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MAJOR ARTICLE
31 TAPROBANICA VOL. 09: NO. 01
urn:lsid:zoobank.org:pub:103CB32E-CE9C-417B-B23A-09E7840A7400
ON TWO POORLY KNOWN, INSULAR SKINKS, Eutropis tytleri
(THEOBALD, 1868) AND E. andamanensis (SMITH, 1935) (REPTILIA:
SCINCIDAE), ENDEMIC TO THE ANDAMAN ISLANDS, INDIA
Section Editor: Jakob Hallermann Submitted: 28 March 2020, Accepted: 30 April 2020
S. R. Chandramouli1 & A. A. Thasun Amarasinghe2,3
1 Department of Ecology and Environmental Sciences, School of Life Sciences, Pondicherry University,
Puducherry, India
2 Research Center for Climate Change, University of Indonesia, Gd. PAU Lt. 8.5, Kampus UI, Depok 16424,
Indonesia
3 Association of Asian Herpetology (Asosiasi Herpetologi Asia), Jl. BSD Bintaro No. 88, Pondok Aren 15228,
Tangerang Selatan, Indonesia
1 Corresponding author. E-mail: findthesnakeman@gmail.com
Abstract
We examined the holotype of Eutropis tytleri and one of the syntypes of E. andamanensis collected
from the Andaman Islands, currently deposited at the Zoological Survey of India (ZSI). Information
on taxonomy, morphology, distribution, behaviour and natural history of these two poorly-known,
insular endemic skinks are presented here based on individuals recorded and examined in-situ, in the
field as well as preserved material. Their types are illustrated and redescribed. A distribution map is
presented based on records from the present study; threats and conservation status are discussed.
Key words: Andaman grass-skink, distribution, Mabuya, natural history, Sincus, taxonomy.
Introduction
The genus Eutropis Fitzinger, 1843 in the
Andaman archipelago is represented by only two
species: E. tytleri (Theobald, 1868) and E.
andamanensis (Smith, 1935). These species are
endemic to the Andaman Islands (Das 1999,
Amarasinghe et al. 2020). Although described in
the 1860s and 1930s, Eutropis tytleri and E.
andamanensis remain among the most poorly
known species within this genus, possibly
because of their narrow distribution, and the
difficulty of access to the Andaman Islands. The
Andaman archipelago, located south of the
Burmese peninsula is believed to be a
submerged extension of the Arakan Mountains,
which date back to the middle Miocene (15
Mya) with a general faunal affinity with the
Indo-Chinese region (Hall 1997, Das 1999).
Theobald (1868) described Scincus tytleri
based on the specimen ZSI 2273 from
‗Andamans, India‘. This was the first skink to be
named from the Andaman Islands.
Subsequently, Boulenger (1887) transferred it to
Mabuia. Later, Smith (1935) in his monograph
TAPROBANICA, ISSN 1800–427X. April, 2020. Vol. 09, No. 01: pp. 31–38, pls. 12–14.
© Research Center for Climate Change, University of Indonesia, Depok, Indonesia
www.taprobanica.org
ON Eutropis tytleri AND E. andamanensis FROM ANDAMAN ISLANDS
32 TAPROBANICA VOL. 09: NO. 01
on lizards of the Indian subcontinent used the
combination Mabuya tytleri, with the corrected
generic spelling Mabuya. He also described a
new species, Mabuya andamanensis, from the
Andaman Islands based on two syntypes, one
deposited in London: BMNH 1946.8.3.62 and
the other in Kolkata: ZSI 15084. The
comprehensive generic revision of skinks by
Mausfeld et al. (2002) and Mausfeld and
Schmitz (2004) resulted in the transfer of
Mabuya tytleri and M. andamanensis to the
genus Eutropis, the currently accepted
combination.
Recent literature on Indian lizards such as
Daniel (2002), Das (2002), and Das & Das
(2018) could not provide any reliable novel data
on either species, but only provide anecdotal
information owing to a lack of field data based
on recent surveys. Hence, the present paper aims
to provide additional information on various
aspects of these poorly known, insular endemic
species.
Material and methods
Field surveys were carried out on several islands
of the Andaman Archipelago to record the target
species. Individuals were observed in the field,
gently restrained, measured and released at the
point of capture.
We compared specimens (including types),
original descriptions, and other specimens of
Eutropis tytleri and E. andamanensis, as well as
specimens of their congeners (Appendix 1).
Museum acronyms are those of Uetz et al.
(2019). We examined the external morphology
of specimens by using a Wild M3Z
stereomicroscope. Sex was not determined from
specimens unless the hemipenes in males were
everted. All locality records are based on our
personal observations and data recorded in
museums. Altitudes are given in meters above
sea level (a.s.l.). The following measurements
were taken to the nearest 0.1 mm on the left side
of the body with a Mitutoyo digital caliper:
snout–vent length (SVL, from tip of snout to
anterior margin of vent), axilla–groin distance
(AG, from the posterior margin of the forelimb
at its insertion point on the body to the anterior
margin of the hind limb at its insertion point on
the body), thigh length (FEL, from the anterior
margin of the hind limb at its insertion point on
the body to the knee while flexed 90 degree),
shank length (TBL, from the posterior surface of
the knee while flexed 90 degree to the base of
the heel), head length (HL, from posterior edge
of the retro-articular process of the mandible to
tip of snout); head width (HW, width of head at
the temporo-mandibular articulation / angle of
the jaws), orbit diameter (ED, the greatest
horizontal diameter of the orbit); tympanum–eye
length (TYE, from posterior border of orbit to
anterior border of tympanum), snout length (ES,
from anterior border of orbit to tip of snout),
eye–nostril length (EN, from anterior border of
orbit to the midpoint of posterior border of
nostril), and toe lengths (from tip of toe, to the
junction with the adjacent digit, excluding the
claw). Supralabial and infralabial scales were
counted from the gape of the lips to the rostral
and mental scales respectively. Ventrals
included all scales from the scale posterior to the
postmental to the last scale bordering the vent,
counted along the ventral midline. Paravertebral
scales are between postparietal / nuchal
(included) to the level of the posterior margin of
the thigh in a straight line immediately left of the
vertebral column. Subdigital lamellae on toe IV
from the first proximal enlarged lamellae wider
than the width of the largest palm scale to the
distal-most lamella at the base of the claw. Total
number of longitudinal scale rows was counted
around the midbody.
Geographic coordinates of the skink‘s
localities were recorded with a GPS (WGS 84
datum) and mapped with ARC MAP 10.
Taxonomy
Eutropis tytleri (Theobald, 1868)
(Figs. 1–3, 6, 7; Tables 1–3)
Scincus tytleri Theobald, 1868
Mabuia tytleri — Boulenger 1887
Mabuya tytleri — Smith 1935
Eutropis tytleri — Mausfeld and Schmitz 2003
Holotype. Adult male, ZSI 2273 [fide Das et al.
(1998)], SVL 153.7 mm, collected from
Andamans, India, by Lt. Col. Robert Christopher
Tytler [a handwritten note ―confirmed as a type
by Dr. Malcolm Smith‖ on the original label is
present].
Other specimens (n=4). ZSI 2371, 2296,
4624; CSPT/L-33a, collected from Andamans,
India: collector and date unknown.
Diagnosis. A large species of Eutropis from the
Andaman Islands, SVL 125.0–153.7 mm; tail
long, about 1.5–2.2 times as long as the body;
prefrontals and supranasals in contact with each
other; temporals smooth; nuchals in a single
pair; tympanic opening relatively large; as large
CHANDRAMOULI & AMARASINGHE 2020
33 TAPROBANICA VOL. 09: NO. 01
as an adjacent lateral scale, lacking visible
auricular lobules; 24–30 midbody scale rows;
40–43 paravertebrals; dorsals tricarinate;
anterior genials in contact; posterior genials
separated by a single scale; cheeks swollen in
adult males; 50–53 ventrals; 27–30 subdigital
lamellae on toe IV; dorsum uniform bronze
brown with a greenish yellow venter.
Redescription of holotype. Meristic characters
of the holotype (ZSI 2273). Male, SVL 153.7
mm. Head moderately large (HL 26.1% of
SVL), narrow (HW 62.8% of HL, HW 16.4% of
SVL), indistinct from neck; snout short (ES
32.7% of HL, ES 52.0% of HW), slightly
concave in lateral profile; rostral shield large,
hemispherical, visible from above, posterior
margin of midpoint concave; frontonasal not
contacting rostral; frontonasal wide, lateral
border touching first loreal; prefrontals in broad
contact, separating frontal and frontonasal,
length equals maximum frontonasal length,
laterally contacting both loreal scales, posterior
border contacting first supraciliary, first two
supraoculars and frontal; frontal large, elongate,
subtriangular, bluntly pointed posteriorly, length
equal to frontoparietals and interparietal
combined; frontoparietals two, in contact, larger
than interparietal; parietals large and completely
separated by interparietal, contacting
pretemporal scales anterolaterally; single pair of
nuchals, not overlapping behind interparietal.
Nostril large and placed in centre of nasal;
supranasal single; loreals two, anterior
contacting nasal, supranasal, frontonasal,
prefrontal, posterior loreal, and second & third
supralabials; posterior slightly longer than
anterior loreal in the longitudinal axis,
contacting prefrontal, first supracilliary, third
supralabial, and forth supralabials slightly;
presuboculars two; eye large (ED 21.2% of HL;
considerably smaller than TYE), pupil rounded;
interorbital distance broad; postoculars four,
small; Supraoculars five, all wide, second
longest in the longitudinal axis and widest in the
transverse axis, 1st supraocular in contact with
prefrontal, 2nd in contact with prefrontal, frontal,
and frontoparietal, 3rd in contact with
frontoparietal, 4th in contact with frontoparietal
and parietal; 5th in contact with parietal;
supraciliaries six; eyelid moveable, window
divided into several tall rectangular scales.
Supralabials seven, fifth largest, at the mid orbit
position, and contacting granular scales of lower
eyelid; pretemporals three; primary temporals
three, secondary temporals four; infralabials
seven; ear opening small (approximately one
quarter ED), deep, near round. Mental large;
postmental single, large; two pairs of chin
shield, first pair slightly meeting in midline, first
chinshield in contact with first and second
infralabial scales, the second pair in contact with
second and third infralabials.
With the exception of head shields and
nuchals, all dorsal scales are tricarinate, on some
scales the median keel is not prominent; all
scales imbricate; scales on dorsal surface of
thigh moderately tricarinate; body slender,
elongate; midbody scale rows 24; paravertebral
scales 40; ventrals 51; preanal scales enlarged,
four.
Forelimbs short, hind limbs relatively long
(FEL 14.2% of SVL, TBL 16.0% of SVL); thigh
shorter (FEL 89.0% of TBL); dorsal surfaces of
fore and hind limbs moderately tricarinate;
subdigital lamellae of toes I-V: 11, 18, 25, 28
and 15 respectively; relative length of fingers IV
> III > II > V > I; those of toes IV > III > V > II
> I. Tail broken, median scale row of subcaudals
of original tail horizontally elongate and entire.
Variation. See Tables 1 & 2.
Coloration. In the preserved holotype,
dorsal head, body and limbs appear uniform
olive green, limbs darker. Venter lighter. In life
(not collected), dorsal head, body and limbs are
uniform bronze brown, with scattered black
spots on the neck, lateral body, on the limbs, and
tail. Lateral body lighter than the dorsum in
colour, temporal area pinkish. Throat lemon
green and venter light yellow.
Distribution. During the present study, E.
tytleri was recorded from different localities
throughout the Andaman Islands (Table 3).
Natural History. E. tytleri is the largest
skink species found within the Indian
subcontinent. It is crepuscular in habits and is
often seen emerging in the evening hours.
Diurnal activity is relatively less and is often
restricted to basking in the morning and foraging
until mid-day.
Based on personal observations, it feeds on
insects, frogs and smaller reptiles; and tends to
be bulkier on certain small islands, possibly due
to the lack of competition from other similar
sized lizards.
Unlike other Eutropis species, it is arboreal
to some extent and can often be seen on tree
trunks till a height of about 3 m above the
ground. Unlike other skinks, it can be seen
moving around actively until about 20:00 h, well
ON Eutropis tytleri AND E. andamanensis FROM ANDAMAN ISLANDS
32 TAPROBANICA VOL. 09: NO. 01
after the dark. Individuals of E. tytleri were
observed to suffer mortality due to road traffic in
reserve forests on three occasions.
Eutropis andamanensis (Smith, 1935)
(Figs. 4–7; Tables 1–3)
Mabuya andamanensis Smith, 1935
Mabuya andamanensis — Greer et al. 2000
Eutropis andamanensis — Mausfeld et al. 2002
Syntypes (ex. 2). Adult female, ZSI 15084, SVL
102.1 mm, collected from Andamans, India, by
C.G. Rogers; BMNH 1946.8.3.62 (formerly ZSI
14658), SVL not measured, other data same as
ZSI 15084.
Diagnosis. A medium sized species of Eutropis
from the Andaman Islands, SVL 75.0–115.0
mm; tail relatively short, about 1.4 times as long
as the body; the two prefrontals and supranasals
separated from each other; temporals keeled;
nuchals in a single pair; tympanic opening
relatively large; larger than an adjacent lateral
scale, with three small, rounded auricular
lobules; 28–30 midbody scale rows; 36–40
paravertebrals; dorsals pentacarinate; anterior
genials in contact; posterior genials separated by
a single scale; 48–56 ventrals; 26–28 subdigital
lamellae on toe IV; dorsum uniform bronze
brown with a greenish yellow venter.
Redescription of a syntype. Meristic and
morphometric characters of the syntype (ZSI
15084) are presented, the other syntype (BMNH
1946.8.3.62) not examined. Female, SVL 102.1
mm. Head moderately large (HL 25.6% of
SVL), narrow (HW 59.4% of HL, HW 15.2% of
SVL), indistinct from neck; snout short (ES
31.8% of HL, ES 53.5% of HW), slightly
concave in lateral profile; rostral shield large,
hemispherical, visible from above, posterior
margin of midpoint concave; frontonasal
contacting rostral; frontonasal wide, lateral
border touching first loreal; prefrontals not in
contact, not separating frontal and frontonasal,
length less than frontonasal length, laterally
contacting both loreal scales, posterior border
contacting first supraciliary, first supraocular
and frontal; frontal large, elongate,
subtriangular, bluntly pointed posteriorly, length
greater than frontoparietals and interparietal
combined; frontoparietals two, in contact, larger
than interparietal; parietals large and posteriorly
not separated by interparietal, contacting
pretemporal scales anterolaterally; single pair of
nuchals, overlapping middorsally. Nostril large
and placed in centre of nasal; supranasal single;
loreals two, anterior contacting nasal,
supranasal, frontonasal, prefrontal, posterior
loreal, and second supralabial; posterior longer
than anterior loreal in the longitudinal axis,
contacting prefrontal, first supracilliary, third
and fourth supralabials; presuboculars two; eye
large (ED 25.7% of HL; slightly smaller than
TYE), pupil rounded; interorbital distance
broad; postoculars three, small; Supraoculars
four, all wide, second longest in the longitudinal
axis and widest in the transverse axis, 1st
supraocular in contact with prefrontal and
frontal, 2nd in contact with frontal, and
frontoparietal, 3rd in contact with frontoparietal,
4th in contact with frontoparietal and parietal;
supraciliaries six; eyelid moveable, window
divided into several tall rectangular scales.
Supralabials nine, sixth largest, at the mid
orbit position, and contacting granular scales of
lower eyelid; pretemporals three; primary
temporals two, secondary temporals three;
infralabials eight; ear opening small
(approximately one sixth ED), deep, near round.
Mental large; postmental single, large; two pairs
of chin shield, first pair broadly meeting in
midline, first chinshield in contact with second
and third infralabial scales, the second pair in
contact with third and fourth infralabials.
Nuchals carinate; all dorsal scales are
strongly quinquecarinate; all scales imbricate;
body slender, elongate; midbody scale rows 30;
paravertebral scales 37; ventrals 56; preanal
scales enlarged, four.
Forelimbs short, hind limbs relatively long
(FEL 17.0% of SVL, TBL 17.7% of SVL); thigh
slightly shorter (FEL 96.1% of TBL); dorsal
surfaces of fore and hind limbs strongly
quinquecarinate; subdigital lamellae of toes I-V:
9, 17, 23, 29 and 17 respectively; relative length
of fingers IV > III > II > V > I; those of toes IV
> III > V > II > I. Tail complete, median scale
row of subcaudals of original tail entire.
Variation. See Tables 1 & 2.
Coloration. In the preserved syntype, dorsal
head, body and limbs uniform dark greyish
brown, limbs darker. A narrow chestnut brown
broken pair of lateral lines starting from back of
the eyes until midbody, disappear afterwards.
Venter light yellow.
In life (not collected), dorsum rich bronze
brown with two dark brown lateral bands along
the body from post orbital region till the tail.
Intensity of the lateral bands varies with age,
34
CHANDRAMOULI & AMARASINGHE 2020
33 TAPROBANICA VOL. 09: NO. 01
with the bands being distinct and dark in young
individuals while broken with intermittent white
spots in aged individuals. Upper lip, lower
lateral body and venter cream coloured.
Distribution. During the present study, E.
andamanensis was recorded from different
localities throughout the Andaman Islands
(Table 3).
Natural History. A diurnal species which
has, at times, been seen active till 18:30 h at the
dusk. Predominantly terrestrial and often seen on
the ground in leaf-litter, but occasionally seen on
low-lying tree trunks to a height of about 1.5 m
above the ground. Based on personal
observations, this skink feeds on insects, worms,
small frogs and crustaceans.
Discussion
The information presented here on taxonomy,
morphology, behaviour and natural history of E.
tytleri and E. andamanensis adds significantly to
our existing knowledge on these species, which
were described in the 1860s and 1930s
respectively. Until now, only anecdotal notes
have been available on these species based on
older literature. Stoliczka (1870) mentions
certain unusually large specimens of ―Tiliqua
carinata‖ collected from the Andaman Islands
by Roepstorff with a long tail, which were
referred to E. tytleri by Smith (1935).
These species still remain poorly known due
to the lack of sufficient field records and data,
and hence, their conservation status assessments
remain yet to be assessed (IUCN 2020). Das
(2002) erroneously illustrated E. andamanensis
as E. tytleri in his species accounts. The identity
of the individuals and specimens described and
illustrated here were confirmed by comparing
their respective type specimens (Fig. 6).
Both these species seem to be equally
widespread within the Andaman archipelago
(Fig. 7 & Table 3).
Table 1. Morphometric (in mm) and meristric character comparisons of Scincus tytleri Theobald, 1868 holotype
and Mabuya andamanensis Smith, 1935 synype, and other specimens from Andamans; ―––‖ = not measured.
E. tytleri
E. andamanensis
Character
Holotype
(ZSI 2273)
Other
preserved (n=4) & live (n=7)
Syntype
(ZSI 15084)
Other
live (n=6)
Sex
male
both
female
both
SVL
153.7
125.8–153.0
102.1
75.0–115.0
Head length (HL)
40.1
21.3–25.6
26.1
12.5–20.1
Head width (HW)
25.2
18.7–22.3
15.5
11.9 – 20.8
Axilla–groin distance (AG)
81.2
57.5–80.0
42.9
30.0–50.0
Eye–nostril length (EN)
9.2
9.2–11.2
5.7
6.1–8.5
Snout length (ES)
13.1
10.6–14.3
8.3
8.2–12.4
Tympanum–eye (TYE)
12.0
10.1–13.4
7.1
7.2–9.9
Orbit diameter (ED)
8.5
5.4–7.2
6.7
3.5–6.8
Thigh length (FEL)
21.9
18.2–23.9
17.4
12.2–22. 3
Shank length (TBL)
24.6
19.8–22.4
18.1
12.5–23.5
Tail length (TAL)
––
220.0–290.0
––
120.0–175.0
Midbody scale rows
24
24–26
30
28–30
Nuchal pairs
1
1
1
1
Ventrals
51
48–53
56
48–55
Lamellae on 4th toe
28
24–30
29
26–28
Paravertebral scales
40
40–45
37
36–40
Table 2. Diagnostic characters of Eutropis tytleri and E. andamanensis (live and preserved) in Andaman Islands
E. tytleri (n=12)
E. andamanensis (n=7)
Dorsal scales
tricarinate
quinquecarinate
Temporal scales
smooth
keeled
Lateral band
invisible
visible
Supranasals
in contact
separated
Prefrontals
in broad contact
separated
Prefrontals in contact with
1st & 2nd supraocular
1st supraocular only
Frontal in contact with
2nd supraocular only
1st & 2nd supraocular
Parietals
completely separated
In contact behind interparietal
Midbody scale rows
24–26
28–30
35
ON Eutropis tytleri AND E. andamanensis FROM ANDAMAN ISLANDS
32 TAPROBANICA VOL. 09: NO. 01
Figure 6. Head in dorsal and lateral views of (A, C)
the holotype of Eutropis tytleri (ZSI 2273) and (B, D)
syntype of E. andamanensis (ZSI 15084). Illustration
© A.A.T. Amarasinghe
E. andamanensis is more common and can
be seen abundantly whereas E. tytleri is
relatively rare and less abundant. E.
andamanensis is relatively more tolerant to
human mediated habitat disturbances than E.
tytleri, which is a more forest dependent species.
There seems to be some degree of ecological
niche partitioning between these two relatively
large bodied skinks.
Figure 7. Current distribution of Eutropis tytleri (in
circles) and E. andamanensis (in squares) in the
Andaman Islands based on known locality records.
Among them, E. tytleri has a larger body
size in comparison to E. andamanensis and is
more arboreal in habit than the latter. Likewise,
E. tytleri shows a crepuscular activity by being
more active during twilight hours than during
the day. On the other hand, E. andamanensis
seldom ventures out after sunset.
E. andmanensis seems to undergo a gradual
ontogenetic colour change as the intensity of the
dark lateral bands seem to fade with age: young
individuals have much darker and more legible
bands than old ones, which have intermixed
white spots on the dark bands along the lateral
sides of the body. Such a phenomenon has not
36
CHANDRAMOULI & AMARASINGHE 2020
33 TAPROBANICA VOL. 09: NO. 01
been recorded in E. tytleri. Road traffic in forest
reserves has been identified as a potential threat
to these skinks (Fig. 3). A precise distribution
map for these species presented here based on
current and confirmed past records will aid in
their conservation status assessments. Locality
details are listed in Table 3.
Table 3. Locality records of Eutropis tytleri and E. andamanensis in the Andaman Islands based on the present
study and literature.
Species
Locality, Island
Lat.
Long.
Alt. (m)
Source
both species
Mt. Harriet NP, South Andaman
11.7207
92.7336
362
present study
both species
Chidiyatapu, South Andaman
11.4917
92.7088
77
present study
both species
Wandoor, South Andaman
11.6154
92.6194
3
present study
both species
Bada Khadi, Rutland
11.4291
92.6649
2
present study
both species
Kyd, Kyd Island
11.9663
92.7546
6
present study
both species
Sitapur, Neil Island
11.8302
93.0342
11
present study
both species
Radhanagar, Havelock
12.0085
92.9638
43
present study
both species
Harminder Bay, Little Andaman
10.5334
92.5353
24
present study
both species
Krishnanallah, Little Andaman
10.6776
92.5087
153
present study
both species
Dinghi Ghat, Little Andaman
10.7375
92.5673
16
present study
both species
Long Island
12.3693
92.9193
26
present study
both species
Cutbert Bay, Middle Andaman
12.6359
92.9560
22
present study
both species
Mayabunder, Middle Andaman
12.8474
92.8565
20
present study
both species
Kishori Nagar, North Andaman
13.1485
92.9427
99
present study
both species
Lamia Bay, North Andaman
13.1983
93.0367
6
present study
both species
Interview
12.9425
92.6978
64
present study
E. andamanensis
Tarmugli
11.5734
92.5519
29
present study
E. andamanensis
Alexandra
11.5785
92.6129
46
present study
E. andamanensis
Redskin
11.5709
92.5929
36
present study
E. andamanensis
Chester
11.5832
92.5782
0
present study
E. andamanensis
Grub
11.5894
92.5936
0
present study
E. andamanensis
Paget
13.4301
92.8324
34
present study
E. andamanensis
North Reef
13.0899
92.7013
18
present study
E. tytleri
Narcondam
13.4491
94.2638
13
Raman et al. (2013)
E. tytleri
Inglis
12.1399
93.0940
7
Sivaperuman (2014)
E. tytleri
John Lawrance
12.0975
93.0411
5
Sivaperuman (2014)
Acknowledgments
We thank the Department of Environment and
Forests, Andaman and Nicobar Islands for
permission to SRC (permit no:
CWLW/WL/134/(J)/Folder/417) for conducting
this study and for the infrastructure provided;
K.V. Devi Prasad and the faculty of the
Department of Ecology and Environmental
Sciences and the Department of Ocean studies
and Marine Biology, Pondicherry University for
the support extended, the Mohamed bin Zayed
Species Conservation fund for a grant
(#14058387) which partly facilitated this study.
We also thank the former Director, K.
Venkataraman (Zoological Survey of India) for
granting research permission to AATA. In
particular we thank K. Chandra (Director) K. C.
Gopi (officer-in-charge, FPS Building, ZSI,
Kolkata), K. A. Subramanian (officer-in-charge,
Technical Section, ZSI) for their help throughout
the permitting application process. K. Deuti, S.
Raha, P.G.S. Sethy, P. Bag, and S. Debnath are
also acknowledged for assisting us while
examining collections in ZSI; Officers at the
Chennai Snake Park Trust facilitated the study
of preserved specimens under their care. We
thank Jakob Hallermann (Universität Hamburg,
Germany) and Philip Bowles (IUCN) for the
review of the manuscript and critical comments.
Finally, we thank J. Supriatna and the staff of
the Research Center for Climate Change,
University of Indonesia, for their support.
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