Taxonomy and distribution of Liopeltis calamaria (Günther, 1858) (Reptilia: Colubridae), including redescription of the syntypes

Abstract

Liopeltis calamaria, a rare non-venomous colubrid snake of South Asia, is redescribed. Its syntypes and all the available type specimens of its recognized synonyms are examined, including information about the respective populations found across India and Sri Lanka. Our literature compilation and mapping analyses reveal three distinct populations-(I) Sri Lankan (probably also present in some parts of South India as well), (II) Peninsular Indian, and (III) Himalayan / Nepalese, separated by the Palk Strait and the Indo-Gangetic plains respectively.

Full text

MAJOR ARTICLE
39 TAPROBANICA VOL. 09: NO. 01
urn:lsid:zoobank.org:pub:B2D867F0-009C-4D4D-91E1-A86AE89B059F
TAXONOMY AND DISTRIBUTION OF Liopeltis calamaria (GÜNTHER,
1858) (REPTILIA: COLUBRIDAE), INCLUDING REDESCRIPTION OF
THE SYNTYPES
Section Editor: Ivan Ineich Submitted: 01 December 2019, Accepted: 11 March 2020
A. A. Thasun Amarasinghe1,6, Suranjan Karunarathna2, Patrick D. Campbell3,
S. R. Ganesh4 & Gernot Vogel5
1 Research Center for Climate Change, University of Indonesia, Gd. PAU Lt. 8.5, Kampus UI, Depok 16424,
Indonesia
2 Nature Explorations & Education Team, No: B-1 / G-6, De Soysapura Flats, Moratuwa 10400, Sri Lanka
3Department of Life Sciences, Darwin Centre, Natural History Museum, Cromwell Road, South Kensington,
London SW7 5BD, England
4 Chennai Snake Park, Rajbhavan Post, Chennai 600 025, Tamil Nadu, India
5 Society for Southeast Asian Herpetology, Im Sand 3, D-69115 Heidelberg, Germany
6 Association of Asian Herpetology (Asosiasi Herpetologi Asia), Jl. BSD Bintaro No. 88, Pondok Aren 15228,
Tangerang Selatan, Indonesia
1 Corresponding author. E-mail: thasun@rccc.ui.ac.id
Abstract
Liopeltis calamaria, a rare non-venomous colubrid snake of South Asia, is redescribed. Its syntypes
and all the available type specimens of its recognized synonyms are examined, including information
about the respective populations found across India and Sri Lanka. Our literature compilation and
mapping analyses reveal three distinct populations – (I) Sri Lankan (probably also present in some
parts of South India as well), (II) Peninsular Indian, and (III) Himalayan / Nepalese, separated by the
Palk Strait and the Indo-Gangetic plains respectively.
Key words: Reed snake, Sri Lanka, India, population variation, synonyms, sexual dimorphism.
Introduction
The Calamaria Reed Snake, Liopeltis calamaria,
was described by Günther (1858) as Cyclophis
calamaria, based on two adult specimens
collected from ―Ceylon‖ (=Sri Lanka,
catalogued as types: BMNH 1946.1.21.64 and
1946.1.5.60). Even though Günther (1858) used
two specimens for the description he did not
provide specific data for any one of them. Later,
Jan (1862) described, then (1865) illustrated a
morphologically similar species, Homalosoma
baliolum from an unknown locality. Günther
(1864) later described another species,
Cyclophis nasalis, also from an unknown
locality, which he regarded as similar to his
previous species. In the description he stated
―This species agrees in almost every aspect with
Cyclophis calamaria, but it may be readily
TAPROBANICA, ISSN 1800–427X. April, 2020. Vol. 09, No. 01: pp. 39–49, pls. 15–18.
© Research Center for Climate Change, University of Indonesia, Depok, Indonesia
www.taprobanica.org

TAXONOMY AND DISTRIBUTION OF Liopeltis calamaria
40 TAPROBANICA VOL. 09: NO. 01
distinguished by the presence of two
preoculars…‖.
Boulenger (1890) nonetheless synonymized
Homalosoma baliolum and Cyclophis nasalis
with Liopeltis calamaria. Smith (1943) followed
this decision, and though he did not examine the
holotype (by monotypy) specimen of
Homalosoma baliolum, he regarded it as a
subjective synonym of Liopeltis calamaria and
this scheme has been followed by later workers.
Boulenger (1890, 1894) provided detailed
meristic data for Liopeltis calamaria (as Ablabes
calamaria), and reported its geographic range as
Madras Presidency and Bombay (=Mumbai),
India, aside from its type locality in Sri Lanka.
Wall (1921) described the variation of
ventral and subcaudal counts within Sri Lankan
populations, commenting that the snake found
on that island is rather distinct from Indian
populations. In contrast, Smith (1943) contended
that the meristic character variation observed
between Indian and Sri Lankan populations was
due to sexual dimorphism rather than
geographical variation.
Deraniyagala (1955) disagreed with Smith‘s
suggestion, and following Wall‘s (1921)
assumption, named a new subspecies Liopeltis
calamaria indicus (sic) for the Indian population
without designating a type specimen [only
referred to the Indian specimens examined in
Wall (1921)], and the name, Liopeltis calamaria
calamaria remained with the Sri Lankan
population. Subsequent authors have not
accepted Deraniyagala‘s subspecific separation
and have considered Liopeltis calamaria a rare
but widely distributed monotypic species in Sri
Lanka and India (De Silva 1969, 1980, de Silva
1990, Das 1994, Batuwita 2001, Somaweera
2006).
In much of the historical literature, Liopeltis
calamaria is considered uncommon throughout
its range (Willey 1906, Sarasin 1910, Cazaly
1914, Wall 1919, 1921). More recent work on
Indian and Sri Lankan snakes (Daniel 2002, Das
2002, Das & de Silva 2005, Whitaker & Captain
2008) have not shed any further light on this
species. Recently it has been observed in Sri
Lanka (Karunarathna et al. 2004, Karunarathna
& Perera 2010). To provide more clarity on this
taxon, we hereby illustrate and redescribe one of
the two syntypes of Cyclophis calamaria and all
other available types of all its subjective
synonyms.
We also map its distribution in Sri Lanka
and India based on published records, observed
specimens, and museum collections.
Material and methods
Specimens were examined in the collections of
the Natural History Museum, London UK
(BMNH); Ecological Resource Monitoring Lab,
Sri Sailam India (ERM); Madras Government
Museum, India (MAD); Naturhistorisches
Museum Wien, Vienna, Austria (NMW);
Naturalis Biodiversity Center, Nationaal
Natuurhistorische Museum (Rijksmuseum),
Leiden, The Netherlands (RMNH); Vietnam
National University, Hà Nội, Vietnam (VNUH);
Zoological Research Museum Alexander
Koenig, Bonn, Germany (ZFMK); Zoological
Museum of Moscow University, Moscow,
Russia (ZMMU); and Zoological Survey of
India, Kolkata, India (ZSI). Museum acronyms
follow Uetz et al. (2019). Morphometric and
meristic data for species comparisons were
obtained from examined specimens (see
Appendix I). Natural history data were taken
from our own field observation notes made
during the last ten years, as well as published
literature.
The following characters were measured
with a digital caliper (±0.1 mm): eye diameter
(ED, horizontal diameter of eye); eye–nostril
length (distance between anterior most point of
eye and middle of nostril); snout length (ES,
distance between anterior most point of eye and
snout); nostril diameter (horizontal diameter of
nostril); internarial distance (least distance
between nostrils); mandible–posterior eye
distance (distance between posterior edge of
mandible and posterior most edge of eye);
interorbital width (IO, least distance between
upper margins of orbits); head length (HL,
distance between posterior edge of mandible and
tip of snout); head width (HW, maximum width
of head); snout–vent length (SVL, measured
from tip of snout to anterior margin of vent); tail
length (TL, measured from anterior margin of
vent to tail tip). Meristic characters were taken
as follows: supralabials and infralabials (first
labial scale to last labial scale bordering gape);
costal scales (counted around the body from one
side of ventrals to the other in three positions, on
one head length behind neck, at mid body and at
one head length prior to anal plate); when
counting the number of ventral scales, we scored
specimens according to the method described by
Dowling (1951). We counted subcaudal scales
from first subcaudal scale to the scale before the
tip of the tail.

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41 TAPROBANICA VOL. 09: NO. 01
Taxonomy
Liopeltis calamaria (Günther, 1858)
(Figs. 1–4, 6; Tables 1, 2)
Cyclophis calmaria Günther, 1858
Homalosoma baliolum Jan, 1862
Cyclophis nasalis Günther, 1864
Cyclophia calamaria (sic)— Phipson 1888
Ablabes calamaria — Boulenger 1890
Liopeltis calamaria — Wall 1921, Smith 1943
Opheodrys calamaria — Constable 1949
L. calamaria indicus (sic)— Deraniyagala, 1955
Liopeltis calamaria indica Deraniyagala, 1955
Liopeltis calamarius (sic)— Wallach et al. 2014
Syntypes (n=2). BMNH 1946.1.5.60 (adult
male) and BMNH 1946.1.21.64 (subadult male);
collected from ―Ceylon‖ (=Sri Lanka) by an
unknown collector [according to the museum
registry].
Other specimens (n=30). See Appendix I
Diagnosis. Liopeltis calamaria can be separated
from its congeners by the following combination
of morphological characters: 130–136 ventrals
in males; 66–76 subcaudals in males; longer
snout, ED 60.6–65.5% of ES; no loreal; single
nasal shield; dorsal scales in 15-15-15 rows, all
smooth; single preocular, 2 postoculars and 1+2
temporals; 7 supralabials, of which 3rd and 4th in
contact with the eye; 7 infralabials; nasal
contacting internasal and prefrontal; prefrontal
separated from supralabials by the preocular and
the nasal; subcaudal scales paired, and dorsally
light brown, greenish-brown or olive green
coloration.
Redescription of syntypes. Meristic characters
of the adult male syntype (BMNH 1946.1.5.60)
are followed, where appropriate, by those of the
subadult male syntype (BMNH 1946.1.21.64) in
square brackets. Adult male [subadult male],
SVL 227.0 mm [192.0 mm]; tail length 150.0
mm [85.0 mm]; head elongate (HL 4.4% of
SVL), twice as long as wide (HW 60.4% of HL),
slightly flattened, distinct from neck; snout
elongate (ES 37.6% of HL), moderate, blunt in
dorsal view, rounded in lateral profile, forming
an oval shape, rather depressed.
Rostral shield large, hemispherical,
distinctly visible from above, rounded
posteriorly; interorbital width broad (IO 63.9%
of HW); internasals semicircular; nostrils rather
small; nasals undivided, elongate, replacing the
position of loreal, in anterior contact with
rostral, internasal and prefrontal dorsally, 1st and
2nd supralabial ventrally; prefrontal rather large,
broader than long, and subhexagonal; frontal
large, subhexagonal, elongate posteriorly and
longer than its width; supraoculars wide,
elongated, subrectangular, posteriorly wider;
parietals large, butterfly wing-like in shape,
bordered by supraoculars, frontal, upper
postocular anteriorly, anterior and upper
posterior temporals, and five dorso-nuchal scales
posteriorly; no loreal scale; one preocular (both
sides), vertically elongated, hexagonal, in
contact with prefrontal and nasal anteriorly,
supraocular dorsally, and 2nd & 3rd supralabial
ventrally; eye large (ED 20.8% of HL), round,
nearly the same size as snout length (ED 50% of
ES), pupil rounded; two postoculars, upper
postocular larger, pentagonal, in contact with
supraocular, parietal and anterior temporal
broad, in narrow contact with lower postocular;
lower postocular crescent in contact with 4th and
5th supralabial ventrally, anterior temporal
posteriorly; temporals 1+2, elongated,
hexagonal; anterior temporal larger and slightly
longer than posterior temporals, in contact with
parietal dorsally, 5th and 6th supralabial ventrally;
posterior temporals smaller, lower one in contact
with 6th and 7th supralabials ventrally.
Supralabials 7 (on both sides), 4th–7th larger
in size, 6th being the largest; 1st supralabial in
contact with rostral anteriorly, nasals dorsally,
2nd supralabial with nasal and preocular dorsally,
3rd supralabial with preocular and orbit dorsally,
4th supralabial with orbit and the lower
postocular dorsally, 5th supralabial with lower
postocular and anterior temporal dorsally, 6th
supralabial with anterior temporal and lower
posterior temporal dorsally, and 7th supralabial
with lower posterior temporal dorsally and body
scales posteriorly.
Mental of moderate size, triangular, wider
than its length; first infralabial pair larger than
mental plate and in broad contact with each
other, in contact with anterior chin shield
posteriorly; seven infralabials, 1st–4th in contact
with first chin shield, 4th infralabial largest in
size in narrow contact with the anterior chin
shield and in broader contact with the posterior
chin shield; 5th infralabial in narrow contact with
posterior chin shield, 6th–7th infralabials in
contact with gular scales; two larger anterior
chin shields in broad contact, and two smaller
posterior chin shields in narrow contact;
posterior chin shields bordered posteriorly by
five gular scales.
Body robust, elongate and subcylindrical;
costal scales in 15-15-14 rows [15-15-15], all

TAXONOMY AND DISTRIBUTION OF Liopeltis calamaria
40 TAPROBANICA VOL. 09: NO. 01
smooth and bluntly pointed; 130 ventral scales
[134]; anal plate divided. Tail comparatively
short (TL 66.1% of SVL), robust and thick; 76
[72] paired subcaudals.
Colouration. In preservation, dorsally
yellowish chestnut brown; the dorsolateral scales
(5–6 rows above the ventrals) edged with black
colour, create two longitudinal lines on each side
of the vertebral column; these lines break up
rapidly into spots anteriorly and continues well
onto the mid body and disappear towards the
posterior body and tail; ventral side yellow [pale
in colour; the black longitudinal line absent,
instead dark brown patches break up into spots
anteriorly; ventral side cream].
In life (based on live specimens, not
collected), dorsally light brown, greenish-brown
or olive green; the dorsal scales (5–6 rows above
the ventrals) edged with black colour creating
paired longitudinal lines on each side of the
vertebral column, which break into spots
anteriorly and continue well onto the tail; the
area between these two lines is sometimes
darker in colour; another less distinct line runs
along the confines of the 3rd–4th rows; a series of
black spots occur on each side of the head;
ventral side pale yellow or cream. The black line
is more prominent among juveniles than adults
(Fig. 1); an ill-defined but distinct dark patch
can be seen on the parietal region. Also, some
juveniles exhibit lateral white linear stripes that
begin on the neck and finish at the level of vent.
Redescription of Liopeltis calamaria indica
Deraniyagala, 1955
L. calamaria indicus (sic) Deraniyagala, 1955
Lectotype (designated herein). BMNH
1922.5.25.22 (male); collected from Waynad
(=Wayanad), Kerala, India, by Frank Wall
[Registered on 25 May 1922 according to the
museum registry]. See the discussion for details
related to the syntype series.
Paralectotypes. ZSI 18608 (male);
collected from Waynaad (=Wayanad), Nilgiris,
Kerala, India, by Frank Wall. Note: The other
paralectotypes most probably assorted with L.
calamaria, and others might be lost, misplaced
or destroyed.
Description of lectotype. Male, SVL 165.0 mm;
tail length 70.0 mm; head elongate (HL 5.5% of
SVL), twice as long as wide (HW 47.2% of HL),
slightly flattened, distinct from neck; snout
elongate (ES 28.6% of HL), moderate, blunt in
dorsal view, rounded in lateral profile, forming
an oval shape, rather depressed.
Rostral shield large, hemispherical,
distinctly visible from above, rounded
posteriorly; interorbital width broad (IO 65.1%
of HW); internasals semicircular; nostrils rather
small; nasals undivided, elongate, replacing the
position of loreal, in anterior contact with
rostral, internasal and prefrontal dorsally, 1st and
2nd supralabial ventrally; prefrontal rather large,
broader than long, and subhexagonal; frontal
large, subhexagonal, elongate posteriorly and
longer than its width; supraoculars wide,
elongated, subrectangular, posteriorly wider;
parietals large, butterfly wing-like in shape,
bordered by supraoculars, frontal, upper
postocular anteriorly, anterior and upper
posterior temporals, and six dorso-nuchal scales
posteriorly; no loreal scale; one preocular (both
sides), vertically elongated, hexagonal, in
contact with prefrontal and nasal anteriorly,
supraocular dorsally, and 2nd & 3rd supralabial
ventrally; eye large (ED 19.8% of HL), round,
smaller than the size of snout length (ED 69.2%
of ES), pupil rounded; two postoculars, upper
postocular larger, pentagonal, contact with
supraocular, parietal and anterior temporal
broad, in narrow contact with lower postocular;
lower postocular crescent in contact with 4th and
5th supralabial ventrally, anterior temporal
posteriorly; temporals 1+2, elongated,
hexagonal; anterior temporal larger and slightly
longer than posterior temporals, in contact with
parietal dorsally, 5th and 6th supralabial ventrally;
posterior temporals smaller, lower one in contact
with 6th and 7th supralabials ventrally.
Supralabials 7 (on both sides), 4th–7th larger
in size, 6th being the largest; 1st supralabial in
contact with rostral anteriorly, nasals dorsally,
2nd supralabial with nasal and preocular dorsally,
3rd supralabial with preocular and orbit dorsally,
4th supralabial with orbit and the lower
postocular dorsally, 5th supralabial with lower
postocular and anterior temporal dorsally, 6th
supralabial with anterior temporal and lower
posterior temporal dorsally, and 7th supralabial
with lower posterior temporal dorsally and body
scales posteriorly.
Mental of moderate size, triangular, wider
than length; first infralabial pair larger than
mental plate and in broad contact with each
other, in contact with anterior chin shield
posteriorly; seven infralabials, 1st–4th in contact
with first chin shield, 4th infralabial largest in
size in narrow contact with the anterior chin
42

AMARASINGHE ET AL. 2020
41 TAPROBANICA VOL. 09: NO. 01
shield and in broader contact with the posterior
chin shield; 5th infralabial in narrow contact with
posterior chin shield, 6th–7th infralabials in
contact with gular scales; two larger anterior
chin shields in broad contact, and two smaller
posterior chin shields in narrow contact;
posterior chin shields bordered posteriorly by six
gular scales.
Body robust, elongate and sub-cylindrical;
costal scales in 16-15-13 rows, all smooth and
bluntly pointed; 126 ventral scales; anal plate
divided. Tail comparatively short (TL 42.4% of
SVL), robust and thick; 62 paired subcaudals.
Colouration. In preservation, dorsally
yellowish chestnut brown; the dorsolateral scales
(5–6 rows above the ventrals) edged with black
colour creating two longitudinal lines on each
side of the vertebral column, this line breaks up
and disappears on the anterior part of the body;
ventral side cream.
Dentition. Maxillary 24, Palatine 15,
Pterygoid 16, Mandibular 18 teeth (Wall 1919).
Table 1. Morphometric (in mm) and meristric character comparisons of Liopeltis calamaria (Günther, 1858)
syntypes; Liopeltis calamaria indica Deraniyagala, 1955 lectotype; Cyclophis nasalis Günther, 1864 holotype;
and other examined specimens―––‖ = not measured/counted.
Character
L. calamaria
L. calamaria indica
C. nasalis
Syntype
BMNH
1946.1.5.60
Syntype
BMNH
1946.1.21.64
Others
n=4
Lectotype
BMNH
1922.5.25.22
Others
n=3
Holotype
BMNH
1946.1.5.36
Locality
Ceylon (Sri Lanka)
Wayanad (India)
Unknown
snout–vent length (SVL)
227.0
192.0
195.0–218.0
165.0
150.0–335.0
285.0
tail length (TAL)
150.0
85.0
82.0–90.0
70.0
55.0–90.0
105.0
head length (HL)
10.1
9.3
10.2–10.8
9.1
10.5–12.5
10.2
head width (HW)
6.1
5.6
5.2–6.0
4.3
5.5–9.0
5.4
internarial distance
2.3
2.2
2.3–2.5
2.0
2.1–3.2
2.4
interorbital width (IO)
3.9
3.8
3.8–4.0
2.8
2.5–5.2
3.6
eye–nostril length
2.2
1.8
1.9–2.2
1.5
1.5–2.7
1.9
eye–snout length (ES)
3.8
3.0
2.9–3.1
2.6
2.4–3.8
2.9
mandible–eye distance
5.5
4.9
3.9–4.1
5.6
5.5
5.4
nostril diameter
0.4
0.4
0.4
0.2
0.4
0.3
eye diameter (ED)
2.1
1.8
1.9
1.8
1.9–2.3
2.2
costals
15, 15, 14
15, 15, 15
15, 15, 15
16, 15, 13
15, 15, 15
15, 15, 15
subcaudals
76
72
66–70
62
59–73
72
ventrals
130
134
131–135
126
121–158
151
supralabials
7
7
7, 8
7
6, 7
7
infralabials
7
7
7
7
6, 7
7
The following data was published in previous
literature for Liopeltis calamaria sensu lato.
Variation. See Tables 1 and 2.
Hemipenis. The hemipenis of Liopeltis
calamaria extends to the 10th caudal plate; distal
half calyculate; calyces smaller, deeply
scalloped, closely packed, only the papillae are
visible on the surface; the spines in the spinous
region are shorter, thicker and numerous (De
Silva 1980); a fold is present (Smith 1943).
Dentition. Maxillary 24–26, Palatine 15–16,
Pterygoid 15–16, Mandibular 18–21 teeth (Wall
1919, 1921, Smith 1943).
Natural history. A terrestrial, diurnal snake
usually preferring dry mixed deciduous habitats
near water bodies, and also mostly found
underneath rotten logs on the forest floor
covered with a thick layer of leaf litter. Notably,
found frequently near the forest edge and rarely
inside the forest itself or in deep forest. It
appears to be a highly-adaptable species found
between low-lying coastal scrub belts such as
Puttalam Bay and the adjacent Wilpattu coast to
tall mountainous sub-Alpine forest tracts such as
Kullu Manali in Himalaya and the montane
sholas of the Kannan Devan Hills of the Western
Ghats. This species is insectivorous, having a
preferred diet of smooth-bodied caterpillars,
crickets, grasshoppers and spiders (Ditmars
1910). It is known to be oviparous (Deoras
1965). It does not try to bite when disturbed and
prefers to hide or escape. When defending, it
puffs up the anterior portion of the body to
display the darker interstitial skin in the neck
region (Karunarthna & Perera 2010). Natural
predators are assumed to be ophiophagous birds
such as crested serpent eagles (Spilornis cheela),
shikras (Accipiter badius), white-breasted king
43

TAXONOMY AND DISTRIBUTION OF Liopeltis calamaria
40 TAPROBANICA VOL. 09: NO. 01
fishers (Halcyon smyrnensis), and grey hornbills
(Ocyceros gingalensis). It is known to inhabit
mainly mountainous tracts including low (< 300
m see Karunarathna et al. 2004, Karunarathna &
Perera 2010, Ganesh et al. 2018), mid (900 m
see Chikane & Bhosale 2012) and high
elevations (1600–2,000 m, see Ferguson 1902,
Wall 1919, Malhotra & Davis 1991).
Distribution. Even though Liopeltis
calamaria is considered a widely distributed
species throughout Sri Lanka and India
(Somaweera 2006, Whitaker & Captain 2008,
Smith 1943), it is also considered to be rare in
both countries, which are separated by the 32 km
wide Palk Strait (see Ferguson 1877, Wall 1919,
1921). Wall (1921) and Taylor (1950) did not
examine any specimens from Sri Lanka.
In Sri Lanka: Subsequent to its original
description by Günther in 1858, Deraniyagala
(1955) recorded this subspecies from Badulla,
Uva patanas, Haputale (Uva Province),
Varahena (Southern Province), Marai Vila
(=Marawila, North-western Province), Sigiri
(=Sigiriya, North-central Province). De Silva
(1990) reported one specimen from Opatha
(Southern Province) and Batuwita (2001) found
one from Kottawa (Southern Province). The
latter author further recorded two specimens
present at National Museum of Sri Lanka
(NMSL), which had been collected from
Periyankulam (Madu Road) and Polonnaruwa
(North Central Province). Karunarathna et al.
(2004) observed this species at Kukulugala
(Sabaragamuwa Province) and Ritigala (North-
central Province). Later, Karunarathna & Perera
(2010) and Karunarathna & Amarasinghe (2010)
recorded it from Nilgala (Uva Province) and
Eluwankulama (Northwestern Province). We
additionally observed L. calamaria at Paalaviya-
Puttalam (Northwestern Province), Duviliella-
Kaltota (Uva Province), Wasgomuwa (North-
central Province), Udawalawa (Uva Province),
and Minneriya (North-central Province). Since
its original description in 1858 (over 160 years
ago), this species has been sighted
approximately only 20 times, indicating that it
is one of the rarest snakes in Sri Lanka (see Fig.
4 for current distribution records of this species).
In India: Beddome (1863) was perhaps the
first to record this species from this country
when he reported one specimen from the
Shevaroy Hills in the Eastern Ghats. He
explicitly states that he had recorded the species
which Günther had described from Ceylon (=Sri
Lanka). Subsequent treatises state its distribution
in India to be ―peninsular India‖ (Günther 1864),
―South India‖ (Theobald 1876) and ―Madras
Presidency, Bombay‖ (Boulenger 1890).
Phipson (1888) reports specimens from Ceylon
and Mahabaleshwar (in Western Ghats) housed
in the Bombay Natural History Society Museum.
Sclater (1891) lists specimens from Sirgaja,
Chota Nagpur and from Tinnevelley (ZSI 4421),
Madras Presidency (ZSI 8734 and 8735; both
coll. by R.H. Beddome). Boulenger (1894)
examined one additional specimen from Sri
Lanka and five specimens from India: Kotagiri –
Nilgherries, Sevagherry Ghat, Madras
Presidency, and Matharan, all localities in the
Western Ghats. Ferguson (1902) reports of
having observed one atop the Kannan Devan
Hills, near Munnar in the wet, montane regions
of the Southern Western Ghats. Wall (1919)
mentions it as being ―evidently an uncommon
snake‖, he reports it from Wynad and remarks
that its scalation matches Boulenger‘s
description well. Until this period, the species
had been known only from Sri Lanka and the
peninsula of India. But soon afterwards, records
from the Himalayan region started to emerge.
Wall (1921) did not observe any specimen from
Sri Lanka, but reported Indian specimens from
Mahableshwar, Wynaad, Nilgiris, and
Bangalore. Later he (Wall 1924) mentions this
species from Buxa Dooras, Songara of Gond
District, Khurkhana in Philibit District,
Melanghat in Almorah District (all in the far
North of India), Surguja in Chota Nagpur
District, Mahabaleshwar, Mysore plateau and
Tinnevelley Hills. Mullan (1927) recorded it
from Panchgani in the Satara ranges of the
Western Ghats. Smith (1943) stated the
distribution to be ―Western Ghats, as far north as
Matheran, Tinnevelley Hills, Mysore Plateau,
Bangalore, United Provinces, Chota Nagpur‖
while Whitaker (1978) located it from ―Hilly
areas throughout India, except extreme
northwest‖. Constable (1949) reported one male
and two females collected by R.H. Beddome
from ‗near Madras‘ lodged at the Museum of
Comparative Zoology, Harvard, USA. Very few
recent workers have reported the species from
India. Malhotra & Davis (1991) report sighting
of three specimens in rock piles situated within
the montane grasslands atop the Srivilliputhur
Hills, at 1290–1690 m. Recent reports originated
from Nallamalai Hills in Andhra Pradesh (Rao et
al. 2005), Kambakkam Hills in Andhra Pradesh
(Ganesh & Asokan 2010), Kaas plateau in
Maharashtra (Chikane & Bhosale 2012),
44

AMARASINGHE ET AL. 2020
41 TAPROBANICA VOL. 09: NO. 01
Meghamalai Hills in Tamil Nadu (Bhupathy &
Sathishkumar 2013), Kalakadu Mundanthurai
Tiger Reserve in Tamil Nadu (Narayanan 2016),
and Vellore as well as the Gingee Hills of the
Eastern Ghats in Tamil Nadu (Ganesh et al.
2018). Bhattarai et al. (2018) also reported the
species from Nepal (Chitwan, Makwanpur). We
suggest a careful study of those specimens and
the adjoining Himalayan population (also see
Wall 1924) that are fully geographically
separated from the Peninsular Indian population
by the mighty Indo-Gangetic plains (see Fig. 4).
Threats and Conservation. This species is
found mostly in habitats where forest fires are a
distinct threat, so is exposed to illegal logging
and chena cultivation (shifting cultivation).
Clearing for agriculture, expanding human
settlements, direct persecution by humans,
because of mythical beliefs are some other
common threats to this snake. In some habitats
(e.g. Eluwankulama, Sri Lanka) mining for
cement is identified as a threat (Karunarathna &
Perera 2010). The specimens recorded from
Minneriya and Nilgala (Sri Lanka) were road
kills, which is another threat to the species. In
India this species has also been reported as road
kills (Ganesh et al. 2018) and had featured in
snake rescue data of animals affected by
urbanisation (Nande & Deshmukh 2007). In
Nepal, this species has been reported to suffer
from wanton killing out of fear by locals
(Bhattarai et al. 2018).
Table 2. Ventral and subcaudal scale count comparisons between Sri Lankan and Indian populations of
Liopeltis calamaria based on published literature and new data (this study): SL, Sri Lanka; IN, India; ―–― Not
applicable.
Discussion
Phipson (1888) misspelt the generic name in his
work, as Cyclophia. Constable (1949)
challenged Smith‘s generic allocation and
represented this species as Opheodrys
calamaria, remarking that he follows most
American herpetologists in considering Liopeltis
a synonym of the unrelated, Nearctic genus
Opheodrys Fitzinger, 1843. Wallach et al.
(2014) changed the widely used Latin species
epithet ―calamaria‖ which is feminine in gender,
to ―calamarius‖ which is masculine in gender.
The generic name Liopeltis Fitzinger, 1843
is feminine, meaning ―having smooth shields‖. It
is derived from the Ancient Greek ―péltē‖,
which means a small crescent-shaped shield
(peltis in plural) and ―leióō‖, which means
smooth. Thus, the specific epithet calamaria is
correctly placed, which also is feminine in the
nominative case (originated from calamarius),
derived from the Greek word ―kalamos‖,
meaning ―tube, reed‖. Therefore, according to
the Article 31.2 of the International Code of
Zoological Nomenclature (ICZN 1999), we
refute the amended epithet ―calamarius‖ used by
Wallach et al. (2014), and reinstate the correctly
used previous species epithet ―calamaria‖ as the
current nomenclatural combination of this taxon.
At the same time the subspecific epithet given
by Deraniyagala (1955), is also corrected
herewith as Liopeltis calamaria indica (not
indicus) following the same article of the ICZN.
Günther‘s (1864) assignment of Liopeltis
nasalis from an unknown locality (probably
Source
Ventrals
Subcaudals
Males
Females
Males
Females
SL
IN
SL
IN
SL
IN
SL
IN
Günther (1858) syntypes
(n=2)
129, 134
–
–
–
72, 76
–
–
–
Boulenger (1894) (n=6)
134
–
–
130–154
68
–
–
64–70
Wall (1919) (n=5)
–
126–132
–
135–139
–
59–70
–
53–57
Constable (1949) (n=3)
–
–
–
133–146
–
–
–
60–71
De Silva (1969) (n=6)
129–132
–
134, 139
–
65–73
–
61, 64
–
Batuwita (2001) (n=1)
133
–
–
–
–
–
–
–
Karunarathna et al.
(2004) (n=2)
–
–
148, 156
–
–
–
52, 58
–
Karunarathna & Perera
(2010) (n=2)
–
–
159, 162
–
–
–
61, 65
–
This study (n=7)
–
121–136
–
140, 158
–
60–73
–
65–73
Summary
129–134
121–136
134–162
130–158
65–76
59–73
52–65
53–73
45

TAXONOMY AND DISTRIBUTION OF Liopeltis calamaria
40 TAPROBANICA VOL. 09: NO. 01
from India) is based on a similar diagnosis of
having 149 ventrals and 77 subcaudals.
Therefore, we presume that Günther‘s nomen C.
nasalis as available for the Indian population
even though the locality is stated as unknown.
The morphological and meristic characters of
the holotype of C. nasalis, however, fit within L.
calamaria, we hence hereby follow Boulenger‘s
view that C. nasalis is a junior subjective
synonym of L. calamaria. The attempt to trace
the type specimen of Jan‘s (1862, 1865)
description of Homalosoma baliolum was
unsuccessful. According to the curator of the
collection it is missing, misplaced or has been
destroyed (Pers. comm. Stefano Scali, Museo
Civico di Storia Naturale di Milano, Italy; and
Richard Gemel, Naturhistorisches Museum
Wien, Vienna, Austria). However, based on the
description of Jan (1862) it has 159 ventrals and
67 subcaudals (most probably a female
specimen) and thus sits within the range known
for this species [see Fig. 5 of this publication for
the illustration in Jan (1865) respectively], but
probably represents an unidentified population
either from Sri Lanka or India. At this time, we
therefore consider it a junior subjective synonym
of L. calamaria, thus supporting the view of
Boulenger (1890).
De Silva (1969) stated that this species
shows slight sexual dimorphism. Deraniyagala
(1955) assigned Indian population to the name
Liopeltis calamaria indicus (sic) based on 126–
163 ventrals and 53–76 subcaudals (vs 121–138
ventrals and 61–76 subcaudals in the Sri Lankan
population). In Table 1 and 2, for both sexes, we
show that the Sri Lankan population has a
minimum of 130 ventrals (6 specimens
examined + 11 specimens from literature), and
Indian populations have a minimum of 134
ventrals (20 specimens examined). Even though
neither a type specimen nor a series of syntypes
for L. c. indica has been formally designated,
Deraniyagala (1955) clearly referred to the
specimens examined by Wall, stating the
following ―…ventrals 121 to 138 in Ceylon
specimens, and 126 to 163 in Indian ones
(Wall),…‖. Therefore, according to the Article
72.1.1, in the absence of a type designation, we
consider all the specimens from India available
to Wall (1919, 1921, 1924) as type specimens,
hence a type-series (syntypes). Wall (1921)
stated that he had seen several examples
including specimens from Mahableshwar,
Wynaad, Nilgiris, and Bangalore, so there must
have been in his possession at least four
specimens. That being said, and unfortunately,
we were able to locate only two among them
(BMNH 1922.5.25.22 and ZSI 18608), which
were both collected from Wynaad, India by
Frank Wall himself. Furthermore, the ventral
counts of those specimens are 126 and 121
respectively. Therefore, we regard BMNH
1922.5.25.22 as one of Wall‘s specimens stated
by Deraniyagala (syntypes, 126–163 ventrals),
and furthermore designate it as the lectotype of
Liopeltis calamaria indica, in order to stabilize
the subspecific name with a name bearing type
specimen (onomatophore).
Interestingly the lectotype of L. c. indica,
along with the paralectotype (ZSI 18608) also
collected from Nilgiri-Wynad, is on the lower
margin of ventral counts 121 and 126 (vs 130–
134) and subcaudal counts 59 and 62 (vs 68–76)
compared to the Sri Lankan specimens (see
Table 1). Also, if such a non-overlapping
scalation range holds true for a larger series of
specimens, the subspecies L. c. indica will turn
out to be endemic to Western Ghats (probably
around the area of Wayanad), while the forma-
typica L. c. calamaria would refer to the
population in Sri Lanka (and perhaps some parts
of South India). However, there might be
several, more cryptic, species isolated in narrow
ranges, especially in the Indian region.
Specifically, we suggest a careful study of the
Himalayan / Nepalese population (also see Wall
1924) that are fully separated from the
Peninsular Indian population by the mighty
zoogeographical barrier of the Indo-Gangetic
plains, where there are no records to date.
Molecular data from different populations all
over the geographic range of this ‗species‘ are
required to further understand its evolutionary
history and dispersal route, and this might also
give evidence for raising the status from either
synonyms or subspecies to a full species level.
Acknowledgments
We thank the former Director, K. Venkataraman
(ZSI) for granting research permission. 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. Silke Schweiger
and Georg Gassner (NMW) gave us the
opportunity for examining specimens under their
46

AMARASINGHE ET AL. 2020
41 TAPROBANICA VOL. 09: NO. 01
care. Dushmantha Kulathunga is acknowledged
for photographs. SK thanks P. Antony, K.
Gunawardena and V. Weeratunga for the travel
grant to visit ZSI (India). SRG thanks the
Executive Chairman and Trustees of Chennai
Snake Park Trust for encouraging his research
activities; and the Field Director NSTR, the
Principal Commissioner, and the Secretary
Madras Government Museum are also
acknowledged for access to material under their
care. Ivan Ineich (Muséum national d'Histoire
naturelle), Philip Bowles (IUCN), and two
anonymous reviewers are acknowledged for
valuable comments and reviewing the
manuscript. Finally, we thank J. Supriatna and
the staff of the Research Center for Climate
Change, University of Indonesia; and Anslem de
Silva for their support.
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Appendix I. Other specimens examined
Liopeltis calamaria (30 ex.): Sri Lanka (n=6): Ceylon (Sri Lanka): BMNH 1946.1.5.60 & 1946.1.21.64
(syntypes), 1846.12.2, NMW 26966:1–2; Diyatalawa, Uva Province: BMNH 1933.12.6.13.
Peninsular India (n=16): Madras Presidency: ZSI 8734, 8735, BMNH 1866.12.15.8; Tinnevelly
Hills: ZSI 4421; Hosur, Mysore: BMNH 1925.4.2.51; Kotagiri, Nilgherries: BMNH 1891.11.27.12;
Catounni, Bangalore: BMNH 1931.6.15.16–18; Matheran: BMNH 1869.8.28.146a–c; Wynaad:
BMNH 1922.5.25.532 (lectotype of Liopeltis c. indica), ZSI 18608 (paralectotype of Liopeltis c.
indica). Nallamalai, Andhra Pradesh: ERMR29a; Andhra Pradesh, Kambakkam: MAD 8-31-VII-22.
Himalaya / Nepal (n=7): Songara, Gonda: ZSI 16385; Kurkhana, Uttar Pradesh: ZSI 16437;
Melaghat Almorah: ZSI 16438; BMNH 1940.3.4.22–23; Punjab: BMNH 1948.1.7.5, 1922.6.14.1.
Unknown locality (probably India) (n=1): BMNH 1946.1.5.36 (holotype of Cyclophis nasalis).
Liopeltis frenata (1 ex): India: Khasi Hills: BMNH 1946.1.1.72 (holotype).
Liopeltis pallidonuchalis (4 ex): Vietnam: Gia Lai: ZMMU R-15682 (holotype); Thua Thien–Hue: ZFMK
83105 (paratype), DTU 307 (paratype); Quang Binh: VNUH 9.7.06–1.
Liopeltis rappi (5 ex): India: Sikkim: BMNH 1946.1.5.61 (holotype); Simla: NMW 26963:1; Kulu at
Kategarh: NMW 26963:2; Myanmar: Rangoon: NMW 26962:1–3.
Liopeltis tricolor (14 ex): Indonesia: Java: BMNH 1946.1.5.34, 69.12.4.127 (Syntypes), BMNH uncat.
(holotype of Ablabes schlegelii), NMW 26967:1–4; Sumatra: NMW 26968:2, SMF 81197, ZFMK
33533; Kalimantan: NMW 26968:3; Banka: RMNH 554; East Indian Archipelago: RMNH 4036;
Peninsular Malaysia: Penang: NMW 26968:1.
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40 TAPROBANICA VOL. 09: NO. 01
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