Journal of Threatened
Taxa | www.threatenedtaxa.org | 26 April 2025 | 17(4): 26803–26813
ISSN 0974-7907 (Online) | ISSN 0974-7893 (Print)
https://doi.org/10.11609/jott.9436.17.4.26803-26813
#9436 | Received 26 September 2024 | Final received 26 February 2025 |
Finally accepted 06 March 2025
Comments on the systematics and
morphology of Smithophis bicolor (Blyth, 1855) (Reptilia: Squamata:
Natricidae) based on topotypical specimens from Meghalaya, India
Jayaditya Purkayastha 1,
Bipin Meetei Asem 2, Hmar Tlawmte Lalremsanga 3,
Madhurima Das 4 ,
Holiness Warjri 5,
Goldenstar Thongni 6 &
Sanath Chandra Bohra 7
1,4,7 Help Earth, 16, Raghunath
Choudhury Path, Lachitnagar, Guwahati, Assam 781007, India.
2 Zoological Survey of India,
Northeastern Regional Centre, Shillong, Meghalaya 793014, India.
3 Developmental Biology and
Herpetology Laboratory, Department of Zoology, Mizoram University, Tanhril,
Mizoram 796004, India.
4,5 Department of Zoology, Assam Don
Bosco University, Sonapur, Assam 781027, India.
6 Rohbah, South West Khasi Hills
District, Mawkyrwat, Meghalaya 793114, India.
1 mail.jayaditya@gmail.com, 2 abmeetei@gmail.com,
3 htlrsa@yahoo.co.in, 4 madhurima.das@dbuniversity.ac.in,
5 holiness12345@gmail.com, 6 golden2394@gmail.com, 7
sreptilian6@gmail.com (corresponding author)
Editor: S.R. Ganesh, Kalinga Foundation, Agumbe,
India. Date of publication: 26 April 2025
(online & print)
Citation:
Purkayastha, J., B.M. Asem, H.T. Lalremsanga, M. Das, H. Warjri, G. Thongni
& S.C. Bohra (2025). Comments on the systematics and morphology of Smithophis
bicolor (Blyth, 1855) (Reptilia: Squamata: Natricidae) based on topotypical specimens from
Meghalaya, India. Journal of Threatened Taxa 17(4): 26803–26813. https://doi.org/10.11609/jott.9436.17.4.26803-26813
Copyright: © Purkayastha et al. 2025. Creative Commons Attribution 4.0 International License.
JoTT allows unrestricted use, reproduction, and distribution of this article in
any medium by providing adequate credit to the author(s) and the source of
publication.
Funding:
Rufford Small Grants for Nature Conservation.
Competing interests:
The authors declare that there are no conflicts of interest regarding the
publication of this paper. All authors have participated in the research,
design and drafting of this manuscript, and have reviewed and approved the
final version. They have no financial or personal relationships that could
inappropriately influence or bias the content of this work.
Author details: Jayaditya Purkayastha is a wildlife biologist
at Help Earth NGO. Bipin Meetei Asem
is an assistant zoologist at Zoological Survey of India, North Eastern Regional
Centre. Hmar Tlawmte Lalremsanga is
a professor at the department of Zoology, Mizoram University. Madhurima Das is an assistant professor
at the department of Zoology, Assam Don Bosco University. Holiness Warjri is a
research scholar at the department of Zoology, Assam Don Bosco University. Goldenstar Thongni is a naturalist with
interest in biodiversity of Meghalaya.
Sanath Chandra Bohra is a wildlife biologist at Help Earth.
Author contributions: JP—conceptualization
and paper writing; BMA—provided morphological data; HTL—provided morphological
data and helped in paper writing; MD—molecular analysis; HW—collection of
specimens and field study; GT—collection of specimens and field study;
SCB—paper writing and securing funds.
Acknowledgements: We
would like to thank chief wildlife warden (Environment, Forest and Climate
Change) and Government of Meghalaya (Permit No.FOR.7/2021/216) for issuing the
collection permit. We are also very thankful to the local community for
supporting us in our field surveys. We are also thankful to MegLIFE and Rufford
Small Grants Foundation for supporting this work. Finally we would like to
thank the reviewers and the editor for substantially improving the manuscript
through their inputs.
Abstract: The Natricid Snake Smithophis
bicolor has been poorly studied and understood since its initial
description in the mid-1800s. This study presents a detailed morphological
description of S. bicolor based on five examined specimens including a
freshly collected mutilated specimen from Meghalaya State, India. For the first
time, genetic data has been obtained and analysed, providing novel insights
into the species’ phylogenetic position within its genus Smithophis.
Morphological comparisons and molecular analyses using cytochrome b
mitochondrial gene reveal subtle variations that contribute to a clearer
understanding of the species’ taxonomic identity. This study allows us to
conclude that currently S. bicolor is restricted to the higher
elevations of Meghalaya indicating that literature records mentioning the
presence of this species from Mizoram actually represents a distinct, new
unnamed lineage.
Keywords: Cytochrome b, hemipenis,
herpetofauna, Khasi hills, Mairang, northeastern india, phylogeny,
re-description, shillong plateau, taxonomy, type locality.
Introduction
Blyth (1855) described Smithophis
bicolor in the genus Calamaria Boie, 1827 based on a single specimen
(holotype ZSI 7030) provided by Mr. Robinson with the type locality recorded as
Assam, India. In the same piece of work, Blyth (1855: p. 287) mentioned that
the specimens provided by Mr. Robinson might be from Khasya Hills (present day
Khasi Hills, Meghalaya State, India) or other upland territories. The holotype
ZSI 7030 was mentioned to be with a dusky-plumbeous dorsal surface and a buffy
white ventral surface, 17 mid-dorsal scale rows, 210 ventrals and 75 pairs of
subcaudals, snout-vent length of 495 mm and a tail length of 111 mm.
Boulenger (1893) allocated bicolor to the genus Rhabdops
Boulenger and stated its distribution from Khasi Hills, Assam and Yunnan. Rhabdops
olivaceus (Beddome 1863) with distribution in the Western Ghats, India is
the type species of Rhabdops along with the recently described Rhabdops
aquaticus Giri et al. 2017 also known from the Western Ghats, India
whereas Rhabdops bicolor was the only species known from northeastern
India. Based on genetics and disjunct distribution in Western Ghats and
northeastern side, Giri et al. (2019) erected the genus Smithophis to
accommodate northeastern populations (bicolor) and also described a new
species in the genus namely Smithophis atemporalis noting that the genus
Smithophis is more closely related to Opisthotropis and Sinonatrix
than to Rhabdops. Furthermore, Giri et al. (2019) in their genetic
assessment of Smithophis bicolor used data based on materials from
Mizoram State, northeastern India. Since then, three more species were added to
the genus, namely, Smithophis arunachalensis Das et al., 2020; S.
linearis Vogel et al., 2020 and S. mizoramensis Mirza et al.,
2024 (Mirza et al. 2024a). Chandramouli et al. (2021) provided a morphological
description of an adult male topotypic specimen of S. bicolor from
Shillong, in the Khasi Hills. Giri et al. (2019) in their genetic assessment of
S. bicolor used data based on materials from Mizoram, India but missed
out the typical population from the Khasi Hills of Meghalaya. In this study,
based on fresh collection and museum specimens (Zoological Survey of India,
Northeastern Regional Centre, Shillong, Meghalaya State, India) we provide
additional morphological and molecular insights for Smithophis bicolor
from its type locality and point to the possibility of existence of an
undescribed Smithophis from Mizoram State, India, treated as Smithophis
cf. bicolor by Giri et al. (2019).
Materials
and Methods
The study was conducted in Khasi
hills of Meghalaya and throughout Mizoram after obtaining permission for
collection within both the states from the chief wildlife warden of
Environment, Forests and Climate Change Department, Government of Meghalaya and
Mizoram (No. FOR./7/2021/216 for Meghalaya and A.33011/2/99-CWLW/225 for
Mizoram). A road killed male specimen of Smithophis bicolor was
collected from Mairang (ADBU-HN/HW0210), Eastern West Khasi Hills district
(25.559°N, 91.635°E; altitude 1,564 m; see Image 1), Meghalaya, India by
Holiness Warjri on 12 August 2024 at around 1230 h and has been deposited
in the herpetological museum maintained by the Department of Zoology, Assam Don
Bosco University, Sonapur, Assam, India. Furthermore, four samples
(V1/ERS/ZSI-444, V1/ERS/ZSI-2592, V1/ERS/ZSI-3052 from Shillong, Meghalaya and
VR/ERS/ZSI-725 from Umiam, Meghalaya) were also studied from the collection of
Zoological Survey of India, Eastern Regional Station, Shillong, Meghalaya,
India. Genomic DNA was extracted from ethanol (100%) preserved liver tissue of
the specimen ADBU-HN/HW0210 using tissue kit (Qiagen) following manufacturer’s
instructions. Partial sequence of the mitochondrial cytochrome b gene was
generated using the primer pair L14910 (GACCTGTGATMTGAAAAACCAYCGTTGT), H16064
(CTTTGGTTTACAAGAACAATGCTTTA)
(Burbrink et al. 2000). Sequence chromatograms were quality checked, edited and
assembled into contigs using Chromas and Sequence Scanner v1.0 (Applied
Biosystems). Comparative cytochrome b (Table 1) sequences were used following
Mirza et al. (2024) and were obtained from GenBank. Sequence alignment was done
using MUSCLE (Edgar 2004) in MEGA7 (Tamura & Nei 1993; Kumar et al. 2016)
with default parameter settings. Maximum likelihood (ML) phylogenetic tree was
reconstructed using un-partitioned dataset in IQ-TREE (Nguyen et al. 2015) with
the substitution model GTR+F+I+G4 selected based on the BIC scores by
ModelFinder (Kalyaanamoorthy et al. 2017) implemented in the IQ-TREE (Nguyen et
al. 2015). The ML analysis was run with an ultrafast bootstrap option (Minh et
al. 2013) for 1000 iterations to assess clade support. The un-corrected
pairwise p-distance was calculated in MEGA7 (Kumar et al. 2016) with pairwise
deletions of missing data and gaps.
Morphometrics and meristics are
taken as follows: HL (Head length taken from the retro articular process of the
jaw to the tip of the snout), HW (Head width measured at the widest point of
the head), HD (Head depth measured at the greatest depth of the head), ED (Eye
diameter taken horizontally at the outer margins of the orbit), ND (Greatest
nostril diameter), NS (Nostril to snout distance taken from the anterior
portion of nostril to the tip of the snout), EN (Eye to nostril distance taken
from the anterior portion of orbit to the posterior margin of nostril
openings), IND (Internarial distance measured between the two nostril
openings), IOD (Inter orbital distance measured between the minimum distance of
two outer orbital margins), PL (Greatest length of the parietal scale), SVL
(Snout-vent length measured from the tip of the snout to the vent ventrally),
TaL (Tail length measured from the vent to the tip of the tail ventrally), TL
(Total length measured from the tip of the snout to the to the tip of the tail ventrally
or dorsally), DSR (Dorsal scale rows counted in a row transversely in three
sections, i.e., exactly one head length behind the neck, exactly in the midbody
and exactly one head length before the vent dorsally), V (Number of ventral
scales counted ventrally from the 1st scale just behind pre-ventrals
till the last scale just before the anal plate), SC (Number of subcaudal scales
beginning just behind the cloaca ventrally till the last scale in the tail
excluding the terminal scute), SL (Number of supralabial scales counted on the
upper lips), IL (Number of infralabial scales counted on the lower lips), COS
(Number of circumorbital scales surrounding the orbit), Temp (Number of
anterior plus posterior temporal scales in the lateral portion of the head),
and Anal (Condition of the anal plate whether its divided or single).
Results
Based on Cyt b (1109 bp) data
generated in this study, the Smithophis bicolor from Mairang, Eastern
west Khasi Hills, Meghalaya State was seen to show a sister relationship to S.
mizoramensis differing in a low to moderate uncorrected p-distance of 4%
whereas S. cf. bicolor from Mizoram State (Giri et al. 2019) was
seen to be sister to the clade comprising S. bicolor (Mairang) + S.
mizoramensis. Smithophis cf. bicolor from Mizoram differs
by an uncorrected p-distance of at least 11.5% from the S. bicolor s.
str. of Mairang, Eastern West Khasi Hills, Meghalaya State (nearly thrice as
high as the divergence between S. bicolor s. str. and S. mizoramensis)
(Figure 1, Table 2). Genetic divergence for the cytochrome b (cyt b) gene
reveals that true S. bicolor is 4% to 14.3% divergent from other Smithophis
species. Morphologically, the species S. bicolor differs from its sister
S. mizoramensis mainly in body coloration as well as carination
condition of the sacral scales.
Morphology of topotypical,
Meghalaya specimens
(V1/ERS/ZSI-2592, V1/ERS/ZSI-725,
V1/ERS/ZSI-3052, V1/ERS/ZSI-444, ADBU-HN/HW0210; Images 2–4): Smithophis
bicolor is mainly characterized by having 17 smooth dorsal scale rows
across the body without having any keeled sacral scales in males. It possesses
temporal scales (1+1) and features 5 circum-orbital scales along with a single
internasal and a single prefrontal scale. Tail is relatively short and ends
with a short spine like scale. The examined specimens from Meghalaya exhibit an
elongated body with a head slightly distinct from neck, little longer than
broad (HW/HL 0.58–0.75), slightly flattened dorsoventrally (HL/HD 0.41–0.57).
The eyes are moderately sized (ED/HL 0.12–0.19); nostrils are placed closer to
snout than to the eye (NS/EN 0.69–0.88), relatively small in size (ND/ED
0.50–0.71); a pair of parietals, each parietal scale is relatively large, half
or slightly more than half of head length (Parietal/HL 0.50–0.61); SVL ranges
from 360–535 mm; TaL ranges from 110–160 mm; tail relatively short (TaL/SVL
0.30–0.44; TaL/TL 0.22–0.25); TL ranges from 470–690 mm. The meristic characteristics
of the specimens are consistent across the examined specimens. All individuals
possess 17–17–17 dorsal scale rows; Ventrals 209–212 in males and 192 in
females; subcaudals 75–80 in males and 68 in females; SL usually 5 and IL
usually 6 or 7. One unsexed individual has 212 ventrals and 71 subcaudals, SL 5
on each side and IL 6 on each side. The circum-orbital scale count is uniform
across all specimens, with five scales surrounding each eye. The temporal
scales are arranged in a 1+1 pattern in all individuals and anal plate is
divided in all specimens.
Sexual dimorphism
Based on our data, males of this
species have ventrals ranging from 209 to 212 (versus 192 in a female), Sc
ranging from 75 to 80 (versus 68 in a female) and Tal/TL ratio of 0.23
to 0.25 (versus 0.22 in a female).
Coloration in preservative (Image 2)
The specimens in ZSI, especially
V1/ERS/ZSI 444 and 3052, have become completely faded in comparison to their
coloration in life (read below), such that there is barely much difference between
their dorsal and ventral colours, indicating bleaching of dorsal colour. Eyes
became paler and pupil became light grey.
Coloration in life (based on live uncollected
snakes; Image 3)
Smithophis bicolor we encountered had a dorsal
region with uniformly shiny black above with a slightly violet tinge, whereas
the ventral is uniformly (mostly) gambodge yellow in life. In the posterior
half of the lateral region, a prominent, broad gambodge yellow stripe runs
along the body, covering the ventrals. The dorsal and ventral coloration meet
somewhat at the middle of the lateral region, giving it a typical bicoloured
appearance which begins from the snout and extends up to the tip of the tail in
life.
Hemipenial description (based on ADBU-HN/HW0210; Image
4)
Short and stout, characterized by
heavy spinosity over the distal three-quarters, with the proximal one-quarter,
near the base, exhibiting a somewhat calyculate texture. The spines are more
pronounced and larger towards the intermediate region and the base, gradually
becoming smaller and conspicuous, somewhat hair or serrations like towards the
tip. The spines in hemipenis of Smithophis bicolor are much more
pronounced than its sister taxa S. mizoramensis.
Comparisons
Smithophis bicolor has well defined temporal scales
(1+1) vs. absent in S. atemporalis; Smithophis bicolor does
not have keeled sacral scales in males vs. presence of keeled sacral scales in S.
arunachalensis and S. mizoramensis. The dorsum is immaculate in Smithophis
bicolor vs. either blotched or striped in all the other congeners.
Furthermore, Smithophis bicolor has 4–5 circumorbital scales vs. 6–7 in S.
linearis.
Natural history and distribution (Image 5)
Based on our current data, Smithophis
bicolor is a terrestrial to semi-aquatic species which mainly prefers to
live in close proximity to streams and other water bodies surrounded by
semi-evergreen, evergreen, moist deciduous, montane forests and at times even
moderately disturbed hilly areas. Based on the specimens, the species is found
in areas of higher elevation ranging 1,000–1,700 m. All the freshly observed
specimens from Meghalaya State were seen near streams, boulders with or without
mosses, loose soils or roadside edges with or without mosses within wet montane
or wet semi-evergreen or evergreen forests, particularly during the day. It has
been observed that this species (just like other congeners) is often seen or
becomes active during heavy rainfall. The current understanding based on the
available data allows us to conclude that Smithophis bicolor is
currently restricted to the higher elevations of Meghalaya State, particularly
in the Khasi Hills and its adjacent areas, i.e., Garo hills whereas those
reported as S. bicolor s. lat. from regions outside Meghalaya State
might represent a different species.
Discussion
Despite being one of the earliest
described species of the genus Smithophis, S. bicolor lacked any
new information until recent times (Chandramouli et al. 2021). This study not
only gives genetic data from the type locality of S. bicolor but also
provides expanded morphological description of the species based on multiple
specimens. The examination of additional topotypic specimens of S. bicolor
from Meghalaya resulting in this expanded morphological description makes the
species better characterised in terms of its morphology. This provides a
concrete dataset for researchers to describe as well as compare new and
previously described species with the nominal taxon in the future. Furthermore,
the population previously identified as S. bicolor from Mizoram (Giri et
al. 2019) which represents a genetically and morphologically distinct lineage,
will be formally described as a new species in our forthcoming publication in
preparation.
Meghalaya State, which is rich in
terms of biodiversity forms a part of the Indo-Burma biodiversity hotspot which
lies on the southern bank of river Brahmaputra and has been relatively well
surveyed historically, particularly during the time of British India. Since
then, after a long time the state has witnessed a significant increase in the
number of new herpetofaunal species descriptions in the past two decades
alongside range extensions regarding certain species (e.g., Das et al. 2010;
Mahony et al. 2011, 2013, 2018, 2020; Purkayastha & Matsui 2012; Datta-Roy
et al. 2013; Kamei et al. 2013; Agarwal et al. 2018b; Purkayastha et al.
2020a,b; 2021; 2022; Rathee et al. 2022; Mirza et al. 2024b). Despite being a
described species, further research and field work are necessary to properly
understand the accurate distribution range, ecology, reproductive biology and
conservation biology of S. bicolor. The Khasi Hill is the “type
locality” for snake species like Stoliczkia khasiensis Jerdon, 1870
which is known just from a single specimen and since its description in 1870,
the species was never reported for the second time as a result of which it has
been considered as a lost species. This highlights the importance of conducting
continuous as well as comprehensive herpetofaunal surveys throughout Khasi
Hills and its adjoining hill ranges before the depletion of forest cover due to
anthropogenic pressures.
We suggest further field
investigations involving wide samplings throughout the given range of the genus
Smithophis to understand the potential reason behind the diversification
of these natricids that will definitely help to uncover further new unnamed
populations in the region.
Table 1. List of cytochrome b sequences used in this study. New sequence
entry in bold.
|
Species |
Voucher |
Genbank accession number |
Location |
|
Opisthotropis andersonii |
SYS r001383 |
KY594735 |
Mt. Maofeng, Guangzhou,
Guangdong, China |
|
Opisthotropis cheni |
YBU 071040 (=GP383) |
GQ281779 |
Nanling NR, Guangdong, China |
|
Opisthotropis daovantieni |
FMNH 252009 |
OK315831 |
An Ke, Gia Lai, Vietnam |
|
Opisthotropis durandi |
NCSM 80739 |
MK941137 |
Nam Lan, Phosaly, Laos |
|
Opisthotropis guangxiensis |
GP746 |
GQ281776 |
Guangxi, China |
|
Opisthotropis haihaensis |
SYS r000537 |
MN890017 |
Quang Ninh, Vietnam |
|
Opisthotropis hungtai 1 |
SYSr000538 |
MN890018 |
Mt. Wuhuang, Guangxi, China |
|
Opisthotropis hungtai 2 |
SYS r000946 |
KY594748 |
Heishiding NR, Fengkai,
Guangdong, China |
|
Opisthotropis jacobi |
ZFMK 100818 |
MG545602 |
Tam Dao NP, Vinh Phuc, Vietnam |
|
Opisthotropis kuatunensis |
SYS r000998 |
KY594745 |
Qixiling NR, Yongxin County,
Jiangxi, China |
|
Opisthotropis lateralis |
ZMMU NAP-08678 |
OK315832 |
Xuan Son NP, Phu Tho, Vietnam |
|
Opisthotropis latouchii |
GP647 |
GQ281783 |
Fujian, China |
|
Opisthotropis laui |
SYS r001161 |
KY594738 |
Shangchuan Í., Taishan,
Guangdong, China |
|
Opisthotropis maxwelli |
SYS r000841 |
KY594736 |
Nan'ao Is., Nan'ao, Guangdong,
China |
|
Opisthotropis shenzhenensis |
SYS r001032 |
KY594729 |
Mt. Tiantou, Shenzhen,
Guangdong, China |
|
Opisthotropis typica |
HT0794 |
LC325343 |
Malaysia |
|
Opisthotropis voquyi |
ZMMU R-16681 |
OK315833 |
Tay Yen Tu NR, Bac Giang,
Vietnam |
|
Opisthotropis zhaoermii |
CIB110000 |
MG012801 |
Guzhang, Hunan, China |
|
Smithophis atemporalis |
BNHS 2366 |
MK350262 |
Mizoram, India |
|
Smithophis cf. bicolor |
BNHS 2369 |
MK350261 |
Mizoram, India |
|
Smithophis cf. bicolor |
MZMU 1798 |
PP996090 |
Aizawl, Mizoram, India |
|
Smithophis bicolor |
ADBU-HN/ HW0210 |
PQ727125 |
Mairang, Eastern West Khasi
Hills, Meghalaya, India |
|
Smithophis linearis |
KIZ 059110 |
MT185677 |
Yunnan, China |
|
Smithophis mizoramensis |
BNHS 3766 |
PP996092 |
Suangpuilawn, Saitual, Mizoram,
India |
|
Smithophis mizoramensis |
BNHS 3767 |
PP996093 |
Suangpuilawn, Saitual, Mizoram,
India |
|
Smithophis mizoramensis |
BNHS 3768 |
PP996094 |
Suangpuilawn, Saitual, Mizoram,
India |
|
Smithophis mizoramensis |
MZMU 2602 |
PP996091 |
Suangpuilawn, Saitual, Mizoram,
India |
|
Trimerodytes percarinata |
ZMMU R-16444 |
OK315846 |
Pu Mat NP, Nghe An, Vietnam |
Table 2. Uncorrected p-distance between the members of the genus Smithophis.
|
PQ727125_S._bicolor |
|
|
|
|
|
|
|
|
|
PP996092_S. mizoramensis |
0.040 |
|
|
|
|
|
|
|
|
PP996093_S._mizoramensis |
0.040 |
0.000 |
|
|
|
|
|
|
|
PP996094_S._mizoramensis |
0.040 |
0.000 |
0.000 |
|
|
|
|
|
|
PP996091_S._mizoramensis |
0.040 |
0.000 |
0.000 |
0.000 |
|
|
|
|
|
MK350262_S._atemporalis |
0.107 |
0.096 |
0.096 |
0.096 |
0.096 |
|
|
|
|
MK350261_S._cf. bicolor |
0.115 |
0.098 |
0.098 |
0.098 |
0.098 |
0.125 |
|
|
|
PP996090_S._ cf. bicolor |
0.115 |
0.098 |
0.098 |
0.098 |
0.098 |
0.125 |
0.000 |
|
|
MT185677_Smithophis_linearis |
0.143 |
0.138 |
0.138 |
0.138 |
0.138 |
0.156 |
0.169 |
0.169 |
Table 3. Morphometric and meristic data of Smithophis bicolor
from eastern and western Khasi Hills, Meghalaya State, India (-- indicates
missing data). Measurements in mm.
|
Museum no. |
V1/ERS/ ZSI-2592 |
V1/ERS/ ZSI-725 |
V1/ERS/ ZSI-3052 |
V1/ERS/ ZSI-444 |
ADBU-HN/ HW0210 |
|
Location |
Fruit garden, Shillong |
Tripura Castle Road, Shillong |
Risa Colony, Shillong |
Umiam, Umsaw, NEPA Campus |
Mairang, Eastern West Khasi
hills |
|
Sex |
Unsexed |
Male |
Female |
Male |
Male |
|
HL |
12.54 |
10.9 |
14.74 |
9.8 |
11 |
|
HW |
8.28 |
7.52 |
9.79 |
7.48 |
7.5 |
|
HD |
7.09 |
5.8 |
7.51 |
5.5 |
5.1 |
|
ED |
1.8 |
1.9 |
1.51 |
1.62 |
1.7 |
|
ND |
1.27 |
0.9 |
0.9 |
1.18 |
0.9 |
|
NS |
1.94 |
1.6 |
1.9 |
1.43 |
1.8 |
|
EN |
2.17 |
1.87 |
2.6 |
2.15 |
2.2 |
|
IND |
3.48 |
3.4 |
3.45 |
3.2 |
3.4 |
|
IOD |
5.8 |
5.34 |
5.77 |
5.2 |
4.8 |
|
PL |
7.63 |
6 |
7.21 |
6.8 |
6.5 |
|
SVL |
520 |
385 |
535 |
360 |
360 |
|
TaL |
160 |
113 |
155 |
120 |
110 |
|
TL |
680 |
498 |
690 |
480 |
470 |
|
DSR |
17/17/17 |
17/17/17 |
17/17/17 |
17/17/17 |
17/17/17 |
|
V |
212 |
209+ |
192 |
212 |
209 |
|
SC |
71 |
75 |
68 |
80 |
76 |
|
SL (L/R) |
5/5 |
5/5 |
5/5 |
5/5 |
5/-- |
|
IL (L/R) |
6/6 |
6/6 |
6/6 |
7/6 |
-- |
|
COS |
5 |
5 |
5 |
5 |
5 |
|
Temp |
1+1 |
1+1 |
1+1 |
1+1 |
1+1 |
|
Anal |
Divided |
Divided |
Divided |
Divided |
Divided |
For
figures & images - - click here for full PDF
References
Agarwal, I.,
S. Mahony, V.B. Giri, R. Chaitanya & A.M. Bauer (2018). Six new Cyrtodactylus
(Squamata: Gekkonidae) from northeast India. Zootaxa 4524(5): 501–535. https://doi.org/10.11646/zootaxa.4524.5.1
Beddome, R.H.
(1863). Further
notes upon the snakes of the Madras Presidency; with descriptions of new
species. Madras Quarterly Journal of Medical Science 6: 41–48 (Reprint: Journal
of the Society for the Bibliography of Natural Science London, 1: 306–314,
1940).
Blyth, E.
(1855). Notices and
descriptions of various reptiles, new or little known (part 2). Journal of
Asiatic Society of Bengal, Calcutta 23(3): 287–302 (1854).
Boulenger,
G.A. (1893). Catalogue
of the Snakes in the British Museum. (Natural History), London (Taylor
& Francis), 448 pp.
Burbrink,
F.T., R. Lawson, & J.B. Slowinski (2000). Mitochondrial DNA
phylogeography of the polytypic North American Rat Snake Elaphe obsoleta:
a critique of the subspecies concept. Evolution 54(6): 2107–2118.
Chandramouli,
S.R., P. Karthik, R.S. Naveen, S. Babu, P.V. Karunakaran & H.N. Kumara
(2021). A
Two-colored Forestsnake, Smithophis
bicolor (Blyth 1855) (Reptilia: Natricidae), from the Khasi Hills, Meghalaya, India. Reptiles
& Amphibians 28(1): 24–25. https://doi.org/10.17161/randa.v28i1.15285
Das, A., V.
Deepak, A. Captain, E.O. Wade & D.J. Gower (2020). Description of a new species of
Smithophis Giri et al. 2019 (Serpentes: Colubridae: Natricinae) from
Arunachal Pradesh, India. Zootaxa 4860(2): 267–283. https://doi.org/10.11646/zootaxa.4860.2.8
Das, I.,
R.K.L. Tron, D. Rangad & R.N. Hooroo (2010). A new species of Leptolalax
(Anura: Megophryidae) from the sacred groves of Mawphlang, Meghalaya,
north-eastern India. Zootaxa 2339(1): 44–56. https://doi.org/10.11646/zootaxa.2339.1.2
Datta-Roy,
A., I. Das, A.M. Bauer, R.K.L. Tron & P. Karanth (2013). Lizard wears shades. A
spectacled Sphenomorphus (Squamata: Scincidae), from the sacred forests
of Mawphlang, Meghalaya, north-east India. Zootaxa 3701(2): 257–276. https://doi.org/10.11646/zootaxa.3701.2.7
Edgar, R.C.
(2004). MUSCLE:
multiple sequence alignment with high accuracy and high throughput. Nucleic
Acids Research 32: 1792–1797. https://doi.org/10.1093/nar/gkh340
Giri, V.B.,
R. Chaitanya, S. Mahony, S. Lalrounga, C. Lalrinchhana, A. Das, V. Sarkar, P.
Karanth & V. Deepak (2019). On the systematic status of the genus Oriocalotes
Günther, 1864 (Squamata: Agamidae: Draconinae) with the description of a new
species from Mizoram state, northeast India. Zootaxa 4638(4): 451–484. https://doi.org/10.11646/zootaxa.4638.4.1
Giri, V.B.,
V. Deepak, A. Captain, A. Das, S. Das, K.P. Rajkumar, R.L. Rathish & D.J.
Gower (2017). A new
species of Rhabdops Boulenger, 1893 (Serpentes: Natricinae) from the
northern Western Ghats region of India. Zootaxa 4319(1):
27–52. https://doi.org/10.11646/zootaxa.4319.1.2
Giri, V.B.,
D.J. Gower, A. Das, H.T. Lalremsanga, S. Lalronunga, A. Captain & V. Deepak
(2019). A new genus
and species of natricine snake from northeast India. Zootaxa 4603:
241–264. https://doi.org/10.11646/zootaxa.4603.2.2
Kalyaanamoorthy,
S., B.Q. Minh, T.K. Wong, A.V. Haeseler & L.S. Jermiin (2017). ModelFinder: fast model
selection for accurate phylogenetic estimates. Nature Methods 14:
587–589. https://doi.org/10.1038/nmeth.4285
Kamei, R.G.,
D.J. Gower, M. Wilkinson & S.D. Biju (2013). Systematics of the caecilian
family Chikilidae (Amphibia: Gymnophiona) with the description of three new
species of Chikila from northeast India. Zootaxa 3666(4):
401–435. http://doi.org/10.11646/zootaxa.3666.4.1
Kumar, S., G.
Stecher & K. Tamura (2016). MEGA7: molecular evolutionary genetics analysis
version 7.0 for bigger datasets. Molecular Biology and Evolution 33:
1870–1874. https://doi.org/10.1093/molbev/msw054
Lanfear, R.,
P.B. Frandsen, A.M. Wright, T. Senfeld & B. Calcott (2017). PartitionFinder 2: new methods
for selecting partitioned models of evolution for molecular and morphological
phylogenetic analyses. Molecular Biology and Evolution 34: 772–773. https://doi.org/10.1093/molbev/msw260
Mahony, S.,
R.G. Kamei, E.C. Teeling & S.D. Biju (2018). Cryptic diversity within the Megophrys
major species group (Amphibia: Megophryidae) of the Asian Horned Frogs:
phylogenetic perspectives and a taxonomic revision of south Asian taxa, with
descriptions of four new species. Zootaxa 4523(1): 1–96. https://doi.org/10.11646/zootaxa.4523.1.1
Mahony, S.,
R.G. Kamei, E.C. Teeling & S.D. Biju (2020). Taxonomic review of the Asian
horned frogs (Amphibia: Megophrys Kuhl & Van Hasselt) of northeast India
and Bangladesh previously misidentified as M. parva (Boulenger), with
descriptions of three new species. Journal of Natural History 54(1–4):
119–194. https://doi.org/10.1080/00222933.2020.1736679
Mahony, S.,
S. Sengupta, R.G. Kamei & S.D. Biju (2011). A new low altitude species of Megophrys
Kuhl and van Hasselt (Amphibia: Megophryidae), from Assam, northeast India. Zootaxa
3059: 36–46. https://doi.org/10.11646/zootaxa.3059.1.2
Mahony, S.,
E.C. Teeling & S.D. Biju (2013). Three new species of Horned
Frogs, Megophrys (Amphibia: Megophryidae), from northeast India, with a
resolution to the identity of Megophrys boettgeri populations reported
from the region. Zootaxa 3722(2): 143–169. https://doi.org/10.11646/zootaxa.3722.2.2
Minh B.Q.,
M.A.T. Nguyen & A.V. Haeseler (2013). Ultrafast approximation for
phylogenetic bootstrap. Molecular Biology and Evolution 30: 1188–1195. https://doi.org/10.1093/molbev/mst024
Mirza, Z.A.,
V.K. Bhardwaj, J.C. Lalmuanawma, G. Choure, H.T. Lalremsanga, M.
Vabeiryureilai, A. Captain, A. Zagade & H. Patel (2024a). A new species of Smithophis
Giri et al. 2019 from the Indo-Burma Region. Diversity 16(8):
480. https://doi.org/10.3390/d16080480
Mirza, Z.A.,
S. Pattekar, S. Verma, B.L. Stuart, J. Purkayastha, P.P. Mohapatra & H.
Patel (2024b). A new
Long-snouted Vine Snake species in the genus Ahaetulla Link, 1807 (Colubridae:
Chrysopeleinae) from India. Journal of Asia-Pacific Biodiversity 17(4):
696–703. https://doi.org/10.1016/j.japb.2024.04.017
Nguyen, L.T.,
H.A. Schmidt, A. Von Haeseler & B.Q. Minh (2015). IQ-TREE: a fast and effective
stochastic algorithm for estimating maximum likelihood phylogenies. Molecular
Biology and Evolution 32: 268–274. https://doi.org/10.1093/molbev/msu300
Purkayastha,
J. & M. Matsui (2012). A new species of Fejervarya (Anura: Dicroglossidae) from
Mawphlang, northeastern India. Asian Herpetological Research 3(1):
31–37. https://doi.org/10.3724/SP.J.1245.2012.00031
Purkayastha,
J., S.C. Bohra & M. Das (2020a). First record of the Urban
Bent-toed Gecko, Cyrtodactylus urbanus Purkayastha, Das, Bohra, Bauer,
and Agarwal 2018 (Squamata: Gekkonidae), from Meghalaya, India. ICRF Reptiles
& Amphibians Conservation and Natural History 27(3): 512–513. https://doi.org/10.17161/randa.v27i3.14901
Purkayastha,
J., M. Das, S.C. Bohra, A.M. Bauer & I. Agarwal (2020b). Another new Cyrtodactylus
(Squamata: Gekkonidae) from Guwahati, Assam, India. Zootaxa 4732(3):
375–392. https://doi.org/10.11646/zootaxa.4732.3.2
Purkayastha,
J., H.T. Lalremsanga, S.C. Bohra, L. Biakzuala, H. Decemson, L. Muansanga, M.
Vabeiryureilai, S. Chauhan & Y.S. Rathee (2021). Four new Bent-toed Geckos (Cyrtodactylus
Gray: Squamata: Gekkonidae) from northeast India. Zootaxa 4980(3):
451–489. https://doi.org/10.11646/zootaxa.4980.3.2
Purkayastha,
J., H.T. Lalremsanga, B. Litho, Y.S.
Rathee, S.C. Bohra, M. Vabeiryureilai, L. Biakzuala & L. Muansanga (2022). Two new Cyrtodactylus
(Squamata, Gekkonidae) from northeast India. European Journal of Taxonomy
794: 111–139. https://doi.org/10.5852/ejt.2022.794.1659
Rathee, Y.S.,
J. Purkayastha, H.T. Lalremsanga, S. Dalal, L. Biakzuala, L. Muansanga &
Z.A. Mirza (2022). A new cryptic species of Green Pit Viper of the genus Trimeresurus Lacépède,
1804 (Serpentes, Viperidae) from northeast India. PLoS One 17(5):
e0268402. https://doi.org/10.1371/journal.pone.0268402
Tamura, K.
& M. Nei (1993). Estimation of the number of nucleotide substitutions in the control
region of mitochondrial DNA in humans and chimpanzees. Molecular Biology and
Evolution 10: 512–526. https://doi.org/10.1093/oxfordjournals.molbev.a040023
Vogel, G., Z.
Chen, V. Deepak, D.J. Gower, J. Shi, L.I. Ding & M. Hou (2020). A new species of the genus Smithophis
(Squamata: Serpentes: Natricidae) from southwestern China and northeastern
Myanmar. Zootaxa 4803(1): 51–74. https://doi.org/10.11646/zootaxa.4803.1.3