tenaster 382 Pig, Eurasian Wild 56, 339 395
Noctule, Asian 23, 262 Pig, Vietnamese Warty 340
Nomascus annamensis see Pika, Forrest’s 400 Rabbit, Annamite Striped 81, 401
N. gabriellae Pika, Moupin 81, 400 Rat, Annandale’s 73, 376
Nomascus concolor 37, 301 Pipistrelle, Anthony’s 265 Rat, Berdmore’s 74, 378
Nomascus gabriellae 37, 302 Pipistrelle, Benom Gilded 23, 266 Rat, Bowers’s 74, 378
Nomascus hainanus 301 Pipistrelle, Big-eared 21, 264 Rat, Crump’s Soft-furred 74, 387
Nomascus leucogenys 37, 302 Pipistrelle, Black Gilded 23, 266 Rat, Edward’s Giant 76, 386
Nomascus nasutus 303 Pipistrelle, Cadorna’s 21, 265 Rat, Fea’s Tree 383
Nomascus siki 37, 302 Pipistrelle, Chinese 21, 263 Rat, Greater Bandicoot 73, 376
Nyctalus cf. labiata 23, 262 Pipistrelle, Chocolate 21, 265 Rat, Grey Tree 74, 389
Nyctalus noctula see N. cf. labiata Pipistrelle, Coromandel 20, 260 Rat, Herbert’s Giant 385
Nyctereutes procyonoides 38, 304 Pipistrelle, Indochinese Thick- Rat, House 72, 372
Nycteris javanica see N. tragata thumbed 22, 263 Rat, Indochinese Forest 73, 374
Nycteris tragata 9, 226 Pipistrelle, Japanese 20, 260 Rat, Indochinese Tree 384
Nycticebus bengalensis 29, 289 Pipistrelle, Javan 20, 259 Rat, Lao Limestone 74, 379
Nycticebus coucang 29, 289 Pipistrelle, Joffre’s 265 Rat, Lesser Bandicoot 377
Nycticebus pygmaeus 29, 289 Pipistrelle, Kelaart’s 20, 261 Rat, Lesser Ricefield 72, 375
Pipistrelle, Least 20, 261 Rat, Long-tailed Giant 76, 385
Ochotona forresti 400 Pipistrelle, Long-toothed 264 Rat, Losea 375
Ochotona thibetana 81, 400 Pipistrelle, Mount Popa 21, 260 Rat, Mackenzie’s 378
Orcaella brevirostris 52, 329 Pipistrelle, Myanmar 264 Rat, Malayan Woolly Tree 74, 389
Orcinus orca 50, 326 Pipistrelle, Narrow-winged 20, 261 Rat, Malaysian Wood 72, 373
Otomops wroughtoni 28, 288 Pipistrelle, Sunda Thick-thumbed Rat, Manipur 74, 379
Otter, Eurasian 42, 311 262 Rat, Millard’s Giant 73, 385
Otter, Hairy-nosed 42, 311 Pipistrellus abramus 20, 260 Rat, Millet’s Giant 386
Otter, Oriental Small-clawed 42, 312 Pipistrellus ceylonicus 20, 261 Rat, Müller’s 73, 380
Otter, Smooth 42, 312 Pipistrellus coromandra 20, 260 Rat, Neill’s 76, 386
Pipistrellus javanicus 20, 259 Rat, Norway 73, 374
Paguma larvata 44, 315 Pipistrellus paterculus 21, 260 Rat, Osgood’s 72, 376
Panda, Red 39, 306 Pipistrellus stenopterus 20, 261 Rat, Pacific 72, 374
Pangolin, Chinese 29, 192 Pipistrellus tenuis 20, 261 Rat, Popa Soft-furred 74, 387
Pangolin, Sunda 29, 192 Pithecheir melanurus see P. parvus Rat, Ricefield 72, 373
Panthera pardus 47, 320 Pithecheir parvus 74, 389 Rat, Savile’s Bandicoot 73, 377
Panthera tigris 47, 320 Porcupine, Brush-tailed 81, 399 Rat, Sundaic Mountain 386
Paracoelops megalotis 247 Porcupine, Long-tailed 81, 399 Rat, Thomas’s Tree 75, 384
Paradoxurus hermaphroditus 44, 314 Porcupine, Malayan 81, 399 Rat, Tonkin Limestone 74, 379
Parascaptor leucura 197 Porpoise, Indo-Pacific Finless 52, 330 Rat, White-footed Indochinese 73,
Pardofelis marmorata 48, 320 Presbytis femoralis 30, 290 375
Penthetor lucasi 6, 218 Presbytis siamensis 30, 290 Rattus andamanensis 73, 374
Peponocephala electra 50, 325 Prionailurus bengalensis 48, 322 Rattus annandalei 73, 376
Petaurillus hosei see P. kinlochii Prionailurus planiceps 48, 322 Rattus argentiventer 72, 373
Petaurillus kinlochii 71, 368 Prionailurus viverrinus 48, 322 Rattus exulans 72, 374
Petaurista elegans 69, 364 Prionodon linsang 44, 319 Rattus hoxaensis see R. argentiventer
Petaurista petaurista 68, 362 Prionodon pardicolor 44, 319 Rattus koratensis see
Petaurista philippensis 68, 363 Pseudois nayaur 61, 350 R. andamanensis
Petaurista sybilla 68, 364 Pseudorca crassidens 50, 325 Rattus losea 375
Petaurista yunanensis 68, 363 Pseudoryx nghetinhensis 61, 349 Rattus nitidus 73, 375
Petinomys genibarbis 71, 366 Pteromyscus pulverulentus 70, 365 Rattus norvegicus 73, 374
Petinomys setosus 71, 366 Pteropus giganteus 5, 216 Rattus osgoodi 72, 376
Petinomys vordermanni 71, 366 Pteropus hypomelanus 5, 216 Rattus rattus 72, 372
Philetor brachypterus 20, 263 Pteropus intermedius see Rattus remotus see R. andamanensis
Phoca largha 49, 324 P. giganteus Rattus sakeratensis 72, 375
Phoniscus atrox 27, 285 Pteropus lylei 5, 216 Rattus sikkimensis see
Phoniscus jagorii 27, 284 Pteropus vampyrus 5, 215 R. andamanensis
Physeter catadon see Ptilocercus lowii 4, 209 Rattus tanezumi see R. rattus
P. macrocephalus Pygathrix cinerea 34, 296 Rattus tiomanicus 72, 373

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 Ratufa affinis 63, 353 Rousettus amplexicaudatus 7, 214 Shrew, Wuchih 206
Ratufa bicolor 63, 353 Rousettus leschenaultii 7, 215 Shrew, Zaitsev’s 208
Rhinoceros, Asian Two-horned 55, Rucervus eldii see Cervus eldii Shrew-mole, Slender 1, 195
337 Rucervus schomburgki 59, 343 Siamang 36, 299
Rhinoceros, Lesser One-horned 55, Rusa unicolor see Cervus unicolor Sika 59, 343
338 Sorex bedfordiae 2, 198
Rhinoceros sondaicus 55, 338 Saccolaimus saccolaimus 8, 225 Sorex cylindricauda 198
Rhinolophus acuminatus 11, 232 Sambar 59, 342 Soriculus nigrescens 199
Rhinolophus affinis 12, 235 Saola 61, 349 Sousa chinensis 52, 329
Rhinolophus borneensis see Saxatilomys paulinae 74, 379 Sousa plumbea see S. chinensis
R. chaseni Scaptonyx fuscicaudus 1, 198 Sousa sahulensis see S. chinensis
Rhinolophus chaseni 12, 236 Scotomanes ornatus 23, 270 Sphaerius blanfordi 6, 221
Rhinolophus chiewkweeae 238 Scotophilus heathii 23, 271 Squirrel, Black Giant 63, 353
Rhinolophus coelophyllus 11, 232 Scotophilus kuhlii 23, 270 Squirrel, Cambodian Striped 66, 358
Rhinolophus convexus 11, 234 Seal, Northern Fur 49, 323 Squirrel, Cream-coloured Giant 63,
Rhinolophus francisi 10, 229 Seal, Spotted 49, 324 353
Rhinolophus francisi thailandicus see Serotine, Asian 23, 267 Squirrel, Eastern Striped 66, 358
R. francisi Serotine, Thick-eared 267 Squirrel, Grey-bellied 64, 357
Rhinolophus hirsutus 231 Serow, Indochinese 62, 350 Squirrel, Horse-tailed 66, 359
Rhinolophus lepidus 233 Serow, Red 62, 350 Squirrel, Indochinese Ground 66,
Rhinolophus luctoides 230 Serow, Southern 62, 350 361
Rhinolophus luctus 10, 230 Sheep, Blue 61, 350 Squirrel, Inornate 64, 357
Rhinolophus macrotis 11, 230 Shrew, Annamite 208 Squirrel, Irrawaddy 63, 355
Rhinolophus malayanus 13, 236 Shrew, Asian Grey 205 Squirrel, Low’s 66, 359
Rhinolophus marshalli 11, 232 Shrew, Bailey’s Brown-toothed 200 Squirrel, Orange-bellied 67, 360
Rhinolophus microglobosus 235 Shrew, Chinese 207 Squirrel, Pallas’s 64, 355
Rhinolophus monticolus 233 Shrew, Cranbrook’s 208 Squirrel, Perny’s Long-nosed 67,
Rhinolophus morio see R. luctus Shrew, Greater Stripe-backed 198 360
Rhinolophus paradoxolophus see Shrew, Guy’s 209 Squirrel, Phayre’s 63, 355
R. rex Shrew, Hill’s 207 Squirrel, Plantain 63, 354
Rhinolophus pearsonii 13, 238 Shrew, Himalayan Water 3, 202 Squirrel, Prevost’s 63, 354
Rhinolophus pusillus 11, 233 Shrew, Hodgson’s Brown-toothed Squirrel, Red-cheeked 67, 360
Rhinolophus refulgens 11, 234 2, 200 Squirrel, Red-hipped 67, 360
Rhinolophus rex 11, 231 Shrew, House 3, 203 Squirrel, Red-throated 67, 360
Rhinolophus robinsoni 12, 237 Shrew, Indochinese 3, 206 Squirrel, Shrew-faced Ground 67,
Rhinolophus robinsoni thaianus see Shrew, Indochinese Short-tailed 2, 361
R. robinsoni 199 Squirrel, Slender 66, 359
Rhinolophus rouxii 12, 237 Shrew, Ke Go 3, 206 Squirrel, Stripe-bellied 64, 357
Rhinolophus sedulus 10, 229 Shrew, Lesser Stripe-backed 2, 198 Squirrel, Sunda Black-banded 63,
Rhinolophus shameli 11, 232 Shrew, Long-tailed Brown-toothed 355
Rhinolophus shortridgei 234 2, 200 Squirrel, Swinhoe’s Striped 66, 358
Rhinolophus siamensis 11, 231 Shrew, Lowe’s Brown-toothed 2, Squirrel, Three-striped Ground 67,
Rhinolophus sinicus 12, 237 200 361
Rhinolophus stheno 13, 235 Shrew, Malayan 204 Squirrel, Variable 65, 356
Rhinolophus subbadius 234 Shrew, Malayan Pygmy 203 Squirrel, Western Striped 66, 358
Rhinolophus thailandensis 13, 238 Shrew, Malayan Water 202 Stenella attenuata 51, 327
Rhinolophus thomasi 12, 236 Shrew, Myanmar Short-tailed 199 Stenella coeruleoalba 51, 328
Rhinolophus trifoliatus 10, 228 Shrew, Peninsular 204 Stenella longirostris 51, 328
Rhinolophus yunanensis see Shrew, Phan Luong’s 207 Stenella longirostris roseiventris see
R. thailandensis Shrew, Phu Quoc 208 S. longirostris
Rhinopithecus avunculus 33, 296 Shrew, Pygmy White-toothed 3, 203 Steno bredanensis 52, 328
Rhinopithecus strykeri 33, 296 Shrew, Sa Pa 209 Suncus etruscus 3, 203
Rhinopoma hardwickii 8, 223 Shrew, Sokolov’s 208 Suncus malayanus 203
Rhinopoma microphyllum 222 Shrew, South-east Asian 3, 204 Suncus murinus 3, 203
Rhinosciurus laticaudatus 67, 361 Shrew, Styan’s Water 3, 202 Sundamys muelleri 73, 380
Rhizomys pruinosus 80, 397 Shrew, Sunda 3, 205 Sundasciurus hippurus 66, 359
Rhizomys sinensis 80, 397 Shrew, Taiwanese Grey 205 Sundasciurus lowii 66, 359
Rhizomys sumatrensis 80, 398 Shrew, Van Sung’s 2, 201 Sundasciurus tenuis 66, 359
Rousette, Geoffroy’s 7, 214 Shrew, Voracious 3, 207 Sus barbatus 56, 339
Rousette, Leschenault’s 7, 215 Shrew, Web-footed Water 3, 202 Sus bucculentus 340

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 Sus scrofa 56, 339 Tragulus kanchil 57, 340 Viverra tangalunga 43, 313
Sus scrofa vittatus see S. scrofa Tragulus napu 57, 340 Viverra zibetha 43, 313
Symphalangus syndactylus 36, 299 Tragulus versicolor 57, 341 Viverricula indica 43, 313
Tragulus williamsoni see T. kanchil Vole, Clarke’s 79, 396
Tadarida latouchei 28, 286 Tree-mouse, Indomalayan Pencil- Vole, Forrest’s Mountain 79, 396
Tadarida teniotis see T. latouchei tailed 78, 393 Vole, Kachin 79, 396
Takin 61, 349 Treeshrew, Common 4, 210 Vole, Père David’s 79, 396
Tamiops maritimus 66, 358 Treeshrew, Feather-tailed 4, 209 Volemys sp. see Microtus clarkei
Tamiops mcclellandii 66, 358 Treeshrew, Lesser 4, 211 Vulpes vulpes 38, 304
Tamiops rodolphii 66, 358 Treeshrew, Northern 4, 210
Tamiops swinhoei 66, 358 Treeshrew, Northern Slender-tailed Weasel, Malay 41, 308
Taphozous longimanus 8, 224 4, 211 Weasel, Siberian 41, 308
Taphozous melanopogon 8, 224 Trichys fasciculata 81, 399 Weasel, Stripe-backed 41, 309
Taphozous nudiventris 8, 225 Tupaia belangeri 4, 210 Weasel, Tonkin 41, 308
Taphozous theobaldi 8, 224 Tupaia glis 4, 210 Weasel, Yellow-bellied 41, 308
Tapir, Asian 56, 337 Tupaia minor 4, 211 Whale, Antarctic Minke 334
Tapirus indicus 56, 337 Tursiops aduncus 52, 329 Whale, Blainville’s Beaked 53, 332
Thainycteris aureocollaris 23, 266 Tursiops truncatus 329 Whale, Blue 54, 333
Tiger 47, 320 Tylonycteris fulvida 22, 271 Whale, Bryde’s 54, 334
Tonkinomys daovantieni 74, 379 Tylonycteris malayana 22, 271 Whale, Cuvier’s Beaked 53, 331
Trachypithecus barbei 31, 292 Tylonycteris pachypus see T. fulvida Whale, Dwarf Sperm 53, 331
Trachypithecus crepusculus 31, 293 Tylonycteris robustula see Whale, False Killer 50, 325
Trachypithecus cristatus 30, 291 T. malayana Whale, Fin 54, 333
Trachypithecus delacouri 32, 294 Tylonycteris tonkinensis 272 Whale, Ginkgo-toothed Beaked 53,
Trachypithecus ebenus 32, 294 Typhlomys capensis see T. cinereus 332
Trachypithecus francoisi 32, 293 Typhlomys cinereus 78, 395 Whale, Great Sperm 54, 330
Trachypithecus germaini 30, 291 Whale, Humpback 54, 334
Trachypithecus hatinhensis 32, 294 Uropsilus gracilis 1, 195 Whale, Killer 50, 326
Trachypithecus laotum 32, 294 Ursus thibetanus 39, 306 Whale, Long-finned Pilot 326
Trachypithecus margarita 30, 292 Urva auropunctata 318 Whale, Longman’s Beaked 332
Trachypithecus obscurus 31, 292 Urva brachyura 46, 317 Whale, Melon-headed 50, 325
Trachypithecus phayrei 31, 293 Urva edwardsii 46, 318 Whale, Minke 54, 334
Trachypithecus pileatus 31, 294 Urva javanica 46, 317 Whale, Pygmy Killer 50, 325
Trachypithecus poliocephalus 32, Urva urva 46, 318 Whale, Pygmy Sperm 53, 331
293 Whale, Sei 54, 333
Trachypithecus selangorensis see Vandeleuria oleracea 77, 393 Whale, Short-finned Pilot 50, 326
T. cristatus Vernaya fulva 78, 394 Whale, Sittang see Whale, Bryde’s
Trachypithecus shortridgei 31, 295 Viverra megaspila 43, 313
Tragulus javanicus see T. kanchil Viverra tainguensis see V. zibetha Ziphius cavirostris 53, 331

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