A New Species of the Genus Murina (Chiroptera: Vespertilionidae) from the Central Highlands of Vietnam with a Review of the Subfamily Murininae in Vietnam

Abstract

The subfamily Murininae has high species diversity in Vietnam, but taxonomic studies are limited. In this paper, we describe a new species of the genus Murina based on a specimen collected from Ngoc Linh Nature Reserve, Kon Turn Province in the Central Highlands of Vietnam. It is a medium-sized species with 'sitilla-type' dentition. A taxonomic review of Murininae from Vietnam was also conducted based on combined morphological, DNA, and karyological characteristics. Molecular phylogenetic analyses based on the mitochondrial cytochrome c oxidase subunit (COI) gene supported the subfamily Murininae, while the genus Murina proved to be paraphyletic in relation to the genera Harpiocephalus and Harpiola. Fourteen species of the genus Murina, one species of Harpiocephalus, and one species of Harpiola are recognized from Vietnam. Murina tiensa is regarded as a junior synonym of M.harrisoni; strong sexual dimorphism was observed in M. harrisoni. Relations between forearm length and total length of skull showed different trends among species and sexes. Karyotypes of Murina huttoni, M. cyclotis, M. lorelieae, M. beelzebub, M. feae, and Harpiola isodon were 2n = 44, FN = 50, while that of Harpiocephalus harpia was 2n = 44, FN = 52.

Full text

INTRODUCTION
Species of the subfamily Murininae are small to
medium-sized bats of the family Vespertilionidae.
They are distributed throughout Asia from north-
eastern Pakistan, eastern Siberia, Korea to Japan to
northeastern Australia. Diagnostic characters are the
combination of: tubular nostrils, thick and woolly
fur, furred forearms, hind limbs, proximal parts of
wing membranes, and upper surface of interfemoral
membrane; and two premolars in both the upper and
lower toothrows, where the first premolars are
unusually well-developed (Tate, 1941; Corbet and
Hill, 1992; Koopman, 1994; Simmons, 2005; Kuo et
al., 2006, 2009; Francis and Eger, 2012).
Simmons (2005) listed 17 species of the ge-
nus Murina Gray, 1842 and subsequent studies de-
scribed further 21 species, mostly from southern Asia
(Csorba and Bates, 2005; Bumrungsri et al., 2006;
Csorba et al., 2007, 2011; Kruskop and Eger, 2008;
Furey et al., 2009; Kuo et al., 2009; Eger and Lim,
2011; Francis and Eger, 2012; Ruedi et al., 2012;
Soisook et al., 2013a, 2013b; Tu et al., 2015).
Inventory studies have been recently conducted in
many Southeast Asian countries such as Thailand
(10 species — Bumrungsri et al., 2006; Soisook,
2011; Soisook et al., 2013a, 2013b), Laos (eight
species — Francis and Eger, 2012), Cambodia (five
species — Csorba and Bates, 2005; Matveev and
Csorba, 2007; Csorba et al., 2011; Ith et al., 2011),
and Myanmar (four species — Bates et al., 2000),
indicating that the genus Murina includes interest-
ing forest bats with high species diversity in Asia.
The subfamily Murininae also includes further
two genera, Harpiocephalus Gray, 1842 and Har -
piola Thomas, 1915. The validity of the genus
Acta Chiropterologica, 17(2): 201–232, 2015
PL ISSN 1508-1109 © Museum and Institute of Zoology PAS
doi: 10.3161/15081109ACC2015.17.2.001
A new species of the genus Murina (Chiroptera: Vespertilionidae) from the Central
Highlands of Vietnam with a review of the subfamily Murininae in Vietnam
NGUYEN TRUONG SON1, 8, GABOR CSORBA2, VUONG TAN TU1, VUDINH THONG1, YIWU3,
MASASHI HARADA4, TATSUO OSHIDA5, HIDEKI ENDO6, and MASAHARU MOTOKAWA7
1Institute of Ecology and Biological Resources, Vietnam Academy of Sciences and Technology, 18 Hoang Quoc Viet St., Caugiay,
Hanoi, Vietnam
2Department of Zoology, Hungarian Natural History Museum, H1088 Budapest, Baross u.13, Hungary
3College of Life Science, Guangzhou University, Guangzhou 510006, China
4Graduate School of Medicine, Osaka City University, Osaka 545-8585, Japan
5Laboratory of Wildlife Ecology, Obihiro University of Agriculture and Veterinary Medicine, Obihiro 080-8555, Japan
6The University Museum, The University of Tokyo, 7-3-1 Hongo, Bunkyo-ku, Tokyo 113-0033, Japan
7The Kyoto University Museum, Kyoto University, Sakyo, Kyoto 606-8501, Japan
8Corresponding author: E-mail: ntson@iebr.vast.vn
The subfamily Murininae has high species diversity in Vietnam, but taxonomic studies are limited. In this paper, we describe a new
species of the genus Murina based on a specimen collected from Ngoc Linh Nature Reserve, Kon Tum Province in the Central
Highlands of Vietnam. It is a medium-sized species with ‘suilla-type’ dentition. A taxonomic review of Murininae from Vietnam
was also conducted based on combined morphological, DNA, and karyological characteristics. Molecular phylogenetic analyses
based on the mitochondrial cytochrome coxidase subunit (COI) gene supported the subfamily Murininae, while the genus Murina
proved to be paraphyletic in relation to the genera Harpiocephalus and Harpiola. Fourteen species of the genus Murina, one species
of Harpiocephalus, and one species of Harpiola are recognized from Vietnam. Murina tiensa is regarded as a junior synonym of
M. harrisoni; strong sexual dimorphism was observed in M. harrisoni. Relations between forearm length and total length of skull
showed different trends among species and sexes. Karyotypes of Murina huttoni, M. cyclotis, M. lorelieae, M. beelzebub, M. feae,
and Harpiola isodon were 2n = 44, FN = 50, while that of Harpiocephalus harpia was 2n = 44, FN = 52.
Key words: DNA barcode, karyotype, morphology, taxonomy, tube-nosed bats

Harpiocephalus has been widely accepted, whereas
the taxonomic rank of Harpiola has been controver-
sial and considered either a separate genus (Tate,
1941; Bhattacharyya, 2002; Kuo et al., 2006) or
a subgenus of Murina (Ellerman and Morrison-
Scott, 1951; Corbet and Hill, 1992; Koopman, 1994;
Simmons, 2005). The phylogenetic reconstruction
of Francis et al. (2010) based on sequences of the
barcoding cytochrome coxidase subunit I (COI)
demonstrated that, contrary to the morphological
distinctiveness, genetic analyses did not separate
Harpiola and Harpiocephalus from the species of
Murina.
Vietnam possesses the highest known species
diversity of the Murininae in the world. Kuznetsov
(2006) and Can et al. (2008) listed six species of
Muri ninae in Vietnam, and Kruskop (2013) recog-
nized 12 Murina, one Harpiola, and one Harpio -
cephalus species from Vietnam after incorporating
information from recent studies (Csorba et al., 2007,
2011; Kruskop and Eger, 2008; Furey et al., 2009).
Son et al. (2015) analysed morphometric variation
of the skull of Vietnamese Murina and documented
considerable interspecific and sexual variation in
size and shape of the skull, possibly reflecting food
adaptations from interactions of sympatric species.
In addition, the existence of sexual dimorphism and
the extent of differences among species were docu-
mented by Son et al. (2015), indicating that the com-
plicated patterns of sexual differences can be the
cause of taxonomic confusion among the species.
During the investigation of recently acquired ma-
terials, we identified a specimen collected from the
Central Highlands of Vietnam that is clearly differ-
ent from all currently known species of Murina. In
this study, we describe a new species based on the
combined morphological, DNA, and karyological
evidences, and review the taxonomic status of all
Vietnamese species belonging to the subfamily
Muri ninae. The species diversification of the sub-
family and distribution patterns of tube-nosed bats
within Vietnam are also discussed.
MATERIALS AND METHODS
Bats were collected through surveys conducted in 43 pro-
tected areas of 28 provinces in Vietnam from 2001 to 2014
(Fig. 1, Table 1, and Appendix I). Three-bank harp traps and
mist nets were set at ground level, frequently across trails,
streams, and rivers in different habitat types in secondary and
primary forests. Mist nets and harp traps were checked every 20
min before dusk from 17:30 until 23:00. Harp traps were left
open until around 06:00. Most captured bats were released at the
capture site after recording standard measurements. Selected
specimens were prepared as voucher specimens. These were
202 N. Truong Son, G. Csorba, V. Tan Tu, V. Dinh Thong, Y. Wu
fixed in 95% ethanol, followed by preservation in 70% ethanol
for about 12 hrs; tissue samples (usually liver or muscle) were
preserved in 95% ethanol.
A total of 252 specimens were examined in this study (Ap -
pendix II). Voucher specimens are kept in the Institute of
Ecology and Biological Resources, Hanoi, Vietnam (IEBR);
Hun garian Natural History Museum, Budapest, Hungary
(HNHM); Harrison Institution, Sevenoaks, United Kingdom
(HZM); Royal Ontario Museum, Toronto, Canada (ROM); Na -
tural History Museum, London, United Kingdom (BMNH);
Zool ogical Museum at Moscow University, Moscow, Russia
(ZMMU); and Muséum National d’Histoire Naturelle, Paris,
France (MNHN). Chinese specimens deposited in the College
of Life Science, Guangzhou University, Guangzhou, China
(IBHG) were also investigated.
Morphological Examination
Terminology of external, skull, and dental morphology fol-
lowed Bates and Harrison (1997), Csorba and Bates (2005),
Csorba et al. (2007, 2011), and Furey et al. (2009). Abbrevia -
tions used for the dental nomenclature were incisor (I/in), canine
(C/c), premolar (P/pn), and molar (M/mn), with premaxillary
and maxillary teeth denoted by uppercase and mandibular teeth
by lowercase letters. Only adult specimens (age was determined
following An thony, 1988) were used for statistical analyses.
The following external measurements were taken to the
nearest 0.1 mm: HB, head and body from the tip of nose to the
FIG. 1. Map of localities for specimens of the subfamily Murininae
from Vietnam. Locality names are given in Appendix I

New species of Murina from Vietnam 203
TABLE 1. Distribution of the subfamily Murininae from Vietnam. Locality numbers correspond to those in Fig. 1 and Appendix I. Species are arranged following the overall skull size
(in decreasing order; species names abbreviated to the first 3-4 letters). Number of provinces where the species occurs is shown in parentheses
Locality Number of Species (Number of provinces)
number Province species H.har M.harr M.leu M.fio M.hut M.cyc H.iso M.lor M.ann M.bee M.wal M.fea M.kon M.ele M.harp M.chr
(10) (8) (1) (7) (9) (23) (2) (1) (10) (4) (4) (18) (1) (10) (1) (2)
1 Lao Cai 5 x x x x x
2 Ha Giang 3 x x x
3 Tuyen Quang 4 x x x x
4, 6 Bac Kan 5 x x x x x
5 Cao Bang 5 x x x x x
7 Thai Nguyen 1 x
8 Lang Son 1 x
9 Quang Ninh 1 x
10 Hai Phong 2 x x
11 Vinh Phuc 4 x x x x
12 Phu Tho 4 x x x x
13, 14, 15 Son La 5 x x x x x
16, 17, 19, 20 Thanh Hoa 5 x x x x x
18 Ninh Binh 2 x x
21, 22 Nghe An 7 x x x x x x x
23 Ha Tinh 1 x
24 Quang Binh 5 x x x x x
25 Quang Tri 6 x x x x x x
26 Thua Thien-Hue 2 x x
27, 28 Quang Nam 5 x x x x x
29, 31 Kon Tum 11 x x x x x x x x x x x
30 Quang Ngai 5 x x x x x
32 Gia Lai 4 x x x x
33 Binh Dinh 1 x
34, 35 Dak Lak 3 x x x
36 Lam Dong 4 x x x x
37 Khanh Hoa 1 x
38 Ninh Thuan 1 x
39, 40 Dong Nai 4 x x x x
41 Binh Phuoc 2 x x
42 Ba Ria-Vung Tau 1 x
43 Kien Giang 1 x

base fundament; FA, forearm length from the extremity of the
elbow to the extremity of the carpus with the wings folded;
T, tail length from the tip of tail to its base fundament; HF, hind
foot from the tip of the longest digit, excluding claw, to the ex-
tremity of the heel, behind the os calcis; TIB, tibia length from
the knee joint to ankle; E, ear length from the lower border of
external auditory meatus where it joins with the body to the tip
of pinna; TRAGUS, tragus length from the lower posterior
emargination to the tip of the tragus; and body mass (in gram).
Craniodental and mandibular measurements were taken to
the nearest 0.01 mm following Son et al. (2015): STOTL, total
length of skull from the anterior rim of the alveolus of the first
upper incisor to the most projecting point of the occipital region;
CCL (condyle-canine length), from the exoccipital condyle to
the most anterior part of the canine; C1C1W, greatest width
across the outer borders of the upper canines; M3M3W, greatest
width across the outer crowns of the last upper molars; ZYW
(zygomatic width), greatest width of the skull across the zygo-
matic arches; MAW (mastoid width), greatest distance across
the mastoid region; IOW (interorbital width), least width of the
interorbital constriction; BCW (braincase width), greatest width
of the braincase; BCH (braincase height), from the basisphenoid
at the level of the hamular processes to the highest part of the
skull, including the sagittal crest (if present); CM3L (maxillary
toothrow length), from the front of upper canine to the back of
the crown of the third molar; CP4L (upper canine-premolar
length), from the front of the upper canine to the back of the
crown of the posterior premolar; ML (mandible length), from
the anterior rim of the alveolus of the first lower incisor to the
most posterior part of the condyle; cm3L (mandibular toothrow
length), from the front of the lower canine to the back of the
crown of the third lower molar; cp4L (lower canine-premolar
length), from the front of the lower canine to the back of the
crown of the posterior premolar; CPH (least height of the coro-
noid process), from the tip of the coronoid process to the apex
of the indentation on the inferior surface of the ramus adjacent
to the angular process.
Multivariate Analyses
To study individual variation in M. harrisoni, we used all
15 craniodental and mandibular measurements to conduct two
PCA analyses with the software PAST (Hammer et al., 2001)
using (1) raw data to assess size factors using the PC1 score that
represents overall size (Barlow et al., 1995; Lindenfors et al.,
2007), and (2) standardized data (raw score/geometric mean) to
assess the shape factor (Jungers et al., 1995) using each of the
PC scores that have eliminated size factors. Both the raw data
and standardized data were log-transformed (Blackith and Rey -
ment, 1971; Reyment, 1971).
DNA Barcoding
Total genomic DNA was extracted from muscle samples
using the DNeasy Blood and Tissue Kit (Qiagen, California)
following the manufacturer’s protocol. A 657 bp fragment of
the mitochondrial COI gene was amplified and sequenced with
the primers LCO1490 and HCO2198 (Hebert et al., 2003). The
polymerase chain reactions (PCR) were carried out in a volume
of 10 mml of HotStarTaq mastermix (Qiagen, California), 5 ml
of water, 2 ml of each primer at 10 pmol/µl and 2 ml of DNA.
PCR condition was: 95°C for five minutes to active Hot Star
Taq; with 40 cycles at 95°C for 30s, 45° for 45s, 72°C for 60s;
and the final extension at 72°C for six minutes. PCR products
were purified using GeneJETTM PCR Purification kit (Fer -
mentas, Canada) and then were sent to Macrogen Inc. (Seoul,
South Korea) for sequencing. Sequences were edited and as-
sembled using Codoncode Alignment ver. 5.0.2 (Codon Code
Cor poration).
The new COI sequence (GenBank accession number
KT820760) was compared to those of 25 identified species of
the subfamily Murininae in the EMBL, GenBank, DDBJ, and
BOLD nucleotide databases (Appendix III). Phylogeny of the
subfamily Murininae was reconstructed using the Bayesian
method. Based on previous studies (Hebert et al., 2003), Myotis
muricola and Kerivoula hardwicki were used as outgroups.
DNA sequences were aligned manually on PhyDE v0.9971
(Müller et al., 2010). The COI dataset represents a total align-
ment of 657 nucleotides and 120 taxa. The best-fitting model of
sequence evolution was selected under jModelTest using the
Akaike information criterion. Bayesian analyses were then con-
ducted using the selected GTR+I+G model on MrBayes v3.2
(Hebert et al., 2003). Post erior probabilities (PP) were calcu-
lated using four independent Markov chains run for 10,000,000
Metropolis-coupled MCMC generations, with tree sampling
every 1,000 generations, and a burn-in of 25%. Mean pairwise
distances were calculated with PAUP version 4b10 (Swofford,
2003) using Kimura’s two-parameter (K2P) model.
Karyotype Analyses
Chromosomal preparations were made with culture cells of
tail bone or ear tissue following the method of Harada and
Yosida (1978). The tissue sample was cultured with Eagle’s
MEM medium supplemented with 12% calf serum, 3% calf
serum and a few non-essential amino acids (l-glutamine, l-se-
rine, sodium pyruvate). Diploid chromosome number (2n) and
fundamental number (FN, as the total number of autosomal
arms) was then calculated.
RESULTS
The Bayesian tree reconstructed from the nu-
cleotide alignment of COI sequences (Fig. 2) sup-
ported the monophyly of the subfamily Murininae
(PP = 1) that included the genera Murina, Harpio -
cephalus, and Harpiola; however, the genus Murina
was proved to be paraphyletic in relation to the gen-
era Harpiocephalus and Harpiola. The species rela-
tionships within Murininae were poorly supported
(PP < 0.7), similar to findings of Francis et al.
(2010) and Tu et al. (2015). Only the following sis-
ter relationships were supported with posterior prob-
abilities > 90%: Harpiocephalus harpia and Murina
sp. nov.(IEBR-M5697; to be described in this pa -
per); M. suilla and M. walstoni; M. cf. cyclotis from
India and M. guilleni; M. cf. cyclotis from India / M.
guilleni / M. fionae / M. cyclotis / M. peninsularis;
M. gracilis and M. recondita; M. eleryi / M. gra-
cilis / M. recondita / M. balaensis; M. harrisoni / M.
huttoni / M. ussuriensis; M. chrysochaetes and
204 N. Truong Son, G. Csorba, V. Tan Tu, V. Dinh Thong, Y. Wu

M. harpioloides; M. leucogaster and M. shuipuensis;
M. hi lgendorfi / M. leucogaster / M. shuipuensis.
Although the monophyly of the genus Murina
was not supported, the morphological distinctness
of the three genera (Harpiocephalus, Harpiola, and
Murina) was clear, and was therefore recognized
as valid.
TAXONOMIC ACCOUNTS
Genus Murina Gray, 1842
Diagnostic characters
Small to medium-sized bats with tubular nostrils
and dense woolly pelage. I2/3 C1/1 P2/2 M3/3 = 34.
Height of upper incisors is distinctly less than that of
the corresponding canines. I3 is not contact with C.
M3 is developed.
Taxonomic notes
Two morphogroups, namely ‘suilla-group’ and
‘cyclotis-group’, differing in the relative size and
position of the upper incisors, canines, premolars,
and ratio of upper canine and second upper premolar
have been widely accepted (Corbet and Hill, 1992;
Koopman, 1994; Csorba and Bates, 2005; Csorba et
al., 2007; Furey et al., 2009; Kuo et al., 2009;
Soisook et al., 2013b; Son et al., 2015). As an im-
portant characteristic, crown area of the upper
canine is less than that of P4 in the ‘suilla-group’,
New species of Murina from Vietnam 205
FIG. 2. Bayesian tree of the subfamily Murininae reconstructed based on 120 sequences of the mitochondrial cytochrome coxidase
subunit I (COI) gene, with posterior probabilities (PP) values. Murina feae and M. beelzebub are based on data from Francis
et al. (2012), collectively reported as M. feae

and is equal to or exceeds that of P4 in the ‘cyclotis-
group’. Although we recognize the usefulness of
these characteristics for identification of the species,
these morphogroups do not represent separate phy-
logenetic lineages. Therefore, morphological char-
acteristics are referred hereafter as ‘suilla-type’ den-
tition and ‘cyclotis-type’ dentition without using the
term ‘group’.
Murina kontumensis Son, Csorba, Tu
and Motokawa, sp. nov. (Fig. 3)
Holotype
IEBR-M5697, field number B20140920.14.
Adult female, skin and skull, collected from Ngoc
Linh Nature Reserve, Xop commune, Dak Glei dis-
trict, Kon Tum province, Vietnam (15°05’30N,
107°51’35E; Fig. 1, No. 22), 1,780 m a.s.l.; collect -
ed by Vuong Tuan Tu, Masaharu Moto kawa and
Nguyen Truong Son on 20 September 2014. Meas -
urements (in mm) are as follows, HB: 40.00, FA:
32.28, T: 38.50, HF: 7.60, TIB: 16.20, E: 18.70,
TRA GUS: 8.74, STOTL: 14.98, CCL: 13.29,
C1C1W: 3.66, M3M3W: 5.30, ZYW: 8.53, MAW:
7.55, IOW: 3.94, BCW: 7.32, BCH: 6.65, CM3L:
5.01, CP4L: 2.28, ML: 10.10, cm3L: 5.56, cp4L:
2.18, CPH: 3.47, and body mass: 5 g (Table 3).
Diagnosis
Medium-sized, with ‘suilla-type’ dentition. Head
is with a distinct facemask, dark brown around the
eyes, with a contrasting whitish collar around the
neck. General impression of dorsal pelage is brown-
ish grey with scattered golden hairs; ventral fur is
light brown. Plagiopatagium is attached to the base
of the first claw of the outer toe. Skull is domed, and
sagittal and lambdoid crests are evident. I2 clearly is
anterior to I3. Crown area of P2 is one-half of P4;
crown area of upper canine is less than that of P4;
P2 is antero-posteriorly compressed. The mesostyles
of M1 and M2 are well developed.
Description of the holotype
Medium-sized, with ‘suilla-type’ dentition (Figs.
3 and 4, Tables 2 and 3). FA 32.28, STOTL 14.98,
and body mass 5.0 g. On the dorsum, the basal por-
tions of the hairs are dark brown and followed by
a light brown section before terminating in a dis-
tinctly darker brownish-grey tip. Guard hairs with
shiny golden tips are scattered on the head and
behind the ears, and more sparsely over the back.
Ventrally, hairs are grey for the proximal one-third,
whereas the remaining upper portion is light brown.
Head is with a distinct facemask, dark brown
around the eyes, with a contrasting whitish collar
around the neck. Ear is 18.70 mm long and 11.25
mm wide, having a slight emargination along its
posterior border; the tragus is 8.74 mm in length, the
widest point near the base is 2.10 mm. The upper
surface of hind limbs, feet and uropatagium are
densely covered in short, uniformly shiny golden
brown hairs. The ventral surface of the uropatagium
is covered in uniformly golden white hairs, some of
which are also present on the plagiopatagium adja-
cent to the body. Plagiopatagium attached to base
of the claw of the first toe.
Skull and dentition (Figs. 3, 5, 6, and Table 3).
Skull is medium-sized. The rostrum is slightly short-
ened and domed, and from the top of skull to nasal
bone, is relatively steep. Braincase is domed. Zy go -
ma is not strong. Length of second upper premolar
is half the length of the canine. The upper tooth row
is convergent anteriorly (C1C1W/M3M3W, 0.70).
One half of I2 is obscured in the lateral view. I3 is
not in contact with upper C. The basal area of upper
C is less than that of P4, whereas its height is greater.
In the occlusal view, upper C is relatively circular. P2
is strongly compressed, wider than long, with a small
cusp on the anterior inner cingulum. Its basal area
and height are approximately half or slightly less
than that of P4. Mesostyles of M1 and M2 are well
developed comparable in height to metastyle and
parastyle, giving a distinct W-shape to the surface.
Lower c exceeds p4 in height and is equal or
greater in basal area; p2 has less than half the basal
area of p4 and attains more than two-thirds of its
height. Talonids of m1 and m2 are wider than their
trigonids. Hypoconid of m2 distinctly exceeds its
entoconid in height, whereas entoconid of m2 is
lower than hypoconid and metaconid.
Etymology
The specific epithet kontumensis refers to
Kon Tum Province, from where the holotype was
collected.
COI sequences
The Bayesian tree reconstructed from the nu-
cleotide alignment of COI sequences determined
that M. kontumensis sp. nov. was distantly related to
25 examined species of the subfamily Murininae,
forming sister relationship with H. harpia (Fig. 2).
The mean pairwise distances between M. kontumen-
sis sp. nov. and other species of Murininae varied
between 16.2% to 20.5%, supportive as representing
a distinct species.
206 N. Truong Son, G. Csorba, V. Tan Tu, V. Dinh Thong, Y. Wu

New species of Murina from Vietnam 207
Comparative material
Murina annamitica: IEBR-M3167, M2181,
M4131, M2997, M3650, HNHM TJO 423, HNHM
BHH 12; M: beelzebub: IEBR-M5760/HNHM.
2007.50.6; M. chrysochaetes: ZMMU S186699;
M. cyclotis: IEBR-M3128, M3316, M3646; M. feae:
IEBR-M4214, M3154, HNHM QT 003, HNHM
PHV 23; M. eleryi: HNHM. 2007. 51.1, HNHM.
2007.28.2, IEBR-M4070; M. gracilis: HNHM 2005.
1.1; M. harpioloides: ZMMU-S-173401, IEBR-
M5718; M. lorelieae: ROM 116171; MNHN 2013-
1078, VN11-1120, VN11-1223, VN11-1161, IEBR-
M5648, M5651, M5656, M5662; HNHM. B2014
0915.5, 20140915.7; M. recondita: HNHM 2005.
36.1; M. suilla: HNHM 2000.13.2; M. walstoni:
HNHM 2010.20.1, IEBR-M4592, M2920, M2479.
Comparison with other taxa
Murina kontumensis sp. nov. differs from all
known Southeast Asian species having ‘suilla-type’
dentition by the combination of the brownish dorsal
pelage, scattered shiny golden guard hairs on the
head, the dorsum and the dark brown colouration
around the eyes, a contrasting whitish collar around
the neck, more domed skull, and narrow nasal sinus
(Figs. 4–6).
Compared to other dorsally brownish-red-
dish species with golden-tipped guard hairs,
FIG. 3. Murina kontumensis sp. nov. (holotype, IEBR-M5697): A — face; B — ear and tragus; C — dorsal pelage; D — ventral fur;
E — dorsal fur; F — upper surface of uropatagium; G — ventral surface of uropatagium; H — plagiopatagium and the first claw;
I — dorsal view of skull; J — ventral view of skull; K — lateral view of mandible

208 N. Truong Son, G. Csorba, V. Tan Tu, V. Dinh Thong, Y. Wu
M. kontumensis sp. nov. further differs from M. ele -
ryi by its brown-tipped ventral fur (white tipped in
M. eleryi), and from M. gracilis by its well-furred
uropatagium (almost naked in M. gracilis).
Compared with the dorsally reddish and brown-
ish species with no shiny guard hairs, M. kontumen-
sis sp. nov. is smaller than M. lorelieae (Tables 2 and
4), M. recondita (FA mean 30.9, GLS mean 15.48 —
FIG. 4. Face (left), ventral fur (middle), and dorsal fur (right) of specimens from species in the subfamily Murininae from Vietnam:
A — M. kontumensis sp. nov. (holotype, IEBR-M5697); B — M. harrisoni (IEBR-M3299); C — M. fionae (IEBR-M3080);
D — M. huttoni (IEBR-M5693); E — M. cyclotis (IEBR-M4223); F — M. lorelieae (IEBR-M5656); G — M. annamitica
(IEBR-M3034); H — M. beelzebub (IEBR-M3904); I — M. walstoni (IEBR-M4592); J — M. feae (IEBR-M3264)); K — M. eleryi
(IEBR-M4070); L — M. harpioloides (IEBR-M5806); M — M. chrysochaetes (ROM MAM 116181); N — H. harpia (IEBR-M6037
left, IEBR-M5661 middle and right); O — H. isodon (IEBR-M5436)

New species of Murina from Vietnam 209
TABLE 2. Mean, range, and sample size (in italics) for external measurements (mm) and body mass (g) of bats of the subfamily Murininae from Vietnam. The measurements for
M. leucogaster follow Hendrichsen et al. (2001)
Species Sex FA HB T HF E TIB Body mass
0Range 0Range 0Range 0Range 0Range 0Range 0Range
M. kontumensis ♀ 32.3 – 40.0 – 38.5 – 7.6 – 18.7 – 16.2 – 5.0 –
sp. nov.
M. leucogaster ♀ 41.8 – – – 40.0 – 9.8 – 14.2 – 18.8 – 8.8 –
M. harrisoni ♂ 35.30 34.0–36.7 11 44.27 38.9–50.0 339.50 35.8–45.5 38.20 7.4–9.0 314.60 13.5–15.5 319.33 19.0–19.5 38.4 –
♀ 38.18 35.6–40.1 10 40.3 44.9 40.1 41.7 8.1 10.2 15.4 18.8 18.6 21.0 10.5 –
M. fionae ♂ 35.32 34.2–36.4 645.62 42.1–48.2 639.95 37.0–44.5 68.17 6.8–9.3 614.32 13.7–14.6 619.20 18.5–20.8 67.87 7.5–9.0 6
♀ 37.9 – 51.3 – 41.7 – 9.1 – 16.0 – 20.7 – 12.0 –
M. huttoni ♂ 33.36 32.0–36.0 846.21 44.0–50.0 737.57 31.0–41.5 77.30 6.6–8.6 716.32 14.0–17.0 816.71 16.7–16.7 85.89 5.3–6.3 5
♀ 34.34 33.1–35.5 10 46.26 40.0–53.0 10 36.02 32.0–42.5 10 7.89 7.0–8.9 10 16.30 15.9–17.0 10 16.74 16.0–17.7 10 6.36 5.3–8.0 10
M. cyclotis ♂ 30.33 28.9–32.0 33 40.98 38.0–48.0 33 35.13 31.8–38.7 33 7.27 6.3–8.5 33 14.55 12.3–17.0 33 17.01 15.9–17.8 36 5.01 4.1–6.0 19
♀ 34.37 32.1–36.3 26 44.07 40.0–51.0 26 38.33 32.1–43.5 26 7.53 6.3–8.60 26 15.01 12.5–18.6 26 18.27 15.0–20.1 26 6.82 4.7–8.2 16
M. lorelieae ♂ 33.72 33.0–34.5 739.14 37.0–41.0 735.00 31.0–38.0 76.71 6.0–7.5 715.10 14.0–16.0 733.72 33.0–34.5 74.87 4.2–6.6 x
♀ 35.25 34-9–35.6 344.40 43.3–45.5 339.70 38.4–41.0 38.45 8.4–8.5 315.80 15.6–16.0 319.20 19.1–19.3 36.00 5.5–6. 3
M. annamitica ♂ 30.91 29.4–32.1 11 40.32 36.2–48.0 10 35.72 32.5–38.8 10 6.74 5.9–8.1 10 12.86 11.9–14.2 10 16.58 15.4–17.4 11 5.66 4.3–7. 8
♀ 32.12 27.0–34.6 10 43.70 38.2–55.0 10 36.80 32.4–40.3 10 7.15 6.2–8.1 10 13.09 11.9–15.0 10 17.19 16.0–18.0 10 6.75 5.3–8.0 10
M. beelzebub ♂ 34.43 34.4–34.5 342.67 40.0–44.0 338.90 36.8–41.4 37.13 6.6–8.0 314.13 14.1–14.2 318.57 18.1–19.4 33.0 5.5
♀ 36.45 36.0–37.3 544.37 41.6–49.0 539.60 33.0–44.8 56.80 5.5–8.0 513.73 13.2–14.0 518.84 18.0–19.6 53.0 5.5
M. walstoni ♂ 33.16 32.7–34.1 335.41 34.8–35.9 330.93 29.8–32.5 36.52 6.1–7.0 313.01 12.5–14.0 314.08 13.4–15.0 34.40 4.2–4.6 3
♀ 33.40 32.6–33.5 340.22 35.2–43.5 328.79 26.6–30.0 36.36 6.1–6.5 312.90 12.1–14.1 314.66 13.9–15.6 34.6 –
M. feae ♂ 30.12 27.5–33.4 17 38.29 33.7–43.0 13 34.89 31.4–39.5 13 6.86 6.0–8.4 14 12.99 11.5–14.3 14 16.57 15.7–18.1 17 4.22 3.5–5.3) 8
♀ 31.40 28.1–34.3 16 38.69 32.8–43.5 16 35.46 30.0–41.6 16 6.83 6.1–8.0 16 12.90 11.0–15.0 16 16.95 15.6–17.8 16 4.20 3.9–4.4 6
M. eleryi ♂ 28.23 27.3–29.4 733.19 31.5–36.2 329.52 26.5–32.3 76.40 5.2–7.7 712.20 11.5–13.3 714.05 13.0–15.1 74.03 2.5–5.0 6
♀ 29.87 28.6–31.3 835.8 39.0 29.87 27.3–32.1 86.75 6.0–7.4 812.57 11.7–13.3 813.96 13.2–15.0 84.79 4.0–5.5 7
M. harpioloides ♀ 29.7 29.8 34.5 35.0 27.0 30.5 6.5 – 12.3 – 13.0 14.4 4.2 –
M. chrysochaetes ♀ 28.6 29.8 40.0 41.0 24.0 26.0 5.5 5.6 12.0 12.6 12.6 12.7 4.0 4.4
H. harpia ♂ 49.4 49.5 55.0 57.0 40.0 50.0 11.0 11.2 17.0 18.0 22.0 24.0 16.0 24.0
♀ 52.8 50.9–54.9 359.7 59.0–61.0 348.6 47.7–50.0 311.5 11.0–12.0 317.7 17.0–18.0 322.7 22.0–23.5 327.1 25.0–30.2 3
H. isodon ♂–––– ––––––––––
♀ 37.3 – 44.0 – 36.0 – 7.5 – 15.0 – 15.0 – 7.0 –

Parameter
M. kontumensis M. lorelieae M. harrisoni H. harpia H. isodon
sp. nov.
♀ (n = 1) ♂♂ (n = 7) ♀♀ (n = 3) ♂♂ (n = 10) ♀♀ (n = 11) ♂♂ (n = 3) ♀♀ (n = 2) ♀ (n = 1)
STOTL 14.98 16.12 ± 0.19 (15.83–16.39) 16.54 ± 0.29 (16.33–16.74) 17.85 ± 0.46 (16.95–118.39) 18.64 ± 0.63 (17.46–19.43) 22.26 ± 0.07 (21.45–23.07) 22.82, 23.25 17.09
CCL 13.29 13.96 ± 0.20 (13.55–14.14) 13.96 ± 0.20 (13.55–14.14) 15.92 ± 0.25 (15.46–16.25) 16.62 ± 0.49 (15.96–17.16) 19.42 ± 0.73 (18.90–19.42) 20.01, 20.26 14.95
C1C1W 3.66 3.86 ± 0.06 (3.74–3.92) 3.86 ± 0.06 (3.74–3.92) 4.64 ± 0.21 (4.16–4.94) 5.00 ± 0.19 (4.66–5.31) 6.74 ± 0.41 (6.45–7.03) 6.74, 7.17 4.26
M3M3W 5.30 5.43 ± 0.09 (5.29–5.58) 5.43 ± 0.09 (5.28–5.58) 5.81 ± 0.18 (5.63–5.98) 6.12 ± 0.21 (5.78–6.44) 6.57 ± 0.46 (6.24–6.89) 7.23, 7.75 5.65
ZYW 8.53 8.97 ± 0.14 (8.78–9.17) 9.25 ± 0.28 (9.05–9.45) 10.32 ± 0.25 (9.97–10.62) 9.29 ± 0.38 (10.13–11.47) 13.73 ± 1.15 (12.92–14.54) 14.48, 15.09 9.61
MAW 7.55 7.68 ± 0.10 (7.76–7.89) 7.92 ± 0.22 (7.76–8.07) 8.85 ± 0.29 (8.31–9.31) 9.29 ± 0.38 (8.34–9.61) 11.23 ± 0.74 (10.70–11.75) 11.45, 11.98 8.12
IOW 3.94 4.21 ± 0.04 (4.17–4.29) 4.29 ± 0.15 (4.17–4.39) 4.36 ± 0.12 (4.23–4.58) 4.48 ± 0.09 (4.30–4.58) 5.57 ± 0.21 (5.42–5.72) 5.57, 6.05 5.12
BCW 7.32 7.46 ± 0.11 (7.33–7.62) 7.68 ± 0.32 (7.45–7.90) 7.81 ± 0.22 (7.46–8.11) 7.94 ± 0.29 (7.76–8.22) 9.42 ± 0.34 (9.18–9.66) 9.60, 10.23 7.72
BCH 6.65 6.65 ± 0.26 (6.20–6.94) 6.53 ± 0.08 (6.45–6.54) 6.49 ± 0.18 (6.16–6.78) 6.74 ± 0.29 (6.22–7.23) 9.51 ± 0.36 (9.25–9.76) 9.74, 10.12 7.77
CM3L 5.01 5.30 ± 0.06 (5.21–5.40) 5.55 ± 0.13 (5.46–5.64) 6.11 ± 0.17 (5.82–6.31) 6.53 ± 0.25 (6.29–7.14) 6.95 ± 0.19 (6.81–7.08) 7.08, 7.12 5.61
CP4L 2.28 2.52 ± 0.06 (2.40–2.56) 2.82 ± 0.05 (2.78–2.85) 3.04 ± 0.11 (2.84–3.19) 3.25 ± 0.08 (3.16–3.37) 4.19 ± 0.09 (4.12–4.25) 4.05, 4.33 2.74
ML 10.10 10.53 ± 0.18 (10.21–10.78) 11.10 ± 0.13 (10.87–11.32) 12.44 ± 0.34 (11.95–12.89) 13.08 ± 0.37 (12.55–13.62) 15.43 ± 0.45 (15.11–15.74) 16.28, 16.45 11.63
cm3L 5.56 5.74 ± 0.06 (5.62–5.81) 6.04 ± 0.22 (5.88–6.19) 6.69 ± 0.21 (6.30–6.95) 7.07 ± 0.19 (6.81–7.43) 7.84 ± 0.26 (7.65–8.02) 7.83, 8.10 5.91
cp4L 2.18 2.37 ± 0.04 (2.31–2.42) 2.59 ± 0.13 (2.50–2.68) 3.03 ± 0.13 (2.84–3.22) 3.19 ± 0.09 (3.07–3.36) 4.20 ± 0.16 (4.08–4.331) 4.13, 4.24 2.47
CPH 3.47 3.67 ± 0.17 (3.50–4.00) 3.95 ± 0.01 (3.94–3.96) 4.70 ± 0.28 (4.26–5.04) 5.14 ± 0.27 (4.61–5.52) 8.43 ± 0.50 (8.07–8.78) 8.80, 9.66 3.74
210 N. Truong Son, G. Csorba, V. Tan Tu, V. Dinh Thong, Y. Wu
Kuo et al., 2009), M. ryukyuana (FA mean 36.83,
GL 18.65 — Maeda and Matsumura, 1998), M. shu -
i puensis (FA 30.55, GLS 15.90 — Eger and Lim,
2011), and M. walstoni (Tables 2 and 4), and slightly
larger than M. suilla (FA mean 30.70, GTL 14.93 —
Hill, 1964; FA mean 30.6, STOTL mean 14.58 —
Furey et al., 2009). Murina kontumensis sp. nov.
also differs from these species in having golden-
tipped guard hairs; from M. shuipuensis by its pla-
giopatagium, which attaches to the base of the claw
(compared to the base of the toe); from M. recondita
by its short and bluntly rounded tragus (Fig. 3) (tra-
gus reaches at least half the length of pinna and is
sharply pointed in M. kontumensis sp. nov.).
Compared with species having predominantly
greyish-black pelage, M. kontumensis sp. nov. is
smaller than M. beelzebub, and similar in size to
M. feae (Tables 2 and 4) and M. jaintiana (FA 29.1–
31.5, STOTL 14.75–15.25 — Ruedi et al., 2012);
M. kontumensis sp. nov. has a distinct sagittal crest
and developed mesostyles on M2 and M3 (reduced
in M. beelzebub, M. jaintiana and M. feae).
The general size of the skull and dentition of
M. kontumensis (Table 4) is similar to those of
M. chrysochaetes (Eger and Lim, 2011), M. harpio-
loides (Kruskop and Eger, 2008; IEBR-M5718),
M. eleryi (Furey et al., 2009), M. feae (Thomas,
1891), and M. walstoni (Csorba et al., 2011).
However, CM3L (Table 4) could be useful to sepa-
rate M. kontumensis from the latter species; M. kon-
tumensis sp. nov. (5.01) is larger than M. aurata
(4.35–4.60 — Maeda, 1980; Furey et al., 2009;
Kuo et al., 2009; Csorba et al., 2011), M. balaensis
(4.66), M. eleryi (4.48–4.89), M. harpioloides
(4.68–4.78), and similar to M. feae (4.89–5.38),
M. gracilis (4.92–5.05 — Kuo et al., 2009), M. re-
condita (4.91–5.10 — Kuo et al., 2009), M. shui -
puensis (5.23: Eger and Lim, 2011), M. suilla (4.54–
5.08 — Hill, 1964; Kuo et al., 2009; Csorba et al.,
2011), and M. ussu riensis (4.95–5.20 — Maeda,
1980), and smaller than M. walstoni (6.66–7.14) and
M. beelzebub (5.47–5.70).
In M. kontumensis sp. nov. the ratio of braincase
height (BCH) and braincase width (BCW) is 90.8%,
whereas all other species of Murina from Vietnam,
except M. fionae and M. cyclotis, have ratios less
than 90% (Table 4). In comparison of BCH with the
other species in Asia, BCH in M. kontumensis sp.
nov. (6.65) is greater than M. balaensis (5.33 — Soi -
sook et al., 2013a), M. tubinaris (5.55–5.86 —
Csorba et al., 2011), M. pluvialis (6.43 — Ruedi et
al., 2012), and M. jaintiana (5.96–6.17 — Ruedi et
al., 2012).
TABLE 3. Craniodental measurements (0± SE, range; in mm) of M. kontumensis sp. nov., M. lorelieae, M. harrisoni, H. harpia, and H. isodon (specimens of M. harrisoni from Vietnam,
Cambodia, China, and Thailand, and those for other species from Vietnam)

New species of Murina from Vietnam 211
FIG. 5. Left lateral view of skull of specimens from species in the subfamily Murininae in Vietnam: A — M. kontumensis sp. nov.
(holotype, IEBR-M5697); B — M. leucogaster (GZU 10122); C — M. harrisoni (IEBR-M3299); D — M. fionae (IEBR-M3635);
E — M. huttoni (IEBR-M5693); F — M. cyclotis (IEBR-M4223); G — M. lorelieae (IEBR-M5656); H — M. annamitica (IEBR-
M3639); I — M. beelzebub (IEBR-M5645); J — M. walstoni (IEBR-VTTu 15-0033); K — M. feae (IEBR-M5719); L — M. eleryi
(IEBR-M5718); M — M. harpioloides (ZMMU S173401); N — M. chrysochaetes (ZMMU S186699); O — H. harpia (IEBR-M422);
P — H. isodon (IEBR-M5436)

212 N. Truong Son, G. Csorba, V. Tan Tu, V. Dinh Thong, Y. Wu
FIG. 6. Lateral and crown view of upper (left) and lower (right) dentition of specimens from species in the subfamily Murininae in
Vietnam: A — M. kontumensis sp. nov. (holotype, IEBR-M5697); B — M. leucogaster (IBHG 10122); C — M. harrisoni (IEBR-
M3299); D — M. fionae (IEBR-M3635); E — M. huttoni (IEBR-M5693); F — M. cyclotis (IEBR-M4223); G — M. lorelieae (IEBR
M5656); H — M. annamitica (IEBR-M3639); I — M. beelzebub (IEBR-M5645); J — M. walstoni (IEBR VTTu 15-00033); K —
M. feae (IEBR-M5719); L — M. eleryi (IEBR-M4070); M — M. harpioloides (ZMMU S173401); N — M. chrysochaetes
(ZMMU S186699); O — H. harpia (IEBR-M422); P — H. isodon (IEBR-M5436)

New species of Murina from Vietnam 213
TABLE 4. Morphological comparison among species of the subfamily Murininae. Sample sizes in parentheses
Species Sex STOTL (mm) CM3L (mm) ML (mm) BCH/BCW IOW/BCW Dentition Braincase Sagittal crest Lambdoid crest M1/M2
(%) (%) mesostyles
M. kontumensis sp. nov. ♂ – – – – – suilla-type Well domed Developed Developed Well developed
♀ 14.98 (1) 5.61 (1) 10.1 (1) 90.8 53.8
M. leucogaster ♂ 19.27 (1) 6.27 (1) 13.17 (1) 84.7 65.7 suilla-type Domed Weakly developed Weak Weakly developed
♀ 18.49 (1) 6.33 (1) 13.32 (1) 80.8 61.1
M. harrisoni ♂ 16.95–18.39 (12) 5.82–6.31 (12) 11.95–12.89 (12) 77.9–89.1 53.5–58.3 cyclotis-type Flat Well developed Well developed Weakly developed
♀ 17.46–19.72 (11) 6.29–7.14 (11) 12.55-13.75 (11) 79.0–92.9 54.2–58.6
M. fionae ♂ 17.86–18.23 (6) 6.10–6.38 (6) 12.05–12.44 (6) 84.1–94.4 54.3–58.3 cyclotis-type Well domed Well developed Weakly developed Absent
♀ 19.08 (1) 6.39 (1) 12.85 (1) 91.5 57.0
M. huttoni ♂ 16.61–18.52 (14) 5.56–6.23 (14) 11.35–12.30 (14) 80.1–93.1 54.7–60.3 cyclotis-type Flat Weakly developed Weakly developed Well developed
♀ 16.61–18.81 (14) 5.66–6.30 (14) 11.59–12.70 (14) 79.8–87.9 55.0–59.6
M. cyclotis ♂ 16.08–17.66 (40) 5.16–5.59 (40) 10.44–11.87 77.7–96.2 51.7–57.8 cyclotis-type Domed Weakly developed Weakly developed Absent
♀ 16.70–18.08 (33) 5.55–5.89 (33) 10.90–12.34 (33) 80.5–96.4 51.7–58.3
M. lorelieae ♂ 15.83–16.39 (7) 5.21–5.40 (7) 10.21–10.78 (7) 86.8–94.1 55.5–57.0 suilla-type Domed Absent Weak Well developed
♀ 16.33–16.74 (3) 5.46–5.64 (3) 10.79–11.32 (3) 82.8–89.8 55.6–57.8
M. annamitica ♂ 15.63–16.47 (11) 5.19–5.60 (11) 10.13–10.98 (11) 82.4–91.6 53.6–58.9 cyclotis-type Domed Weakly developed Weak Well developed
♀ 16.40–17.16 (11) 5.23–5.69 (11) 10.69–11.47 (11) 80.5–90.5 54.8–60.5
M. beelzebub ♂ 16.40–16.69 (4) 5.27–5.54 (4) 10.73–11.11 (4) 80.8–83.3 56.5–59.0 suilla-type Domed Absent Weak Weakly developed
♀ 16.73–16.99 (6) 5.40–5.70 (6) 10.88–11.48 (6) 79.8–83.7 59.0–61.1
M. walstoni ♂ 15.39–15.73 (3) 5.34–5.39 (3) 10.62–10.74 (3) 79.3–90.9 54.7–57.0 suilla-type Domed Well developed Well developed Developed
♀ 15.15–15.92 (6) 5.27–5.48 (6) 10.40–10.95 (6) 86.1–94.5 56.7–62.6
M. feae ♂ 14.91–16.30 (20) 4.62–5.25 (20) 9.63–10.64 (20) 76.4–91.1 53.6–62.0 suilla-type Domed Absent Weak Weak
♀ 15.13–16.11 (18) 4.89–5.47 (18) 9.80–10.86 (18) 76.4–88.0 55.2–63.3
M. eleryi ♂ 13.79–15.18 (9) 4.53–4.79 (8) 9.25–9.60 (8) 79.5–84.2 56.7–62.6 suilla-type Domed Absent Weak Well developed
♀ 14.21–15.15 (10) 4.48–4.89 (10) 9.28–9.97 (10) 76.6–87.8 58.4–60.7
M. harpioloides ♂ 14.62 (1) 4.70 (1) 9.52 (1) 90.3 58.6 suilla-type Domed Absent Weak Weak
♀ 14.53 (1) 4.68 (1) 9.31 (1) 88.2 57.0
M. chrysochaetes ♂– – – – –suilla-type Domed Absent Weak Reduced
♀ 14,57, 14.72 (2) 4.66, 5.44 (2) 9.30, 9.94 (2) 87.0, 89.0 (2) 57.3, 59.5
H. harpia ♂ 21.45, 23.07 (2) 6.81, 7.09 (2) 15.11, 15.74(2) 100.8, 101.0 59.0, 59.2 – Domed Well developed Well developed Absent
♀ 22.82–23.25 (3) 7.08–7.12 (3) 16.28–16.45 (3) 97.6, 105.4 55.8, 59.6
H. isodon ♂ – – – – – – Domed Absent Weakly developed Weak
♀ 17.09 (1) 5.61 (1) 11.63 (1) 100.6 66.3

214 N. Truong Son, G. Csorba, V. Tan Tu, V. Dinh Thong, Y. Wu
The ratio of interorbital width to braincase width
(IOW:BCW) for M. kontumensis sp. nov. was
53.8%, while other Murina species, except M. cyclo-
tis, had more than 54% (Table 4). This is related to
the narrower nasal sinus of M. kontumensis sp. nov.
than that in the other Murina species.
Karyotype
Karyotype (Fig. 7A and Table 5) of the holotype
was 2n = 44, FN = 50. Autosomes consisted of three
large metacentric pairs, one small submetacentric
pair, and 17 medium-sized to small acrocentric pairs,
gradually decreasing in size. The X chromosome
was identified as a large submetacentric element.
Distribution and ecological notes
Holotype was collected by mist nets set along
a stream in relatively undisturbed primary forest at
1,780 m a.s.l. The nearby Ngoc Linh Mountain (ca.
2,598 m a.s.l.) is situated at the northwestern margin
of Kon Tum province, and is the highest point of the
Truong Son (Annamite) Range in Central Highlands
of Vietnam. The primary vegetation at the collection
site was moist evergreen forest, with a mixture of
abundant bananas and bamboos on the slopes.
Ngoc Linh is split into two nature reserves: one
in Quang Nam Province and another in Kon Tum
Province. Ngoc Linh (Kon Tum) Nature Reserve
was established in 1986 for the conservation of
the evergreen forest and covers 41,240 ha in Dak
Glei and Dac To districts. This part of the Central
High lands includes a complex mosaic of vol-
canic basalts, granites and sedimentary substrates
with wet evergreen hardwood and conifer forests.
A total of 55 mammal species were recorded for
Ngoc Linh (Kon Tum) Nature Reserve (Trai et al.,
FIG. 7. Karyotypes of specimens from species in the subfamily Murininae from Vietnam: A — M. kontumensis sp. nov. (holotype,
IEBR-M5697); B — M. huttoni (IEBR-M5407); C — M. cyclotis (IEBR-M4071); D — M. lorelieae (HNHM B20140915.7);
E —M. beelzebub (IEBR-M4842); F — M. feae (IEBR-M4214); G — H. harpia (IEBR-M5661); H — H. isodon (IEBR-M5436)

New species of Murina from Vietnam 215
1999, 2000; Tor doff et al., 2000; Abramov et al.,
2006).
Murina leucogaster Milne-Edwards, 1872
Murina leucogaster Milne-Edwards, 1872: 252; Type locality:
Moupin, Sichuan, China.
Murina leucogaster: Hendrichsen et al., 2001: 102.
Distribution
Northeast India, Nepal, western Thailand, Viet -
nam, and southern China (Milne-Edwards, 1872;
Tate, 1941; Hill, 1964; Corbet and Hill, 1992; Bates
and Harrison, 1997; Hendrichsen et al., 2001;
Simmons, 2005; Smith and Xie, 2008). In Vietnam
(Fig. 1): Nghe An (Pu Mat NP) [22] (Hendrichsen et
al., 2001).
Diagnostic descriptions
Large species with ‘suilla-type’ dentition (Table
4). Wing membranes are greyish-brown, the muzzle
is dark, and the ears are relatively narrow and short.
There is a basal notch in the outer margin of ears.
Pelage is thick and woolly, with hairs covering the
interfemoral membrane and toes. The dorsum is fer-
ruginous red with dark brown roots; ventrum is pale
yellow with dark grey roots on the flanks. Wing is
attached at the base of the toe claw and the tail is
hairy.
Skull and dentition (Figs. 5 and 6). Rostrum is
not inflated and the braincase is domed. Sagittal and
lambdoid crests are relatively weak. I2 is partly an-
terior to I3. C is slightly higher than or subequal to
P4. C basal area is smaller than that of P4. P2 is
small with a basal area one-third that of P4. M1 and
M2 mesostyles are weakly developed. In the
mandible, talonids on the lower molars are smaller
than the corresponding trigonids. We measured a fe-
male skull (HZM 1.31758) from Vietnam and a male
skull (IBHG 10122) from Sichuan Province, China
(Table 4).
Taxonomic note
Milne-Edwards (1871) originally described
M. leu cogaster from Moupin, Szechwan, China.
In Vietnam, Hendrichsen et al. (2001) only record-
ed one specimen from Pu Mat National Park,
Nghe An Province and no further specimens were
found.
Murina harrisoni Csorba and Bates, 2005
Murina huttoni: Hendrichsen et al., 2001: 103 (part).
Murina harrisoni Csorba and Bates, 2005: 2; Type locality:
O Tuk Chehn, Kirirom National Park, Kompong Speu Prov -
ince, Cambodia; Wu et al., 2010: 277; Francis and Eger,
2012: 29; Thomas et al., 2013: 231.
Murina tiensa Csorba, Thong, Bates and Furey, 2007: 3;
Type locality: An Tinh commune, Na Ri district of Kim
Hy Na ture Reserve, Bac Kan Province, Vietnam, about
750 m a.s.l.
Distribution
Vietnam, Laos, Cambodia, Myanmar, Thailand,
southern China including Hainan Island (Csorba and
Bates, 2005; Csorba et al., 2007; Wu et al., 2010;
Francis and Eger, 2012).
In Vietnam (Fig. 1): Son La (Co Ma and Thuan
Chau district) [13], Phu Tho (Xuan Son NP) [12],
Bac Kan (Kim Hy NR) [6], Hai Phong (Cat Ba NP)
[10], Vinh Phuc (Tam Dao NP) [11], Thanh Hoa
(Xuan Lien) [19], Nghe An (Pu Mat NP) [22], Dak
Lak (Yok Don NP) [34] (This study; Csorba et al.,
2007; Thong et al., 2011; Francis and Eger, 2012).
Description
Large species with ‘cyclotis-type’ dentition (Fig.
4, Tables 2, 3, and 4). The ear is round and large. The
hairs of the dorsal fur are pale-based, light, yellow-
ish-red, gradually darkening towards the tip. The
ventral pelage is uncoloured whitish or light grey.
Plagiopatagium attached to the base of the toe claw.
Skull and dentition (Figs. 5 and 6). Rostrum not
inflated and the braincase is flat. Sagittal and lamb-
doid crests are well developed. I2 is lateral to I3.
C is much higher than P4. C basal area equals or
slightly larger than that of P4. P2 height is subequal
to P4; its basal area is two-third or almost equals that
of P4. M1 and M2 mesostyles are weakly devel-
oped. Lower canines are well developed. The height
of p2 is about equal to that of the p4; m1 and
m2 have well-developed talonids, and have a well-
defined hypoconids and entoconids.
Taxonomic note
Csorba and Bates (2005) described M. harrisoni
as a new species from Cambodia. Subsequently,
Csorba et al. (2007) described another species,
M. tiensa from Vietnam, and mentioned that M. tien -
sa differs from M. harrisoni by the insertion point of
the wing membrane, and rostral characteristics
where the anterior part of the rostrum is almost
straight in M. tiensa, and more bulbous in M. har-
risoni. Francis and Eger (2012) regarded M. tiensa
as a junior synonym of M. harrisoni in examining
specimens from Laos, Thailand, China and Vietnam,
and concluded that the morphological characteris-
tics separating the two species were actually in-
traspecies variation. DNA sequence variation among
these specimens indicated two distinct clades, but
the divergence of all specimens was less than 5–6%.

216 N. Truong Son, G. Csorba, V. Tan Tu, V. Dinh Thong, Y. Wu
FIG. 8. Scatter plots between size PC1 and shape PC1 based
on craniodental measurements of M. harrisoni and ‘M. tiensa’
specimens. Open and closed circles represent females and
males, respectively
Thereafter, Wu et al. (2010) reported the species
as M. harrisoni from Hainan Island, China, and Son
et al. (2015) recognized M. tiensa instead of M. har-
risoni for specimens from Vietnam.
To clarify the relationship between M. harrisoni
and M. tiensa, we examined specimens from the col-
lections of HZM, HNHM, IEBR, ROM, and IBHG.
Plots between size PC1 (PCA of log-transformed
raw data) and shape PC1 (PCA of log-transformed
standardized data) are given in Fig. 8. In size PC1,
females were larger than males with little overlap.
Size PC1 and shape PC1 (based on log-transformed
standardized data) had a negative correlation. Size
PC1 explained 76.2% of the variance, and all char-
acters had positive loading factors between 0.090–
0.459. The shape PC1 explained 33.6% of the vari-
ance, and CPH (-0.586, negative), BCW (0.472,
positive), and IOW (0.442, positive) showed high
factor loadings. Holotypes of M. harrisoni and
M. tiensa (both females) plotted closely. This result
does not support the existence of two distinct
species. We, therefore, suggest that the observed
morphological variation is intraspecific variation
found in M. harrisoni and involves strong sexual
dimorphism; consequently, we regard M. tiensa as
a junior synonym of M. harrisoni. Because the num-
ber of specimens was not sufficient at each locality,
we cannot provide conclusions about geographic
variation and the possible intraspecific divergence
within M. harrisoni.
TABLE 5. Comparison of karyotypes of the subfamily Murininae. Abbreviations: M: metacentrics; SM: submetacentrics; ST: mubtelocentrics; A: acrocentrics
Species Locality 2n FN M SM ST A X Y References
M. kontumensis sp. nov. Vietnam 44 50 3 1 0 17 M – This study (n = 1)
M. leucogaster China 44 58 3 1 4 13 M A Gu (2006)
M. hilgendorfi Japan 44 56 3 1 3 14 SM A Harada (1973), Ando et al. (1977), Harada et al. (1987)
M. harrisoni China 44 50 3 1 0 17 M – Wu et al. (2010)
M. harrisoni Thailand 44 50 3 1 0 17 M A McBee et al. (1986), see also Francis and Eger (2012)
M. huttoni Vietnam 44 50 3 1 0 17 M A This study (n = 5)
M. cyclotis Vietnam 44 50 3 1 0 17 M A This study (n = 5)
M. lorelieae Vietnam 44 50 3 1 0 17 SM A This study (n = 2)
M. beelzebub Vietnam 44 50 3 1 0 17 M – This study (n = 1)
M. feae Vietnam 44 50 3 1 0 17 M – This study (n = 1)
M. feae China 44 50 3 1 0 17 M A Zhou et al. (2011)
M. puta Taiwan 44 50 3 1 0 17 M A Lin et al. (2002)
M. suilla Malaysia 44 58 3 1 4 13 SM ST Volleth (2006)
M. ussuriensis Japan 44 56 3 0 4 14 M A Ono and Obara (1994)
M. ussuriensis Japan 44 56 3 1 3 14 SM A Ando et al. (1977), Harada et al. (1987)
H. harpia Vietnam 44 52 4 1 0 16 M A This study (n = 1)
H. harpia Taiwan 44 52 4 1 0 16 M – Lin et al. (2006)
H. harpia China 44 52 4 1 0 16 M A Zhou et al. (2014)
H. harpia Thailand 40 – – – – – – – McBee et al. (1986)
H. isodon Vietnam 44 50 3 1 0 17 M – This study (n = 1)

Difference in the braincase region between males
and females was relatively small, as shown in the
area connecting the posterior end of the interorbital
region, outer margin of the braincase and the poste-
rior end of the skull in dorsal view (Fig. 9). On
the other hand, the area of the nasal capsule was
expanded more laterally and anteriorly in females
than males, with a line connecting the anterior end
of the interorbital region, the anterior of the zygo-
mata, the posterior margin of the canines, and the
anterior tip of the skull. In addition, females have
well-developed canines (C) and a robust rostrum as
compared to males (Fig. 9).
Size and shape of the braincase and nasal capsule
may be related to functional limitation for size and
shape change throughout various regions. Therefore,
sexual dimorphism is suggested to have achieved
under these functional limitations different among
skull regions (e.g., braincase and nasal capsule), and
not necessarily with simple overall size change and
accompanied allometric shape changes. As a result,
sexual dimorphoism is to become so complicated
pattern. Previously observed profile variation in
M. harrisoni and M. tiensa should be re-evaluated
with consideration of these complicated morpholog-
ical variations among the sexes (Fig. 9). In addition,
the difference in length of the interorbital region
(elongated in females) likely results in a differ-
ence in bite force between sexes. To understand the
sexual dimorphism, three-dimensional geometric
mor phometric or CT scanning reconstruction meth-
ods are needed.
Francis and Eger (2012: Fig. 9) provided photos
of five individuals of M. harrisoni and M. tiensa.
The smallest depicted female specimen (HZM
1.31525) has caused confusion, suggesting that
females and males widely overlap in overall size.
Our STOTL measurement of the same specimen
(19.33 mm), however, is actually closer to the two
larger female specimens, EGD 24974 (19.87 mm)
and HZM 1.36316 (18.39 mm; erroneously labelled
as ‘1.36136’ in the caption). We assume that scaling
or size adjustment error must have occurred during
the preparation of the figure in Francis and Eger
(2012).
Murina fionae Francis and Eger, 2012
Murina CMF sp. B: Francis et al., 2010: 6.
Murina peninsularis: Matveev and Csorba, 2007: 100.
Murina fionae Francis and Eger, 2012: 32; Type locality: Pha
Deng, 8 km E of Ban Navang, Khammouan Province, Laos,
1,140 m a.s.l.; Thomas et al., 2013: 231.
Distribution
Vietnam, Laos, and Cambodia (Francis and Eger,
2012; Soisook et al., 2013b).
In Vietnam (Fig. 1): Quang Binh (Phong Nha-Ke
Bang NP) [24], Quang Tri (Bac Huong Hoa NR)
New species of Murina from Vietnam 217
FIG. 9. Dorsal and left lateral views of skulls from male (B190913.7, top) and female (PM28, bottom) specimens of M. harrisoni

[25], Quang Nam (Ngoc Linh NR, Song Thanh NR)
[27, 28], Quang Ngai (Ba To area) [30], Kon Tum
(Chu Mom Ray NP) [31], Gia Lai (Kon Cha Rang
NR, Kon Ka Kinh NP) [32], Dong Nai (Cat Tien
NP) [39] (This study; Francis and Eger, 2012; Soi -
sook et al., 2013b).
Description
Large species with ‘cyclotis-type’ dentition (Fig.
4, Tables 2 and 4). Ears are moderately large and
rounded. The fur of the dorsum is long with pale
buff bases and orange-brown tips. Scattered longer
guard hairs are pale to the tip, creating a frosted ap-
pearance. The hairs of the ventrum are unicoloured,
pale buff orange over most of the venter, but more
whitish near the chin. The interfemoral membrane,
legs, feet and tail are covered with long orange-
brown hairs, which are relatively dense on the legs
and feet. Plagiopatagium is attached to the base of
the toe claw.
Skull and dentition (Figs. 5 and 6). Rostrum is
not inflated and the braincase is well domed.
Sagittal crest is strong and the lambdoid crest is
weakly developed. I2 is lateral to I3, C is much
higher than P4. C basal area is slightly larger or
equal to P4. P2 height is subequal to P4, its basal
area is two-thirds to nearly equal to P4. No meso -
styles are on M1 and M2. In the mandible, talonids
of m1 and m2 are reduced relative to trigonid. In the
lateral view, posterior cusps are slightly more than
half the height of the anterior cusps. When viewing
from above, the length of the talonid is less than half
the length of trigonid.
Taxonomic note
Francis and Eger (2012) described this species
based on specimens from Laos and Vietnam.
Murina huttoni (Peters, 1872)
Harpiocephalus huttoni Peters, 1872: 257; Type locality: Dehra
Dun, Kumaon, northwestern India.
Murina huttoni: Hendrichsen et al., 2001: 103; Francis and
Eger, 2012: 20; Thomas et al., 2013: 231.
Distribution
Northwest India, Tibet to Thailand, Vietnam,
northeastern and southern China, and west Malaysia
(Tate, 1941; Hill, 1964; Corbet and Hill, 1992; Bates
and Harrison, 1997; Simmons, 2005; Smith and Xie,
2008; Zhou et al., 2011).
In Vietnam (Fig. 1): Lao Cai (Hoang Lien NP)
[1], Cao Bang (Phia Oac-Phia Den NR) [5], Nghe
An (Pu Mat NP) [22], Quang Tri (Bac Huong Hoa
NR) [25], Quang Nam (Ngoc Linh NR) [28], Kon
Tum (Ngoc Linh NR) [29], Dak Lak (Chu Yang Sin
NP) [35], Lam Dong (Bi Dup-Nui Ba NP) [36],
Khanh Hoa (Hon Ba NR) [37] (This study; Can et
al., 2008; Kruskop, 2013).
Description
Medium-sized species with ‘cyclotis-type’ denti-
tion (Fig. 4, Tables 2 and 4). Both dorsum and ven-
tral fur are dark. The fur of the dorsum is long and
fluffy, and is slate grey with a pale buffy band which
darkens gradually into a darker orange-brown band.
The ventral fur is similar but somewhat paler. The
interfemoral membrane is extensively covered with
reddish-brown hairs, which are longer near the
body and shorter near the edge. Plagiopatagium is
attached to base of the toe claw.
Skull and dentition (Figs. 5 and 6). Rostrum is
not inflated and the braincase is flat. Sagittal and
lambdoid crests are weakly developed. I2 is lateral
or partly anterior to I3. C is much higher than P4.
C basal area equals that of P4. P2 height is subequal
to P4, and its basal area is two-thirds that of P4.
M1 and M2 mesostyles are well developed. In the
mandible, talonids of m1 and m2 are well devel-
oped; viewed from above, the length of talonids on
the lingual side is only slightly less than that of
trigonid; in lateral view, the posterior cusps are
about two-thirds the height of the anterior cusps.
Karyotype
Karyotype (Fig. 7B and Table 5) was 2n = 44,
FN = 50. Autosomes consisted of three large meta-
centric pairs, one small submetacentric pair, and
17 medi um-sized to small acrocentric pairs that
gradually decrease in size. The X chromosome pair
could be identified as a medium-sized metacentric
element.
Taxonomic note
Hill (1964) and Corbet and Hill (1992) recog-
nised two subspecies: M. huttoni rubella (Thomas,
1914) (from southeast China and probably in north
Thailand) and M. h. huttoni (Peters, 1872) (from the
rest of the area) in having different colour patterns.
In Vietnam, we found variation in colour among two
specimens from Hoang Lien National Park (Lao Cai
province): a more reddish orange specimen that
is most similar to M. huttoni rubella (Hill, 1964;
Hendrichsen et al., 2001; Francis and Eger, 2012),
and a more greyish brown individual in concordance
with the description of M. h. huttoni. Additional
studies covering the whole distribution range are
needed to understand the possible cryptic diversity.
218 N. Truong Son, G. Csorba, V. Tan Tu, V. Dinh Thong, Y. Wu

Murina cyclotis Dobson, 1872
Murina cyclotis Dobson, 1872: 210; Type locality: Darjeeling,
NE India; Hendrichsen et al., 2001: 104; Ith et al., 2011: 97;
Francis and Eger, 2012: 17; Thomas et al., 2013: 230;
Soisook et al., 2013b: 274.
Distribution
Sri Lanka, India, Myanmar, Laos, Vietnam,
south ern China in Guangdong Province and Hainan
Island, south to West Malaysia, Borneo, Sumatra,
Philippines, and Lesser Sunda (Tate, 1941; Eller-
man and Morrison-Scott, 1951; Hill, 1964; Cor-
bet and Hill, 1992; Bates and Harrison, 1997;
Francis et al., 1999; Simmons, 2005; Francis, 2008;
Smith and Xie, 2008; Ith et al., 2011; Francis and
Eger, 2012; Soisook et al., 2013b; Thomas et al.,
2013).
In Vietnam (Fig. 1): Son La (Xuan Nha NR) [15],
Phu Tho (Xuan Son NP) [12], Cao Bang (Phia Oac-
Phia Den NR) [5], Vinh Phuc (Tam Dao NP) [11],
Quang Ninh (Bai Tu Long NP) [9], Hai Phong (Cat
Ba NP) [10], Ninh Binh (Cuc Phuong NP) [18],
Thanh Hoa (Pu Hu NR, Pu Luong NR, and Xuan
Lien NR) [16, 17, 19], Nghe An (Pu Huong NR, Pu
Mat NP) [21, 22], Quang Binh (Phong Nha-Ke Bang
NP) [24], Quang Tri (Dakrong NR) [25], Thua
Thien-Hue (Bach Ma NP) [26], Kon Tum (Ngoc
Linh NR) [29], Quang Ngai (Ba To area) [30], Gia
Lai (Kon Ka Kinh NP) [32], Binh Dinh (Phu Yen
district) [33], Lam Dong [36], Binh Phuoc [41], Ba
Ria-Vung Tau [42] (This study; Can et al., 2008;
Kruskop, 2013).
Description
Medium-sized species with ‘cyclotis-type’ denti-
tion (Fig. 4, Tables 2 and 4). The ears are relatively
long. The hairs of the dorsum have grey to dark grey
bases, and a buffy area that gradually darkens to
orange near the tips, but does not create strongly
contrasting bands of colour. The hairs on the ventral
surface have dark grey bases with buffy white tips,
which may have a slight orange colouration.
However, orange covers most of the venter, and is
more whitish near the tips for specimens in the
south. Plagiopatagium is attached to the base of the
toe claw.
Skull and dentition (Figs. 5 and 6). Rostrum not
inflated and the braincase is slightly domed in
females, and more domed in males. Sagittal and
lambdoid crests are weak. I2 is lateral to I3, but I3
partly anterior to I2. C is much higher than P4. The
C basal area is equal or is slightly larger than P4.
P2 height is subequal to or slightly shorter than P4,
and its basal area is two-third that of P4. No meso -
style is found on M1 and M2. In the mandible, the
talonid of m1 and m2 is greatly reduced relative to
the trigonid. Viewed from above, the length of the
talonid is less than half the length of the trigonid. In
the lateral view, the posterior cusps are no more than
half the height of the anterior cusps. In the mandible,
both premolars are similar in height.
Karyotype
Karyotype (Fig. 7C and Table 5) was 2n = 44,
FN = 50. Autosomes consisted of three large meta-
centric pairs, one small submetacentric pair, and
17 medi um-sized to small acrocentric pairs that
gradually decrease in size. The X chromosome
was a medium-sized metacentric element, and the Y
chromosome was a small acrocentric element.
Taxonomic note
Ellerman and Morrison-Scott (1951), Hill
(1964), Eisenberg and McKay (1970), Corbet and
Hill (1992), Koopman (1994), Simmons (2005) rec-
ognized three subspecies: M. cyclotis eileenae
(Ceylon), M. cyclotis cyclotis (Northeastern India to
Hainan and Vietnam), and M. cyclotis peninsularis
(Malaysia). Francis and Eger (2012) elevated
M. peninsularis to full species, and suggested that
M. cyclotis sensu stricto is a complex of cryptic
taxa; therefore, future taxonomic studies covering
its wide distribution are desired. Soisook et al.
(2013b) studied this species’ complex and consid-
ered eileenae to be a synonym of M. cyclotis, and
M. peninsularis to be a valid species. In addition,
Soisook et al. (2013b) described a new species,
M. guilleni, from Thailand and further discussed the
taxonomic problems in M. cyclotis that should be
addressed in future studies.
Murina lorelieae Eger and Lim, 2011
Murina lorelieae Eger and Lim, 2011: 234; Type locality: Did -
ing Headwater Forest Nature Reserve, Jing Xi County,
Guangxi Zhuang Autonomous Region, China.
Murina lorelieae ngoclinhensis Tu, Cornette, Utge and Has sa -
nin, 2015: 209; Type locality: Ngoc Linh Nature Reserve,
Vietnam.
Distribution
Southern China, Vietnam (Tu et al., 2015).
In Vietnam (Fig. 1): Kon Tum (Ngoc Linh NR)
[29] (This study; Tu et al., 2015).
Description
Medium-sized species with ‘suilla-type’ denti-
tion (Fig. 4, Tables 2, 3, and 4). The ear is round and
New species of Murina from Vietnam 219

both sides of the muzzle are dark brown. The pelage
is characterized by long shiny hairs (8 mm ventrally
and 13–15 mm dorsally), with distinct copper red-
dish-brown and dirty white colourations on dorsal
and ventral surfaces. Dorsal hairs are dark grey
basally, pale in the middle and reddish brown at the
tip. Ventral hairs are dark grey to about two-thirds
of the length and whitish at the tip.
Skull and dentition (Figs. 5 and 6). The skull
is domed. The lateral profile of the anterior part of
the skull gradually rises from the rostrum to the
forehead. The sagittal crest is lacking; lambdoid
crest is visible but very weak. Maxillary toothrows
are convergent anteriorly. The dentition is quite
robust. I2 is anterior to I3, and I2 is visible lateral-
ly. I2 and I3 are subequal in height and are
much less than half the height of upper C. Upper
C slightly but clearly exceeds the height of
P4, the basal area of C is less than that of P4.
The crown area of P2 is slightly more than half
that of P4. M1 and M2 have well developed
mesostyles and curved labial faces. Paracone, meta-
cone, and protocone of M1 and M2 are distinctly
defined.
Karyotype
Karyotype (Fig. 7D and Table 5) based on a spec-
imen from Kon Tum province (Ngoc Linh Nature
Reserve) was 2n = 44, FN = 50. Autosomes consist -
ed of three large metacentric pairs, one small sub-
metacentric pair, and 17 medium-sized to small
acro centric pairs that gradually decrease in size. The
X chromosome was a medium-sized submetacen-
tric, and the Y chromosome was a small acrocentric
element.
Taxonomic note
Eger and Lim (2011) described M. lorelieae
based on a single specimen from Diding Head-
water Forest Nature Reserve, Jing Xi County,
Guangxi Zhuang Autonomous Region, China; close
to the border with northeast Vietnam. Tu et al.
(2015) recorded this species from Ngoc Linh
Mountain in the Central Highlands of Vietnam at
over 1,600 m elevation (Tu et al., 2015). Based on
DNA barcoding and morphological data, Tu et al.
(2015) described a new subspecies M. lorelieae
ngoclinhensis. This species has been recorded
only from two localities at high elevations in south-
ern China and Vietnam, but additional surveys
are expected to find new localities and to further
describe the geographic boundary of the two
subspecies. Although Tu et al. (2015) regarded
M. lorelieae as a member of the M. cyclotis-
group (on the basis of three specimens from Viet -
nam), a closer examination of additional speci-
mens proved that M. lorelieae shows ‘suilla-type’
dentition.
Murina annamitica Francis and Eger, 2012
Murina CMF sp. D: Francis et al., 2010: 6.
Murina annamitica Francis and Eger, 2012: Type locality: near
Nam Pan in the Annamite Mountains, Bolikhamxai Prov -
ince, Laos, about 1,300 m a.s.l.; Thomas et al., 2013: 231.
Distribution
Vietnam and Laos (Francis and Eger, 2012).
In Vietnam (Fig. 1): Son La (Xuan Nha NR) [15],
Lao Cai (Hoang Lien NP) [1], Tuyen Quang (Na
Hang NR) [3], Thanh Hoa (Pu Luong NR) [17],
Nghe An (Pu Huong NR and Pu Mat NP) [22],
Quang Tri (Bac Huong Hoa NR) [25], Quang Nam
(Ngoc Linh) [28], Kon Tum (Ngoc Linh) [29],
Quang Ngai (Ba To area) [30], Binh Phuoc (Bu Gia
Map NR) [41] (This study; Francis and Eger, 2012;
Kruskop, 2013).
Description
Medium-sized species with ‘cyclotis-type’ denti-
tion (Fig. 4, Tables 2 and 4). The interfemoral mem-
brane is extensively covered with hairs. There
are very short hairs on the forearm and leading edge
of the wing. Plagiopatagium is attached to the base
of the toe claw. The ear is round without a notch
on its posterior border. Dark bases extend to the fur
of the dorsum. The fur of the dorsum is long and
fluffy. The hairs have slate grey bases followed by
a buffy band, then darker brown to orange-brown
tips. The overall appearance is orange-brown to
brown. The fur of the underside has slate grey bases
followed by buffy tips, giving an overall greyish
buff appearance.
Skull and dentition (Figs. 5 and 6). Rostrum not
inflated and the braincase is domed. Sagittal and
lambdoid crests are weak. I2 is lateral to I3. C basal
area equals that of P4. P2 is less in height than P4
and its basal area is two-thirds that of P4. M1 and
M2 mesostyles are well developed, comparable in
height to the metastyle and parastyle, giving a dis-
tinct W-shape to the surface. In the mandible, both
premolars are similar in height, and lower molars
have well-developed talonids.
Taxonomic note
Francis and Eger (2012) described M. annami-
tica as a new species from Laos; and also referred
material from the centre of Vietnam.
220 N. Truong Son, G. Csorba, V. Tan Tu, V. Dinh Thong, Y. Wu

Murina beelzebub Son, Furey and Csorba 2011
Murina tubinaris: Hendrichsen et al. 2001: 103 (part).
Murina beelzebub Son, Furey and Csorba, 2011 in Csorba
et al., 2011: 899; Type locality: Bac Huong Hoa Nature
Reserve, Huong Hoa District, Quang Tri Province, Vietnam,
400 m a.s.l.
Distribution
Recorded only from Vietnam. In Vietnam (Fig.
1): Quang Tri (Bac Huong Hoa NR) [25], Kon Tum
(Ngoc Linh NR) [29], Quang Ngai (Ba To area)
[30], Gia Lai (Kon Ka Kinh NP) [32] (This study;
Csorba et al., 2011).
Description
Medium-sized species with ‘suilla-type’ denti-
tion (Fig. 4, Tables 2 and 4). On the dorsal surface,
the proximal sixth of individual hairs is very
dark brown (almost black), whereas the remain-
ing distal portion is initially light grey and termi-
nates in a distinctly darker tip. Longer silver
guard hairs are scattered over all of the dorsum.
The upper surface of the hind limbs, feet, and
uropatagium are densely covered in long, uniformly
dark brown hairs. Ventrally, hairs are very dark
brown (almost black) for the proximal two-thirds,
whereas the remaining upper portion is white.
Ventral surface of the uropatagium is covered in uni-
formly white hairs, some of which are also present
on the plagiopatagium adjacent to the body. The ear
has a slight emargination along its posterior border,
and the plagiopatagium is attached to the base of the
claw on the outer toe.
Skull and dentition (Figs. 5 and 6). Rostrum is
not inflated and the braincase is domed. No sagittal
crest, and the lambdoid crest is weak. Rostrum
is slightly elongated but is not inflated. The depth
of the narial emargination exceeds its width.
The zygoma is strong and possesses a slight dorsal
process. A medial process is present in the pos-
terior palatal region. The medial ridge separating
the basioccipital pits is relatively narrow, and the
anterior borders of the pits are weakly defined.
I2 is partly anterior to I3. C is slightly higher
than P4. C has a basal area smaller than that of
P4. P2 is much shorter than P4; its basal area
is half that of P4. M1 and M2 mesostyles are weakly
developed. In the mandible, p2 has less than
one-half the basal area of p4 and attains more than
two-thirds its height. Talonids of m1 and m2
equal their trigonids in the crown area and ento-
conids of the teeth distinctly exceed their hypo -
conids in height.
Karyotype
Karyotype (Fig. 7E and Table 5) based on a fe-
male specimen from Quang Tri province (Bac Huong
Hoa Nature Reserve) was 2n = 44, FN = 50. Auto -
somes consisted of three large metacentric pairs, one
small submetacentric pair, and 17 medium-sized to
small acrocentric pairs that gradually decrease in
size. The X chromosome pair could be identified as
a medi um-sized submetacentric element.
Taxonomic note
The first specimen (HZM) was collected around
Kon Cha Rang and Kon Ka Kinh Nature Reserve
(1,600 m a.s.l.) by Benjamin Hayes (Trai et al.,
2000), which Hendrichsen et al. (2001) identified as
M. tubinaris. Thereafter, Csorba et al. (2011) de-
scribed M. beelzebub based on specimens from cen-
tral Vietnam (Quang Tri province) and the HZM
specimen from Kon Ka Kinh Nature Reserve.
Murina walstoni Furey, Csorba and Son, 2011
Murina walstoni Furey, Csorba and Son, 2011 in Csorba et al.,
2011: 900; Type locality: Veun Sai Protected Forest, Veun
Sai District, Cambodia, 110 m a.s.l.; Francis and Eger,
2012: 29; Thomas et al., 2013: 232.
Distribution
Vietnam, Laos, and Cambodia (Csorba et al.,
2011; Francis and Eger, 2012; Kruskop, 2013;
Thom as et al., 2013). In Vietnam (Fig. 1): Dak Lak
(Yok Don NP) [34], Ninh Thuan (Nui Chua NP)
[38], Dong Nai (Vinh Cuu NR) [39], Kien Giang
(Phu Quoc NP) [43] (This study; Csorba et al.,
2011; Kruskop, 2013).
Description
Medium-sized species with ‘suilla-type’ denti-
tion (Fig. 4, Tables 2 and 4). On the dorsal surface,
the fur is warm brown. Ventrally, basal two-thirds of
the hairs are uniformly white, the tips are brown or
orange-brown, and the ventral surface is pure white.
The plagiopatagium is attached to the base of the
first toe claw.
Skull and dentition (Figs. 5 and 6). Rostrum is
not inflated and the braincase is domed. Sagittal and
lambdoid crests are well developed. I2 is partly an-
terior to I3. Upper C is higher than P4. C basal area
is smaller than that of P4. P2 is much less in height
than P4, and its basal area is half that of P4. M1 and
M2 mesostyles are moderately developed. In the
mandible, c exceeds p4 in height and basal area. The
basal area of p2 varies from one-half to less than that
of p4 and attains more than two-thirds its height.
The talonids of m1 and m2 exceed their trigonids
New species of Murina from Vietnam 221

in crown area, and their entoconids are equal to or
slightly higher than their respective hypoconids.
Murina feae (Thomas, 1891)
Harpiocephalus feae Thomas, 1891: 884; Type locality: Biapo,
Karin Hills, Burma.
Murina tubinaris: Corbet and Hill, 1992: 151 (part); Koopman,
1994: 132 (part); Simmons, 2005 (part); Francis et al.,
1999: 233 (part); Francis, 2008: 253 (part).
Murina cineracea Csorba and Furey, 2011: 896 (in Csorba et
al., 2011); Type locality: Cambodia, Mondulkiri Province,
Sei ma Biodiversity Conservation Area.
Distribution
Myanmar, Thailand, Vietnam, Laos, and Cam -
bodia (Thomas, 1891; Osgood, 1932; Francis et al.,
1999, Can et al., 2008; Csorba et al., 2011, Francis
and Eger, 2012; Kruskop, 2013).
In Vietnam (Fig. 1): Lai Chau (Co Ma) [13], Son
La (Phu Yen) [13], Phu Tho (Xuan Son NR) [12], Ha
Giang (Duc Xuan area) [2], Tuyen Quang (Na Hang
NR) [3], Bac Kan (Ba Be NP, Kim Hy NR) [4, 6],
Thai Nguyen (Than Sa NR) [7], Vinh Phuc (Tam
Dao NP) [1], Thanh Hoa (Pu Luong NR, Pu Hu NR,
Xuan Lien NR) [16, 17, 19], Ninh Binh (Cuc
Phuong NP) [18], Nghe An (Pu Hoat NR, Pu Mat
NP, Pu Huong NR) [19, 21, 22], Ha Tinh (Vu Quang
NP) [23], Quang Binh (Phong Nha-Ke Bang NP)
[24], Quang Tri (Bac Huong Hoa NR, Dakrong NR)
[25], Thua Thien-Hue (Bach Ma NP) [26], Quang
Nam (Ngoc Linh NR) [28], Kon Tum (Ngoc Linh
NR) [29], Gia Lai (Kon Ka Kinh NP) [32], and
Dong Nai (Cat Tien NP) [40] (This study; Can et al.,
2008; Csorba et al., 2011; Kruskop, 2013).
Description
Small to medium-sized species with ‘suilla-type’
dentition (Fig. 4, Tables 2 and 4). The ear is evenly
rounded and without an emargination. On the dorsal
surface, the lower portion of individual hairs is dark
brown, whereas the upper portion is light grey and
terminates in a distinctly darker tip. Darkening of
hair tips is more evident on the nape and the head,
with an overall impression of dark greyish-brown
and darker brown toward the head. The upper sur-
face of the hind limbs, feet, and uropatagium are
sparsely covered in short, uniformly dark brown
hairs. On the ventral surface of the body, hairs are
dark brown basally, whereas the upper portion is
white. The ventral surface of the uropatagium is
covered in relatively short uniformly white hairs,
which are also present on the plagiopatagium adja-
cent to the body. The plagiopatagium is attached to
the base of the toe claw.
Skull and dentition (Figs. 5 and 6). Rostrum is
not inflated and the braincase is domed. There is no
sagittal crest and the lambdoid crest is very weak. I2
is partly anterior to I3. C is slightly but evidently
higher than P4, P2 is much less in height than P4.
C basal area is smaller that of P4. M1 and M2 meso -
styles are very weak. In the mandible, cslightly ex-
ceeds p4 in height and is equal or greater in basal
area; p2 has less than one-half the basal area of p4
and attains more than two-thirds its height. The
talonids of m1 and m2 equal their respective trigo-
nids in crown area, and the entoconids of these
teeth exceed their hypoconids in height. The
postristid connects the hypoconid with the tip of
the entoconid.
Karyotype
Karyotype (Fig. 7F and Table 5) based on a spec-
imen from Thanh Hoa Province (Xuan Lien Nature
Re serve) was 2n = 44, FN = 50. Autosomes con-
sisted of three large metacentric pairs, one small
submetacentric pair, and 17 medium-sized to small
acrocentric pairs that gradually decrease in size. The
X chromosome was identified as a medium-sized
meta centric element.
Taxonomic note
Thomas (1891) described Harpiocephalus feae
from Burma (Myanmar), but the species was consid-
ered a synonym of M. aurata since then (Tate 1941,
Maeda, 1980; Corbet and Hill, 1992). Recently,
Csorba et al. (2011) split M. tubinaris (Scully, 1881)
into two species and restricted the distribution of
M. tubinaris sensu stricto to Pakistan and northwest
India. They described a new species, M. cineracea,
that occurred from West Bengal and Arunachal Pra -
desh in India through Myanmar, Thailand, Laos to
Vietnam. Francis and Eger (2012), however, based
on the morphological study of the holotype, con-
cluded that M. feae is actually a conspecific with
M. cineracea, and therefore the name M. feae has
priority over M. cineracea.
Murina eleryi Furey, Thong, Bates and Csorba, 2009
Murina aurata: Francis et al., 1999: 233 (part); Francis, 2008:
253 (part); Francis et al., 2010: 6 (part).
Murina eleryi Furey, Thong, Bates and Csorba, 2009: 226; Type
locality: Kim Hy Commune, Na Ri district of Kim Hy
Nature Reserve, Bac Kan province, Vietnam, 525 m a.s.l.;
Francis and Eger, 2012: 28; Thomas et al., 2013: 231.
Distribution
Vietnam, Laos, and southern China in the
provinces of Guizhou, Hunan, Guangdong, Guangxi
222 N. Truong Son, G. Csorba, V. Tan Tu, V. Dinh Thong, Y. Wu

(Furey et al., 2009; Francis and Eger, 2012; Liu et
al., 2014; Xu et al., 2014).
In Vietnam (Fig. 1): Ha Giang [2], Cao Bang
(Phia Oac-Phia Den NR) [5], Bac Kan (Kim Hy NR)
[6], Phu Tho (Xuan Son NP) [12], Son La (Muong
Do area) [13], Thanh Hoa (Xuan Lien NR) [19],
Quang Binh (Phong Nha-Ke Bang NP) [24]; Quang
Nam (Ngoc Linh NR) [28]; Kon Tum (Ngoc Linh
NR) [29], Quang Ngai (Ba To area) [30] (This study;
Furey et al., 2009; Francis and Eger, 2012; Kruskop,
2013).
Description
Small species with ‘suilla-type’ dentition (Fig. 4,
Tables 2 and 4). Longer, shiny golden hairs with
darker bases are scattered over the back, nape and
head. On the dorsal surface, the lower portion of
under hairs are very dark brown and are followed by
a pale grey-yellow mid-section which progressively
darkens to copper-reddish before terminating in
a distinctly darker tip. The superficial impression
is copper-reddish mottled with underlying dark
brown and overlain by individual shiny gold hairs.
Ventrally, the fur is black on the basal half and
creamy white on the remainder, except the sides of
the ventrum and upper chest, where hair tips gradu-
ate toward very light brown. The plagiopatagium is
attached to the base of the first claw.
Skull and dentition (Figs. 5 and 6). Rostrum is
not inflated and the braincase is moderately domed.
No sagittal crest and the lambdoid crest is weak, but
present. I2 is anterior to I3. C is higher than P4.
C basal area is much smaller than that of P4. P2
basal area is less than half that of P4. M1 and M2
meso styles are well developed. In the mandible,
cdistinctly exceeds p4 in height and is equal or
slightly greater in basal area. The talonids of m1 and
m2 are clearly separated from their trigonids and ex-
ceed these in crown area, and the entoconid clearly
exceeds the hypoconid in height.
Taxonomic note
Francis et al. (1999), Furey and Tu (2006) and
Francis (2008) reported specimens of M. aurata
Milne-Edwards, 1872 from and around Vietnam.
Furey et al. (2009), based on many specimens from
northern Vietnam, described a new species, M. el-
eryi, and provided diagnostic characters to separate
it from M. aurata. Thereafter, Francis and Eger
(2012) suggested that all specimens from Laos and
Vietnam formerly identified as M. aurata, to be
referred to as M. eleryi or M. harpioloides. There -
fore, we conclude that M. aurata does not occur in
Vietnam and Laos. Xu et al. (2014) studied the ge-
netic structure among specimens from southern
China, Vietnam, and Laos, and suggested that
a com plicated phylogeographic pattern may exist.
Murina harpioloides Kruskop and Eger, 2008
Murina harpioloides Kruskop and Eger, 2008: 215; Type
locality: Da Lat plateau, 30 km north-east from Da Lat, Lam
Dong Province, Vietnam, about 1800 m a.s.l.
Distribution
Known only from Vietnam (Kruskop and Eger,
2008). In Vietnam (Fig. 1): Lam Dong (Bi Dup-Nui
Ba NP and Da Lat plateau) [36] (This study;
Kruskop and Eger, 2008; Abramov et al., 2009).
Description
Small species with ‘suilla-type’ dentition (Fig. 4,
Tables 2 and 4). The fur of the dorsum is bicoloured
in having dark brown under fur and a bright golden
tip on the guard hairs. The dorsal guard hairs are
dark brown at the base. The distal halves are tri-
coloured, with pale brown and darker brown rings
and bright orange gold tips. Hair on the ventrum is
dark brown at the base and tipped with pale silver
grey. The entire tail membrane from the proximal to
distal edge, and about one-third of the plagiopatag-
ium next to the body, are covered dorsally with fur
similar to guard hairs covering the body. Toes (up to
the bases of claws), thumb, the upper side of the
forearm and proximal part of the fifth metacarpal are
covered with bright golden hairs.
Skull and dentition (Figs. 5 and 6). Rostrum is
gradually sloped and the braincase is moderately
rounded and domed. No sagittal crest and the lamb-
doid crest is very weak, but present. I2 is anterior to
I3; C is similar in height as P4. C basal area is much
smaller than that of P4. P2 much less in height than
P4, and its basal area is less than half that of P4. M1
and M2 mesostyles are very weak. In the mandible,
the lower canine possesses a small but distinct addi-
tional anterior cusp; c distinctly equals p4 in height
and is equal or slightly greater in the basal area. The
talonids of m1 and m2 are moderately separated
from their trigonids, and the hypoconid distinctly
exceeds the entoconid in height.
Taxonomic note
Kruskop and Eger (2008) described this species
from Lam Dong province. Subsequently, Kruskop
and Shchinov (2010) recorded M. cf. harpioloides
from the Hoang Lien mountain range in north-
ern Vietnam, far from the only previously known
location of the species. Recently, Kruskop (2013)
New species of Murina from Vietnam 223

re-identified this specimen as M. chrysochaetes, and
indicated that M. harpioloides has more orange
colouration of the guard hairs and a less domed
braincase as compared to M. chrysochaetes. During
the present study, two specimens were collected
from the Bi Dup-Nui Ba National Park in the Lam
Dong province, extending the known distribution of
this species.
Murina chrysochaetes Eger and Lim, 2011
Murina chrysochaetes Eger and Lim, 2011: 228; Type local-
ity: Diding Headwater Forest Nature Preserve, Jing Xi
County, Guangxi Zhuang Autonomous Region, China, 978
m a.s.l.
Distribution
Known to occur in Vietnam and the Guangxi
Zhuang Autonomous Region of southern China
(Eger and Lim, 2011; Kruskop, 2013). In Vietnam
(Fig. 1): Lao Cai (Hoang Lien NP) [1] (Kruskop,
2013), and Cao Bang (Phia Oac-Phia Den NR) [5]
(This study).
Description
Small species with ‘suilla-type’ dentition (Fig. 4,
Tables 2 and 4). Ear is small, broad and round with
little emargination on it is posterior edge. Tubular
nostrils are proportionally long and the nostrils and
the tip of the muzzle have a mid-brown pigmenta-
tion. The dorsum is a mix of black and gold bands,
with gold mid-bands and dark tips, overlaid by long,
gold-tipped guard hairs. The ventral pelage is dark at
the base and the tips of the guard hairs are golden in
colour.
Skull and dentition (Figs. 5 and 6). Rostrum is
short and narrow. Braincase is highly domed, the
slope of the forehead is abrupt. No sagittal crest and
the lambdoid crest is very weak. I2 is anterior to I3;
C equals P4 in height, and only half of it in basal di-
mensions; P2 is small, about half the height and one-
third the crown area of P4. M1 and M2 mesostyles
are reduced. The lower canine is the same height as
p2, but exceeds it in basal area; p2 is small.
Taxonomic note
Eger and Lim (2011) described M. chrysochaetes
from China, close to the border of northeast Viet -
nam. Kruskop (2013) recorded a single specimen
from Hoang Lien Son Mountain in northern Viet -
nam. During the present study, a single specimen
was collected from Phia Oac-Phia Den Nature Re -
serve, Cao Bang province. These known localities
are restricted to high mountains around the borders
of Vietnam and China.
Genus Harpiocephalus Gray, 1842
Diagnostic characters
A medium-sized vespertilionid bat with tubular
nostrils. Total length of skull is over 20.0 mm.
Dental formula: I2/3 C1/1 P2/2 M3/3 = 34. M3 is
very reduced and peg-like. The incisors are shorter
than the first upper premolar.
Harpiocephalus harpia (Temminck, 1840)
Vespertilio harpia Temminck, 1840: 219; Type locality: Mt.
Gede, Java.
Harpiocephalus mordax Thomas, 1923:88; Type locality: Mo -
gok, N Burma.
Harpiocephalus harpia: Hendrichsen et al., 2001: 105; Lunde et
al., 2007: 160; Abramov et al., 2009: 67.
Distribution
Known to occur in India, Vietnam, Cambodia,
Myanmar, Thailand, China, Malaysia, and Indo-
nesia (Corbet and Hill, 1992; Koopman, 1994; Hen -
drich sen et al., 2001; Matveev, 2005; Kruskop,
2013).
In Vietnam (Fig. 1): Lao Cai (Hoang Lien NP)
[1], Tuyen Quang (Na Hang NR) [3], Bac Kan (Ba
Be NP) [4], Cao Bang (Phia Oac-Phia Den NR) [5],
Lang Son (Huu Lien NR) [8], Hai Phong (Cat Ba
NP) [10], Nghe An (Pu Huong NR) [21], Quang
Binh (Phong Nha–Ke Bang NP) [24], Kon Tum
(Ngoc Linh NR) [29], Lam Dong (Bi Dup-Nui
Ba NP) [36] (This study; Hendrichsen et al.,
2001; Lunde et al., 2007; Thong and Furey, 2008;
Abramov et al., 2009; Kruskop and Shchinov,
2010).
Description
A medium-sized vespertilionid bat, but the
largest species of the subfamily Murininae
(Fig. 4, Tables 2, 3 and 4). The dorsal pelage has
dark grey bases with a light reddish buff, and
a rich, dark red tip. The ventral pelage is pale grey
or reddish with dark bases. The wings are dark
brown.
Skull and dentition (Figs. 5 and 6). Rostrum is
short and broad and the braincase is domed. Sagittal
and lambdoid crests are well developed. The denti-
tion is robust. Upper canines are very strong. I2
height clearly exceeds that of I3. Both upper incisors
are much lower in height than C. P2 smaller than
P4 in basal dimensions, and are subequal in height.
M1 and M2 have no mesostyle. M3 is reduced. In
the mandible, the coronoid process is very promi-
nent, c is well developed; p2 is less than p4 basally
but equal in height.
224 N. Truong Son, G. Csorba, V. Tan Tu, V. Dinh Thong, Y. Wu

Karyotype
The karyotype (Fig. 7G and Table 5) of the spec-
imen from Kon Tum Province (Ngoc Linh Nature
Re serve) was 2n = 44, FN = 52. Autosomes con-
sisted of three large and one small metacentric pairs,
one submetacentric pair, and 16 medium-sized to
small acrocentric pairs, gradually decreasing in size.
The X chromosome was a medium-sized metacen-
tric, and the Y chromosome was a small acrocentric.
Taxonomic note
Harpiocephalus had been considered to include
two species, H. harpia and H. mordax (Thomas,
1923; Hill and Francis, 1984; Corbet and Hill, 1992;
Hendrichsen et al. 2001; Simmons, 2005), where
H. mordax is greater and more robust in skull size
and shape than H. harpia. Matveev (2005) reviewed
the literature and specimens from Cambodia and in-
dicated that male and female specimens differed in
size and shape. These specimens purely fit with the
view of Hill and Francis (1984) on H. harpia (male)
and H, mordax (female), and molecular markers
(Inter-SINE-PCR) clearly demonstrated specimens
to be conspecifics (Matveev, 2004). Lin et al. (2006)
reported the genus from Taiwan for the first time and
considered Harpiocephalus a monotypic genus and
H. mordax to be a synonym of H. harpia in refer-
ence to size differences between sexes. Sexual di-
morphisms were also confirmed in the population in
China (Zhou et al., 2014; Chen et al., 2015), and it
is currently thought that only H. harpia exists.
There is no study on geographic variation of
the widely distributed H. harpia. The karyotype of
‘H. mordax’ from Thailand (McBee et al., 1986) is
2n = 40. This is different from karyotypes reported
in Taiwan and Guangdong Province, China (Lin et
al., 2006; Zhou et al., 2014). Further morphological
and molecular biological studies throughout the
whole distribution area of Harpiocephalus should
be performed.
Genus Harpiola Thomas, 1915
Diagnostic character
Small-sized vespertilionid bats with tubular nos-
trils: STOTL < 18.0. The dental formula is I2/3 C1/1
P2/2 M3/3=34. Both upper premolars (P2 and P4)
and canine are similar in shape and size. I3 is large,
robust and in contact with the upper canine. I3 is
slightly larger than I2. The upper incisors exceed the
half the height of the corresponding canines. P2
height is more than that of P4, and the lower canine
is bicuspid.
Taxonomic note
Harpiola was described by Thomas (1915) based
on Murina grisea Peters, 1872 as type species.
Although, Ellerman and Morrison-Scott (1951),
Cor bet and Hill (1992), Koopman (1994) and Sim -
mons (2005) treated Harpiola as a subgenus of
Murina, Tate (1941), Bhattacharyya (2002), Kuo et
al. (2006) and Kruskop et al. (2006) emphasized the
considerable differences in skull and dentition be-
tween Murina and Harpiola, and accepted the valid
generic status of Harpiola, which view is followed
herewith.
Harpiola isodon Kuo, Fang, Csorba and Lee, 2006
Harpiola isodon Kuo, Fang, Csorba and Lee, 2006: 13; Type
locality: Hualien County, Jhuosi Township, Yuli Wildlife
Re fuge, Taiwan, 23°32’N, 121°15’E, 2,000 m elevation;
Kru skop, 2013: 176.
Harpiola cf. isodon; Kruskop et al., 2006: 14.
Distribution
Vietnam and Taiwan (Kuo et al., 2006; Kruskop
et al., 2006; Kruskop and Shchinov, 2010; Kruskop,
2013). In Vietnam: (Fig. 1). Lao Cai (Hoang Lien
NP) [1] and Kon Tum (Ngoc Linh NR) [29]
(Kruskop et al., 2006, Kruskop and Shchinov, 2010;
Kruskop, 2013; this study).
Description
Medium-sized species of the subfamily (Fig. 4,
Tables 2, 3 and 4). On the dorsal surface, hairs are
very long and woolly; the basal part of underfur is
dark brown with a bright yellow subterminal band
and a dark brown tip. Guard hairs scattered all over
the back are dark brown at the basal four-fifths with
shiny golden yellow tips. From the ventral aspect,
the fur is shorter, dark brown at the base and light
brown in the terminal one-third. The dorsal side of
the tail membrane, the tibia and the foot are all
densely and evenly furred, including the last caudal
vertebra, which is free from the uropatagium. The
whole area of the tail membrane is also covered with
dense, stiff, silvery grey hairs. The ear conch is pos-
sesses a very distinct emargination at the upper third
of its posterior border. The tragus is moderately long
(7.60 mm around), but wide at its base and gradually
tapering to the backward-curved tip, which just
reaches the level of the notch. The base of the tra-
gus is with a small tooth-like projection at its outer
margin.
Skull and dentition (Figs. 5 and 6). Rostrum is
not inflated and the braincase is moderately domed.
No sagittal crest and the lambdoid crest is weakly
developed. Narial emargination is much longer than
New species of Murina from Vietnam 225

wide. Basioccipital pits are well-defined, usually
elongated and especially narrower posteriorly. I2 is
a bit longer than the outer upper incisor (I3). Both
upper incisors are about two-thirds that of C in
height, and the basal area of the second upper inci-
sor is more than two-thirds that of C. The basal area
of C, P2, and P4 are subequal and they are gradually
decreasing in height. The mesostyle of M1 and M2
is very weak, but recognizable. In the mandible, the
lower canine has a well-developed additional cusp.
The lower canines and premolar teeth are similar
in bulk, and c is slightly less than p2 in height.
Entoconid in m1 and m2 is lower than hypoconid,
and formed a distinct cusp widely separated from
metaconid.
Karyotype
Karyotype (Fig. 7H and Table 5) based on
one specimen from Lao Cai province (Hoang Lien
National Park) (IEBR-M5436) was 2n = 44,
FN = 50. Auto somes consisted of three large meta-
centric pairs, one small submetacentric pair, and
17 medium-sized to small acrocentric pairs gradu-
ally decreasing in size. The X chromosome was
a medium-sized metacentric.
Taxonomic note
This species was described as a second species of
the genus Harpiola from Taiwan by Kuo et al.
(2006). Kruskop et al. (2006) and Kruskop and
Shchi nov (2010) reported this species from Viet -
nam, and considered it to be conspecific with the
population in Taiwan.
DISCUSSION
Present and previous studies for karyotypes of
the subfamily Murininae (Table 5; Harada, 1973;
Ando et al., 1977; McBee, 1986; Harada et al.,
1987; Ono and Obara, 1994; Lin et al., 2002, 2006;
Gu, 2006; Volleth, 2006; Wu et al., 2010; Zhou et
al., 2011, 2014) emphatically indicated that all
species karyotyped had 2n = 44, except for Harpio -
cephalus harpia (as H. mordax) reported by McBee
(1986) from Thailand. Species of Murina and Har -
piola share similar karyotypes, characterized by
three large metacentric and one small submetacen-
tric autosomal pairs, while Harpiocephalus differs
in having an additional small submetacentric pair
(no. 5). This feature is also shared with H. harpia
populations in Taiwan and Guangdong, southern
China (Lin et al., 2006; Zhou et al., 2014), and
is suggested to have evolved by the inversion of
the largest acrocentric pair in the Murina karyotype
during the evolution of the genus Harpiocephalus.
Except for H. harpia, conservative trends in chro-
mosome rearrangements in the subfamily Murininae
have been confirmed in this study, including addi-
tional species in the genus Murina. Cytological iso-
lation mechanisms may not be responsible for the
diversification of the genus Murina. Molecular data
did not suggest clear relationships among lineages,
but this may be interpreted as many lineages of the
subfamily Murininae having been separated simul-
taneously and diverged thereafter. Future studies to
examine the evolutionary history and mechanisms
that enable such diversification within a short time
period are needed.
Son et al. (2015) suggested that the important
role of morphological diversification among the
species of Murina in Vietnam is in the interaction
among sympatric species pairs and between sexes.
This is due to the observations from species that
have diverged in the combination of size and shape
of skulls, and sexual size dimorphism (i.e., larger fe-
males) with different dimorphic characteristics
among species. Detailed analyses in this study of the
sexual differences in M. harrisoni showed more
complicated patterns of diversification. Sexual size
dimorphism has not simply been achieved through
differences in size and allometric-based shape, but
through complicated size and shape differentia-
tion due to the functional limitations and compensa-
tion of the skull. Continued observations of skulls
(Fig. 9) may suggest that limiting factors for mor-
phological variability exist in the nasal capsule or
braincase, possibly in relation to echolocation
function.
Distribution data and overall body size provide
another interesting view for the interaction among
sympatric species pairs. In each locality (Table 1),
there were from one to seven species, except in the
Kon Tum where 11 species were recorded, indicat-
ing a high species diversity among localities. This is
probably because high mountains and primary
forests enable the co-occurrence of multiple spe-
cies through differences in altitudinal distribution
(Fig. 10).
Among the 16 Murininae species in Vietnam,
Harpiocephalus harpia, M. cyclotis, M. annamitica,
M. feae, and M. eleryi were recorded from more
than 10 localities (Table 1). Similar-sized species
tend to separate by elevation as shown in Table 1
and Fig. 10. On the other hand, two medium-sized
species, M. cyclotis and M. feae, are both found in
lowland and overlap localities (Fig. 10 and Table 1).
226 N. Truong Son, G. Csorba, V. Tan Tu, V. Dinh Thong, Y. Wu

They provide an interesting example in that these
two species clearly separate by STOTL in spite of
overlapping FA (Fig. 11). We suggest that a morpho-
logical shift in skull size contributed to decreased
interspecific competition among similar-sized
species with overlapping FA. FA-STOTL plots (Fig.
11) also indicate that other species are well sepa-
rated and only overlap in FA or STOTL among sim-
ilar-sized species. In addition, the patterns of FA-
STOTL shift for each species might not be parallel
between males and females.
Forearm length and overall skull size have often
been considered an indicator of overall size of bats,
but we suggest that these two characteristics are re-
lated with completely different adaptations among
species and between sexes. Wu et al. (2015) re-
ported that echolocation call frequencies in Rhino -
lophus bats correlates with nasal capsule size in the
skull, but did not relate to forearm length. In concor-
dance with Son et al. (2015), we suggest that skull
size and shape factors are affected by food habits
and the echolocation function, whereas the FA
and external morphology are more related to flight
behaviour and aerial niche use.
Vietnam possesses the highest number of species
of the subfamily Murininae in the world. Although
New species of Murina from Vietnam 227
FIG. 11. Scatter plots between FA and STOTL of specimens from species in the genera Murina and Harpiola from Vietnam: male
(A) and female (B)
FIG. 10. Elevation distribution of specimens from species in the subfamily Murininae from Vietnam. Species names abbreviated to
the first 3-4 letters

this diversity may be a result of greater sampling ef-
fort within Vietnam, the complex geological history
and biogeographic features of the country suggest
a greater likelihood of complex speciation within
this group of bats than perhaps exists in other coun-
tries of mainland southeast Asia. Our study also
suggests that ecological adaptations, such as interac-
tions among sympatric species pairs and intraspec -
ific relationships between males and females, played
important roles in the formation of taxonomic and
morphological diversity. Understanding the reasons
for the high species diversity as well as the evolu-
tionary processes of the subfamily will require addi-
tional study on the taxonomy, distribution, ecology,
and behavior of this group in Vietnam, and in South -
east Asia in general.
ACKNOWLEDGEMENTS
We are grateful to the Director, Nguyen Xuan Dang, Ngu -
yen Quang Truong, Le Duc Anh, Tring Quang Phap (IEBR), Bui
Tuan Hai (Vietnam National Museum of Nature), Hoang Trung
Thanh, Le Duc Minh (Hanoi University of Sci ence), and
Nguyen Vu Khoi, Thomas O’Shea (Wildlife-at-Risk), Takafumi
Nakano, Satoru Arai, Dai Fukui, Shin-ichiro Kawada for sup-
port with field survey, specimens, photo credit, and preparing
distribution map. We are grateful to Sergei Kru skop (ZMMU),
Judith Eger (ROM), Tamás Görföl, Péter Estók (HNHM) for
kind support in loan of specimens and photos. We thank the
Ministry of Agriculture, and staffs of Nature Reserves and Na -
tional Parks in Vietnam for providing support to conduct field
survey. This study was supported by the Japan Society for the
Promotion of Science (JSPS) Ronpaku Program (VAST-11125),
JSPS Core-to-Core Program B. Asia-Africa Scientific Plat forms,
VAST program for young researcher (IEBR.CBT. TS08.14/
-05.2015), NSFC Major International Regional Joint Research
Project Grant (31110103910), SYNTHESYS Project financed
by the European Community Research Infrastructure Action
under the FP7 ‘Capacities’ Program, Hungarian Sci entific
Research Fund (OTKA) K112440, and JSPS Grants-in-Aid for
Scientific Research (26304009).
LITERATURE CITED
ABRAMOV, A. V., V. V. ROZHNOV, and P. N. MOROZOV. 2006.
Notes on mammals of the Ngoc Linh Nature Reserve (Viet -
nam, Kon Tum province). Russian Journal of Theriology, 5:
85–92.
ABRAMOV, A. V., S. V. KRUSKOP, and A. V. SSCHINOV. 2009.
Small mammals of the Dalat Plateau, southern Vietnam.
Rus sian Journal of Theriology, 8: 61–73.
ANDO, K., T. TAGAWA, and T. A. UCHIDA. 1977. Considerations
of karyotypic evolution within Vespertilionidae. Experien -
tia, 33: 877–879.
ANTHONY, E. L. P. 1988. Age determination in bats. Pp. 47–58,
in Ecological and behavioral methods for the study of bats
(T. H. KUNZ, ed.). Smithsonian Institution Press, Washing -
ton, D.C., 479 pp.
BARLOW, K. E., G. JONES, and E. M. BARRATT. 1995. Can skull
morphology be used to predict ecological relationships be-
tween bat species? A test using two cryptic species of pip-
istrelle. Proceedings of the Royal Society of London, 264B:
1695–1700.
BATES, P. J. J., and D. L. HARRISON. 1997. Bats of the Indian
sub continent. Harrison Zoological Museum, Sevenoaks,
UK, 258 pp.
BATES, P. J. J., TIN NWE, M. J. PEARCH, KHIN MAUNG SWE, SISI
HLA BU, and THANDA TUN. 2000. A review of bat research
in Myanmar (Burma) and results of a recent survey. Acta
Chiropterologica, 2: 53–82.
BHATTACHARYYA, T. P. 2002. Taxonomic status of the genus
Har piola Thomas, 1915 (Mammalia: Chiroptera: Vesper ti -
lio nidae), with a report of the occurrence of Harpiola gri sea
(Peters, 1872) in Mizoram, India. Proceedings of the Zool -
ogical Society, Calcutta, 55: 73–76.
BLACKITH, R. E., and R. A. REYMENT. 1971. Multivariate mor-
phometrics. Academic Press, London, UK, ix + 412 pp.
BURMUNGSRI, S., D. L. HARRISON, C. SARASOOK, A. PRAJUKJITR,
S. THONG-AREE, and P. J. J. BATES. 2006. A review of bat
research in Thailand with eight new species records for the
country. Acta Chiropterologica, 8: 325–359.
CAN, N. D., H. ENDO, N. T. SON, T. OSHIDA, L. X. CANH, D. H.
PHUONG, D. P. LUNDE, S. KAWADA, A. HAYASHIDA, and M.
SASAKI. 2008. Checklist of wild mammal species of Viet -
nam. Primate Research Institute, Inuyama, Japan, 400 pp.
[In English and Vietnamese].
CHEN, B., W. YU, Y. WU, F. LI, Z. XU, Q. ZHANG, M. HARADA,
M. MOTOKAWA, H. PENG, Y. WANG, and Y. LI. 2015. New
record and sexual dimorphism of Harpiocephalus harpia in
Guangxi and Jiangxi, China. Sichuan Journal of Zoology,
34: 211–215. [In Chinese with English abstract].
CORBET, G. B., and J. E. HILL. 1992. The mammals of the
Indo malayan Region. Natural History Museum, London,
488 pp.
CSORBA, G., and P. J. J. BATES. 2005. Description of a new spe -
cies of Murina from Cambodia (Chiroptera: Vespertilio ni -
dae: Murininae). Acta Chiropterologica, 7: 1–7.
CSORBA, G., V. D. THONG, P. J. J. BATES, and N. M. FUREY.
2007. Description of a new species of Murina from Vietnam
(Chiroptera: Vespertilionidae: Murininae). Occasional Pa -
pers, Museum of Texas Tech University, 268: 1–12.
CSORBA, G., N. T. SON, I. SAVENG, and N. FUREY. 2011. Reveal -
ing cryptic bat diversity: three new Murina and redescrip-
tion of M. tubinaris from Southeast Asia. Journal of Mam -
malogy, 92: 891–904.
DOBSON, G. E. 1872. Notes on some bats collected by Captain
W. G. Murray, in the North-Western Himalaya, with de-
scription of new species. Proceedings of the Asiatic Society
of Bengal, 1872: 208–210.
EGER, J. L, and B. K. LIM. 2011. Three new species of Murina
from Southern China (Chiroptera: Vespertilionidae). Acta
Chi ropterologica, 13: 227–243.
EISENBERG, J. F., and G. M. MCKAY. 1970. An annotated check-
list of the recent mammals of Ceylon with keys to the
species. Ceylon Journal of Science (Biological Sciences),
8: 69–99.
ELLERMAN, J. R., and T. C. S. MORRISON-SCOTT. 1951. Chec-
klist of Palaearctic and Indian mammals 1758 to 1946.
Trustees of the British Museum (Natural History), London,
810 pp.
FRANCIS, C. M. 2008. A guide to the mammals of Southeast
Asia. Princeton University Press, Princeton, 392 pp.
228 N. Truong Son, G. Csorba, V. Tan Tu, V. Dinh Thong, Y. Wu

FRANCIS, C. M, and J. EGER. 2012. A review of tube-nosed bats
(Murina) from Laos with a description of two new species.
Acta Chiropterologica, 14: 15–38.
FRANCIS, C. M., A. GUILLEN, and M. F. ROBINSON. 1999.
Order Chiroptera: Bats. Pp. 225–235, in Wildlife in Lao
PDR: 1999 status report (J. W. DUCKWORTH, R. E.
SALTER, and K. KHOUNBOLINE, eds.). IUCN, Vientiane, Lao
PDR, 275 pp.
FRANCIS, C. M., A. V. BORISSENKO, N. V. IVANOVA, J. L. EGER,
B. K. LIM, A. G. SERVENT, S. V. KRUSKOP, I. MACKIE, and
P. D. N. HEBERT. 2010. The role of DNA barcodes in under-
standing and conservation of mammal diversity in Southeast
Asia. PLoS ONE, 5: e12575.
FUREY, N. M., and V. T. TU. 2006. The bat fauna of Khau Ca
area, Vi Xuyen and Bac Me District, Ha Giang Province,
northern Vietnam. Report to the Fauna and Flora Inter na -
tional Vietnam Programme, 18 pp.
FUREY, N. M., V. D. THONG, P. J. J. BATES, and G. CSORBA. 2009.
Description of a new species belonging to the Murina ‘suil -
la-group’ (Chiroptera: Vespertilionidae: Murininae) from
north Vietnam. Acta Chiropterologica, 11: 225–236.
GRAY, J. E. 1842. Description of some new genera and fifty un-
recorded species of Mammalia. Annals and Magazine of
Natural History, 10: 255–267.
GU, X. 2006. The Karyotypes of six species of bats from Gui -
zhou. Chinese Journal of Zoology, 41(5): 112–116. [In Chi -
nese with English abstract].
HAMMER, Ø., D. A. THARPER, and P. D. RYA N. 2001. PAST:
Paleontological statistics software package for education
and data analysis. Palaeontologia Electronica 4: 9 pp. Avail -
able at http://palaeo-electronica.org/2001_1/past/ issue1_01.
htm. Downloaded on 10 September 2012.
HARADA, M. 1973. Chromosomes of nine Chiroptera species in
Japan (Chiroptera). La Kromosoma, 91: 2885–2895.
HARADA, M., K. ANDO, T. A. UCHIDA, and T. H. YOSIDA. 1978.
Karyological study of four Japanese Myotis bats (Chiro -
ptera, Mammalia). Chromosoma, 5: 283–291.
HARADA, M., K. ANDO, T. A. UCHIDA, and S. TAKADA. 1987.
A karyological study on two Japanese species of Muri-
na (Mammalia: Chiroptera). Journal of the Mammalogical
So ciety of Japan, 12: 15–23.
HEBERT, P. D. N., A. CYWINSKA, S. L. BALL, and J. R. DEWAARD.
2003. Biological identifications through DNA barcodes.
Pro ceedings of the Royal Society, 270B: 313–321.
HENDRICHSEN, D. K., P. J. J. BATES, B. HAYES, and J. L. WAL -
STON. 2001. Recent records of bats (Mammalia: Chiroptera)
from Vietnam with six species new to the country. Myotis,
39: 35–122.
HILL, J. E. 1964. Notes on some tube-nosed bats, genus Murina,
from southeastern Asia, with descriptions of a new spe-
cies and a new subspecies. Federation Museums Journal, 8:
48–59.
HILL, J. E., and C. M. FRANCIS. 1984. New bats (Mammalia:
Chiroptera) and new records of bats from Borneo and Ma -
laya. Bulletin of British Museum of Natural History (Zool -
ogy), 46: 305–329.
ITH, S., G. CSORBA, P. J. J. BATES, and N. M. FUREY. 2011. Con -
firmation of seven bat species for Cambodia. Cambo dian
Journal of Natural History, 2011: 92–103.
JUNGGERS, W. L., A. B. FALSETTI, and C. E. WALL. 1995. Shape,
relative size, and size-adjustments in morphometrics. Year -
book of Physical Anthropology, 38: 137–161.
KOOPMAN, K. F. 1994. Chiroptera: systematics. Handbook of
Zoology. Mammalia, Part 60. Walter de Gruyter, Berlin,
217 pp.
KRUSKOP, S. V. 2005. Towards the taxonomy of the Russian
Murina (Vespertilionidae, Chiroptera). Russian Journal of
Theriology, 4: 91–99.
KRUSKOP, S. V. 2013. Bats of Vietnam. Checklist and an identi-
fication manual. Joint Russian-Vietnamese Sciences and
Technological Tropical Centre and Zoological Museum of
Moscow M. V. Lomonosov State University, Moscow,
299 pp.
KRUSKOP, S. V., and J. L. EGER. 2008. A new species of tube-
nosed bat Murina (Vespertilionidae, Chiroptera) from Viet -
nam. Acta Chiropterologica, 10: 213–220.
KRUSKOP, S. V., andA. V. SHCHINOV. 2010. New remarkable bat
records in Hoang Lien Son mountain range, northern Viet -
nam. Russian Journal of Theriology, 9: 1–8.
KRUSKOP, S. V., M. V. KALYAKIN, and A. V. ABRAMOV. 2006.
First record of Harpiola (Chiroptera, Vespertilionidae) from
Vietnam. Russian Journal of Theriology, 5: 13–16.
KUO, H.-C., Y.-P. FANG, G. CSORBA, and L.-L. LEE. 2006. The
definition of Harpiola (Vespertilionidae: Murininae) and the
description of a new species from Taiwan. Acta Chiroptero -
logica, 8: 11–19.
KUO, H.-C., Y.-P. FANG, G. CSORBA, and L.-L. LEE. 2009. Three
new species of Murina (Chiroptera: Vespertilionidae) from
Taiwan. Journal of Mammalogy, 90: 980–991.
KUZNETSOV, G. V. 2006. Mammals of Vietnam. KMK Scientific
Press Ltd., Moscow, 420 pp. [In Russian].
LIN, L.-K., M. MOTOKAWA, and M. HARADA. 2002. Karyology
of ten vespertilionid bats (Chiroptera: Vespertilionidae)
from Taiwan. Zoological Studies, 41: 347–354.
LIN, L.-K., M. HARADA, M. MOTOKAWA, and L.-L. LEE. 2006.
Updating the occurrence of Harpiocephalus harpia (Chiro -
ptera: Vespertilionidae) and its karyology in Taiwan. Mam -
malia, 70: 170–172.
LINDENFORS, P., J. L. GITTEMAN, and K. E. JONES. 2007. Sex-
ual size dimorphism in mammals. Pp. 16–26, in Sex, size,
and gender roles: evolutionary studies of sexual size
dimorphism (D. J. FAIRBAIRN, W. U. BLANCKENHORN,
and T. SZÉKELY, eds.). Oxford University Press, Oxford,
278 pp.
LIU, Z., Y. ZHANG, J. ZHANG, and L. ZHANG. 2014. Murina eleryi
discovered in Hunan Province. Chinese Journal of Zoology,
49(1): 132–135. [In Chinese with English abstract].
LUNDE, D. P., N. T. SON, and G. G. MUSSER. 2007. A survey of
small mammals from Huu Lien Nature Reserve, Lang Son
Province, Vietnam. Mammal Study, 32: 155–168.
MAEDA, K. 1980. Review on the classification of little tube-
nosed bats, Murina aurata group. Mammalia, 44: 531–551.
MAEDA, K., and S. MATSUMURA. 1998. Two new species of
vespertilionid bats, Myotis and Murina (Vespertilionidae:
Chiroptera) from Yanbaru, Okinawa island, Okinawa Pre -
fecture, Japan. Zoological Science, 15: 301–307.
MATVEEV, V. A. 2004. Systematics of the Old World Chiroptera
derived from the insertions of DNA interspersed re-peats.
PhD. Thesis, Moscow State University, Moscow, 293 pp.
[In Russian].
MATVEEV, V. A. 2005. Checklist of Cambodian bats (Chiro -
ptera), with new records and remarks on taxonomy. Russian
Journal of Theriology, 4: 43–62.
MATVEEV, V. A., and G. CSORBA. 2007. Bat fauna of Cambodia.
Bat Research News, 48: 100–101.
MCBEE, K., J. W. BICKHAM, S. YENBUTRA, J. NABHITABHATA,
New species of Murina from Vietnam 229

and D. SCHLITTER. 1986. Standard karyology of nine species
of vespertilionid bats (Chiroptera: Vespertilionidae) from
Thai land. Annals of Carnegie Museum, 55: 95–116.
MILNE-EDWARDS, A. 1868–74. Memoire sur la faune mammal -
ogique du Tibet oriental et principalement de la principauté
de Moupin. Pp. 231–379, in Recherches pour servir à l’his-
toire naturelle des mammifères: comprenant des considéra-
tions sur la classification de ces animaux (H. MILNE-ED -
WARDS), 2 volumes. G. Masson, Paris.
MÜLLER, J., K. MÜLLER, C. NEINHUIS, and D. QUANDT. 2010.
PhyDE – Phylogenetic data editor, version 0.9971. Released
November 23, 2010, http://www.phyde.de.
ONO, T., and Y. OBARA. 1994. Karyotypes and Ag-NOR varia-
tions in Japanese vespertilionid bats (Mammalia: Chiro -
ptera). Zoological Science, 11: 473–484.
OSGOOD, W. H. 1932. Mammals of the Kelley-Roosevelts and
Delacour Asiatic Expeditions. Publications of Field Muse -
um of Natural History, Zoological Series, 18: 193–339.
PAYNER, R. W. 1987. Genstat 5 reference manual. Clarendon
Press, Oxford, 749 pp.
PETERS, W. 1872. Mittheilung über neue Flederthiere. Monats -
bericht der Königlich Preussischen Akademie der Wissen -
schaften zu Berlin, 1872: 256–264.
POSADA, D. 2008. jModelTest: Phylogenetic model averaging.
Molecular Biology and Evolution, 25: 1253–1256.
REYMENT, R. A. 1971. Multivariate normality in morphometric
analysis. Mathematical Geology, 3: 357–368.
RONQUIST, F., M. TESLENKO, P. VAN DEER MARK, D. L. AYRES,
A. DARLING, S. HOHNA, B. LARGET, L. LIU, M. A. SUCHARD,
and J. P. HUESENBECK. 2012. MrBayes 3.2: efficient Bayes -
ian phylogenetic inference and model choice across a large
model space. Systematic Biology, 61: 539–542.
RUEDI, M., J. BISWAS, and G. CSORBA. 2012. Bat from the wet:
two new species of tube-nosed bats (Chiroptera: Vespertilio -
nidae) from Meghalaya, India. Revue Suisse de Zoologie,
119: 111–135.
SCULLY, J. 1881. On the mammals of Gilgit. Proceedings of
the Zoological Society of London, 1881: 197–209.
SIMMONS, N. B. 2005. Order Chiroptera. Pp. 312–529, in Mam -
mal species of the World: a taxonomic and geographic ref-
erence, 3rd edition (D. E WILSON and D. M. REEDER, eds.).
Johns Hopkins University Press, Baltimore, MD, 2142 pp.
SMITH, A. T. and Y. XIE. 2008. A guide to the mammals of Chi -
na. Princeton University Press, Princeton, 544 pp.
SOISOOK, P. 2011. A checklist of bats (Mammalia: Chiroptera) in
Thailand. Journal of Wildlife in Thailand, 18: 121–151.
SOISOOK, P., S. KARAPAN, C. SATASOOK, and P. J. J. BATES.
2013a. A new species of Murina (Mammalia: Chiroptera:
Vespertilionidae) from peninsular Thailand. Zootaxa, 3746:
567–579.
SOISOOK, P., S. KARAPAN, C. SATASOOK, V. D. THONG, F. A. A.
KHAN, I. MARYANTO, G. CSORBA, N. FUREY, B. AUL, and P.
J. J. BATES. 2013b. A review of the Murina cyclotis complex
(Chiroptera: Vespertilionidae) with descriptions of a new
species and subspecies. Acta Chiropterologica, 15: 271–292.
SON, T. N., M. MOTOKAWA, T. OSHIDA, V. D. THONG, G. CSORBA,
and H. ENDO. 2015. Multivariate analysis of the skull size
and shape in tube-nosed bats of the genus Murina (Chiro -
ptera: Vespertilionidae) from Vietnam. Mammal Study, 40:
79–94.
SWOFFORD, D. L. 2003. PAUP*: phylogenetic analysis using
par simony (*and other methods), version 4. Sinauer
Associates, Sunderland, Massachusetts.
TATE , G. H. H. 1941. Results of the Archbold expeditions no.
40: notes on vespertilionid bats of the subfamilies Minio -
pterinae, Murininae, Kerivoulinae and Nyctophilinae.
Bulle tin of the American Museum of Natural History, 78:
567–597.
TEMMINCK, C. J. 1835–1841. Monographies de mammalogie ou
description de quelques genres de mammifères, dont les es-
pèces dont été observées dans les différens musées de
l’Europe. Volume 2. C.C. van der Hoek, Leiden, 392 pp.
THOMAS, N. M., J. W. DUCKWORTH, B. DOUANGBOUBPHA, M.
WIL LIAMS, and C. M. FRANCIS. 2013. A checklist of bats
(Mam malia: Chiroptera) from Lao PDR. Acta Chiroptero -
logica, 15: 193–260.
THOMAS, O. 1891. Diagnoses of three new mammals collected
by Signor L. Fea in the Carin Hills, Burma. Annali del Mu -
seo Civico di Storia Naturale di Genova, 30 (2, 10): 884.
THOMAS, O. 1914. New Asiatic and Australian bats and a new
bandicoot. Annals Magazine Natural History, 13: 439–444.
THOMAS, O. 1915. A special genus for the Himalayan bat known
as Murina grisea. Annals and Magazine of Natural History,
8, 16: 309–310.
THOMAS, O. 1923. Scientific results from the mammal survey.
XLI. On the forms contained in the genus Harpiocephalus.
Journal of the Bombay Natural History Society, 29: 8889.
THONG, V. D. and N. M. FUREY. 2008. The bat fauna of Cat Ba
Biosphere Reserve. Journal of Biology, 30: 73–77 [In Viet -
namese with English summary].
THONG, D. V., D. C. DIETZ, A. DENZINGER, P. J. J. BATES, N. M.
FUREY, G. CSORBA, G. HOYE, L. D. THUY, and H.-U.
SCHNITZ LER. 2011. Further records of Murina tiensa from
Vietnam with first information on its echolocation calls.
Hystrix, the Italian Journal of Mammalogy, 22: 129–138.
TORDOFF, A. W., T. M. HIEU, T. Q. NGOC, E. STERLING, P. SWEET,
T. C HESSER, B. K. KEVIN, J. EGER, B. LIM, D. HARDER, et al.
2000. A feasibility study for the establishment of Ngoc Linh
Nature Reserve, Quang Nam Province, Vietnam. Birdlife
Inter national Vietnam Programme, Hanoi, Vietnam, 74 pp.
TRAI, L. T., W. J. RICHARDSON, B. D. TUYEN, L. V. CHAM, N. H.
DUNG, H. V. HOACH, A. L. MONASTYRSKII, and J. C. EAMES.
1999. An investment plan for Ngoc Linh Nature Reserve,
Kon Tum Province, Vietnam: a contribution to the manage-
ment plan. BirdLife International Vietnam Programme,
Hanoi, Vietnam, 106 pp.
TRAI, T. L., L. V. CHAM, T. Q. NGOC, T. M. HIEU, N. V. SANG, A.
L. MONASTYRSKII, B. D. HAYES, and J. C EAMES. 2000. An
investment plan for Kon Ka Kinh Reserve, Gia Lai prov -
ince, Vietnam: A contribution to the management plan.
Bird Life International Vietnam Programme, Hanoi, 77 pp.
TU, T. V., R. CORNETTE, J. UTGE, and A. HASSANIN. 2015. First
records of Murina lorelieae (Chiroptera: Vespertilionidae)
from Vietnam. Mammalia, 79: 201–213.
VOLLETH, M. 2006. Karyotype analysis of Murina suilla and
Phoniscus atrox from Malaysia (Chiroptera: Murininae,
Keri voulinae). Lynx (Praha), 37: 275–284.
WU, H., T.-L. JIANG, R. MÜLLER, and J. FENG. 2015. The allom-
etry of echolocation call frequencies in horseshoe bats:
nasal capsule and pinna size are the better predictors than
forearm length. Journal of Zoology (London), 297: 211–219.
WU, Y., M. MOTOKAWA, Y. LI, M. HARADA, Z. CHEN, and W. YU.
2010. Karyotype of Harrison’s tube-nosed bat Murina
harrisoni (Chiroptera: Vespertilionidae: Murininae) based on
the second specimen recorded from Hainan Island, China.
Mammal Study, 35: 277–279.
230 N. Truong Son, G. Csorba, V. Tan Tu, V. Dinh Thong, Y. Wu

New species of Murina from Vietnam 231
XU, Z., W. YU, Y. WU, F. LI, B. CHEN, M. HARADA, M. MOTA -
KAWA, Y. GONG, and Y. LI. 2014. Preliminary study on pop-
ulation genetic structure and taxonomy of Elery’s tube-
nosed bat (Murina eleryi). Acta Theriologica Sinica, 34:
270–277. [In Chinese with English abstract].
ZHOU, Q., Y. ZHANG, M. MOTOKAWA, M. HARADA, Y. GONG,
Y. L I, and Y. WU. 2011. A new record bat Murina hutto-
ni from Guangdong, China and its morphology, karyotypes,
echolocation calls. Chinese Journal of Zoology, 46(1): 109–
114. [In Chinese with English abstract].
ZHOU, Q., Z. XU, W. YU, F. LI, B. CHEN, Y. GONG, M. HARADA,
M. MOTOKAWA, Y. LI, and Y. WU. 2014. The occurrence of
bat Harpiocephalus harpia from Nanlin, Guangdong and its
karyotypes, echolocation calls. Chinese Journal of Zoology,
49(1): 41–45. [In Chinese with English abstract].
Received 29 July 2015, accepted 05 November 2015
1. Lao Cai (Hoang Lien NP); 2. Ha Giang (Duc Xuan area;
Vi Xuyen area; Khau Ca NR); 3. Tuyen Quang (Na Hang NR);
4. Bac Kan (Ba Be NP); 5. Cao Bang (Phia Oac-Phia Den NR);
6. Bac Kan (Kim Hy NR); 7. Thai Nguyen (Than Sa NR);
8. Lang Son (Huu Lien NP); 9. Quang Ninh (Bai Tu Long NP);
10. Hai Phong (Cat Ba NP); 11. Vinh Phuc (Tam Dao NP; Me
Linh and Tam Dao NP); 12. Phu Tho (Xuan Son NP); 13. Son
La (Co Ma area; Ta Sua NR); 14. Son La (Thuan Chau); 15. Son
La (Xuan Nha NR); 16. Thanh Hoa (Pu Hu NR); 17. Thanh Hoa
(Pu Luong NR); 18. Ninh Binh (Cucc Phuong NP); 19. Thanh
Hoa (Xuan Lien NR); 20. Thanh Hoa (Ben En NP); 21. Nghe
An (Pu Huong NR); 22. Nghe An (Pu Mat NP); 23. Ha Tinh
(Vu Quang NP); 24. Quang Binh (Phong Nha-Ke Bang NP);
25. Quang Tri (Bac Huong Hoa NR; Dak Rong NR); 26. Thua
Thien-Hue (Bach Ma NP); 27. Quang Nam (Song Thanh NR);
28. Quang Nam (Ngoc Linh NR); 29. Kon Tum (Ngoc Linh
NR); 30. Quang Ngai (Ba To area); 31. Kon Tum (Chu Mom
Ray NP); 32. Gia Lai (Kon Ka Kinh NP); 33. Binh Dinh and
Phu Yen (Hoa Son area); 34. Dak Lak (Yok Don NP); 35. Dak
Lak (Chu Yang Sin NP); 36. Lam Dong (Bi Dup-Nui Ba NP);
37. Khanh Hoa (Hon Ba NR); 38. Ninh Thuan (Nui Chua NP);
39. Dong Nai (Cat Tien NP); 40. Dong Nai (Vinh Cuu NR);
41. Binh Phuoc (Bu Gia Map NP); 42. Ba Ria-Vung Tau (Con
Dao NP); 43. Kien Giang (Phu Quoc NP).
APPENDIX I
Localities for the specimens of the subfamily Murininae from Vietnam used in this study. Locality numbers correspond with Fig. 1
APPENDIX II
Specimens of the subfamily Murininae used in this study. Asterisk (*) indicates specimens examined for karyotype
M. kontumensis sp. nov. (n = 1) — Kon Tum [29]: IEBR-
M5697* (holotype) (F)
M. leucogaster (n= 2) — Nghe An (Pu Mat NP) [22]:
HZM1.31758 (F). China: Sichuan: IBHG10122 (M).
M. harrisoni (n = 21) — Son La [15]: HNHM 2010.42.1
(M); Phu Tho [12]: Thong Coll.T2, IEBR T.290708.7 (M); Bac
Kan [6]: HZM.2.38178 (holotype of tiensa) (F), HNHM
2007.28.1 (paratype of tiensa), HZM NF 301006.1 (M); Vinh
Phuc [11]: HNHM 2009.6.2 (F), IEBR-M4998 (M); Hai Phong
[10]: IEBR T.220408.2 (F); Thanh Hoa [19]: IEBR-M6033 (F),
Nghe An [22]: IEBR-M3299; HZM 1.31525 (F); Dak Lak [34]:
ROM 107750 (F), 107739, 107749 (M); Cambodia: HZM
1.36316 (holotype), CBC 01290 (F); China: Hainan: IBGH
8295 (F); China: Guangxi: ROM 116463 (F), ROM 116468
(M); Thailand: SMF 53218 (M).
M. fionae (n= 7) — Quang Nam [27]: IEBR-M3080 (M);
Kon Tum [31]: IEBR-M5075 (M); Gia Lai [32]: IEBR-M3588
(M); Quang Ngai [30]: IEBR-M3635 (F), M3902, M3917 (M);
Dong Nai [40]: HNHM 22858 (M).
M. huttoni (n= 27) — Lao Cai [1]: ZMMU S186525, IEBR-
M5434, M5435*, M5437* (F), ZMMU S186700, IEBR-
M5428*, M5480*, M5482 (M); Cao Bang [5]: IEBR-M6023;
Quang Tri [25]: IEBR-M3153 (F); Kon Tum [29]: IEBR-
M5644, M5693* (F), VN11 1543, M5640*, M5641, M5643,
M5653, M5696 (M); Dak Lak [35]: HNHM 22885 (F); Lam
Dong [36]: IEBR-M5407*, M5413, M5415*, M5416 (F),
M4718, M5419 (M); Khanh Hoa [37]: ZMMU S175150,
ZMMU S175151 (M).
M. cyclotis (n= 75) — Son La [14, 15]: IEBR-M3046,
M4678 (M); Phu Tho [12] IEBR-M4052*, M4071* (M); Ha
Giang [2]: IEBR-M4753* (M); Cao Bang [5]: IEBR M5631,
M5630, M6001 (F); IEBR M5627, M5628, M5629, M6000
(M); Bac Kan [4] IEBR-M1210, M4561 (F), M4562 (M); Tuyen
Quang [3]: IEBR-M0492, M5394 (F), M0505, M1887, M1891,
M4976, M5296, M5309, IEBR VN11-1562 (M); Vinh Phuc
[11]: IEBR-M4560 (F); Quang Ninh [9]: IEBR-M2855, M3128,
M3131 (F), M3129, M3132, M3133, M3134 (M); Ninh Binh
[18]: HNHM22919/2008.23.1 (F); Thanh Hoa [16, 17, 19]
IEBR-M3725, M3726, M4223*, M6034, M6039, M6040 (F),
M3736, M3738, M4126, M6036, M6038, M4172 (M); Nghe An
[21, 22]: HNHM 22925, IEBR-M1359, M1632, M3316, M3330
(F), M1390, M2180, M4119, M4120, M3054, M3069, M3320
(M); Quang Binh [24]: IEBR-M3868, M3872, M3876 (F),
M3874, M3875, M5333 (M); Quang Tri [25]: IEBR-M4953 (F),
M4942* (M); Kon Tum [29]: IEBR-M5338 (M); Gia Lai [32]:
IEBR-M3607 (M); Lam Dong [36]: IEBR-M5776 (M); Quang

232 N. Truong Son, G. Csorba, V. Tan Tu, V. Dinh Thong, Y. Wu
Ngai [30]: IEBR-M3645, M3646, M3651 (F), M3632 (M); Phu
Yen [33]: IEBR-M4187 (M); Dong Nai [40]: IEBR-M4591 (M);
Binh Phuoc [41]: ZMMU S184674 (F).
M. lorelieae (n= 10) — Kon Tum [29]: VN11-1161, 1223,
IEBR-M5656 (F); VN11-1220, IEBR-M5648*, M5651,
M5662, M5663, HNHM B20140915.5, 20140915.7* (M)
M. annamitica (n= 22) — Son La [15]: IEBR-M3034 (F),
M2997 (M); Lao Cai [1] IEBR-M5429 (F), ZMMU S184673
(M); Tuyen Quang [3] IEBR-M4508 (M); Thanh Hoa [17]
IEBR-M4122, M4124 (M); Nghe An [21, 22] IEBR-M2181 (F),
HNHM 22929, IEBR-M1600, M3327 (M); Quang Tri [25]
IEBR-M3148, M3167 (F); Kon Tum [29] IEBR-M4131 (F);
Quang Ngai [30] IEBR-M3630, M3639, M3640, M3643,
M4718 (F), M3633, M3650, M3652 (M).
M. beelzebub (n= 10) — Quang Tri [25]: IEBR-M3636,
M4842*, M5645, M5760 (paratype), VN11-1586, HNHM
2007.50.7 (paratype) (F), HNHM 2007.50.24 (holotype) (M);
Kon Tum [29]: IEBR Tu071211.1, M5645 (F), IEBR-M4149,
M5646 (M); Quang Ngai [30] IEBR-M3636 (F), M3904 (M).
M. walstoni (n= 9) — Yok Don [34]: IEBR-M1480, HNHM
22933 (F), IEBR-M1481 (M); Ninh Thuan [38]: IEBR-M6030,
M6031, M6032 (F); Dong Nai [40]: IEBR-M4592 [F); Kien
Giang: IEBR-M2479, M2920 (M).
M. feae (n= 38) — Son La [14]: HNHM 2010.42.2, IEBR-
M4679 (F); Ha Giang [2] IEBR-M3264 (M); Tuyen Quang [3]:
IEBR-M0495 (F), IEBR-M504, M5350, (M); Bac Kan [4]:
IEBR-M4563 (F); IEBR-M323, M4510, HNHM 2000.84.7
(M); Vinh Phuc [11]: IEBR-M4991, M5056 (M); Ninh Binh
[18]: IEBR-M5054 (F); Thanh Hoa [16, 17, 19]: HNHM
2000.84.4, IEBR-M3718, M3728, M4123, M4127, M4214*,
VN110495 (F), HNHM 2000.84.7, IEBR-M3722, M4121,
M4125 (M); Nghe An [21, 22]: IEBR-M1360, M1387.1/22868
(F), IEBR-M1363, M1364, M3068 (M); Ha Tinh [23]: IEBR-
VN110001, 0007; Quang Binh [24]: IEBR-M3867, M3870,
M3871 (F), M3869, M3873 (M); Quang Tri [25] IEBR-M3154,
M4116 (M); Kon Tum [29]: IEBR-M5719 (M); Dong Nai [39]:
IEBR-M323/22860 (M).
M. eleryi (n= 19) — Son La [14]: IEBR-T.241107.1 (M);
Phu Tho [12] IEBR-M4070 (M); Ha Giang [2]: NF.250506.1
(M); Cao Bang [5]: IEBR-M5622 (F); M6024 (M); Bac Kan [6]:
BMNH 2008.25, ROM-NF.240507.1, HZM.1.39006,
NF.230707.1, 240707.2, HNHM 2007.28.2 (paratype) (F),
HNHM 2007.51.1 (holotype), NF.170906.3, 030707.1 (M);
Thanh Hoa [19]: IEBR-M6035 (M); Quang Binh [30]: IEBR-
M3866 (F); Kon Tum [29]: IEBR-M5718 (F); Quang Ngai [35]:
IEBR-M4511 (F), IEBR-M3644 (M).
M. harpioloides (n= 3) — Lam Dong [44]: ZMMU
S173401 (F), IEBR-M5806 (F), IEBR-M5860 (M).
M. chrysochaetes (n= 2) — Cao Bang [5]: IEBR-M6020
(F), Lao Cai [13]: ZMMU S186699 (F).
Harpiocephalus harpia (n= 5) — Cao Bang [5]: IEBR-
M6037 (M); Tuyen Quang [3]: IEBR-M422 (F); Nghe An [21]:
IEBR-M1362, M1391 (F); Kon Tum (Ngoc Linh NR): IEBR-
M5661* (M).
Harpiola isodon (n= 1) — Lao Cai [1]: IEBR-M5436* (F).
APPENDIX II. Continued
Murina harrisoni: Vietnam: HM540980, 540981, 540982;
Laos: HM540983; China: HM540984. M. leucogaster: China:
HM540987. M. fionae: Vietnam: HM540966; Laos:
HM540965. M. huttoni: Vietnam: JQ601542, KF772782; Laos:
HM540976; China: HM540978, 540979, JQ601452, 601454,
601455. M. cf. cyclotis: Vietnam: JQ601536, HM540953,
JQ601535, JQ601538, 601544, KF772775, 772776; Laos:
HM540940, 540941, 540944, 540945, 540946, 540947,
540952; China: HM540948, 540949, 540950, 540951; India:
HM540939. M. lorelieae: Vietnam: KF772780; China:
JN082179. M. annamitica: Laos: HM540967, 540968, 540971,
JQ601528, 601530, 601531, 601532, 601533, 601534. M. wal-
stoni: Laos: HM540958. M. feae and M. beelzebub (examined
by Francis et al. [2012] as M. feae): Vietnam: HM541000,
JQ601539, 601540, 601541, KF772777; Laos: HM540993,
540994, 540995, 540997, 540999; China: JQ601463, 601519,
601526. M. eleryi: Vietnam: HM540933; Laos: HM540931;
China: HM540935, 540937. M. harpioloides: Vietnam:
HM540975. M. chrysochaetes: China: HM540986, JQ601461,
601464, 601468, 601469, 601478, 601496, 601518, 601524,
601525. M. aenea: Malaysia: ADQ50994, HM540929. M. bal-
aensis: Thailand: PSUZC MM2012-214. M. gracilis: Taiwan:
KJ198513, 198514, 198527, 198538, 198540, 198542, 198556,
198565, 198570. M. guilleni: Thailand: HM540955. M. hilgen-
dorfi: China: JF442833; Russia: JF442834, 442835, 442836,
442839, 442840, 442841. M. peninsularis: Malaysia:
HM540972, 540973. M. recondita: Taiwan: KJ198578, 198583,
198592, 198603, 198619. M. shuipuensis: China: JN082180. M.
suilla: Malaysia: HM540989, 540990, 540991. M ussuriensis:
Russia: JF442842, 442843, 442844, 442848, 442850.
Harpiocephalus harpia: Laos: HM540283. Harpiola isodon:
Vietnam: HM540286. Myotis muricola: Vietnam: HM914942.
Kerivoula cf. hardwickii: Vietnam: HM540687.
APPENDIX III
GenBank/BOLD accession numbers for published COI sequences used in this study