A summary of the taxonomy and distribution of the red giant flying squirrel, Petaurista petaurista (Sciuridae, Sciurinae, Pteromyini), in mainland Southeast Asia with the first record from Lao PDR

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DOI: 10.1515/mammalia-2014-0005
Cite this publication
Abstract
The occurrence of the red giant flying squirrel, Petaurista petaurista, in Lao LAO is confirmed on the basis of a single adult male specimen obtained from an informal food market in Thatlouang village, Xekong Province in the south of the country. This individual was reported to have been collected from close-by Thatlouang in the mixed deciduous forest or dry dipterocarp forest on the Bolaven Plateau. The record extends the known distribution of this species by approximately 700 km eastwards. Information is provided on the external, cranial, dental, and bacular characters of the new Lao specimen. It is compared with the holotypes and/or type descriptions of seven taxa, namely, barroni, candidula, cicur, melanotus, penangensis, taylori, and terutaus, described from mainland Southeast Asia, all of which are currently included in the synonymy of P. petaurista. On the basis of its external pelage colour, the Lao specimen is referred to P. p. barroni, which was described from southeast Thailand and is considered here to be a valid subspecies.
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Mammalia 2014; aop
Daosavanh Sanamxay * , Bounsavane Douangboubpha , Sara Bumrungsri , Chutamas Satasook
and Paul J.J. Bates
A summary of the taxonomy and distribution of
the red giant flying squirrel, Petaurista petaurista
(Sciuridae, Sciurinae, Pteromyini), in mainland
Southeast Asia with the first record from Lao PDR
Abstract: The occurrence of the red giant flying squir-
rel, Petaurista petaurista , in Lao PDR is confirmed on
the basis of a single adult male specimen obtained from
an informal food market in Thatlouang village, Xekong
Province in the south of the country. This individual was
reported to have been collected from close-by Thatlouang
in the mixed deciduous forest or dry dipterocarp forest on
the Bolaven Plateau. The record extends the known distri-
bution of this species by approximately 700 km eastwards.
Information is provided on the external, cranial, dental,
and bacular characters of the new Lao specimen. It is
compared with the holotypes and/or type descriptions of
seven taxa, namely, barroni , candidula , cicur , melanotus ,
penangensis , taylori , and terutaus , described from main-
land Southeast Asia, all of which are currently included in
the synonymy of P. petaurista. On the basis of its external
pelage colour, the Lao specimen is referred to P. p. barroni,
which was described from southeast Thailand and is con-
sidered here to be a valid subspecies.
Keywords: local market; red giant flying squirrel; South-
east Asia; taxonomy.
DOI 10.1515/mammalia-2014-0005
Received January 10 , 2014 ; accepted July 10 , 2014
Introduction
Lao People s Democratic Republic (Lao PDR) is home to a
rich variety of rare and endemic species and is considered
as a biodiversity hotspot ( Myers etal. 2000 ). In recent years,
the number of mammal species recorded from the country
has increased with the discovery of a range of unusual taxa,
including amongst the rodents, the Kha-nyou, Laonastes
aenigmamus Jenkins etal., 2005 ; the Lao limestone rat, Sax-
atilomys paulinae Musser etal., 2005 ; and, most recently,
the Laotian giant flying squirrel, Biswamoyopterus laoen-
sis Sanamxay etal., 2013 . The discovery of this latter taxon
increased to seven the number of flying squirrels recorded
from the country, the other six being Petaurista elegans
(Müller), P. philippensis (Elliot), Belomys pearsonii (Gray),
Hylopetes alboniger (Hodgson), Hylopetes phayrei (Blyth),
and Hylopetes spadiceus (Blyth) ( Thorington etal. 2012 ). In
addition, it has been suggested that two more species may
possibly occur in the north of the country, namely, P. petau-
rista ( Francis 2008 , Thorington etal. 2012 ) and Petinomys
setosus (Temminck) ( Duckworth etal. 1999 ).
Petaurista petaurista is currently known from Afghani-
stan, Pakistan, India, Nepal, South China, Myanmar, Thai-
land, Malaysia, Sumatra, Java, and Borneo ( Thorington
etal. 2012 ). Previously, Askins (1977) had considered the
range of P. petaurista to include all of Lao PDR, but this sup-
position was based on a different interpretation of the tax-
onomy, one which included philippensis and allied forms
as synonyms. Subsequently, Srikosamatara et al. (1992)
reported that P. petaurista had been provisionally identi-
fied in the That Louang fresh food market in the capital
*Corresponding author: Daosavanh Sanamxay, Faculty of
Environmental Sciences, National University of Laos, Dong Dok
Campus, P.O. Box 7322, Xaythany District, Vientiane Capital,
Lao PDR, e-mail: daosavanhsnx@gmail.com ; and Department of
Biology, Faculty of Science, Prince of Songkla University, Hat Yai,
Songkla 90112, Thailand
Bounsavane Douangboubpha: Faculty of Environmental Sciences,
National University of Laos, Dong Dok Campus, P.O. Box 7322,
Xaythany District, Vientiane Capital, Lao PDR ; and Department of
Biology, Faculty of Science, Prince of Songkla University, Hat Yai,
Songkla 90112, Thailand
Sara Bumrungsri: Department of Biology, Faculty of Science, Prince
of Songkla University, Hat Yai, Songkla 90112, Thailand
Chutamas Satasook: Department of Biology, Faculty of Science,
Prince of Songkla University, Hat Yai, Songkla 90112, Thailand ; and
Princess Maha Chakri Sirindhorn Natural History Museum, Prince of
Songkla University, Hat Yai, Songkla 90112, Thailand
Paul J.J. Bates: Harrison Institute, Centre for Systematics and
Biodiversity Research, Bowerwood House, St. Botolph s Road,
Sevenoaks, Kent, TN13 3AQ, UK
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2D. Sanamxay etal.: Red giant flying squirrel from Lao PDR
city, Vientiane. However, the authors most likely also fol-
lowed the taxonomic interpretation of Askins (1977) .
Corbet and Hill (1992) mapped the distribution of
Petaurista petaurista and included northwest Lao PDR but
without details. This record was probably based on that
of Askins (1977) , who included the forms candidula and
barroni (currently included in P. petaurista ) in the synon-
ymy of P. alborufus (Milne-Edward) and mapped the range
of this latter taxon as including northwest Lao PDR. Oshida
et al. (1992, 2004) suggested that the taxon occurred in
northern Lao PDR on the basis of four individuals, which
they referred to P. petaurista melanotus Gray, 1837 . However,
this latter record is not credible because this material was
obtained from a pet dealer in Japan and melanotus is a
distinct Sundaic form from peninsular Malaysia ( Figure
1 ). It should be noted that Timmins and Duckworth (2008)
stated that Lao PDR was known to be a source of export for
various Sundaic species that do not occur naturally in the
country. As such, P. petaurista is omitted from the faunal
list of the country by Duckworth etal. (1999) , Thorington
and Hoffmann (2005) , Francis (2008) , Timmins and Duck-
worth (2008), and Thorington etal. (2012) .
However, on the 18 April 2013 a dead, adult male
Petaurista was observed for sale in a local market in That-
louang village, Xekong Province, south Lao PDR. This
specimen is referred to P. petaurista and is here consid-
ered to be the first authenticated record from the country.
Materials and methods
Material
An adult male was obtained from Thatlouang village,
market in Xekong Province, south Lao PDR and is held in
the zoological collection of the Faculty of Environmental
Sciences (FES), National University of Laos, Lao PDR.
Measurements
Unfortunately, the skull was damaged, which restricted
the number of cranial measurements that could be taken.
Measurements (in mm) were taken with digital calipers
and followed Hayashida etal. (2006) , Helgen etal. (2009) ,
and Koyabu etal. (2009) . Mass (in grams) was taken using
a spring balance. Measurements comprise the following
characters: M: body mass; HB: head and body length
from tip of the nose to the anus, ventrally; T: tail length
from anus to tip of the tail, not including hairs; E: ear
length from the intertragal notch to the crown; HF:
hindfoot length from the extremity of the heel behind
the os calcis to the extremity of the longest digit, not
including the claw or the hair; ONL: occipitonasal length
( = greatest length of skull) from the tip of nasals to the
supraoccipital; CBL: condylobasal length from the tip of
gnathion to the occipital condyle; ZB: zygomatic breadth
the greatest breadth across the zygomatic arches; ZH:
zygomatic height from the bottom edge of the jugal to
the tip of the superior process of the jugal; BB: breadth
of braincase the greatest breadth of the braincase at
the posterior roots of the zygomatic arches; RB: rostrum
breadth the greatest breadth across the rostrum, includ-
ing the bony capsules enclosing the nasolacrimal canals;
NL: nasal length from the anterior tip of the nasal
bones to the most posterior suture between the nasal and
frontal bones; IOB: interorbital breadth taken across
the supraorbital notch; POB: postorbital breadth the
narrowest width across the constriction posterior to the
orbits; POPB: postorbital processes breadth from tip of
the right postorbital process to the tip of the left postor-
bital process; LIF: length of the incisive foramina from
the anterior border to the posterior border of the incisive
foramina; MWN: maximum width of nasals bone the
maximum width at the anterior end of the nasal bones;
P
3 M
3 : upper toothrow length from the front of the
third upper premolar to the back of the crown of the third
upper molar; M
2 M
2
: greatest palatal breadth from the
labial wall of the left second upper molar to labial wall of
the right second upper molar, across the palate; M
1 M
1 :
width of the bony palate at the first upper molar from
lingual wall of left first upper molar to the lingual wall
of right first upper molar; ML: mandible length from
the most posterior part of the condyle to the most ante-
rior part of the body of mandible, not including the lower
incisors; P
4 M
3 : lower toothrow length from the front of
the fourth lower premolar to the back of the crown of the
third lower molar; BL: baculum length from the base to
the tip of the baculum.
Results
Systematic description
Petaurista petaurista Pallas, 1766
Red Giant Flying Squirrel
Sciurus petaurista Pallas, 1766; west Java, Indonesia.
Restricted to Preanger Regencies by Robinson and Kloss
(1918)
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D. Sanamxay etal.: Red giant flying squirrel from Lao PDR3
Figure 1  The subspecies of Petaurista petaurista in mainland southeast Asia with a description of the dorsal and ventral pelage and tail
of each taxon. Black triangle represents the new locality in Lao PDR. Black circles are type localities of the taxa that are currently referred
to P. petaurista ( sensu Corbet and Hill 1992 , Thorington and Hoffmann 2005 ). Open circles are localities of P. petaurista collected from this
region, which are held in the Natural History Museum, London [BM(NH)] and Natural History Museum, National Science Museum Thailand
[NHM]. Numbers from 1 to 23 refer to localities listed in Appendix 1.
Synonyms from mainland Southeast Asia, as illus-
trated in Figure 1.
Pteromys melanotus Gray, 1837: 584; Selangor; origi-
nally Nepal in error.
Petaurista terutaus Lyon, 1907 : 17; Terutau (= Tarutao)
Island, Straits of Malacca, Thailand.
Petaurista candidula Wroughton, 1911 : 1014. Kindat,
western Myanmar.
Petaurista taylori Thomas, 1914 : 205; Bankasun,
southern Tennasserim, Myanmar.
Petaurista annamensis barroni Kloss, 1916 : 33; Hup
Bon, Sriracha, southeast Thailand.
Petaurista nitida cicur Robinson and Kloss, 1914 : 223;
Ban Kok Klap, Bandon, Thailand.
Petaurista petaurista penangensis Robinson and
Kloss, 1918 : 224; Penang Island, Malaysia.
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4D. Sanamxay etal.: Red giant flying squirrel from Lao PDR
New material
FES.MM.13.001, adult , skull extracted, dry skin, obtained
from a vendor at Thatlouang village, Lamam District,
Xekong Province, Lao PDR, approximately 15 ° 21 ′ N, 106 °
43 E (Figure 1). According to the vendor, the specimen was
obtained in the local area, probably in or adjacent to the
forests of the three former Proposed National Biodiver-
sity Conservation Areas (PNBCAs) , which comprise Phou
Theung, Bolaven northeast, and Phou Kathong.
External and bacular characters
The specimen is a large flying squirrel with a head and
body length of 435 mm, a tail length of 475 mm, and a
body mass of 1500 grams ( Table 1
). The pelage of the
upper surface comprises predominantly an admixture of
dark rust-coloured and black hairs, variably grizzled with
whitish grey. Individual hairs are variable in colour but
usually include the following components: metallic ash
at the base for up to 15 mm, rust coloured in the mid-part
for about 7 mm, and a black tip of about 2 mm. A distinc-
tive whitish-grey band, some 5mm in length, is present
in hairs that reside in areas of the upper surface that are
extensively grizzled. These areas include the upper neck
and back (except for the crown of the head), the flanks,
and on the rump extending on to the base of the tail.
The upper forearms are pale orange with some grizzling
( Figure 2 A).
The upper surface of each patagium is thickly haired
and is comparable in colour to the body but with little or
no grizzling. The upper surface of the interfemoral mem-
brane is also well haired, especially adjacent to the tail,
where it is heavily grizzled with white. The fore- and hind-
feet are covered in black hairs on their upper surfaces
(Figure 2A). The pads are dark. The claws form a half circle
with very sharp tips.
The ventral pelage is whitish grey. The throat is white
and the chin has a dark metallic-ash coloured patch. The
anterior margin of the forearms is deep orange, the colour
extending to the sides of the neck and the chin. The wrist,
pro-forearm, and styliform cartilage are black. The under-
sides of the patagia, forearms, and hindlegs are relatively
sparsely covered in brownish-orange hairs, especially in
comparison to the upper surface. This brownish-orange
colour extends to the interfemoral membrane, except for
the area of the tail, which is covered with whitish-grey
Table 1 Body mass (in grams), external, cranial, dental, and bacular measurements (in mm) of a single male Petaurista petaurista speci-
men from Lao PDR. Measurements of the comparative taxa are taken from the type descriptions.
Characters P. petaurista (from Lao
PDR) new specimen
P. p. candidula
Wroughton ()
P. p. barroni
Kloss ()
P. p. cicur Robinson
and Kloss ()
P. p. terutaus
Lyon ()
P. p. penangensis Robinson
and Kloss ()
M 
HB     
T    
E  
HF    
ONL  . . 
CBL  . 
ZB .   . . .
ZH .
BB .
RB .
NL . . . .
IOB .   . 
POB . . .
POPB . 
LIF .
MWN . 
P
M . . . .
M
M .
M
M .
ML .
P
M . 
BL .
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D. Sanamxay etal.: Red giant flying squirrel from Lao PDR5
hairs. The lateral margins of the patagia have hairs with
whitish-grey tips and brownish-grey bases. The interfem-
oral membrane has a well-developed fringe of pale grey
hairs above and whitish below. It is connected to about
one quarter of the basal part of the tail. The tail is cylin-
drical, not distichous, and from the outer border of the
interfemoral membrane towards the tip is entirely yellow-
ish mixed with some black hairs. The individual hairs are
brownish at the base, whitish grey in the mid-part, and
yellowish at the tip (Figures 2B and 2D).The tail tip is dark
brown for about 100 mm.
On the head, each ear is covered by short white hairs
anteriorly, including the anterior margin. The posterior
part of the ear is black. The base of the ear is thickly
haired; these individual hairs are white at the tip and
blackish basally. The eyes are ringed with black. The
cheeks are whitish grey; the hair bases are brownish grey.
The upper surface of the head is similar to the upper part
of the body. The nose is surrounded by short dark brown
hairs (Figure 2C).
A
B
CD
Figure 2  Views of skin of Petaurista petaurista (FES.MM.13.001)
from Lao PDR. (A) Dorsal view; (B) ventral view; (C) head, left lateral
view; and (D) ventral view of the left patagium and interfemoral
membrane to show edge of patagium and interfemoral membrane
(scale = 300 mm).
The baculum of Petaurista petaurista is essentially
straight and long, with the greatest length of 26.5 mm. It
has a large simple base and is slightly narrower at the tip,
where there is a spiral plate ( Figures 3 C, 3D, and 3E).
Cranial and dental characters
Unfortunately, the skull of the specimen was damaged
by the hunter and, as a consequence, some cranial char-
acters could not be observed. It is large with a zygomatic
breadth of 47.9mm and a breadth of braincase of 31.7 mm.
When viewed in lateral profile, it is relatively flat with
a low braincase. The orbital regions are large. In lateral
view, the zygomatic arches are low and the jugal bone is
greatly expanded anteriorly. When viewed from above,
the interorbital region is broad, exceeding the breadth
of the postorbital constriction. The rostrum is short and
wide. The frontal is deeply depressed. The postorbital pro-
cesses are large and relatively long (Figure 3A, Table 1).
The coronoid process of each half mandible is well
developed. The condylar process has a large dorsal
articular surface, and the post-condylar process is well
developed. The angular process is large and downwardly
projecting.
In the dentition, the upper incisors are orthodont. The
anterior surface of each tooth is pigmented with orange.
The small upper premolar (P
3 ) is relatively well developed
and is situated on the anterior border of the second upper
premolar (P
4 ). This latter tooth is similar in size to the
first (M
1 ) and second upper molars (M
2 ). The third upper
molar (M
3 ) is the smallest molar and circular in shape.
P
4 has three well-defined cusps on the labial side and one
large cusp on the lingual side. The three labial cusps are
connected to the large lingual cusp by transverse ridges,
which are separated by deep valleys. Both M
1 and M
2 have
three defined cusps on the labial side and one large cusp
on the lingual side; each labial cusp is connected to the
prominent lingual cusp by a transverse ridge; the postero-
lingual diagonal flexus is very well developed. M
3 has two
cusps on the labial side, the parastyle and paracone, of
which the parastyle is much reduced. On the lingual side
it has one large cusp; as with the other cheek teeth, the
cusps on the labial side are connected to the large lingual
cusp by low transverse ridges; the metacone is absent
(Figure 3B).
In the lower dentition, the incisor is moderately long
and is brownish orange on its anterior surface. The first
premolar (P
4 ) is the smallest tooth; it has two cusps on
the lingual side and three cusps on the labial side; the
anteroconid, protoconid, and entoconid are low crested;
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6D. Sanamxay etal.: Red giant flying squirrel from Lao PDR
the metaconid is the highest cusp; the hypoflexid is
very well developed and opens labially. All the molars
are essentially similar in structure. However, the third
lower premolar (M
3 ) is more elongated; it has two cusps
on the lingual side and three cusps on the labial side;
the anteroconid and entoconid are almost absent; there
is also a small cusp (mesoconid) between the two main
cusps on the labial side (protoconid and hypoconid)
(Figure 3B).
Comparison with other similar taxa
Petaurista petaurista is similar to P. elegans in terms of
external morphology, but differs in its larger size (in
P. elegans : HB: 340 365mm and T: 350 370 mm;Francis
2008). The ears of P. elegans are predominantly black and
A
B
CDE
Figure 3  (A) Views of skull, mandible (scale = 50 mm) and (B) dorsal
views of the left maxillary (left) and left mandibular (right) toothrows
(scale = 5 mm) of Petaurista petaurista (FES.MM.13.001); (C) dorsal
view, (D) lateral view, and (E) ventral view of bacula of Petaurista
petaurista (FES.MM.13.001), P. philippensis , and P. elegans , from left
to right, respectively (scale = 10 mm).
hairless, with dark tawny hairs at the base. In P. petau-
rista , the ears are white anteriorly and black posteriorly.
The dorsal pelage of P. elegans has prominent white spots
( Figure 4 A). The cheek and throat of P. elegans are pale
orange, but are white in P. petaurista . The skull of P. petau-
rista (ONL: 66 73 mm) is larger than that of P. elegans
(ONL: < 65 mm) ( Corbet and Hill 1992 ).
Petaurista petaurista is more similar in size to
P. philippensis (HB: 400 490 mm and T: 400 550 mm;
Francis 2008 ) but differs in pelage colour. Within the
Indochinese region, P. philippensis has a much darker
dorsal pelage that is extensively grizzled with white
(Figure 4B), whereas it is paler brown in P. petaurista and
the grizzling is less extensive (Figure 1A). The anterior
half of the ears is brownish orange in P. philippensis but
white in P. petaurista . In P.philippensis , the tail is either
all-dark or all-grey but in P.petaurista , it is yellowish with
a prominent dark brown or black tip (Figures 4B and 4C).
The skull of the P.petaurista specimen from Lao PDR is
damaged. However, it appears to be essentially similar in
size to that of P. philippensis (P 3 – M 3 : 15.0 – 17.8 mm; Corbet
and Hill 1992 ).
A
B
C
Figure 4 Views of skins. (A) Dorsal view of Petaurista elegans (FES.
MM.12.161); and (B) dorsal and (C) ventral views of P. philippensis
(FES.MM.12.152) from Lao PDR (scale = 300 mm).
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Taxonomic notes
The Lao specimen was compared with the holotypes and/
or original descriptions of seven taxa, which are currently
referred to Petaurista petaurista ( sensu Corbet and Hill
1992 , Thorington and Hoffmann 2005 ) and which origi-
nate from mainland southeast Asia (Figure 1). They are
listed above as synonyms of P. petaurista and comprise P.
p. candidula Wroughton, 1911 from northwest Myanmar;
P. p. barroni Kloss, 1916 from southeast Thailand; P. p.
taylori Thomas, 1914 from peninsular Myanmar; P. p. cicur
Robinson and Kloss, 1914 from peninsular Thailand; P. p .
terutaus Lyon, 1907 from Terutau Island, off the west coast
of peninsular Thailand; P. p. penangensis Robinson and
Kloss, 1918 from Penang Island, off the west coast of pen-
insular Malaysia; and P. p. melanotus Gray, 1837 from pen-
insular Malaysia. The Lao specimen differs from all these
taxa with the exception of P. p. barroni.
The taxon candidula is significantly paler, with exten-
sive grizzling on the upper surface of the body, the chest-
nut hairs on the back are greatly interspersed with white;
the undersurface is pure white; the external surface of
the ear is white anteriorly and dark reddish-brown pos-
teriorly; the tail is buffy grey, with a well-marked black
tip. The taxon taylori is essentially deep reddish brown
above, with some grizzling, whereas a large patch behind
the ears and the edge of the interfemoral membrane are
prominently black. The taxon cicur is a rich chestnut
above, without grizzling; the margin of the antebrachial
membrane and the muzzle are black; the distal half of the
tail except for the black tip is orange brown. The taxon
terutaus is a bright cinnamon-rufous on the upper body
and patagia; there is some light grizzling, especially on
the nape of the neck and shoulders; the undersurface is
salmon-buff tending towards dull orange-rufous on the
patagia; the crown of the head is a mixture of whitish and
cinnamon-rufous; the distal half of the tail except for the
black tip is dull orange-rufous. The taxon penangensis is a
rich chestnut above, without any grizzling; the undersur-
face is a creamy cinnamon; the margins of the patagia are
whitish grey; the forehead is a mixture of some whitish
and orange brown; the tail is deep orange brown with a
short black tip. The taxon melanotus has a bright reddish
brown dorsal pelage and is without grizzling; it is pale on
the cheeks and the undersurface.
In contrast to the other taxa, which differ in a range of
characters from the new specimen from Lao PDR, barroni
from eastern Thailand is essentially similar. Its dorsal
fur is comparable in colour; it is chestnut brown grizzled
with white above. The muzzle, cheeks, and throat are
also white. The ears are dull white anteriorly and black
posteriorly. The fringe of each patagium is white. The
undersurface of the body is light rusty brown, with a cin-
namon-brown extending on to the sides of the neck and
membrane. Therefore, the Lao specimen is assigned to
barroni , which is also the most geographically proximate
taxon. Thorington and Hoffmann (2005) did not recognise
barroni as a valid subspecies. However, on the basis of the
pelage colour of three Thai specimens from localities 4,
5, and 6 in Figure 1 (specimen data listed in the Appen-
dix) and the new specimen from Lao PDR, barroni is here
considered to be a distinct geographical race of Petaurista
petaurista . The new specimen from Lao PDR is the first
record of P. petaurista ( sensu Corbet and Hill 1992 , Thor-
ington and Hoffmann 2005 ) from east of the Mekong River.
On the basis of colour, barroni appears most similar to
Petaurista petaurista candidula , which is recognised as a dis-
crete subspecies by Thorington and Hoffmann (2005) . Other
subspecies from the study area recognised by these authors
are cicur , melanotus , penangensis , taylori , and terutaus . It is
possible that future studies may show that penangensis and
cicur are synonyms of P. p. melanotus. The taxon terutaus ,
although found in the Sundaic subregion, exhibits charac-
ters, such as grizzling on the dorsal surface, more associated
with specimens from Indochina. The taxon taylori in penin-
sular Myanmar, on the border of the Sundaic/Indochinese
subregions, also shares this character trait.
Conservation status
Petaurista petaurista is considered of Least Concern in
the IUCN Red List of Threatened Species ( Walston etal.
2008 ).
Ecological notes
Petaurista petaurista is found in a variety of forests: wet
tropical lowlands, mountain temperate forests, hardwood
forests, evergreen broadleaf forests, coniferous forests,
plantations, and orchards ( Thorington etal. 2012 ). In south-
eastern China, P. petaurista occurs in subtropical forest to
mixed deciduous and coniferous forest, with an altitudinal
range of 200 m 4000m elevation ( Yu etal. 2006 ).
Discussion
Kloss (1916) described the taxon barroni from Hup Bon
Village, Sriracha District, Chonburi Province, southeast
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8D. Sanamxay etal.: Red giant flying squirrel from Lao PDR
Thailand, approximately 13 ° 7 ′ N, 101 ° 6 ′ E. Originally
referred to Petaurista annamensis (which is today con-
sidered to be a synonym of P. philippensis ), it was subse-
quently included in alborufus by Askins (1977) . Later, it
was considered to be a synonym of P. petaurista by Corbet
and Hill (1992) and Thorington and Hoffmann (2005) , but
it was omitted from the synonymy list by Thorington etal.
(2012) . Both Francis (2008) and Thorington et al. (2012)
in their range maps omitted P. petaurista from eastern
Thailand and thereby also omitted the type locality of P.
p. barroni.
In addition to the holotype of barroni , five specimens
of Petaurista petaurista in the Natural History Museum,
National Science Museum Thailand (NHM), which had
been collected from Chaiyaphum (Figure 1, locality 4) and
Nakhon Ratchasima Provinces (Figure 1, locality 5), north-
east Thailand, were examined. The pelage colour of these
specimens essentially corresponds to the description of P.
p. barroni .
The recent discovery of P. petaurista in southern
Lao PDR indicates that this species is more widespread
than previously reported ( Corbet and Hill 1992 , Thoring-
ton and Hoffmann 2005 , Francis 2008 , Thorington etal.
2012 ). As noted above, the market from where it was
obtained is near the forests of the three formerly PNBCAs,
which comprise Phou Theung, Bolaven northeast, and
Phou Khanthong. This area is mostly covered with mixed
deciduous forest and dry dipterocarp forest. The locality
of the market, the hunting behaviour of the local people,
together with information obtained from the vendor, the
price of the specimen (equal to about $ US 40), and the fact
that it was already dead but still fresh, suggest that most
probably it was captured in this area. Other bush meat
in the market included local species, such as Malayan
porcupines ( Hystrix brachyura Linnaeus), Red cheeked
squirrels ( Dremomys rufigenis Blanford), and Indochi-
nese ground squirrels ( Menetes berdmorei Blyth). Further
surveys in the National Biodiversity Conservation Areas
(NBCAs) near to this new locality should be conducted.
These include Xe Sap NBCA, Dong Amphan NBCA, and
Xe Piane NBCA. In order to clarify the actual distribu-
tion range of this species, it would be also of interest to
conduct research elsewhere in Lao PDR and in areas of
similar habitat in Cambodia and Vietnam.
Finding this species as bush meat in a local
market supports the view of Jenkins et al. (2005) , who
noted that ongoing market surveys in the country will
provide an informative overview of the exploitation of
animals occurring in the surrounding area and occasion-
ally provide interesting species records. It also comple-
ments the very recent discovery of the new taxon of flying
squirrel Biswamoyopterus laonensis in an informal market
in central Lao PDR ( Sanamxay etal. 2013 ).
The finding of this species as bush meat is also con-
sistent with the view of Duckworth etal. (1999) , who con-
sidered that one of the major threats to wildlife in Lao PDR
is harvesting for food and trade.
Acknowledgments: In Lao PDR, we thank D. Sanamxay s
family for their support and encouragement. Thanks are
also due to the staff of the Faculty of Environmental Sci-
ences, National University of Laos for their help, sup-
port, and encouragement. We are grateful to Nouansi
Sisompheng for her help in obtaining the specimen. In
Thailand, we thank the staff of the Princess Maha Chakri
Sirindhorn Natural History Museum, Prince of Songkla
University for their help. We are also grateful to the staff
of the Natural History Museum, National Science Museum
Thailand (NHM) for their assistance. In the UK, we thank
the staff of the Mammal and Library sections of the Natu-
ral History Museum, London, especially Paulina Jenkins
and Roberto Portela-Miguez. At the Harrison Institute,
Sevenoaks we are most grateful to David Harrison, Mal-
colm Pearch, Nikky Thomas, and Beatrix Lanzinger for
their advice, access to specimens, and provision of lit-
erature. Finally, we acknowledge the 2011 Thailand On-
Place Scholarship under the ASEA-UNINET Programme
for Cambodia and Lao PDR ; Department of Biology,
Faculty of Science and Graduate School, Prince of Song-
kla University for their financial support; the Systematics
Association, UK for their contribution towards setting up a
zoological research collection in the National University of
Laos and the Darwin Initiative, UK (Project No: 18002) for
their ongoing support of taxonomic training and research
in Southeast Asia.
Appendix 1
Specimens of Petaurista petaurista held in the Natural
History Museum, London (BM[NH]) and the Natural
History Museum (NHM), National Science Museum Thai-
land (NHM).
Numbers from 1 to 23 in parentheses refer to localities
included in Figure 1.
(1) BM(NH).10.10.19.4; Kindat, upper Myanmar,
approximately 23 ° 44 ′ N, 94 ° 26 E. (2) BM(NH), not reg-
istered; Mogaung, Kachin, Myanmar, approximately
25 ° 18 ′ N, 96 ° 56 E. (3) BM(NH).15.5.5.40; Kin, Sagaing,
Myanmar, approximately 22 ° 45 ′ N, 94 ° 41 ′ E. (4) THNHM-
M-2344; Khon San, Pa Phu Khieo, Chaiyaphum, Thailand,
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D. Sanamxay etal.: Red giant flying squirrel from Lao PDR9
approximately 16 ° 31 ′ N, 101 ° 45 ′ E. (5) THNHM-M-2340;
Sakae Rat, Nakhon Ratchasima, Thailand, approximately
14 ° 36 ′ N, 102 ° 2 E. (6) BM(NH).47.1472; Hup Bon, Srira-
cha, Chonburi, Thailand, approximately 13 ° 7 ′ N, 101 ° 6 ′ E.
(7) BM(NH).14.12.1.5; Bankasun, Tenasserim, Myanmar,
approximately 12 ° 4 ′ N, 99 ° 1 E. (8) BM(NH).55.1656 and
BM(NH).55.1657; Koh Lak, Prachuap Khilikhan Prov-
ince, Thailand, approximately 11 ° 47 ′ N, 99 ° 48 ′ E. (9)
BM(NH).49.431; Khok Klap, peninsular Thailand, approxi-
mately 8 ° 53 ′ N, 99 ° 17 E. (10) BM(NH).10.10.1.79; Nong
Thale Song Hong, Trang Province, approximately 7 °
51 ′ N, 99 ° 29 E. (11) BM(NH).55.1649; Koh Klang, Trang
Province, Thailand, approximately 7 ° 21 ′ N, 99 ° 34 ′ E. (12)
NMNH.123.934; Terutau Island, Straits of Malacca, Thai-
land, approximately 6 ° 37 ′ N, 99 ° 40 ′ E. (13) BM(NH).49.433;
Penang Island, Malaysia, approximately 5 ° 27 ′ N, 100 °
13 E. (14) BM(NH).5.6.15.1; Larut hills, Perak, Malaysia,
approximately 5 ° 1 ′ N, 100 ° 53 ′ E. (15) BM(NH).1939.2373;
Perak, Malaysia, approximately 4 ° 48 ′ N, 100 ° 47 ′ E. (16)
BM(NH), not registered; Selangor, Malaysia, approxi-
mately 3 ° 38 ′ N, 101 ° 30 E. (17) BM(NH).28.7.20.62; Hutan
Rizab Semangkok, Selangor-Pahang Boundary, Malaysia,
approximately 3 ° 39 ′ N, 101 ° 45 ′ E. (18) BM(NH).61.1164;
Bukit Pelindong, Kuantan, Pahang, Malaysia, approxi-
mately 3 ° 50 ′ N, 103 ° 22 E. (19) BM(NH).22.8.12.2; Kuala
Selangor, Malaysia, approximately 3 ° 20 ′ N, 101 ° 14 ′ E. (20)
BM(NH).61.1163; Ulu Gombak Forest Reserve, Malaysia,
approximately 3 ° 18 ′ N, 101 ° 46 ′ E. (21) BM(NH).8.1.25.6,
BM(NH).8.1.25.7, BM(NH).55.1652, BM(NH).71.2714; Tioman
Island, Mersing Johor, Malaysia, approximately 2 ° 47 ′ N,
104 ° 10 E. (22) BM(NH).62.192 and BM(NH).85.8.1.118;
Gunung Belumut Recreational Forest, Johor, Malaysia,
approximately 2 ° 1 ′ N, 103 ° 32 ′ E. (23) BM(NH).118.8.14.2;
Singapore, approximately 1 ° 22 ′ N, 103 ° 48 ′ E.
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