Biodi e si y Da a Jou nal 13: e165516
doi: 10.3897/BDJ.13.e165516
Resea ch A icle
S udy o ba di e si y (Mammalia, Chi op e a) in
Xuan Nha Na u e Rese e, Son La P o ince, no h-
wes e n Vie nam, based on in eg a i e insigh s
om mo phology, gene ics and echoloca ion da a
Yen H Vu , Hai T Bui , Toan T Giang , Luong K Vu , Luong T Nguyen , Masaha u Mo okawa ,
Son T Nguyen
‡ Uni e si y o Science, Vie nam Na ional Uni e si y (VNU), Hanoi, Vie nam
§ Ins i u e o Biology, Vie nam Academy o Science and Technology (VAST), Hanoi, Vie nam
| Vie nam Na ion Uni e si y o Fo es y, Hanoi, Vie nam
¶ The Kyo o Uni e si y Museum, Kyo o Uni e si y, Kyo o, Japan
# G adua e Uni e si y o Science and Technology, Vie nam Academy o Science and Technology (VAST), Hanoi, Vie nam
Co esponding au ho : Son T Nguyen ( uongsonieb @gmail.com)
Academic edi o : K izle Tanalgo
Recei ed: 17 Jul 2025 | Accep ed: 24 Oc 2025 | Published: 04 No 2025
Ci a ion: Vu Y, Bui H, Giang T, Vu L, Nguyen L, Mo okawa M, Nguyen S (2025) S udy o ba di e si y
(Mammalia, Chi op e a) in Xuan Nha Na u e Rese e, Son La P o ince, no h-wes e n Vie nam, based on
in eg a i e insigh s om mo phology, gene ics and echoloca ion da a. Biodi e si y Da a Jou nal 13: e165516.
h ps://doi.o g/10.3897/BDJ.13.e165516
Abs ac
This s udy p esen s he esul s o ba di e si y su eys in Xuan Nha Na u e Rese e,
no h-wes e n Vie nam. A o al o 114 indi iduals, ep esen ing 19 species belonging o
ou amilies, we e eco ded. The Rhinolophidae amily was he mos species- ich,
con ibu ing eigh species o he o al di e si y, ollowed by Hipposide idae, wi h ou
species. Eigh species, including Rhinolophus episcopus,R. siamensis, R. c . episcopus,
R. pe nige , Hipposide os g i ini, Megae ops niphanae, Tylonyc e is onkinensis and
Myo is mu icola, we e newly eco ded o Xuan Nha NR. Species ichness was high, bu
e enness was low, wi h Hipposide os pou ensis and Rhinolophus pea sonii domina ing
cap u es. Ele en species we e obse ed, ep esen ed by only one indi idual. The
mo phological examina ions, wi h suppo om echoloca ion calls and p elimina y
gene ic analysis, e ealed he p esence o mo phologically conse ed and po en ially
‡,§ ‡ | ‡ § ¶
§,#
© Vu Y e al. This is an open access a icle dis ibu ed unde he e ms o he C ea i e Commons A ibu ion License (CC BY 4.0),
which pe mi s un es ic ed use, dis ibu ion, and ep oduc ion in any medium, p o ided he o iginal au ho and sou ce a e
c edi ed.
c yp ic axa. Rep oduc i e da a indica ed ea ly we -season b eeding o se e al species.
Compa ed o p e ious su eys in he egion, ou s udy subs an ially expands he
knowledge o chi op e an auna in Xuan Nha. Gi en he high p opo ion o habi a -
specialis and mon ane-associa ed species and he documen ed p esence o a
conse a ion-p io i y axon, Hipposide os g i in, in Xuan Nha NR, con inued biodi e si y
su eys inco po a ing molecula and acous ic me hods a e essen ial o e ine species
in en o ies and o p o ide in o ma ion o conse a ion s a egies o his biologically
impo an landscape.
Keywo ds
biodi e si y, Chi op e a, cy och ome b, species ichness
In oduc ion
Vie nam, loca ed in Indochina, is ecognised as a ho spo coun y and amongs he
wo ld's iches egions o mammal di e si y. Such biodi e si y is a ibu ed o he
coun y's complex opog aphy, di e se clima ic condi ions and dis inc ecosys ems
(S e ling e al. 2006, To do e al. 2012). No he n Vie nam, in pa icula , wi h nume ous
p o ec ed a eas, se es as impo an e ugia o bo h widesp ead and endemic ba
species. Xuan Nha Na u e Rese e (NR), in Xuan Nha Commune o Son La P o ince
includes bo h p ima y and seconda y e e g een o es s, wi h limes one a eas
in e spe sed amongs hem (The People’s Commi ee o Son La P o ince 2019).
Rega ding ba di e si y, Nguyen e al. (2012) eco ded a o al o 19 species a his a ea;
ne e heless, da a on en i onmen al condi ions, as well as de ailed mo phological
cha ac e is ics and gene ic in o ma ion o he eco ded species, emain una ailable.
In Augus 2024, wi hin he amewo k o he Nagao NEF P ojec , we conduc ed an 11-day
ield su ey in Xuan Nha NR. The esul s signi ican ly expand cu en knowledge o ba
di e si y in he egion by inc easing he numbe o eco ded species and p o iding
unde s andings o undocumen ed axa. This pape aims o upda e he chi op e an auna
o Xuan Nha NR by p o iding in o ma ion ela ed o mo phological cha ac e is ics,
echoloca ion calls and gene ic in o ma ion; and hen we discuss species composi ion
and di e si y in no h-wes e n Vie nam o suppo u u e ecological and axonomic
esea ch in he egion.
Ma e ial and me hods
S udy a ea
Xuan Nha NR (20°36'–20°48'N, 104°29'–104°50'E) is posi ioned in he sou h-wes e n
a ea o Son La P o ince, no h-wes e n Vie nam, bo de ing Laos o he wes (Fig. 1) and
is one o ou na u e ese es in Son La (The People’s Commi ee o Son La P o ince
2019). The Rese e is domina ed by opical o es s and in e spe sed wi h subs an ial
2Vu Y e al
a eas o limes one o es ecosys ems (To do e al. 2004). Pha Luong Moun ain (1970 m
a.s.l.) is he highes peak and loca ed on he moun ainous idge ha delinea es he
Vie nam–Laos bo de , se ing as an impo an ecological co ido o mon ane auna
(Cen e o Na u e Conse a ion and De elopmen (CEV) 2005).
Along he mid-ele a ion ansec s (500–800 m a.s.l.), he su eyed ou es c ossed a eas
hea ily al e ed by an h opogenic ac i i y. Remnan pa ches o p ima y o es we e
agmen ed and embedded in a ma ix o cul i a ed lands and seconda y ege a ion (Fig.
2c). Dense hicke s o low- unk ac s p edomina e be ween 700 and 900 m a.s.l., while
limes one ou c ops was obse ed nea a ba ca e a app ox. 910 m a.s.l. Abo e 1000 m
a.s.l., undis u bed o es pa ches pe sis along he slopes (Fig. 2a). These o es s ha e a
canopy heigh o 20–30 m wi h a dense unde s o y o saplings, climbing ines and dense
sh ubs. A ele a ions abo e 1500 m a.s.l., nea ly p is ine mon ane moss o es s emain,
cha ac e ised by high canopy ees (20–50 m), abundan epiphy es and closed canopy
co e . The egion has coole empe a u es and high humidi y. S eams o med by
p ecipi a ion pa e ns and seasonal uno a e ypically 1.5–4.5 m wide and less han 0.7
m deep.
Su ey me hods and sampling
The main ield su ey was conduc ed om 4-15 Augus 2024, wi h addi ional da a om a
wo-day su ey on 25-26 Ma ch 2010. Ba s we e su eyed using mis ne s and ha p aps
(Tidemann and Woodside 1978) se along ou ele a ional bands (ca. 600, 700, 900 and
1100 m a.s.l.) (Fig. 1). Six een apping si es we e es ablished, each ope a ed o 1–2
nigh s depending on wea he and accessibili y. Ha p aps (1.8 m high × 1.2 m wide) wi h
Figu e 1.
Map o Vie nam showing he loca ion o Xuan Nha NR wi hin Son La P o ince. Red do s
indica e ba sampling si es. The map was o ganised using QGIS 3.38.3 (h ps://www.qgis.o g).
S udy o ba di e si y (Mammalia, Chi op e a) in Xuan Nha Na u e Rese e, ... 3
ou pa allel ows o mono ilamen lines (2.5 cm spacing) we e deployed a each si e
(Tidemann and Woodside 1978). T aps we e s a egically placed along ba lyways such
as: o es ails, idgelines, s eams and ca e en ances (Fig. 3a, b and d). A leas one
ha p ap was ins alled pe si e and se be o e dusk, emaining open un il midnigh o
la e depending on ba ac i i ies. T aps we e checked e e y 30–45 minu es,
synch onised wi h mis ne inspec ions, o minimise s ess and inju y o cap u ed
indi iduals. Mis ne s we e deployed a g ound le el ac oss p esumed lyways. Two ne
sizes we e used: 9 × 3 m and 12 × 3 m, cons uc ed om black nylon wi h ou shel es
(Fig. 3c). Ne s we e moun ed on ibe glass o in combina ions wi h locally sou ced
bamboo poles. Sampling was conduc ed om 18:00 h o 23:00 h and 04:00 h o 05:00 h,
wi h 1–3 ne s ope a ed pe si e pe nigh . Ne s we e closed du ing he day o a oid
acciden al cap u es o bi ds o li es ock. Geog aphic coo dina es and ele a ions we e
eco ded using he Gaia GPS (WGS84 da um).
Echoloca ion eco dings and analyses
Echoloca ion calls we e eco ded using an Echo Me e Touch 2 ul asonic de ec o
(Wildli e Acous ics, Inc., Mayna d, Massachuse s, USA) connec ed o he sma phone
de ice. Reco dings we e ob ained unde wo condi ions: (1) handheld eco dings made
immedia ely a he mis ne si e o cap u ed indi iduals; (2) con olled eco dings inside a
empo a y ligh en (4 × 4 × 2 m). All eco dings we e analysed isually using
spec og ams o cha ac e ise call s uc u e and equency pa ame e s. Echoloca ion calls
we e compa ed wi h e e ences om p e ious s udies in Vie nam (Nguyen e al. 2021,
Győ össy e al. 2024) o assis species iden i ica ion, pa icula ly o c yp ic axa o
species di icul o cap u e.
Figu e 2.
Habi a na u es in Xuan Nha NR. (a) T opical e e g een o es a 1000 m a.s.l.; (b) Mixed
habi a including o es edges and open a eas a a ound 600 m a.s.l.; (c) Dis u bed seconda y
o es a 900 m a.s.l.
4Vu Y e al
Specimen examina ion
Mos specimens (109 indi iduals) we e collec ed du ing he main su ey in 2024, while
only i e indi iduals we e cap u ed on Ma ch 2010, wi h de ailed in o ma ion o each
specimen p esen ed in Suppl. ma e ial 1. Pho og aphs o li e specimens we e aken
using a Canon EOS Kiss X7 digi al came a wi h an EF-S 18–55 mm /3.5–5.6 ki lens.
S anda d mo phome ic measu emen s we e ollowed Wilson and Mi e meie (2019).
The ep oduc i e condi ion o emales was assessed ollowing Racey (2009) and age
class was de e mined by he deg ee o phalangeal epiphyseal usion (B une -Rossinni
and Wilkinson 2009). P egnancy in emales was checked by gen le abdominal palpa ion.
Specimens we e p ese ed in 95% e hanol du ing bo h he 11-days su ey in 2024 and
2-days in 2010. Upon ans e o he labo a o y, e hanol concen a ion was educed o
70% o long- e m s o age. Skulls we e ex ac ed and cleaned and one o wo indi iduals
pe species we e selec ed o gene ic analyses. C anioden al cha ac e s we e measu ed
unde a s e eoscopic mic oscope (SMZ 745, Nikon) using an elec onic digi al calipe
(Mi u oyo NTD12-15PMX, 0.01 mm p ecision). A o al o 20 me ics we e measu ed
ollowing Vu e al. (2024) (Fig. 4; Suppl. ma e ial 2). Specimens we e iden i ied, based on
ex e nal and c anial mo phology, supplemen ed by DNA analysis and echoloca ion da a,
using compa a i e e e ences (Bo issenko and K uskop 2003, Vu 2012,K uskop 2013,
Vuong e al. 2015, Vuong e al. 2017a, Vuong e al. 2017b, Vuong e al. 2018, Mo a elli e
al. 2019, Vu e al. 2024). Vouche specimens and associa ed issue samples a e cu en ly
deposi ed in he Depa men o Zoology, Ins i u e o Biology (IB), VAST, Hanoi, Vie nam.
Figu e 3.
Ba apping deploymen : (a, b, d) Double-ha p aps se along o es ails and o e s eams;
(c) Se up o mis ne s.
S udy o ba di e si y (Mammalia, Chi op e a) in Xuan Nha Na u e Rese e, ... 5
Molecula da a and phylogene ic analyses
DNA was isola ed om issue samples p ese ed in 99% e hanol using he DNeasy®
Blood & Tissue Ki (Qiagen, Hilden, Ge many). The mi ochond ial cy och ome b (Cy b)
gene was a ge ed o species-le el iden i ica ion. PCR ampli ica ion employed p ime
pai So iF/So iR (Bui e al. 2020a) wi h he ollowing he mal cycling condi ions: ini ial
dena u a ion a 95°C o 5 min; 35 cycles o 95°C o 30 s, 55°C o 50 s and 72°C o 2
min; and a inal ex ension a 72°C o 10 min. PCR p oduc s we e pu i ied and
sequenced by 1s BASE (Selango , Malaysia) using Sange sequencing.
Ch oma og ams we e edi ed and assembled in Ch omas P o (Technelysium P y L d.,
Aus alia) and MEGA 11 (Tamu a e al. 2021). Sequences we e aligned wi h MUSCLE in
MEGA 11. The inal alignmen o he Cy b agmen was 1140 bp. Phylogene ic analyses
we e pe o med using Maximum Likelihood (ML) unde he GTR+G+I model, selec ed by
he Bayesian In o ma ion C i e ion (BIC) in ModelFinde implemen a ion in IQ-TREE .
1.6.12 (Nguyen e al. 2015). Node suppo was e alua ed wi h 10,000 ul a as boo s ap
eplica es (Hoang e al. 2018). The esul ing phylogene ic ees we e isualised using
FigT ee .1.4.4 and edi ed using Adobe Pho oshop 2023.
Da a esou ces
Da a package i le: Occu ence da ase o ba s (Mammalia, Chi op e a) om Xuan Nha
Na u e Rese e, Son La P o ince, no h-wes e n Vie nam.
Resou ce link: h ps://doi.o g/10.15468/7d54s5
Figu e 4.
(a) Do sal, (b) en al, (c) la e al iews o he c anium; (d) la e al iews o mandible displaying
c anioden al measu emen s. The diag am is cons uc ed, based on he skull mo phology o P.
enuis species.
6Vu Y e al
Numbe o da ase s: 1
Da a se name: Occu ence da ase o ba s (Mammalia, Chi op e a) om Xuan Nha
Na u e Rese e, Son La P o ince, no h-wes e n Vie nam.
Da a o ma : Da win Co e E en
Desc ip ion: This da ase p esen s species occu ence o ba s collec ed om Xuan Nha
NR, Son La P o ince, no h-wes e n Vie nam. Field su eys we e conduc ed in
Augus
2024 as pa o he biodi e si y esea ch p og amme wi h he suppo o he Nagao
Na u al En i onmen Founda ion (NEF). Fo e e y occu ence eco d, he da ase
p o ides in o ma ion on loca ion (GPS coo dina es), da e, sex, ep oduc i e condi ion and
ap ype.
Resul s
Species ichness, di e si y and dis ibu ion
A o al o 114 ba indi iduals we e collec ed du ing wo ield su eys in Xuan Nha NR,
ep esen ing 19 species om nine gene a and ou amilies: Hipposide idae,
P e opodidae, Rhinolophidae and Vespe ilionidae. The Rhinolophidae was he mos
species- ich amily, wi h eigh species o one genus (R. pea sonii, R. pe nige , R. a inis,
R. pusillus, R. episcopus, R. siamensis, R. homasi and R. c . episcopus). This was
ollowed by Hipposide idae, wi h ou species o wo gene a (A. s oliczkanus, H. a mige ,
H. g i ini and H. pou ensis) and Vespe ilionidae, wi h i e species o ou gene a (K. c .
dongduongana, M. al ic ania us, M. mu icola, P. enuis and T. onkinensis). The
P e opodidae amily was ep esen ed by wo species o wo gene a (C. sphinx and M.
niphanae) (Table 1). H. pou ensis was he mos abundan species, wi h 39 indi iduals
(34.2%). The second mos common was R. pea sonii (32 indi iduals, 28.1%). O he
equen ly cap u ed species included R. homasi (7), C. sphinx (8) and M. niphanae (7). In
con as , ele en species we e ep esen ed by a single indi idual, including R. pusillus, R.
a inis, R. episcopus, R. siamensis, R. c . episcopus, R. pe nige , H. g i ini, M.
al ic ania us, M. mu icola, T. onkinensis and T. enuis.
Cap u e success a ied by ap ype: ha p aps accoun ed o 88 indi iduals (77.2%),
p o ing e ec i e o Rhinolophus and Hipposide os species, while mis ne s (26
indi iduals, 22.8%) exclusi ely cap u ed ugi o ous axa, such as C. sphinx and M.
niphanae. Species e enness was low, wi h cap u es domina ed by a ew abundan axa.
Signs o ep oduc i e ac i i y, including p egnancy, lac a ion o he p esence o
newbo ns, we e obse ed in 15 emales ac oss i e species, while 54 emales showed
no e idence o b eeding.
S udy o ba di e si y (Mammalia, Chi op e a) in Xuan Nha Na u e Rese e, ... 7
No. Speci ic name n
(M♂,
F♀)
Rep oduc i e
in o ma ion
Ele a ion (m) Habi a
na u e
Species
eco ded
in his
s udy
P e ious
Reco d
(Nguyen e
al. 2012)
IUCN
S a us
Hipposide idae Lydekke , 1891
1Hipposide os a mige
(Hodgson, 1835)
3 (2♂,
1♀)
1♀ (No
ep oduc i e)
2 (650 m), 1
(750 m)
2✓ ✓ LC
2Hipposide os g i ini
Vu, Puechmaille,
Denzinge , Die z,
Cso ba, Ba es, Teeling
& Schni zle , 2012
1 (1♂,
0♀)
— 1 (650 m) 2 ✓* — NT
3Hipposide os pou ensis
Allen, 1906
39
(21♂,
18♀)
3♀ (3 Lac a ing),
15♀ (No
ep oduc i e)
1 (600 m), 4
(650 m), 8 (700
m), 15 (750 m),
7 (800 m), 2
(850 m), 2 (910
m)
1, 2, 3,
4
✓ ✓ LC
4Hipposide os gen ilis
Ande sen, 1918
—✓LC
5Aselliscus s oliczkanus
(Dobson, 1871)
3 (0♂,
3♀)
3♀ (No
ep oduc i e)
1 (750 m), 2
(800 m)
1, 2 ✓ ✓ LC
P e opodidae B isson, 1762
6Cynop e us sphinx
(Vahl, 1797)
8 (2♂,
6♀)
3♀ (2 Lac a ing, 1
P egnan , 1 wi h
newbo n), 3♀ (No
ep oduc i e)
1 (600 m), 5
(700 m), 2 (750
m)
2, 4 ✓ ✓ LC
7Megae ops niphanae
Yenbu a & Fel en,
1983
7 (3♂,
4♀)
2♀ (2 Lac a ing),
2♀ (No
ep oduc i e)
3 (600 m), 2
(700 m), 2 (750
m)
2, 4 ✓* — LC
8Sphae ias blan o di
(Thomas, 1891)
—✓LC
Rhinolophidae G ay, 1825
9Rhinolophus a inis
Ho s ield, 1823
1 (0♂,
1♀)
1♀ (No
ep oduc i e)
1 (750 m) 2 ✓ ✓ LC
Table 1.
Lis o ba species eco ded om Xuan Nha NR. n (♂, ♀) = sample size (♂ = male, ♀ = emale);
Ele a ion: n (m) = numbe o indi iduals wi h ele a ion in me es. Reco ded habi a : 1 = e e g een
o es , 2 = dis u bed seconda y o es , 3 = ca e a eas, 4 = s eam alley. Species eco ds: ✓ =
eco ded, ✓* = newly eco ded in he 2024, — = No eco ded.
8Vu Y e al
No. Speci ic name n
(M♂,
F♀)
Rep oduc i e
in o ma ion
Ele a ion (m) Habi a
na u e
Species
eco ded
in his
s udy
P e ious
Reco d
(Nguyen e
al. 2012)
IUCN
S a us
10 Rhinolophus episcopus
Allen, 1923
1 (1♂,
0♀)
— 1 (600 m) 2 ✓* — LC
11 Rhinolophus siamensis
Gyldens olpe, 1917
1 (1♂,
0♀)
— 1 (750 m) 2 ✓* — LC
12 Rhinolophus pea sonii
Ho s ield, 1851
32
(5♂,
27♀)
6♀ (5 Lac a ing, 1
P egnan ), 21♀
(No ep oduc i e)
1 (500 m), 4
(600 m), 1 (650
m), 1 (700 m),
20 (750 m), 5
(800-850 m)
2, 4 ✓ ✓ LC
13 Rhinolophus pusillus
Temminck, 1834
1 (1♂,
0♀)
— 1 (650 m) 2 ✓ ✓ LC
14 Rhinolophus c .
episcopus Allen, 1923
1 (1♂,
0♀)
— 1 (750 m) 2 ✓* — LC
15 Rhinolophus homasi
K. Ande sen, 1905
7 (5♂,
2♀)
1♀ (1 Lac a ing),
1♀ (No
ep oduc i e)
2 (600 m), 5
(800 m)
2, 4 ✓ ✓ LC
16 Rhinolophus ouxii
Temminck, 1835
—✓LC
17 Rhinolophus pe nige
Hodgson, 1843
1 (0♂,
1♀)
1♀ (No
ep oduc i e)
1 (600 m) 2 ✓* — LC
Vespe ilionidae G ay, 1821
18 Ke i oula c .
dongduongana Vuong
e al. 2018
4 (0♂,
4♀)
4♀ (No
ep oduc i e)
4 (900 m) 1 ✓ ✓ LC
19 Mu ina cyclo is Dobson,
1872
—✓LC
20 Myo is al ic ania us
Osgood, 1932
1 (0♂,
1♀)
1♀ (No
ep oduc i e)
1 (800 m) 2 ✓ ✓ LC
21 Myo is mu icola (G ay,
1864)
1 (0♂,
1♀)
1♀ (No
ep oduc i e)
1 (950 m) 2 ✓* — LC
22 Pipis ellus enuis
(Temminck, 1840)
1 (1♂,
0♀)
— 1 (800 m) 2 ✓ ✓ LC
23 Pipis ellus ab amus
(Temminck, 1840)
—✓LC
S udy o ba di e si y (Mammalia, Chi op e a) in Xuan Nha Na u e Rese e, ... 9
Cha ac e Species
(n)
H. a mige
(3)
H. g i ini
(1)
H. pou ensis
(39)
A.
s oliczkanus
(3)
C. sphinx
(8)
M. niphanae
(7)
M M W 7.14–7.34
7.25±0.07
11.18 5.06–5.2
5.14±0.06
4.55 5.79 5.02 5.38
ML 12.57–
12.79
12.68±0.08
22.98 9.13–9.49
9.31±0.16
8.68 10.43 8.54 8.81
CPH 2.61–2.73
2.66±0.05
5.72 2.83–3.05
2.95±0.1
2.06 3.11 2.24 2.42
cm L 7.33–7.69
7.54±0.13
13.48 5.42–5.51
5.47±0.04
4.66 5.65 4.45 4.23
cp L 2.32–2.79
2.55±0.17
5.38 2.33–2.39
2.36±0.03
1.87 2.07 1.43 1.21
p m L 5.67–6.07
5.86±0.16
10.03 — 3.27 4.02 3.41 3.12
m m L 4.93–5.18
5.05±0.1
8.16 — 2.95 3.37 2.85 2.81
Hipposide os pou ensis Allen, 1906
In he s udy, 39 indi iduals o H. pou ensis (21♂, 18♀) we e eco ded (Table 1, Suppl.
ma e ial 1). Amongs emales, only h ee showed ac i e ep oduc i e s a us as lac a ing,
3 3
3
4
4 3
1 3
Figu e 6.
(Le ) C anium o H. pou ensis (A–C) and A. s oliczkanus (a–c) in do sal (A, a), la e al (B, b)
and en al (C, c) iews; mandible in do sal (D, d) and la e al (E, e) iews; (Righ ) Phylogeny
based on Cy b sequences o Hipposide idae ba s. Boo s ap suppo alues (BS) a e shown
a nodes.
16 Vu Y e al
while he emaining 15 we e non- ep oduc i e. H. pou ensis is a medium-sized lea -
nosed ba (Fig. 7a), cha ac e ised by a slende skull compa ed o H. a mige o H. g i ini,
wi h a de eloped sagi al c es , a na ow b aincase and a sligh ly swollen os um.
The specimen clus e ed wi hin he H. pou ensis lineage on he phylogene ic ee, wi h
s ong suppo and showed he closes gene ic ela ionship wi h specimens collec ed
om Bai Tu Long Island (OP142137), ollowed by popula ion om Ca Ba Island
(OP142143–OP142145) (Fig. 6, igh ). Pai wise gene ic di e gence anged be ween 2%
and 3.6%. These indings suppo he ecen axonomic dis inc ion o H. pou ensis om H.
la a us as epo ed by Yuze o ich e al. (2022), wi h bo h mo phological and molecula
analyses con i ming i s s a us as he dis inc species. Based on eco dings o a single
male indi idual, his species emi ed na owband CF-FM calls (Fig. 5, igh ) wi h a peak
equency o 85.6 ± 0.7 kHz, s a ing a 88.7 kHz and ending a 70.7 kHz. Mean call
du a ion was 6.2 ± 0.2 ms, wi h he longes call las ing 6.5 ms.
Aselliscus s oliczkanus (Dobson, 1871)
A. s oliczkanus is widely dis ibu ed ac oss no h-wes e n and cen al Vie nam (Vuong e
al. 2015). In his s udy, h ee specimens we e iden i ied as A. s oliczkanus, based on
diagnos ic ex e nal and c anial ai s. These ba s a e small-sized, wi h a icuspid
pos e io nose-lea , sho bodies and bicolou ed do sal u (whi e bases and b own ips),
while he en al u is pale (Fig. 7b). C anial ea u es include a sligh ly swollen os um
and a ela i ely elonga ed snou compa ed o membe s o Hipposide idae (Fig. 6, le ).
The newly-ob ained specimens clus e ed wi h e e ence A. s olickzanus sequences (BS =
89–98%) (Fig. 6, igh ). Gene ic di e gence be ween he s udy specimens and known
e e ences anged om 0.5% o 1.5%.
Figu e 7.
Selec ed Hipposide idae species eco ded on Xuan Nha NR: (a) H. pou ensis; (b) A.
s olickzanus.
S udy o ba di e si y (Mammalia, Chi op e a) in Xuan Nha Na u e Rese e, ... 17
Family P e opodidae
Two species o ui ba s we e eco ded du ing his su ey (Table 1). C. sphinx, p e iously
epo ed om he Rese e, was e-con i med, while M. niphanae ep esen s a new eco d
o his locali y. Bo h species exhibi ed signs o ep oduc i e ac i i y du ing Augus .
Cynop e us sphinx (Vahl, 1797)
C. sphinx is common, occu ing ac oss lowland and edge habi a s. Adul s show sho
o ange-b own do sal pelage, a g eyish en al side and a da ke man le egion, mo e
p onounced in males. Ju eniles a e pale wi h an o e all g eyish one. The ea s a e
b own wi h a dis inc i e whi ish ma gin and he in e emo al memb ane is na ow, bu
e iden , wi h a sho ail ex ending sligh ly beyond i . Compa ed wi h Megae ops, C.
sphinx is la ge -bodied, mo e obus , wi h a mo e de eloped in e emo al memb ane and
a diagnos ic whi e ea ma gin (absen in Megae ops). C anially, he skull is elonga e,
na owing an e io ly (Fig. 8), wi h expanded zygoma ic a ches and a b oad, elonga ed
pala e. The den i ion consis s o obus mola s wi h sligh ly ounded cusps and elonga ed
uppe canines.
Megae ops niphanae Yenbu a & Fel en, 1983
This is a small p e opodid ba . The species is easily ecognised by i s so , ligh b ownish-
g ey pelage and absence o a ail. The ligh memb anes a e pale g ey wi h weak
pigmen a ion, while he ea s, muzzle and limbs a e pale b ownish, gi ing a sub ly
anslucen appea ance. C anially, he skull is sho (Fig. 8), wi h na ow zygoma ic
a ches and a cons ic ed pos o bi al egion. The pala e is smoo h and lacks s ong idging
and he mola s a e less obus han hose o C. sphinx. O ou adul emales cap u ed in
mid-ele a ion o es s, wo we e lac a ing.
Figu e 8.
Do sal (A, a), la e al (B, b), en al (C, c) iews o he c anium and Do sal (D, d), la e al (E, e)
iews o he mandible o C. sphinx and M. niphanae, espec i ely.
18 Vu Y e al
Family Rhinolophidae
Eigh species o Rhinolophidae we e documen ed. Fou species (R. episcopus, R.
siamensis, R. c . episcopus and R. pe nige ) a e newly eco ded, while R. a inis, R.
homasi, R. pea sonii and R. pusillus had p e iously been epo ed by Nguyen e al.
(2012). R. ouxii, lis ed in ea lie su eys, was no de ec ed. This species is now
conside ed es ic ed o eas e n Asia; he e o e, i s p e ious eco d om Xuan Nha NR is
likely a misiden i ica ion. Rep oduc i e e idence was obse ed in R. homasi and R.
pea sonii.
Rhinolophus a inis Ho s ield, 1823
Only a emale R. a inis was eco ded showing no signs o ep oduc i e ac i i y. This
species is a medium-sized ho seshoe ba . Ex e nally, i has a b oad ho seshoe wi h a
deep median ema gina ion, ounded connec ing p ocess and a mode a ely con ex sella
lacking basal lappe s; he lance is sub iangula wi h an un educed ip. The pelage is
so , da k g eyish-b own. The skull is small, wi h well-de eloped la e al nasal
compa men s and na ow in e o bi al cons ic ion. The den i ion shows a sligh ly educed
P wi hin he oo h ow. In compa ison wi h R. homasi (Fig. 9, le ), al hough bo h species
sha e skull shape ypical o he “R. megaphyllus” g oup (I h e al. 2015), R. a inis has a
sligh ly la ge c anium, a p opo ionally longe mandible, b oade os um and mo e
la e ally expanded a ches. Thei den al dimensions (C C W, M M W) a e o he wise
simila . Molecula da a place he specimen wi hin he R. a inis clade, wi h pai wise
gene ic dis ances o 2.5–2.75% (Fig. 9, igh ).
2
1 1 3 3
Figu e 9.
(Le ) Do sal (A, a, a’), la e al (B, b, b’), en al (C, c, c’) iews o he c anium and Do sal (D,
d, d’), la e al (E, e, e’) iews o he mandible o R. a inis, R. homasi and R. pusillus; (Righ )
ML ee based on Cy b sequences o Rhinolophydae ba s.
S udy o ba di e si y (Mammalia, Chi op e a) in Xuan Nha Na u e Rese e, ... 19
Rhinolophus homasi K. Ande sen, 1905
One lac a ing emale o R. homasi was obse ed. Compa ed o R. a inis, R. homasi has
sligh ly smalle zygoma ic, b aincase b ead hs and i s mandible is hinne and sho e .
Ex e nally, i shows a na ow, ec angula ho seshoe and a b oad, blun ly poin ed lance .
The pelage is uni o mly ligh g ey wi h a me allic sheen, dense and el e y, wi h pale
bases and da ke ips (Fig. 10a).
Echoloca ion calls o a emale exhibi ed b oadband FM–CF–FM s uc u e domina ed by
CF componen s, wi h a peak equency o 77.4 ± 0.2 kHz, s a ing a 78.6 kHz and ending
a 66.7 kHz. Calls a e aged 19.6 ± 0.3 ms in du a ion, wi h he longes call las ing 20.3
ms (Fig. 10d).
Rhinolophus pea sonii Ho s ield, 1851
A o al o 32 indi iduals o R. pea sonii we e cap u ed and we e he mos common
species o Rhinolophus in he su ey. Six emales we e ep oduc i ely ac i e ( i e
lac a ing, one p egnan ). This species has a b oad os um and a well-de eloped
b aincase. C anial dimensions include GTL 22.78–23.85 mm, CM L 8.97–9.78 mm and
ML 16.27 ± 0.28 mm. The mola s a e ma ked by high, sha p cusps (Fig. 10c’).
Echoloca ion calls o a male eco ded in a en exhibi ed b oadband FM-CF-FM s uc u e
(Fig. 10d), wi h peak equency 56.9 ± 0.7 kHz, anging om 39.6 o 58.9 kHz. Calls
a e aged 21.6 ± 0.3 ms in du a ion, wi h he longes call las ing 22.3 ms.
Rhinolophus pe nige Hodgson, 1843
P e iously, R. pe nige was conside ed a subspecies wi hin he R. luc us complex;
howe e , s udies o Volle h e al. (2017) demons a ed di e ences be ween R. pe nige
3
Figu e 10.
Selec ed Rhinolophydae species eco ded: (a) R. homasi; (b) R. pusillus; (c) R. pea sonii;
(c’) Fi e aspec s o he c anium and mandible o R. pea sonii; (d) Ul asonic echoloca ion calls
o Rhinolophids.
20 Vu Y e al
and R. luc us sensu s ic o. In his s udy, a single emale was cap u ed and clus e ed wi h
e e ence R. pe nige sequences, showing 1.2% gene ic di e gence (Fig. 9, igh ). The
indi idual was non- ep oduc i e. R. pe nige is a la ge ho seshoe ba (FA 73.4 mm, W 48
g) wi h a massi e skull (GTL 33.54 mm). The ho seshoe-shaped nose-lea is b oad,
co e ing he uppe lip wi h a deep median no ch. The u is hick, da k g ey do sally and
en ally. Den i ion is obus , wi h well-de eloped oo h ows. Echoloca ion calls eco ded
om he cap u ed emale we e b oadband FM–CF–FM (Fig. 10d), wi h peak equency
31.9 ± 0.7 kHz, s a ing a 32.4 kHz and ending a 22.7 kHz. Calls a e aged 33.4 ± 0.7 ms
in du a ion, wi h he longes call las ing 35.3 ms.
Rhinolophus pusillus Temminck, 1834
This species is amongs he smalles ho seshoe ba s (W 4.2 g, FA 35.4 mm, CCL 11.54
mm), wi h p opo ionally small ea s and nose-lea and weakly de eloped supplemen a y
lea le s (Fig. 10b). The lance is elonga ed wi h a sligh o wa d bend a he ip. The
pelage is ine and silky, anging om ligh b own o g eyish-b own do sally wi h a pale
unde side. The skull is small and delica e (Fig. 9, le ), wi h a sho , na ow os um,
in la ed b aincase and slende zygoma ic a ches. The den i ion is compac , wi h small,
closely-spaced canines and p emola s; he mandible is gen ly cu ed wi h diminu i e
lowe ee h. In he phylogene ic analysis (Fig. 9, igh ), he specimen g ouped wi hin he
R. pusillus clade; howe e , pai wise di e gence om Vie namese R. pusillus popula ions
is high, anging om 2.0% o 2.5%. This di e gence shows a ce ain gene ic
di e en ia ion o he s udied indi idual ela i e o Vie namese R. pusillus’s popula ions.
Rhinolophus episcopus Allen, 1923
The male R. episcopus has mo phome ic measu emen s: HB 41.8 mm, TL 21.9 mm, HF
8.5 mm, EL 23.4 mm, FA 42.1 mm and W 6 g. The ba has ligh b own pelage, la ge ea s
and a well-de eloped nose-lea . The ho seshoe is b oad, co e ing he muzzle, wi h small
la e al lea le s and a isible median no ch. The lance is elonga ed wi h con ex ma gins
and a ounded ip (Fig. 11). The an e io median swellings a e p ominen and elonga ed,
while he pos e io swellings a e sho . The sagi al c es is weakly de eloped and he
on al dep ession is shallow. Sup ao bi al c es s a e well-de ined wi h sha p idges.
In he R. mac o is complex, c anioden al di e ences can be de ec ed e en amongs
sympa ic axa. Compa a i e measu emen s be ween R. episcopus and R. c . episcopus
specimens collec ed om he same locali y e ealed ha R. episcopus exhibi s smalle
CCL (13.79 mm s. 14.16 mm) and BCH (6.73 mm s. 6.95 mm) ela i e o R. c .
episcopus. Den al measu emen s such as CM L (6.38 mm s. 6.71 mm) and CP L (2.85
mm s. 3.05 mm) a e smalle in R. episcopus. Con e sely, R. episcopus displays a
b oade in e o bi al wid h (IOW). In his s udy, Cy b sequencing was a emp ed o he
specimen, bu he ob ained agmen (~ 500 bp) was insu icien in leng h and quali y o
be included in phylogene ic analyses. Howe e , based on ex e nal mo phology and
c anioden al ai s, he iden i ica ion o he specimen as R. episcopus is conside ed
eliable. The echoloca ion call o his male was a b oadband FM-CF-FM s uc u e (Fig. 10
d). The maximum s a equency was 65.6 kHz and he minimum alue was 58.2 kHz,
3 4
S udy o ba di e si y (Mammalia, Chi op e a) in Xuan Nha Na u e Rese e, ... 21
wi h peak ene gy a 63.4 ± 0.5 kHz. The mean call du a ion was 25.2 ± 0.6 ms, wi h he
longes call las ing 26.8 ms.
Rhinolophus siamensis Gyldens olpe, 1917
R. siamensis ini ially desc ibed as a subspecies o R. mac o is, has since been ele a ed
o species le el, wi h a wide dis ibu ion in Sou heas Asia (Vuong e al. 2017b, Hu son e
al. 2019). This indi idual ep esen s a small-sized membe o he mac o is complex. The
FA is 37.3 mm, HB is 40.7 mm and he W is only 3.07 g, all lowe han co esponding
alues o R. c . episcopus. The skull o R. siamensis is no only sho e , bu also mo e
g acile, shows a na owe b aincase, educed os um wid h and a sho e mandible
(Table 3). Mo phologically, R. siamensis has ela i ely la ge ea s, abou hal he o ea m
leng h and a b oad ho seshoe co e ing he muzzle wi h a dis inc median no ch. The
sella p ojec s o wa ds and he connec ing p ocess is b oad and ounded, gi ing he
nose-lea a s uc u e dis inc om o he small ho seshoe ba s. I s pelage is so and
woolly, b own do sally and pale en ally.
Rhinolophus c . episcopus Allen, 1923
In his s udy, R. c . episcopus was ep esen ed by a single adul male. Ex e nal
measu emen s included FA o 43.2 mm, E o 23.1 mm and W o 4.51 g. The ea s we e
la ge, app oxima ely hal he leng h o he o ea m and he pelage was so , woolly and
b own do sally wi h a sligh ly pale en al su ace. The c anial p o ile is b oad, wi h MAW
o 8.91 mm and ZYW o 8.17 mm. The mandible is well-de eloped, wi h a ML o 10.12
mm. The skull mo phology shows a mo e hea ily buil c anial s uc u e compa ed o o he
membe s o he mac o is g oup (Fig. 11). Molecula analysis placed he specimen wi hin
he R. mac o is complex clade. I clus e ed mos closely wi h indi iduals p o isionally
iden i ied as R. c . episcopus om Vie nam in he s udy o Vuong e al. 2017b, wi h a
Figu e 11.
Do sal (A, a, a’), la e al (B, b, b’), en al (C, c, c’) iews o he c anium and Do sal (D, d, d’)
iews o he mandible o R. episcopus, R. c . episcopus and R. siamensis, espec i ely.
22 Vu Y e al
gene ic di e gence o app oxima ely 2.5–3%. This le el o di e gence o e s a close
ela ionship, while also showing he possibili y o egional di e en ia ion wi hin R. c .
episcopus (Liu e al. 2019).
Family Vespe ilionidae
Se e al species epo ed by Nguyen e al. (2012), including P. ab amus, P. co omand a,
P. ja anicus, Ia io,T. ul ida ( e e ed o as T. pachypus) and Mu ina cyclo is, we e no
de ec ed in he p esen su ey. Ne e heless, ou s udy expands he species in en o y o
Xuan Nha NR by p o iding new eco ds o T. onkinensis and M. mu icola.
Ke i oula c . dongduongana Vuong, Hassanin, Fu ey, Nguyen & Cso ba, 2018
Acco ding o Vuong e al. (2018) and o he s udies, K. ha dwickii sensu la o in Vie nam
has been di ided in o ou smalle , closely- ela ed species ha a e mo phologically
di icul o dis inguish. Based on ex e nal and c anioden al measu emen s, ou specimen
is en a i ely assigned o K. dongduongana; howe e , molecula analysis is necessa y o
con i m i s p ecise species iden i y. Mo phologically, i shows so , long u , wi h smoky-
b own do sally and ligh e g eyish-b own en ally. HB anges 34.3–40.4 mm, FA 33.5–
35.4 mm, wi h b ownish-black anslucen wing memb anes. K. c . dongduongana
p esen s a domed b aincase wi h a conca e on al p o ile. C anial measu emen s (GTL
14.03–14.14 mm, ML 6.65–6.75 mm) poin ou a small and compac c anial s uc u e
compa ed o la ge Ke i oula species (Vuong e al. 2018, Liang e al. 2023).
Myo is al ic ania us Osgood, 1932
M. al ic ania us was once conside ed a subspecies o M. siligo ensis in Vie nam, bu
mo e ecen axonomic s udies by Ruedi e al. (2021) ha e cla i ied i s s a us as a dis inc
species. Acco ding o hese s udies, M. siligo ensis is es ic ed o he Cen al and
Eas e n Himalayas, whe eas ba s occu ing u he eas , including popula ions in China
and Indochina, a e now assigned o M. al ic ania us. A emale specimen ep esen s a
well-p ese ed example o his small espe ilionid. I has a delica e skull wi h a low
os um, domed b aincase, s eep on al p o ile and sligh ly swollen occipi al egion. The
zygoma ic a ches a e hin and inwa dly conca e. Do sal u is bu o da k b own, wi h
ligh e g eyish-b own en ally. Ea s a e long and na ow, eaching o su passing he
muzzle ip. The uppe p emola s ha e a p ominen size di e ence, pa icula ly be ween P
and P (Fig. 12). The ba has a delica e mandible wi h small lowe p emola s (p ) and i s
lowe mola s a e semi-nyc alodon ypes.
Myo is mu icola (G ay, 1864)
A emale was collec ed a 950 m a.s.l. The pelage is so and dense, pale b ownish-g ey
do sally and di y whi e en ally wi h da k hai oo s (Fig. 13, le ). Ea s a e long and
na ow, he agus slende and o wa d-ben and he ee small wi h sho claws. The skull
is small wi h an elonga ed os um, la ened b aincase and de eloped sagi al c es (Fig.
13, igh ). The zygoma ic a ch is e enly cu ed and massi e. C a e wide and obus ,
exceeding he heigh o P . P a e well-de eloped and aligned in he oo h ow. P a e
2 4 4
1
4 2 3
S udy o ba di e si y (Mammalia, Chi op e a) in Xuan Nha Na u e Rese e, ... 23
small, displaced lingually and s and inwa ds on he inside o he uppe oo h ow. c a e
la ge, poin ed and sligh ly highe han p . Lowe mola s a e o he myo odon ype.
Gene ic analysis o Cy b con i med he specimen wi hin he M. mu icola clade, wi h 1–
1.5% di e gence om o he conspeci ic sequences (Fig. 14, le ).
1
4
Figu e 12.
Do sal (A), la e al (B), en al iew (C) o c anium; Do sal (D), la e al (E) o mandibles,
Occlusal iew o le uppe (G) and igh lowe (H) oo h ows o M. al ic ania us.
Figu e 13.
(Le ) M. mu icola eco ded on Xuan Nha NR; (Righ ) Do sal (A), la e al (B), en al iew
(C) o c anium; Do sal (D), la e al iew (E) o mandibles, Occlusal iew o le uppe (G) and
igh lowe (H) oo h ows o M. mu icola.
24 Vu Y e al
Pipis ellus enuis (Temminck, 1840)
Ex e nally, P. enuis has da k b own do sal pelage wi h a sligh ly pale en al side and a
sho b oad agus (Fig. 15a). The skull is small and delica e, wi h a na ow os um,
in la ed b aincase, weak c es s and simple bicuspid uppe canines (Fig. 15b).
Sequencing o he Cy b gene posi ioned he specimen i mly wi hin he enuis clade.
Pai wise gene ic dis ances anged om 1.0% o 1.7% compa ed o P. enuis indi iduals
p e iously collec ed om he Tay Con Linh Moun ain by K uskop e al. (2024) (Fig. 14,
le ).
Figu e 14.
(Le ) ML phylogene ic ee, based on Cy b sequences o he eco ded Vespe ilionidae ba s;
nodes wi h BS < 70% a e no displayed; (Righ ) Ul asonic echoloca ion call cha ac e is ics o
T. onkinensis.
Figu e 15.
(a) P. enuis eco ded on Xuan Nha NR; (b) Fi e aspec s o he c anium and mandible o P.
enuis.
S udy o ba di e si y (Mammalia, Chi op e a) in Xuan Nha Na u e Rese e, ... 25
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Supplemen a y ma e ials
Suppl. ma e ial 1: Ba da ase able
Au ho s: YHV, STN, HTB
Da a ype: occu ences
B ie desc ip ion: Ba da ase able om he ield su ey in Xuan Nha NR, Son La, Vie nam
(Ma ch 2010 & Augus 2024). Dash (–): No de e mined. Habi a na u e: 1 = E e g een o es , 2
= Dis u bed seconda y o es , 3 = Ca e a eas, 4 = S eam alley.
Download ile (44.13 kb)
Suppl. ma e ial 2: Lis o c anioden al measu emen s used in his s udy
Au ho s: YHV, STN, HTB
Da a ype: mo phological
Download ile (19.07 kb)
S udy o ba di e si y (Mammalia, Chi op e a) in Xuan Nha Na u e Rese e, ... 33
