Content uploaded by H.T. Lalremsanga
Author content
All content in this area was uploaded by H.T. Lalremsanga on May 18, 2022
Content may be subject to copyright.
Content uploaded by Amit Kumar Bal
Author content
All content in this area was uploaded by Amit Kumar Bal on May 16, 2022
Content may be subject to copyright.
101
Systematics of two lesser known colubrid snakes of Northeast India
Open access at hps://www.salamandra-journal.com
© 2022 Deutsche Gesellscha für Herpetologie und Terrarienkunde e.V. (DGHT), Germany
15 May 2022 ISSN 0036–3375
SALAMANDRA 58(2): 101–115 SALAMANDRA
German Journal of Herpetology
Molecular phylogenetic analyses of lesser known colubrid snakes
reveal a new species of Herpetoreas (Squamata: Colubridae: Natricinae),
and new insights into the systematics of Gongylosoma scriptum
and its allies from northeastern India
H T L, A K B, G V L B
1) Developmental Biology and Herpetology Laboratory, Department of Zoology, Mizoram University, 796004 Tanhril,
Mizoram, India
2) Amity Institute of Forestry and Wildlife, Amity University, 201313 Noida, Uttar Pradesh, India
3) Society for South East Asian Herpetology, Im Sand 3, Heidelberg, Germany
Corresponding author: , e-mail: bzachawngthu123@gmail.com
Manuscript received: 9 November 2021
Accepted: 11 April 2022 by
Abstract. A new species of the genus Herpetoreas is described from Murlen National Park, Mizoram, northeastern India,
based on morphological dierences and molecular evidence inferred from the mitochondrial cytochrome b and S rRNA
genes. e new taxon represents the seventh nominal species in the genus Herpetoreas, out of which three species, i.e.,
H.pealii, H. platyceps and H. xenura, have been recorded from northeastern India as well. An updated taxonomic key to
the species of Herpetoreas is provided. Our phylogenetic reconstructions of selected natricine and colubrine snakes from
Mizoram state, northeastern India, also contribute to the systematics and known distribution of the little known species
Gongylosoma scriptum, and corroborate its taxonomic placement into the subfamily Colubrinae. Moreover, the rst mo-
lecular based identications for Liopeltis stoliczkae and Sibynophis collaris from Mizoram are presented.
Key words. Serpentes, Amphiesma, Hebius, Herpetoreas sp. n., Liopeltis, Mizoram, Murlen National Park, Sibynophis, taxo-
no my.
Introduction
The taxonomy of the natricine snake genus Amphiesma
sensu lato
revised by
the genus Herpetoreas Hebius
Amphiesma sensu stricto. The genus
Herpetoreas currently comprises six nominal taxa, namely
H. burbrinki -
H. pealii (H. platyceps (,
H. sieboldii H. xenura (,
H. tpser
of Hebius and one species, A. stolatum
snakes from the last decades have yielded the discovery
of several new species of Hebius (see
et al.
-
ten attributable to their morphological resemblance and
-
though the true diversity of this large and complex group
remains poorly investigated, the existence of multiple
cryptic lineages within these genera seems plausible be-
cause paraphylies among several taxa of these groups have
been demonstrated in a few recent studies (
reconstructions based both upon morphological charac-
specimens from Mizoram belongs to the genus Herpeto-
reas, but it could not be assigned to any of the described
species.We consider this specimen to represent a new spe-
cies, and thus formally describe it herein.
In this study, we also investigated the systematic status
of the poorly known colubrid snake species Gongylosoma
scriptum (T, ). Currently, the genus Gongylo-
soma F, accommodates ve nominal species,
namely G. baliodeira (B, ), G. scriptum (T,
), G. nicobariensis (S, ), G. longicauda
(P, ), and G. mukutense G, D L,
102
H T L et al.
. Although it was speculated to be a genus closely re-
lated to Liopeltis based on morphology (e.g., G et
al. , L , P et al. ), there is no
published evidence that would resolve the precise system-
atic position of this genus, and it thus remained incertae
sedis as of now (U et al. ). Nevertheless, Gongylo-
soma scriptum is one of the least known species among the
congeners, and is the only Gongylosoma species known
from India so far. Altogether, it is known from India (Mi-
zoram) (L et al. ), Cambodia (N et
al. ), Myanmar (D J , T
), and ailand (G et al. , I C-
, S ). Earlier works suggest this species
to be diurnal and secretive in its habits, as all specimens
were encountered during the day, usually below the soil
surface, under rocks, bushes and leaves, and sometimes
exposed on leaf litter, rocks or soil next to forest paths
in evergreen forest, semi-evergreen mixed with bamboo,
tropical wet evergreen forest, and in montane sub-tropi-
cal forest at approximate altitudes between and ,
m asl. (I C , L et al. ,
N et al. ). In addition, the species is poorly stud-
ied and only few specimens are available in museum col-
lections globally (L et al. ). erefore,
we here attempt to ll some knowledge gaps for the ge-
nus as well as to provide new insights into the morphol-
ogy, distributional records, and we reassess the phyloge-
netic status of G.scriptum. In addition, we provide molecu-
lar based identications for morphologically allied species
of G.scriptum, such as Liopeltis stoliczkae (S, )
and Sibynophis collaris (G, ) based on mt S and
Cytb gene sequences generated from new materials from
Mizoram, northeastern India.
Material and methods
Sampling
Specimens analysed in this study were obtained during
eldwork carried out between and in the state
of Mizoram, India. Collected specimens were euthanized
with MS-, following C et al. (); liver tissues
were subsequently recovered by means of dissections and
stored in ethanol for genetic analysis. Specimens were
then xed in formalin, subsequently preserved in
ethanol, and deposited in the reptile section of the De-
partmental Museum of Zoology, Mizoram University. e
sampling sites’ geographical coordinates and altitudes were
recorded using a Garmin Montana- GPS unit, and the
map (Fig. ) was produced using QGIS version ...
Morphology
Morphological measurements were taken with dial calli-
pers accurate to . mm (Mitutoyo –). Snout–vent
(SVL), tail (TaL) and total lengths (TL) were taken to the
nearest millimetre using a measuring tape. e following
morphometrics and meristics were taken: SVL; TaL; TL;
HL, head length; HW, head width; HD, head depth; ED, eye
diameter; E–Sn, eye to snout distance; E–Ns, eye to nostril
distance; SW, snout width; INL, length of internarial scales;
PFL, length of prefrontals; FL, length of frontal; PL, length
of parietals; DSR, dorsal scale rows; PVe, pre-ventrals; Ve,
ventrals; Sc, subcaudals; As, condition of anal scute; SL, su-
pralabials; IL, infralabials; SLe, supralabials contacting the
eye; AG, anterior genials; PG, posterior genials; Tem, tem-
porals; PrO, preoculars; PoO, postoculars; and Lor, lore-
al. Ventral scales were counted following D ().
Dorsal scales were counted at one head length posterior
to the neck, at the midpoint of snout–vent length, and at
one head length anterior to the As. Paired Sc were counted
from immediately posterior to the anal scute to the tail tip,
but excluding the terminal scute.
Figure 1. Digital topographic map of the Indo-Myanmar region:
Type locality of Herpetoreas murlen sp. n. in the Murlen National
Park, Mizoram, India (1); type locality of Gongylosoma scrip-
tum at Mottama, Mon State, Myanmar (2); previous records of
G.scriptum from India, at Saithah village (3) and Suangpuilawn
village (4); new records (this study) at Suangpuilawn village (4),
Baktawng village (5), Dampui village (6), and Vanhne village (7).
103
Systematics of two lesser known colubrid snakes of Northeast India
Museum acronyms
Abbreviations used for natural history collections are:
AMNH, American Museum of Natural History (USA);
BMNH, British Museum of Natural History (Great Brit-
ain); CAS, California Academy of Science (USA); CBC,
Centre for Biodiversity Conservation (Cambodia); CHS,
Collection Huang Song (China); DTU, Duy Tan Univer-
sity, Da Nang (Vietnam); DL, private catalogue of Ding Li
(China); GP, private catalogue of Guo Peng (China); KU,
Biodiversity Institute, University of Kansas (USA); KUZ,
Kyoto University, Department of Zoology Museum (Ja-
pan); LSUHC, La Sierra University Herpetological Col-
lection (USA); MNHN, Muséum national d’Histoire na-
turelle, Paris (France) MZMU, Departmental Museum
of Zoology, Mizoram University (India); NCBS, Nation-
al Centre for Biological Sciences (India); NCSM, North
Carolina Museum of Natural Sciences (USA); NMBE,
Naturhistorisches Museum Bern (Switzerland); NMW
Naturhistorisches Museum Wien (Austria); ROM, Royal
Ontario Museum (Canada); WII, Wildlife Institute of In-
dia (India); YBU, Yibin University (China); ZFMK, Zoo-
logisches For schungs museum Alexander Koenig (Ger-
many), ZMB, Zoo logisches Museum für Naturkunde der
Humboldt-Universität zu Berlin (Germany); ZMMU, Zoo-
logical Museum of Moscow (Russia); ZSM, Zoologische
Staatssammlung München (Germany).
Molecular analyses
Genomic DNA was extracted from the liver tissues using
the QIAamp DNA Mini Kit following the manufacturer’s
protocol. e fragment of S was amplied using for-
ward L (P ) and reverse H prim-
ers (R ) at thermal conditions of °C for
min., followed by cycles of °C for min., .°C for
sec., °C for min., and a nal extension step at °C
for min.; and Cytb using forward and reverse primers
(Snk) (D et al. ) at thermal conditions of °C
for min., followed by cycles of °C for sec., °C
for sec., °C for sec., and a nal extension at °C
for min. e amplied products were puried and se-
quenced using Sanger’s dideoxy method at Barcode Bio-
Sciences, Bangalore, India.
e datasets of natricine and colubrine snakes com-
prised the newly generated sequences of S and Cytb
along with congeneric sequences and outgroups (Am-
phiesma stolatum for natricines; Elaphe cantoris for colu-
brines) obtained from the NCBI database (see Supplemen-
tary Material for sequences utilized). Both datasets were
aligned with the MUSCLE algorithm (E ) us-
ing default parameters in MEGA X (K et al. ).
Both datasets were concatenated in SequenceMatrix v..
(V et al. ), and were partitioned by gene and by
codon positions. PartitionFinder v (L et al. )
was employed for selecting the best partitioning schemes
and for evolutionary model searching under the Bayesian
Information Criterion (Table ). For both datasets, Bayes-
ian Inference (BI) and Maximum Likelihood (ML) phylo-
genies were performed in Mr.Bayes v.. (R et al.
) and in IQ-TREE (N et al. ), respectively.
In the BI analyses, four independent runs with one cold
and three hot chains were conducted for million genera-
tions and sampled every generations aer discarding
the rst of samples as burn-in. e trace plots gener-
ated by the MCMC runs were viewed in Tracer v. (R-
et al. ). Since some of the nucleotide evolution-
ary models selected by PartitionFinder v (L et al.
) were unavailable in IQ-TREE (N et al. ),
the ML analyses were performed using the previously se-
lected partitioning schemes with the models selected un-
der BIC scores by ModelFinder (K et al.
) implemented in the IQ-TREE (N et al. ) at
, Ultrafast Bootstrap (UFB) (M et al. ). e
uncorrected p-distances were estimated in MEGA X (K-
et al. ).
Nomenclatural acts
e electronic edition of this article conforms to the re-
quirements of the amended International Code of Zoo-
logical Nomenclature, and hence the new names contained
herein, are available under that Code from the electronic
edition of this article. is published work and the nomen-
Table 1. Partitions and models of sequencial evolutionary models used in the Bayesian Inference (BI) and Maximum Likelihood (ML)
phylogenetic analyses.
Datasets Partitions Models
BI (PartitionFinder) ML (ModelFinder)
Natricine 16S GTR+I+G TIM2+F+I+G4
Codon 1st pos of Cytb TRN+I+G TIM2+F+I+G4
Codon 2nd pos of Cytb K81UF+I+G TPM2u+F+I+G4
Codon 3rd pos of Cytb TIM+G TIM+F+G4
Colubrine 16S, Codon 1st pos of Cytb GTR+G TIM2+F+I
Codon 2nd pos of Cytb HKY+I HKY+F+I
Codon 3rd pos of Cytb TIM+G TIM+F
104
H T L et al.
clatural acts it contains have been registered in ZooBank,
the online registration system for the ICZN. e LSID (Life
Science Identier) for this publication is: urn:lsid:zoobank.
org:pub: FFD-DA-A-BCDB-BFDADF.
e electronic edition of this work was published in a jour-
nal with an ISSN, has been archived, and is available from
the following digital repositories: www.salamandra-jour-
nal.com, zenodo.org.
Results
e concatenated datasets (S and Cytb) of the natri-
cine and the colubrine groups contained and bp
aligned characters, respectively. Two types of phylogenet-
ic analyses (BI and ML) were performed, which inferred
consistent tree topologies (Fig. ), with the Cytb gene tree
(BI) also being congruent with the concatenated trees (see
Supplementary Figure S). In our phylogenetic analyses of
the natricine group, the unnamed individual from Mizo-
ram, India, is nested with the putative Herpetoreas clade
and forms a sister species to H. burbrinki with well-sup-
ported Bayesian posterior probabilities (PP) and UFB val-
ues. It is distinct from the other Herpetoreas species by
considerable genetic divergences of . (to H. burbrinki)
to . (to H.pealii), while the genetic distances to Hebius
species ranged from . (to H. parallelus CHS de L
et al. ()) to . (to H. concelarus), and . to the
Figure 2. Bayesian inference (BI) phylogenetic tree of the genera Herpetoreas and Hebius, based on the concatenated mitochondrial
cytochrome b and 16S rRNA genes. Bayesian posterior probabilities (PP) are provided before the slash, and Ultrfast Bootstrap (UFB)
values from the Maximum Likelihood (ML) analysis are given aer the slash. e sequence generated in this study is outlined in bold.
105
Systematics of two lesser known colubrid snakes of Northeast India
outgroup (A. stolatum). e morphological features of the
specimen agree with the generic characters of Herpetoreas
provided by G et al. (), but exhibit strong genetic
divergence from the ve nominal species of Herpetoreas.
erefore, based on morphological and molecular data,
the specimen from Mizoram is considered a distinct evo-
lutionary lineage of Herpetoreas, and is described as fol-
lows.
Herpetoreas murlen sp. n.
ZooBank LSID: zoobank.org:pub:FFD-DA-A-BCDB-
BFDADF
N,
Champhai District, Mizoram, northeastern India, collected
Etymology. e specic epithet refers to the type local-
ity, Murlen National park, Champhai District, Mizoram,
India. We propose as common names, Murlen Keelback
Snake (English) and Murlen-Wassernatter (German).
Diagnosis. Herpetoreas murlen sp. n. is diagnosable by
the following morphological characters (based on the holo-
type, male; Figs. –): ) TaL/TL .; ) two PVe, Ve,
and paired Sc; ) As divided; ) nine IL, with the rst
four contacting the AG, and the fourth and h contact-
ing the PG; ) eight SL, the third to h contacting the eye,
the seventh largest; ) Tem +; ) two PrO, and three PoO;
) dorsal scales in :: rows, distinctly keeled, those in
the rst row feebly keeled; ) internasal scales sub-trian-
gular; ) scales on posterior part of head and temporal
region smooth; ) the everted hemipenes bilobate near the
tip; sulcus spermaticus bifurcated just beneath the crotch;
weakly developed apical naked area at the crotch and invis-
ible from the asulcate face; numerous small spines present,
increasing in size distally, and a distinct basal hook present
(Fig. ).
Description of the holotype. Body elongated and slender;
SVL mm; TaL mm; TaL/TL .; head distinct
from neck, longer than wide (. of the SVL); snout
modestly long ( of HL); nostrils small; eyes rather
large ( of HL), pupils round; a single loreal, wider
than long; prefrontals wider than long, extending later-
ally in contact with internasals, nasals, loreal, preoculars,
and frontal; frontal pentagonal, anterior part a bit wider;
parietals longer than wide; two PrO, and three PoO; max-
illary teeth on one side, the last two distinctly enlarged
without diastema between the last two and the anterior
teeth.
In life, body dark olive-grey, most scales randomly
speckled with black on the margins; inter-scales sparse-
Figure 3. Holotype of Herpetoreas murlen sp. n. (MZMU2041) in life (A–C). Head of an uncollected individual in dorsolateral view (D).
106
H T L et al.
ly speckled with white; a dorsolateral stripe extending
from the neck to the tail, covering the upper part of the
th row along the anterior part of the body (where there
are DSR), the whole of the th row, and the lower part
of the th row, covering the upper part of the th row along
the posterior part of body (where there are DSR), the
whole of the th row, and the lower part of th row, con-
spicuous maroon, lighter in the inter-scales, with dark
speckles on the margins of the lower part of the th row
and the upper part of the th row along the anterior part
and in the lower part of the th row and the upper part
of the th row along the posterior contrast the stripe; tail
colored and patterned as the posterior part of the body,
with the maroon dorsolateral stripe progressively vanish-
ing towards half the length of the tail, ill-dened and dif-
fuse towards the tail tip.
Dorsal face of head and nape dark olive-brown; a me-
dian lighter brown stripe on the mid-dorsal scales of the
nape starting from behind the posterior interparietals,
anked by irregular dark speckling on either side, bifur-
cating at the th scale, partially interrupted by small dark
speckles before becoming the dorsolateral stripes on both
sides; frontal with irregular and thin dark speckles on the
lateral and posterior margins. Lower region of the snout
and the lateral faces of the head pale ivory cream, demar-
cated from the upper olive-brown coloration by a distinct
dark streak on the upper borders of the rostral and ex-
tending posteriorly through the nasal, loreal and the low-
er pre-ocular; a continuous streak on the middle postocu-
lar, upper part of the lower postocular, st temporal, upper
parts of th and th SLs, and extending posteriorly as an
oblique band in the th SL on both sides of the head; an-
terior supralabials pale ivory cream and edged with dark
brown on their posterior borders; a dark spot on the up-
per edge of the th SL, an oblique black streak from the
posterior upper margin of the th SL interrupted in the
lower anterior part of the th SL; anterior IL the same col-
our as SL, thinly framed with darker [igment on their pos-
terior borders.
Venter uniformly light sand-brown, the outer part of
each scale brick-red, becoming darker olive-brown on the
Figure 4. Dorsal (A) and ventral (B) views of the preserved holo-
type (MZMU2041) of Herpetoreas murlen sp. n.
Figure 5. Dorsal (A), ventral (B), and lateral (C) views of the
head of the holotype (MZMU2041) of Herpetoreas murlen sp. n.
107
Systematics of two lesser known colubrid snakes of Northeast India
scales’ outer tips; each ventral with a thin, yet well dened,
blackish brown spot between the brick-red and light brown
parts, these spots become progressively smaller posteriorly
from mid-body and do not connect with each other. Ven-
tral part of tail same as the venter, bordered on either side
by a blackish brown stripe as described above; the black-
ish brown ventral spots progressively vanish in the ante-
rior half of the tail length. Chin, throat, and ventral part of
the neck uniform whitish ivory cream, lighter than venter.
Posterior infralabials each with a blackish spot on the rear
margin (Figs A–C).
Comparison with other species. Among the known
Herpeto reas species, the new species is genetically closest
to H. burbrinki by forming a well supported sister lineage
in all the phylogenetic analyses. In the Cytb gene-based
genetic divergence (p-distance) estimation (Table ), the
new species likewise has the least genetic divergence from
H.burbrinki (.). However, the new species is morpho-
logically distinct from H. burbrinki by having: smooth
scales in the posterior part of the head and in the temporal
region (vs. keeled); more Ve ( vs. –); fewer Sc (
vs. –); a lower TaL/TL ratio (. vs. .–.); fewer
maxillary teeth ( vs. ); fewer Ate and Pte (+ vs. + or
+); lesser IL ( vs. ), and a lower number of IL contact-
ing the AG (– vs. –) (Table ).
Herpetoreas murlen sp. n. is distinguished from
H. pealii
Herpetoreas murlen H. xenura in the
Herpetoreas murlen H. platyceps by
-
-
fers from Tropidonotus rthi (-
nym of H. platyceps, by having
Herpetoreas murlen sp. n. is also distinct from H. sie-
boldii -
-
Taxonomic remarks. During the eldwork for this study,
another individual was encountered near the type locality
within the Murlen National Park at ca. hrs on No-
vember . Unfortunately, this individual (approx. TL
mm) escaped aer it was photographed (Fig. D). Its
head scalation largely agreed with that of the holotype ex-
cept that some variation was seen with regard to the le
temporals, i.e., +/+ vs. +/+ in the holotype.
Distribution and natural history. Herpetoreas murlen sp. n.
is as yet known only from the type locality in the Murlen
National Park, located in Champhai District, Mizoram
(.–.° N, .–.° E), in the northeastern por-
tion of Mizoram state. e holotype was collected from leaf
litter along a forest trail (Fig. ). e area is characterized
by dense tropical to subtropical evergreen mixed forests,
covering ca. km² at an altitude of to ,m a.s.l.
It receives an annual rainfall of ,–, mm with tem-
perature ranging between °C in winter and °C in sum-
mer (S et al. ). According to C S
(), the area falls into the category of Assam Subtropical
Pine Forest (/C).e vegetation is dominated by Quercus
sp., Schima wallichii, Betula sp., Michelia champaca, Pinus
khasiana, Prunus, Myrica sp., Rhododendron sp., Arundi-
nania callosa, canes,and a rich variety of orchids. Other
snakes encountered during the survey at the type local-
Figure 6. Everted hemipenis, sulcal side (A) and asulcal side (B) of the holotype (MZMU2041) of Herpetoreas murlen sp. n.
108
H T L et al.
Table 2. Morphometric and meristic characters of Herpetoreas murlen sp. n., and comparative data of its congeners obtained from
published data. For H. xenura, a total of 18 specimens examined in this study and published data were combined. Unavailable data
are indicated as “–”.
Taxa H. murlen sp. n. H. burbrinki H. tpser H. pealii H. xenura H. platyceps H. sieboldii
Vouchers MZMU
2473
YBU
071128
ANU
20210006
– BMNH
1946.1.13.43
WII
ADR547
– – –
Sex Male
Holotype
Male
Holotype
Female
(n=1)
Sex pooled
n=6
Male (n=1)
Lectotype
Female
(n=1)
Sex pooled
(n=18+)
Sex pooled
(n=?)
Sex pooled
(n=?)
SVL 355 459 462 279–491 334 511 364–550 570–655 –
TaL 106 130 184 108–207 117 150 176–210 165– 225 –
TaL/TL 0.23 0.26 0.29 0.26–0.32 0.26 0.23 0.26–0.31 0.23–0.29 0.24–0.32
DSR 19:19:17 19:19:17 19:19:17 19:19:17 19:19:17 19:19:17 19:19:17 19:19:17 19:19:19
Ve 179 172 169 153–167 142 136 156–165 191–234 190–216
PVe 2 2 2 – 3 2 – – –
Sc 78
(paired)
96
(paired)
94
(paired)
79–97 (paired or
Sc 7–9 single)
77
(paired)
69
(paired)
87–114
(entire)
78–98
(paired)
81–111
As 2 2 2 2 1 1 1 or 2 – 2
SL 8/8 8 9/8 8 or 9 9/9 9/9 9/9 8 8
IL 9/9 10 10/10 9–10 (8) 10/10 10/10 10/10 8 or 9 10
SLe 3–5/3–5 3–5 3–5 3–5 or 4–6 4–5/4–5 4–5/4–5 4–6/4–6 3–5 3–5
IL contacting AG 1–4/1–4 1–5 1–5 1–4 or 1–5 1–5/1–5 1–5/1–5 – – –
IL contacting PG 4–5/4–5 – – – 5–6/5–7 5–7/5–7 – – –
Tem 1+1/1+1 3+2/3+2 2+2/2+2 1–2 [or 1+1/1] +
2, 1–2+1
[or 1/(1+2)]
2+2/2+2 2+2/2+2 2+2 or 2+1
or 1+2
(rarely 1+1)
1+1
(rarely 2+2)
1 or 2
anterior
PrO 2/2 2/1 1/1 1 or 2 1/1 1/1 1/1 1 1 or 2
PoO 3/3 3 3/3 3 or 2 3/3 3/3 3/3 2 or 3 2
Lor 1/1 1 2/2 1/1 in holotype 1/1 1/1 1/1 1 1
HL 13.96 – 16.5 9.60–13.56 15.5 22.4 12.44–16.60 – –
HW 6.80 – 9.2 6.91–9.54 7.3 11.9 8.50–12.98 – –
HD 4.60 – 6.1 – 5.2 8.9 – – –
ED 2.48 – 3.8 – 2.5 3.3 2.44–3.32 – –
E–Sn 3.28 – – – 4.1 6.1 – – –
E–Ns 1.70 – – – 2.7 3.7 2.70–4.22 – –
INL 1.60 – – – 1.4 2.0 – – –
PFL 1.90 – – – 1.9 2.8 – – –
FL 3.72 – – – 3.8 5.9 – – –
PL 5.46 – – – 6.3 9.2 – – –
Maxillary teeth 13, gradually
enlarged
posteriorly, last
two teeth (larger)
aer a small
gap, but without
diastema
–21 in a
continuous
series, last
15 enlarged.
20–21, last two
enlarged, a small
gap present, with
diastema
13–21,
gradually
enlarged
posteriorly.
Last two
teeth aer
a small gap,
but without
diastema
22–23,
gradually
enlarged
posteriorly.
Last two
teeth (large)
aer a small
gap, but
without
diastema
18–22 in a
continuous
series,
last 16
enlarged
17–21 + 2
with a wide
diastema,
last two
enlarged
Hemipenes Bilobate ar tip,
sulcus distinct
and bifurcated
just beneath the
crotch area of the
two lobes. Basal
hook present
– – Sulcus single,
extends to crotch
Sulcus
single,
bilobate
– Sulcus
single,
bilobate
at tip
Sulcus single,
extends to tip,
unilobate
rarely
bilobate
Sulcus single,
extends to
tip,
bilobate
at tip
Source is study G et al.
(2014)
P et al.
()
R et al. () M
(1960); D
et al. (2020)
D et al.
(2020); is
study
S
(1943);
M
(1966);
D et al.
(2015)
G
(1860); M-
(1966);
G et al.
(2014); D
et al. (2015)
109
Systematics of two lesser known colubrid snakes of Northeast India
ity were Ahaetulla prasina, Boiga ochracea, Dendrelaphis
cyanochloris, Gonyosoma prasina, Hebius khasiense, Lyco-
don fasciatus, Oligodon dorsalis, Oreocryptophis porphyra-
ceus, Orthriophis taeniurus, Ovophis monticola, Pareas an-
dersoni, Pareas monticola, Psammodynastes pulverulen-
tus, Pseudo xenodon macrops, Rhabdophis helleri, Sibyno-
phis collaris, Sinomicrurus macclellandi, and Tri meresurus
erythrurus. Since the two individuals of H. murlen sp. n.
were encountered during dusk (– hrs), we sug-
gest the species to be crepuscular in activity. Although the
park has several small seasonal and perennial streams, and
is also characterized by an undulating topography and rug-
ged mountains (S et al. ), there was no open
water apparent in the vicinity of the collection site. It seems
plausible that the species may occur in the neighbouring
Manipur state (India) and some parts of the adjacent Chin
state (Myanmar).
Systematic reassessment
In the scenario of the colubrine dataset, both the BI and
ML phylogenetic analyses inferred ) with signicant PP
and UFB support the clustering of the new specimen of
G.scriptum (MZMU) from Mizoram with a conspecif-
ic specimen from Cambodia (NCSM, and both spec-
imens clearly nested within a clade of Liopeltis consisting
of L. frenatus + L. pallidonuchalis + Liopeltis sp. (Fig.A).
From the Cytb gene based p-distance estimation, G. scrip-
tum (MZMU) from Mizoram has an intraspecic dis-
tance of . from the specimen (NCSM) from Teuk
Chou, Kampot Province (Cambodia), while the smallest
interspecic divergence (.) is seen vis-à-vis L. frenatus
(CAS) and the greatest (.) vis-à-vis S. collaris
(CHS). e two specimens of L. stoliczkae (MZMU,
MZMU) from Mizoram exhibit . divergence be-
tween them, and . intraspecic genetic distance vis-à-
vis the Chinese specimen (CHS); with the other conge-
ners, genetic divergences of . (L. philippina) to .
(L. pallidonuchalis) are manifest. Only the S fragment for
S. collaris was generated in this study so that the estimation
of genetic distances was performed using the S dataset.
Sibynophis collaris (MZMU) from Mizoram (India)
showed . and . genetic distances compared with a
conspecic specimen from China (CHS) and one spec-
imen of unknown provenance (ROM), respective-
ly. Our specimen also exhibited . interspecic genet-
ic divergences vis-à-vis the two specimens of S. chinensis
(CHS, CHS). Although the present molecular phy-
logenetics and DNA barcoding may suggest an inclusion of
G. scriptum in the genus Liopeltis, we took a conservative
approach and le the generic reallocation of G. scriptum
pending due to insucient data. However, we contribute
herein additional data for the species that may be crucial
for a future nomenclatural re-evaluation of the genus.
Figure 7. Microhabitat at the collection site of Herpetoreas murlen
sp. n. in the Murlen National Park, Mizoram, northeastern India.
Table 3. Uncorrected p-distances for members of the putative Herpetoreas clades. GenBank accession numbers are provided aer the
species names.
Species 1 2 3 4 5 6 7 8 9 10
1. Herpetoreas murlen sp. n. ON204025
2. Herpetoreas burbrinki GQ281781 0.087
3. Herpetoreas platyceps MT571587 0.111 0.109
4. Herpetoreas platyceps KJ685690 0.115 0.123 0.043
5. Herpetoreas platyceps MW111464 0.115 0.125 0.043 0.003
6. Herpetoreas xenura MN993850 0.130 0.125 0.126 0.144 0.140
7. Herpetoreas xenura MN993851 0.130 0.125 0.126 0.144 0.140 0.000
8. Herpetoreas pealii MT571586 0.145 0.120 0.133 0.144 0.144 0.113 0.113
9. Herpetoreas tsper MW111476 0.099 0.089 0.096 0.108 0.104 0.135 0.135 0.130
10. Hebius parallelus MK201567 0.099 0.092 0.099 0.111 0.108 0.135 0.135 0.133 0.003
110
H T L et al.
Gongylosoma scriptum (T, 1868)
Holotype. ZSI , Martaban (= Mottama), Mon State,
Myanmar.
-
-
-
et
by and
“East India”, collected by
W. .
Figure 8. (A) Bayesian inference (BI) phylogenetic tree of Gongylosoma scriptum and allied species, based on the concatenated mi-
tochondrial cytochrome b and 16S rRNA genes. Bayesian posterior probabilities (PP) are provided before the slash, and Ultrafast
Bootstrap (UFB) values from the Maximum Likelihood (ML) analysis are given aer the slash, the value for the node not recovered in
the ML tree is given as “–”. Sequences generated for this study are outlined in bold; (B) e newly collected specimen of G. scriptum
(MZMU2041) in life; (C) sulcal (le) and asulcal (right) views of the everted hemipenis of G. scriptum (MZMU2041).
111
Systematics of two lesser known colubrid snakes of Northeast India
Description. Based on the material examined, body slen-
der and elongate; tail thin; head moderately distinct from
the neck; eye rather large, pupil circular; snout blunt;
rostral slightly visible from above; loreal small; preocu-
lar higher than wider. Measurements: SVL – mm
in males (n=), –. mm in females (n=); TaL
– mm in males, mm in one female (excluding
the specimen with the damaged tail tip); ratio TaL/TL
.–. in males, . in one female; HL: –. mm
in males, .–. mm in females; HW .–. mm in
males, .–. mm in females; ratio HW/HL .–.
in males, .–. in females; ED .–. mm in males,
.–.mm in females; ratio ED/HL .–. in males,
.–. in females. Meristics: dorsal scale rows ::,
all smooth; Lor single; one nasal and two internasals; two
prefrontals and a single frontal; Ve – in males, –
in females; paired Sc – in males, in one female;
anal scutedivided; SL eight on both sides, rarely seven;
eight IL on both sides; SLe –, rarely –. Morphological
data taken from the new specimens and published values
are given in Table .
collected specimen shows a simple cylindrical structure
-
face is covered with smooth epithelium. The distal third of
the organ is covered with fairly evenly distributed small
-
mal area with medium-sized spine-line, but less densely
spermaticus single and running up to the crotch of the api-
Taxonomic remarks. This species is diagnosable from
its congeners by having thin and light coloured vertical
morphological characters of the newly collected specimen
, ) largely agree with
the species’ original description (-
er specimens ( et al.
et al.
-
et al.
Table 4. Morphological data of Gongylosoma scriptum obtained in this study and published data. Asterisk indicates tail tip damaged,
and unavailable data and revised data aer re-examination of the same specimens are indicated as “–” and “#”, respectively.
Source is study L et al.
(2018)
N et al. (2015) S (1943)
Locality India
(Mizoram)
ailand
(Khao Yai)
East India India
(Mizoram)
Cambodia
(Pursat)
Cambodia
(Preah Sihanouk)
–
Vouchers MZMU
2041
ZMB
50677
ZMB
5286
MZMU
892
MZMU
914
CBC
01365
CBC
02543
–
Sex Male Male Female Female Male Female Male –
SVL 280.0 228 171 224.2# 230.0# 256.1 252.2 465 (male),
495 (female)
TaL 155.2 161 79 58.0#* 123.0# 145.6 172.2 155 (male)
TaL/TL 0.36 0.41 0.32 * 0.35 0.36 0.41 –
DSR 13:13:13 13:13:13 13:13:13 13:13:13 13:13:13 13:13:13 13:13:13 13:13:13
Ve 128 123 136 139 127 138 122 126–145
Sc 89 98 77 45* 88 93 101 87–98
As 2 2 2 2 2 2 2 2
SL 7/8 8/8 8/8 8/8 8/8 8/8 8/8 8/8
IL 8/8 8/8 8/8 8/8 8/8 8/8 8/8 –
SLe 3–4/3–5 3–5/3–5 3–5/3–5 3–5 3–5 3–5 3–5 3–5
Tem 1+2/1+2 – – 1+2/1+2 1+2/1+2 1+2/1+2 1+2/1+2 1+2/1+2
PrO 1/1 – – 1/1 1/1 1/1 1/1 1/1
PoO 2/2 – – 2/2 2/2 2/2 2/2 2/2
HL 11.0 13.4 10.8 9.9 10.0 – – –
HW 6.8 6.7 4.8 4.6 7.2 – – –
ED 2.6 2.4 1.8 2.1 2.2 – – –
E–Ns 1.9 1.4 1.0 0.9 1.1 – – –
SW 3.4 – – 2.6 3.0 – – –
112
H T L et al.
Distribution and natural history. In addition to the known
distributional records from India (Mizoram) (L-
et al. ), we documented three new distributional re-
cords during this study: the specimen MZMU was col-
lected at hrs on October , from Suangpuilawn
village, Aizawl District (.° N, .°E; ,
m a.s.l.) by J. C. L, the same locality where
L et al. () collected one of their speci-
mens; an uncollected individual was encountere d at hrs
on May on the way to Dampui village from en-
hlum village, Lunglei District (.° N, .°
E; m a.s.l.) by P S; on May , one indi-
vidual was observed at Baktawng village, Serchhip District
(.° N, .° E; m a.s.l.; and another in-
dividual was documented on June , at Vanhne vil-
lage, Lunglei District (.° N, . °E; m
a.s.l.) by M K; All specimens were encoun-
tered during daytime in habitats similar to the previous re-
cords (I C , L et al. ,
N et al. ).
Discussion
In this study, we provide molecular evidence supporting
the morphology-based hypothesis of a close relationship
between the genera Gongylosoma and Liopeltis (see G-
et al. , L , P et al. ).
Our phylogenetic reconstructions ascertain the taxonom-
ic placement of Gongylosoma in the subfamily Colubri-
nae. Although the genus was resurrected from the syn-
onymy of Liopeltis by L () and most recent
publications consider the genus Gongylosoma valid (e.g.,
G et al. , L et al. , N et
al. , P et al. , S et al. ); evidence
from the present molecular phylogenies instead warrants
designating its generic position to Liopeltis de M
G (), S (), and W ().
Yet, the precise determination of the generic position of
G. scriptum must remain unresolved as of now due to the
unavailability of genetic data from either the type speci-
men, topotypical specimens, or congeneric specimens.
However, the morphology-based phylogeny proposed by
G et al. () suggests a probably close relation-
ship between G. scriptum and G. baliodeirum (type spe-
cies); thus, it seems plausible that the specimens of G.
scriptum examined in this work reect the generic po-
sition of this species to some extent. Consequently, this
work has put forward a taxonomic recommendation for
the genus Gongylosoma. Further research by sampling
more genetic and morphological information is needed
to resolve the nomenclatural uncertainty of this genus by
and large. Nevertheless, we trust that the so far very limit-
ed knowledge of this taxon is improved to a certain degree
by the newly generated molecular data, distributional re-
cords, and morphological data. Currently, the IUCN con-
siders G. scriptum “least concern”, but it is also known to
suer from habitat loss and degradation in ailand and
Myanmar (G et al. ). More extensive surveys
are recommended to establish the exact range of distribu-
tion and the population status of this little-known snake,
which will be important for implementing knowledge-
based conservation strategies.
Phylogenetic analyses of natricine snakes in this study
largely showed similar topologies to those arrived at by
previous workers (G et al. , L et al. , Z et
al. ). Considering the fact that due to paucity of data
many of the nominal Hebius species are not covered in
phylogenetic studies (D et al. ), and even the plau-
sible placement of certain species of Hebius in either Am-
phiesma sensu stricto or Herpetoreas (G et al. ), the
present phylogenetic inferences, however, clearly depict a
monophyletic clade of Hebius with well-supported root-
ing. On the other hand, Herpetoreas is recovered as para-
phyletic and comprising three monophyletic sub-clades in
the shapes of the H. platyceps lineage, the species group
lineage of H. murlen + H. burbrinki + H. tpser, and the
H.xenura + H. pealii species group lineage. In this study,
although we could not examine the type species of the ge-
nus Herpeto reas, i.e., H. sieboldii (G, ), we in-
cluded H. platy ceps, which was previously synonymized
with H. sieboldii, but later removed from synonymy and
considered a closely related taxon (M ). In the
latest review of the genus Herpetoreas, R et al. ()
mentioned misidentied specimens of H. parallelus from
Medog, Tibet, China, including the specimen KIZ
de X et al. (), and subsequently re-allocated the Ti-
betan specimens along with KIZ to their newly de-
scribed H. tpser. Moreover, the evidence from the present
phylogenetic study depict H.parallelus (CHS) de L
et al. () clustering with H. tpser (KIZ), which
clearly refers specimen CHS to H. tpser. e descrip-
tion of the new species brings the number of recognized
species in the genus Herpetoreas to seven, of which four of
them (H. pealii, H. xenura, H. platyceps, and H. sieboldii)
have already been recorded from India (U et al. ),
and the remaining ones (H. burbrinki and H. tpser) are re-
garded as endemic to China as of now (G et al. ;
R et al. ). Moreover, the specimen of “H. venningi“
recorded earlier from Mizoram by B L-
() has since been referred to an undescribed
species (D et al. ) so that the systematic status of
this Hebius species is also waiting to be resolved in future
studies. Given that the northeastern Indian Herpeto reas,
i.e., H.xenura and H.pealii, have been recorded at altitudes
from m a.s.l. to mid-altitudes of , m a.s.l. (D et al.
), we consider H. murlen a more montane zone dwell-
er as the only known specimens were found in the high-
altitude zone of Mizoram state at , m a.s.l.
Given the lack of information on the possible threats,
reproductive biology, feeding habits, population struc-
ture, distribution range, and other ecological data of the
new species, we suggest H. murlen to be regarded as a data
decient (DD) species under the categorization of the
IUCN Red List of reatened Species (IUCN Standards
and Petitions Subcommittee ).
113
Systematics of two lesser known colubrid snakes of Northeast India
Updated key to the genus Herpetoreas
1A Subcaudals single .......................................... H. xenura
1B Subcaudals divided ...................................................... 2
2A Anal scute single ............................................. H. pealii
2B Anal scute divided ....................................................... 3
3A Ventrals fewer than 168 ................................... H. tpser
3B Ventrals no less than 168 ............................................ 4
4A Tail relatively long, TaL/TL > 0.300 ....... H. burbrinki
4B Tail relatively short, TaL/TL ≤ 0.300 ......................... 5
5A Ventrals < 190 ..................................... H. murlen sp. n.
5B Ventrals ≥ 190 ............................................................... 6
A A high proportion of the length of the tail with
supracaudal scale rows to that with supracaudal
scale rows high, SC4/SC6 = 1.43 ............ H. platyceps
B A low proportion of the length of the tail with
supracaudal scale rows to that with supracaudal
scale rows high, SC4/SC6 = 0.53 .............. H. sieboldii
Acknowledgements
We are grateful to the Chief Wildlife Warden of the Environ-
ment, Forests and Climate Change Department, Government
of Mizoram, for the collection permit of herpetological speci-
mens within Mizoram (No. A.//-CWLW/). is re-
search work is supported by DST-SERB, New Delhi (DST No:
EMR//); DBT, New Delhi (DBT-NER/AAB//);
National Mission for Himalayan Studies (NMHS), Uttarakhand
(GBPNI/NMHS-/MG-/); DRDO, New Delhi (DGTM/
DFTM/GIA/-/); and DST-SERB, New Delhi (DST No.
EEQ//). e senior author (HTL) expresses his grati-
tude to the Herpetological Symposium, Inc. (IHS), USA; AKB ac-
knowledges the National Mission for Himalayan Studies (NMHS),
Uttarakhand (GBPI/IERP/-/) for funding; and LB is also
grateful to e Ruord Foundation, UK, for the nancial grant
No. -. Our thanks and appreciation extend to M V-
, L M, M V, R
M, and L R for their assistance in this study.
We deeply appreciate the valuable comments from J L. L
on the initial version of the manuscript. We are grateful to P -
C (BMNH), D L (DL), N V, A
D, and A O (MNHN), S S
and G G (NMW), D R and W
B (ZFMK), and M-O R and F T
(ZMB), and to F G and M F (ZSM) for
letting us examine specimens in the collections of their respective
institutions.
References
A, L. R., T. B. Q, F. G. G, M. L. A,
M. M, M. V H. Z (): Diversication
in vipers: Phylogenetic relationships, time of divergence and
shis in speciation rates. – Molecular Phylogenetics and Evo-
lution, : –.
B, L. H. T. L (): Geographic distri-
bution. Hebius venningi (Chin Hills Keelback). – Herpetologi-
cal Review, : .
B, E. (): Notices and descriptions of various reptiles, new
or little known [part ]. – Journal of the Asiatic Society of Ben-
gal, : –.
B, F. (): Bemerkungen über Merrem‘s Versuch eines Sy-
stems der Amphibien, . Lieferung: Ophidier. – Isis von Oken,
: –.
C, S. H. G. S. K. S (): A Revised Survey of the
Forest Types of India. – e Manager of Publication, Govt. of
India, New Delhi.
C, C. J., T. P, J. P N. E. H (): Use
of tricainemethanesulfonate (MS) for euthanasia of reptiles.
– Journal of the American Association for Laboratory Animal
Science, : –.
D, A., D. J. G V. D (): Lost and found: redis-
covery and systematics of the Northeast Indian snake Hebius
pealii (S, ). – Vertebrate Zoology, : –.
D, P. I. D (): A new species of the snake genus Am-
phiesma (Serpentes: Colubridae: Natricinae) from western Su-
matra, Indonesia. – Raes Bulletin of Zoology, : –.
D, P. G. Vogel (): A new species of the natricine snake
genus Amphiesma from Borneo (Squamata: Natricidae). – Rus-
sian Journal of Herpetology, : –.
D, P., R. H. B, N. Q. T, N. L. O, G. V, V.
N. T T. Z (): A new species of the natricine
snake genus Amphiesma from the Indochinese Region (Squa-
mata: Colubridae: Natricinae). – Zootaxa, : –.
D, P., I. A, R. A, R. M, G. V V.
K. M (): Revalidation of Natrix clerki Wall, , an
overlooked species in the genus Amphiesma Duméril, Bibron
& Duméril, (Squamata: Natricidae). – Zootaxa, : –
.
D, P., G. V, T. Q. N, N. L. O, O. S. P,
A. T T. Z (): A revision of the dark-bellied,
stream-dwelling snakes of the genus Hebius (Reptilia: Squama-
ta: Natricidae) with the description of a new species from Chi-
na, Vietnam and ailand. – Zootaxa, : –.
D, H. G. (): A proposed standard system of counting
ventrals in snakes. – British Journal of Herpetology, : –.
D, H. G. J. V. J (): Snakes of Burma: checklist
of reported species and bibliography. – Smithsonian Herpeto-
logical Information Service .
D, B., P. R. M I. H (): Multiplex PCR
assay for rapid identication of three endangered snake species
of India. – Conservation Genetics, : –.
D, A. M. C., G. B A. H. A. D (): Er-
pétologie générale ou histoire naturelle complète des reptiles.
Tome septième Premiere partie. Comprenant l’histoire des ser-
pents non venimeux. – Librairie Encyclopédique de Roret, Pa-
ris.
E, R. C. (): MUSCLE: a multiple sequence alignment
method with reduced time and space complexity. – BMC Bio-
informatics, : –.
F, A., A. D. MK, L. L. G, C. D. B S. P.
L (): A species-level phylogeny of extant snakes
with description of a new colubrid subfamily and genus. –
PLOS One, : p.e.
F, L. J. (): Systema reptilium. Fasciculus primus,
Amblyglossae. – Braumüller et Seidel, Vindobonae.
G, J. E. (): Descriptions of some undescribed species of
reptiles collected by Dr. Joseph Hooker in the Khassia Moun-
114
H T L et al.
tains, East Bengal, and Sikkim Himalaya. – e Annals and
Magazine of Natural History, : –
G, L. L., I. D T. M. L (): A new species of
Gongylo soma (Squamata: Colubridae) from Pulau Tioman,
West Malaysia. – Herpetologica, : –.
G, L., T. C-A, N. R M. B. ():
Gongylo soma scripta. e IUCN Red List of reatened Spe-
cies : e.TA. http://dx.doi.org/./
IUCN.UK.-.RLTS.TA.en, downloaded on
October .
G, A. (): Contributions to a knowledge of the reptiles
of the Himalaya mountains. – I. Descriptions of the new spe-
cies. II. List of Himalayan reptiles, with remarks on their hori-
zontal distribution. – Proceedings of the Zoological Society of
London, –.
G, P., F. Z, Q. L, L. Z, J. X. L, Y. Y. H R. A.
P (): A taxonomic revision of the Asian keelback
snakes, genus Amphiesma (Serpentes: Colubridae: Natricinae),
with description of a new species. – Zootaxa, : –.
I, R. F. R. K. C (): Organization of contiguous
communities of amphibians and reptiles in ailand. – Eco-
logical Monographs, : –.
IUCN Standards and Petitions Committee (): Guidelines for
Using the IUCN Red List Categories and Criteria. Version .
Prepared by the Standards and Petitions Committee. Down-
loadable from http://www.iucnredlist.org/documents/RedList-
Guidelines.pdf.
K, T. T. M (): e biogeographical history of
Asian keelback snakes of the genus Hebius (Squamata: Colu-
bridae: Natricinae) in the Ryukyu Archipelago, Japan. – Bio-
logical Journal of the Linnean Society, : –.
K, S., B. Q. M, T. K. W, A. V H-
L. S. J (): ModelFinder: fast model selection
for accurate phylogenetic estimates. – Nature Methods, :
–.
K S., G. S, M. L, C. K K. T ():
MEGA X: Molecular Evolutionary Genetics Analysis across
computing platforms. – Molecular Biology and Evolution, :
–.
L, H. T., L, M. V, V-
G. V (): First record of the species
Gongylo soma scriptum (eobald, ) (Squamata: Colubri-
dae) from India. – Hamadryad, : –.
L, S., C. L, V, D. J. G-
V. D (): A multilocus molecular perspective on
the systematics of the poorly known Northeast Indian colubrid
snakes Blythia reticulata (Blyth, ), B. hmuifang Vo gel, L-
V, , and Hebius xenura (Wall,
). – Zootaxa, : –.
L, R., P. B. F, A. M. W, T. S B.
C (): PartitionFinder : new methods for selecting
partitioned models of evolution for molecular and morpholog-
ical phylogenetic analyses. – Molecular Biology and Evolution,
: –.
L, A. E. (): Contributions to a review of Philippine
snakes, II. e snakes of the genera Liopeltis and Sibynophis. –
e Philippine Journal of Science, : –.
L, J. N., D. L, Y. Y. W, P. G, S. H P. Z
(): A large-scale systematic framework of Chinese snakes
based on a unied multilocus marker system. –Molecular Phy-
logenetics and Evolution, : .
L, Q., G. H. Z, P. W, Y. L P. G (): A new
species of the genus Hebius (Squamata: Colubridae) from Si-
chuan, China. – Zootaxa, : –.
M, R., U. M, G. V, P. H J. K-
(): Amphibians and Reptiles of Mount Kinabalu
(North Borneo). – A.R.G. Gantner Verlag Kommanditgesell-
scha, Ruggell (Liechtenstein).
M, E. V. (): Systematic division and evolution of the
colubrate snake genus Natrix, with comments on the subfam-
ily Natricinae. – Proceeding of Academy of Natural Science,
Philadelphia, : –.
M, E. V. (): Amphiesma platyceps (Blyth) and Am-
phiesma sieboldii (Günther); sibling sp. (Reptilia: Serpentes). –
Journal of Bombay Natural History Society, : –.
M, B. Q., M. A. T. N A. H (): Ultra-
fast approximation for phylogenetic bootstrap. – Molecular Bi-
ology and Evolution, : –.
N, T., L. L. G, S. H C. P (): New her-
petofauna records and range extensions for Daboia siamensis
(Smith, ) and Gekko petricolus Taylor, from Cambo-
dia. – Cambodian Journal of Natural History, : –.
N, L. T., H. A. S, A. H B. Q. M
(): IQ-TREE: A fast and eective stochastic algorithm for
estimating maximum likelihood phylogenies. – Molecular Bio-
logy and Evolution, : –.
P, S. R. (): Nucleic acids II: the polymerase chain re-
action. – pp. in: H, D. M., C. M B. K. M
(eds): Molecular Systematics. – Sinauer Associates Inc., Sun-
derland.
P, L., S. H, F. T. B, Y. Z, P. G H.
W (): Morphological description of a new specimen of
Herpeto reas burbrinki Guo et al. (Serpentes: Colubridae).
Zootaxa, : –.
P, W. (): Uber neue Reptilien aus Ostafrika und Sarawak
(Borneo), vorzüglich aus der Sammlung des Hrn. Marquis J.
Doria zu Genua. – Mber. k. preuss. Akad. Wiss., Berlin.
P , N. A., T. V. N G. V (): A new spe-
cies of the genus Liopeltis F from Vietnam (Squa-
mata: Colubridae). – Journal of Natural History, : –.
P, J. P. David (): A new species of the snake ge-
nus Hebius ompson from Northeast India (Squamata: Natri-
cidae). – Zootaxa, : –.
P, R. A., H. D. K, C. R. H, V. P, F.
T. B R. S (): Genus-level phylogeny
of snakes reveals the origins of species richness in Sri Lanka. –
Molecular Phylogenetics and Evolution, : –.
R, A., A. J. D, D. X, G. B M. A. S-
(): Posterior summarization in Bayesian phyloge-
netics using Tracer .. – Systematic Biology, : –.
R, K. (): e microevolution of the Galápagos ma-
rine iguana Amblyrhynchus cristatus assessed by nuclear and
mitochondrial genetic analyses. – Molecular Phylogenetics and
Evolution, : –.
R, J. L., K. J, J. J. H, P. D J. T. L (): Taxo-
nomic Review of the Genus Herpetoreas (Serpentes: Natrici-
dae), with the Description of a New Species from Tibet, China.
– Diversity, : .
R, F., M. Teslenko, P. V D M, D. L. A, A.
D, S. H, B. L, L. L, M. A. S J. P.
H (): MrBayes .: ecient Bayesian phyloge-
115
Systematics of two lesser known colubrid snakes of Northeast India
netic inference and model choice across a large model space. –
Systematic Biology, : –.
S, W. L. (): Notes on a collection of snakes in the Indi-
an Museum, with descriptions of several new species. – Journal
of Asiatic Society of Bengal, LX, –.
S, S., B. S. K, R. K A. K (): Pterido-
phytic diversity in human-inhabited buer zone of Murlen Na-
tional Park, Mizoram, India. – Check List, : .
S, M. A. (): e Reptilia and Amphibia of the Malay Pen-
insula. – Bulletin of the Raes Museum, : –.
S, M. A. (): e Fauna of British India, Ceylon and Bur-
ma, including the whole of the Indo-Chinese Sub-Region. Rep-
tilia and Amphibia. (Serpentes). – Taylor and Francis, Lon-
don.
S, H. E., L. G, J. L. G, P. L. W J., E. S. Q,
R. M. B, A. C. D, J. L. W B. L. S
(): A new Liopeltis Fitzinger, (Squamata: Colubridae)
from Pulau Tioman, Peninsular Malaysia. – Zootaxa, :
–.
S, F. (): Observations of some Indian and Malayan
Amphibia and Reptilia (abstract). – e Annals and Magazine
of Natural History, : –.
T, W. (): Catalogue of the reptiles of British Birma,
embracing the provinces of Pegu, Martaban, and Tenasserim;
with descriptions of new or little-known species. – Zoological
Journal of the Linnean Society, : –.
T, J. C. (): Contributions to the anatomy of the
Ophidia. – Proceedings of the Zoological Society of London,
–.
U, P., P. F J. H (): e Reptile Database. (
March ). – Electronic database accessable at http://reptile-
database.reptarium.cz.
V, G., D. J. L R. M (): SequenceMatrix:
concatenation soware for the fast assembly of multigene
datasets with character set and codon information. – Cladis-
tics, : –.
W, F. (): Some new Asian snakes. – Journal of the Bombay
Natural History Society, : –.
X, W., W. J. D, T. T. F, W. G, C. Q. L, F. Y, Y. H. W ,
K. J, J. Q. J, H. M. C Y. P. Z (): Her-
petological phylogeographic analyses support a Miocene focal
point of Himalayan upli and biological diversication. – Na-
tional Science Review, : nwaa.
Z, T. Q. K. L (): A new natricine snake of the genus
Amphiesma (Squamata: Colubridae: Natricinae) from the cen-
tral Truong Son, Vietnam. – Zootaxa, : –.
Z, Z., Z. S, S. Q, Y. L, Z. L, Y. W, P. L J. M
(): A new species of the genus Hebius (Squamata: Colu-
bridae: Natricinae) from Hunan Province, China. – Zootaxa,
: –.
Appendix
Specimens examined
Amphiesma stolatum: India: Mizoram: Aizawl District, Sawleng
village, MZMU; India: Mizoram: Kolasib District, Buh-
changphai village, MZMU, , ; India: Mizoram: Cham-
phai District, Champhai vengthar, MZMU; China: Ning Po,
NMBE.
Gongylosoma scriptum: India: Mizoram: Aizawl District: Su-
angpuilawn village, MZMU; India: Mizoram: Mamit District:
Saithah village, MZMU; ailand: Khao Yai, ZMB ; East
India, ZMB .
Herpetoreas xenura: India: Mizoram, MZMU–, ,
, , , , , .
Herpetoreas platyceps: India: Darjeeling, NMW., India:
Kulu Simla, NMW, ., . (syntypes Zamenis hima-
layanus).
Herpetoreas sieboldii: India: Darjeeling, NMW., .,
. .; India: Kulu Simla, NMW, ., .
(syntypes Zamenis himalayanu s) ; Nepal, Kulu valley, ZSM..
Hebius khasiense: India: Mizoram, MZMU, ; India:
Meghalaya, BMNH ...–...; Myanmar: Kachin
State, “Huton, Bhamo District ( miles north-east of Bhamo;
circa , feet; Lat. circa ° .; Long. circa ° .)”, now Hu-
tung, BMNH ... (holotype of Natrix gilhodesi Wall, ),
BMNH ..., BMNH ...–
Hebius cf. venningi: India: Mizoram: Aizawl District, Hmui-
fang Community Reserved Forest, MZMU, , ; India:
Mizoram: Aizawl District, Durtlang locality, MZMU; India:
Mizoram: Aizawl District, Tlangnuam locality, MZMU; In-
dia: Mizoram: Mamit District, Reiek Community Reserved For-
est, MZMU, , , , , , .
Hebius spp.: India: Mizoram: Mamit District, Dampa Tiger Re-
serve, MZMU.
Liopeltis frenatus: India: Mizoram, MZMU–; India:
Meghalaya: Khasi Hills, BMNH ... (holotype); China:
Yunnan: Jangcheng City, DL ...
Liopeltis stoliczkae: India: Mizoram, Aizawl District,
MZMU–, ; India: Mizoram, Lunglei District, .
Sibynophis collaris: India: Mizoram, Aizawl District, Tanhril
locality, MZMU; India: Khasi Hills, BMNH...A,
BMNH...B (syntypes); India: Uttar Pradesh Bimtal,
ZFMK; India: Darjeeling, NMW:; India: Sikkim,
NMW:; Myanmar: Rangoon valley, NMW:, :,
:, :; India: Kategarh, NMW; Vietnam: Annam,
NMW:; China, Taiwan, Suishargo, NMW:, :,
:.
Sibynophis chinensis: China, MNHN-RA .; China:
Szechuan: Panzihua, collection of Ding Li (no catalogue num-
ber), China: no exact locality, MNHN-RA ., China: Up-
per Jang Tschang Tse Kiang BMNH..., China: Ichang, Up-
per Jang Tschang Tse Kiang, BMNH... (holotype); China:
Tianchuan County, Sichuan, NMW; China Beijing, Hairou
ZFMK .
Sibynophis chinensis grahami: China: Jang Tse Kiang, BMNH
... (holotype).
Supplementary data
e following data are available online:
Supplementary Table S1. Newly generated sequences and Gen-
Bank sequences used in this study.
Supplementary Figure S1. Bayesian inference phylogenetic tree of
the genera Herpetoreas and Hebius, based on the mitochondrial
cytochrome b gene.
