INTRODUCTION
Of the 73 species of Rhinolophus cur-
rently described (Cotterill, 2002; Fahr et al.,
2002; Csorba et al., 2003), 30 are known
from mainland southern and South-East
Asia (Bates et al., 2004). Of these, 17 are
recorded from Thailand, or 18 if R. siamen-
sis is treated as a distinct taxon from R. ma-
crotis (Simmons, 2005). Csorba et al.
(2003) included 13 species of Rhinolophus
from Vietnam. Borissenko and Kruskop
(2003) listed 17, of which one, R. acumina-
tus, was the first country record. Five others
were based on provisional or doubtful
records and one, R. sinicus, which was 
included by Csorba et al. (2003), was omit-
ted. Of the others, Borissenko and Kruskop
(2003) included R. cf. rouxi, on the basis of
specimens from Phong Nha-Ke Bang Na-
tional Park but the identification was ?pro-
visional? and they stated that specimens
may prove to represent another taxon. The
inclusion of R. subbadius was based on
Huynh et al. (1994) and Corbet and Hill
(1992), which in turn was based on Osgood
(1932). However, Borissenko and Kruskop
(2003) and Csorba et al. (2003) suggest that
both records may represent R. pusillus. Two
species, R. yunanensis and R. shameli, are
included on the basis of the unpublished re-
port entitled ?Preliminary report on the bats
Acta Chiropterologica, 8(1): 83?93, 2006
PL ISSN 1508-1109 ? Museum and Institute of Zoology PAS
New records of Microchiroptera (Rhinolophidae and Kerivoulinae)
from Vietnam and Thailand
VU DINH THONG1, SARA BUMRUNGSRI2, DAVID L. HARRISON3, MALCOLM J. PEARCH3,
KRISTOFER M. HELGEN4, 5, and PAUL J. J. BATES3, 6
1Institute for Ecology and Biological Resources (IEBR), Vietnamese Academy of Science and Technology,
18 Hoang Quoc Viet Road, Cau Giay, Hanoi, Vietnam
2Department of Biology, Faculty of Science, Prince of Songkla University Hat Yai, Songkla, Thailand 90112
3Harrison Institute, Centre for Systematics and Biodiversity Research, Bowerwood House, St. Botolph's Road,
Sevenoaks, Kent, TN13 3AQ, Great Britain
4School of Earth and Environmental Sciences, University of Adelaide, Adelaide, SA 5005, Australia
5South Australian Museum, North Terrace, Adelaide, SA 5000, Australia
6Corresponding author: E-mail: hzm@btinternet.com
The diversity of Rhinolophidae in Thailand and Vietnam is briefly discussed and the taxonomy of Asian
Kerivoulinae, with particular reference to the genus Phoniscus, is reviewed. Four new country records are
included: Rhinolophus shameli and Kerivoula kachinensis from Vietnam and Phoniscus jagorii from Vietnam
and Thailand. A second record of Phoniscus atrox from Thailand is also discussed. 
Key words: Rhinolophus shameli, Phoniscus jagorii, P. atrox, Kerivoula kachinensis, Thailand, Vietnam, 
systematics, distribution, ecology
of Pu Mat Nature Reserve. Fauna and Flora
International, Indochina Programme, Ha
Noi? of B. Hayes and T. Howard (in litt.).
The status of such records, which have been
included in the ?grey literature? but not in
peer-reviewed publications, is problemati-
cal. It is therefore of value to include here
the first published record of R. shameli
based on specimens from the central high-
lands.
The taxonomy of the Kerivoulinae in
Asia is also problematical. Hill (1965) not-
ed that bats of the genera Kerivoula (Gray,
1842) and Phoniscus (Miller, 1905) are rel-
atively uncommon in reference collections.
To some extent, this is still the case today,
although the advent of the harp trap has 
led to a recent and dramatic increase in 
the frequency of their collection in the field
(Kingston et al., 2003; Struebig et al.,
2005). Hill (1965) also noted that the taxo-
nomic status of Phoniscus was uncertain.
Some bat taxonomists treated Phoniscus as
congeneric with Kerivoula (Ryan, 1965;
Koopman, 1993, 1994). Others considered
the differentiating characters to be incon-
clusive but maintained them as separate
genera (Troughton, 1929) or as subgenera
(Laurie and Hill, 1954) whilst others con-
sidered Phoniscus to be clearly separate
from Kerivoula (Le Souef and Burrell,
1926; Miller, 1931; Tate, 1941; Hill, 1965;
Medway, 1969; Lekagul and McNeely,
1988; Corbet and Hill, 1992; Flannery,
1995; Simmons, 2005). 
A detailed review of the characters used
to distinguish Phoniscus from Kerivoula is
included in Hill (1965) and summarized in
Corbet and Hill (1992). It includes for
Phoniscus: the tragus with a deep notch
near the base (it is shallow in Kerivoula ?
Fig. 1); upper canine (C1) with longitudinal,
lateral grooves on its outer face (it is
smoothly rounded in Kerivoula ? Fig. 2);
second upper incisor (I3) much reduced
with its greatest (antero-posterior) diameter
84 V. D. Thong, S. Bumrungsri, D. L. Harrison, M. J. Pearch, K. M. Helgen, and P. J. J. Bates
about one half, and its height one half or
less, that of the first (I2) (in Kerivoula the
greatest diameter is equal or subequal to
that of I2 and the height one half or more ?
Fig. 3). In addition to the characters of Hill
(1965), it should be noted that in Phoniscus,
the second upper incisor (I3) is also marked-
ly reduced in size (both area and height) in
comparison to C1 (in Kerivoula, I2 is not so
reduced relative to C1 ? Figs. 2 and 3).
Tate (1941) included six species in Phonis-
cus (P. rapax, P. atrox, P. javanus, P. agnel-
la, P. myrella and P. papuensis). Hill (1965)
recognized four species (P. atrox, P. papu-
ensis, P. jagorii and P. aerosa); he treated 
P. rapax and P. javanus as junior synonyms
of P. jagorii and transferred P. agnella and
P. myrella to Kerivoula. This arrangement
has been followed by Corbet and Hill
(1992) and Simmons (2005). 
The current paper includes further
records of Phoniscus from mainland South-
East Asia. Currently, the only records 
from this area (here defined as Myanmar,
FIG. 1. Tragus of the left ear of Phoniscus (left) and
Kerivoula (right); P. jagorii, XS.53, Vietnam and 
K. kachinensis, CMR-28, Vietnam. Scale = 2 mm
Thailand, Lao PDR, Cambodia and Thai-
land) are a single specimen of P. atrox
recorded from southern Thailand by Kloss
(1916) and P. jagorii, and possibly P. atrox,
reported from Lao PDR (Francis et al.,
1999). In addition to the data on Phoniscus,
this paper includes a significant new record
of Kerivoula kachinensis, which is based on
three specimens from Vietnam. This is only
the second record of this recently described
species (Bates et al., 2004) and the first re-
cord of its occurrence outside Myanmar. 
This paper represents one of the first
outputs of a programme of studies, co-ordi-
nated by the Harrison Institute and support-
ed by the UK Government?s Darwin Initia-
tive programme, to promote collaborative
research between bat taxonomists from dif-
ferent countries and institutions in mainland
South-East Asia. 
MATERIALS AND METHODS
Study Areas
In Vietnam, a number of field surveys was con-
ducted at Xuan Son National Park between November
2003 and September 2004. The National Park is situ-
ated at Thanh Son District, Phu Tho Province, about
75 km north-west of Hanoi (21?03?? 21?12?N,
104?51??105?01?E). The elevation ranges from ca.
200 m to 1,386 m at the highest point, which is the
mountain peak of Dinh Voi (Anon, 1995b). Limestone
karst, which contains a large number of caves, covers
about one third of the Park?s area. Some of the caves
contain river systems and many of them are diurnal
roosting sites for bats that forage within the Park
and/or its surrounding areas. The vegetation types
comprise lowland and lower montane evergreen for-
est. Primary forest still covers the core zone of the
Park, where there is little serious disturbance from lo-
cal communities (Anon, 1995b). Prior to this study,
no comprehensive bat surveys had been undertaken
(Anon, 1995b; Thong, 2003).
Chu Mom Ray National Park is situated in West-
ern Kon Tum Province in the highlands of cen-
tral Vietnam (14?21??14?36?N, 107?29??107?47?E). 
Elevations range from around 200 m to the high-
est mountain peak, Chu Mom Ray, at 1,773 m (Anon,
1995a). The Park has several vegetation types 
including lowland evergreen forest, semi-evergreen
forest, and a large area of bamboo. The buffer zone of
the park is covered by secondary forest with various
vegetation types including shrubs and grasslands.
There are also two medium-sized rivers, Dak Hodrai
and Krong Po Ko, which are tributaries of the Me-
kong River system.
Huai-Thab Tan-Huai Samran Wildlife Sanctuary,
which is located in Surin Province along the Thai-
land-Cambodia border, covers an area of 48,000 ha
and is predominantly a plateau with an elevation of
between 200 to 500 m a.s.l. The vegetation is charac-
terized by dry evergreen forest mixed with dry dipte-
rocarp and deciduous forest. Most of this forest is dis-
turbed. It is less disturbed along the border. The cli-
mate is seasonal, with a rainy season from May to
September and a dry season from November to April. 
Bala Forest covers an area of 17,000 ha and has
an altitudinal range of about 100?963 m a.s.l. The for-
est is ?Malesian type? tropical rain forest (Whitmore,
1984). It is surrounded by fruit orchards and rubber
plantations. Numerous small streams are found in the
narrow valleys. Small patches of peat swamp forest
are also present. 
Specimens and Measurements
All of the recent specimens from Vietnam and
Thailand, which are listed below, were collected in
harp traps. Specimens were measured with digital
calipers, weighed and preserved in 70% ethanol. The
skulls were subsequently extracted, prepared and
measured.
The following external, cranial and dental meas-
urements were taken using digital calipers. HB: head
and body length, from the tip of the snout to the base
of the tail, dorsally; TAIL: tail length, from the tip of
the tail to its base adjacent to the anus; HF: foot
length, from the extremity of the heel behind the os
calcis to the extremity of the longest digit, not includ-
ing the claws; TIBIA: length of tibia, from the knee
joint to the ankle; FA: forearm length, from the ex-
tremity of the elbow to the extremity of the carpus
with the wings folded; E: ear length, from the lower
border of the external auditory meatus to the tip of the
pinna; GTL: greatest length of the skull, the greatest
antero-posterior length of the skull, taken from the
most projecting point at each extremity; SL: skull
length, from the alveolus of the anterior canine to the
most projecting posterior point of the skull; CBL:
condylo-basal length, from an exoccipital condyle to
the alveolus of the anterior incisor; CCL: condylo-
canine length, from an exoccipital condyle to the
alveolus of the anterior canine; ZB: zygomatic
breadth, the greatest width of the skull across the 
New records of Microchiroptera from Vietnam and Thailand 85
(VDT05005/..05006 and ..05008), 1 X
(VDT05007) collected by Vu Dinh Thong
and Pham Duc Tien.
This is the first published record for
Vietnam, although B. Hayes and T. Howard
(in litt.) included it in an unpublished re-
port on Pu Mat Nature Reserve. Previously
known from Myanmar, Thailand, Lao PDR,
Cambodia, and peninsular Malaysia (Sim-
mons, 2005). 
Description and taxonomic notes
Externally, the four specimens from
Vietnam, with a forearm length of 45.8?
47.3 mm (Table 1), are large in comparison
to those included in Csorba et al. (2003),
for which the values are 42.0?46.5 mm.
However, they are comparable in all mor-
phological characters including the con-
necting process of the noseleaf, which is in-
serted into a fissure in the thickened lancet
and covered with short, dense hairs. Skull
length (SL) is 20.4?21.2 mm (Table 2),
which is within the range (19.3?21.6 mm)
given by Csorba et al. (2003). The rostrum
is high with the anterior and median
swellings prominent. The width across the
lateral swellings of the rostrum (RW) is 6.0
mm (6.0?6.1 mm, n = 4). In the smaller, but
morphologically similar R. coelophyllus,
the width is less than 5.5 mm (Csorba et al.,
2003).
Ecological notes and conservation status
All four specimens were captured in 
a harp trap, which was set across a foot-
path next to a narrow stream and a small
cave. The study site was located in the core
zone of the Chu Mom Ray National Park 
in an area of essentially undisturbed, low-
land evergreen forest. Light rain was falling
at the time of capture. Kerivoula kachinen-
sis and Murina tubinaris were also captur-
ed at this site. The conservation status of 
R. shameli is lower risk, near threatened
(Hutson et al., 2001).
86 V. D. Thong, S. Bumrungsri, D. L. Harrison, M. J. Pearch, K. M. Helgen, and P. J. J. Bates
zygomatic arches; BB: breadth of braincase at the
posterior roots of the zygomatic arches; GBB: great-
est width of the braincase; BH: braincase height, tak-
en from the basisphenoid to the highest part of the
skull; RW: rostral width taken across the anterior lat-
eral swellings of the rostrum; PC: post orbital con-
striction; C?M3: maxillary toothrow length, from the
alveolus of the upper canine to the back of the crown
of the third upper molar; M3?M3: palatal width, taken
across the outer borders of the third upper molar, at
the widest part; C?M3: mandibular toothrow length,
from the alveolus of the lower canine to the back 
of the crown of the third lower molar; MDL: man-
dible length, from the most posterior part of the con-
dyle to the most anterior part of the mandible. These
measurements are illustrated in Figs. i?v in Bates and
Harrison (1997). Body mass (MASS) was taken using
a 60 g Pesola scale. Teeth size was measured using 
a graticule in a Leica MZ8 stereo-microscope.
Measurements included in Tables 1 and 2 are of
recent specimens collected in Vietnam and Thailand
and deposited at the Institute for Ecology and Bio-
logical Resources, Hanoi and the Prince of Songkla
University, Hat Yai, respectively. In addition, there is
one Vietnamese specimen of K. kachinensis, which is
deposited at the Field Museum of Natural History,
Chicago.
Echolocation calls of voucher specimens were
recorded with a Tranquillity II (David Bale, UK) bat
detector connected to a Sony Walkman Professional
(WM D6C). Calls were recorded after bats were re-
leased in a large room (6 ? 8 ? 3 m) and allowed to
fly freely. Calls were analysed using sound analysis
software (Bat Sound Pro, Pettersson Electronik AB,
Sweden) on a personal laptop. Call characteristics in-
cluding call types, minimum frequency, most energy
frequency, call duration and number of harmonics
were examined. At least five calls from each individ-
ual were analysed.
SYTEMATIC REVIEW OF SPECIES
Rhinolophus shameli Tate, 1943
Shamel?s horseshoe bat
Rhinolophus shameli Tate, 1943: 3; Koh Chang Is-
land, Thailand
New material, previous records and distri-
bution
Vietnam: Chu Mom Ray National Park,
Kon Tum Province (14?28?N, 107?47?E,
750 m a.s.l.), 9 June, 2005, 3 YY
Phoniscus jagorii (Peters, 1866)
Peter?s trumpet-eared bat
Vespertilio (Kerivoula) jagorii Peters, 1866: 399; 
Samar Island, Philippine Islands
New material, previous records and distri-
bution
Vietnam: Xom Du study site, Xuan Son
National Park, Phu Tho Province (21?06?N,
104?57?E), 24 November 2003, 1 X (XS
53) collected by Vu Dinh Thong and Pham
Duc Tien. Thailand: Tamearn-Thom Castle,
Huai-Thab Tan-Huai Samran Wildlife 
Sanctuary, Surin Province, 400 m a.s.l.,
(14?21?N, 103?50?E), 18 January 2000, 1 Y
(PSU-M 05.10) collected by S. Bum-
rungsri.
These are the first records for Vietnam
and Thailand. Previously known from Lao
PDR, peninsular Malaysia, Borneo, Java,
Bali, Sulawesi, Lesser Sunda Islands and
Samar Island in the Philippines (Simmons,
2005).
Description and taxonomic notes
The recent specimen from Vietnam is
preserved in ethanol and it is difficult to de-
termine the pelage colour with certainty.
However, its hairs appear to have four dis-
tinct colour bands, namely dark brown or
blackish-brown at the base, followed by
buff, then brown, and finally golden or
whitish-yellow tips; the paler tips are more
pronounced on the ventral surface. The
hairs of the wet specimen from Thailand
have become ?foxed? after being stored 
in daylight and have no diagnostic value.
The skulls have a condylo-basal length of
15.4 and 15.6 mm (Table 2). This corre-
sponds well with measurements (in mm) for
P. j. jagorii (15.3) and P. j. javanus (15.3,
15.9) included in Hill (1965). Skull shape is
essentially similar to the description given
in Hill (1965). Both specimens have well-
defined basioccipital pits. These are deeper
than those of P. atrox, which has a smaller
skull, but this character is only useful if 
New records of Microchiroptera from Vietnam and Thailand 87
Sp
eci
es
Se
x
HB
TA
IL
HF
TI
BI
A
FA
E
MA
SS
Rh
ino
lop
hu
s s
ha
me
li
3Y
Y
47
.0?
48
.1
19
.4?
21
.0
9.4
?1
0.2
22
.7?
23
.7
46
.5?
47
.3
19
.6?
20
.7
8.4
?9
.5
47
.5,
 0.
5
20
.4,
 0.
9
9.7
, 0
.4
23
.1,
 0.
5
47
.0,
 0.
5
20
.3,
 0.
7
8.8
, 0
.6
1X
47
.7
22
.4
9.2
21
.9
45
.8
16
.9
8.6
Ph
on
isc
us 
jag
ori
i
1Y
45
.2
36
.5
8.5
18
.0
35
.3
12
.3
?
1X
39
.8
40
.2
9.1
20
.3
38
.0
12
.5
5.8
P. 
atr
ox
1Y
41
.3
29
.5
7.6
16
.3
34
.3
13
.3
5.0
Ke
riv
ou
la 
ka
ch
ine
nsi
s
2Y
Y
47
.0,
 47
.0
53
.0,
 59
.0
8.5
, 1
1.0
22
.7,
 24
.0
40
.4,
 43
.4
11
.0 
(1)
7.5
 (1
)
2X
X
35
.5,
 46
.7
50
.5,
 56
.1
8.5
, 8
.7
22
.8,
 23
.5
41
.6,
 43
.2
10
.0,
 12
.9
7.4
, 8
.3
TA
BL
E
1. 
Six
 ex
ter
na
l m
eas
ure
me
nts
 (i
n m
m)
 an
d t
he
 bo
dy
 m
ass
 (i
n g
) o
f 1
1 s
pe
cim
en
s o
f f
ou
r b
at 
spe
cie
s r
ece
ntl
y c
oll
ect
ed
 in
 V
iet
na
m 
an
d T
ha
ila
nd
, in
clu
din
g
the
 le
ng
th 
of 
he
ad
 an
d b
od
y (
HB
), t
ail
 (T
AI
L)
, fo
ot 
(H
F),
 tib
ia 
(T
IB
), f
ore
arm
 (F
A)
 an
d e
ar 
(E
). F
or 
ma
les
 of
 R
. s
ha
me
li,
the
 ra
ng
e, 
me
an
 an
d s
tan
da
rd 
de
via
tio
n
are
 gi
ve
n. 
In 
K.
 ka
ch
ine
nsi
s, 
sam
ple
 si
zes
 w
he
n d
iff
eri
ng
 fr
om
 n
are
 gi
ve
n i
n p
are
nth
ese
s
material of both species is available for
comparison at the same time. Upper tooth-
row length (C?M3) of 6.7 and 6.8 mm is
comparable to those of 6.7, 6.7 and 7.1 mm
listed by Hill (1965). The first upper incisor
(I2) has a single cusp and is pointed. The
second incisor (I3) is distinctly small and
short (0.5?0.6 mm). The canine (C1) has
two longitudinal, lateral grooves on its out-
er face (Fig. 2). The first upper premolar
(P2) is broader than the second (P3) but
shorter antero-posteriorly (Fig. 3). P3 is ap-
proximately rectangular in shape but with
an antero-internal elongation. In the lower
dentition, the premolars are narrow, espe-
cially the second (P3 ? see Fig. 3).Since so few voucher specimens are
available for study, it is difficult to deter-
mine if the taxa javanus and rapax are valid
subspecies, as provisionally indicated by
Hill (1965), or whether they are simply syn-
onyms of P. j. jagorii. For this reason, the
subspecific status of both of the current
specimens cannot be determined and they
are referred simply to P. jagorii.
Echolocation
Based on the recent Thai material, the
echolocation calls (in kHz) of P. jagorii are
characterized by a broadband FM sweep
with a start frequency of 137?140 (0 =
137.55, SD = 1.66, n = 4) and an end fre-
quency of 82.9 ? 3.45 (77.0?85.7). Peak
frequency is 88 (87.5?88.2). Calls are of
low intensity and short duration (1.41?1.58
ms). When calls are emitted in series, the
interval is about 13 ms. In Malaysia, King-
ston et al. (1999) reported a start frequency
of 169.3 (154.4?184.8, n = 18), and end fre-
quency of 70.4 kHz (61.6?76.0, n = 18), 
a peak frequency of 94.4 (79.2?117.6, n =
18), and a call duration of 2.2 ms (1.6?3.2
ms, n = 18).
Ecological notes and conservation status
In Vietnam, P. jagorii was collected in 
a harp trap set across a narrow path close to
88 V. D. Thong, S. Bumrungsri, D. L. Harrison, M. J. Pearch, K. M. Helgen, and P. J. J. Bates
TA
BL
E
2. 
Ele
ve
n c
ran
ial
 an
d d
en
tal
 m
eas
ure
me
nts
 (i
n m
m)
 of
 11
 sp
eci
me
ns 
of 
fou
r b
at 
spe
cie
s r
ece
ntl
y c
oll
ect
ed
 in
 V
iet
na
m 
an
d T
ha
ila
nd
, in
clu
din
g t
he
 gr
eat
est
len
gth
 of
 sk
ull
 (G
TL
), s
ku
ll l
en
gth
 (S
L)
, c
on
dy
lo-
ba
sal
 le
ng
th 
(C
BL
), c
on
dy
lo-
can
ine
 le
ng
th 
(C
CL
), z
yg
om
ati
c b
rea
dth
 (Z
B)
, b
rea
dth
 of
 br
ain
cas
e (
BB
), p
ost
orb
ita
l
co
nst
ric
tio
n (
PC
), m
ax
illa
ry 
too
thr
ow
 le
ng
th 
(C
?M
3 ),
 pa
lat
al 
bre
ad
th 
(M
3 ?M
3 ),
 m
an
dib
ula
r to
oth
row
 le
ng
th 
(C
?M
3) 
an
d m
an
dib
ula
r le
ng
th 
(M
DL
). I
n R
. s
ha
me
li,,
the
 m
eas
ure
me
nts
 of
 G
TL
an
d C
BL
are
 om
itte
d s
inc
e t
he
 pr
em
ax
illa
e a
re 
fre
qu
en
tly
 da
ma
ge
d o
r m
iss
ing
 an
d t
he
se 
me
asu
rem
en
ts 
are
 co
nse
qu
en
tly
 no
t re
lia
ble
; in
ma
les
 of
 R.
 sh
am
eli
, th
e r
an
ge
, m
ean
 an
d s
tan
da
rd 
de
via
tio
n a
re 
giv
en
. In
 K
. k
ac
hin
en
sis
, s
am
ple
 si
zes
 w
he
n d
iff
eri
ng
 fr
om
 n
are
 gi
ve
n i
n p
are
nth
ese
s
Sp
eci
es
Se
x
GT
L
SL
CB
L
CC
L
ZB
BB
PC
C?
M3
M3
?M
3
C?
M 3
MD
L
Rh
ino
lop
hu
s s
ha
me
li
3Y
Y
?
20
.9?
21
.2
?
18
.3?
18
.8
10
.2?
10
.4
8.7
?8
.9
1.8
?2
.1
8.1
?8
.5
7.4
?7
.9
8.6
?9
.0
14
.2?
14
.5
21
.1,
 0.
2
18
.6,
 0.
2
10
.4,
 0.
2
8.9
, 0
.1
1.9
, 0
.2
8.3
, 0
.2
7.6
, 0
.2
8.8
, 0
.2
14
.4,
 0.
2
1X
?
20
.4
?
18
.0
9.9
8.6
2.1
8.0
7.4
8.3
13
.8
Ph
on
isc
us 
jag
ori
i
1Y
16
.4
16
.3
15
.4
15
.1
9.9
7.7
4.5
6.8
5.7
7.3
12
.0
1X
17
.0
16
.8
15
.6
15
.3
9.6
7.7
4.0
6.7
5.8
7.2
12
.2
P. 
atr
ox
1Y
15
.4
15
.1
14
.0
13
.6
9.0
7.2
3.9
5.9
5.2
6.4
10
.6
Ke
riv
ou
la 
ka
ch
ine
nsi
s
2Y
Y
17
.9 
(1)
16
.9,
 17
.4
15
.2,
 16
.7
15
.5,
 16
.3
10
.1,
 10
.7
7.6
, 8
.2
3.5
, 3
.5
6.8
, 6
.9
6.4
, 6
.4
7.4
, 7
.4
12
.6,
 12
.7
2X
X
17
.5,
 17
.8
16
.8,
 17
.3
16
.4,
 16
.6
15
.6,
 16
.1
10
.3,
 10
.7
7.8
, 7
.9
3.6
, 3
.7
6.6
, 6
.9
6.4
, 6
.4
7.2
, 7
.5
12
.2,
 12
.5
the small village of Xom Lap, which is 
situated in Xuan Son National Park. There
is considerable limestone karst and primary
forest in the vicinity. Murina cyclotis was
collected from the same locality. In Thai-
land, a male specimen was caught in a harp
trap at 19:30 hours on the edge of a moder-
ately disturbed, semi-evergreen forest in 
a flat area with an elevation of 400 m a.s.l.
This forest is on a plateau that connects with
a large forest patch in Cambodia. P. jagorii
was found to be very manoeuvrable, some-
times flying close to the ground. Previously,
specimens from Malaysia were recorded
from the primary forest (Kingston et al.,
2003). Its conservation status is lower risk,
least concern (Hutson et al., 2001).
Phoniscus atrox Miller, 1905
Groove-toothed trumpet-eared bat
Phoniscus atrox Miller, 1905: 230; Vicinity of the
Kateman River, eastern Sumatra
New material, previous records and distri-
bution
Thailand: Wildlife Research Station,
Bala Forest, Hala-Bala Wildlife Sanctu-
ary, Wang District, Narathiwat Province
(05?48?N, 101?50?E), 21 May 2003, 1 Y
(PSU-M 05.11) collected by S. Bumrungsri.
In Thailand, the previous locality record
is Klong Bang Sai, Patiyu District, Chum-
phon Province (10?43?N, 99?20?E ? Kloss,
1916). It is also known from peninsular
Malaysia, Sumatra and Borneo (Simmons,
2005) and possibly Lao PDR (Francis et al.,
1999).
Description and taxonomic notes
In general, the pelage of P. atrox is com-
parable to that of P. jagorii but slightly
paler. Kloss (1916) noted that of his two
Thai specimens, the female was duller and
darker than the male. With a forearm length
of 34.3 mm (Table 1), the current specimen
slightly exceeds in size those (32.3, 32.6
mm) listed in Hill (1965). However, the
condylobasal length of 14.0 mm (Table 2) is
within the range (0 = 13.6, 13.1?14.1 mm,
n = 7) of specimens of P. atrox in The Nat-
ural History Museum, London. The mor-
phology of the skull is similar to that of 
P. jagorii, except that the basioccipital pits
are shallower (see above). In the dentition,
the upper canine is very large, with its outer
side having two deep grooves. The upper
New records of Microchiroptera from Vietnam and Thailand 89
FIG. 2. Lateral view of the left anterior dentition (I2?P4 and I1?P4) of Phoniscus (left) and Kerivoula (right); P. jagorii, XS.53, Vietnam and K. kachinensis, VN014-S411, Vietnam. Scale = 2 mm
premolars are significantly smaller than
those of P. jagorii (Fig. 3). This is evident 
in the first (P2) and especially the second
(P3) premolar. Furthermore, in comparison
to P. jagorii, P3 is essentially round in out-
line, rather than rectangular, and without 
an antero-internal elongation. The second 
lower premolar (P3) is particularly narrow
(Fig. 3).
90 V. D. Thong, S. Bumrungsri, D. L. Harrison, M. J. Pearch, K. M. Helgen, and P. J. J. Bates
FIG. 3. Left maxillary (above) and mandibular 
(below) toothrows of P. jagorii, XS.53 (left), 
P. atrox, PSU-M 05.11 (centre), and K. kachinensis,
CMR-28 (right). (i): I3, much reduced in Phoniscus;
(ii): antero-internal elongation of P3 in P. jagorii; (iii):
P3 larger and rectangular in P. jagorii in comparison
to P. atrox; (iv): P3 narrow in Phoniscus, broader inKerivoula. Scale = 2 mm
Echolocation
Based on recent material from Thailand,
like P. jagorii, the echolocation calls of 
P. atrox are characterized by broad FM
sweeps. The initial frequency (0 ? SD) is
125.53 ? 7.31 kHz (n = 6) and the terminal
frequency is 61.80 ? 3.76 kHz. The mean
peak frequency is 77.37 ? 6.55 kHz (range
64.6?83.5). Generally, energy is distributed
fairly evenly throughout the emitted call.
The calls have low intensity and a relative-
ly short duration of 2.97 ? 0.79 ms. Occa-
sionally, calls are emitted in series with an
interval of 14?18 ms. In Malaysia, King-
ston et al. (1999) reported a start frequency
of 166.1 kHz (145.6?183.2, n = 42), an end
frequency of 60.0 kHz (51.2? 72.0, n = 42),
a peak frequency of 86.9 kHz (62.4?122.4,
n = 42), and a call duration of 2.8 ms
(1.9?3.9, n = 42).
Ecological notes and conservation status
In Bala Forest, Thailand, a single indi-
vidual was captured in the early evening
(19:00 hrs) in disturbed forest interspersed
with orchards and rubber plantations. This
area, at an elevation of 150 m a.s.l., is adja-
cent to a large patch of pristine lowland
evergreen forest. Phoniscus atrox appears
to favour birds? nests, particularly those of
the broad bill (Eurylaimus sp.) as a diurnal
roost. In eastern Sumatra, the holotype and
paratype, both females, were collected on
9th September, 19:03 hrs from an aban-
doned nest of a broad bill in dense forest on
the banks of the Kateman River. Individuals
belonging to this species were also captured
in birds? nests in Malaysia (T. Kingston,
personal comm.). The first record from Ma-
laysia was of a subadult collected at 615 m
a.s.l. from the Ulu Gombak Forest Reserve,
Selangor (Medway, 1969). The first record
from Thailand was an adult male and fe-
male caught in January, 1916 (Kloss, 1916).
Its conservation status is lower risk, least
concern (Hutson et al., 2001).
i i
ii
iii
iviv
Ic 
1L 
New records of Microchiroptera from Vietnam and Thailand 91
Kerivoula kachinensis Bates et al., 2004
Kachin woolly bat
Kerivoula kachinensis (Bates et al., 2004): 220;
Namdee Forest, Bhamo Township, Kachin State,
Myanmar, 24?34?N, 97?08?E
New material, previous records and distri-
bution
Vietnam: Chu Mom Ray National Park,
Kon Tum Province (14?28?N, 107?47?E,
750 m a.s.l.), 28 June, 2005, 1 Y (CMR-28)
and 2 XX (CMR-25, VN014-S411) collect-
ed by Vu Dinh Thong and Pham Duc Tien;
Muong Mo, Lai Chau Province (22?13?N,
102?55?E, 229 m a.s.l.), 14 March, 1929, 
1 Y (FMNH 33209) collected by R. W.
Hendee. These are the first published re-
cords for Vietnam. Previously, K. kachinen-
sis was known only from southern Kachin
State, Myanmar (Bates et al., 2004).
Description and taxonomic notes
With forearm lengths of 40.4?43.4 mm
(see Table 1), the four Vietnamese speci-
mens of K. kachinensis compare favourably
in size to the holotype (41.3 mm ? Bates et
al., 2004). Since the holotype was a wet
specimen, the pelage colour could not be
determined accurately. It is therefore of in-
terest to describe for the first time the
pelage of a prepared skin (VN014-S411).
The hairs on both the dorsal and ventral as-
pects have grey-brown roots, the mid-parts
are pale whitish grey and the tips are buff to
mid-brown; the ventral surface is slightly
paler than the dorsal. Notably, the overall
pelage coloration is darker brown than the
fur of Kerivoula papillosa, with which K.
kachinensis is most likely to be confused
when examined in the field. Other charac-
ters such as the ears, wings and tail are as
described for the Myanmar specimen by
Bates et al. (2004). The skulls of the Viet-
namese specimens, with condylo-canine
lengths of 15.5?16.3 mm (see Table 2), 
likewise agree in size with the holotype
(15.5 mm). Their most obvious feature is
the flattened braincase. This is similar to the
holotype, with a relative height of braincase
(BH/GBB ? 100%) of 66.5% (62.9?69.6%,
SD = 2.8, n = 4) compared to 64.0% in the
holotype and 86.4% (80.6?91.6%, n = 22)
in K. papillosa (Bates et al., 2004). The hol-
otype of K. kachinensis was an old adult
with worn teeth. The current specimens
agree with the description of the dentition
by Bates et al. (2004) and in addition con-
firm that the upper canine has a well-de-
fined cingulum on its internal border and
that the third lower incisor (I3) has a well-defined central cusp and lateral cusps. Addi-
tional measurements (in mm) not included
in Table 2 are BH: 5.4 (5.1?5.6, n = 4);
GBB: 8.2 (8.1?8.4, n = 4). The Muong Mo
specimen, previously included in the Field
Museum collection as K. papillosa, was col-
lected more than seven decades prior to the
description of K. kachinensis from Myan-
mar (Bates et al., 2004) but never received
critical study.
Ecological notes and conservation status
The three recent specimens were col-
lected on 28 June, 2005 in Chu Mom Ray
National Park. For further details of the
study site see Methods and also Ecological
notes for R. shameli. The conservation sta-
tus of K. kachinensis has not been assessed
(Hutson et al., 2001).
ACKNOWLEDGEMENTS
In Vietnam, we would like to thank Mr Pham Duc
Tien of IEBR for his assistance in the field surveys.
We are grateful for the support and encouragement of
the Director of IEBR, Ass. Prof. Dr Le Xuan Canh,
the Vice Directors, Dr Tran Minh Hoi and Ass. Prof.
Dr Ta Huy Thinh, and of the Director of Xuan Son
National Park, Dr Tran Dang Lau. We are also very
grateful to Prof Vu Quang Con (Programme Manag-
er) and all committee members of the Vietnam Na-
tional Flora and Fauna Programme for their financial
support of the field surveys in Xuan Son National
Park. Many thanks to Prof Dr Le Vu Khoi, leader of
the bat and small mammal project, and to members of
the Basic Research Programme in Natural Sciences
for their financial support of a field survey in Chu
Mom Ray National Park. We would also like to thank
Ass. Prof. Tran Hong Viet (Hanoi University of Edu-
cation), Dr Huynh Thi Kim Hoi (Head of Terrestrial
Environment Department, IEBR) and Neil Furey
(FFI-Vietnam Programme) for their valuable encour-
agement and consideration. In Thailand, we are grate-
ful to Ass. Prof. Chutamas Pholpunthin and the staff
and students of the Biology Department of Prince of
Songkla University, especially those involved in the
fieldwork in the Hala-Bala and Huai-Thab Tan-Huai
Samran Wildlife Sanctuaries. We are also most grate-
ful to the directors and staff of these sanctuaries for
their permissions and logistical support and to the
Biodiversity Research and Training Programme,
Thailand for its financial support of the fieldwork. In
the UK, many thanks are due to the Mammal Section
and the Library of The Natural History Museum, Lon-
don for their kind co-operation (especially to Paula
Jenkins); and to Prof. Paul Racey of Aberdeen Uni-
versity for his encouragement. In the US, we wish to
thank Bill Stanley and Rebecca Banasiak of the Field
Museum of Natural History, Chicago and Don Wilson
of the Smithsonian Institution. Finally, we would like
to thank the British Government, which, through the
Darwin Initiative biodiversity programme, has most
generously supported our South-East Asian bat study
programme. 
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New records of Microchiroptera from Vietnam and Thailand 93
Received 16 December 2005, accepted 28 February 2006