identification keys to the mammals of borneo

BBEC Publication No.7
IDENTIFICATION KEYS TO THE MAMMALS OF BORNEO
Written by:
Dr. Shigeki Yasuma,
Mr. Mahedi Andau,
Mr. Ludi Apin,
Mr. Fred Tuh Yit Yu,
Ms. Lucy Kimsui,
JICA Short-term Expert
Director, Sabah Wildlife Department
Manager, Crocker Range Park, Sabah Parks
Assistant Research Officer, Zoology Unit,Kinabalu Park, Sabah Parks
Research Assistant, Institute for Tropical Biology & Conservation, Universiti Malaysia Sabah
Preface by:
Datuk Lamri Ali, Director of Sabah Parks
Mr. Takahisa Kusano, Chief Advisor for BBEC Programme, JICA
Copyright © 2003, Sabah Parks and JICA
ISBN 983-2770-01-7
Published by:
Park Management Component
BBEC Programme
c/o Sabah Parks
Lot 1-3, Block K, Sinsuran Complex
P. O. Box 10626
88806 Kota Kinabalu, Sabah
Tel : +6 - 088 - 211881 / 212719
Fax : +6 - 088 - 221001 / 211585
C O N T E N T S
P R E F A C E
K E Y
T O
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O R D E R S
P A R T
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I N S E C T I V O R A
S C A N D E N T I A
R O D E N T I A
P A R T
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I I
C H I R O P T E R A
43
P R E F A C E
N
ine organizations of the Sabah State
For the purpose of introducing the survey and
Government, Universiti Malaysia Sabah
idenfification techniques, and to be able to assist in
(UMS) and several NGOs have been
the study of mammal distribution in Sabah, JICA has
implementing a joint endeavor of “Bornean
dispatched an expert on mammal identification, Dr.
Biodiversity and Ecosystems Conservation (BBEC)
Shigeki Yasuma from April until November 2002. He
Programme in Sabah, Malaysia” with the technical
had previous experiences working with Sabah Wildlife
cooperation of the Japanese Government through Japan
Department and published a manual titled
International Cooperation Agency (JICA) since
“Identification Key to the Mammals of Sabah”
February 2002 until January 2007, for a period of 5
together with Mr. Mahedi Andau, the director of the
years.
Wildlife Department. Based on this book, Dr. Yasuma
gave training to the staff of Sabah Parks, Sabah Wildlife
To conserve the endangered and precious biodiversity
Department and UMS and jointly worked and studied
and ecosystems of Sabah, the BBEC Programme is
the mammals of Crocker Range Park together with
working towards establishing “Comprehensive and
Mr. Ludi Apin Fred and Mr. Tuh Yit Yu of Sabah Parks
sustainable conservation approaches of biodiversity
and Ms. Lucy Kimsui of ITBC, UMS, then updated
and ecosystems,” by integrating the efforts of various
the information, especially insectivora, scandentia,
governmental organizations, a university and NGOs.
rodentia and chiroptera.
The BBEC Programme has been planned, implemented
and monitored by the Sabah state agencies namely;
We are pleased to present one of the results of this
Sabah Parks, Wildlife Department, Science &
highly meaningful technical cooperation and joint work
Technology Unit, Forestry Department, the eight
of Japanese and several Sabahans. We believe this book
district offices related to Crocker Range Park, Land &
will serve as a reference for many researchers, rangers
Surveys Department, Environmental Conservation
and students in Borneo to pursue and perform their
Department and Drainage & Irrigation Department,
work more efficiently and effectively.
Universiti Malaysia Sabah, Environmental Action
Committee and other NGOs with the assistance of
March 2003
Japanese advisors and experts dispatched by JICA.
Datuk Lamri bin Ali
Identifying wildlife and making their inventory are the
Director of Sabah Parks
very basic and essential conducts that lead us to
(Head of Park Management Component,
understand the existing biodiversity in Sabah and
BBEC Programme)
Borneo. The status of ecosystems, the needs of
protection and where and how to conserve them are
Takahisa Kusano
our immediate concerns. The mammals in Borneo,
Chief Advisor for BBEC Programme
especially relatively smaller species are to be studied
Japan International Cooperation Agency (JICA)
more and we need to understand their habits,
population, distribution, ecologies and consequently
present knowledge on the status of biodiversity and
ecosystems in Borneo and their conservation needs.
1
Key to orders
1
Body covered with imbricate scales. Hair on the belly only.
Pangolin Fig. 1-1 ............................................................................. PHOLIDOTA
1’
Body hairy or naked .................................................................................................... 2
2 (1’)
With a flying membrane between limbs ...................................................................... 3
2’
With no flying membrane ............................................................................................ 5
3 (2)
Forearm adapted for sustained flight; second through fifth digits are greatly extended
and wholly enclosed in flying membrane, only thumb being free and normal.
Bats Fig. 1-2 .................................................................................. CHIROPTERA
3'
Flying membrane between limbs only. Forepaws normal, clawed, although the front of
membrane may be supported on an extended wrist bone.
Gliders Fig. 2 ......................................................................................................... 4
4 (3)
Flying membrane enclosing limbs and tail to the tip. First two lower incisors widened
transversely with numerous elongate cusps, each tooth comb-shaped.
Colugo Fig. 2- right, Fig. 3-1 ....................................................... DERMOPTERA
4’
Flying membrane enclosing, at most, the base of well-developed tail. Canines absent,
chisel shaped incisors.
Flying squirrels Fig. 2- left, center ......................RODENTIA (In Part) 19 page
5 (2')
An elephant Fig. 1-3; with nose and upper lip prolonged into a long trunk. Ears large
and flattened against the side of head ............................................... PROBOSCIDEA
5’
Not an elephant ............................................................................................................ 6
6 (5’)
With hoofs (i.e. enlarged nad, surrounding the toe, and providing walking surface).
Hair sometimes sparse or absent. Hoofed animals .................................................... 7
6’
Writh claws or flat nails. Body thickly hairy............................................................... 8
7 (6)
A rhinoceros Fig. 1-4: large, naked. Feet with three toes and the center toes larger than
others ......................................................................................... PERISSODACTYLA
2
7’
With the two center toes equal and with hooves, giving the appearance of a single hoof
split down the middle ("cloven hoofed"). Usually hairy.
Pig, Mouse-deer, Deer, Cattle (Fig. 1-5) ................................ ARTIODACTYLA
8 (6’)
Both fore and hind-feet “hand” -shaped, with an opposable “Thumb” which usually
has a flat nail................................................................................................................ 9
8’
Feet never so modified for grasping .......................................................................... 10
9 (8)
Tree-mice: very small with teeth of rodent type (chisel shaped incisors and no canines)
Orbit open. Fig. 3-2 ..................................................RODENTIA (In Part) 19 page
9’
Apes, Monkeys, Tarsier, Slow Loris
Fig. 1 -6: canines present, orbit closed .................................................PRIMATES
10 (8’) Teeth of rodent type, i.e. incisors chisel-shaped (Fig. 3-2). And usually large, canines
absent, leaving a large gap (Diastema) between incisors and the grinding cheek teeth
into which the corners of lips are tucked, thus separating off the front of mouth.
Rats, Squirrels and Pomupines Fig. 4-2, 3 ....... RODENTIA (In Part) 19 page
10’
Teeth not so, canines present so that the toothrow is complete ..................................11
11 (10’) Middle-sized mammals, canines large, with incisors forining a transverse row of six
small teeth which are equal, or of which the outer ones are slightly larger (Fig. 3-3).
Muzzle not projecting beyond lower jaw.
Bear, Mustelids, Civets, Mongooses and Cats ...............................CARNIVORA
11
Small mammals, front teeth variable, the central incisors typically the largest. Muzzle
projecting far beyond the lower jaw .......................................................................... 12
12 (11’) Squirrels like: tail thickly hairy, at least towards the tip, and tail usually as long as or
longer than body. Eye encircled by bone. A complete set of teeth, all of which are of
small, many pointed.
Treeshrews Fig. 1-7, Fig. 4-1 ....................................... SCANDENTIA 15 page
12’
3
Rat-like: tail, which may be short, and is never longer than body, bearing only a sparse
covering of short hairs. Eye not encircled by bone. Canine or ca nine-form tooth (most
anterior tooth in maxilia) small and not clearly differentiated from cheek teeth.
Shrews and Moonrat Fig. 1-8 ................................... INSECTIVORA 12 page
Fig. 2 Characteristics features of mammals of Borneo
1. Pangolin: body covered with scales
2. Bat: forearm adapted for sustained flight
3. Elephant
4. Rhinoceros
5. Pig, Mouse-deer, Deer, Cattle
6. Monkey
7. Treeshrew: squirrel-like animal
8. Shrew: rat-like animal
4
Fig. 2 Silhouttes of three gliders
Left. Giant Flying Squiml: larger species, gliding membrane extending to the neck and to the
base of tail, tail round.
Center. Small Flying Squirrel: small species, gliding membrane from front to hind limbs only,
not including the base of tail, tail flat.
Right. Colugo: gliding membrane extends between front and hind legs, and encloses the tail.
Fig. 3 Different types of teeth
I1
I1
I2
I2
C
C
left
right
1 Colugo
Chisel-shaped incissors
Diasrema
(large gap)
Grinding cheek teeth
(premolars and molars)
2 Rodent
3 Carnivore
1. Colugo: comb-like incisors of lower jaw. 2. Rodent: incisors chisel-shaped, canines absent,
leaving a large gap (Diastbma). 3. Carnivore: incisors forming a transverse row of six teeth.
5
Fig. 4 Treeshrew, Squirrel and Rat
1 Treeshrew
2 Squirrel
3 Rat
6
Measurements
1. Total length (abbreviated as L):
The distance in a straight line, from the tip of the nose to the end of the last tail vertebra, exclusive
of the hairs. If rigor mortis (the stiffness, which sets in shortly after death,) has contracted the
muscles, the body must be stretched and the limbs pulled into a natural position.
Head and Body length (abbreviated as HB):
From the tip of the nose to the anus, not measured directly but calculated by measuring the total
length of body and tail, and subtracting the tail length.
Head and Body length = Total length - Tail length
2. Tail length (abbreviated as T):
The distance from the joint of the tail with the sacrum to the tip of the last vertebra, exclusive of
the hairs those project beyond the end. But it is also useful to record the length of hair separately.
3. Hindfoot length (abbreviated as HF):
The distance from the end of heel bone to the end of the longest toe, excluding the claw. This is
sometimes recorded as “length of hind foot, sine unguis”, to distinguish it from the American
measurement of to the end of claw on the longest toe, i.e. “length of hind foot, cum unguis”. The
length of the hindfoot is in many ways the most useful measurement of all. Because this length
is subject to less individual variation in specimens of the same species and age than any other
superficial measurement that can be made, and what is of practical importance, the measurement
is fairly uniform as recorded by different collector.
4. Ear length (abbreviated as E):
The distance from the base of the external opening to the tip, excluding the ear tuft, if present.
Bats should also have “height of tragus” (the flattened eminence of the auricular front of the
opening of the external ear). It is a good thing to make a pencil sketch of the outline of a bafs ear,
on the back of the label, as the ear is apt to shrivel up in drying if not carefully preserved.
7
5. Forearm length (abbreviated as FA): for bats only
The distance from the outside of the elbow to the outside of the wrist in the bent wing. This is
quoted in place of the Head and Body length for other mammals.
6. Weight (abbreviated as W):
Weight is usually recorded in grams, but sometimes in kilograms in the case of big animal.
7. Shoulder height:
The distance from bottom of the forefoot to the highest point of the shoulder in standing posture.
Usually measured in elephant, rhinoceros and deer.
8. Antler length:
The distance from the joint of antler and pedicel to the tip of the longest branch along the curve.
9. Dental formula (abbreviated as D):
For some species, the teeth are required to confirm the identity. The dental formula is a shorthand
way of indicating the number of teeth on one side of the upper and lower jaws. The teeth are
always given in the order of incisors (abbreviated as I), canines (abbreviated as C), premolars
(abbreviated as PM), and molars (abbreviated as M). For example, the dental formula for a
mammal, which has the maximum numbers of teeth, showed as 3143/3143=44. It is indicating 3
incisors, 1 canine, 4 premolars and 3 molars on each side of the upper jaw, and 3 incisors, 1
canine, 4 premolars and 3 molars on each side of the lower jaw for a total of 44 teeth. But many
mammals have less numbers than 44.
10. Mammillae (abbreviated as M):
Sometimes the number of mammillae may be of value in deciding the species. They are usually
arranged in pairs. The mammillae formula indicates the number of pairs of manimillae on the
chest, front-abdomen and rear-abdomen. For example, the mammillae formula for the Moonrat
Echinosorex Symnurms, is 1+1+1=6, indicating I pair on the chest, I pair on the front-abdomen
and I pair on the rear-abdomen giving a total of 6 mammillae. But sometimes it is expressed as
1+2=6, it indicating I pair on the chest and 2 pairs on the abdomen giving a total of 6 mammillae.
8
1
Total length
3
Total length
(T)
2
Ear length
(E)
4
Tragus
9
Forearm length
(FA)
Hindfoot length
(HF)
5
le
ng
th
A
nt
le
r
7
Shoulder height
8
Chest
incissors
canines
premolars
molars
9
Fore-abdomen
Rear-abdomen
Dental formula (D)
10
Mammillae (M)
Fig. 1 Measurements of External body
1. Total length (L), for calculating Head and Body length (HB).
2. Tail length (T).
Head and Body length (BB) is calculated by measuring the Total length (L), and subtracting
the Tail length (T). i.e. HB = L - T
3. Hindfoot length (M).
4. Ear length (E) and Tragns length.
5. Forearm length (FA).
6. Weight (W). Not illustrated.
7. Shoulder height.
8. Antler length.
9. Teeth = Dental formula (D). In this figure. D is 3 142/3 143=42.
10. Mammillae (M). In this figure, M is 1+1+2=8.
10
Fig. 2 Measurements of Skull
Greatest length (gl): the longest distance from the back of the skull to the front.
Condylobasal length (cbl): distance from the back of the occipital condyle to the front of the
premaxilla.
Inter-orbital breadth (iob): the least width across the top of the skull between the position of
orbits.
Maxillary toothrow (mt): the length of the upper toothrow from the back of the molars to the
front of the canine (mtC-M). For rodents that have no canines, this includes only the molars and
premolars (3 teeth in rats, 4 teeth in squirrels - the tiny premolar at the front is excluded). For
shrews, and usually for bats the whole toothrow is measured, including the incisors (mtl-M).
Molar width (m-m): the width across the outside offfic upper molars.
Canine width (c-c): the width across the outside of the base of the canines.
Zygomatic arch width (z-z): the width across the outside of the zygomatic arches.
Canine-occipital length (col): distance from the back of the skull to the front of the canine.
Applied for Rhinolophidae bats.
mtl-M
cbl
gl, cbl
mt
mt I-M
C-M
cbl
gl
gl
cbl
z-z
mtc-M
c-c
cbl
col
z-z
m-m
gl
iob
gl
11
mt
col
P A R T
I
I N S E C T I V O R A
S C A N D E N T I A
R O D E N T I A
12
Order INSECTIVORA
Moonrats and Shrews
Key to families
1
1’
Totally 44 teeth (I I teeth in each side of the upper jaw) ........................... Erinaceidse
8 or 9 teeth in each side of the upper jaw. 6 teeth in each side of the lower jaw.
Totally 28-30 teeth. first incisor large, protruding forward and hooked, small cusp present
behind main cusp ........................................................................................... Soricidae
Family Erinaceidae
Moonrat and Lesser Gymnure
Key to genera and species
1
A fairly large animal. HB 300 mm or more. HF over 50 mm,
fur nearly white ........................................................................................ Echinosorex
Echinosorex gymnurms
Moonrat
(HB 320-396, T 207-292, T/HB = 60010, HF 65-75, E 26.5, Wt 700-1,480g, D 3143/
3143=44, M 1+1+1=6. Skull: gl 75)
As big as cat. Has a distinct, pungent odor with strong ammonia content.
1’
Tail very short and naked, T/HB = about 10% ............................................... Hylomys
Hylomys suillus
Lesser Gymnure
(HB 116-150, T 10-24, T/HB= about 10-15%, HF 20-29, E 16, Wt 50-80g, D 3143/
3143=44. Skull: gl 37)
Upperparts brownish, underparts paler. Look like a large shrew. Momtains, above 1000m
(Common).
Family Soricidae
Shrews
Fig. 1 Difference of upper teeth
Crocidura (left): 2nd teeth twice as big as the 3rd.
Suncus (center): 2nd, 3rd, 4th and 5th teeth are unicuspid, the 5th teeth are much smaller than
the other teeth.
Chimarrogale (right): 2nd and 3rd teeth about equal.
13
Key to genera
1
9 teeth in each side of the upper jaw, of which four, the 2nd, 3rd, 4th and 5th are
unicuspid (only one cusp). The 5th teeth is much smaller dm the other teeth.
D 3123/1113 = 30 .............................................................................................. Suncus
1’
8 teeth in each side of the upper jaw, of which three are unicuspid.
D 3113/1113 = 28 ........................................................................................................ 2
2 (1’)
Feet with fringes of white stiff hairs. Ears very small, hidden among the fur of the head.
Tail thickly hairy. 2nd and 3rd teeth of upper jaw about equal.
Aquatic ................................................................................................... Chimarrogale
Chimarrogale himalayica
Himalayan Water Shrew
(HB 90-110, T 75-95, T/HB = 81-90%, HF 19-22, E 6-8.2, Wt 24-40g, D 3113/1113=28.
Skull: gl 25-27, cbl 23.0-26.0, mt 11.5-11.9) Lives in mountain streams (Very rare).
2’
Feet not fringed. Ears fairly conspicuous. Tail only thinly hairy. 2nd upper teeth twice
as big as the 3rd ............................................................................................ Crocidura
Genus Suncus
Key to species
1
1’
1’
Fairly large, HB over 90 mm, T/HB=50-70010. I-IF 14-24 mm. Usually in or near
houses (Common)............................................................................... Suncus murinus
House Shrew
(HB 92-146, T 46-86, T/HB = 50-70%, HF 14-24, Wt 60g, D 3123/1113=30. Skull: gl
32, cbl 33.1-33.4, iob 5.9-6.0, mt 13.7-14.4)
Entire body and tail uniform mid-gray to brownish gray. Utters from time to time a
shrill whistling chirp, almost like the clinking of two coins together.
HB 75 mm, T 57 mm, T/HB = about 76%, HF 12 mm, D 3123/1113=30.
Mountains. Known only from G. Kinabalu (1700m point) ........................ Suncus ater
Black Shrew
(Skull: cbl 21.3, greatest width of braincase 9.1, mtI-M 9.2)
Very small, HB 50-56 mm, T 24-32 mm, T/HB = 48-56%, HF 10 mm or less. D 3123/
1113=30 ............................................................................................... suncus etruscus
Savi’s Pigmy Shrew
(Skull: cbl 14.5, iob 4.1, mt 6.4)
14
Genus Crocidura
Key to species
15
1
Very small, HB less than 65 mm, T 37 mm. Tail, with sparse, long, pale hairs on basal
10 mm .......................................................................................... Crocidura monticola
Sunda Shrew
(Skull: cbl 15.2-17.4, mt 6.6-7.3)
1'
HB 52-112 mm, T 54.1-97 mm, T/HB=54-90%, HF 12.5-18.5 mm, Tail brownish, with
few or no long hairs ..................................................................... Crocidura fuliginosa
South-east Asian White-toothed Shrew
(HB 52-112, T 54.1-97, T/HB = 54-90%, HF 12.5-18.5, E 8.7-10.5, Wt 10.8-13.3 g,
D 3113/1113=28. Skull: cbl 21.3-25.0, mt 9.4-11.5)
Upperparts dark brownish gray. Underparts slightly paler. Tail brownish, with few or
no long hairs.
Order SCANDENTIA
Treeshrews
Key to genera
1
Tail naked at the base, but with a horizontally compressed fringe of long hairs near the
tip, giving the tail appearance of a feather. Nocturnal ................................. Ptilocercus
Ptilocercus lowii
Pentail Treeshrew
(HB 115-150, T 155-210, T/HB = 135-150%, HF 24-30, E 19-21, Wt 40g. D 2133/
3133=38. Skull: gl 33-36)
Quite unlike any other treeshrews, and resembling a true shrew. Upperparts is graybrown, and the underparts yellow-gray. Eyes and ears tend to protrude more than do
those of a treeshrew. The tail is long and much of its length naked and rat like. However,
half or one third of its length near the tip is clothed in long hair which remains in a
horizontal position, giving the tail the appearance of a large quill feather.
1’
Tail hairy to the base usually bushy. Diurnal .............................................................. 2
2 (1’)
Small species. Tall long appears thin and rat like. Mountains ................... Dendrogale
Dendrogale melanura
Smooth-tailed Treeshrew
(HB 103-137, T 135-149, T/HB = 109-122%, HF 27-30, E 13-18, D 2133/3133=38.
Skull: gl 30-31)
2’
Tail hairs long, at least 10 mm, so that the tail is bushy ..................................... Tupaia
Genus Tupaia
Key to species
L
With black stripe down the middle of the back.
1’
With no such black stripe, although the fur of dorsal crest of hair may be somewhat
darker than that of the sides ......................................................................................... 3
2 (1)
Black stripe down the back continues to the root of the tail.
Rather smaller species .......................................................................... Tupaia dorsalis
Striped Treeshrew
(HB 160-220, T 140-162, T/HB = 68-95%, HF 39-46, E 12-14, D 2133/3133=38. Skull:
gl 45-47, cbl 43.6-48.0, iob 11.9-14.3, mtC-M 16.8-18.2 ). Rare.
2’
Black stripe continues for only about half the length of the back ............................... 4
16
17
3 (1’)
Larger species. Tail not noticeable longer than UB, usually much shorter. Skull length
over 45 mm .................................................................................................................. 5
3')
Smaller, and slender species. Tail always longer than HB (T/HB = usually more than
110%). Skull length under 40 mm. HF under about 40 mm ....................................... 6
4 (2’)
Very large species. Upperparts paler towards the front of the body, and blacker towards
the rump. HF 40-57. Center of eye to the tip of muzzle more than 37 mm in adults.
................................................................................................................... Tupaia tana
Large Treeshrew
(HB 165-321, T 130-220, T/HB = 75-96.2%, HF 40-57, E 9.8-19, Wt 154-305 g, D 2133/
3133=38, M 0+2+0=4. Skull: gl 53-60, cbl 50.8-57.5, iob 13.3-14.9, mtC-M 18.5-21.0)
The largest treeshrew, slightly larger than Tupaia glis. The dorsal fur is reddish brown
shading to ahnost black to the rear. There is a black stripe running from the back of
neck about half or two thirds of the way down the back where it disappears in dark
dorsal color. The under part reddish-buff. The center of the eye to the tip of muzzle is
more than 37 mm in adults. Common.
4’
Upperparts generally brown. Underside and distal end of tail with a strong orange or
reddish tinge .............................................................................................. Tupaia picta
Painted Treeshrew
(HB 159-240, T 137.2-175, T/HB = 64-103.8%, HF 39-46, E 12.5-17, Wt 100-195g, D
2133/3133=38, M 0+2+0=4. Skull: cbl 43.6-48.5, iob 13.7-14.5, mtC-M 17.4-19.0)
Upperparts generally brown, with a black central stripe on the front half of the body
Underparts dull orange. Underside and distal end of the tail with a strong orange or
reddish. No record in Sabah.
5 (3)
Very large. Shoulder stripe and underparts bright buff or reddish brown in contrast to
dark brown fur of back. HF 45-56 mm. Lowlands and hills ....................... Tupaia glis
Common Treeshrew
(HB 170-237, T 160-203, T/HB = 93.2-112.2%, HF 45-56, E 12.1-16.4, Wt 148-235g, D
2133/3133=38, M 0+2+0=4. Skull: gl 48-52, cbl 48.0-51.0, mtC-M 19.1-21.0) The hair
on the upper part is banded dark and pale, appearing finely speckled reddish brown and
the tail is of uniform coloring. Usually has a pale stripe on each shoulder. The under part
is dull buff to reddish buff and the underside of the tail is generally grayish. Common.
5’
A plain reddish treeshrew. Tail hairs pure dark red.
HF 37.3-44 mm .............................................................................. Tupaia splendidula
Ruddy Treeshrew
(HB 164-200, T 130-164, T/HB = 72-95.1%, HF 37.3-44, E 13.7-15.1, Wt 123.5-190g,
D 2133/3133=38, M 0+2+0=4. Skull: cbl 42.2-48.8, iob 13.2-13.8, mtC-M 17.0-20.2)
A plain reddish treeshrew, with pale orange shoulder stripe. The upper part tends to be
darker in the midline and brighter on sides, underparts dark reddish with an orange
throat. The hair on the tail pure dark red above, orange below. No record in Sabah.
5’’
Smaller. Shoulder stripe and underparts not very much paler than dull brown dorsum,
so that usually appears entirely dark brown when glimpsed in the field. Mountains.
Common ............................................................................................. Tupaia montana
Mountain Treeshrew
(HB 156-227, T 135-190, T/HB = 62.5-100 %, less than 100%, BF 37-45, E 10-18,
W 95-219g, D 2133/3133=38, M 0+2+0=4. Skull: gl 42-47, cbl 43.1-46.1, iob 12.715.3, mtC-M 16.2-18.0)
6 (3’)
HB 130-165 mm, T 162-190 mm, T/HB 110-150%, HF 36-43 mm,
Wt 60-98 g .............................................................................................Tupaia gracilis
Slender Treeshrew
(HB 130-165, T 162-190, T/HB = 110-150%, HF 36-43, E 10.7-13.3, Wt 60-98g. D 2133/
3133=38. M 1+1+0=4. Skull: gl 34-38, cbl 34.6-36.6, iob 11.6-12.7, mtC-M 13.3-14.3)
Upperparts speckled olive-brown, underparts buff white with no reddish tinge.
6’
HB 11 0- 1 40 mm, T 131-172 mm, T/HB = 115-130%, FIF 28-34 mm,
Wt 30-71 g ............................................................................................... Tupaia minor
Lesser Treeshrew
(HB 110-140, T 131-172, T/HB = 115-130%, HF 28-34, E 10-14, Wt 30-75g. D 2133/
3133=38. Skull: gl 33-37, cbl 31.0-34.9, iob 10.5-12.1, mtC-M 11.4-12.7)
The hair on the upper part is banded light and dark giving an overall speckled olivebrown
appearance. The under part is buff, often with a reddish tinge towards the rear. The
dorsal and tail are of the same brownish color, but the tail is darker than the body.
Fig. 1 Size and color patterns of six Treeshrews.
Tupaia minor, Tupaia gracilis: small and long, thin tail, the dorsum and tail are of the same
olive brown. Hindfoot length 28-34 mm (Tupaia minor), 36-43 mm (Tupaia gracilis).
Tupaia glis: upperparts brown or reddish-brown, an orange stripe on each shoulder.
Tupaia tana: the largest, upperparts is reddish brown shading to almost black to the rear, a black
midline runs towards the rump but it disappears in the dark dorsal color, an oblique buff stripe
on each shoulder
Tupaia picta: middle size, upperparts reddish brown, with shoulder stripe pale buff, tail bright
red,black central stripe on the front half of the body.
18
Tupaia splendidula: middle size, upperparts pure dark red, with shoulder stripe pale orange, tail
bright red. Very similar to Tupaia picta but lack dorsal stripe.
pale buff
Tupaia minor
Tupaia gracilis
olive brown
pale orange
reddish brown
Tupaia glis
dark red
black
Tupaia tana
reddish brown
pale buff
bright red
black
Tupaia picta
reddish brown
pale orange
bright red
Tupaia splendidula
dark red
19
Order RODENTIA
Squirrels, Rats and Porcupines
Key to families
1
Hair of upperparts replaced by hard spines or quills. and with a tuft of hairs or spines at
the tip of the tail.
.................................................................................................................... Hystricidae
1’
Back covered with soft hair, or if spiny (as in some rats) the spines are mingled with
soft hair.
..................................................................................................................................... 2
2 (1’)
Squirrels, with a bushy tail covered with long hairs from the base.
........................................................................................................................ Sciuridae
2’
Rats, tail naked and scaly, sometimes with sparse hairs towards the tip.
......................................................................................................................... Muridae
Family Sciuridae
Squirrels
Key to subfamilies
1
1’
Gliding membrane present between the limbs, but the tail is not enclosed by a membrane.
Nocturnal.
................................................................................................................. Petauristinae
No gliding membrane.
....................................................................................................................... Sciurinae
Subfamily Sciurinae
Tree squirrels and Ground squirrels
Key to genera
1
1’
2 (1’)
2’
Upperparts with three or four black stripes.
......................................................................................................................... Lariscas
One black stripe on each flank, or no stripes
..................................................................................................................................... 2
White or buff stripe in each flank usually underlined with a black stripe.
..................................................................................................................................... 3
No stripes at all.
..................................................................................................................................... 5
20
3 (2)
Very small, Head & Body length less than 150 mm. Hindfoot length less than 33 mm.
Underparts reddish so that the animal looks similar to a miniature Callosciurus adamsi.
....................................................................................................................... Glyphotes
Glypholes simus
Red-bellied Sculptor Squirrel
(HB 94-144, T 95-106, T/HB=105%, HF 27-33, W 50g. Skull: gl 25-30)
3’
Larger than above.
..................................................................................................................................... 4
Very large, Head & Body are 300 mm. Hindfoot length over 80 mm. Upperparts very
dark brown, with a blackish and pale stripe on each side of the body. Tufts of hair on the
ears. Tail large.
............................................................................................................. Rheithrosciurus
Rheithrosciurus macrotis
Tufted Ground Squirrel
(HB 335-352, T299-342, T/HB=90-100%, HF 81-88, Wt 1, 170-1, 280g. Skull: gl 70)
4 (3’)
4’
5 (2’)
5’
6 (5’)
6’
7 (6)
21
Medium sized, Head & Body length less than 270 mm, Hindfoot length less than
65 mm. Ears without tufts.
.................................................................................................................. Callosciurus
Very large, Head & Body length 225-380 mm, Hindfoot length over 60 mm. Tail 110150% of Head & Body length. Upperparts usually dark in the middle, underparts pale.
............................................................................................................................ Ratufa
Ratufa affinis
Giant Squirrel
(HB 225-380, T 317-444, T/HB=110-150%, HF 50-87, E 23.5, Wt 875-1,500 g, D1013/
1013=20, M 2+3+0=10. Skull: gl 63-68) The largest tree squirrel.
No so as above
..................................................................................................................................... 6
Very small, Head & Body length usually less than 90 mm, Hindfoot length less
than 25 mm.
..................................................................................................................................... 7
Larger species.
..................................................................................................................................... 8
Buff or creamy white stripe from nose across cheek to behind the ear.
................................................................................................................ Nannosciurus
Nannosciurus metanotis
Black-eared Pigmy Squirrel
(HB 61-87, T 60-71, T/HB=85-100%, HF 19-209 W 20g. Skull: gl 20-40)
Upperparts grayish brown, underparts orange brown. Tail black and gray, head with
black spot behind each ear. Distinguished from other pigmy squirrels by broad buff line
across face with black above. No record in Sabah.
7’
8 (6’)
8’
9 (8)
9’
Without such stripe across cheek.
................................................................................................................... Exilisciurus
Muzzle long and rather shrew-like. Length of nasal bones greater than the width of
skull between the eyes. Incisors reduced. Tail short, bushy, nearly always less than 75%
of Head & Body length.
..................................................................................................................................... 9
Muzzle short, typically squirrel-like. Length of nasal bones shorter than width of skull
between eyes. Incisors well developed
................................................................................................................................... 10
Smaller animals, Head & Body length usually less than 10 mm. Upperparts gray or
grayish buff. Muzzle slightly pointed. Mountains.
...................................................................................................................... Dremomys
Dremomys everetti
Bornean Mountain Ground Squirrel
(HB 160-190, T 92-130, T/HB=usually 55-60%, HF 30-42, E 15-18, Wt 145-185 g,
D1023/1013=22, M 0+1+2=6. Skull: gl 45)
Larger species, Head & Body length usually over 195 mm. Upperparts cream or orange.
Narrow pointed head like a treeshrew.
................................................................................................................. Rhinosciurus
Rhinosciurus laticaudatus
Shrew-faced Ground Squirrel
(HB 195-233, T 131-170, T/HB=45-70%, HF 40-46, W 250g. Skull: gl 58-60)
10 (8’) Large squirrels, Head & Body length usually over 200 mm, Hindfoot length usually
over 45 mm.
....................................................................................................................................11
10’
Smaller squirrels, Head & Body usually less than 160 mm, Hindfoot length less
than 40 mm.
................................................................................................................. Sundasciurus
11 (10) Upperparts to black, underparts chestnut red.
..................................................................................................................Callosciurms
(Callosciurus prevostii)
black phase only, Occurs in Sabah and the northern-most of Sarawak.
11’
Upperparts brown. Tail gray or black. Shoulders and head usually gray. Underparts
white or chestnut. tail bushy.
................................................................................................................ Sundasciurms
(Sundasciurus hippurus only)
22
Genus Callosciurus
Key to species
1
1’
2 (1)
2’
3 (1’)
3’
4 (3)
4’
23
Large squirrels, Head & Body length over 200 mm, Hindfoot length over 45 mm.
..................................................................................................................................... 2
Medium sized squirrels. White or buff stripe on each flank, underlined with a black
stripe.
..................................................................................................................................... 3
Upperparts gray-black with bright buffy-red speckling especially on the face and legs.
Short white and black side-stripes. Underparts reddish. Mountains.
.................................................................................................. Callosciurus baluensis
Kinabalu Squirrel
(HB 210-255, T 215-260, HF 48-55)
Upperparts black, underparts chestnut red (Black phase). Or upperparts black, grizzled
or grizzled olive. White or, gray stripe, or white and black stripes on each side, but all
have reddish underparts.
................................................................................................... Callosciurus prevostii
Prevost’s Squirrel
(HB 200-270, T 202-273, T/HB = 90-110%, HF 45-65, E 17.1-20, Wt 250-500 g.
Skull: gl 53-57)
Pale spot behind ear.
..................................................................................................................................... 4
No pale spot, underparts reddish.
..................................................................................................... Callosciurus notatus
Plantain Squirrel
(HB 160-238, T 160-21 0, T/HB = 70-110%, HF 42-52, E 16.3-18, Wt 150-280 g,
D 1023/1013=22. Skull: gl 50.0-51.0, cbl 45.7, iob 18.6, mt 18.6)
Upperparts and tail brown. Under part pale red or orange, never gray. Side-stripes buff
and black.
Underparts gray.
...................................................................................................... Callosciurus orestes
Bornean Black-banded Squirrel
(HB 132-170, T 128-170, HF 32-39)
Underparts reddish.
...................................................................................................... Callosciurus adamsi
Ear-spot Squirrel
(HB 166-183, T 152-167, HF 36-40, Wt 115-154 g)
Genus Sundasciurus
Key to species
1
1’
2 (1’)
2’
3 (2’)
3’
4 (3’)
4’
Very large, Head & Body length over 210 mm, Hindfoot over 50 mm. Upperparts
brown to dark-brown. Tail gray to dark gray. Tail bushy. Head and shoulders gray to
dark gray. Underparts white or chestnut red.
................................................................................................. Sundasciurus hippurus
Horse-tailed Squirrel
(HB 213-250, T 176-257, T/HB = 110-120 %, HF 52-61, Wt 260-365 g. Skull: gl 54-56)
Upperparts reddish brown with gray head and shoulders. Tail entirely dark. Underparts
dark reddish.
Small squirrels. Head & Body length less than 160 mm, hindfoot length less than 40 mm
..................................................................................................................................... 2
Underparts gray. Reddish patch between hind legs (not visible from side). Medium
sized, Hindfoot length over 32 mm. Mountains.
................................................................................................... Sundasciurus brookei
Brook’s Squirrel
(HB 140-205, T 108-170, T/HB=70-106 %, HF 32-38, Wt 103-128g. Skull: gl 38.040.2, cbl 34.4, iob 14.2, mt 6.9)
Underparts cream, buff, or grayish-buff. Smaller, Hindfoot length less than 40 mm.
..................................................................................................................................... 3
Tail very short and bushy. Underparts cream with the color sharply defined on the flanks.
........................................................................................................ Sundasciurus lowii
Low’s Squirrel
(HB 132-157, T 77-106, T/HB = about 60-70 %, HF 27-35, E 12.4-14, Wt 60-120g,
D 1023/1013=22. Skull: gl 35.0-40.0, cbl 33.5, iob 11.7, mt 6.4)
Upper part speckled brown, under part cream with the color sharply defined on the
flanks. Tail very short and bushy.
Tail longer and thin, usually about 70 % of Head & Body length
..................................................................................................................................... 4
Creamy-white eye-ring, ear margins and mustache-like stripe. Mountains.
................................................................................................... Sundasciurus jentinki
Jentink’s Squirrel
(HB 120-144, T 110-133, T/HB = 96-100 %, HF 30-34)
Ears and eyes without such patterns.
...................................................................................................... Sundasciurus tenuis
Slender Squirrel
(HB 135-155, T 125-132, T/HB = 85-95 %, HF 32-359 W 70g. Skull: gl 35)
24
Genus Lariscus
Key to species
1
1’
Three dark stripes along midline of upperparts. One in the center separated by bands of
the body color.
........................................................................................................... Lariscus insignis
Three-striped Ground Squirrel
(HB 170-230, T 117-138, T/HB = 59.1-63.6%, HF 33-46, E 17, W 216-245 g, D 1023/
1013=22. Skull: gl 51)
Four black stripes, the two in the center separated by a central buff line, and the inner
and outer stripes on each side separated by a very pale line.
................................................................................................................ Lariscas hosei
Four-striped Ground Squirrel
(HB 172-192, T 110-142, T/HB = 60-72.5%, HF 42-46, E 13.6, Wt 145-215 g. Skull: gl
45.0-47.7, cbl 42.9, iob 12.5, mt 8.0)
Genus Exilisciurus
Key to species
1
1
Ears with long gray tufts of hair. Mountains.
................................................................................................ Exilisciurus whiteheadi
Whitehead’s Pigmy Squirrel
(HB 83-93, T 65-73, T/HB = 80-90%, HF 22-24, W 20g. Skull: gl 25)
Without such a tufts of hair. Very small.
......................................................................................................... Exilisciurus exiles
Plain Pigmy Squirrel
(HB 62-82, T 42-62, T/HB = 50-90%, HF 18-23, Wt 12-16 g. Skull: gl 21-23)
Upperparts speckled olive-brown. Underparts pinkish-buff. Call is a single high-pitched
squeak.
Subfamily Petauristinae
Flying squirrels
Key to genera
1
25
Large species. Gliding membrane extending to the neck and to the base of the tail. Tail
round
..................................................................................................................................... 2
1’
2 (1)
Smaller species. Gliding membrane from front to hind limbs only, not including the
base of the tail. Tail flat that is with hairs all in one plane like a feather.
..................................................................................................................................... 4
Upperparts almost uni-color, gray-black or reddish. No distinct mark on the upperparts.
......................................................................................................................... Aeromys
2’
Upperparts gray-brown to reddish, but different color or speckles on somewhere of
dorsum.
..................................................................................................................................... 3
3 (2’)
Smaller forms, Head & Body length less than 290 mm. Upperparts very dark-brown
with fine pale grayish speckling. Underparts creamy with some gray. Tail rusty colored.
.................................................................................................................. Pteromyscus
Pteromyscus pulvermientus
Smoky Flying Squirrel
(HB 221-290, T 215-235, HF 41-44, E 20, Wt 232-305 g. Skull: gl 46)
3’
Larger forms, Head & Body length over 330 mm. Upperparts reddish.
...................................................................................................................... Petaurista
4 (1’)
Upperparts brown to dark gray. Underparts orange-buff or whitish, without gray. Gliding
membrane fringed with rusty brown. Tail dark brown above, reddish below. Ears rather
large, 22-25 mm, long and almost naked.
............................................................................................................................. Iomys
Iomys horsfieldi
Horsfield’s Flying Squirrel
(HB 165-231, T 160-207, HF 33-44, E 22-25, W 135-215. Skull: gl 41-43)
4’
Tail uniformly colored. Ears smaller less than 20 mm, and hairy.
..................................................................................................................................... 5
Bullae of skull flattened, so that their underside is roughly level with the rest of the base
of the skull.
...................................................................................................................... Petinomys
Bullae normal, projecting below the general level of the base of the skull.
..................................................................................................................................... 6
Very small species, Head & Body length less than 90 mm. Cheek teeth with cusps and
ridges poorly marked. Last premolar smaller than first molar.
..................................................................................................................... Petaurillus
Larger species, Head & Body 110 mm. Cheek teeth with well-mairked cusps and ridges.
Last premolar not smaller than the first molar.
....................................................................................................................... Hylopetes
5 (4’)
5’
6 (5’)
6’
26
Genus Petaurillus
Key to species
1
Upperparts very dark gray with pale buff streaks, especially in the midline. Underparts
off-white on gray under-fur. Tail buffy at base, becoming blacker near the end, with a
white tip. Cheeks buffy white with a strong gray tinge beneath the eye.
........................................................................................................... Petaurillus hosei
Hose’s Pigmy Flying Squirrel
(HB 80-87, T 80-98, HF 19-20, M 0+2 = 4)
1’
Similar to above, but smaller. Cheeks entirely buffy white without gray below the eyes.
........................................................................................................ Petaurillus emiliae
Lesser Pigmy Flying Squirrel
(HB 68,72, T 67,62, HF 16,17)
Genus Aeromys
Key to species
1
Upperparts, tail and cheeks dark gray-brown, almost black. Underparts slightly paler.
Fluffy hair.
.................................................................................................... Aeromys tephromelas
Black Flying Squirrel
(HB 355-426, T 410-470, HF 67-78, E 35, Wt about 900 g. Skull: gl 67)
1’
Entirely reddish-brown.
........................................................................................................... Aeromys thomasi
Thomas’s Flying Squirrel
(HB 350-403, T 340-430, HF 60-70, E 30, Wt 1,380-1,490 g. Skull: gl 55-66)
Entirely reddish-brown.
Genus Petinomys
Key to species
1
27
Whitish hairs at the base of each ear. Distinct tuft of long whiskers on the cheek behind
the eye. Gliding membrane with a white margin. Upperparts reddish-brown with gray
under-fur, speckled with gray anteriorly, and with reddish posteriorly, giving an unusual
1’
2 (1’)
2’
3 (2’)
2
pinkish tinge. Underparts cream or dull orange-buff.
.................................................................................................... Petinomys genibarbis
Whiskered Flying Squirrel
(HB 160-180, T 155-188, HF 31-32)
No prominent tuft of whiskers behind the eye. Rump not pinkish.
..................................................................................................................................... 2
Upperparts dull reddish-brown, with buffy tips to hairs. Underparts off-white. A broad
band or red-brown extends between and around eyes to ears. Long, stiff hairs behind
and in front of ears.
.......................................................................................................... Petinomys hageni
Hagen’s Flying Squirrel
(HB about 280, T about 230, HF about 45)
Not so as above.
..................................................................................................................................... 3
Cheeks orange. Gliding membrane with a buff (not white) margin. Upperparts blackish
with rusty colored tips. Underparts buffy white.
................................................................................................ Petinomys vordermanni
Vordermann’s Flying Squirrel
(HB 92-105, T 98-103, HF 21. Skull: cbl 28.2-30.0, iob 5.9, mt 5.5-6.2)
Cheeks grayish. Margin of gliding membrane not distinctly pale. Upperparts dark gray
or black with pale buff tips. Underparts white with gray under-fur.
.......................................................................................................... Petinomys selosas
Temminck’s Flying Squirrel
(HB 105-132, T 93-145, T/HB = 109.8%, HF 21-31.5, E 19, Wt 54 g. Skull: gi 27-30,
cbl 27.6-29.6, iob 5.0-6.7, mt 5.0-5.2)
Genus Hylopetes
Key to species
1
Upperparts blackish or dark gray-brown with rust-colored markings. Underparts white
or buffy white on gray under-fur. Tail dark brownish-gray, somewhat broader at the
base than at the tip. Cheeks and patch on each side of the base of the tail pale gray.
.......................................................................................................... Hylopetes lepidus
Grey-cheeked Flying Squirrel
(HB 117-135, T 118-120, HF 29-30, M 0+1+2=6. Skull: cbl 29.8, iob 8.0-8.1,
mt 7.1- 7.2)
28
1’
Similar to above. Tail dark, slightly orange brown with buffy under-fur, distinctly orange
at base. Cheeks orange brown on gray.
...................................................................................................... Hylopetes spadiceus
Red-cheeked Flying Squirrel
(HB 157-184, T 152-166, T/HB = 85.7%, HF 29-35, Wt 80-157 g, M 0+1+2=6. Skull:
gl 35-39, cbl 31.4-36.4, iob 7.8-8.7, mt 6.8-7.6)
Genus Petaurista
Key to species
1
Entire body dark reddish except for black on the nose, chin, eye-ring, behind the ears,
feet and tail tip.
..................................................................................................... Pelaurista petaurista
Red Giant Flying Squirrel
(HB 370-450, T 365-490, HF 65-80, E 40, Wt 1,000-2,900 g. Skull: gi 65-70)
The largest flying squirrel. Entire body reddish brown with black markings on nose and
chin, around the eyes, behind the ears, at the tips of the feet and tail.
1’
Upperparts black in the middle with many white speckles. Reddish laterally and on the
gliding membrane. Underparts reddish. Tail black.
......................................................................................................... Petaurista elegans
Spotted Giant Flying Squirrel
(HB 338-365 mm, T 340-365 mm, HF 60-66 mm, E 30, Wt 840-1,240 g. Skull:
gl 65- 70)
Family Muridae
Rats and Mice
Key to genera
1
1’
2 (1)
29
Large species, Head & Body length more than 100 mm, Hindfoot length more than
21 mm.
..................................................................................................................................... 2
Small species, Head & Body length less than 80 mm, Hindfoot length less than 20 mm
(possible to distinguish from juvenile of Rattus species by hindfoot length).
..................................................................................................................................... 7
Tail uni-color.
..................................................................................................................................... 3
2’
Tail bicolored.
..................................................................................................................................... 6
3 (2)
Tail uniformly dark, at least 125 % of Head & Body length, slightly tufted with hair
at the tip.
....................................................................................................... Niviventer Fig. 1-3
3’
Tail brown, hairy and with a brush of hair at the tip. Hallux (first digit of hindfoot) is
semi-opposable, has a nail (not a claw).
............................................................................................................... Chiropodomys
(Chiropodomys major only)
3’’
Tail not tufted.
..................................................................................................................................... 4
4 (3’)
Tail very long, at least 134% of Head & Body length. Upperparts reddish brown
.................................................................................................. Leopoldamys Fig. 1-5
Leopoldamys sabanus
Long-tailed Giant Rat
(HB 204-273, T 327-402, T/HB = 134.2-174 %, HF 40-52, E 24.3-30.4, Wt 216-532g,
D 1003/1003=16, M 2+0+2=8. Skull: gl 50-58, cbl 46.0-52.3, iob 8-8.5, mt 9.1-10.0)
Upperparts bright brown with gray under-fur and numerous soft hair-like spines. Under
part creamy-white. Tail, which is very long, sometimes with pale, irregular blotches.
4’
Tail less than 125% of Head & Body length. Upperparts olive-brown or dark grayish.
..................................................................................................................................... 5
5 (4)
Spines numerous on upperparts, scare on underparts but generally soft, stiff hair.
............................................................................................................. Rattus Fig. 1-1
5’
Large. Prominent long black guard hair on upperparts.
.......................................................................................................Sundamys Fig. 1-2
5’’
Fur long, dense and soft, body fur extending well onto base of tail.
................................................................................................. Pithecheirops Fig. 1-7
Pithecheirops Orion
Danum Tree Rat
Tail robust, slickly naked, and at least partially prehensile; hind feet broad, with large
pads, hallux separated by a gap from other digits, with a claw forefeet also broad, with
large palmar pads, pollux with a broad shiny nail. Strongly arboreal-adapted rat. Only
a juvenile male was collected at Danum Valley.
30
6 (2)
Tail entirely white at the end distal from the body.
..................................................................................................... Lenothrix Fig. 1-6
Lenothrix canus
Grey Tree Rat
(HB 165-220, T 200-265, T/HB = 90-130%, HF 30-37, Wt 80-220 g, M 3+1=8. Skull:
cbl 42.0-44.5, job 6.1-6.6, mt 8.2-8.6)
Dense, woolly hair without spines. Tail entirely white at the end distal from the body.
Confined to forested areas, and are arboreal.
6’
Tail, dark brown or black above, and white beneath, separated by a sharp line.
........................................................................................................ Maxomys Fig. 1-4
Hallux is semi-opposable.
..................................................................................................................................... 8
Hallux not so as above. An incision at the rear ridge of each upper incisor. First molar
always longer than half the toothrow( MI > M2+M3), Mammillae 3+2=10 (cf. Rattus:
M 2+2=8, M 2+0+3=10 or M 3+0+3=12), urine bad smell and it is dyed in body.
...........................................................................................................Mus Fig. 1-8, 8a
Hallux with a nail. Tail hairy and with a brush of hair at the tip.
................................................................................................ Chiropodomys Fig. 1-9
Hallux with a tiny claw. tail very long with tiny stiff hair.
..................................................................................................... Haeromys Fig. 1-10
Haeromys margarettae
Ranee Mouse
(HB 76-77, T 123-144, HF 17-20, M 0+3=6. Skull: cbl 20.7-23.9, iob 4.3-4.5,
mt 3.3-3.7)
Hallux is semi-opposable with a tiny claw. Tail very long, dark brown, with tiny
stiff hair.
7 (1)
7’
8 (7)
8’
Fig. 1 Size and distinctive mark of nine genera of Muridae
1 Ratims: tail entirely dark, spines numerous but generally soft.
2 Sundamys: large, tail entirely dark, prominent long black guard hairs.
3 Niviventer: numerous stiff spines, tail uniformly dark, and slightly tufted. 4 Maxomys: tail
short and bicolored.
5 Leopoidamys: large, very long tail, spines numerous but soft.
6 Lenothrix: woolly hair without spines, tail entirely white at the end distal from the body.
7 Pithecheirops: woolly hair without spines, tail uni-color, robust, slickly naked, and at least
partially prehensile.
8 Mus: small, incision on each upper incisor (7a).
9 Chiropodomys: hallux is semi-opposable (9a) with a nail, tail hairy and with a brush of hairs
at the tip.
10 Haeromys: hallux is semi-opposable (9a), tail with tiny stiff hairs.
31
1
Rattus
2
Sundamys
entirely dark
long guard hairs
Nivivender
3
tufted
Maxomys
4
upperside: brown or black
underside: white
5
Leopoldamys
6
Lenothrix
very long tail,
unicolor
white
32
7
Pithecheirops
woolly hair without spines,
base of the tail covered with bod fur,
both hand and foot wide and have big pads
8
hairy
Mus
incision of upper incisor
a nail (not a claw on the hallux)
9
Chiropodomys
10
Haeromys
brush of hairs
a tiny claw on the hallux
0
2
4
6
8
10 cm
Genus Rattus
Tail entirely dark. Spines numerous on upperparts, scarce on underparts but generally soft, like
stiff hair. Hindfoot over 20 mm. First molar always less than half the toothrow (M1 < M2+M3).
Key to species
1
1’
33
Hair soft, 30-35 mm long. Upperparts dark grayish brown with pale reddish brown
speckling. Underparts gray-buff. Mountains
............................................................................................................ Rattus baluensis
Summit Rat
(HB 150-188, T 145-206, T/HB = 128%, HF 21-38, E 19, Wt 85-135 g) Recorded from
Gunung Kinabalu (2150-3350m) and Trus Madi.
Shorter, coarser hair. Occur at lower altitudes.
..................................................................................................................................... 2
2 (1)
2’
3 (2’)
3’
4 (3’)
4 (3’)
Smaller species, Hindfoot length less than 28 mm, Weight less than 65 g. Upperparts
grayish to reddish brown. Underparts grayish-white.
............................................................................................................... Rattus exulans
Polynesian Rat
(HB 101-138, T 118-159, T/HB = 104.6-131.9%, HF 22-28, E 15-17.5, Wt 42-65 g, D
1003/1003=16, M 2+0+2=8. Skull: gl 30-34, cbl 26.8-29.2, iob 4.6-4.9, Mt 4.7-5.6)
Upper part grayish-brown, under part grayish-white. Spines white with dark brown
tips. Tail uniformly black.
Larger species, Hindfoot length more than 28 mm. Weight more than 75 g.
..................................................................................................................................... 3
Tail shorter, Less than half of Total length. Upperparts mid-brown, underparts gray
brown. Town only.
.......................................................................................................... Rattus norvegicus
Norway Rat
(HB 163-265, T 170-230, T/HB = 79-97%, HF 34-47, Wt 150-400 g, D 1003/1003=16,
M 3+3=12. Skull: gl 50, cbl 41.4-50.9, iob 6.3-7.4, Mt 7.0-7.4)
Upperparts brown, underparts gray-brown. Tail being slightly darker above and
paler below.
Tail relatively longer. Upperparts grizzled olive-brown, or pale brown with fine black
speckling.
..................................................................................................................................... 4
Underparts silver-gray, often with a dark streak along the middle.
...................................................................................................... Rattus argentiventer
Ricefield Rat
(HB 110-220, T 130-192, T/HB = 80-125%, HF 28-38, E 15.8 (young), Wt 85-190 g, D
1003/1003=16, M 3+3=12. Skull: gl 41, cbl 35.1-39.7, iob 5.6-6.6, Mt 7.0-7.1)
Upperparts pale brown with fine black speckling, very similar to House Rat, Rattus rattus,
but more speckled appearance, and underparts wholly silvery-gray, usually with a dark
streak across the middle chest. Young has an orange-colored tuft in front of each ear.
Underparts white or pale gray with buffy-white tips. Distinct pattern of fine ridges on
the pads of the feet.
......................................................................................................... Rattus tiomanicus
Malaysian Field Rat
(HB 140-188, T 120-211, T/HB = 75-120%, HF 28-36, Wt 78-158 g. D 1003/1003=16,
M 2+3=10. Skull: gl 41, cbl 34.3-36.9, iob 5.9-6.5, Mt 6.0-6.8)
Upperparts finely grizzled olive-brown, underparts variable but usually whitish. Fur
soft and close, with black guard hairs of moderate length distributed evenly through the
pelage. Tail entirely dark brownish. Feet relatively broad with a distinct pattern of fine
ridges on the pads of the underside.
34
4’
Upperparts finely grizzled olive-brown, underparts slightly paler, buffy-brown with
gray bases. Confined to human settlements.
.................................................................................................................. Rattus rattus
House Rat
(HB 110-219, T 121-220, T/HB = 85-120%, HF 27.5-39, E 16.5-21.6, Wt 80-210 g,
D 1003/1003=16, M 2+3=10 rarely 3+3=12. Skull: gl 41-45.9, cbl 33.8-42.9,
iob 5.8-7.4, mt 6.2-8.6)
Upperparts dull-brown, underparts usually slightly paler. Tail entirely brownish.
Genus Sundamys
Large. Tail entirely dark, usually 110-120 % of Head & Body length. Prominent long black
guard hair on upperparts.
Key to species
1
Very large, Hindfoot length more than 50 mm. Upperparts almost black, underparts
gray with a strong orange tinge. Mountains.
................................................................................................... Sundamys infraluteus
Mountain Giant Rat
(HB 226-295, T 260-343, T/HB = 114.8-120%, HF 52-61, E 29, Wt 237-600 g, D 1003/
1003=16, M 1+2=6. Skull: gl 55.9-63.3, iob 7.8-9.1, mt 10.6-11.6)
Recorded from Gunung Kinabalu (900-2920m) and Trus Madi.
1’
Hindfoot length less than 50 mm. Upperparts grizzled brown with coarse fur and
prominent long black guard hairs. Underparts distinctly paler.
........................................................................................................ Sandamys muelleri
Muller’s Rat
(HB 173-244, T 191-285, T/HB = 108.9-142.5%, HF 37-49, E 20.4-22.7, Wt 160-385 g,
D 1003/1003=16, M 2+0+2=8. Skull: gl 44.1-54.2, iob 6.8-8.0, mt 8.2-10.0)
Upperparts dark brown with a coarse fur and prominent long black guard hair. Under
part distinctly paler. Tail entirely dark brown.
Genus Niviventer
Upperparts brown with numerous rather stiff spines. Under part whitish. Tail uniformly dark, at
least 125 % of Head and Body length, and slightly tufted with hair at the tip.
35
Key to species
1
Upperparts reddish brown, darker and duller along the midline, with inconspicuous
gray under-fur and numerous rather stiff spines. Underparts white or creamy-white.
............................................................................................................. Niviventer rapit
Long-tailed Mountain Rat
(HB 122-163, T 183-233, T/HB=140-159.6%, HF 27-34, D 1003/1003 =16, M 2+2=8.
Skull: gi 41, cbl 30.9-36.9, iob 5.7-6.5, mt 5.8-6.1)
More dominant in lower montane forests.
1
Upperparts pale brown Underparts whitish, usually with a yellow or buff tinge. Distinct
long, black guard hairs.
.............................................................................................. Niviventer cremoriventer
Dark-tailed Tree Rat
(HB 99-160, T 124-219, T/HB = at least 125%, HF 23-28, E 16.3-19, Wt 37-100 g,
D 1003/1003=16, M 2+2=8. Skull: gl 31.5-35.4, iob 5.3-6.1, mt 5.6-6.6)
Upperparts pale brown, usually with a yellowish tinge. Under part whitish, usually
with a yellow or buff tinge. Distinct long, black guard hair and numerous rather stiff
spines in both upper and under part. Tail uniformly dark brown. Tail covered in short
hairs with longer, more prominent hairs on the terminal 2 cm.
Genus Maxomys
Upperparts brown with very stiff and prominent spines. Underparts white or buff. Tail short and
bicolored, being dark brown or black above and white beneath, separated by a sharp line.
Key to species
1
1
2 (1’)
Upperparts dark gray-brown along the midline, paler along the sides of the body.
Underparts whitish to creamy-buff without yellow tinge. Very spiny. Mountains.
.......................................................................................................... Maxomys alticola
Mountain Spiny Rat
(HB 139-176, T 128-180, T/HB = 90-120%, HF 29-37, E 19, Wt 90 g, D 1003/1003=16.
Skull: gl 37.0-41.7, iob 7.0-8.1, mt 5.4-6.1)
Known only from Gunung Kinabalu (1050-3350m) and Trus Madi.
Upperparts brown to reddish brown.
..................................................................................................................................... 2
Underparts white.
..................................................................................................................................... 3
36
2’
3 (2)
3’
Underparts grayish or buffy.
..................................................................................................................................... 4
Upperparts distinctly orange or reddish-brown. Coloration of upperparts extends around
part or all of the neck, forming "collar"'. and around the inner side of the leg above the
ankle.
........................................................................................................... Maxomys surifer
Red Spiny Rat
(HB 160-202, T 187-215, T/HB = 100-120%, HF 35-439 E 31.6, Wt 150-200 g,
D 1003/1003=16, M 2+2=8. Skull: gl 39.4-46.1, iob 5.6-6.9, mt 5.8-6.8)
Upperparts distinctly orange or reddish-brown, slightly darker along the middle, with
numerous short, stiff, dark spines. Under part white with soft, white spines. Coloration
of upper part usually extends around part or all of the neck and around the inner side of
the leg above the ankle. Tail brown above, pale below. Young grayer and duller like
those of Maxomys rajah.
Upperparts brown. Underparts usually with a dark brown streak along the middle. White
on inner side of thigh normally extends unbroken to the feet.
............................................................................................................. Maxomys rajah
Brown Spiny Rat
(HB 138.1-218, T 142-210, T/HB = 102.9-109.3%, HF 33.8-43, E 21.9-22.3, Wt 71218 g, D 1003/1003=16, M 2+2=8. Skull: gl 40.9-48.6, iob 6.6-7.4, mt 6.9-8.1)
Upper parts brown, darker in the midline and with numerous stiff gray-brown spines.
Under part white, with many short, white spines, usually with a dark brown streak
along the middle in adults, but never with an orange throat patch. White of the under
part extends down in a narrow line to the feet. Tail brown above, pale below, and thinly
haired. Young grayer and duller.
Maxomys surifer
Maxomys rajah
Fig. 2 Difference of Maxomys surifer and Maxomys rajah
37
Maxomys surifer: Coloration of upperparts extends around part or all of the neck, forming
“collar”, and around the inner side of the leg above the ankle.
Maxomys rajah: A dark brown streak along the middle. White on inner side of thigh normally
extends unbroken to the feet.
4 (2’)
Larger, Head & Body length 140-171 mm, Hindfoot length 29-35 mm. Upperparts
reddish brown. Underparts grayish with a strong yellow or buffy- red tinge, sometimes
appear entirely yellowish. Numerous stiff spines on entire body.
.............................................................................................. Maxomys ochraceiventer
Chestnut-bellied Spiny Rat
(HB 140-171, T 128-175, T/HB = 111.1-116.3%, HF 29-36, E 21-22, W 92.5-101.5,
D 1003/1003=16. Skull: gl 35.3-39.6, iob 6.2-7.4, mt 5.4-6.0)
4’
Smaller, Head & Body length less than 150 mm, Hindfoot length less than 30 mm.
Underparts not yellowish.
..................................................................................................................................... 5
5 (4’)
Upperparts brown, dark along the midline. Progressively paler towards the underparts.
Underparts buffy. Small. Entire body very spiny.
...................................................................................................... Maxomys baenodon
Small Spiny Rat
(HB 125-140, T 119-141, T/HB = 85-110.2%, HF 25-29, E 19.0-22, D 1003/1003=16.
Skull: gl 33.2-35.2, iob 6.7-6.9, mt 4.1-5.0)
5
Upperparts reddish-brown with gray under-fur. Underparts orange-buff with gray underfur. Tail always shorter than Head & Body length.
.................................................................................................... Maxomys whiteheadi
Whitehead’s Rat
(HB 103-150, T 87-125, T/HB=95.8-98.2%, always shorter than HB, HF 23-30, E 18.319.9, Wt 30-83 g, D 1003/1003=16)
Upper part reddish-brown with gray under-fur and numerous stiff, pale gray spines
with black tips. Under part orange-buff with gray under-fur and numerous soft, pale
spines. Tail dark above, pale below.
Genus Mus
Small. Hindfoot less than 20 mm. An incision at the rear ridge of each upper incisor. First molar
always longer than half the toothrow (MI > M2+M3), hair soft, urine bad smell and it is dyed
in body.
38
nasal
incision
Mus castaneus
nasal
incision
Mus caroli
Fig. 3 Difference of skull of Mus species
Mus castaneus: above, Nasal bones long so that the tip of incisors not be seen in dorsal view.
Mus caroli: below, Nasal bones short so that the tip of incisors be seen in dorsal view.
Key to species
1
Upperparts brown with gray under-fur, underparts slightly paler. Tail entirely dark
brownish. Nasal bones long so that the tip of incisors not is seen in dorsal view (Fig. 3
above). Restricted to buildings in towns
............................................................................................................... Mus castaneas
Asian House Mouse
(HB 50-79, T 80-84, T/HB=90-120%, HF 14-17, E 5.5, Wt 7-12 g, D 1003/1003=16,
M 3+2 =10. Skull: gl 10-20, cbl 18.5, iob 4.1, mt 3.2)
1’
Upperparts brownish-gray, underparts whitish. Tail dark almost black above, pale below.
Nasal bones short so that the tip of incisors be seen in dorsal view (Fig. 3-below). Front
of upper incisors are brown, it’s color more stronger than Mus castaneus. Occurs in
rice-field and grassland.
..................................................................................................................... Mus caroli
Ricefield Mouse
(HB 76, T 78, HF 18, Wt 12 g, D 1003/1003=16, M 3+2=10)
Genus Chiropodomys
Hallux (first digit of the hindfoot) is semi-opposable with a nail. Tail brown, hairy and with a
brush of hair at the tip.
39
Key to species
1
1’
2 (1’)
2’
Large, Head & Body length more than 90 mm, Hindfoot length more than 20 mm.
Upperparts pale gray with a yellow tinge, underparts white.
..................................................................................................... Chiropodomys major
Large pencil-tailed Tree-Mouse
(HB 94-114, T 109-144, HF 21-28, E 16, W 42.0. Skull: gl 27.5-30.2, iob 4.9-5.4,
mt 4.2-5.1)
Confined to forested areas, and are arboreal.
Small, Head & Body length less than 85 mm, Hindfoot length less than 18 mm.
..................................................................................................................................... 2
Upperparts pale gray, underparts white.
................................................................................................ Chiropodemys gliroides
Common Pencil-tailed Tree-mouse
(HB 65-85, T 81-96, T/HB = 110-150%, HF 16-18, Wt 14-22 g, D 1003/1003=16.
Skull: gl 19.7, 22.3, iob 4.4, mt 3.3)
Upperparts pale gray with a yellow tinge, underparts white. Fur short and soft. tail
brown, hairy and with a brush of hairs at the tip. The hallux (big toe) of the hind foot is
semi-opposable, with a nail. Arboreal.
Upperparts reddish-brown, base of hairs gray. Underparts gray with a buff tinge.
............................................................................................... Chiropodomys muroides
Grey-bellied Pencil-tailed Tree-mouse
(HB 66-80, T 85-91, HF 15-17. Skull: cbl 18.0-19.9, iob 4.6-4.9, mt 3.0-3.1)
Confined to forested areas, and are arboreal.
Family Hystricidae
Porcupines
The porcupines are like gigantic rats that are covered with characteristic hard spines or quills
over most of the upper part. The spines of quills normally lie flat on the back, but can be erected
so that they point out in all directions. All three species found in Borneo are normally terrestrial
and usually nocturnal. They sleep during the daytime in underground holes or burrows.
Key to genera and species
1
Tail short, about 25 % of Head & Body length or less, and often hidden beneath the
long spines of the back. Large animals, Head & Body length over 500 mm, Hindfoot
length over 80 mm.
..................................................................................................................................... 2
40
41
1’
Tail long, about half as long as Head & Body length, and clearly visible beyond the
body spines. Tail with a terminal tuft of hairs.
........................................................................................................................... Trichys
Trichys fasciculata
Long-tailed Porcupine
(HB 375-437, T 152-240, HF 61-67, Wt 1.5-2 kg. Skull: gl 81-91)
Gives the overall impression of a large rat. Upper part brown, under part whitish. The
tail is long and scaly, ending in a tuft of large hair, and the quills are quite short, at most
5 cm, and flat. The individual quills are brown at the tip, but pale the base.
2 (1)
General colors black and white. White parts of the quills prominent from a distance.
Nasal bones very large forming nearly half the total length of the skull.
........................................................................................................................... Hystrix
Hystrix brachyura
Common Porcupine
(HB 590-630, T 95-130, HF 80-95, E 38, Wt 8-9.8 kg, D 1013/1013=20, M 0+3+0=6.
Skull: bl 100-200)
Generally black. Long spines or quills white with a black band towards the tip, short
spines on front parts of body, mostly blackish.
2’
General color brown and white. White quill tips not prominent form a distance.
........................................................................................................................Thecurus
Thecurus crassispinis
Thick-spined Porcupine
(HB 500-665, T 90-135, HF 80-90, Wt 8 kg. Skull: gl 110-125)
Generally dark brown. Long spines or quills, dark brown with white tip and base, short
spines in front part of body, gray-brown with a pale tip and base. Hollow quills on the
tail, which are shaken to make a noise.
P A R T
I I
C H I R O P T E R A
42
Order CHIROPTERA
Bats
Bats fall into two suborders. One of them is the Fruit-bats, which feed on fruit or on the nectar of
flowers, use eyes for finding their way, and are often large. Another is the Insectivorous-bats
which feed largely on insects, although they have eyes but they usually use a mechanism for
finding their way about echolocation, and are usually small.
The size is emphasized in the suborder names of MEGACHIROPTERA, i.e. “Big bats”, for
the fruit bats (Pteropodidae) and MICROCHIROPTERA, i.e. “Little bats”, for all the remaining
insectivorous bats (Emballonuridae, Megadermatidae, Nycteridae, Rhinolophidae,
Hipposideridae, Vespertilionidae and Molossidae).
Fig. 1 Bats showing principal features.
Fruit bat (Suborder MEGACHIROPTERA): ear margins forming a continuous ring round the
ear, never with nose leaves, usually with a claw on the second digit (except Eonycteris),
interfemoral membrane is narrow and tail is short or absent.
Insectivorous bat (Suborder MICROCHIROPTERA): ear margins beginning and ending on
the head, noseleaves present or absent, never with a claw on the second digit, interfemoral
membrane well developed, tail usually long.
M: Metacarpal, I: lst digit (thumb). II: 2nd digit. III: 3rd digit, IV: 4th digit, V: 5th digit,
lPh: 1st phalange. 2Ph: 2nd phalange.
Fruit bat
I
Eye
Nose
Claw
1Ph
Ear
M
1Ph
M
2Ph
M
II
1Ph
2Ph
M
Forearm
Claw
M
3Ph
1Ph
Upper arm
Wing membrane
1Ph
IV
2Ph
Tail
Interfemoral
membrane
Hind foot
43
III
2Ph
V
Insectivorous bat
Nose
Eye
I
1Ph Claw
M
M
Ear
M
M
M
Forearm
Upper arm
1Ph 2Ph II
3Ph
2Ph
III
1Ph
2Ph
1Ph
IV
Wing membrane
2Ph
V
Hind foot
Tail
Interfemoral
membrane
Calcar
Eight families can be distinguished as follows. Key to families
Key to families
1. With noseleaf (frills of skin around the nostril) -1 With no tail or very short although the interfemoral membrane is well developed, tragus is
long and forked ............................................................................. Megadermatidae Fig. 2-3
2 Tail is long with a T-shaped tip and fully enclosed in the interfemoral membrane.
................................................................................................................ Nycteridae Fig. 2-4
3 Tail is long and almost completely enclosed within the interfemoral membrane; no tragus,
but ears with antitragus; anterior and posterior noseleaf joined by a connecting process in a
longitudinal vertical plane.
.......................................................................................................... Rhinolophidae Fig. 2-5
4 Tail is long and almost completely enclosed within the interfemoral membrane; no tragus,
but ears with low antitragus; no connecting process.
........................................................................................................ Hipposideridae Fig. 2-6
44
2. No noseleaf --1 Large conspicuous eyes; ear margins forming a continuous ring around the ear, usually
with a claw on the second digit.
................................................................................................... Pteropodidae Fig. 1-former
2 Tail is shorter than the interfemoral membrane and emerges from the middle of it.
....................................................................................................... Emballonuridae Fig. 2-1
3 Tail is long, completely enclosed in the interfemoral membrane.
....................................................................................................... Vespertilionidae Fig. 2-7
4 Tall projecting for at least half its length beyond the interfemoral membrane.
................................................................................................................Molossidae Fig. 2-2
Fig. 2 Distinctive shape of seven families of insectivorous bat
Showed the relation between tail and interfemoral membrane, nose with noseleaves, left ear
with tragus or antitragus, nostrils of Murininae and characteristic third digit of Miniopterinae.
Noseleaves: frills of skin around the nostril.
Tragus: a cartilaginous projection in the form of a rod or leaf in front of the funnel-shaped ear.
Antitragus: a lobe of the outer edge of the ear.
1
Emballonuridae: tail emerges from the middle of interfemoral membrane.
2
Molossidae: tail projecting for at least half its length.
3
Megadermatidae: with no tail, tragus is long and forked.
4
Nycteridae: tail is long with T-shaped tip.
5
Rhinolophidae: ears with antitragus.
6
Hipposideridae: ears with antitragus.
7
Vespertilionidae: tail is long, completely or almost completely enclosed in interfemoral
membrane.
7-(1) Nostrils of Murininae: nostrils extended into short tubes that protrude either side of the
muzzle.
7-(2) Ears of Kerivoulinae and Vespertilioninae: Ears of Kerivoulinae are funnel-like with a
large flap on the outside, tragus long, narrow and pointed. Ears of Vespertilioninae shape
with tragus of common bats. Myotis (ears moderately long and triangular at the tip, tragus
tapered), Pipistrellus (ears generally rounded, tragus shorter and not tapered).
45
7-(3) the third digit of Miniopterinae: the third digit, which has a short first phalanx and very
long terminal phalanx. 2Ph+3ph =3 X lPh or lPh < 1/2 X 3Ph.
Cf: other species 2Ph + 3Ph 2 X lPh or lPh 3Ph.
2 Molossidae
1 Emballonuridae
Tail
Interfemoral membrane
Interfemoral
membrane
Tail
3 Megadermatidae
Tailess
Ear
Nose
Interfemoral membrane
Tragus
4 Nycteridae
Ear
Nose
Tragus
Interfemoral membrane
Tail
Ear
5 Rhinolophidae
Tail
Nose
Interfemoral membrane
Antitragus
Hair
6 Hipposideridae
Tail
Nose
Interfemoral membrane
Antitragus
46
7 Vespertilionidae
Tail
Interfemoral membrane
7-(1) Murininae
distinct nose shape
Murininae
nostlis
mouth
nostlis
Common type
Murininae
(under view)
7-(2) Kerivoulinae
distinct ear shape
Vespertilioninae
Myotis
Kerivoulinae
Funnel-like ear
2 Ph
7-(3) Miniopterinae
3 Ph
1 Ph
III
the third finger has very long terminal phalanx
47
Metacarpal
Pipistrellus
Family Pteropodidae
Fruit Bat
Fruit bats have large, conspicuous eyes because they lack echolocation (except Rousettus) and
they rely instead on eyesight and smell to find their way. The ears are relatively small and simple
with a complete unbroken margin. The teeth are simple, rather degenerate, because they feed
mainly in nectar and fruit.
The second digit is well-developed independent on the third digit and usually with a claw at the
end (except Eonycteris), and using the claws as well as their feet to grip branches, they often
climb through trees.
Key to genera and species
1
With no claw on the second digit.
.................................................................................................................... Eonycteris
1’
With a claw on the second digit.
................................................................................................................................... 2
2 (1’) Very long forearm over 120 mm. First digit very long. Tail absent.
....................................................................................................................... Pteropus
2’
Smaller forearm less than 100 mm. Tail absent or present.
................................................................................................................................... 3
3 (2’) Small bats with a long narrow muzzle and very long tongue.
.............................................................................................................. Macroglossus
Macroglossus minimus
Long-tongued Nectar Bat
(FA 37.6-45, T none or very short 4.1-5.7, HF 8.1-10.1, E II.-16, W 11.2-16,8 g, pregnant female
15-20 g, D 2132/2133=34, M 1+0+0=2. Skull: gl 25.0-27.0) Small bats with a long narrow
muzzle and a very long tongue, tail minute or lacking. Lower jaw thin and weak.
3’
Tongue shorter, muzzle shorter.
................................................................................................................................... 4
4 (3’) Muzzle nearly a long as in Macroglossus. Well developed tail about 10-15 mm.
5 upper and 6 lower cheek teeth (premolar + molar).
..................................................................................................................... Rousettus
4’
Muzzle short giving a “dog faced” appearance. 3-5 upper and 5 lower cheek teeth.
................................................................................................................................... 5
5 (4’) Only 3 upper cheek teeth.
............................................................................................................... Dyvacopterus
48
Dyacopterus spadiceus
Dayak Fruit Bat
(FA 77-82, T 19-28.6, HF 18-19.6, E 17.5-21.4, W 75-100g, D 2121/2132=28,
M 1+0+0=2. Skull: gl 37.0-40.0)
Similar to Cynopterus, with a distinct tail, short muzzle and thick jaws.
5’
6 (5)
With 4-5 upper cheek teeth.
................................................................................................................................... 6
Small dark bat, pale spots on the wings. Only one pair of lower incisors.
D 2132/1132=30.
................................................................................................................Balionycteris
Balionycteris maculata
Spotted-winged Fruit Bat
(FA 37.6-45, T none, HF 6.9-9.5, E 9-12, W 10-15g, D 2132/1132=30, M 1+0+0=2.
Skull: gl 21.0-23.0)
A small dark fruit bat easily recognized by pale spots on the wings. Upper incisors close together
and angled inwards.
6’
No spots on the wings.
................................................................................................................................... 7
7 (6’) Tail present.
................................................................................................................................... 8
7’
Tail present.
................................................................................................................................... 9
8 (7) Medium to large bats with a short stout muzzle. White wing bones and rims to the
ears. D 2131/2132=30.
.................................................................................................................. Cynopterus
8’
No white wing bones and rims to the ears. Only one pair of lower incisors.
D 2131/1132=28.
..................................................................................................................... Penthetor
Penthetor lucasi
Dusky Fruit Bat
(FA 56.0-63.4, T 7.9-14, HF 11.2-13.9, E 14-17.1, W 30-44g, D 2131/1132=28, M 1+0+0=2.
Skull: gl 27.0-31.0)
Medium sized bats, very similar to Cynopterus, but coloration generally dark gray-brown.
Upperparts dark gray-brown, underparts pale buff gray. Top of head often distinctly darker down
the center and paler near the eyes. Ears have dark edges. Only one pair of lower incisors.
9(7’) Small bat, two pairs of lower incisors. D 2131/2132=30.
......................................................................................................................Chironax
49
Chironax melanocephalus
Black-capped Fruit Bat
(FA 42.8-45.6. T none, E 13, W 15.7g, D 2131/2132=30)
Small bat very similar to Cynopterus, but lacking a tail.
9’
One pair of lower incisors. D 2131/1132=28.
................................................................................................................................. 10
10 (9’) Small to medium bats, FA more than 46 mm. Muzzle short. Upperparts uniform
mid-brown; posterior margin of interfemoral membrane more or less naked.
................................................................................................................... Megaerops
10’
Small bat, FA less than 46 mm. Muzzle narrow. Upperparts gray, crown distinctly
darkened, posterior margin of interfemoral membrane thick furred to the ankle.
..................................................................................................................... Aethalops
Aethalops alecto
Grey Fruit Bat
(FA 41.3-46, T none, W 9.1-11.1, E 9.9-13, W 13.5-25.3 g. D 2131/1132=28, M 1+0+0=2.
Skull: gl 25.0)
Small tailless bat similar to Balionycteris, but without the second upper molar. Interfemoral
membrane narrows but thickly furred. D 2131/1132=28. Mountains.
Genus Rousettus
Large with a fairly long muzzle, a claw on the second digit and a well-developed tail. Produces
a distinctive clicking call with the tongue for echolocation. D 2132/2133=34.
Key to species
1
Wings attached to sides of back, separated by a broad band of fur.
........ Rousettus amplexicaudatus
Geoffroy’s Rousette
(FA 71-87, T 11-21, HF 16.4-17.7, E 16.1-19, W 59-82g, D 2132/2133=34, M 1+0+0 =2.
Skull: gl 35.0-40.0)
Upperparts gray-brown to brown, darker on top of head. Underparts paler gray-brown.
1
Wing membranes joined along middle of the back and merging with tail membrane
over legs.
........ Rousettus spinalatus
Bare-backed Rousette
(FA 83-89, T 12-18, E 15.5-18, W Male 88-94 g, Female 66-88 g, D 2132/2133=34)
Adult males have yellow tufts of fur on the sides of the neck.
50
Genus Pteropus
Very large bats include the largest in the world. First digit very long, second digit has a
well-developed claw. D 2132/2133=34.
Key to species
1
Forearm 177-200 mm, Wt 565-1,100 g.
........ Pteropus vampyrus
Large Flying Fox
(FA 177-200, T none, HF 46.3-50, E 39.7-41.7, W 565-1,100 g, D 2132/2133=34,
M 1+0+0=2. Skull: gl 72.0-78.0)
The largest of all the bats. Black back with gray streaking, backs of head, neck and shoulders
reddish, underparts blackish brown.
1’
Forearm not more than 140 mm. Islands off the coast.
........ Pyeropus hypomelanus
Island Flying Fox
(FA 121-140, T none, E 25, W 213 g, D 2132/2133=34. Skull: gl 62.0-68.0)
Genus Cynopterus
Medium to large bats with a short stout muzzle. Most species have brown fur with a yellowish or
reddish tinge and contrasting whitish wing bones and rims to the ears. D 2131/2132=30.
Key to species
1
Forearm 51.6-67.2 mm, Tail 6.2-15.5 mm, Ear not more than 17 mm.
........ Cynopterus brachyotis
Short-nosed Fruit Bat
(FA 51.6-67.2, T 6.2-15.5. HF 10.5-17,7, E 14-17. Wt 21-42.5g, pregnant female 47-50g,
D 2131/2132=30. M 1+0+0=2)
A medium sized fruit bat with a short muzzle, brown with brighter collar, which is dark
orange in adult males, yellowish in females. Ears and wing bones are edged in white.
Molar teeth plain.
1’
Forearm 65 76 mm, T 18-22 mm, E more than 18 mm.
........ Cynopterus sphinx
Greater Short-nosed Fruit Bat
(FA 65-76, T 18-22, E more than 18. Skull: cbl 30.1-32.0)
Lower cheek teeth rounded without accessory cusps.
1”
51
Forearm 68-76 mm, T 13-14 mm, E 17-20 mm. Cheek teeth broader and squarer than
other Cynopterus, a small distinct cusp present on the occlusal surfaces of lastpremolar
and first molar of lower jaw.
........ Cynopterus horsfieldi
Horsfield’s Fruit Bat
(FA 68-76, T 13.1-18, HF 14.9-16.5, E 17-20, W 41.0-70 g. D 2131/2132=30, M 1+0+0=2.
Skull: gl 32.0-35.0, cbl 31.0-31.7)
Similar to the Cynopterus brachyotis but slightly larger. The last premolar and first molar of
lower jaw gave peg-like projections in the center of each. Feeds mainly on fruit.
Fig. 3 The last premolar and first molar of Cynopterus
Cynopterus brachyotis
Cynopterus horsfieldi
Genus Megaerops
Small to medium bats, similar to Cynopterns, but with a slightly shorter nose, no visible tail.
Only one pair of lower incisors. D 2131/1132=28.
Key to species
1
FA 50-58 mm.
........ Megaerops ecaudatus
Tailless Fruit Bat
(FA 50-58, T none, MF 8-12.6, E 13-15.4, W 18.2-38 g, D 2131/1132=28, M 1 +0+0=2.
SWI: gl 27.0, cbl 24.0-26.2)
Upperparts yellowish brown to reddish brown with gray bases to fur; underparts paler and
grayer. Muzzle short with broad slightly tubular nostrils.
1’
FA 46-51 mm. White shoulder tufts. First phalanges of both IV and V digit whitish.
........ Megaerops wetmorei
White-collared Fruit Bat
(FA 46-51, T none, HF 9-12.9, E 11.7-14.4, W 14-21.6 g, D 2131/1132=28, M 1+0+0=2.
Skull: cbl 21.2-23.2)
Body fur pale gray brown, lower back darker. Fifth metacarpal, and first phalanges of both IV
and V digit have whitish part. some of them have white shoulder tufts. There are two types: one
has large white tufts on side of neck extending onto back to form a broken collar; another type
lacks white collar. Muzzle short and thick.
52
1’
FA 46-51 mm. White shoulder tufts. First phalanges of both IV and V digit whitish.
........ Megaerops wetmorei
White-collared Fruit Bat
(FA 46-5 1, T none, HF 9-12.9, E 11.7-14.4, W 14-21.6 g, D 2131/1132=28, M 1+0+0=2.
Skull: cbl 21.2-23.2)
Body fur pale gray brown, lower back darker. Fifth metacarpal, and first phalanges of both IV
and V digit have whitish part. some of them have white shoulder tufts. There are two types: one
has large white tufts on side of neck extending onto back to form a broken collar; another type
lacks white collar. Muzzle short and thick.
Fig. 4 Identifiable key to the Megaerops bats.
Megaerops ecaudatus: thicker muzzle with broad slightly tubular nostrils. Megaerops wetmorei:
fifth metacarpal, and first phalanges of both IV and V digit have whitish part. some of them have
white shoulder tufts.
slightly thicker muzzle
Megaerops ecaudatus
Megaerops wetmorei
pale grey brown
M
1Ph
pale white
upperside
white shoulders tufts
(lacking in some individuals)
1Ph
bone and both sides white
Megaerops wetmorei
53
Genus Eonycteris
Medium to large fruit bats with a long narrow muzzle and a very long sticky tongue. Distinguished
from all other Bornean fruit bats by the absence of a claw on the second digit. D 2132/2133=34.
Key to species
1
Forearm 62-70 mm. Pale in color with a shorter muzzle.
........ Eonycteris spelaea
Cave Nectar Bat
(FA 62-70, T 14.8-19.1, HF 10.1-16.6, E 17-20. W 44-62.8 g, D 2132/2133=34. M 1+0+0=2.
Skull: gl 32.0-37.0)
Upperparts gray-brown, underparts slightly paler. Fur short. Lacks a claw on the second
digit. Muzzle elongate.
1’
Forearm 71-80 mm. Longer more decurved muzzle and darker fur.
........ Eonycteris major
Greater Nectar Bat
(FA 71-84.2, T 18-24.2, HF 15.7-20, E 20.1-21.1, W 91-114 g, D 2132/2133=34)
Fur uniform dark blackish brown. Muzzle elongate and slightly decurved.
Family Emballonuridae
Sheath-tailed Bats
Small to medium sized bats with a distinctive tail, which is enclosed until about half way along
the interfemoral membrane, and the tip of the tail lies freely on the surface of the interfemoral
membrane. When the legs are outstretched the membrane completely encloses the tail. The
muzzle is simply with no noseleaf.
Key to genera
1
1’
Two pairs of upper incisors. Rather small bats. Forearm 50 mm or less.
D 212/33123=34
................................................................................................................. Emballonura
Only one pair of upper incisors. Rather larger bats. Forearm 55 mm or more.
D 1123/3123=32
...................................................................................................................... Taphozous
54
Genus Emballonara
Small and dark brown. Two distinct pairs of upper incisors, anterior upper premolar very small.
D 2123/3123=34.
Key to species
1
Forearm 45-49.5 mm. A short gap (Diastema) between the first and second upper
premolars and a longer palate in front of the molars, condylobasal length of skull more
than 14.0 mm, maxillary toothrow more than 5.7 mm.
........ Emballonura alecto
Greater Sheath-tailed Bat
(FA 45-49.5, T 13-19.5, BF 6.3-9.5, E 9.7-15, W 4-9 g, D 2123/3123=34, M 1+0+0=2.
Skull: condylocanine length (ccl), from the back of the occipital condyle to the front of
the canine, 12.7-13.8)
Uniformly dark brown to reddish-brown. Roosting posture, supported by the wrists, is distinctive,
as is their alarm call - a shrill squeak.
1’
Forearm 42.4-45.6 mm. No gap between the upper premolars, condylobasal length of
skull less than 13.2 mm, maxillary toothrow less than 5.5 mm.
........ Emballonura monticola
Lesser Sheath-tailed Bat
(FA 42, 4-45.8, T 10.9-19.3, HF 6.2-8.4, E 10-13.6, W 4.5-7.5 g. D 2123/3123=34, M 1+0+0=2.
Skull: gl 12.0-14.0, condylocanine length (ccl), 11.8-12.8)
Very similar to the Greater Sheath-tailed Bat, but slightly smaller.
Genus Taphozous
Medium to large bats variably colored. Often have a glandular throat pouch or a pocket in the
wing at the wrist. Only one pair of upper incisors which are very small. D 1123/3123=32.
Key to species
1
1’
Wing pouch (A pocket on the wing formed by a membrane, which connecting the forearm
and the last fingers) well developed.
..................................................................................................................................... 2
FA over 66 mm. No wing pouch (poorly developed), wings very white, chin pouch in
both sexes.
........ Taphozous saccolaimus
Pouched Tomb Bat
(FA 70.4-78, T 27.5-34.4, HF 16.7-18, E 19-20.8, W 39.5-54.5 g, D 1123/3123=32, M 1+0+0=2)
55
2 (1) FA 60-65 mm. Chin furred in both sexes. Male with a patch of black hair under the
chin. No chin pouch. Tail thickened at tip.
........ Taphozous melanopogon
Black-bearded Tomb Bat
(FA 60-65, T 23.3-28.7. BF 9.6-12.3, E 18.8-22, W 21.7- 26 g, D 1123/3123=32, M 1+0+0=2.
Skull: gl 19.0)
2’
FA 52-58 mm. Chin naked with a distinct pouch in males. Tail tapered at tip.
........ Taphozous longimanus
Long-winged Tomb Bat
(FA 52-58, D 1123/3123=32. Skull: gl 19.0)
Fig. 5 Wing pouch and Chin pouch of Taphozous
wing pouch
Taphozous
saccolaimus
no wing pouch
(poorly
developed)
well developed
no wing pouch
(poorly
developed)
well developed
Taphozous
well developed
Lomgimanus
well developed
Taphozous
melanopogon
chin
glandular
pouch
glandular
pouch
covered with fur
patch of black
hair, no pouch
glandular pouch,
chin naked
covered with fur
no pouch
no pouch,
chin naked
I
Forearm
II
III
Chin pouch
Wing Pouch
IV
V
56
Family Megadermatidae
False Vampires
Medium to large bats with large rounded ears joined across the top of the head, and with a large
erect noseleaf. The tragus is long and forked. The tail is very short, not visible externally.
Genus Megaderma
Premaxillaries minute with no upper incisors. Canines project well forward with a distinct
secondary cusp.
Megaderma spasma
Lesser False Vampire
(FA 52-61, HF 17.1, E 30.9-40, Tragus 17.3, W 20-28 g. D 0123/2123=28. Skull: gl 25.0-27.0)
Fur pale gray to gray-brown. Noseleaf has long dorsal lobe with stiffened central ridge. Ears
very large joined at bases. Tragus is long and forked. No visible tail although interfemoral
membrane is well developed.
Family Nycteridae
Hollow-faced Bats
Medium to large bats with very large rounded ears, which do not meet across the top of the head.
The tail is long with a T-shaped tip and fully enclosed in the interfemoral membrane.
D 2113/3123=32.
The skull is distinguished by the deep frontal depression with broad ridges on either side. The
family contains only one genus.
Genus Nycteris
Nycteris javanica
Hollow-faced Bat
(FA 45-51, T 65-73.1, HF 6.8-8.9, E 27-31, W 12-17 g, D 2113/3123=32, M 1+0+0=2.
Skull: gl 21.0)
Deep hollow groove in the middle of face fringed with large flaps. Tail is long with T-shaped tip
and fully enclosed in the interfemoral membrane.
57
Family Rhinolophidae
Horseshoe Bats
Small to medium sized bats with an elaborate noseleaf. The anterior section is rounded and
roughly horseshoe-shaped. In the middle, a raised portion behind the nostrils is called a sella.
Behind this is the posterior noseleaf that rises to a long lancet-shaped point. The ears are large
with a prominent fold on the outside edge, the antitragus. The tail is fully developed in all the
Bornean species, and is wholly enclosed in the interfemoral membrane. D 1123/2133=32.
The family contains only one genus Rhinolophus (Fig. 6, Fig. 7).
Posterior noseleaf
Connecting process
Sella
Ear
Lappet
Nostril
Internal septum
Antitragus
Eye
Anterior noseleaf
Fig. 6 Noseleaf of Horseshoe Bat, Rhinolophus.
Showing names of parts.
Key to species
1
Sides of the horizontal base of the sella much expanded make Tappets, fur rather long
and fluffy.
..................................................................................................................................... 2
1’
Sides not expanded, scarcely concealing the nasal aperture, fur thick but not fluffy.
..................................................................................................................................... 4
58
Posterior noseleaf
Sella
Connecting process
Rhinolophus creaghi
Anterior noseleaf
Rhinolophus arcuatus
Rhinolophus affinis
Rhinolophus philippinensis
Rhinolophus borneensis
Rhinolophus pusillus
Rhinolophus acuminatus
Lappet
Lappet
Lappet
Rhinolophus trifoliatus
Rhinolophus sedulus
Rhinolophus luctus
Fig. 7 Noseleaves of Horseshoe Bat (side view).
Difference of noseleaves is important to identify the species.
Rhinolophus philippinensis : Sella long with cup-like base.
Rhinolophus creaghi
: conical tuft of hairs instead of connecting process.
Rhinolophus acuminatus : connecting process pointed.
Rhinolophus arcuatus
: connecting process rounded, originating at tip of Sella.
Rhinolophus borneensis
: connecting process varies from slightly rounded to bluntly pointed.
Rhinolophus pusillus
: connecting process triangular and slightly pointed.
Rhinolophus affinis
: connecting process rounded, originating from back of Sella.
Rhinolophus luctus
: lateral lappets at base of Sella, noseleaf dark.
Rhinolophus sedulus
: lateral lappets at base of Sella.
Rhinolophus tnfoliatus
: lateral lappets at base of Sella, noseleaf yellowish.
59
2 (1)
Forearm 58-67 mm. Dark gray in color, almost black. Noseleaf and ears gray.
........ Rhinolophus luctus
Great Woolly Horseshoe Bat
(FA 58-67, T 38-50, E 31-35, Wt 29-37g, D 1123/2133=32. Skull: gl 27.5)
Largest Bornean Horseshoe Bat. Hairs pale-tipped. Fur long and woolly. Nose leaf has lateral
Tappets at base of sella.
2’
Forearm under 55 mm.
................................................................................................................................... 3
3 (2’) Forearm 47-52.1 mm. Fur pale. Noseleaf and ears yellowish.
........ Rhinolophus trifoliatus
Trefoil Horseshoe Bat
(FA 47-52.1, T 27-38.6, HF 11.4-13.3, E 21-27, W 10.5-18g. D 1123/2133=32. Skull:
gl 19.0-23.0)
Fur long and woolly; pale buffy brown to brownish gray. Noseleaf pale yellow, has lateral lappets
at base of sella.
3’
Forearm 40-44 mm. Small and dark brown, hairs tipped pale gray. Noseleaf and
ears gray.
........ Rhinolophus sedulus
Lesser Woolly Horseshoe Bat
(FA 40-44, T 20-25, HF 9.8-9.9, E 22-23.1, W 8.3-11 g, D 1123/2133=32)
4 (1’) Sides of the base of the sella turned up so as to form a cup-Re structure. Ears very
long. Forearm 48-53.1 mm.
........ Rhinotophus philippinensis
Philippine Horseshoe Bat
(FA 48-53.1, T 25-33, HF 8.1-9.5, E 27-31.1, W 7-13 g. D 1123/2133=32, M 1+0+1=4)
Fur usually uniformly dark-brown. Noseleaf and ears dark gray. Ears very large. Posterior noseleaf
long and triangular. Sella long and protruding expanded at base into a cup-like structure between
the nostrils.
4’
Sides of the base of the sella not cup-like structure.
................................................................................................................................... 5
5 (4’) Conical tuft of hairs instead of connecting process. Forearm 46-51 mm.
........ Rhinolophus creaghi
Creagh’s Horseshoe Bat
(FA 46-52, T 19.1-21.9, HF 8.2-11.7, E 19-23, W 10.5-13.5g. D 1123/2133=32, M 1+0+1=4)
Fur color variable, ranging from gray-brown to yellowish-brown to reddish. Ears and noseleaf
moderately large. A distinctive conical tuft of stiff hairs on anterior face of posterior noseleaf
replaces connecting process.
60
5’
Connecting process there.
................................................................................................................................... 6
6 (5’) Connecting process rounded.
................................................................................................................................... 7
6’
Connecting process pointed.
................................................................................................................................... 8
7 (6)
Connecting process originating at tip of sella. Forearm 46-49 mm.
........ Rhinolophus arcuatus
Arcunte Horseshoe Bat
(FA 46-49.0, T 22.3-28.4, HF 8.8-9.9, E 18.4-21.4, W 7.5-12.5g, D 1123/2133=32)
Uniformly dark brown. Ears large. Noseleaf broad, coverings muzzle, lacking lateral lappets.
Connecting process broadly rounded, originating from tip of sella. Sella broad and slightly
expanded at base.
7’
Connecting process originating from back of sella. Forearm 49-54 mm.
........ Rhinolophus affinis
Intermediate Horseshoe Bat
(FA 49-54, T 22-26.4, HF 8.8, E 20.5-25, W 12.5-14.0 g, D 11 23/2133=32)
Upperparts dark brown to reddish brown, underparts slightly paler. Noseleaf simple without
extra lappets on sella. Connecting process broadly rounded, originating from below tip of sella.
8 (6’) Connecting process pointed, triangular in outline. Wing arise from near the base of
the toes. Forearm 48-50 mm.
........ Rhinolophus acuminatus
Acuminate Horseshoe Bat
(FA 48-50, T 21-31, HF 7-9.8, E 18.4-21, W 11-14 g, D 1123/2133=32, M 1+0+1=4.
Skull: gl 22)
8’
Wing arise from the ankles. Forearm 40-44.3 mm.
........ Rhinolophus borneensis
Bornean Horseshoe Bat
(FA 40-44.3, T 21-29, HF 6.8-8.4, E 15.6-19, W 6.5-9.4 g. D 1123/2133=32, M 1+0+1=4.
Skull: gl 19-21)
Gray-brown to reddish, ears relatively small.
61
8’’
Very small. Forearm 37-40.5 mm.
........ Rhinolophus pusillus
Least Horseshoe Bat
(FA 37-40.5, T 13-21.1, HF 5.3-6.3, E 12.5-16.1, W 4-5.5 g, D 1123/2133=32. Skull:
condylocanine length (ccl) 13, iob 2.1)
Smallest Bomean horseshoe bat, but with relatively large ears. Upperparts light brown, underparts
slightly paler. Noseleaf simple and small, connecting process triangular and slightly pointed,
sella parallel sided.
Family Hipposideridae
Roundleaf Bats
Vary in size from quite small to moderately large with an elaborate noseleaf. The anterior noseleaf
is rounded and somewhat horseshoe-shaped. The median leaf is low cushion-like structure
expanded laterally without a sella. While the posterior leaf is low and rounded (Fig. 8).
D 1123/2123=30.
Vertical septum
Pocket
Ear
Frontal sac
Antitragus
Posterior noseleaf
Eye
Nostril
Intermediate noseleaf
Lateral leaflet
Anterior noseleaf
Internal septum
Fig. 8 Noseleaf of Roundleaf Bat, Hipposideros.
Showing naming of parts (Hipposideros diadema).
62
Key to genera
1
Horseshoe-shaped anterior noseleaf. Tail there.
................................................................................................................Hipposideros
1’
Lateral leaflets large and expanded forwards. No tail.
........ Coelops
Coelops robinsoni
Lesser Tailless Roundleaf Bat
(FA 34-37, T-none, HF 5.8, E 12-14, W 3 g, D 1123/2123=30. Skull: mtI-M 4.7)
Fur long and soft, brown to blackish with paler tips, underparts slightly grayer. Ears rounded and
lacking supporting ridges. Narrow interfemoral membrane with no visible tail. Anterior margin
of noseleaf deeply notched, forming two lobes.
Genus Hipposideros
Typical horseshoe-shaped anterior nose-leaf. Tail short to moderate, largely enclosed in
interfemoral membrane. D 1123/2123=30.
Key to species
1
Large bats. Forearm over 75 mm, 3-4 lateral leaflets. Pale spots on shoulder.
........ Hipposideros diadema
Diadem Roundleaf Bat
(FA 76-89, T 41.0-55.1, HF 12.5-16.7, E 27-30.3, W 30-49 g, D 1123/2123=30. Skull:
gl 28.0-32.0)
Largest Bornean horseshoe bat. Fur of upperparts dark brown with pale bases, white patches on
the shoulders and sides. Underparts grayish white. Noseleaf with 3 or 4 lateral leaflets. Posterior
noseleaf large and rounded.
1’
Small or middle bats. Forearm less than 70 mm.
................................................................................................................................... 2
2 (1’) Three lateral leaflets. Forearm 56-65.4 mm.
........ Hipposideros larvatus
Intermediate Roundleaf Bat
(FA 5,6-65.4, T 36.2-41.1, HF 9.9-10.8, E 22.1-23.9, W 15.5-24.5 g, D 1123/2123=30,
M 1+0+1=4)
Upperparts dark gray-brown or reddish-brown. Underparts slightly paler. Noseleaf has 3 lateral
leaflets.
2’
63
Two lateral leaflets.
................................................................................................................................... 3
Hipposideros dyacorm
Hipposideros ater
Hipposideros bicolor
Hipposideros cineraceus
Hipposideros sabanus
Hipposideros larvatus
Hipposideros ridleyi
Hipposideros cervinus
Hipposideros coxi
Hipposideros galeritus
Coelops robinsoni
Fig. 9 Difference of noseleaves of Hipposideridae.
: No lateral leaflets, noseleaf pinkish, internal septum slightly swollen
at base, narrow in middle.
Hipposideros bicolor
: No lateral leaflets, internal septum fairly uniform width.
Hipposideros cineraceus : No lateral leaflets, internal septum swollen in middle.
Hipposideros dyacorum : No lateral leaflets, noseleaf dark, ears triangular.
Hipposideros ridleyi
: No lateral leaflets, disk-like internal septum.
Hipposideros coxi
: two lateral leaflets, noseleaf very large.
Hipposideros larvatus : Three lateral leaflets.
Hipposideros sabanus : No lateral leaflets, no supporting septa on posterior noseleaf.
Hipposideros cervinus : Two lateral leaflets, intermediate noseleaf narrower than posterior
noseleaf, tail less than 30 mm.
Hipposideros galeritus : Two lateral leaflets, intermediate noseleaf wider than posterior
noseleaf, tail over 30 mm.
Hipposideros diadema : (see Fig. 8). Three or four lateral leaflets.
Coelops robinsoni
: Anterior noseleaf forms two lobes, lateral leaflets expanded
forwards.
Hipposideros ater
64
2’
No lateral leaflets.
................................................................................................................................... 5
3 (2’) Forearm 53-55 mm. Noseleaf very large (13 X 10 mm).
........ Hipposideros coxi
(FA 53-55, W10g, D1123/2123=30)
3’
Cox’s Roundleaf Bat
Forearm less than 51 mm.
................................................................................................................................... 4
4 (3’) Forearm 45-51 mm. Tail over 30 mm. Intermediate noseleaf wider than posterior
noseleaf.
........ Hipposideros galeritus
Cantor’s Roundleaf Bat
(FA 45.2-51, T 30.0-43, HF 5.2-7.4, E 12.2-15.2, W 6.3-8.5 g, D 1123/2123=30. Skull: gl 16.0)
Fur usually dark gray-brown. Noseleaf pinkish gray. Ears broad, rounded at base, triangular at
tip. Noseleaf with two lateral leaflets. Median noseleaf broader than posterior noseleaf.
4’
Forearm 44-50.2 mm. Tail less than 30 mm. Intermediate noseleaf narrower than
posterior noseleaf.
........ Hipposideros cervinus
Fawn Roundleaf Bat
(FA 44-50.2, T 21-30.6, HF 5.4-9.3, E 13.5-17.2, W 5.5-10.2 g, D 1123/2123=30, M 1+0+1=4)
Fur color varies from gray-brown or yellowish-brown to bright red-brown or orange. Noseleaf
grayish pink. Noseleaf simple with two lateral leaflets. Median noseleaf narrower than posterior
noseleaf.
5 (2”) Very small. Forearm 34-37 mm. No supporting septa (vertical septa or ridges) on
posterior noseleaf.
........ Hipposideros sabanus
Least Roundleaf Bat
(FA 34-37, T 21.2, W 4.2-6 g, D 1123/2123=30. Skull: condylocanine length (ccl) 4.4-5.1)
5’
Supporting septa on posterior noseleaf dividing four cells.
................................................................................................................................... 6
6 (5’) Noseleaf and ears large. Disk-like internal septum.
........ Hipposideros ridleyi
Ridley’s Roundicaf Bat
(FA 47-49, T 25-29, HF 22.1, E 20-27, W 6.5-9.5 g, D 1123/2123=30, M 1+0+0=2)
Fur uniformly dark brown. Ears very large and rounded. Noseleaf large, no lateral leaflets. Internal
septum expanded into a large disk.
65
6’
Internal septum no disk-like shaped.
................................................................................................................................... 7
7 (6’) Long forearm 45-48 mm. Internal septum fairly uniform width.
........ Hipposideros bicolor
Bicolored Roundleaf Bat
(FA 45-48, T 27-3 1, E 17.5-20, W 7-8.5g, D 1123/2123=30. Skull: condylocanine length
(ccl) 16.4)
7’
Internal septum not uniform width.
................................................................................................................................... 8
8 (7’) Ear triangular. Noseleaf dark. Forearm 38-43 mm. First lower premolar at least half
height of the second.
........ Hipposideros dyacorum
Dayak Roundleaf Bat
(FA 38-44.2, T 19-24.2, HF 5.8-7.9, E 14.6-18. W 4.5-7.6g, D 1123/2123 =30)
Fur uniformly dark brown. Noseleaf, ears and wing membranes dark gray-brown. Ears broadly
triangular and pointed. Noseleaf simple without lateral leaflets.
8’
Noseleaf pinkish. Ears rounded.
................................................................................................................................... 9
9 (8’) Internal septum swollen at base, narrow in middle. Forearm 39-43 mm.
........ Hipposideros ater
Dusky Roundleaf Bat
(FA 39-43, T 22-26, HF 5, E 15-17.5, W 4.5-7.5 g, D 1123/2123=30. Skull: condylocanine
length (ccl) 15.3)
9’
Internal septum swollen in middle. Forearm 34.1-40.5 mm. First lower premolar less
than half height of the second.
........ Hipposideros cineraceus
Ashy Roundleaf Bat
(FA 34.1-40.5, T 22.2-30, HF 5.5-6, E 15-21, W 3.5-7.3 g, D 1123/2123=30. Skull: condylocanine
length (ccl) 13.2-13.9)
66
Family Vespertilionidae
Common Bats
The largest, most diverse and most widespread family of bats. The nose is simply without any
noseleaf. The ears are small to large with a well-developed tragus. The tail is long, completely
enclosed in the interfemoral membrane.
Key to subfamilies
1
Very long terminal phalanx on the third digit, total length of 2Ph & 3Ph about three
times as long as the first phalanx (2Ph+3Ph=3 X lPh) or the first phalanx less than
half as long as the third phalanx (lPh < 1/2 X 3Ph).
............................................................................................................. Miniopterinae
1’
Total length of 2Ph & 3Ph on the third digit not more flian twice as long as the first
phalanx.
................................................................................................................................... 2
2 (1’) Nostrils extended into short tubes, which protrude either side of the muzzle.
Murininae
mouth
nostlis
Common type
Murininae
........................................................ Murininae
(under view)
2’
Nostrils not tube shaped.
................................................................................................................................... 3
3 (2’) Ears funnel like with a large flap on the outside. Tragus long, narrow and pointed
(Fig. 2-7-2).
(Left ear)
67
............................................................................................. Murininae
3’
Another common bats.
............................................................................................................ Vespertilioninae
Subfamily Vespertilioninae
Key to genera
1
Only one pair of upper incisors that are large, conical and well developed.
D 1113/3123=30.
(Upper incisor) ......................................................................................... Scotophilus
Scotophilus kuhlii
Yellow House Bat
(FA 47-52, T 44.6-52, HF 8.9-9, E 11.4-15, Tragus 6.9-7, W 16-23 g, D 1113/3123=30.
Skull: gl 17.0)
Ears moderate, tragus very long and curved forwards.
1’
Two pairs of upper incisors.
................................................................................................................................... 2
2 (1’) Ears moderately long and triangular at the tip. Tragus tapered, bluntly pointed and
bentslightly forwards (Fig. 2 7-2).
.
(Left Ear) .......................................................................................................... Myotis
68
2’
Ears and tragus not as above.
................................................................................................................................... 3
3 (2’) Eight upper teeth. D 2123/3123=34.
................................................................................................................................... 4
3’
Seven upper teeth. D 2113/3123=32.
................................................................................................................................... 5
4 (3)
Externally similar to Myotis, but ears more rounded. Tragus shorter and not tapered,
sometimes quite broad (Fig. 2 7-2). First premolar usually displaced inwards.
.
.............................................. Pipistrellus
(Left ear)
(First upper molar)
4’
Small pink thumb and foot pads. Second upper incisor displaced outwards from tooth
row. FA about 30 mm.
................................................................................................................. Glischropus
Glischropus tylopus
Thick-thumbed Pipistrelle
(FA 28-30.2, T 28-37.2, HF 4.8-7.9, E 7.4-11.5, Tragus 6.5, W 3.2-5.5 g, D 2123/3123=34,
M 1+0+0=2. Skull: gl 11.0-12.0)
Upperparts dark brown, underparts paler buffy brown. Short broad face with rounded head.
Very similar to Pipistrellus, but base of thumb and sole of foot have thickened unpigmented
whitish or pink pads.
(Left ear)
69
(pink thumb and foot pad)
(Second upper incisor)
5 (3’) Body and skull extremely flattened. Large, dark brown disk-shaped thumb and foot
pads.
(Left ear)
(dark brown thumb and foot pad)
................................................................................................................. Tylonycteris
5’
Ears fairly short and rounded. Tragus slightly hatchet-shaped. First incisor very large
and conical about half the size of the canine. Second incisor small.
(Left ear and tragus)
(First upper incisor)
............................................................................................................. Hesperoptenus
5”
Externally similar to Pipistrellus. Fifth finger very short. First upper incisors long
and narrow with two cusps.
(First upper incissor)
........................................................................................................................ Philetor
Philetor brachypterus
Narrow-winged Brown Bat
(FA 30-36, T 30-38, E 13-16, W 8-13 g, D 2113/3123=32) External genitalia quite elaborate.
Only one upper premolar.
70
Pipistrellus
Glischropus
Hesperoptenus
Philetor
Scotophilus
Fig. 10 Upper left toothrows of Vespertilioninae.
Pipistrellus
: only two premolars, of which the anterior is usually displaced inwards.
Glischropus : second upper incisor displaced outwards from tooth row.
Hesperoptenus : first incisor very large and conical about half the size of the canine.
Philetor
: first incisors long and narrow with two cusps.
Scotophilus
: only one pair of incisors that are large and well developed.
Genus Myotis
Ears moderately long and triangular at the tip. Tragus tapered, bluntly pointed and bent slightly
forwards. D 212-33/312-33=34-38.
Key to species
1
Small feet with the wing membrane usually attached at the base of the toes.
Subgenus Selysius ................................................................................................... 2
(note: Myotis ridleyi is included in this Subgenus because of it has small feet, but the wings
attached at the side of foot)
1’
71
Large feet with the wing attached at the side of the foot.
Subgenus Leuconoe ................................................................................................ 6
Interfemoral membrane
Wing
Selysius
Myotis muricola
Myotis ater
Myotis montivagus
Myotis siligorensis
Myotis ridleyi
Leuconoe
Myotis hasseltii
Myotis adversus
Myotis macrotarsus
Myotis horsfieldii
Fig. 11 Hindfoot and associated membrane.
Subgenus Selysims : wing attached at base of toes.
Subgenus Leuconoe : wings attached at ankle or at side of foot.
Note: Myotis ridleyi is included in the subgenus Selysius, but the wings attached at the side of
foot.
Myotis adversus
Myotis horsfiedii
Myotis siligorensis
Myotis hasseltii
Myotis ater
Myotis montivagus
Myotis ridleyi
Myotis muricola
Fig. 12 Upper left toothrows of Myotis Bats, Myotis.
Myotis adversus, Myotis horsfleldii, Myolis siligorensis: second premolars not displaced
inwards.
Myolis hasseltii, Myotis ater, Myotis montivagus: the first and third premolars are in contact or
nearly so.
Myotis ridleyi: only two pair of premolars.
Myotis muricola: second premolar small and slightly intruded from the tooth row.
72
Subgenus Selysius
2 (1)
Forearm longer than 40 mm.
................................................................................................................................... 3
2’
Forearm less than 40 mm.
................................................................................................................................... 4
3 (2)
Fur dark often with golden-brown patch in the center of belly. Upper and lower second
premolars very small and displaced inwards so that the first and third premolars are
touching. Third upper premolar as long as canine.
Myotis ater
Black Myotis
(FA 40-43, T 37-41, HF 8-9, E 13-15, W 6.5-8 g, D 2133/3133=38. Skull: cbl 14.5, mt 5.5-5.8,
m-m 6.3-6.5)
3’
Dark blackish brown with brown tips to the hairs.
Myotis montivagus
Large Brown Myotis
(FA 42-45.3, T 43-48.4, HF 7-9, E 13.7-16, Tragus 6.5-7.5, W 9-14 g, D 2133/3133=38.
cbl 16.4-16.9, mt 6.7-7.0, m-m 7.4-7.7)
4 (2’) Only two pair of upper and lower premolars. Short wings but relatively heavy body,
4-6 g. Forearm 26.8-32 mm.
Myotis ridleyi
Ridley’s Myotis
(FA 26.8-32, T 30-36, HF 5.1-5.9, E 11.2-11.5, W 4-6 g, D 2123/3123=34. M 1+0+0=2)
A small dark bat with short wings but a relatively heavy body and a typical Myotis ear and tragus
shape. Feet small with wing membrane attached to side of foot. Differs from other Myotis in
having only two upper and lower premolars.
4’
Three pair of premolars.
................................................................................................................................... 5
5 (4”) Ears reach post tip of nose when folded forward. Reddish brown fur. Relatively long
forearm but otherwise very small and light, 2-3 g. Forearm 30-33.5 mm.
Myotis siligorensis
Small-toothed Myotis
(FA 30-33.5, W 2.3-2.6 g, D 2133/3133=38. Skull: condylocanine length (ccl) 10.4, mt 4.4)
73
5’
Upper canine much longer than third premolar. Second premolar usually not especially
reduced. Forearm 33-37 mm.
Myotis muricola
Whiskered Myotis
(FA 33-37, T 34.2-42, HF 6.1-7.1, E 9.1-14.5, Tragus 7.8-8.5, W 3.5-7,4 g, D 2133/3133=38,
M 1+0+0=2. Skull: gl 12, condylocanine length (ccl) 12.0-12.5)
Upperparts brown to gray with dark bases. Underparts with broad buffy-white to pale gray tips.
Feet small, with wing membrane attached at base of toes. Middle upper premolar small and
slightly intruded from the tooth row. Upper canine much longer than posterior upper premolar.
Subgenus Leuconoe
6 (1)
Feet very large. White and pale gray fur. Forearm 45-49 mm.
Myotis macrotarsus
Pallid Large-footed Myotis
(FA 45-49, T 42-52, HF 12-14, E 18-20, W 11-16 g, D 2133/3133=38. Skull: gl 19)
6’
Dark brown to dark gray. Forearm much less than 40 mm.
................................................................................................................................... 7
7 (6’) Ear large. Feet large. NVing attached to side of foot at least I mm from base of toes.
Forearm 35-38.8 mm.
Myotis horsfieldii
Horsfield’s Myotis
(FA 35-38.8, T 37-40.4, HF 9.8-1 1, E 13.5-17, W 5-8 g, D 2133/3133=38. Skull: cbl 14.3-14.7,
iob 3.5-3.6, mt 5.8-6.0)
7’
Wing attached at ankle.
................................................................................................................................... 8
8 (7’) Second upper premotar displaced inwards, so first and third premolar are touching.
Forearm about 37 mm.
Myotis hasseltii
Hasselt’s Large-footed Myotis
(FA 37, D 2133/3133=38. Skull: cbl 14.1-14.6, iob 3.9-4.5, mt 5.6-8.0)
8’
Second premotar not displaced inwards. Forearm 39-40 mm.
Myotis adversus
Grey Large-footed Myotis
(FA 39-40, T 75, E 15, D 2133/3133=38. Skull: gl 18, cbl 14.9-15.2, iob 3.9-4.3, mt 5.9-6.0)
74
Genus Pipistrellus
Externally similar to Myotis, but ears generally more rounded, tragus shorter and not tapered.
D 2123/3123=34.
Key to species
1
Wings whitish. Ears large with slightly hatchet-shaped tragus. Forearm about 30 mm.
Pipistrellus vordermanni
White-winged Pipistrelle
(FA 33-40, T 33-75, E 15, W 6 g, D 2123/3123=34. Skull: cbl 11.1, condylocanine length (ccl)
10.8, iob 3.5, mt 3.8)
1’
Dark wings.
................................................................................................................................... 2
2 (1’) Fur long and thick with black bases and red-orange tips. Ears and tragus dark brown
with buff rims. Ear moderately large with distinct lobe at base near eye.
Pipistrellus cuprosus
Coppery Pipistrelle
(FA 34.5-36.5, T 38-39, E 12.5, W 5.3-5.6 g, D 2123/3123=34. Skull: cbl 12.9-13.0, mt 4.8-4.9)
2’
Fur dark brown to reddish brown.
................................................................................................................................... 3
3 (2’) Tragus narrow and long but not tapered with rounded tip.
(Left ear with tragus)................................................................................................. 4
75
3’
Tragus broad.
(Left ear with tragus)................................................................................................. 5
4 (3)
Forearm 34-36 mm. First upper premotar small and slightly displaced inwards.
Pipistrellus javanicus
Javan Pipistrelle
(FA 33-36, T 34-41, HF 7.2, E 9.8-12.5, Tragus 5.6-6.9, W 3.8-5 g, D 2123/3123=34. Skull:
cbl 12.4, mt 4.6)
4’
Forearm 29-32.1 mm. Tail longer than 95% of HB (Head and Body length)
Pipistrellus tenuis
Least Pipistrelle
(FA 29-32.1, T 32-37.2, HF 5.2-5.4, E 8-11, Tragus 4.2, W 4.2-6 g, D 2123/3123=34. Skull:
gl 10-12, cbl 11.1, mt 4.0-4.2)
Upperparts uniformly dark brown, underparts similar or slightly paler. Second upper premolar
slightly displaced inwards.
5 (3’) Fifth finger very short, not much longer than metacarpals of third and fourth fingers.
Muzzle broad and fairly heavy. Fur short. Forearm 38-43 mm.
(Fifth finger)
Pipistrellus stenopterus
5’
(Left upper teeth)
Narrow-winged Pipistrelle
Fifth finger is longer than metacarpal of fourth finger.
................................................................................................................................... 6
6 (5’’) Forearm more than 40 mm. Fur very long and woolly. Muzzle long and narrow.
Pipistrellus petersi
Woolly Pipistrelle
(FA 40-42, T 39-4 1, E 13-15.5, W 6-7 g, D 2123/3123=34. Skull: cbl 15.3, mt 5.6)
76
6’’
Forearm less than 40 mm.
................................................................................................................................... 7
7 (6’) Fur uniform dark brown. Relatively large. Forearm about 38 mm.
Pipistrellus ceylonicus
Dark Brown Pipistrelle
(FA 38, D 2123/3123=34. Skull: cbl 14.1-14.9, mt 5.4-5.9)
7’
Fur red-brown with blackish bases. Tragus short. First premolar small. Forearm
35-38 mm.
Pipistrellus kitcheneri
Red-brown Pipistrelle
(FA 35-38, T 38, D 2123/3123=34. Skull: cbl 13.1-13.6, mt 4.6-4.9)
Genus Hesperoptenus
Ears fairly short and rounded, tragus slightly hatchet-shaped. Only one upper premolar, distinctive
upper incisors, first incisor very large and conical about half the size of the canine, second
incisor small. D 2113/3123=32.
(Left upper teeth)
(First incisor and canine, second incisor not visible from outside)
Key to species
1
Second upper incisor small and slightly displaced inwards. First upper incisor large
and conical. Forearm 38-41 mm.
Hesperoptenus doriae
False Serotine
(FA 38-41, T 40, E 15, D 2113/3123=32. Skull: gl 15, cbl 13.3-13.6, mt 4.9-5.0)
77
1’
Second upper incisor behind first incisor.
................................................................................................................................... 2
2 (1’) Very small. Dark reddish brown to brown. Base of thumb and sole of foot have
thickened dark brown pads.
Hesperoptenus blanfordi
Least False Serotine
(FA 24-26.5, T 27-30, E 11, W 6.1-6.4 g, D 2113/3123=32)
2’
Forearm 50-53 mm. First upper incisor large and conical, touching the canine.
Hesperoptenus tomesi
Tome’s False Serotine
(FA 50-53.2, T 49-539 HF 12.9, E 17-18, Tragus 7.9, W 30-32 g, D 2113/3123=32. Skull:
cbl 20.4, iob 5.8, mt 8.5)
(Left ear and tragus)
(First upper incisor and canine)
Genus Tylonycteris
Body and skull extremely flattened, base of thumb and foot bears large dark brown disk-shaped
pads. D 2113/3123=32.
(Left ear)
(dark brown thumb and foot pad)
78
Key to species
1
Fur shiny dark brown, very smooth and sleek. Forearm 26-30 mm.
Tylonycteris robustula
Greater Bamboo Bat
(FA 26-30, T 29-36, HF 5.8-6.3, E 8.8-12.1, Tragus 2.3, W 6.5-8.5 g, D 2113/3123=32.
Skull: gl 11.8, condylocanine length (ccl) 12.4-12.7, mt 4.3-4.5)
Upperparts dark brown to dark grayish brown.
1’
Fur fluffier and redder. Underparts slightly paler and strongly tinged orange.
Forearm 24-28.5 mm.
Tylonycteris pachypus
Lesser Bamboo Bat
(FA 24-28.59 T 27-32.5, HF 5.8, E 8-9, W 2.7-5 g, D 2113/3123=32. Skull: gl 11.6,
condylocanine length (ccl) 10.4-10.6, mt 3.1-3.5)
Fur rather short and fluffy.
Subfamily Murininae
Key to genera
1
Nostrils expanded into short tubes that protrude either side of the muzzle. First upper
premolar large, similar in size to the second premolar. Upper molars with well
developed W-shaped cusps.
......................................................................................................................... Murina
1’
Similar to Murina with tubular nostrils and the same dental formula. The third upper
molar reduced to a tiny flake, sometimes lacking. Anterior upper molars with reduced
cusps except the central one. Upperparts bright orange with dark bases.
........................................................................................................... Harpiocephalus
Harpiocephalus harpia
Hairy-winged Bat
(FA 40-54, T 37-55, D 2123/3123=34. Skull: gl 22.1, mt 6.6, c-c 6.0, m-m 7.1)
Genus Murina
Nostrils extended into short tubes, which protrude either side of the muzzle. Ears rounded, not
funnel shaped, tragus long and pointed. D 2123/3123=34.
79
Murininae
mouth
nostlis
Common type
Murininae
(under view)
Key to species
1
Moderately large. Upperparts pale orange mixed with gray or gray-brown. Underparts
paler and grayer. Tragus long, pointed and white. Tail membrane thinly covered in
long reddish hairs.
Murina cyclotis
Orange Tube-nosed Bat
(FA 34-41, T 38-50, E 13-16, W 6.5-12 g, D 2123=3123=34. Skull: cbl 16.8-16.9, mt 5.9,
c-c 4.7)
1’
Color of upperparts differ to above.
................................................................................................................................... 2
2 (1’) Small. Upperparts buffy brown to gray brown. Fur without shiny tips. Underparts
grayish white.
Murina suilla
Lesser Tube-nosed Bat
(FA 28-31, T 26-35, HF 7.5, E 10.5-13, W 3-3.5 g, D 2123/3123=34. Skull: gl 15, cbl 12.6-13.3,
mt 4.7-4.9, c-c 3.6-3.7)
2’
Fur of upperparts dark brown bases with orange-brown tips. Underparts dark brown.
Murina aenea
Bronzed Tube-nosed Bat
(FA 35-38, T 35-41, E 13.5-15.5, W 6-8.5 g, D 2123/3123=34. Skull: cbl 15.3-16.1, mt 5.7-6.0,
c-c 4.7-4.8)
2”
Upperparts dark brown bases with shiny yellow or golden brown tips. Underparts
whitish.
Murina rozendaali
Gilded Tube-nosed Bat
(FA 31.5-33.5, T 35-41, E 13-15, W 4.3-4.8 gg D 2123/3123=34. Skull: cbl 14.1-14.6, mt 5.25.6, c-c 3.8-4.1)
80
Subfamily Kerivoulinae
Key to genera
1
Ears funnel-like with a large flap on the outside. Tragus long, narrow and pointed,
Fur long and woolly, often covering much of the face.
..................................................................................................................... Kerivoula
1’
Very similar to Kerivoula, but tragus white with distinct notch in its posterior margin
near the base. Fur banded, often with pale tips. Upper canines have longitudinal grooves
on the outer faces.
.................................................................................................................... Phoniscus
Genus Kerivoula
Ears funnel-like with a large flap on the outside, tragus long, narrow and pointed. Fur long and
woolly, often covering much of the face. D 2133/3133=38.
Key to species
1
Wing membranes and ears pale brown, almost translucent. Ears very large. Upperparts
pale orange-brown with paler gray bases. Underparts grayish white.
Kerivoula pellucida
Clear-winged Woolly Bat
(FA 29.5-32, T 39-53, E 14.5-17, W 3.5-4.8 g, D 2133/3133=38)
1’
Wing membranes and ears not translucent.
................................................................................................................................... 2
2 (1’) Larger species. Forearm length over 38 mm. Upperparts brown to buffy-brown,
sometimes with buff near the wings. Underparts paler.
Kerivoula papillosa
Papillose Woolly Bat
(FA 38-49, T 48.5-56, HF 8.1-9.1, E 14-17, W 6-13 g, D 2133/3133=38)
2’
Smaller species, Forearm length less than 35 mm.
................................................................................................................................... 3
3 (2’) Ears relatively small. Upperparts orange-brown, underparts paler.
................................................................................................................................... 4
3’
Ears moderately large. Underparts grayish.
................................................................................................................................... 5
81
4 (3)
Upperparts orange-brown with dark bases. Underparts paler. Premolar small and
rounded. Forearm length 26.5-31.3 mm. Weight 2.6-4.2 g. Skull (Condylobasal length)
11.1-11 .8 mm.
Kerivoula intermedia
Small Woolly Bat
(FA 26.5-31.3, T 32-43.2, HF 6.3-7.2, E 9-11.5, Tragus 8.1-9.6, W 2.6-4.5 g, D 2133/3
133=38. Skull: cbl 11.1-11.8, mt 4.6-5.0)
Ears relatively small.
4’
Upperparts orange-brown with brown bases to the fur. Underparts slightly paler.
Forearm length 25-29.5 mm. Weight 1.9-2.3g. Skull (Condylobasal length)
10.0-11.1 mm.
Kerivoula minuta
Least Woolly Bat
(FA 25-29.5, T 36, HF 5.5-5.7, E 9.8-10, W 1.9-2.3 g, D 2133/3 133=38. Skull: cbl 10.0- 11.1,
mt 4.1-4.6)
Smallest Bornean bat. Ear small and rounded. Very similar to Kerivoula intermedia, but smaller
skull and lighter body.
5 (3’) Upperparts gray-brown with gray bases to the fur. Underparts paler and grayer. Forearm
length 32-34 mm.
Kerivoula hardwickii
Hardwicke’s Woolly Bat
(FA 31-34, T 38-479 E 12-14, W 3.5- 5 g, D 2133/3133=38)
5’
Upperparts brown with dark gray bases. Underparts grayish white. Anterior two upper
and lower premolars elongate and oval in cross-section.
Kerivoula whiteheadi
Whitehead’s Woolly Bat
(FA 28-29.4, T 22-3 1, W 3-6.2 g, D 2133/3133=38)
Genus Phoniscus
Very similar to Kerivoula, but tragus unpigmented (white) with a distinct notch in its posterior
margin near the base. Fur banded. D 2133/3133=38.
Key to species
1
Fur grayish black-hairs banded with black and white, sometimes with a shiny white
tip. Short shiny yellow hairs along the forearm and fingers.
Phoniscus jagorii
Frosted Groove-toothed Bat
(FA 37-39, D 2133/3133=38. Skull: cbl 15.3-15.9, mt 6.7-7.1)
82
1’
Upperparts overall golden brown and black, hairs banded with gray and brown bases,
black centers and orange-brown or buff tips. Underparts paler and grayer.
Phoniscus atrox
Gilded Groove-toothed Bat
(FA 31-33, T 39-40, HF 6.3, E 12.5-14, Tragus 8.9, W 2.6-5 gg D 2133/3133=38. Skull: cbl
12.9-13.2, mt 5.6-5.7)
Subfamily Miniopterinae
The subfamily contains the genus Miniopterus
Genus Miniopterus
Distinctive wing shape, especially the third digit, which has a short first phalanx and very long
terminal phalanx.
Key to species
1
Larger species. Forearm length more than 44 mm.
................................................................................................................................... 2
1’
Smaller species. Forearm less than 43 mm.
................................................................................................................................... 3
2 (1)
Head wider, with especially wide palate. Forearm length 47-52 mm. Skull
(Condylobasal length) 15.8-16.8 mm.
Miniopterus magnaterLarge Bent-winged Bat
(FA 47-52, T 54-64.2, HF 9.2-9.4, E 11.4-13.8, Tragus 4.5-6.5, W 9-16 g, D 2123/3123 =34>
Skull: cbl 15.8-16.8, m-m 7.4-8.0)
Largest of the Miniopterus, with and especially wide palate. Head is wider than the Common
Bent-winged Bat. Almost black color.
2’
Forearm length 44-49 mm. Skull (Condylobasal length) 15.1-16 mm.
Miniopterus schreibersi
Common Bent-winged Bat
(FA 44-49, T 51.6-58.5, HF 7.9-9.5, E 9.9-11.9, Tragus 4.6-5.8, W 8-12.5 g, D 2123/3123=34.
Skull: gl 16, cbl 15.1-16.0, m-m 6.6-7.0)
Forearm length slightly overlaps the range of the Large Bent-winged Bat, but it is usually
shorter, and the body and skull are smaller and narrower.
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3 (1’) Forearm length 27.8-39 mm. Skull (Condylobasal length) 11.8-12.6 mm. Fur usually
very dark or black sometimes with reddish patches.
Miniopterus australis
Lesser Bent-winged Bat
(FA 27.8-39.3, T 28.6-42.2, HF 6.1-7.6, E 9.4-9.7, Tragus 3.4-5.9, W 4.0-7 g, D 2123/3123=34.
Skull: gl 14-16, cbl 11.8-12.6)
Smallest of the bent-winged bats.
3’
Forearm length 41-42 mm. Skull (Condylobasal length) 13.8-14.5 mm. Fur tends to
be more gray-brown and less blackish dm the Miniopterus australis.
Miniopterus medius
Medium Bent-winged Bat
(FA 41-42, T 47.4, HF 7.7, E 9.5, Tragus 5, W 7.3-8.3 g, D 2123/3123=34. Skull: cbl 13.8-14.5)
3’’
Forearm length 40-43 mm. Skull (Condylobasal length) 12.8-13.3 mm. Relatively
short skull and long forearm.
Miniopterus pusillus
Small Bent-winged Bat
(FA 40-43, D 2133/3123=34. Skull: cbl 12.8-13.3)
Family Molossidae
Free-tailed Bats
Medium small to large bats distinguished by their thick 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, with fleshy ears.
Key to genera
1
Large and naked with only scattered hairs on the skin. Ears separate.D 1113/2123=28
................................................................................................................. Cheiromeles
1’
Fur thick and short. Ears joined across top of head by narrow flap of skin.
....................................................................................................................... Tadarida
Genus Cheiromeles
Large and naked with only scattered hairs on the skin. Ears separate. D 1113/2123=28.
Cheiromeles torquatus
Naked Bat
(FA 74-83.4, T 50-66.7, HF 24.8-27.5, E 26.3-31.4, W 145-182.5 g. D 11 13/2123=28.
Skull: gl 40)
Bare skin of body dark gray.
84
Genus Tadarida
Fur thick and short. D 111-23/2123=28-30.
Key to species
1
Upperparts and underparts uniform dark brown to reddish brown, crown of head
nearly naked. Only one pair of premolars. Upper incisors long and well-developed.
D 1113/2123=28.
........................................................................................................... Subgenus Mops
Tadarida (Mops) mops
Free-tailed Bat
(FA 41.6-46, T 37-37.8, HF 11.5-12.3, E 19.1-20.7, W 16.1-21.8 g, D 1113/2123=28. Skull:
cbl 19.1-19.4, mt 7.0-7.4, c-c 5.2-5.7)
1’
Upperparts dark brown, underparts paler with gray tips to the fur. Two upper premolars,
the anterior quite small. D 1123/2123=30.
................................................................................................ Subgenus Chaerephon
Tadarida (Chaerephon) plicata
Wrinkle-lipped Bat
(FA 39.9-43.5, T 30.2-35.9, HF 8.3-8.9, E 16-18.1, W 10.5-18 g, D 1123/2123=30. Skull: gl 19,
cbl 16.5-17.7, mt 6.5-6.8, c-c 4.4-5.0)
Left upper teeth (Tadarida mops)
(Tadarida plicata: two upper premolars, the
anterior quite small)
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BBEC Publication No. 7
I D E N T I F I C AT I O N K E Y S T O T H E M A M M A L S O F B O R N E O
This publication is one of the results born by the “Bornean Biodiversity and Ecosystems
Conservation (BBEC) Programme in Sabah, Malaysia.” The BBEC Programme is a joint
endeavor to integrate the various efforts and to enhance comprehensive and sustainable
conservation approaches of biodiversity and ecosystems in the state of Sabah, Malaysia by
Sabah State Government Agencies, Universiti Malaysia Sabah, and NGOs. The activities
under the BBEC Programme have been implemented with the cooperation of technical and
financial supports by the Government of Japan through its agency; Japan International
Cooperation Agency (JICA) since February 2002 until January 2007, for a period of 5 years.
URL: http://www.bbec.sabah.gov.my
Japan International Cooperation Agency
Sabah State
Government Agencies
Universiti Malaysia Sabah
JAPAN
Official Development Assistance