Journal of Threatened
Taxa | www.threatenedtaxa.org | 26 March 2026 | 18(3): 28479–28494
ISSN 0974-7907 (Online) | ISSN 0974-7893 (Print)
https://doi.org/10.11609/jott.10309.18.3.28479-28494
#10309 | Received 17 December 2025 | Final received 15 February 2026|
Finally accepted 04 March 2026
New record of two natricine
snakes, Hebius gilhodesi (Wall, 1925) and Herpetoreas davidi Nguyen
et al., 2024 (Reptilia: Squamata: Colubridae), from India
Sourav Dutta 1 , Bitupan Boruah 2
& Abhijit Das 3
1–3 Wildlife Institute of India,
Chandrabani, Dehradun, Uttarakhand 248001, India.
1 souravdutta4048@gmail.com, 2
bitupan.kaz@gmail.com, 3 abhijit@wii.gov.in (corresponding
author)
Abstract: Two natricine snakes, Hebius
gilhodesi and Herpetoreas davidi, are reported for the first time
from India based on mitochondrial DNA and morphological data. This study
extends the northwestern-most distribution of H. gilhodesi by 107 km and
H. davidi by 577 km, both of which were thus far known only from
adjacent Myanmar. Additionally, a detailed description of the hemipenial
morphology of H. gilhodesi is provided.
Keywords: Arunachal Pradesh, keelback, Mizoram,
morphology, Myanmar, natural history, northeastern India, phylogeny, range
extension.
Editor: S.R. Ganesh, Kalinga
Foundation Agumbe, India. Date of publication: 26 March 2026 (online & print)
Citation: Dutta,
S., B. Boruah & A. Das (2026). New record of two natricine snakes, Hebius
gilhodesi (Wall, 1925) and Herpetoreas davidi Nguyen et al., 2024
(Reptilia: Squamata: Colubridae), from India. Journal of Threatened Taxa 18(3): 28479–28494. https://doi.org/10.11609/jott.10309.18.3.28479-28494
Copyright: © Dutta et al. 2026. 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: 1—National Geographic Society Explorer Grant (NGS-74044R-20);
2—SERB-DST (CRG/2018/000790).
Competing interests: The authors declare no competing interests.
Author details: Sourav Dutta is a
researcher at Wildlife Institute of India (WII) with interest in taxonomy of
reptiles. Bitupan Boruah is a PhD scholar and senior
project associate at WII. He is currently working on systematics of amphibians
and reptiles with special focus in Northeast
India. Abhijit Das is a scientist and faculty at WII
involved in teaching, training and research in herpetology.
Author contributions: SD conducted field work, studied specimens and prepared the manuscript
draft. BB conducted field work, did the phylogenetic
analyses and reviewed the manuscript. AD conceptualized the work, conducted
field work and reviewed the manuscript.
Acknowledgements: We thank the National Geographic
Society for the Explorer Grant (NGS-74044R-20) and SERB-DST (CRG/2018/000790)
for financial support. We are thankful to the forest department of Arunachal
Pradesh, Meghalaya and Mizoram for providing research permission (CWL/GEN/355/2021/3178 dated 28 September 2021; WL/FG.31/Technical
Committee/2019, dated 18 July, 2019; Memo No. FWC/Research/1061401-1409 dated
13 July 2022 and Memo No.B.19060/1/2020-CWLW/112 dated 2 February 2020). We
thank Sh. Aduk Paron (former Field Director), Dr Tajum Yomcha (APDF),
Sh. Mayur Variya (Biologist) and all the forest staff of Namdapha Tiger Reserve
for their support. We thank Sh. Harshraj Wathore (DFO), Sh. Bunty Tao (Range
Officer) and Sh. Aditya Das (Biologist), Kamlang Tiger Reserve for their help during field work. We are thankful to Dr G.S. Bhardwaj
(Director), Dr Ruchi Badola (Dean), Wildlife Institute of India, Dehradun for
constant support. Our thanks to Dr Samuel Lalronunga, Dr Santanu Dey, Sh.
Malsawmdawngliana, Sh. Isaac Zosangliana, Sh. K. Lalhmangaiha, Sh.
Naitik G. Patel, Sh. Vijayan Jithin, Sh. Jason D. Gerard, Sh. Vignesh, Sh. Aphu
Yoha Yobin, Sh. Akhida and field assistants for field work.
Introduction
The natricine snake genus Hebius
Thompson, 1913, currently represents over 50 species, widely distributed
throughout eastern and southeastern Asia (Gao et al. 2024; Li et al. 2026; Uetz
et al. 2026). In India, this genus is represented by seven species (Basfore et
al. 2024; Uetz et al. 2026), namely: H. clerki (Wall, 1925); H. khasiensis
(Boulenger, 1890); H. lacrima Purkayastha & David, 2019; H.
modestus (Günther, 1875); H. parallelus (Boulenger, 1890); H.
taronensis (Smith, 1940); and H. venningi (Wall, 1910). Recently,
Bohra et al. (2025) resurrected H. gilhodesi, which was synonymised
under H. khasiensis by Wall (1926). Hebius gilhodesi is currently
known from the type locality, Hutung in Bhamo District and Alangdunhku in Putao
District of Myanmar (Bohra et al. 2025).
Another natricine snake genus Herpetoreas
Günther, 1860, is represented by nine species, distributed across
the foothills of western and eastern Himalaya, including northeastern Pakistan,
northern & northeastern India, Nepal, Bhutan, southwestern China,
Bangladesh, and Myanmar (Ren et al. 2022). In India, five species of this genus
have been reported (Basfore et al. 2024; Nguyen et al. 2024; Uetz et al. 2026),
namely: H. murlen Lalremsanga, Bal, Vogel & Biakzula, 2022; H.
pealii (Sclater, 1891); H. platyceps (Blyth, 1855); H. sieboldii Günther,
1860; and H. xenura (Wall, 1907). Among these five species, H. murlen
and H. pealii are endemic to northeastern India. Recently,
Nguyen et al. (2024) described Herpetoreas davidi from southwestern
Myanmar, which is known only from the type locality Rakhine Yoma Elephant Sanctuary.
During the herpetological surveys
in northeastern India, some unidentified populations of natricine snakes were
encountered that did not fit the definitions of taxa known from India.
Phylogenetic analyses and morphological data suggest two populations from
Arunachal Pradesh are H. gilhodesi and one from Mizoram is H. davidi which
are reported herein as the first record from India.
Materials
and Methods
Sample collection and fixation
Specimens of natricine snakes
were hand-collected from Arunachal Pradesh, Meghalaya, and Mizoram in
northeastern India during 2021 and 2023. Collected samples were euthanized
using Tricaine Methanesulfonate (MS222) and fixed in 3% formalin. Prior to
fixation, liver tissue was collected and stored in molecular-grade ethanol for
DNA extraction. Collected specimens were registered and deposited at the
Wildlife Institute of India, Dehradun.
DNA extraction and phylogenetic
analyses
Genomic DNA was extracted from
liver tissue samples stored in absolute ethanol at –20 °C, using the DNeasy
(Qiagen) blood and tissue kit. Cytochrome b (Cyt-b) gene (~ 1100 base pairs)
was amplified and sequenced using the primers L14910
(5’-GACCTGTGATMTGAAAACCAYCGTTGT-3’) and H16064
(5’-CTTTGGTTTACAAGAACAATGCTTTA-3’) (Burbrink et al. 2000). Polymerase
chain reaction (PCR) condition followed was initial denaturation at 95 °C for 5
min, followed by 35 cycles of denaturation at 95 °C for 45 s, annealing at 54
°C for 45 s, and extension at 72 °C for 55 s. The final extension was at 72 °C for
10 min. Bidirectional sequences were manually checked using CHROMAS v.2.6.6
software (http://technelysium.com.au/wp/chromas) and aligned using ClustalW
(Thompson et al. 1994) with default prior settings implemented in MEGA
v.7 (Kumar et al. 2016). We checked for unexpected stop codons by
translating the sequence to amino acids in MEGA v.7. The new sequences
generated in this study were aligned with 49 sequences downloaded from GenBank
(Benson et al. 2007), and Amphiesma stolatum was used as an
outgroup (Appendix 1). Maximum likelihood (ML) analysis was performed using the
GUI version of the IQ-TREE (Nguyen et al. 2015), implemented in PhyloSuite
(Zhang et al. 2020). The dataset was partitioned by codon position. The
ModelFinder (Kalyaanamoorthy et al. 2017) was used to find the best-fitting
models. The best fit models suggested by
ModelFinder were Partition 1: TN+F+I+G4, Partition 2: HKY+F+I, Partition 3:
TIM+F+G4. We also performed a Bayesian inference (BI) analysis using the
program MrBayes 3.2 (Ronquist et al. 2012) implemented in PhyloSuite (Zhang et
al. 2020). We used PartitionFinder v1.1.1 (Lanfear et al. 2012) with default
settings to find the best-fit model of sequence evolution for the same dataset
used for ML analysis. The best fit models suggested by PartitionFinder were
Partition 1: HKY+I+G, Partition 2: HKY+I, Partition 3: GTR+G. Four separate
runs were set up, each with eight Markov chains, initiated from random trees
and allowed to run for 10 million generations, sampling every 1,000 generations.
Analyses were terminated when the standard deviation of the split frequencies
was less than 0.001, the first 25% of trees were discarded as burn-in, and
trees were constructed using a 50% majority consensus rule. We obtained the ESS
values using TRACER v1.6 (Rambaut et al. 2018) and confirmed greater than 200
for the priors. Support for the internal branches for the ML and BI was
quantified using 10,000 pseudoreplicates (ultrafast bootstrap UFB) and
posterior probability (PP), respectively. The resulting tree was edited in
Figtree v.1.4.4 (Rambaut 2018). Uncorrected pairwise distances (p–distances)
were calculated in MEGA v7.1 with pairwise deletion of missing data and gaps.
Morphological examination
Morphological measurements and
terminologies followed Das et al. (2021) as mentioned in the following:
snout-vent length (SVL); tail length (TL); head length (HL): distance between
posterior edge of last supralabial and tip of the snout; head width (HW): at
angle of jaws; head depth (HD): height at the occipital region; eye diameter
(ED): horizontal diameter; eye to nostril distance (EN): anterior corner of eye
to posterior edge of nostril; eye to snout distance (ES): anterior corner of
eye to tip of snout; interorbital distance (IO): measured at the anterior
corner of orbit. All linear measurements, except SVL and TL, were taken using
Mitutoyo digital callipers (accuracy 0.01 mm). The SVL and TL were measured
using a thread and metal scale. We also counted head and body scales as
follows: SL: number of supralabials; IL: number of infralabials; SL-E: number
of supralabials entering orbit; PreO: numbers of preocular; PostO: number of
postocular; Tmp: number of temporals; PVS: number of preventrals; DSR: dorsal
scales row at one head length behind neck, at midbody, and one head length
before vent; VS: number of ventrals; AN: anal plate divided or entire; SC:
number of subcaudals. Ventral scale counts and hemipenial descriptions followed
Dowling (1951); Dowling & Savage (1960) respectively. Abbreviation used are
WII-ADR: Wildlife Institute of India Abhijit Das
Reptile collection.
Results
Phylogenetic analyses
Both ML and BI analyses yielded
nearly similar tree topologies (Figures 1 & 2). The newly collected materials
(WII-ADR1792, WII-ADR1793, WII-ADR1800, WII-ADR3303, WII-ADR3320) from Namdapha
Tiger Reserve, Gandhigram, and Kamlang Tiger Reserve in Arunachal Pradesh
(Figure 3) clustered with the published sequences of Hebius gilhodesi
from Myanmar. The genetic divergence of these newly collected samples from
northeastern India and samples from Myanmar is 2.2–3.2 % in the Cyt-b gene. One
specimen (WII-ADR1051) collected from Ngengpui Wildlife Sanctuary, Mizoram
(Figure 3) nested with the published type sequences of Herpetoreas davidi
from Myanmar (Figures 1 & 2). The genetic divergence between these two
samples is 0.7% in the Cyt-b gene.
Morphology
The specimens from
Arunachal Pradesh (WII-ADR1792, WII-ADR1793, WII-ADR1800, WII-ADR3303,
WII-ADR3320) (Images 1 & 2) were referred to Hebius gilhodesi based
on the following set of morphological characters: 1) dorsal scale rows 19: 19:
17; 2) scales are moderately keeled; 3) dorsal scale rows reduce from 19–17
between 91st–97th ventrals in males and between 84th–89th
ventrals in females where the third and fourth dorsal scale rows fuse; 4)
ventrals 142–148; 5) subcaudals 71–103; 6) nine supralabials, generally fourth
to sixth supralabials enter the orbit; 7) dorsally dark brown, a very faint,
rusty brown dorsolateral stripe from the neck to the tip of the tail,
interrupted by buff spots.
The specimen (WII-ADR1051) (Image
3) from Mizoram was referred to Herpetoreas davidi based on the
following set of characters: 1) small body size, 490 mm; 2) 19: 19: 17 dorsal
scale rows, scales strongly keeled; 3) ventrals 153; 4) subcaudals undivided;
5) nine supralabials; 6) a white sagittal line just behind the parietal
present; 7) head dorsally rusty brown; 8) dorsum light to dark brown with a
dorsolateral series of white spots from nape to base of the tail; 9) ventral
surface creamish or off white with dark spots along the lateral edge of each
ventral and subcaudal scales.
Thus, this study reports two
Myanmar natricine snakes Hebius gilhodesi and Herpetoreas davidi for
the first time from India (Figure 3) based on phylogenetic and
morphological data. Morphological descriptions of newly collected specimens of
the two species from India are given below.
Hebius gilhodesi (Wall, 1925)
(Table 1; Figure 1–3; Images 1,
2)
Materials examined (n = 5): adult male (WII-ADR1793), adult
female (WII-ADR1792) and subadult female (WII-ADR1800) collected from
Gandhigram (27.26514° N, 96.93704° E, elevation 1,115 m),
Changlang District, Arunachal Pradesh, India on 17–19 September 2022 by Abhijit
Das, Bitupan Boruah, Naitik G. Patel; adult male (WII-ADR3320) collected from
Kamlang Tiger reserve (27.69583° N, 96.44585° E, elevation 1,205
m), Lohit District, Arunachal Pradesh, India on 19 July 2023 by Abhijit Das,
Jason D. Gerard and Rajiv N.V.; juvenile female (WII-ADR3303) collected from
Kamala Valley (27.46127° N, 96.42569° E, elevation 645 m),
Namdapha Tiger Reserve, Changlang District, Arunachal Pradesh, India on 2 June
2023 by Abhijit Das, Sourav Dutta, Jason D. Gerard and Rajiv N.V. (Figure 3).
Description of new Indian
material (Image 1)
A moderate sized snake, SVL
370–410 mm in males and 250–370 mm in females. Body subcylindrical,
widest at midbody, slightly tapering anteriorly and posteriorly; tail
moderately long, TL/SVL= (0.39–0.51) in males and TL/SVL= (0.42–0.51) in
females. Head moderately distinct from neck, longer than wide, HW/HL =
(0.55–0.57) in males and HW/HL= (0.50–0.52) in females; snout nearly rounded,
gradually sloping towards the tip; nostril visible from dorsal aspect; rostral
almost hidden from dorsal aspect, only posterior border is slightly visible;
internasal paired, as long as broad; prefrontal wider than long, laterally
extended towards loreal; frontal posteriorly pointed, longer than broad,
anteriorly wide, slightly longer than the supraocular; supraocular elongated,
widest at posterior part; parietals long, anteriorly wide, posterior margin
extends at the level of more than half of the last supralabial length. In
lateral view tip of snout acute; rostral partially visible; nostril
horizontally elliptical, laterally placed, closer to snout than to orbit; nasal
divided by a vertical slit, posterior nasal larger than anterior one, in
contact with first and second supralabials; loreal region slightly concave; loreal
single, sub-rectangular, dorsally widely contacting with prefrontal, and below
in contact with second and third supralabials; in WII-ADR1793, loreal in
contact with third and fourth supralabials; generally a single preocular
present, but in WII-ADR1792 two preoculars present; preocular vertically
elongated, dorsally wide, below in contact with fourth supralabial, in
WII-ADR1793, preocular in contact with fifth supralabial; eye large with
rounded pupil; orbit semi-circular, orbital diameter less than snout length;
three postoculars, topmost postocular largest and widely connected to parietal, middle postocular connected to
anterior temporal, and third postocular posteriorly connected to anterior
temporal and below in contact with sixth and seventh supralabials, in WII-ADR1793, the bottom postocular in
contact with seventh and eighth supralabials; anterior temporal single but two
in WII-ADR1793; anterior temporal narrow and elongated; two posterior
temporals on each side; 8–10 supralabials, first one smallest, wider than tall;
second to fourth supralabials taller than wide, equal in size, and usually
fourth to sixth supralabials connected to eye, but in WII-ADR1800 and
WII-ADR1793 the third to fifth supralabials and fifth to seventh supralabials
connected to eye on the left side respectively; seventh to ninth supralabials
wider than tall, eighth supralabial largest among all; mental subtriangular,
much wider than long; 10 infralabials on each side, nine on left side of
WII-ADR1800, first to fifth infralabials in contact with first pair of genials,
first pair in midline contact; second pair smallest, second to fourth
infralabials taller than wide, fifth infralabial largest among all; anterior
genials short and elongated; posterior genials long and posteriorly oblique,
anteriorly separated from each other by two small scales and posteriorly separated by a pair of enlarged scales.
Dorsal scales narrow and
posteriorly nearly pointed, arranged in 19: 19: 17 rows; dorsal scales
moderately keeled, except those on first row, which are feebly keeled or nearly
smooth on anterior body; apical pits absent; dorsal scales closer to the vent
comparatively large; all dorsal scale rows on tail moderately keeled; 145–148
ventrals in males and 142–147 in females; 1–3 preventrals, anterior most
preventral separated from posterior pair of genials by one pair of enlarged
scales; subcaudals divided, 71–100 in males and 86–100 in females. Morphometric
and meristic data of the newly collected specimens in this study are provided
in Table 1.
Dorsal scales reduction varies
among the four individuals as follows:
3+4(95) -
WII-ADR3320 (Male): 19 ––––––– 18
–––––––17
- 3+4(97) - 3+4(93)
WII-ADR1793 (Male): 19 –––––– 18
––––––– 17
3+4(91)
-
- 3+4(86)
WII-ADR1792 (Female): 19 –––––––
18 ––––––– 17
3+4(84)
-
- 3+4(89)
WII-ADR1800 (Female): 19 –––––––
18 ––––––– 17
3+4(87)
-
Hemipenis
Organ short, thin and unilobed;
extended up to the level of fourth to fifth subcaudals. The organ with a
slightly extended apical tip on left side of the organ; organ entirely covered
with small spines from base to apical tip on both sulcate and asulcate side;
sulcate side with few enlarged spines at mid-base, and a large basal hook
present at proximal part of truncus; sulcus spermaticus single, shallow (deep
in WII-ADR1793) and simply oblique, extending to inner side of right apical
tip; sulcal lip almost indistinct; apical necked area smooth.
Colouration in preservative
Head on dorsal aspect light
brown, pale rusty brown towards parietals, scattered with small dark spots
throughout (Image 1); laterally cream coloured and posteriorly light brown;
supralabials cream coloured up to sixth scale and bordered with dark brown on
posterior and lower edge of each scale, rest are white on middle and surrounded
by broad dark brown colour; enlarged white spots on supralabials continue to
nape; in WII-ADR1792, these two stripes are disconnected; a short and narrow
faint whitish stripe on posterior dorsal aspect of head. Dorsum on anterior one
third dark greyish, vertebral scales dark greyish, scales towards lateral side
pale reddish-brown with greyish tinge; each dorsal scale scattered with tiny
cream-coloured speckles; a faint dorsolateral stripe along the body,
interrupted by buff spots; tail dorsally dark greyish-brown. Ventral aspect of
head pale cream-coloured, infralabials mottled with dark brown on outer edge;
on ventral side, anterior part of body pale cream-coloured and posteriorly
whitish; tail ventrally whitish; each ventral and subcaudal scales has distinct
or indistinct dark brown spots along the lateral side.
Colouration in life (Image 2)
Nearly the same as in preserved
condition. Head dorsally and laterally light or dark brown, mottled with dark
brown and black throughout; supralabials cream coloured or whitish with dark
edge; on each side, white spots on the supralabials continue to the nape; pupil
entirely dark with light outer margin; iris light golden brown. Dorsum reddish
or dark brown, mottled with dark spots throughout; a very faint, rusty brown
dorsolateral stripe from the neck to the tip of the tail, interrupted by
prominent or obscure buff spots. Each ventral and subcaudal scale has dark spots along the lateral edge.
Natural history and distribution
We recorded H. gilhodesi
along the edge of small streams with shallow water in Gandhigram and at Kamala
Valley in Namdapha Tiger Reserve, and in Kamlang Tiger Reserve, Arunachal
Pradesh, India between 1900–2200 h. The forest type at the locality was
tropical evergreen and semievergreen. The forest floor was covered with ferns
and shrubs. Other reptile species such as Pseudoxenodon macrops, Ptyctolaemus
namdaphaensis, and Sphenomorphus sp. were recorded at that location.
We also observed anurans such as Nasutixalus jerdonii, Raorchestes
orientalis, Gracixalus patkaiensis, Xenophrys sp., and Kurixalus
naso in this locality.
Hepetoreas davidi Nguyen, Lalremsanga,
Biakzuala & Vogel, 2024
(Table 1; Figure 1–3; Image 3)
Material examined (n = 1): adult female (WII-ADR1051)
collected from Ngengpui Wildlife Sanctuary (22.48498° N, 92.75653° E, elevation
215 m), Lawngtlai District, Mizoram, India on 7 September 2021 by Abhijit Das,
Bitupan Boruah, Naitik G. Patel, and Samuel Lalronunga.
Description of new Indian
material (Image 3)
Body and tail subcylindrical,
widest at mid body, substantially narrower towards neck, and slightly tapering
towards vent; tail moderately long (incomplete) (TL/SVL = 0.2*). Head
moderately distinct from neck and longer than broad (HW/HL = 0.4); head widest
at posterior axis of jaw, slightly tapering before mandibular joint; rostral nearly hidden as seen from above,
only posterior border slightly visible; internasal paired, slightly wider than
its height; two prefrontals, laterally extended towards loreal; frontal much
longer than its width, slightly longer than supraocular, anteriorly broad;
parietals elongated, widest at anterior part; in lateral view, parietal region
anteriorly flattened, from frontal to internasal slightly elevated; tip of
snout acute in lateral view; rostral partially visible in lateral aspect;
nostril small and vertically elliptical, laterally oriented, closer to snout
tip than to eye; nasal divided by a vertical slit; loreal region narrowly
concave; a single loreal on each side, sub-rectangular in shape, widely contact
to prefrontal above, and below to second supralabial; single preocular,
vertically elongated, dorsally wide; eye large, with rounded pupil, orbit
horizontally elliptical, eye diameter less than snout length (ED/SL= 0.5);
three postocular on right and two on left, topmost one largest; two anterior
temporal, narrow and elongated, two posterior temporal on right and single on
left; eight supralabials on each side, first six supralabials taller than wide,
seventh and eighth wider than tall, first supralabial smallest and in contact
with nasal, second supralabial connected to post nasal and loreal, third
supralabial connected to preocular and slightly entering to orbit; fourth and
fifth below eye, widely entering to orbit, fifth supralabial posteriorly in
contact with lower postocular; and sixth in contact with second and third
postocular and lower anterior temporal; seventh supralabial largest and widely
connected to lower anterior and posterior temporals; eighth supralabial posteriorly narrow; mental
much wider than long; 10 infralabials on each side, first to fifth in contact
with first pair of genials; first infralabials in mid line contact; second infralabial
smallest; second to fourth taller than the wide; fifth infralabial largest;
anterior genials shorter than posterior paired; posterior genials elongated and
posteriorly oblique.
Dorsal scales strongly keeled,
except the first row, keels are posteriorly serrated, arranged in 19: 19: 17
rows; first dorsal row large and feebly keeled on anterior body, posteriorly
moderately keeled; topmost third and fourth rows on dorsum slightly narrow and
pointed than those on bellow; tail scales strongly keeled; 153 ventral and two
preventrals; anterior most preventral separated from posterior pair of genials
by a pair of small scales; subcaudal scales single, 59 in number (incomplete).
4+5(96) -
Dorsal scale reduction: 19
–––––––– 18 ––––––– 17
- 3+4(98)
Colouration in preservative
Head dorsally rusty brown,
mottled with tiny dark spots, posteriorly darker; supralabials cream-coloured
with irregular dark brown patches; a whitish stripe running behind eye to above
nape on each side and medially connected on nape; an indistinct, narrow, white
streak on posterior part of head; dorsum dark brown; first dorsal scale row
predominantly cream coloured; each dorsal scales marked with tiny cream
coloured spackles; a series of white spots from nape to base of the tail
dorsolaterally present. Ventral surface of head and body cream-coloured;
anterior infralabials with dark brown spots; tail ventrally whitish; ventral
and subcaudals scales with obscure dark spots along lateral aspect.
Natural history and distribution
We recorded the species among the
leaf litter at the edge of a small stream in Ngengpui Wildlife Sanctuary,
Mizoram, India at around 09.30 h. Forest type at the locality was lowland
evergreen forest. We observed Tropidophorus assamensis, Sphenomorphus
sp., Dendrelaphis sp. at the same locality. Anurans such as Limnonectes
khasianus, Ingerana borealis were observed along the stream where
the individual of Hepetoreas davidi was recorded.
Discussion
Northeast India falls under
Tibeto-Yunanese / Indo-Chinese biogeographic zones and is known to have a rich
and unique snake diversity (Das 1996; Das et al. 2009; Malsawmdawngliana et al.
2022; Basfore et al. 2024). In the past as well as in more recent years,
several new records of snakes have been reported in India from the northeastern
region (Captain & Patel 1998; David et al. 2001; Mohapatra et al. 2010;
Biakzuala et al. 2024; Gerard et al. 2024; Ray et al. 2025).
Hebius gilhodesi recently revalidated by Bohra et
al. (2025), and was previously known only from
a few localities in northernmost Myanmar. The present study reported this
species from Namdapha Tiger Reserve and Kamlang Tiger Reserve of Arunachal
Pradesh, India. This record extends the western and northwestern distribution
limit of the species by 103 km and 107 km (areal distance) respectively from
the closest locality of the species in Kachin state, Myanmar. The new locality
records of this species are 280–339 km northwest of the type locality, Huton,
Bhamo District, Myanmar. With this new record, the number of species of the
genus Hebius in India now gets raised to eight.
It is observed that the following
morphological characters in the newly collected specimens of H. gilhodesi which
are incongruent with that of previous literature. According to Wall (1925), the
dorsolateral stripe of H. gilhodesi starts from neck and extends up to
vent, in all the examined specimens this stripe extended to the tip of the
tail. Bohra et al. (2025) mentioned that H. gilhodesi lacks dorsolateral
stripes but has a series of spots whereas in all the examined specimens of H.
gilhodesi, including the subadult, it was observed that light reddish or
rusty brown dorsolateral stripes on which the buff-coloured spots are arranged.
This stripe remains visible even in preserved condition. Notably, Wall (1925)
also stated a reddish dorsolateral stripe is present in all examined specimens.
Furthermore, the holotype of H. gilhodesi depicted in Bohra et al.
(2025) (Figure 11A, page 487) clearly shows the presence of dorsolateral
stripe. Bohra et al. (2025) reported scale reduction involving the fourth and
fifth row which does not correspond to the original description by Wall (1925).
This data of both adult male and female specimens (n = 4) are consistent with
Wall (1925) where the third and fourth rows are involved in the reduction. In
addition to this, the dorsal scale row reduction is observed among our newly
collected materials. The first reduction of dorsal scale rows (19–18) in males
was found at 91st–95th ventral and in females at 84th–87th
ventral. The second reduction (18–17) was found at 93rd–97th
ventral in males and at 86th to 89th ventral in females.
Thus, the reduction in females is more anterior in trunk position compared to
males. However, this reduction is due the merging of third and fourth dorsal
scale rows, which is consistent in all the specimens. This intraspecific
variation of dorsal scale reduction has also been previously recorded in the
species belonging to other Indian colubrid genera such as Ahaetulla and Oligodon
(Deepak et al. 2019; Mirza et al. 2021).
Herpetoreas davidi was originally described from
the Rakhine Yoma Elephant Sanctuary, in Myanmar. In the present study, it is
reported that this species from the southern part of Mizoram, which represents
a range extension of 577 km (areal distance) northwestern from the type
locality. This disjunct distribution could be a result of sampling gap. Further
sampling along Indo-Myanmar border may reveal unknown localities of this
species. With this new record, the number of species of the genus Herpetoreas
increased to six in India. Previously, Nguyen et al. (2025) reported this
species at an elevation range 120–175 m. The present study recorded this
species from Ngengpui Wildlife Sanctuary at an elevation of 215 m, representing
the highest elevation known for the species to date.
The minor morphological
variations mirror the mild genetic divergences noted in Indian vs. Myanmar
specimens of H. gilhodesi. The Indian specimen of H. davidi was
much more congruent with the type series in morphology and genetics.
Ironically, the geographic distances between the known Myanmar distributions
and the new Indian records were lesser for the more divergent H. gilodesi
than H. davidi. It remains to be tested if the sample size being larger
for the Indian material of H. gilodesi than for H. davidi, may be
a reason for this discordance in intra-specific variation patterns. The new
distribution records in recent years from northeastern India, including these
two new records of natricine snakes highlights the urgent need of extensive
faunal surveys in this region.
Table 1. Morphometric and meristic
data of newly collected Hebius gilhodesi and Herpetoreas davidi
from India, with available data. Bilateral characters are given in right and
left order separated by “/”; “n” denotes samples size, “-” indicates data not
provided, and “*” measurement/counts incomplete because of tail missing.
|
Hebius gilhodesi |
Herpetoreas davidi |
|||||||||
|
Voucher specimen |
WII-ADR1793 |
WII-ADR3320 |
WII-ADR1800 |
WII-ADR1792 |
WII-ADR3303 |
Bohra et al. 2025 (n = 15) |
WII-ADR1051 |
Nguyen et al. 2024 (n = 2) |
||
|
Sex |
Male |
Male |
Female |
Female |
Female (juvenile) |
Male |
Female |
Female |
Male |
Female |
|
SVL |
370 |
410 |
250 |
370 |
170 |
288–368 |
156–435 |
490 |
292–384 |
259–482 |
|
TL |
145 |
210 |
105 |
190 |
80 |
152–181 |
68–183 |
140* |
126–168 |
115–178 |
|
HL |
14.80 |
14.50 |
11.30 |
15.80 |
8.90 |
13.4–16.4 |
13.1–17.9 |
21.15 |
- |
- |
|
HW |
8.55 |
8.0 |
5.88 |
8 |
7.70 |
3.2–4.1 |
3.2–4.2 |
10 |
- |
- |
|
HD |
8.60 |
6.10 |
4.68 |
5.85 |
3.50 |
- |
- |
7.46 |
- |
- |
|
ED |
2.56 |
2.60 |
2.18 |
2.61 |
1.85 |
2.2–2.9 |
2.5–2.9 |
3.26 |
- |
- |
|
E-N |
2.52 |
2.45 |
1.78 |
2.52 |
1.47 |
2.3–2.9 |
2.4–3.3 |
3.14 |
- |
- |
|
E-S |
4.45 |
4.25 |
3.10 |
4.20 |
2.50 |
3.2-4.1 |
3.2–4.2 |
5.21 |
- |
- |
|
IN |
2.85 |
2.90 |
2.10 |
2.75 |
1.79 |
- |
- |
3.35 |
- |
- |
|
IO |
4.40 |
4.25 |
3.32 |
4.07 |
2.82 |
- |
- |
5 |
- |
- |
|
PVS |
2 |
2 |
1 |
2 |
3 |
- |
- |
2 |
- |
- |
|
VS |
145 |
148 |
146 |
142 |
147 |
144–153 |
143–151 |
153 |
154–156 |
151–155 |
|
SC |
71, 72 |
99, 100 |
86, 87 |
93, 92 |
102, 103 |
99–111 |
87–98 |
59* |
97–99 |
97–100 |
|
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: 17 |
19: 19: 17 |
|
SL |
10, 9 |
9, 9 |
8, 9 |
9, 9 |
9, 9 |
9, rarely 8 |
9 |
8, 8 |
9 |
9 |
|
SL-E |
5, 6, 7/ 4, 5, 6 |
4, 5, 6/ 4, 5, 6 |
3, 4, 5/ 4, 5, 6 |
4, 5, 6/ 4, 5, 6 |
4, 5, 6/ 4, 5, 6 |
4-6, rarely 4, 5 |
4, 5, 6 |
4, 5/ 3, 4, 5 |
4–6 |
4–6 |
|
IL |
10/10 |
10/10 |
9/ 10 |
10/ 10 |
10/ 10 |
10 |
10, rarely 11 |
10/ 10 |
9 or 10 |
9 or 10 |
|
Tmp |
2+2/ 2+2 |
1+2/1+2 |
1+1 / 1+1 |
1+1/ 1+1 |
1+1/ 1+1 |
1+1, rarely 2+1 |
1+1, rarely 2+1 |
2+2/ 2+1 |
1+2 or 2+2 |
1+2 or 2+2 |
|
PreO |
1/1 |
1/1 |
1/1 |
2/2 |
1/1 |
1, rarely 2 |
1, rarely 2 |
1/1 |
1 |
1 |
|
PostO |
¾ |
3/3 |
3/3 |
3/3 |
3/3 |
|
3, rarely 2 |
3/2 |
3 |
3 |
|
AN |
Divided |
Divided |
Divided |
Divided |
Divided |
Divided |
Divided |
Divided |
Single |
Single |
For
figures & images - - click
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Appendix 1. Details of the DNA
sequences (Cyt-b) used in this study.
|
Species |
Locality |
Voucher no. |
Accession number |
Reference |
|
Hebius vibakari |
Liaoning, China |
GP1345 |
KJ685676 |
Bohra et al. 2025 |
|
Hebius vibakari |
Eastern Asia |
ZCYK Avib1 |
LC640368 |
Bohra et al. 2025 |
|
Hebius sangzhiensis |
Hunan, China |
SYNU 08070350 |
MK340763 |
Bohra et al. 2025 |
|
Hebius maximus |
Sichuan, China |
GP2382 |
KJ685696 |
Bohra et al. 2025 |
|
Hebius pryeri |
Kagoshima, Japan |
NA |
AB989126 |
Bohra et al. 2025 |
|
Hebius concelarus |
Okinawa, Japan |
NA |
AB989268 |
Bohra et al. 2025 |
|
Hebius ishigakiensis |
Okinawa, Japan |
NA |
AB989292 |
Bohra et al. 2025 |
|
Hebius optatum |
Vinh Phuc, Vietnam |
AMNH147155 |
KJ685662 |
Bohra et al. 2025 |
|
Hebius sauteri |
Guangdong, China |
CIB118516 |
OP937178 |
Bohra et al. 2025 |
|
Hebius popei |
Guizhou, China |
GP2386 |
KJ685697 |
Bohra et al. 2025 |
|
Hebius atemporalis |
Guangdong, China |
GP1626 |
KJ685680 |
Bohra et al. 2025 |
|
Hebius atemporalis |
Vietnam |
ZMMU NAP-07877 |
OK315813 |
Bohra et al. 2025 |
|
Hebius bitaeniatus |
Guangxi, China |
GP1940 |
KJ685688 |
Bohra et al. 2025 |
|
Hebius metusia |
Sichuan, China |
GP871 |
KJ685707 |
Bohra et al. 2025 |
|
Hebius craspedogaster |
Guizhou, China |
GP1240 |
KJ685672 |
Bohra et al. 2025 |
|
Hebius johannis |
Guizhou, China |
GP1242 |
KJ685673 |
Nguyen et al. 2024 |
|
Hebius octolineatus |
Yunnan, China |
GP1569 |
KJ685678 |
Nguyen et al. 2024 |
|
Hebius yanbianensis |
China |
LN419 |
OR215499 |
Bohra et al. 2025 |
|
Hebius taronensis |
Myanmar |
GP1618 |
KJ685679 |
Nguyen et al. 2024 |
|
Hebius taronensis |
Myanmar |
CAS224426 |
OK315828 |
Bohra et al. 2025 |
|
Hebius venningi |
Kachin, Myanmar |
CAS233206 |
KJ685670 |
Bohra et al. 2025 |
|
Hebius citrinoventer |
Yingjiang, China |
ANU20230016 |
PP472750 |
Bohra et al. 2025 |
|
Hebius septemlineatus |
China |
KIZ037720 |
MZ570486 |
Bohra et al. 2025 |
|
Hebius bitaeniatus |
Thailand |
AUP-00062 |
OK315816 |
Bohra et al. 2025 |
|
Hebius weixiensis |
Lijiang, Yunnan, China |
HSR19088 |
OQ085074 |
Bohra et al. 2025 |
|
Hebius gilhodesi |
Gandhigram, Changlang,
Arunachal Pradesh, India |
WII-ADR1792 |
PZ160968 |
This Study |
|
Hebius gilhodesi |
Gandhigram, Changlang,
Arunachal Pradesh, India |
WII-ADR1800 |
PZ160970 |
This Study |
|
Hebius gilhodesi |
Gandhigram, Changlang,
Arunachal Pradesh, India |
WII-ADR1793 |
PZ160969 |
This Study |
|
Hebius gilhodesi |
Kamala Valley, Namdapha Tiger
Reserve, Arunachal Pradesh, India |
WII-ADR3303 |
PZ160971 |
This Study |
|
Hebius gilhodesi |
Glaw lake, Kamlang Tiger
Reserve, Arunachal Pradesh, India |
WII-ADR3320 |
PZ160972 |
This Study |
|
Hebius gilhodesi |
Kachin state, Myanmar |
CAS221504 |
KJ685668 |
Bohra et al. 2025 |
|
Hebius gilhodesi |
Kachin state, Myanmar |
CAS221525 |
KJ685669 |
Bohra et al. 2025 |
|
Hebius khasiensis |
Sailam, Aizawl, Mizoram, India |
MZMU2540 |
PQ288048 |
Bohra et al. 2025 |
|
Hebius khasiensis |
Phuldungsei, Dampa Tiger
Reserve, Mizoram, India |
WII-ADR1104 |
PZ160963 |
This Study |
|
Hebius khasiensis |
Mairang, Eastern West Khasi
hills, Meghalaya, India |
MZMU3526 |
PQ288047 |
Bohra et al. 2025 |
|
Hebius boulengeri |
Ha Giang, Vietnam |
AMNH148562 |
KJ685664 |
Bohra et al. 2025 |
|
Hebius boulengeri |
China |
CHS774 |
MK201520 |
Bohra et al. 2025 |
|
Hebius jingdongensis |
Jingdong, China |
CIB 119044 |
OR285310 |
Bohra et al. 2025 |
|
Hebius clerki |
Nujiang, Yunnan, China |
CAS215036 |
KJ685666 |
Bohra et al. 2025 |
|
Hebius modestus |
Yunnan, China |
CAS234262 |
KJ685671 |
Bohra et al. 2025 |
|
Hebius youjiangensis |
Guanxi, China |
HSR22184 |
OQ085073 |
Bohra et al. 2025 |
|
Hebius deschauenseei |
Thailand |
AUP-00182 |
OK315827 |
Bohra et al. 2025 |
|
Hebius chapaensis |
Ha Giang, Vietnam |
AMNH148575 |
KJ685665 |
Bohra et al. 2025 |
|
Hebius annamensis |
Laos |
FMNH 258637 |
OK315812 |
Bohra et al. 2025 |
|
Hebius andreae |
Laos |
R.2017.25 |
MK253674 |
Bohra et al. 2025 |
|
Herpetoreas xenura |
Narpuh Wildlife Sanctuary,
Meghalaya, India |
WII-ADR3185 |
PZ160965 |
This Study |
|
Herpetoreas xenura |
Teirei, Dampa Tiger Reserve,
Mizoram, India |
WII-ADR1158 |
PZ160966 |
This Study |
|
Herpetoreas xenura |
Mizoram, India |
PUCZMX SL1 |
MN993850 |
Nguyen et al. 2024 |
|
Hepetoreas pealli |
Poba Reserve Forest, Arunachal
Pradesh, India |
WII-ADR547 |
MT571586 |
Das et al. 2020 |
|
Hepetoreas davidi |
Gawa, Rakhine, Myanmar |
CAS 220256 |
OK315830 |
Nguyen et al. 2024 |
|
Hepetoreas davidi |
Ngengpui Wildlife Sanctuary,
Mizoram, India |
WII-ADR1051 |
PZ160967 |
This Study |
|
Hepetoreas burbrinki |
Tibet, China |
YBU 071128 |
GQ281781 |
Nguyen et al. 2024 |
|
Hepetoreas murlen |
Mizoram, India |
MZMU2473 |
ON204025 |
Nguyen et al. 2024 |
|
Hepetoreas tpser |
Tibet, China |
CHS849 |
MK201567 |
Nguyen et al. 2024 |
|
Hepetoreas tpser |
Medog, Tibet, China |
CIB107855 |
OM313292 |
Nguyen et al. 2024 |
|
Hepetoreas platyceps |
Uttarakhand |
WII-ADR183 |
MT571587 |
Das et al. 2020 |
|
Hepetoreas platyceps |
Gyirong, Tibet, China |
KIZ YPX26317 |
MW111464 |
Nguyen et al. 2024 |
|
Amphiesma stolatum |
Ha Giang, Vietnam |
CAS215037 |
KJ685667 |
Guo et al. 2014 |