Discovery of Rhinolophus beddomei (Chiroptera: Rhinolophidae) from Thailand with a Brief Comparison to Other Related Taxa

Abstract

Rhinolophus beddomei is reported from mainland Southeast Asia for the first time. It was collected in April, 2008, in Ratchaburi Province, Thailand, in a harp trap set over a stream in a dense evergreen forest. Morphometric and acoustic characters of the specimen are given with a brief comparison to other species of the R. trifoliatus group. Currently, it is not clear whether the Thai specimen represents an eastern extension of the Indian Subcontinent population or whether it is a member of an isolated population which is restricted to continental Southeast Asia. It is suggested that a further review, which would involve material from southern and Southeast Asia, together with R. formosae from Taiwan would be of considerable interest.

Full text

Tropical Natural History 10(1): 67-79, April 2010
©2010 by Chulalongkorn University
Discovery of Rhinolophus beddomei (Chiroptera: Rhinolophidae)
from Thailand with a Brief Comparison to Other Related Taxa
P
IPAT SOISOOK
1, 2*
, PIYAWAN NIYOMWAN
3
, MATTANA SRIKRACHANG
3
,
T
UANJIT SRITHONGCHUAY
4
AND PAUL J. J. BATES
5
1
Princess Maha Chakri Sirindhorn Natural History Museum, Faculty of Science, Prince of Songkla University,
Hat Yai, Songkhla 90112, THAILAND
2
Excellence Centre for Biodiversity of Peninsular Thailand, Faculty of Science, Prince of Songkla University,
Hat Yai, Songkhla 90112, THAILAND
3
Wildlife Research Section, Wildlife Conservation Division, Department of National Park, Wildlife and Plant
Conservation, Pahonyothin Road, Chatuchak, Bangkok 10900, THAILAND
4
Department of Biology, Faculty of Science, Prince of Songkla University, Hat Yai, Songkhla 90112, THAILAND
5
Harrison Institute, Centre for Systematics and Biodiversity Research, Bowerwood House,St. Botolph’s Road,
Sevenoaks, Kent, TN13 3AQ, UK
* Corresponding author. E-mail: pipat.s@psu.ac.th
Received: 21 September 2009; Accepted: 17 March 2010
ABSTRACT.– Rhinolophus beddomei is reported from mainland Southeast Asia for the first time. It was
collected in April, 2008, in Ratchaburi Province, Thailand, in a harp trap set over a stream in a dense
evergreen forest. Morphometric and acoustic characters of the specimen are given with a brief comparison
to other species of the R. trifoliatus group. Currently, it is not clear whether the Thai specimen represents
an eastern extension of the Indian Subcontinent population or whether it is a member of an isolated
population which is restricted to continental Southeast Asia. It is suggested that a further review, which
would involve material from southern and Southeast Asia, together with R. formosae from Taiwan would be
of considerable interest.
K
EY WORDS: Rhinolophus beddomei, R. trifoliatus group, first record, Southeast Asia, Thailand, disjunct
distribution
I
NTRODUCTION
Following a series of intensive bat
surveys by members of the Prince of
Songkla University bat team from 2003 to
July 2008, eleven bat species were recorded
from Thailand for the first time (Bumrungsri
et al., 2006; Thong et al., 2006; Bates et al.,
2007; Soisook et al., 2007). This brought the
total for the country to 122 species of which
18 belong to the genus Rhinolophus,
including R. microglobosus Csorba and
Jenkins, 1998, which has recently been
elevated to specific rank (Soisook et al.,
2008).
In April, 2008, a field survey was
conducted in Ratchaburi Province, western
Thailand. Of particular interest was a single
male specimen of a taxon not recorded
previously from Thailand. The specimen is
here referred to Rhinolophus beddomei
Andersen, 1905 on the basis of size and
morphology. Rhinolophus beddomei is one
of five species which are assigned to the R.
trifoliatus group (sensu Csorba et al., 2003).
They are characterised by the presence of
lateral lappets at base of the sella in the
noseleaf. Three of the five species, R. luctus
Temminck, 1834, R. trifoliatus Temminck,
1834, and R. sedulus Andersen, 1905, have
been recorded previously from mainland

TROPICAL NATURAL HISTORY 10(1), APRIL 2010 68
F
IGURE 1. The known distribution of R. beddomei (circle), R. luctus (squares), R. trifoliatus (triangles) and R.
sedulus (diamonds) in mainland Southeast Asia. Black symbols represent localities from which specimens
have been examined, whereas localities with hollow symbols have not been examined. Locality information
is included in the Appendix.

SOISOOK ET AL. — DISCOVERY OF RHINOLOPHUS BEDDOMEI 69
Southeast Asia (Corbet and Hill, 1992;
Simmons, 2005), whilst the fourth, R.
formosae Sanborn, 1939 is endemic to
Taiwan.
Rhinolophus beddomei is regarded as
distinct from R. luctus on the basis of its
smaller size and the shape of the upper
canine (Topál and Csorba, 1992; Csorba et
al., 2003). Until now, its range was thought
to be restricted to western and southern India
and Sri Lanka (Bates and Harrison 1997).
However, here we report the discovery of R.
beddomei from Thailand with information on
its echolocation and morphometrics. It is
compared to the four other species of the
trifoliatus group.
M
ATERIALS AND METHODS
The specimen of R. beddomei was
collected from Mae Nam Pha Chi Wildlife
Sanctuary, Ratchaburi Province, western
Thailand (13º 15’ N, 99º 21’ E) (loc. B1 in
Fig. 1) at an altitude of 431 m a.s.l on 20
April 2008 during a wildlife diversity
survey by staff of the Department of
National Park, Wildlife and Plant
Conservation (DNP) and the Prince of
Songkla University. The wildlife sanctuary
covers an area of 489.31 km
2
and is
characterised by steeply rugged mountains
which are part of the Tenasserim Mountain
Range. The Pha Chi River runs through the
western part of the sanctuary. The
vegetation comprises four main forest types;
dry evergreen forest, which is the
predominant forest type with additional
primary evergreen forest and mixed
deciduous and dipterocarp forest. The
underlying geology is granite. The wildlife
sanctuary is in the rainshadow of the
Tenasserim Mountain Range and
consequently there is a relatively low annual
average precipitation of 246 mm (DNP,
2006).
The R. beddomei specimen was captured
in a four-bank harp trap (Francis, 1989) set
across a 3 m wide seasonal streamlet in
primary evergreen forest. Five other species
of insectivorous bats were also collected;
Nycteris tragata, Rhinolophus microglobo-
sus, Myotis muricola, Kerivoula papillosa
and Phoniscus jagorii. The specimen of R.
beddomei is preserved in 70% (v/v) alcohol
with its skull extracted and deposited in the
Princess Maha Chakri Sirindhorn Natural
History Museum, Prince of Songkla
University, Hat Yai, Thailand. External and
craniodental characters were measured with
a digital caliper following Bates and
Harrison (1997) and Csorba et al. (2003).
Body mass (MASS) was recorded using a
Pesola 50 g scale and is given in grams.
Additional specimens of R. beddomei, R.
luctus, R. trifoliatus and R. sedulus included
in this study were studied at the Harrison
Institute, Sevenoaks, UK (HZM), The
Natural History Museum, London (BMNH)
and Princess Maha Chakri Sirindhorn
Natural History Museum (PSUZC-MM;
formerly PSU-M) (see Appendix).
The echolocation calls were recorded
using a hand-held Pettersson D-240X time
expansion bat detector, which was
connected to iRiver iHP-120 Multi-Codec
Jukebox recorder. Calls were analysed with
BatSound Pro version 3.31 (Pettersson,
Elektronik AB). A sampling frequency of
44.10 kHz was used and produced a
spectrogram using 1024 samples Fast
Fourier Transform (FFT) with Hanning
window.
Measurements provided in this paper
were taken from specimens examined for
this study only, unless otherwise stated.

TROPICAL NATURAL HISTORY 10(1), APRIL 2010 70
SYSTEMATICS
Rhinolophus beddomei Andersen, 1905
(Figures 1 and 2; Tables 1 and 2)
Rhinolophus beddomei Andersen, 1905:
253; Wynaad, Madras, India.
Rhinolophus beddomei sobrinus Andersen,
1918: 378; Kala Oya, North Central
Province, Sri Lanka.
Material.– Thailand: Huay Pu Nam Ron,
Mae Nam Pha Chi Wildlife Sanctuary, Suan
Pueng, Ratchaburi Province (13º 15’ N, 99º
21’ E, 431 m a.s.l.) (Fig. 1), 20 April, 2008,
1♂ (PSUZC-MM2008.51) collected by
Pipat Soisook and Tuanjit Srithongchuay.
This is the first record from Thailand.
Morphometric characters.– Specimen
PSUZC-MM2008.51 from Ratchaburi
Province has a forearm length of 53.0 mm
(for comparative information on material
from India and Sri Lanka, see Tables 1 and
2, and also Bates and Harrison, 1997). The
noseleaf is dark brown with well developed
lappets at the base of the sella; the
horseshoe is broad (12.2 mm) (Fig. 2a). The
lower lip has one groove. The ears are 23.8
mm in height. In the wing, the shortest
metacarpal is the third, which is
significantly shorter than the fourth and
fifth. The wing membranes are uniformly
dark brown and are attached at the base of
each outer toe. The length of the second
phalanx of the third digit is 1.5 times that of
the first. The fur is soft, woolly in texture
and rather long, ca. 11 - 13 mm. The dorsal
and ventral pelage is uniformly whitish-
brown at the base with greyish-brown tips.
The skull has a condylo-canine length of
21.7 mm. Zygomatic breadth exceeds
mastoid width (Table 2). The anterior rostral
chambers are robust and well inflated when
viewed laterally. The sagittal crest is well
developed; it bifurcates anteriorly to form
two strong supraorbital ridges around a very
deep pit in the interorbital region. The bony
palate is 34.0% of the length of the
maxillary toothrow (C-M
3
). The upper
canine (C
1
) is without a concavity in its
posterior base; it is in contact with the first
upper premolar (PM
2
), which is medium
sized and lies within the toothrow. The
second lower premolar (PM
3
) is absolutely
extruded from the toothrow; the first (PM
2
)
and third (PM
4
) premolars are in contact.
Acoustic characters.– The hand held
echolocation call of R. beddomei (specimen
PSUZC-MM2008.51) from Ratchaburi
Province, Thailand is typical of the genus
F
IGURE 2. Shape of noseleaf and pelage colour of (A) Rhinolophus beddomei from western Thailand, (B) R.
luctus from northeastern Thailand and (C) R. trifoliatus from peninsular Thailand.

SOISOOK ET AL. — DISCOVERY OF RHINOLOPHUS BEDDOMEI 71
Rhinolophus. It starts with a short duration
FM component with an initial frequency of
around 47.9 kHz. This is followed by a 36 -
39 ms CF segment of the peak frequency at
49.3 kHz (n = 1). It finishes with a short FM
sweep with a terminal frequency of 45.6
kHz.
Taxonomic notes.– The external, cranial
and dental morphology and pelage colour of
the Thai specimen is similar to that of
specimens from India and Sri Lanka.
However, the Thai specimen is distinctly
smaller than the nominate subspecies R. b.
beddomei from India and slightly smaller
than R. b. sobrinus from Sri Lanka (Tables 1
and 2, see also Bates and Harrison, 1997).
Since only one individual is known, it is
unclear whether this is typical of the Thai
population or reflects an abnormally small
individual.
An additional specimen (BMNH.84.
1970) collected in Sabah and housed in the
Natural History Museum, London, may also
be referable to R. beddomei. This wet
specimen, with skull not extracted, is
currently labelled ‘R. trifoliatus’. However,
its dark noseleaf and external measurements
(FA 53.1 mm) suggest that it may be a
second specimen of R. beddomei from
Southeast Asia.
Two individuals collected by Struebig et
al. (2006) from Kalimantan (Indonesian
Borneo) are also comparable in size and
morphology to R. beddomei. They were
provisionally assigned to R. luctus but the
authors noted that they were intermediate in
size between R. luctus and R. sedulus and
‘may represent a distinct taxon’. With
forearm lengths of 52.9 mm and 52.6 mm
and tibia lengths of 29.5 mm and 28.7 mm,
they agree in size with the specimen from
Thailand.
Comparison to other related species.– In
comparison to R. luctus, R. beddomei is
distinguished by its smaller size (Tables 1
and 2; see also Topál and Csorba, 1992;
Bates and Harrison, 1997), absence of a
concavity in the posterior base of the upper
canine, and differences in pelage colour and
echolocation call. Both species have long
fur that is woolly in texture. However, the
fur of R. luctus from peninsular Thailand is
dark grey to almost black whereas that of R.
beddomei from Thailand is greyish-brown
(Fig. 2). The hand held frequencies of three
individuals of R. luctus from peninsular
Thailand have CF components of between
31.6 and 32.0 kHz and a duration of 45 - 79
ms (n = 3) (PS, unpublished data). This is
much lower than that of the Thai R.
beddomei (for details see above). The recent
Thai data for R. luctus are comparable to
Francis (2008) who also reported call
frequencies of 32 - 34 kHz for R. luctus
from peninsular Thailand and Lao PDR, but
contrasts with the individuals from Sabah
and peninsular Malaysia, where a constant
frequency of 40 - 42 kHz was recorded
(Kingston et al., 2000; Francis, 2008).
Externally, R. beddomei can be distingui-
shed from R. trifoliatus by its darker pelage
and noseleaf colour (Fig. 2) and cranially by
its broader rostrum (RW) and anterior
rostral chambers (AMSW) (Table 2).
However, the echolocation calls of the two
species are quite similar, with the CF
component of R. trifoliatus
(50.0 - 53.5
kHz, n = 3: PS, unpublished data) from
peninsular Thailand only slightly higher
than that of R. beddomei.
Rhinolophus beddomei from Thailand
greatly exceeds R. sedulus in size, whilst the
call frequency of R. sedulus, 66.8 kHz from
peninsular Malaysia (Kingston et al., 2000)
and 62 and 76 kHz from Sabah (Francis,
2008) exceeds that of Thai R. beddomei.

TROPICAL NATURAL HISTORY 10(1), APRIL 2010 72
Rhinolophus beddomei is indistinguish-
able in size and morphology from R.
formosae, which is currently regarded as an
endemic species to Taiwan (Csorba et al.,
2003). The specific status of R. formosae
was based on its smaller size and differences
in karyology from R. luctus (Ando et al.,
1983; Hood et al., 1988; Yoshiyuki and
Harada, 1995). However, no detailed
comparison was made by these authors with
R. beddomei.
Ecological and conservation notes.– The
specimen of R. beddomei from Thailand was
collected in harp trap which was set across a
seasonal stream at an altitude of 431 metres
(see Methods Section). Since there were
apparently no caves in the vicinity, it is
possible that its diurnal roost is tree hollows
or in a rock crevice. In India and Sri Lanka,
R. beddomei was found roosting singly or
up to three individuals in hollow trees, small
caves and hanging from boulders (Bates and
Harrison, 1997).
Based on previous information from
India and Sri Lanka, the conservation status
of R. beddomei is listed as ‘Least Concern’
(Srinivasulu and Molur, 2008). The new
material from Thailand greatly extends its
known range. However, although Mae Nam
Pha Chi Wildlife Sanctuary is officially a
protected area, the Sanctuary is still subject
to considerable deforestation and hunting
pressure. Currently, the Sanctuary, and also
other nearby protected areas in southwest
Thailand (e.g. Kang Krachan National
Park), are being assessed as part of the
‘Wildlife and Its Habitat Assessment in the
TABLE 1. External measurements (in mm) and mass (in gram) of R. beddomei and R. luctus; FA: forearm
length; EAR: ear length; HB: head and body length; TAIL: tail length; HF: foot length; TIBIA: tibia length;
5MET, 4MET, 3MET: fifth, fourth and third metacarpal length; 3D1P, 3D2P: first and second phalanx of the
third digit; NL: noseleaf width; MASS: body mass. Sample size (n), mean, standard deviation and range are
shown. Sample sizes differing from those reported under n are given in brackets.
n and sex FA EAR HB TAIL HF
Rhinolophus beddomei
(Thailand)
1♂
53.0 23.8 63.4 30.9 14.1
59.8±3.9 27.5, 27.6 67.5 37.0, 38.5 12.1, 13.5
Rhinolophus beddomei
(India and Sri Lanka)
5 (4♂, 1♀)
(55.0-63.4) [2] [1] [2] [2]
Rhinolophus luctus
19 (8♂, 11♀)
70.5±4.3 36.3±2.9 76.8±5.3 51.5±6.1 16.3±2.0
(61.8-77.3) (31.5-39.6)
[11]
(65.4-85.7)
[10]
(41.0-64.3)
[13]
(12.0-18.7)
[14]
Rhinolophus trifoliatus
8 (3♂, 5♀)
51.8±2.0 25.8±0.4 53.8±2.4 34.3±1.7 12.0±0.5
(48.9-54.1) (25.4-26.2)
[4]
(51.1-56.3)
[4]
(32.3-36.4)
[4]
(11.4-12.4)
[4]
Rhinolophus sedulus*
? (?♂, ?♀)
37.0-48.0 17.0-21.0 47.0-51.0 19.0-30.0 -
- - - - -
* After Kingston et al. (2006)

SOISOOK ET AL. — DISCOVERY OF RHINOLOPHUS BEDDOMEI 73
Corridor Zone under the Biodiversity
Conservation Corridors Initiative Pilot Site
in the Tenasserim WEFCOM Thailand’
which is a joint project of WCS Thailand
and DNP (see www.wcsthailand.org ).
D
ISCUSSION
The specimen from Thailand has been
referred to Rhinolophus beddomei on the
basis of its size and morphology. However,
it is not clear whether this specimen
represents an eastern extension of a
population contiguous with those in
southwest India or whether it is a member of
an isolated population which is somehow
restricted to continental Southeast Asia.
This situation is similar to that of the
endangered Molossid, Otomops wroughtoni
(Thomas, 1913), which for many years had
been considered an endemic to western
India, but was subsequently recorded from
Cambodia (Walston and Bates, 2001).
The balance of probability suggests that
the population of R. beddomei is not
contiguous. Unlike Otomops wroughtoni
(Thabah and Bates, 2002), there are no
records from north or eastern India and it
has also not been found in Myanmar despite
many recent faunal surveys (Bates et al.,
2004). If the population is isolated then
there is the possibility that the phenotypic
similarity of the Thai specimen to R.
beddomei from the Indian Subcontinent may
not be reflected in its genotype. This would
mirror the findings of Thabah et al. (2006)
who in their study of the Hipposideros
larvatus species complex discovered an
TABLE 1. Continued.
TIBIA 5MET 4MET 3MET 3D1P 3D2P NL MASS
28.3 44.0 42.5 37.5 18.5 27.8 12.2 16.0
30.2±1.3 45.3, 46.3 41.1, 41.1 37.8, 37.8 19.5, 21.6 27.8, 28.4 11.0 -
(28.9-31.9) [2] [2] [2] [2] [2] [1] -
36.8±2.0 59.5±2.8 57.7±2.4 50.3±2.0 28.5±1.0 40.0±2.2 15.3±2.0 56.2
(33.7-40.2)
[13]
(55.2-62.4)
[6]
(54.1-60.1)
[6]
(47.7-53.4)
[6]
(27.2-29.6)
[6]
(38.0-44.0)
[6]
(11.7-17.2)
[7]
[1]
26.1±1.0 43.3±0.8 41.4±1.0 36.5±1.1 21.1±0.8 29.2±1.6 11.2±0.2 14.0±1.3
(24.9-27.4) (42.3-44.1)
[4]
(40.2-42.4)
[4]
(35.5-38.0)
[4]
(20.3-22.1)
[4]
(27.7-30.9)
[4]
(11.0-11.4) (12.5-15.0)
[3]
19.0-20.5 - - - - - - 5.6-11.5
- - - - - - - -

TROPICAL NATURAL HISTORY 10(1), APRIL 2010 74
interesting but confused pattern involving a
lack of congruence between phenotype,
genotype and acoustic characters. The
situation in ‘R. beddomei’ may prove to be
analogous. Therefore a more thorough
review, which includes R. beddomei from
India and Sri Lanka, the new specimen from
Thailand and previous specimens from
Sabah (BMNH collection) and Kalimantan
(Struebig et al., 2006) would be of
considerable interest. It would also be
valuable to include R. formosae in the study,
since despite its morphological similarity,
its relationship with R. beddomei has never
been clearly elucidated. It is very probable
that future field surveys in forest areas using
harp traps may show that R. beddomei is
quite widespread in Southeast Asia.
In terms of acoustics, it is interesting to
note that as might be predicted by previous
studies (Robinson, 1996; Francis and Haber-
setzer, 1998; Zhang et al., 2000; Soisook et
al., 2008), the call frequency of R. beddomei
from western Thailand (49.3 kHz), with its
smaller body size, is higher than the closely
related larger species R. luctus from penin-
sular Thailand (32.0 kHz). However, it is
distinctly lower than that of the smallest
member of the trifoliatus group, R. sedulus
from peninsular Malaysia, which has a call of
66.8 kHz (Francis, 2008). The hand held
frequency of R. beddomei in Thailand is only
slightly lower than R. trifoliatus from
peninsular Thailand (50.0 - 53.5 kHz), which
averages slightly smaller.
TABLE 2. Cranial and dental measurements (in mm) of R. beddomei and R. luctus; GTL: greatest length of
skull; SL: skull length; CCL: condylo-canine length; ZB: zygomatic breadth; BB: breadth of braincase; MW:
mastoid width; PC: postorbital constriction; C–M
3
: upper toothrow length; PL: palate length; M
3
–M
3
:
posterior palatal width; C
1
–C
1
: anterior palatal width; C–M
3
: lower toothrow length; M: mandible length;
AMSW: anterior median swelling width; RW: greatest rostral width. Sample size (n), mean, standard
deviation and range are shown. Sample sizes differing from those reported under n are given in brackets.
n and sex GTL SL CCL ZB BB MW PC
Rhinolophus beddomei
(Thailand) 1♂ 25.9 24.4 21.7 12.2 9.9 11.3 2.4
Rhinolophus beddomei
(India and Sri Lanka) 26.9±0.9 26.1±1.0 23.4±0.8 13.6±0.7 10.8±0.7 11.8±0.5 2.5±0.3
6 (4♂, 2♀) (26.1-28.5) (25.0-27.2) (22.4-24.5)
[5]
(12.5-14.5) (9.8-11.5) (11.0-12.3) (2.1-3.0)
Rhinolophus luctus
17 (5♂, 12♀)
31.6±1.7 30.5±1.5 27.4±1.0 15.4±0.7 12.2±0.4 13.4±0.4 3.2±0.4
(28.2-33.7)
[13]
(26.8-32.1)
[16]
(25.1-28.6)
[16]
(14.5-16.5)
[17]
(11.6-13.1)
[17]
(12.6-13.9)
[16]
(2.6-3.9)
[17]
Rhinolophus trifoliatus
8 (4♂, 4♀)
23.6±1.0 22.8±0.8 20.3±0.7 11.7±0.4 9.5±0.4 10.6±0.3 2.2±0.2
(22.3-24.6)
[4]
(22.0-23.9)
[6]
(19.5-21.4)
[6]
(11.2-12.5)
[7]
(8.8-10.0)
[7]
(10.1-10.9)
[6]
(1.9-2.5)
[7]
Rhinolophus sedulus*
3 (2♂, 1♀)
20.3±0.6 19.6±0.6 16.9, 17.8 9.9±0.4 8.3±0.2 9.0, 9.6 2.2±0.1
(19.7-21.0) (19.0-20.2) [2] (9.7-10.4) (8.1-8.5) [2] (2.1-2.4)
* Included specimens from Borneo

SOISOOK ET AL. — DISCOVERY OF RHINOLOPHUS BEDDOMEI 75
ACKNOWLEDGEMENTS
In Thailand, we are grateful to the
Director General of the Department of
National Park, Wildlife and Plant
Conservation for a research permit. We also
would like to thank Chairattana
Semasawaddi and all the staff of Mae Nam
Pha Chi Wildlife Sanctuary for their
assistance in field work. At Prince of Songkla
University, we thank Chutamas Satasook and
Sara Bumrungsri for their kind support and
also Ariya Dejtaradol and Bounsavane
Douangboubpha for their help in providing
specimens and measurements. In the UK, we
would like to thank Paula Jenkins, Roberto
Portela Miguez and the staff of the Natural
History Museum, London for their help with
examining specimens. We thank Matt
Struebig for his kind advice. At the Harrison
Institute, we thank David Harrison and
Malcolm Pearch for their advice. At the
University of Bristol, thanks are due to
Gareth Jones and Alice Hughes for their
help, and to the Prime Minister’s Initiative
for International Education 2 (PMI2) for
financial support during a visit by PS to the
UK. We are also most grateful to the Darwin
Initiative of the UK Government (Project
Nos: 14011, 18002) for their on-going
support our taxonomic research and training
programmes in Southeast Asia.
TABLE 2. Continued.
C-M
3
PL M
3
-M
3
C
1
-C
1
C-M
3
M AMSW RW C-M
3
9.3 3.2 8.9 6.1 10.0 17.2 4.8 6.5 9.3
9.8±0.5 3.8±0.3 9.6±0.5 6.7±0.4 10.5±0.5 18.3±0.7 4.9±0.3 6.8±0.4 9.8±0.5
(9.3-10.5) (3.3-4.2) (9.2-10.2) (6.1-7.2)
(9.9-11.1) (17.5-19.1) (4.5-5.2) (6.2-7.2) (9.3-10.5)
11.9±0.5 4.6±0.3 11.0±0.6 7.7±0.3 12.7±0.6 22.1±1.0 5.9±0.3 8.6±0.4 11.9±0.5
(11.1-12.8)
[17]
(4.1-5.3)
[17]
(10.2-12.9)
[17]
(7.2-8.5)
[17]
(11.8-13.8)
[17]
(20.1-23.7)
[17]
(5.4-6.7)
[17]
(7.7-9.1)
[17]
(11.1-12.8)
[17]
8.6±0.3 2.9±0.3 8.3±0.2 5.5±0.3 9.1±0.3 15.7±0.6 4.2±0.1 6.0±0.1 8.6±0.3
(8.2-8.9) (2.5-3.4) (7.9-8.5) (5.1-5.8) (8.6-9.5) (14.9-16.4)
[7]
(4.0-4.3) (6.0-6.1) (8.2-8.9)
7.2±0.5 2.2±0.2 5.6±2.7 4.6±0.1 7.7±0.5 13.4 4.0±0.1 5.3±0.2 7.2±0.5
(6.6-7.6) (2.1-2.5) (2.5-7.2) (4.6-4.7) (7.1-8.1) [1] (3.9-4.1) (5.1-5.5) (6.6-7.6)

TROPICAL NATURAL HISTORY 10(1), APRIL 2010 76
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TROPICAL NATURAL HISTORY 10(1), APRIL 2010 78
APPENDIX
List of examined specimens and localities, including previous records shown in Figure 1.
Rhinolophus beddomei – INDIA: Wynaad, Madras (11º45’N, 76º02’E): BMNH.82.3.3.1.
(type of beddomei); Konkan, Maharashtra (not located): BMNH.11.3.16.1.; Sirsi, Karnataka
(14º40’N, 74º51’E): BMNH.12.11.28.5.; Kannadimazhligai, Courtallum Hills, Tirunelveli
District, Tamil Nadu (08º45N, 77º43’E): HZM.2.36212; Rojampela Range (not located):
BMNH.30.5.24.60.; SRI LANKA: Kala Oya, North Western Province (08º12’N, 80º04’E):
BMNH.18.8.3.3. (type of sobrinus); Mapalagama, Talgaswela, Southern Province (appr.
06
o
15’N, 80
o
15’E): HZM.1.27643.; THAILAND: Huay Pu Nam Ron, Mae Nam Pha Chi
Wildlife Sanctuary, Suan Pueng, Ratchaburi Province (13º15’N, 99º21’E) [loc. B1]: PSUZC-
MM.2008.51.; MALAYSIA: Sabah (not located): BMNH. 84.1970 (labelled as R. trifoliatus).
Rhinolophus luctus – MYANMAR: Nam Tamai, Upper Myanmar (27
o
42’N, 97
o
54’E) [loc.
L1]: BMNH.50.396, BMNH.50.397; Sokleik (not located): BMNH.21.1.6.1.; Chin Hill (appr.
23
o
40’N, 94
o
15’E) [loc. L2]: BMNH.21.1.6.3.; Bankachon, S. Tenasserim (appr. 10
o
12’N,
98
o
37’E) [loc. L3]: BMNH.21.1.6.2.; VIETNAM: Nui Hoang Lien Nature Reserve, Sa Pa
District, Lao Cai Province (appr. 22
o
10’N, 103
o
57’E) [loc. L4]: BMNH.1997.336; Pu Mat
Reserve (20
o
19’N, 105
o
37’E) [loc. L5]: HZM.1.31766; Phong Nha-Ke Bang Prop. NP
(17
o
39’N, 105
o
59’E) [loc. L6]: HZM.2.32212, HZM.2.32213; Bach Ma NP (16
o
12’N,
107
o
45’E) [loc. L7]: T129 (I.E.B.R); LAO PDR: Tam Nong Seng (18
o
05’N, 104
o
28’E) [loc.
L8]; Tam Dinphye (17
o
36’N, 104
o
58’E) [loc. L9]; Khammouan: including Tam Lort and Tam
Lom (17
o
35’N, 104
o
48’E) [loc. L10]: Robinson and Webber (1998); CAMBODIA: Mt.
Samkos Wildlife Sanctuary, Cardomom Mountains (appr. 12
o
22’N, 103
o
02’E) [loc. L11]:
HZM.4.32764; Tatai Lieu (11
o
49’N, 103
o
32’E) [loc. L12]: HZM.6.36475; Cambodia (no
exact locality): HZM.5.36141; THAILAND: Amphoe Samung, Chiang Mai (18
o
55’N,
98
o
40’E) [loc. L13]: Sawada and Harada (1985); Doi Pui, Chiang Mai (appr. 18
o
49’N,
98
o
53’E) [loc. L14]: BMNH.78.2310.; Chiang Mai (not located): BMNH.9.10.11.2.; Khun
Tan, Lampang (appr. 17
o
15’N, 100
o
10’E) [loc. 15]: Shamel (1942); Nam Nao cave,
Petchabun Province (16
o
57’N, 100
o
14’E) [loc. L16]: PP070915.27; KM 79, Highway 23,
Nakhon Ratchasima Province (appr. 14
o
58’N, 102
o
06’E) [loc. L17]: BMNH.70.1463.; Pak
Thong Chai, Sakaerat, Nakhon Ratchasima Province (appr. 14
o
35’N, 101
o
50’E) [loc. L18]:
BMNH.78.2309.; Prachin Buri (appr. 14
o
14’N, 101
o
50’E) [loc. L19]: Yenbutra and Felten
(1986); Khao Ang Runai Wildlife Sanctuary, Chacherngsao Province (13
o
24’N, 101
o
52’E)
[loc. L20]: PSUZC-MM2005.36; Krung Ching, Nakhon Si Thammarat Province (8
o
47’N,
99
o
38’E) [loc. L21]: PP071108.1; Kao Luang, Nakhon Si Thammarat (appr. 8
o
24’N,
99
o
58’E) [loc. L22]: Shamel (1942); Yala (appr. 6
o
06’N, 101
o
18’E) [loc. L23]: Yenbutra and
Felten (1986); MALAYSIA: Tanah Rata, Pahang (4
o
28’N, 101
o
23’E) [loc. L24]: Csorba et al.
(2003); Semangko, Selangor (appr. 3
o
40’N, 101
o
44’E) [loc. L25]: BMNH.1.3.9.3.; Taiping,
Perak (appr. 3
o
41’N, 101
o
45’E) [loc. L26]: BMNH.56.127.; Genting Highland (appr. 3
o
25’N,
101
o
47’E) [loc. L27]: Heller and Volleth (1988); Ulu Gombak (3
o
20’N, 101
o
45’E) [loc. L28]:
Fain (1982); Bangai Forest Reserve, Selangor (2
o
55’N, 101
o
45’E) [loc. L29]: Zubaid (1993);

SOISOOK ET AL. — DISCOVERY OF RHINOLOPHUS BEDDOMEI 79
Rhinolophus trifoliatus – MYANMAR: Murgui, Tenasserim (12
o
26’N, 98
o
34’E) [loc. T1]:
BMNH.85.8.1.110, BMNH85.8.1.111; Bankachon, S. Tenasserim (appr. 10
o
12’N, 98
o
37’E)
[loc. T2]: BMNH.14.12.8.244.; THAILAND: Thung Yai Naresuan WS (15
o
19’N, 98
o
49’E)
[loc. T3]: Robinson et al. (1996); Kao Soi Dao, Chantaburi Province (appr. 13
o
06’N,
102
o
13’E) [loc. T4]: Csorba et al. (2003); Ban Bang Non, Muang Ranong (appr. 9
o
60’N,
98
o
39’E) [loc. T5]: BMNH.78.2311., BMNH78.2312; Ranong Province (appr. 9
o
55’N,
98
o
35’E) [loc. T6]: Yenbutra and Felten (1986); Ban Sichon, Nakhon Si Thammarat (9
o
00’N,
99
o
56’E) [loc. T7]: Shamel (1942); Trang Province (appr. 7
o
04’N, 99
o
36’E) [loc. T8]:
Yenbutra and Felten (1986); Khao Kor Hong, Hat Yai, Songkhla Province (7
o
00’N,
100
o
30’E) [loc. T9]: PSUZC-MM2006.167; Namom, Songkhla Province (6
o
57’N, 100
o
33’E)
[loc. T10]: PSUZC-MM2005.160; Wildlife Education Centre, Hat Yai, Songkhla Province
(6
o
57’N, 100
o
14’E) [loc. T11]: PSUZC-MM2006.140, PSUZC-MM2006.141; Satun
Province (appr. 6
o
40’N, 100
o
00’E) [loc. T12]: Yenbutra and Felten (1986); Narathiwat
Province (appr. 6
o
28’N, 101
o
51’E) [loc. T13]: Shamel (1942); Bangnara, Narathiwat
Province (6
o
28’N, 101
o
51’E) [loc. T14]: Csorba et al. (2003); MALAYSIA: Gunung Tahera,
Pahang (appr. 3
o
35’N, 102
o
15’E) [loc. T15]: BMNH.6.10.4.8., BMNH.65.337.; Ulu Gombak
(3
o
20’N, 101
o
45’E) [loc. T16]: Heller and Volleth (1988); Templer Park (appr. 3
o
17’N,
101
o
37’E) [loc. T17]: Heller and Volleth (1988); Kepong Cubitt Forest Reserve (appr.
3
o
12’N, 101
o
27’E) [loc. T18]: Csorba et al. (2003); Bangai Forest Reserve, Selangor (2
o
55’N,
101
o
45’E) [loc. T19]: Zubaid (1993); SINGAPORE (appr. 1
o
23’N, 103
o
50’E) [loc. T20]:
BMNH74.4.18.1. (type of R. t. edax).
Rhinolophus sedulus – MALAYSIA: Kuala Tekah (appr. 4
o
10’N, 102
o
19’E) [loc. S1]:
Medway (1969); Jenka, Temerloh, Pahang (3
o
27’N, 102
o
25’E) [loc. S2]: BMNH.65.334.; Ulu
Gombak (3
o
20’N, 101
o
45’E) [loc. S3]: Heller et. al. (1993); Kepong Cubitt Forest Reserve
(appr. 3
o
12’N, 101
o
27’E) [loc. S4]: Csorba et. al. (2003); Sarawak (not located):
BMNH7.1.1.292. (type of R. sedulus); BORNEO: Paitan (not located): BMNH.94.7.2.48.