February 1992

Asiatic Herpetological ResearchVol. 4, pp. 23-361

A Preliminary Report on the Reptile Fauna of the Kingdom of Bhutan with
the Description of a New Species of Scincid Lizard (Reptilia:
Scincidae)
Aaron m. Bauer and Rainer Gunther2
1

2^Department of Biology, Villanova University, Villanova, Pennsylvania, USA, 19085
Museumfiir Naturkunde der Humboldt Universitat zu Berlin, Invalidenstrasse 43, Berlin, Germany

Abstract. -The herpetofauna of the Kingdom of Bhutan has been poorly studied and few collections of
Bhutanese reptiles have been made. Reptiles collected by the 1972 expedition of the Naturhistorisches

Museum Basel (Switzerland) are presented as a basis for a preliminary species list for this eastern
Himalayan country. Specimens representing seven families and 18 species were examined. Included is a

new species of scincid lizard of the genus Mabuya. An additional five species have been reported from
Bhutan and numerous other taxa are known from adjacent regions of Sikkim and Assam. Most of the fauna
is pan-oriental in derivation and is
widespread to the east, west and south. A number of species, however,are primarily Indo-Chinese in their affinities and extend only as far west as eastern Nepal. Collections fromeastern Bhutan and from elevations over 1500
are particularly small and additional field work will bemrequired to provide a complete picture of the reptiles of the country.

Key words: Reptilia, Sauria, Scincidae, Mabuya, Bhutan, Himalayas, biogeography, zoogeography.

Introduction

eastern Himalayas (Fig. 1).

To the north it

bordered by Tibet (Xizang Zizhiqu), tothe east by Arunchal Pradesh, to the southby Assam, and to the west by Sikkim andthe Darjeeling district of West Bengal.
isAcharji and Kripilani (1951) reported
the herpetofauna of the western

that

Himalayas was poorly researched incomparison with that of the eastern portionof the range.
Since the time of their
publication this condition may be said tohave reversed, with several major andmany minor expeditions reporting on theherpetofauna of the Kingdom of Nepal,
particularly the region from Annapurna to
the west (Cox, 1985; Dubois, 1974a,
1974b; Leviton et al., 1962; Mrsic, 1980;
Nanhoe and Ouboter, 1987; Sura, 1987,
1989; Swan and Leviton, 1958). In theeastern Himalayas, numerous workers havereported on the reptiles of Sikkim and West
Bengal (e.g., Annandale 1912; Gunther,
1864; Inglis et al., 1920) reviewed theherpetofauna of the Abor district (= central
Arunchal Pradesh), some 225 km east of
Bhutan. Nonetheless, no comprehensive
works on the region have been compiled.

The Kingdom of Bhutan has longremained a gap in the knowledge of eastern
Himalayan zoology, especially that of theherpetofauna (Annandale, 1912; Ouboter
1986; Swan and Leviton, 1962). Bhutan2
occupies approximately 47000 km in the© 1992 by Asiatic Herpetological Research

Government policy severely

limiting

foreign travel and research in the Kingdom
has, until quite recently, left Bhutan as azoological terra incognita in south central
Asia. Fortunately, a great deal of Bhutanlies within national parks, sanctuaries orreserves (Hawkins, 1986) so that there may

yet be opportunities to study its fauna in a
relatively undisturbed state.

To date only four papers have been
published on the herpetofauna of Bhutan.
Bustard (1979, 1980a, 1980b) published aconservation document and two short noteson the status of the gavial (Gavialis
gangeticus) in Bhutan. Biswas (1975)
reported on a small collection of reptiles
from Bhutan and described a new taxon,
Calotes bhutanensis. We here report on amuch more extensive collection of reptiles
and amphibians from Bhutan collected bythe
Zoologische Expedition desNaturhistorischen Museums Basel(NMBA) in 1972. This material issupplemented by additional informationfrom literature sources.

Vol. 4, p. 24

FIG.

1.

Asiatic Herpetological Research

February 1992

Map of the eastern Himalayas and adjacent regions showing the position of Bhutan.

Closedcircles indicate the capitol cities of each country.

Methods
Specimens were collected in theKingdom of Bhutan in April-May 1972 bythedesZoologische
Expedition
Naturhistorischen
Museums Basel(NMBA), primarily by Drs. O. Stemmler
and M. Wurmli.
Specimens were
examined while on loan at the Museum fiir
Other material
Naturkunde, Berlin.
referred to is from the collections of the
Academy of Natural Sciences ofPhiladelphia (ANSP), the U.S. National
Museum (USNM) and the Zoological
Survey of India (ZSI). Live weights forsome specimens were obtained from thefield notes of the collectors. The following
abbreviations are used throughout this= live weight, SC = subcaudal
paper: LW

SVL = snout-vent length, TL = tail= ventral scales. Measurements
length, V
scales,were taken only from representative
specimens of each taxon and samples
generally reflect typical individuals fromeach series.
All of the localities represented by the

NMBA expedition are in southwestern
Bhutan.

Literature records from othercollections also include several localities in
the central and eastern regions of the

NMBA

Localities of thehaveexpedition
already been characterizedcountry.

the literature(Baroni-Urbani et al.,1973).
repetition only thelatitudes, longitudes, and elevations of
those localities at which reptiles
werecollected are provided below, alongin

To avoidFebruary 1992

Asiatic Herpetological ResearchVol. 4, p. 25

28°N
92°E

100 kmFIG. 2. Map of the Kingdom of Bhutan showing the localities referred to in the text. See Methods fornames, elevations and coordinates of numbered localities.

with the same information for literature andother specimen localities. Latitudes and

87 km N of Phuntsholing (1700 m),
27°05'N, 89°35'E.
5.

longitudes are approximate as no adequate
gazetteer exists for Bhutan and several
localities not appearing on any available

6.

maps could only be bracketed between
known places by reference to collectors
field notes.
The following numbered

7.

Chimakothi (2200 m),

27° 10' N 89°34'E.

110 km N of Phuntsholing (2000 m),
27° 12' N 89°34'E.

localities are marked on Figure 2:
8.

125 km N of Phuntsholing (2100 m),

27° 15' N 89°35'E.
1. Samchi (400 m),
26° 54' N 89° 14' E.

2.

Phuntsholing (200-400 m),

26°51'N 89°26'E.
3.

Khala and Balu Jhura (200 m),

9.

Paro (2300 m),

27°26'N 89°25'E.
10. Thimphu (2300-2500 m),
27°29'N 89°3 7'E.

26°50'N 89°26'E.

11. Wangdi Phodrang (1400 m),
27°29'N 89°54'E.

14 km N of Phuntsholing,(850-950 m), 26° 53' N 89° 27' E.

12. Tamji (2450 m),
27° 40' N 89°54'E.

4.

Asiatic Herpetological ResearchVol. 4, p. 26

Biswas (1975) suggest

13. Batase (1500 m),
27° 00' N 90° 37' E.

that thesinglespecimen falls within the range of variation
displayed by C. versicolor. Moody (1980)
in his systematic revision of the Agamidae
was apparently unaware of this description
and did not recognize the species either as
distinct or as a junior synonym. Until suchtime as C. bhutanensis is reevaluated in thecontext of the genus Calotes as a whole we

14. Panjurmane (1525 m),
27° 10' N 90° 43' E.

15. Manas River,26°50'N 90°59'E.
16.

February 1992

recognize it a taxon
distinguishable from C. versicolor on thebasis of the minor scalation and color
features delineated by its describer.

Rongtong (2042 m),

tentatively

27° 16' N 91° 32' E.

17. Samdrup Jhongkhar (300 m),

26°52'N 91°28'E.
Calotes versicolor (Daudin, 1 802)Species Recorded from theKingdom of Bhutan(17 specimens examined): Phuntsholing

NMBA 22582-92, ZMB 48784-85; Balu
Jhura NMBA 22593; Wangdi Phodrang
NMBA 22594-6.

Reptilia

Crocodylia

LW adult males 27.0-36.4 g (x = 31.68= 4), LW females and juveniles 5.8g, n

Family Gavialidae

21.1 g (x = 9.96 g, n = 8).

Gavialis gangeticus (Gmelin, 1789)

Ross (1989) included Bhutan in his listof the recent distribution of the gharial.
Bustard (1979, 1980a, 1980b) reported that
this species had recently been extirpatedfrom its primary habitat in the Kingdom of
The last
Bhutan, the Manas River.
specimens were seen in the 1960's.
Bustard (1980a, 1980b) and Groombridge(1987) have suggested reintroducing thespecies into suitable habitat. At present asmall population survives in the Indian
portion of the Manas River (Whitaker,
1987).
SauriaIn addition to the specimens examined,
Biswas (1975) reported a specimen from
Samdrup Jhongkhar, Bhutan. Calotes

versicolor is widespread throughout theOriental region, including all of thesouthern slopes of the Himalayas. Smith

Cox (1985) recorded thespecies as common up to 1980 m in the(1935) and

Himalayas. It is widespread in most of
Nepal and in adjacent regions of Sikkim
and the Darjeeling District (Acharji, 1961;
Acharji and Kripilani, 1951; Leviton et al.,1956; Mrsic, 1980; Nanhoe and Ouboter,
1987; Rendahl, 1937; Sura, 1987, 1989;
Swan and Leviton, 1962). Barbour (1912)
recorded this species from the Tista Valley
near the Bhutanese frontier of Sikkim.

Family Agamidae
Japalura variegata Gray, 1853
Calotes bhutanensis Biswas, 1975

Biswas (1975) described this endemic
species on the basis of a single specimen(ZSI 22480) from Panjurmane (listed as

Although hemensuralcharactersto distinguishprovided
C. bhutanensis from the very similar C
versicolor, the validity of this taxon mustremain in doubt. The values provided by
Janjurmane by Biswas).

.

(2 specimens examined): 87 km N of
22597-8. Biswas
Phuntsholing(1980) obtained a specimen from Batase,
Bhutan.
Giinther (1865) recorded thistaxon from Sikkim, and Hora (1926) andSmith (1935) reported that this species wascommon throughout the eastern Himalayas
Annandaleat elevations of 330-2970 m.

NMBA(1906) initially reported J. yunnanensis

Asiatic Herpetological Research

February 1992

Vol. 4, p. 27

FIG. 3. Holotype of Mabuya quadratolobus Baur anf Giinther (NMBA 10275). Note general coloration
pattern.

from

Buksa

near
the
(=Buxa),but
later
Bengalese/Bhutan frontier,described the same specimen as J.
bengalensis, now regarded as a synonym

central Nepal.

Annandale (1912) regarded

the species as common at low altitude
throughout the eastern Himalayas.

of J. variegata.

Platyurus platyurus (Schneider, 1797)

Family Gekkonidae(5

Hemidactylus brookii Gray, 1845

specimens examined): Phuntsholing

NMBA 22635; Wangdi Phodrang NMBA
22636-7; no precise locality NMBA 226389.

(1 specimen examined): Samchi NMBA
22599.

Swan and Leviton (1962) did not record

LW 2.5-5.2 g (x = 3.63 g, n = 3).

been reported as locallyAnnandale (1912), Cox (1985), Leviton(1956), Mrsic (1980), and Swan andLeviton (1962) recorded this species from

common at lower altitudes (Mrsic, 1980;

central and eastern Nepal at elevations up to

Nanhoe and Ouboter, 1987). Mitchell and
Zug (1988) found it to be the most1500

this species among the fauna of Nepal, but
it

has

since

abundant gecko at their locality in the Teraiof Nepal as did Cox (1985) at his central

et al.

m and Smith (1935) and Taylor(1962) recordedisitfrom Sikkim.

Thealso known from southeasternspecies
Tibet (Kraig Adler, pers. comm.).

Nepalese sites.

Family Scincidae
Hemidactylus frenatus
Dumeril and Bibron, 1836
(39 specimens examined):

Mabuya quadratilobus
Bauer and Giinther, n. sp.
Figs. 3 and 4.

SamchiNMBA 22600-7; Phuntsholing NMBA
22608-34; ZMB 48786-89.

Holotype.— NMBA 22681, lowest
on the right bank of river valley
Samchi, Bhutan (26°54'N,
elevation
450 m. Collected by
89°14'E),
M. Wiirmli and C. Baroni-Urbani, 12 May
terrace

LW 1.0-6+.2 g (x = 3.16 g, n = 32),
LW males (x = 3.43 g, n = 13), LW
females (x = 2.97 g, n = 19).

west

of

1972.

Leviton et al. (1956) and Swan andLeviton (1962) recorded this species from
Dharan (ca. 330 m) in Eastern Nepal andCox (1985) found it common in Tharu, in

—

NMBA

(11 specimens):
Paratypes.
22682-87, ZMB 48769, 48775-78 all fromSamchi, Bhutan. Collected 8 May 1972 by

Asiatic Herpetological ResearchVol. 4, p. 28

February 1992

48777 = 130% SVL). Axilla-groin length14.2 mm. Hindlimb length 13.8 mm.

—

Frontonasal broader
Scalation (Fig. 4).
than long; prefrontals large, in broad
contact;

frontoparietals

large,

paired;distinct interparietal; parietals each borderedposteriorly by a single nuchal (unilaterally
fragmented in holotype); nostril entirely
within nasal; small supranasals present, innarrow contact dorsally; two loreals,
anterior larger than posterior; lower eyelidwith central transparent disk (diameter
approximately 50% of eye); four
supraoculars, decreasing in size in the orderB.

2>3>4>1;

six

supracilliaries;

seven

supralabials, fifth and sixth under eye; sixth

supralabial separated from orbit by series ofsmall suboculars; fifth supralabial 2.5 x

length of anteriormost supralabials; sevenor eight infralabials, roughly equal in size;
first and second infralabials contact
postmental; two enlarged pairs of chinshields, first pair in narrow contact.

Ears

moderately

large;

tympanum

sunken; anterior margin of ear with three
flattened, distinctly squarish lobules;
remainder of scales on anterior margin ofear slightly raised.
Fig. 4. A) Lateral and B) dorsal views of the head

(NMBA

10275). Notethe ear lobules, enlarged fifth supralabial andcarinate dorsal scales.

of Mabuya quadratolobus

Dorsal scales tricarinate, with lateralcarinations more well developed than
medial; medial ridge limited to posterioredge on many lateral and posterior
on toproximal region of tail; dorsal scales
scale

scales; carinated scales continue

O. Stemmler (NMBA 22682, 22688) and
11 May 1972 by O. Stemmler and M.
Wiirmli (NMBA 22683-87, ZMB 10281-

approximately equal in size, decreasingslightly on flanks; 35 scale rows around
mid-body.

84).

Limbs pentadactyl;—

Diagnosis. Mabuya quadratilobus isdistinguished from all other Asian membersof the genus by the following combinationof characters: lower eyelid with transparent
disk; three large, squared lobules at anteriorscales on dorsal

surface of limbs weakly tricarinate; palmar
scales spinose; 14 unkeeled lamellae underfourth toe.

Color (in preservative).

— Dorsumolive

margin of ear; seven supralabials with fifth
approximately 2.5 x length of first; dorsal

brown with series of dark brown marks on
distal edges of scales, forming irregularscales tricarinate.

spots on nape and broken transverse bands
second scale row on body. Light
dorsolateral stripe, one and one half scale

Description of holotype.
36.0
25.8

— A juvenile,mm SVL, LW 1.3 g. TL (incomplete)
mm (TL of intact paratype NMBA

at everyrows wide, from level of ear to tail. Dark
brown stripe beneath light stripe, extendingAsiatic Herpetological Research

February 1992

from anterior corner of eye on to tail,passing beneath eye as a narrow dark line,
disrupted and diffuse through ear, fleckedwith blue-white scales ventrally and fading
towards flanks. Venter white. Dorsal
surfaces of limbs brown, scales edged withdark brown, forming a diffuse, irregular
reticulate pattern.

Variation.

—The paratypes resembleinfeatures.

all

the

All

holotype
major
specimens appear to be juveniles andpossess yolk scars. Some specimens show
minor fragmentation of some head scales.

LW 0.5-1.3 g (x = 0.85 g, n = 11); SVL
28.9-36.0 mm (x = 32.1 mm, n = 8).

—

Etymology. The name derives from theLatin lobus (lobe) and quadratus (square)
and is in reference to the striking ear
lobules characteristic of this species.

Unfortunately the phylogeny of thelygosomine skinks in general, and Mabuya
in particular, is not well resolved. Greer

Vol. 4, p. 29

Scincella sikkimensis (Blyth, 1853)(45 specimens examined): Phuntsholing

NMBA 22640; 87 km N of Phuntsholing
NMBA 22641; Chimakothi NMBA 2264251, ZMB 48779-83; 110 km N of
Phuntsholing NMBA 22652-60; 125 km N
of Phuntsholing NMBA 22661; Paro
NMBA 22662-67; Thimphu NMBA
22668-76; Tamji USNM 166443; nospecific locality NMBA 22667-80.

LW 0.6-3.5g (x = 1.62g, n = 35), SVL(larger specimens only) 45.8-55.8 mm (x =49.48 mm, n = 6), TL 144-148% SVL (n =3).

Ouboter (1986) reported a maximum
SVL of 55.7 mm and an average of 39.1

mm. Although only larger individuals of
the Bhutanese sample were measured, it
appears that populations from theto other skink lineages and suggested that

northeastern extent of the range may beslightly larger than average.
Specimens
examined in detail revealed at least threedifferent head scale patterns, reflecting
fusions and fragmentation of the standard

the south-east Asian species of the genus

pattern reported by Ouboter (1986).

(1974, 1979) regarded Mabuya as ancestral

exhibitedthegreatest

numberofplesiomorphic features. If this is true, thenthis group must also be paraphyletic. The
only comprehensive treatment of Mabuya
as a whole has been that of Horton (1973),
who provided a key to the species and an
overview of the evolution andbiogeography of the numerous species
groups by geographic region. More than

20 Asian Mabuya were recognized by
Horton (1973), but no unified picture of
relationships among these taxa waspresented. In most existing keys to theherpetofauna of the greater Indian region(e.g., Smith, 1935) the new species falls
out most closely with Mabuya dissimilis, a
species from the western Himalayan
region. Comparison with the types (ANSP
9537-8) and other specimens of the lattertaxon, however, reveal major differences
and the two taxa are clearly distinct, if
closely related at all. With the material athand we are unable to offer any meaningful
suggestion as to the close relationship of

Mabuya quadratilobus.

For

example, NMBA 22668 possessed afragmented frontonasal, NMBA 22663 had
both prefrontals fused to the frontal, and

NMBA 22662 had unilateral prefrontal-

frontal fusion.

Hora (1927) recorded the species from
Sikkim and other neighboring areas ofIndia, and Cox (1985), Mrsic (1980), Sura
(1987), and Swan and Leviton (1962)
reported specimens from central Nepal.
Gruber (1981) and Ouboter (1986)
highlighted the confusion surrounding theidentity of the Himalayan species of
Scincella in general, but demonstrated that
S. sikkimensis is the characteristic speciesof eastern Himalayas.
As defined by
Ouboter (1986) it appears to be limited
chiefly to mesic oak forest regions on thesouthern flanks of the range. Nanhoe andOuboter (1987) characterized this species as
a common inhabitant of forest clearings ormthroughoutedges, generally below 3000
the Himalayan region to the east of Jaljala
Pass in central Nepal. Ouboter (1986)

Asiatic Herpetological ResearchVol. 4, p. 30

February 1992

stated that the species was not known fromforests.

1200 m. However, Mrsic(1980) recorded specimens below 1000 m
in Nepal and the Bhutanese localities

here support this. Smith (1935) mentioned
specimens from Sikkim.

reported here, especially Phuntsholing,

Family Varanidae

clearly indicate that the species may occupy
the entire elevational range of the tropical-

Varanus bengalensis (Daudin, 1802)

localities below

subtropical belt of the foothills.

The Bhutanese localities reportedThe

presence of such low altitude specimens
give some credence to occurrence of
Scincella sikkimensis at Parasnath Hill,
south of the Gangetic Plain in Bihar, thetype locality of Mocoa sacra, a juniorsynonym of S. sikkimensis.

(3 specimens examined):

Phuntsholing

NMBA 22697-8, 22740. Specimens werealso sighted at Samchi.

This speciesiscommon throughout

much of the eastern Himalayas as well as
Assam (Annandale, 1912).

Cox (1985)

reported a specimen from central Nepal at

Sphenomorphus indicus (Gray, 1853)

Patan.
(5 specimens examined): 87 km N of
22689-92, ZMB
Phuntsholing
48768.

NMBA

LW 9.4-26.5g

Serpentes

Family Typhlopidae= 17.58, n =

(5c

5),

Maximum SVL 104.7 mm (NMBA 22689,
a largeRamphotyphlops braminus

female containing embryos), TL(Daudin, 1803)

128-130% SVL (n = 2)
(24 specimens examined):

Hora (1927) reported this species from"the eastern Himalayas below Darjeeling"
and Smith (1935) and Taylor (1962)
mentioned material from Sikkim. Although
literature records in the region are few, theSamchiNMBA 22699-713; Phuntsholing NMBA
22714-17, ZMB 48770-74, 48774.

LW 0.3-1.0 g (x = 0.72 g, n = 21).

Largest specimen ZNB 48771, SVL 146.0

widespread in the eastern
Himalayas (Annandale, 1912; Rendahl,

mm, TL 2.0 mm.

1937).

head scale suture pattern typical of thisspeciesisAll specimens have 20

scale rows around mid-body and show thespecies.

Sphenomorphus maculatus(Blyth, 1853)(4

specimens examined):

SamchiNMBA 22693; Phuntsholing NMBA
22694.

In addition the Basel Expeditioncollected two specimens,

This widespread and easily transported
snake has also been recorded from Nepal(Kramer, 1977; Mrsic, 1980; Nanhoe andOuboter, 1987) and Sikkim (Rendahl,1937).

NMBA 22695-6,

from 23 km N of Siliguri (ca. 150 m) in the

Family Pythonidae

Jaipalguri District of India.

LW 3.0-4.0 g (x = 3.5 g, n = 2), SVL
45.0-58.8 mm (x = 50.6 mm, n = 4), TL
144-200% SVL, n = 4).

Nanhoe and Ouboter (1987) concluded
that this Indo-Chinese species was limited
in its distribution to

lowaltitude riverine

Python molurus (Linnaeus, 1758)

Although we know of no Bhutanese
specimens in collections, the statement ofHarris et al. (1964) that "pythons" werecommon inhabitants of the Duars Plains of
Bhutan, must surely refer to this species. It
has been recorded from comparable habitats

Asiatic Herpetological Research

February 1992

Vol. 4, p. 31

Trachischium guentheri Boulenger, 1890

in Nepal (Kramer, 1977).

Family Colubridae

87(4 specimens examined):

km N

Phuntsholing NMBA 22732-35. Largest

Amphiesma platyceps (Blyth, 1854)(1
Wangdi
specimen examined):
Phodrang NMBA 22741 (475 + 184 mm,
186 V, 98 SC).

specimen (NMBA 22735), 226.3 + 38.6mm.
Kramer (1977) and Swan and Leviton(1962) reported this species from Nepal,

and Smith (1943) from
This species has previously beenrecorded from Sikkim, Assam and Nepal(Kramer, 1977; Mrsic, 1980; Nanhoe andOuboter, 1987; Smith, 1943; Swan andLeviton, 1962) and is widely distributed in

SikkimandDarjeeling.

Xenochrophis piscator(Schneider, 1799)(12 specimens examined): Phuntsholingthe Himalayas.

NMBA 22718-29. The specimens are all

Amphiesma stolata (Linnaeus 1758)

neonates (typical individuals 140.5 + 64.8mm) and were collected along with 16 egg

Biswas (1975) collected a single
specimen from Samdrup Jhongkhar inwidelyshells. Smith (1943) reported clutch sizes
Nanhoe andof 8-87 in this species.
Ouboter (1987) reported this species as

distributed pan-oriental species that appears

common at low altitudes in association with

(Nanhoe andwater and Annandale (1912) reported its
presence at elevations as high as 1450 m in
the western Himalayas. The species range
extends throughout the Oriental region.
Kramer (1977) and Swan and Leviton

eastern

Bhutan.

This

to thrive in disturbed areas

isa

Ouboter, 1987).

Boiga ochracea ochracea(Gunther, 1868)(1962) reported Nepalese localities.
(2 specimens examined):

Phuntsholing

NMBA 22730, juvenile (280.5 + 73.8 mm,
4 g, 234 V, 108 SC); NMBA 22731, adultmale, badly damaged (925 + 284 mm, 241
V, 118 SC).

This species is common in the eastern

Himalayas and is known from the Buksa
Duars along the southern boundary of
Bhutan (Smith, 1937).

Pseudoxenodon macrops (Blyth, 1854)(2 specimens examined):

Zaocys nigromarginatus (Blyth, 1854)
87 km N
specimen examined):
Number
NMBA-Field
Phuntsholing
10258, total length 1980 mm, badly(1

damaged.
This large snake is an eastern form,
ranging from Yunnan to the eastern
Himilayas (Smith, 1943). It is known
from elevations of up to 2500 m.

125 km N of

Family Elapidae

Phuntsholing NMBA 22738, male, headand neck severely damaged (975 + 227
mm, 164 V, 63 SC); Wangdi Phodrang
NMBA 22739 (710 + 183 mm, 169 V, 67
SC).

Bungarus niger Wall, 1908(1

specimen examined):

Phuntsholing

NMBA 22736 (667 + 120 mm, 221 V, 56
SC).

Nanhoe and Ouboter (1987) regarded
taxon has long been

species as an Indo-Chinese form,
extending through the eastern Himalayas as
far west as Jaljala Pass in west central

known from the eastern Himalayas andAssam (Smith, 1943), it has not been

Nepal.

recorded from eastern Nepal by any of the

this

AlthoughthisAsiatic Herpetological ResearchVol. 4, p. 32

recent reviewers of the fauna of that region.

Ophiophagus hannah (Cantor, 1 836)

Biswas (1975) reported a juvenile
specimen from Rongtong in the Manas
Valley. This species has recently beenrecorded from as far west as eastern Nepal(Nanhoe and Ouboter, 1987).

Species Likely to Occur in Bhutan
In addition to the 23 species listed
above, it is certain that a great many more
reptile species are yet to be found in
Bhutan, both in the more well-known, butricher lowlands, and the less-well collected

higher elevations.

No turtles have beenreported from Bhutan, but several species
are likely to occur. Among the batagurine
emydids the range of Kachuga tectabrackets Bhutan, with records from Sikkim

west (Moll, 1987) and the Dihang
River in the east (Annandale, 1912).
Kachuga drongoka has been collected in the
Brahmaputra River in the Kamrup Districtof Assam (Moll, 1986) and was illustrated
in a distribution map as occurring in Bhutan
by Tikader and Sharma (1985). Cuora
amboinensis Melanochelys tricarinata, M.
trijuga, Kachuga smithii, K. tentoria andK. sylhetensis have been reported from theManas Tiger Reserve in Assam (Das, 1988)
and Hardella thutjii, Indotestudo elongata
and Lissemys punctata also approach theborders of Bhutan. The freshwater turtles
are known either from the Tista to the west
in the,February 1992

almost certainly occurs at
lower elevations in Bhutan. Hemidactylus
bowringii was recorded by Barbour (1912)
from the Tista Valley near the Bhutanese
frontier with Sikkim, but no more recent
remarks on this species in the area inquestion have appeared in the literature.
latter species

Among agamids, Japalura tricarinata occurs
ineastern

Nepal,

Sikkim,andtheDarjeeling area, it may be expected to occur
in the southwestern corner of Bhutan,
although the Basel expedition found no
specimens despite their intense collecting
effort in and around Phuntsholing.
Japalura andersoniana was described fromthe Dafla Hills, near the eastern border ofand is another potentiallyindigenous agamid. Annandale (1906)
suggested that Ptyctolaemus gularis might
be found in the Buksa Duars along thesouthern frontier of Bhutan. There are agreat many snake taxa that might be
expected to occur in Bhutan. Over 40
species have been recorded from Nepal

Bhutan(Kramer, 1977). In the family Typhlopidae
jerdoni and R.
extend toandR.
diardioligolepis
possibly
the border regions of Bhutan. The colubrid
fauna of adjacent Sikkim and Assam is
at least Ramphotyphlops

exceptionally rich (Smith, 1943). Welch(1988) listed Ahaetulla p. prasina, Boiga

gokool, Boiga multifasciata andRhabdophis himalayana as well as Boiga o.
ochracea as taxa occurring in Bhutan.
However, these records were purportedly
derived from Smith (1943), who, in fact,
listed no material from Bhutan. Welch's(1988) apparent criterion for inclusion in
was Smith's (1943)
mention of the "eastern Himalayas" in hisor the Brahmaputra to the south (Das,
1985; Iverson, 1986; Smith, 1933; Tikader
and Sharma, 1985) and might be expected
to occur at lower elevations in the Torsa,

distributional

Wong Chu, Sankosh, and Manas drainagesof southwestern and south central Bhutan.

least four species of Oligodon occur in
Sikkim or the Jaipalguri District of India

that additional lizard

and may also be found in Bhutan.
Additional specimens and species almost

It is highly probablespecies may also be found in Bhutan.

The

gekkonid Hemidactylus flaviviridis occurs

the Bhutanese faunacomments.

In addition, at

certainly exist in museum collections, buthave never been reported in the literature.

Nepal (Cox, 1985; Sura, 1987, 1989)
but apparently does not reach the eastern
Himalayas. Hemidactylus garnotii, on theother hand, extends only as far west as
central Nepal (Cox, 1985; Nanhoe andalso likely that the cobra, Naja najaSmithkaouthia, occurs in Bhutan.

Ouboter, 1987; Sura, 1989), where it has
only recently become established. The

subspecies is illustrated as just reaching thesouthwest corner of the country. The true

in

It is

(1943:434) presented a distribution map
showing most of Bhutan within the range
The nominateof the subspecies.

Asiatic Herpetological Research

February 1992

status of these forms relative to one anotherremains unclear. They occur in sympatry
in a number of areas and are probably
specifically distinct (Wiister and Thorpe,1989). To date, however, there have beenno specimens to confirm the presence ofeither form in Bhutan, although there arerecords from Nepal (Acharji, 1961;
Kramer, 1977).

Discussion

Himalayan zoogeography

has

been

reviewed extensively by previous authors(e.g., Annandale, 1912; Blanford, 1901;
Dubois, 1981; Hora, 1948; Nanhoe andOuboter, 1987; Smith, 1933; Swan andAll have used
Leviton,1962).

approximately thesame divisions

inidentifying the affinities of the faunalelements of the region.

Bhutanlies

primarily within the Eastern Himalayan
province of the Indo-Chinese subregion of
the classically-defined Oriental region.

The

herpetology of the eastern Himalayas ingeneral is known from a number of faunalreports from eastern Nepal (e.g., Leviton etal.,1956) and northeastern Bengal(Annandale, 1912), as well as fromdescriptions of new species, especially
amphibians (see Dubois, 1974a). Swanand Leviton (1962) recorded 85 snake andlizard species from Sikkim and the adjacent
Darjeeling area of India. As yet material isinsufficient to compare the similarity andrichness of the Bhutanese fauna with thesefigures.

Bhutan is characterized by north-south flowing rivers that ultimately drain
into the Brahmaputra and thence to the Bayof Bengal.

To the north the higher peaks of

Himalayas form a barrier to reptile
movement. Mountain ridges also form
partial barriers to the west and east. Onlytheto the south is there low altitude access to

Bhutan. The mountains further subdivide
the country into a series of valleys between

which communication is likely only in thesubtropical south. As a consequence, themajority of the species recorded are panoriental in their distribution or, likeVol. 4, p. 33

Eastern Himalayan endemics
Nepal.
include Trachischium guentheri and
Scincella sikkimensis. The distribution of
the two putatively endemic forms describedfrom Bhutan, Calotes bhutanensis and
Mabuya quadratilobus n. sp., remains
poorly known, but it is probable that atleast the latter may be found in adjacent
Sikkim or Assam.
None of the taxa
Tibetan or
here
reflect
reported
Mediterranean influences as reported for the

Nepalese reptile fauna by Swan andLeviton (1962) and Nanhoe and Ouboter(1987). At least in part, this is a reflectionof the limitation of previous collecting
efforts to the valleys and lower elevationsof Bhutan. This same factor precludes ameaningful elevational analysis ofherpetofaunal distribution at this time.

In

Bhutan, as elsewhere in the Himalayas,
there appears to be a strong correlation
between zonation of vegetation and that ofamphibians and reptiles (Dubois, 1974a,
1974b, 1981; Nanhoe and Ouboter, 1987).

The elevational

profilesprovided byBaroni-Urbani et al. (1973) characterized
the native vegetation of the lower elevationsof Bhutan as: moist sal forest (200-800m), evergreen montaine forest (700-1600m), evergreen deciduous forest (1600-2800
conifer forestm), and Rhododendron(2800-3500 m). Dubois (1974b) regarded
elevations below 1000 m in neighboring
Nepal as tropical, those to 2000 m as
subtropical and those to 3000 m as
temperate, with higher elevations subalpine
or alpine in their climate and floral
characteristics.
Although each of thevegetation types is represented by at least
one collecting site, all are within the tropicalor subtropical zones, biasing the fauna
sampled against Tibetan or Mediterranean
taxa which would be expected to occur at
higher, more temperate elevations (Swan et
On the basis of the knownal., 1962).

fauna of other areasof the Eastern

Himalayas only a fraction of the expected
species have as yet been recorded. Further,phylogenies have not been proposed for the

Platyurus platyurus, Sphenomorphus
maculatus, and Pseudoxenodon macrops,
are primarily Indo-Chinese species thatmajority of the known taxa, precluding theapplication of cladistic biogeographic
methods (Humphries and Parenti, 1986) in
the analysis of pattern. It would thus beextend only as far west as eastern or central

premature to speculate about long-term

Asiatic Herpetological ResearchVol. 4, p. 34

historical biogeographical patterns of the
Bhutanese reptile fauna and a more detailed

analysis is defered to a later date.

February 1992

Comparative Zoology (Harvard University)
44:1-203 + 8 pis.

BARONI-URBANI, C, O. STEMMLER, W.
WITTMER, AND M. WURMLI. 1973.

Acknowledgments

We are grateful to the participants of theMuseum BaselNaturhistorisches

Zoologische Expedition des Naturhistorischen
Museums Basel in das KOnigreich Bhutan.

derVerhandlungen

Naturforschenden

Gesellschaft in Basel 83:319-336.

expedition and to the authorities of themuseum, in particular Dr. Eugen Kramer,
for permission to identify and publish areport of their collections. The staff of theUnited States National Museum provided
useful comments and access to literature.
In particular Dr. Bruce Beehler provided
assistance in the location of Bhutanesecollecting

and

sites

AddisonWynn

identification of theDr. Wolfgang
Prov.-Doz.
typhlopids.
Bohme (Museum Koenig, Bonn) also madeconfirmed

thesignificant contributions to the manuscript.

The holotype of Mabuya quadratilobus was
photographed at the University of Calgary
through the efforts of Dr. Tony Russell.

The senior author thanks the authorities ofthe Museum fur Naturkunde der Humboldt
Universitat zu Berlin for extending theinvitation that led to this collaborative
project.

BISWAS, S.

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