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Systematics and ecology of Oligodon sublineatus Duméril, Bibron & Duméril, 1854, an endemic snake of Sri Lanka, including the designation of a lectotype

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The description of Oligodon sublineatus Duméril, Bibron & Duméril, 1854 was based on two syntypes located at Paris Natural History Museum (MNHN). The larger specimen (SVL 254 mm) was described in detail, but erroneously labelled as originating from the Philippines, the second specimen (SVL 150 mm) was labelled as originating from 'Cey-lan' (=Sri Lanka). The smaller specimen, up to this point, has always been considered as the holotype by monotypy. Since recognising the larger specimen in the collection of MNHN as a syntype, we hereby designate it as the lectotype of Oligodon sublineatus and redescribe comprehensively both syntypes. Oligodon sublineatus (SVL 152–310 mm) has 130–161 ventral scales, 23–42 divided subcaudals, a divided anal plate, a loreal, seven supralabials, and 1+2 temporals. Furthermore, we provide a detailed account of the distribution and natural history of this widely distributed Sri Lankan endemic snake.
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museum für naturkunde
Systematics and ecology of Oligodon sublineatus Duméril, Bibron
& Duméril, 1854, an endemic snake of Sri Lanka, including the
designation of a lectotype
A. A. Thasun Amarasinghe1, D. M. S. Suranjan Karunarathna2, Patrick D. Campbell3, Ivan Ineich4
1 Research Center for Climate Change, University of Indonesia, Gd. PAU Lt. 8.5, Kampus UI, Depok 16424, Indonesia
2 Nature Explorations and Education Team, No. B–1 / G–6, De Soysapura Housing Scheme, Moratuwa 10400, Sri Lanka
3 Department of Life Sciences, Darwin Centre, Natural History Museum, Cromwell Road, South Kensington, London SW7 5BD, England
4 Muséum national d’Histoire naturelle, Sorbonne Universités, ISYEB (Institut de Systématique, Évolution et Biodiversité),
UMR 7205 CNRS, EPHE, MNHN, UPMC, 57 rue Cuvier, CP 30 – F-75005 Paris, France
http://zoobank.org/8D420E93-4EA6-4CD7-9885-F522A0296288
Corresponding author: A. A. Thasun Amarasinghe (thasun@rccc.ui.ac.id)
Abstract
The description of Oligodon sublineatus Duméril, Bibron & Duméril, 1854 was based
on two syntypes located at Paris Natural History Museum (MNHN). The larger specimen
(SVL 254 mm) was described in detail, but erroneously labelled as originating from the
Philippines, the second specimen (SVL 150 mm) was labelled as originating from ‘Cey-
lan’ (=Sri Lanka). The smaller specimen, up to this point, has always been considered
as the holotype by monotypy. Since recognising the larger specimen in the collection of
MNHN as a syntype, we hereby designate it as the lectotype of Oligodon sublineatus and
redescribe comprehensively both syntypes. Oligodon sublineatus (SVL 152–310 mm)
has 130–161 ventral scales, 23–42 divided subcaudals, a divided anal plate, a loreal,
seven supralabials, and 1+2 temporals. Furthermore, we provide a detailed account of
the distribution and natural history of this widely distributed Sri Lankan endemic snake.
Key Words
Colubridae
distribution
kukri snake
natural history
systematic
taxonomy
Sri Lanka
Introduction
The colubrid genus Oligodon Fitzinger, 1826 is currently
known to include 75 valid species (Uetz and Hallermann
2014, 1st September 2014). Only four species have been
recorded from Sri Lanka: Oligodon calamarius (Linnae-
us, 1758); Oligodon arnensis (Shaw, 1802); Oligodon
taeniolatus (Jerdon, 1853); and Oligodon sublineatus
Duméril, Bibron & Duméril, 1854. Following the de-
scription of Oligodon sublineatus in 1854, this species
has since been recorded from various locations in Sri
Lanka (Boulenger 1890, 1894; Wall 1921; Smith 1943;
Received 24 March 2015
Accepted 25 March 2015
Published 22 April 2015
Guest academic editor:
Johannes Penner
Deraniyagala 1955; Taylor 1953; De Silva 1980; de Silva
1990; Das and de Silva 2005; Somaweera 2006; Green
2010; Green et al. 2010; David and Vogel 2012; Deep-
ak and Harikrishnan 2013; Pyron et al. 2013). The orig-
inal description of O. sublineatus was clearly based on
two syntypes, the rst and larger of the two having no
collector details associated with it, and labelled as com-
ing from the Philippines, obviously in error. The second
syntype came from Ceylan (= Ceylon) and was collect-
ed by Leschenault according to the original description.
The larger specimen (MNHN 3238) has been correctly
identied as O. sublineatus in the MNHN collections but,
Zoosyst. Evol. 91 (1) 2015, 71–80 | DOI 10.3897/zse.91.4971
Copyright A. A. Thasun Amarasinghe et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0),
which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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Amarasinghe, A.A.T.: Oligodon sublineatus, an endemic snake of Sri Lanka
72
until now, was not recognized as a syntype. However the
smaller specimen (MNHN 3239) has been mistakenly
considered as the holotype by Wallach et al. (2014). We
have identied the large specimen as being one of the
syntypes, so we hereby designate it as a lectotype, and
redescribe it in detail in order to stabilize that name with
a recognised type specimen.
Methods
Museum acronyms follow Sabaj Pérez (2014). Spec-
imens were examined in the collections of the British
Museum of Natural History, UK (BMNH); Muséum na-
tional d’Histoire naturelle, France (MNHN); Naturhis-
torisches Museum Basel, Switzerland (NMB); and Na-
tional Museum of Sri Lanka (NMSL). Morphometric
and meristic data for species comparisons were obtained
from examined specimens (see Appendix 1). We checked
the external morphology of specimens with a Wild M3Z
stereomicroscope and photographed them with a Canon
EOS 7D SLR digital camera. The map was construct-
ed based on Cooray (1967). The conservation status of
the species was evaluated using Red List Categories and
Criteria in IUCN Standards and Petitions Subcommit-
tee (2013: version 10.1) to assess their risk of extinc-
tion. Sex was determined by ventral tail incision of adult
specimens followed by the checking for the presence or
absence of hemipenes. All the natural history data were
taken from our own eld observation notes made during
the last ten years.
The following characters were measured with a digital
caliper (±0.1 mm) on the left side of the body for symmet-
rical characters: eye diameter (ED, horizontal diameter of
eye); eye–nostril length (EN, 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 (ND, horizontal diameter of nostril); in-
ternarial distance (IN, least distance between nostrils);
mandible–posterior eye distance (MPE, 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 (TAL, measured from anterior margin of vent
to tail tip). Meristic characters were taken as follows: su-
pralabials and infralabials (SUP and INF, rst labial scale
to last labial scale bordering gape); costal scales (COS,
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 ventral scale prior to preanal);
when counting the number of ventral scales (MVS), we
scored specimens according to method described by
Dowling (1951). We counted subcaudal scales (SUB)
from rst postcloacal scale to the scale before the tip of
the tail.
Results
Oligodon sublineatus Duméril, Bibron & Duméril, 1854
Figs 1–3; Tables 1, 2
Remarks. Standard morphometric and meristic data of
the two syntypes are presented in Table 1. We hereby rec-
ognise two syntypes: the larger specimen (MNHN 3238)
and the smaller specimen (MNHN 3239). Uncertainties
still exist in Oligodon taxonomy and O. sublineatus may
represent a cryptic species complex in Sri Lanka (see ta-
ble 2 showing the wide range of subcaudal and ventral
counts within O. sublineatus), therefore it is necessary to
stabilize the name with a recognised lectotype. There are
two main reasons for selecting MNHN 3238 as the lec-
totype: (1) it was used in the original description and its
morphometric data has been provided and (2) it is a fully
grown, well-developed and well preserved adult speci-
men in good shape.
Lectotype (here designated). MNHN 3238, adult female
collected from the Philippines (mistakenly so in the origi-
nal description) [from Java (also in error) according to the
museum registry] by an unknown collector [by Bosc (in
error) according to the museum registry].
Paralectotype. MNHN 3239, sub adult male collected
at ‘Ceylan’ [= Sri Lanka] by Leschenault. This specimen
was previously erroneously considered as the holotype by
Wallach et al. (2014).
Diagnosis. Oligodon sublineatus shows sexual dimorphism
in scalation (Table 2) and is distinguished from all conge-
ners by the following characters: SVL 152–310 mm; TAL
20.0–42.0 mm; 130–161 ventrals; 23–42 subcaudals (divid-
ed); anal plate divided; loreal present; seven supralabials;
temporals 1+2; ventral side with three series of dark brown
points forming almost continuous stripes, with the middle
series of points absent on the tail; dorsal coloration (live
or in alcohol) greyish brown, speckled with small elongat-
ed spots irregularly placed; posterior part of the jaws has
a large, oblique spot extending along the neck posteriorly;
dorsally a “˄” shaped marking between the eyes, which
continues laterally across them; an irregular, brown, trans-
versal band from the frontal to the post-parietal region.
English translation of the original French description
in Duméril, Bibron & Duméril (1854: 57). Characters.
Ventral side with three series of points forming stripes.
This species is mostly characteristic, as its specic
name, by having three black stripes along the ventral side,
which are made up of a series of points, meeting together.
The two stripes outside the ventral plates form a continu-
ous line up to the ventral surface of the tail, but the central
one is made up of distinct points in the centre of the ven-
tral plates. These points are quite large, round and wide
posteriorly, and are as notched at the front; the median
stripe does not prolongate onto the ventral side of the tail.
Zoosyst. Evol. 91 (1) 2015, 71–80
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Dorsal coloration grey, speckled with lines or with
small elongated spots irregularly placed; however,
around the anterior third of the body and laterally, three
of those spots appear enlarged with increased width, hav-
ing a circular border. The spots are constricted central-
ly and have white borders. The posterior section of the
jaws has a large, oblique patch along the neck posteriorly
where it forms a tip pointing in the opposite direction to
the characteristic collar of the rst species [note from the
translator: Oligodon sub-quadratum].
Dorsal scales are very smooth, and are close to each
other; they are slightly overlapping, like roof tiles, mostly
around the tail area, and in this respect, very skink-like in
appearance.
Rostral plate is notched, and crescent shaped; other
plates covering the head are large and clearly distinct as
in colubrids.
We were only able to examine one well preserved
specimen, having no clues as to the origin of the speci-
men [the Philippines] and the name ‘Oligodon torquatus’
appears along with the letter “R” on the jar.
Another specimen, younger and obviously added
much later, had a median stripe made up of numerous
spots which were less distinct, was collected from Ceylan
by Mr. Leschenault. This specimen bears all the charac-
ters previously described: the large, brown, post-maxil-
lary mark set posteriorly on the neck forming a croissant
shape; with a laterally set, black mark extending onto the
anterior third of the body.
We counted 15 scale rows on that specimen, 155 ven-
trals and 25 subcaudals.
Total length was 180 cm [sic]; among them 155 for
SVL and 25 for the tail.
Table 1. Morphometric and meristic characters of Oligodon
sublineatus lectotype (MNHN 3238) and paralectotype
(MNHN 3239).
Character MNHN 3238
Lectotype (female)
MNHN 3239
Paralectotype (male)
snout–vent length (SVL) 254.0 150.0
tail length (TAL) 35.0 27.0
head length (HL) 10.8 8.1
head width (HW) 4.7 4.1
internarial distance (IN) 2.1 1.5
interorbital width (IO) 3.7 3.3
eye–nostril length (EN) 1.9 1.7
eye–snout length (ES) 3.4 2.7
mandible–posterior eye
distance (MPE) 5.8 4.1
nostril diameter (ND) 0.3 0.4
eye diameter (ED) 1.7 1.4
costals (COS) 17-15-15 17-15-15
subcaudals (SUB) 28 36
ventrals (MVS) 150 138
supralabials (SUP) 7 (3, 4 touch eye) 7 (3, 4 touch eye)
infralabials (INF) 8 8
Figure 1. A live male of Oligodon sublineatus (not collected) at Sinharaja Forest Reserve, Sri Lanka (photo: H. Jayasinghe).
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Description of the designated lectotype, MNHN 3238.
Adult female, SVL 254 mm; tail length 35 mm; head
elongate (HL 4.3% of SVL), twice as long as wide (HW
43.5% of HL), slightly attened, distinct from neck;
snout elongate (ES 31.5% of HL), moderate, blunt in
dorsal view, rounded in lateral prole, forming an oval
shape, rather depressed.
Rostral shield large, hemispherical, distinctly visible
from above, pointed posteriorly; interorbital width broad
(IO 78.7% of HW); internasals semicircular; nostrils rath-
er large; nasals completely divided by nostrils into two
scales unequal in size; anterior nasal larger, in anterior
contact with rostral, internasal dorsally, 1st SUP ventrally;
posterior nasal in contact with internasal and prefrontal
dorsally, loreal posteriorly, 1st and 2nd SUP ventrally; pre-
frontal rather large, broader than long, and subhexago-
nal; frontal large, subhexagonal, elongate posteriorly and
longer than its width; supraoculars narrow, elongated,
subrectangular, posteriorly wider; parietals large, butter-
y wing-like in shape, bordered by supraoculars, frontal,
upper postoculars anteriorly, anterior and upper posterior
temporals, and six dorso-nuchal scales posteriorly; loreal
large, slightly elongated, subrectangular, in contact with
prefrontal dorsally and preoculars posteriorly, ventrally
only touching the 2nd SUP; one preocular (both sides),
vertically elongated, subrectangular, in contact with pre-
frontal and loreal anteriorly, supraocular dorsally, and 3rd
SUP ventrally; eye moderate (ED 15.7% of HL), ellip-
Table 2. Some sexual dimorphic characters of Oligodon sublineatus based on examined material. See Methods for abbreviations.
sex SVL (mm) TAL (mm) VEN SUB
male 152.0–227.0 (n=12) 25.0–38.0 (n=12) 135–151 (n=18) 30–42 (n=18)
female 156.5–295.0 (n=17) 20.0–42.0 (n=17) 145–159 (n=21) 24–29 (n=21)
Figure 2. Oligodon sublineatus male, NMSL 5161 collected from Nuwara Eliya (1600 m a.s.l.), Sri Lanka: A dorsal aspect of head
B ventral aspect of head C lateral aspect of head D lateral aspect of midbody E dorso-lateral aspect of right hemipenis (scale = 1 mm).
Zoosyst. Evol. 91 (1) 2015, 71–80
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tical, nearly a half of the size of snout length (ED 50%
of ES), pupil rounded; two postoculars, upper postocu-
lar smaller, quadrangular, contact with supraocular and
parietal broad, in narrow contact with anterior temporal;
lower postocular crescent in contact with 4th and 5th SUP
ventrally, anterior temporal posteriorly; temporals 1+2,
elongated, hexagonal; anterior temporal larger and longer
than posterior temporals, in contact with parietal dorsal-
ly, 5th and 6th SUP ventrally; posterior temporals smaller,
lower one in contact with 6th and 7th supralabials ventrally.
Supralabials 7 (on both sides), 4th–7th larger in size;
1st SUP in contact with rostral anteriorly, nasals dorsally,
2nd supralabial with posterior nasal and loreal dorsally, 3rd
SUP with preocular and orbit dorsally, 4th SUP with or-
bit and the lower postocular dorsally, 5th SUP with lower
postocular and anterior temporal dorsally, 6th supralabial
with anterior temporal and lower posterior temporal dor-
sally, and 7th SUP with lower posterior temporal dorsally
and body scales posteriorly.
Mental of moderate size, triangular; rst infralabial
pair larger than mental plate and in broad contact with
each other, in contact with anterior chin shield posteri-
orly; eight infralabials, 1st–5th in contact with rst chin
shield, 5th infralabial largest in size in narrow contact with
the anterior chin shield and in broader contact with the
posterior chin shield; 6th–8th infralabials in contact with
gular scales; two larger anterior chin shields, and two
smaller posterior chinshields all in broad contact; poste-
rior chin shield bordered posteriorly by six gular scales.
Body robust, elongate and sub cylindrical; costal
scales in 17-15-15 rows, all smooth and bluntly pointed;
150 ventral scales; anal plate divided. Tail comparatively
short (TL 13.8% of SVL), robust and thick; subcaudals
28, divided.
Description of the paralectotype, MNHN 3239, and an
additional specimen, NMSL 5161. The values of NMSL
5161 (when different) included within parenthesis. Sub
adult male (adult male), SVL 150.0 (183.3) mm; head
elongate, HL 5.4 (5.6)% of SVL, twice as long as wide,
HW 50.6 (53.9)% of HL, slightly attened, distinct from
neck; snout elongate, ES 31.4 (33.3)% of HL, moderate,
blunt in dorsal view, rounded in lateral prole, forming an
oval shape, rather depressed.
Rostral shield large, hemispherical, distinctly visible
from above, pointed posteriorly; interorbital width broad-
er, IO 80.5% of HW; internasals semicircular; nostrils
rather large; nasals divided into two scales unequal in
size; anterior nasal larger, in contact with the rostral plate
anteriorly, internasal dorsally, 1st SUP ventrally; posterior
nasal in contact with internasal and prefrontal dorsally,
loreal posteriorly, 1st and 2nd SUP ventrally; prefron-
tal rather large, broad, and subhexagonal; frontal large,
subhexagonal, elongate posteriorly and longer than its
width; supraoculars narrow, elongated, subrectangular,
posteriorly wider; parietals large, buttery-like in shape,
bordered by supraoculars, frontal, upper postoculars an-
teriorly, anterior and upper posterior temporals, and six
dorso-nuchal scales posteriorly; loreal large, slightly
elongated, subrectangular, in contact with prefrontal dor-
sally, preoculars posteriorly, posterior nasal anteriorly,
ventrally just meets the 2nd SUP; one preocular in both
sides, vertically elongated, subrectangular, in contact
with prefrontal and loreal anteriorly, supraocular dorsal-
ly, and 3rd SUP ventrally; eye moderate, ED 17.3 (17.6)%
of HL, elliptical, nearly a quarter of the snout length,
ED 51.9 (56.2)% of ES, pupil rounded; two postoculars,
upper postocular smaller, quadrangular, in contact with
supraocular and parietal broad, in narrow contact with an-
terior temporal; lower postocular crescent in contact with
4th and 5th SUP ventrally, anterior temporal posteriorly;
temporals 1+2, elongated, hexagonal; anterior temporal
larger and longer than posterior temporals, in contact
with parietal dorsally, 5th and 6th SUP ventrally; posterior
temporals smaller, lower one in contact with 6th and 7th
SUP ventrally.
Supralabials 7 on both sides, 4th–7th larger in size; 1st
SUP in contact with rostral anteriorly, nasals dorsally, 2nd
SUP with posterior nasal and loreal dorsally, 3rd SUP with
preocular and orbit dorsally, 4th SUP with orbit and the
lower postocular dorsally, 5th SUP with lower postocular
and anterior temporal dorsally, 6th SUP with anterior tem-
poral and lower posterior temporal, and 7th SUP with low-
er posterior temporal dorsally and body scales posteriorly.
Mental moderate, triangular; rst infralabial pair larg-
er than mental and contact with each other broad, in con-
tact with anterior chin shield posteriorly; eight infralabi-
als, 1st–5th in contact with rst chin shield, 5th infralabial
largest in size in narrow contact with anterior chin shield
and contact with posterior chin shield broad; 6th–8th in-
fralabials in contact with gular scales; two larger anterior
chin shields, and two smaller posterior chinshields all in
broad contact; posterior chin shield bordered posteriorly
by six gular scales.
Body robust, elongate and sub cylindrical; costal scales
in 17-15-15 rows, all smooth and bluntly pointed; 138
(142) ventral scales; anal plate divided. NMSL 5161 has
an everted hemipenis covered by lobes, non-bifurcated,
slightly clavate; base naked; sulcus spermaticus single and
deep; spinous ornamentation present on each lobe, shorter
spines at the apex; apex not divided into segments (Fig.
2E); tail comparatively short, TL 18.0 (20.5)% of SVL, ro-
bust and thick; subcaudals 36 in both specimens, divided.
Distribution. This species has never been recorded
outside of Sri Lanka, hence we here restrict terra-typi-
ca to Sri Lanka. Wall (1921), Smith (1943), Deraniya-
gala (1955), De Silva (1980), de Silva (1990), Das and
de Silva (2005), Somaweera (2006), Karunarathna and
Amarasinghe (2010, 2011, 2012), Botejue and Watta-
vidanage (2012), and Karunarathna et al. (2010, 2013)
recorded this species from Bellanwila–Attidiya, Berali-
ya, Colombo, Galle, Gammaduwa (Knuckles), Kitulga-
la, Kotmale, Kukulugala, Matugama, Nilgala, Peradeni-
ya, Ratnapura, Veyangoda, Welimada, and Yatiyantota
(Fig. 3). In addition to the above locations, during our
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Amarasinghe, A.A.T.: Oligodon sublineatus, an endemic snake of Sri Lanka
76
eldwork operations of the last decade we have record-
ed (not collected) O. sublineatus from a 10–1600 m
altitude range, including all vegetational zones of Sri
Lanka: Ambalangoda (6°14’42.35’’N, 80°03’44.56’’E),
Anuradhapura (8°20’46.43’’N, 80°25’43.77’’E), Atwel-
tota (6°31’33.87’’N, 80°18’12.02’’E), Baduraliya
(6°30’53.70’’N, 80°13’41.81’’E), Bibile (7°10’58.02’’N,
81°13’43.61’’E), Chilaw (7°35’11.49’’N, 79°49’16.54’’E),
Deniyaya (6°20’11.54’’N, 80°34’10.44’’E), El-
pitiya (6°17’39.31’’N, 80°08’44.78’’E), Elu-
wankulama (6°20’11.54’’N, 80°34’10.44’’E),
Gampaha (7°05’03.68’’N, 79°58’25.66’’E), Haba-
rana (8°11’12.43’’N, 80°50’17.89’’E), Hora-
na (6°42’24.74’’N, 80°03’02.77’’E), Illukkumbu-
ra (Knuckles) (7°35’46.09’’N, 80°46’14.10’’E),
Kalutara (6°35’13.29’’N, 80°58’21.49’’E), Kan-
neliya (6°12’37.49’’N, 80°24’04.60’’E), Keg-
alle (7°14’10.26’’N, 80°19’57.27’’E), Kotta-
wa-Homagama (6°47’07.00’’N, 79°57’52.17’’E),
Kurunegala (7°30’25.80’’N, 80°23’46.95’’E), Kuru-
wita (6°46’29.02’’N, 80°22’35.50’’E), Maharagama
(6°50’52.54’’N, 79°55’45.54’’E), Mahiyanganaya
(7°20’06.03’’N, 81°00’34.51’’E), Matara (5°57’08.63’’N,
80°31’59.74’’E), Monaragala (6°52’40.25’’N,
80°20’27.39’’E), Naula (7°44’18.42’’N, 80°43’38.22’’E),
Nugegoda (6°51’35.26’’N, 79°53’08.19’’E),
Panadura (6°42’42.76’’N, 79°54’24.44’’E), Pidu-
rutalagala (7°01’08.11’’N, 80°47’23.47’’E), Po-
lonnaruwa (7°56’15.64’’N, 81°01’15.38’’E),
Puttalam (8°02’42.88’’N, 79°51’38.84’’E), Rak-
wana (6°28’03.23’’N, 80°36’32.84’’E), Rit-
igala (8°12’35.71’’N, 80°35’02.78’’E),
Sinharaja (6°24’59.18’’N, 80°24’28.33’’E), Tana-
malwila (6°27’00.66’’N, 81°09’07.66’’E), Tissama-
haramaya (6°16’52.45’’N, 81°16’41.40’’E), Trin-
comalee (8°35’57.38’’N, 81°10’15.73’’E), Udawalawe
(6°26’48.46’’N, 80°52’26.25’’E), Wasgomu-
wa (7°43’23.36’’N, 80°58’06.01’’E), Wilpattu
(8°30’51.13’’N, 79°57’44.67’’E), Yagirala (6°22’47.13’
N, 80°10’23.93’’ E) (see Fig. 3 for the distribution map).
The result of the application of the IUCN (2013) B2
a, b (iii) Red List criteria shows that O. sublineatus as
Least Concern (LC): recorded from an altitude range of
10–1600 m in all vegetation zones of Sri Lanka. Its area
of occupancy is 6,000 km2, and its extent of occurrence
is 40,000 km2.
Natural history. A nocturnal snake, sometimes active
during day time. Temperature, humidity, and light inten-
sities for daytime activity were respectively measured at
24.8–27.2 °C, 67–82%, and 38–365 lux, based on 50 ob-
servations in dense forested areas. It usually does not bite,
but if this does occur then it will lead to soreness, pain
and temporary bleeding in the victim. Biting has been oc-
casionally observed during touching or handling attempts
by the victim. When frightened, the snake either coils up
and hides its head within its coiled up body; or it quickly
tries to escape to a safe hiding place inside the leaf litter.
When the snake coils, it enlarges its body and displays its
vivid skin colours (white, pink and brown), which is visi-
ble between the scales around the mid body. We observed,
on a number of occasions, the snake practicing thanatosis
(death mimicry) for up to 10–15 minutes after carrying out
our own handling attempts. Once the snake had noticed
that threat had disappeared, it quickly escaped and hid it-
self in the leaf litter. We have observed this species living
in sympatry with other snakes of several families such as
Aspidura guentheri Ferguson, 1876 (Natricidae); Hypnale
zara (Gray, 1849) (Viperidae); and Sibynophis subpuncta-
tus (Duméril, Bibron & Duméril, 1854) (Colubridae).
Based on our observations, its diet consists mostly of
lizards (saurophagy) and small snakes eggs (oophagy),
small spiders, beetles, other insects and the larvae of
other invertebrates. More specically, we observed the
snake feeding on ground dwelling skinks (Lankascincus
sp.) and geckos (Hemidactylus frenatus and Cnemaspis
sp.). If the prey is large, the snake wraps itself around
it and squeezes it until it suffocates. In captivity, it was
fed with jumping spiders, small wild cockroaches, anne-
lid worms, meal-worms, small frogs, and the freshly de-
tached tail tips of geckos.
During the breeding season (May–June) 3–5 individ-
uals can be observed close by and we observed sever-
al copulations in the evenings just after dark (18.0–19.0
hrs). The species lays 3–5 eggs at a time on dry, cool,
loose soil or under decaying logs on the ground (soil tem-
perature 26.2–27.9 °C; humidity 58–73%; light intensity
0–27 lux, based on observations of 10 ovipositions). Eggs
are cream in colour and oval in shape (12–14 mm long
and 4–5 mm wide, n = 40). The lectotype MNHN 3238 is
a gravid female with three eggs in its genital tract. The in-
cubation period is 38–45 days (based on observations of
10 incubating clutches). We did not see the parents close
by during the incubation nor shortly afterwards, indicat-
ing the lack of parental care of the eggs or hatchlings. The
new born juveniles were 4–5 cm in total length and their
body colour varied from dark brown to black. We noticed
that ants were their main egg predators on about ten occa-
sions. We also observed on several occasions, this snake
attempting to avoid ant-nests when moving or resting.
We have found this species inside termite mounds
on many occasions, an observation also made by Smith
(1943). This may indicate either a strategy used by the
snake to avoid ants (because we never observed ant nests
in or around termite mounts) or a neat way for the snake
to have instant access to food (may be feeding on termite
eggs). Further studies on habitat ecology would be inter-
esting. Even though this is a ground dwelling species, we
observed it climbing on rock boulders which have crevic-
es, indicating that this snake may be searching for geckos
or their eggs for food. During oods, the snake is usually
found off the oor, in trees at 1–2 m above ground lev-
el. It is also found deep inside forests, and has been ob-
served under old coconut harnesses, decaying logs on the
ground, and inside termite mounds (as mentioned earlier)
set in well maintained home gardens.
Zoosyst. Evol. 91 (1) 2015, 71–80
zse.pensoft.net
77
Road kills are identied as a major growing threat in
addition to forest fragmentation and habitat loss. People
are also a threat, killing the snake out of fear, believing
that it to be venomous, especially because as it displays
such vivid head and body colours. We observed nat-
ural predators including birds: the yellow-billed bab-
bler [Turdoides afnis (Jerdon, 1845)], southern coucal
(Centropus parroti Stresemann, 1913), common mynah
[Acridotheres tristis (Linnaeus, 1766)], white-throat-
ed kingsher [Halcyon smyrnensis (Linnaeus, 1758)],
and the Sri Lankan grey hornbill (Ocyceros gingalensis
Shaw, 1811); ophiophagous snakes including: two el-
apids, the Sri Lankan krait (Bungarus ceylonicus Gün-
ther, 1864), and the Indian krait (Bungarus caeruleus
Schneider, 1801); and amphibians including forest toads
(Duttaphrynus sp.). In addition, Karunarathna and Asela
(2007), and Karunarathna (2009) have observed the com-
mon rat snake (Ptyas mucosus Linnaeus, 1758) feeding
on Oligodon sublineatus and Oligodon calamarius (Lin-
naeus, 1758) in Sri Lanka
Discussion
In the description of Oligodon sublineatus, Duméril et
al. (1854) clearly states the following “We only have ob-
Figure 3. Distribution pattern of Oligodon sublineatus: blue dots, previous records and red dots, our observations; major towns are
displayed on the map (map source: Cooray 1967).
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Amarasinghe, A.A.T.: Oligodon sublineatus, an endemic snake of Sri Lanka
78
served one specimen well preserved …..”, they further
stated “We counted 15 scale rows on that specimen, 155
ventrals and 25 subcaudals”. Those counts are in accor-
dance with MNHN 3238 (respectively 150 and 28) hence;
we hereby designate it as the lectotype. However, the
measurements given in the last line “Total length was 180
cm; among them 155 for SVL and 25 for the tail.” is a
mistake; we believe that the wrong units of measurements
were chosen in error; it should have been in millimeters
and not centimeters! In addition, the newly recognized
syntype (MNHN 3238) had a total length of about 289
mm with 254 mm SVL and 35 mm for the tail. Again
we are making the assumption that Duméril et al. (1854)
must have mistakenly typed the total length as “180cm”
instead of ~280 mm and “SVL 155cm” instead of ~255
mm (typing a ‘1’ instead of a ‘2’). Interestingly, the syn-
type MNHN 3239 (now paralectotype) measured 177
mm total length with SVL 150 mm but its ventral and
subcaudal counts do not match those of the original de-
scription (respectively 138 and 36 [typical of a male] ver-
sus 155 and 25 in the description [typical of a female]).
However, the most probable explanation of this is that
they mistyped, rather than used (which may seem the ob-
vious explanation here) the measurement of the second
specimen, because Duméril et al. (1854) clearly stated
that they had examined only one specimen (the largest
of both syntypes), even though they compared the colour
patterns of both specimens, thus both are here considered
as syntypes. Furthermore, the scale counts in the smaller
specimen (MNHN 3239) do not match the original scale
description of Duméril et al. (1854), and the spots of the
larger specimen (MNHN 3238) are much more narrowed
towards the middle of the body compared to the spots of
the smaller specimen (MNHN 3239), which is in accor-
dance with the details of the examined specimen in the
original description. Thus Duméril et al. (1854) made a
mistake when describing characteristics of the examined
specimens and their ventral and subcaudal counts also
reect the classical mistake often seen when one single,
old and very small specimen is examined by many differ-
ent researchers over time. We have no doubt that MNHN
3238 is the Philippines (in error) specimen of the origi-
nal description and MNHN 3239 the Ceylan specimen of
Leschenault as reported in the original description, both
being the only two syntypes of O. sublineatus.
To be sure that there are no other specimens which
could possibly be a syntype, we examined all the availa-
ble Oligodon sublineatus specimens and all the specimens
of Oligodon collected from Sri Lanka and the Philippines
which were registered on or before 1864 in the MNHN
collection (1864 is the date of the oldest handwritten reg-
ister available for the MNHN herpetological collections).
Among the available specimens (except MNHN 3238–
39), MNHN 0611, 3537, 5768, 1900.0381–385, and
1900.0381, 1900.0381A–B are O. ancorus from the Phil-
ippines, MNHN 1012 is O. modestus from the Philippines
and MNHN 6129 is O. calamarius from Sri Lanka. In the
1864 unpublished catalogue, there were ve specimens of
O. sublineatus registered. However out of the three oth-
er available specimens of O. sublineatus (MNHN 1747,
4234, and 4234A) two do not match the original descrip-
tion. The third specimen MNHN 4234 closely matches
the original description, however we believe that it does
not qualify as the larger syntype based on the following
statement from the original description: “Another speci-
men, obviously younger, …..” which clearly means that
the former specimen is considerably larger (being older
by inference, or fully grown), than the latter. The spec-
imen MNHN 4234 is slightly smaller, than the younger
specimen (MNHN 3239) of Leschenault. It also has a
denite collector’s name (Janssen), and a denite locali-
ty (Sri Lanka) associated with it. Therefore, we can con-
dently reject this specimen MNHN 4234 as a syntype
and can conrm MNHN 3238-39 as the syntypes of O.
sublineatus. The location “Philippine” was never includ-
ed in any of the MNHN registers for O. sublineatus, not
even in the 1864 register, so it could be that subsequent
curators have corrected the error adding “Java, collector
Bosc” in the register (which is also wrong). These were
probably changed sometime between 1854 and 1864.
Therefore, here we correct the locality of the designated
lectotype as “Sri Lanka”.
Louis-Augustin-Guillaume Bosc (1759–1828) was a
French botanist, zoologist, and politician during the Rev-
olution, who described many amphibians and reptiles of
eastern North America. He also described the Savanna
Monitor (Varanus exanthematicus Bosc, 1792) from Sen-
egal in West Africa (Adler et al. 2012). Bosc never visit-
ed Sri Lanka. As MNHN 3238 belongs to an endemic Sri
Lankan species with incorrect collection locality details
(Philippines or even Java), we also agree that the collec-
tors name, Bosc, is a mistake and we follow Duméril et
al. (1854) and believe that the collector details for this
specimen is not known.
Duméril et al. (1854), in the original description states
that “on the jar there was no other indication that the
[data] one on its origin, Philippines and the name Oli-
godon torquatus with the letter “R””; no such indication
was found on any jar that we examined in MNHN col-
lections. It is strange that O. torquatus was mentioned in
the original description, a species which was described by
Boulenger in 1888, 34 years after O. sublineatus. How-
ever, based on the characters mentioned in the original
description of O. sublineatus, we can be sure that both
examined syntypes of Duméril et al. (1854) belong to the
same endemic species of Sri Lanka, O. sublineatus, and
neither match any of the species from the Philippines (see
Leviton 1963). This is also conrmed by the existence
of ventral scales having a series of dark brown spots ar-
ranged in three lines in the syntypes of O. sublineatus [vs.
uniform in O. ancorus (Girard, 1858) and O. notospilus
Günther, 1873], and 25 subcaudals in our lectotype [vs.
28–49 in O. meyerinkii (Steindachner, 1891); 52–54 in
O. maculatus (Taylor, 1918); 27–44 in O. modestus Gün-
ther, 1864; 48–60 in O. octolineatus (Schneider, 1801),
and 34–45 in O. perkinsi (Taylor, 1925); scale counts
Zoosyst. Evol. 91 (1) 2015, 71–80
zse.pensoft.net
79
after Leviton (1963)]. Furthermore, based on the descrip-
tion, it can also be easily distinguished from O. torqua-
tus (Boulenger, 1888) by having ventrals with a series of
dark brown spots in three lines (vs. uniform).
Boulenger (1894) recorded Oligodon sublineatus
from Nicobar Island. Deepak and Harikrishnan (2013)
observed a couple of specimens (ZSI 8899 and 8900) of
O. sublineatus deposited at ZSI-Kolkata, which were la-
belled as “Camorta, Nicobars”. They conrmed that both
the collection locality and the identity were wrong. The
species is denitively absent from Nicobar Islands, as
originally stated by Wall (1921) and has to be considered
as a species strictly endemic to Sri Lanka, but widespread
over the forested areas of the country.
Acknowledgments
We are very grateful to Gernot Vogel, A. de Silva, P. Uetz,
W.M.S. Botejue, D.E. Gabadage, N. Gamage, M.M. Ba-
hir, M.C. De Silva, H. Jayasinghe, V. Weeratunga, M.
Madawala, S. Fernando, S. Wellapulliarachchi, P. Sama-
rawickrama, M. Wickramasighe, D. Kandambi, P. Pieris,
and G.N. Karunarathna who provided valuable support
which enabled us to complete this study. We also thank
N. Wickramasinghe, S. Kasthuriarachchi, M. Nandase-
na, C. Munasinghe, L. Somaratne, R. Dasanayake, M.B.
Vaidyasekara, C. Kotalawala, A. Samanthika, P. Gunasiri
and R. Wickramanayake at National Museum of Sri Lan-
ka (NMSL) for assisting us while examining collections
under their care. Special thanks to D. Vallan for providing
information linked to the specimens at NMB. We thank
J. Supriatna and the staff of the Research Center for Cli-
mate Change of the University of Indonesia (RCCC-UI)
for their support. Finally, we thank the eld staff of Forest
Department and Department of Wildlife Conservation,
Sri Lanka.
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Appendix 1
Other materials examined
O. ancorus: Philippines: MNHN 0611, 3537, 5768,
1900.0381–385, 1900.0381A–B.
O. calamarius: Sri Lanka: MNHN 6129.
O. modestus: Philippines: MNHN 1012, BMNH
1946.1.5.54 (type).
O. sublineatus: Java (in error): MNHN 1747; Sri Lan-
ka: MNHN 4234, 4234A; NMB 21366–7, 1595–9,
10775, 21364–5; BMNH 1841.1.7.12, 1858.10.19.29,
1858.10.19.37–38, 1852.9.13.41, 1845.8.7.6,
1846.12.2, 1853.3.30.53, 1852.2.19.90, 1852.2.19.95,
1969.2769–2771, 1890.11.8.21–22, 1895.7.23.29,
1897.10.20.14, 1915.5.3.6, 1920.5.6.3, 1951.1.8.36,
1955.1.9.81–82, 1862.8.14.31, 1968.875.
Oligodon torquatus: Burma: BMNH 1946.1.4.21–22
(syntypes).
... As this species complex is sexually dimorphic, some of the meristic characters (e.g., ventrals, subcaudals) have to be assessed separately for each sex. It is evident that another Sri Lankan species, O. sublineatus Duméril, Bibron, and Duméril 1854 shows a nonoverlapping distinct range of subcaudals between males and females (Amarasinghe et al. 2015). However, O. arnensis does not show such variation in subcaudals, but indicates dimorphism in ventral count and morphometrics, and it may be the case that it is the same for the other two members of this complex (O. ...
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We review the taxonomic status of Oligodon waandersi sensu lato after examining all the name-bearing types (including synonyms) and morphological evidence. Oligodon waandersi sensu stricto is widely distributed (up to 1200 m above sea level) throughout the southern, some parts of the central, and the northern slopes of Central Sulawesi Island, Indonesia, whereas a similar species, O. propinquus, is only known from its holotype and considered to have originated from Java. Here, we demonstrate that O. propinquus is a close match with the northern population (North and Gorontalo) of O. waandersi and morphologically distinct; hence, we consider the type locality of O. propinquus as North Sulawesi, not Java. Oligodon taeniurus, long considered a junior synonym of O. waandersi, is a distinct species, but here we synonymize it with the morphologically closely matched O. propinquus, which has priority over O. taeniurus. Oligodon waandersi in Southeast Sulawesi (including some populations of Buton Islet) is morphologically distinct from O. waandersi sensu stricto in South Sulawesi; hence, it requires a new name. The new species is distinguished from congeners by having the following combination of characters: maximum snout–vent length of 340 mm, a single postocular, a single cloacal plate, a completely divided nasal, ventrals 150–169, subcaudals 18–26, temporals 1þ2, six supralabials with third and fourth in contact with eye, dorsal scale rows 15-15-15, hemipenes not forked and covered with spines, maxillary teeth 6–7, shorter tail (6.8–11.2% of total length), brownish dorsum with few dark-edged spots on the vertebral line anteriorly, reddish brown vertebral line on the posterior body and tail, mostly a blackish brown blotch below the eye, and whitish collar band interrupted middorsally. We provide a complete redescription for O. waandersi and O. propinquus based on respective holotypes deposited at the Natural History Museum London and the Zoologisches Museum Hamburg.