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On a New Species of Xenophrys (Anura: Megophryidae) from Tale Wildlife
Sanctuary, Arunachal Pradesh, India with Comments on the Earlier Erroneous
Report as X. maosonensis (Bourret, 1937)
Abstract
e Maoson horned frog, Xenophrys maosonensis (Bourret, 1937) was originally described as Megophrys longipes
maosonensis from Vietnam having distribution in China. Saikia et al. (2019) reported its occurrence from Arunachal
Pradesh as a rst country record for India with the inclusion of genetic data and morphological descriptions. ey
arrived at the species-specic identity based on the NCBI BLAST search of molecular sequence data of 12S rRNA and
16S rRNA which showed homology search of 95.4 – 97.5% similarity with X. maosonensis and ‘selective phylogeny’.
However, the earlier genetic dierence of ca. 4.6% – 2.5 % (12S and 16S, respectively) with X. maosonensis sensu stricto
vis-à-vis the specimen from Arunachal Pradesh, India; coupled with the huge biogeographic barrier of ca. 1600 kms.
aerial distance between the two localities, makes it very unlikely of such a disjunct distribution of the species. is
warranted a re-evaluation on the specic identity of the specimen from India. Now, based on comprehensive phylogeny
(including 142 sequences) of the subfamily Megophryinae, the earlier erroneously reported X. maosonensis from India
is described here as a species new to science as Xenophrys apatani sp. nov. based on integrative taxonomic approach.
Interestingly now, the proposed new species is exhibiting 4.4% to 5.5% genetic distance with the X. maosonensis sensu
stricto from Vietnam. Additionally, morphological character dierences between the congeneric species reported from
India and of X. maosonensis sensu stricto with the proposed new species is provided. A discussion on the biogeographic
distribution of Xenophrys in India is made.
Keywords: Xenophrys maosonensis, erroneous report, Xenophrys apatani sp. nov., Northeast India, cryptic species,
Asian horned toads.
Rec. zool. Surv. India: Vol. 124 (iS)/21-40, 2024
DOI: 10.26515/rzsi/v124/i1S/2024/172712
ISSN (Online) : 2581-8686
ISSN (Print) : 0375-1511
Introduction
e megophyrid genus Xenophrys Günther, 1864 sensu lato
(henceforth s.l.) has undergone a number of revisions in
recent years. is necessitated a detailed review to maintain
the taxonomic stability of the taxon. Lyu et al. (2023)
thoroughly revised the group and suggested the splitting of
the erstwhile Xenophrys s.l. into 10 genera. Of these, the genus
Xenophrys sensu stricto (henceforth s.s.) is represented by
28 valid species worldwide of which 16 species are recorded
from India (Frost, 2024). Of these 16 species, Xenophrys
maosonensis (Bourret, 1937) was reported from Arunachal
Pradesh as a new Indian record by Saikia et al. (2019). ey
arrived at the species-specic identity based on the NCBI
BLAST search of molecular sequence data of 12S rRNA and
16S rRNA which showed homology search of 95.37–97.52%
similarity with X. maosonensis and ‘selective phylogeny’.
However, Saikia et al. (2019) overlooked an important
point which makes this report of X. maosonensis from India
erroneous. e authors failed to consider the huge geographic
barriers that exist between Mao Son in Vietnam (the type
locality of X. maosonensis) with Tale Wildlife Sanctuary in
Arunachal Pradesh, Northeast India (the collection locality
of Saikia et al., 2019), as well as the expansive ca. 1600 kms.
aerial distance between the two locations. is, coupled
with the earlier genetic dierence of ca. 4.6% – 2.5 % (12S
and 16S, respectively) with X. maosonensis s.s. vis-à-vis the
Bhaskar Saikia1, Bikramjit Sinha1,*, A. Shabnam2, Ilona Jacinta Kharkongor3 and K. P. Dinesh2
1Zoological Survey of India, North Eastern Regional Centre, Shillong 793 003, Meghalaya, India
2Zoological Survey of India, Western Regional Centre, Vidyanagar, Pune-411044, Maharashtra, India
3Zoological Survey of India, Arunachal Pradesh Regional Centre, Senki Valley, Itanagar 791113, Arunachal Pradesh, India
*Author for correspondence : sinhabj@gmail.com
Article Received on: 23 April 2024 Accepted on: 09 June 2024
22 Vol 124(1S) | iS 2024 | www.recordsofzsi.com
On a New Species of Xenophrys (Anura: Megophryidae) from Tale Wildlife Sanctuary, Arunachal Pradesh, India
with Comments on the Earlier Erroneous Report as X. maosonensis (Bourret, 1937)
specimen from Arunachal Pradesh, makes it very unlikely
for such a disjunct distribution of one single species.
Additionally, Mahony et al. (2018) and Lyu et al. (2023)
suggested that multiple potential new species-level taxa are
present in Vietnam and China that have erroneously been
reported as X. maosonensis s.s. previously.
In the light of these observations, a revised and thorough
study of the specimen reported by Saikia et al. (2019)
was undertaken and a robust phylogenetic analysis was
done based on the complete phylogeny of the subfamily
Megophryinae involving the 142 sequences of mt 16S rRNA
(Supplementary Table S1). is has resulted in the earlier
report of X. maosonensis from India as erroneous and the
specimen from Tale Wildlife Sanctuary in Arunachal Pradesh
is found to belong to an unnamed species of Xenophrys,
which is described herein as a species new to science and
named as Xenophrys apatani sp. nov. based on integrated
taxonomic approach. Interestingly, Xenophrys apatani sp.
nov. is not directly related to any extant species of Xenophrys
from India; but phylogenetically closely related to Xenophrys
mangshanensis (Fei and Ye, 1990) found in China and X.
maosonensis s.s. found in Vietnam and China. Interestingly
the new species is exhibiting 4.4% to 5.5% genetic distance
with X. maosonensis s.s.
Besides the description of this new species, we have also
made the rst comprehensive phylogenetic tree of all the
Indian Xenophrys species which will aid future researchers
for easy comprehension of the genus in India.
Materials and Methods:
e Xenophrys specimen (Museum voucher No. V/A/
NERC/1370) (Fig. 1 – 2) was collected on 5 November, 2015,
between 17:00 to 20:00 hrs. from Pange [27.5485 N; 93.8989
E; 1804 m] in Tale Wildlife Sanctuary, located in the Lower
Subansiri district of Arunachal Pradesh, India (Fig. 3). e
measurements, DNA extraction and amplication are as per
Saikia et al. (2019). e sequence (MH423746.1) for the new
species was published in Saikia et al. (2019).
All measurements (in millimeters) were done with a
MitutoyoTM digital caliper. e measurements used are
SVL (Snout Vent length), HL (Head Length), HW (Head
Width), MN (Mandible Nostril distance), MFE (Distance of
mandible to the front of eye), MBE (Distance of mandible to
the back of eye), IFE (Distance between the front of eyes),
IBE (Distance between the back of eyes), IN (inter-narial
distance), EN (Distance from eye to nostril), SN (Distance
from snout tip to nostril), EL (Eye length), SL (Snout length),
TYD (Horizontal Tympanum Diameter), TYE (distance
from anterior border of tympanum to the posterior orbital
border), IUE (Minimum distance between the upper
eyelids), UEW (maximum width of the upper eyelid), HAL
(Hand length from the base of inner metacarpal tubercle to
the tip of third nger), FLL ( Forearm Length from the elbow
joint to base of the inner metacarpal tubercle), FL (Femur
Length), TL (Tibia Length), TFOL (Length of tarsus and
foot), FOL (Foot length, from inner metatarsal tubercle to
the tip of fourth toe).
For morphological comparisons, the following literatures
were consulted: Bourret (1937); Mahony et al. (2011, 2013,
2018, 2020); Lyu et al. (2023).
A total of 142 sequences were downloaded including our
sequence of interest (MH423746.1) from the NCBI GenBank
(Supplementary Table S1). Sequence alignment and editing
was performed in MEGA 11 (Tamura et al., 2021). Partition
nder v1.0.1 (Lanfear et al., 2012) was used to estimate the
best partition schemes. GTR+G+I nucleotide substitution
model was the best model for the given dataset according
to Bayesian Information Criterion (BIC) using greedy
algorithm. Bayesian analysis was executed using MrBayes
v 3.2. Bayesian analysis was performed in two independent
runs for 5 × 107 generations with four Markov Chains Monte
Carlo (MCMC) chains sampled every 1,000 generations.
Convergence was assessed using Tracer v1.5, rst 25% of
the generations were discarded as burn-in, and the output
tree was visualised in Figure Tree v1.4 treating members of
Leptobrachella as outgroup following Lyu et al. (2023) (Fig.
4).
Result:
Molecular analysis:
At present, 10 genera are considered valid under the
subfamily Megophryinae (Frost, 2024). In our nal single
gene tree for the mt 16S rRNA based on 142 sequences
provided multiple sub clades of Xenophrys with a larger
monophyletic Xenophrys Clade (Fig.4). All the ten genera
were represented in the recovered tree with absolute
monophyly for the genera Grillitschia, Megophrys,
Ophryophryne, Boulenophrys, Pelobatrachus, Sarawakiphrys,
Brachytarsophrys, Atympanophrys and Xenophrys. e genus
Jingophrys showed paraphyly with low bootstrap support.
23
Zoological Survey of India
Bhaskar Saikia, Bikramjit Sinha, A. Shabnam, Ilona Jacinta Kharkongor and K. P. Dinesh
Among the earlier 18 species of Xenophrys s.l. reported for
India (Dinesh et al., 2023), 16 species were recovered under
the genus Xenophrys and two species were recovered under
the genus Jingophrys.
In the larger phylogenetic tree for the subfamily Megophry-
inae, the new lineage is recovered as a sister clade to an un-
known lineage of Xenophrys sp. (KY022213.1) from Viet-
nam, Xenophrys mangshanensis from China and Xenophrys
maosonensis having distribution from China and Vietnam.
Interestingly genetic distance between the new lineage and
the unknown lineage of Xenophrys sp. (KY022213.1) from
Vietnam is 6.2%; Xenophrys mangshanensis from China is
5.8% and Xenophrys maosonensis having distribution from
China and Vietnam is ranging from 4.4% to 5.5%. is con-
rms the wrong identity of the Xenophrys lineage earlier
identied as Xenophrys maosonensis from India by Saikia
et al. (2019). Accordingly, the new lineages of Xenophrys
(MH423746.1) from Tale Wildlife Sanctuary, Arunachal
Pradesh is described herewith as a new species.
New Species Description:
Xenophrys apatani sp. nov.
(Apatani Horned Toad)
(Fig. 1 – 2)
(urn:lsid:zoobank.org:act:E9B4A8D7-399D-4053-B1BF-
0B534B10FD86)
Chresonym: Megophrys maosonensis – Saikia et al., 2019
Holotype: V/A/NERC/1370, adult male (Fig. 1 – 2) collected
from Pange [27.5485 N; 93.8989 E; 1804 m], Tale Wildlife
Sanctuary (WLS), Lower Subansiri district, Arunachal
Pradesh, India on 5 November, 2015 between 17:00 to 20:00
hrs. by Bikramjit Sinha.
Diagnosis: Xenophrys apatani sp. nov. belongs to Xenophrys
due to the following characters vide Lyu et al. (2023): medium
sized frog, body moderate, head depressed, vomerine
and maxillary teeth present, palpebral tubercles medially
bumped, single discontinuous dorso-lateral ridge extending
from shoulder to groin, no ventro-lateral tubercles.
Genetic Relationship: e new species is exhibiting sister
relationship to clade comprising an unknown lineage of
Xenophrys (KY022213.1) from Vietnam with 6.2% genetic
distance; Xenophrys mangshanensis from China with
5.8% genetic distance and Xenophrys maosonensis having
distribution from China and Vietnam with genetic distance
range 4.4% to 5.5%.
Description of the Holotype (V/A/NERC/1370): A medium
sized adult male frog (SVL-54.0 mm; see Table 1); body
slender (Fig. 1 – 2). Head moderately large, broader than
long; snout pointed, obtusely protruding beyond mandible
in lateral view. Snout longer than the upper eyelid width
and inter-orbital distance. Loreal region concave. Canthus
rostralis angular, snout slightly concave. Nostril closer
to eye. Maxillary teeth present. Vomerine teeth present
located on the inner side of choanae. Tongue entire and
rounded. Tympanum, oval and obliquely oriented; upper
margin concealed by the supra-tympanic fold. Tympanum is
separated from the eye by a distance larger than its horizontal
diameter. Eye diameter twice the diameter of tympanum,
but shorter than the snout length. Supra-tympanic fold
prominent. Pupil vertical. Upper eye-lid with weak palpebral
tubercle. A pair of tubercles on the dorso-lateral sides,
extended only to half of the dorsum. A V-shaped tubercle
on the occiput.
Forelimb weak, ngers free, with relative length I<II=IV<III;
I slightly shorter than II. Hindlimbs slender; tibia longer
than femur and foot. Metatarsal tubercles weakly dened.
Webbing rudimentary.
Colouration of the Holotype: (in life) Dorsum darker with
extensive mottling of a lighter shade; dark triangular patch
between the eyes of which the central region is lighter. A
dark blotch on the snout. Limbs banded. Ventrum light in
colour; anteriorly speckled with dark markings (Fig. 1 – 2).
Condition of the type: e type has lost all its colouration in
the preserved condition (Fig. 5). ere has been a marked
change in the colouration of the type since Saikia et al.
(2019). Ventral incision was made to extract liver tissue in
the specimen and was used in the molecular study, and to
determine the sex of the specimen.
Natural history: e single specimen of Xenophrys apatani
sp. nov. was collected from a water tank, ca. ½ km. from the
forest guard camp of Pange inside the Tale WLS in the aer-
dusk hours (Fig. 2). A number of associated herpetofauna
were observed from the collection locality such as
Leptobrachium bompu, Rhacophorus rhodopus, Zhangixalus
burmanus, Ptyas nigromarginata, Sphenomorphus maculates
and Japalura sp. e vegetation of the type locality is
characterised by thick moist mixed tropical evergreen forests
(Fig. 6).
Distribution: Only known from the type locality of Pange in
Tale WLS, Arunachal Pradesh state of India.
24 Vol 124(1S) | iS 2024 | www.recordsofzsi.com
Comparisons: Genetically Xenophrys apatani sp. nov. is not
closely related to any of the extant species of Xenophrys from
India; the closest ones are being X. maosonensis s.s. having
distribution from China and Vietnam with a genetic distance
range 4.4% to 5.5% and X. mangshanensis from China with
5.8% genetic distance. Morphologically, Xenophrys apatani
sp. nov. diers from X. mangshanensis (in parentheses) in
having smaller male SVL (vs. larger), relative nger length
I<II=IV<III (vs. II<I<IV<III), rudimentary toe webbing (vs.
absent), dark blotch on the snout (vs. absent), distribution
in India (vs. China); X. maosonensis s.s. (in parentheses)
in having smaller male SVL (vs. larger), head broader (vs.
longer), relative nger length I<II=IV<III (vs. I<II<IV<III),
dark blotch on the snout (vs. absent), distribution in India
(vs. Vietnam and China).
For the sake of convenience, the new species is compared
with the other species of Xenophrys found in India.
Xenophrys apatani sp. nov. diers from X. himalayana (in
parentheses) in having smaller male SVL (vs. larger), head
broader (vs. as broad as long), snout pointed (vs. bluntly
pointed), eye diameter twice the tympanic diameter (vs.
more than twice), relative nger length I<II=IV<III (vs.
IV<II<I<III), dark blotch on the snout (vs. absent); from
X. periosa (in parentheses) in having smaller male SVL (vs.
larger), snout pointed (vs. bluntly pointed), upper eyelid
width almost equal to inter-orbital distance (vs. upper eyelid
width less than inter-orbital width), relative nger length
I<II=IV<III (vs. IV<II<I<III), femur length less than tibia
length (vs. equal), femur length shorter than foot length
(vs. longer), dark blotch on the snout (vs. absent); from
X. major (in parentheses) in having smaller male SVL (vs.
larger), head broader (vs. longer), snout pointed (vs. bluntly
pointed), eye diameter twice the tympanic diameter (vs.
more than 2.5 times), internarial distance greater than inter-
orbital distance (vs. equal), relative nger length I<II=IV<III
(vs. IV<II=I<III), femur length shorter than foot length
(vs. longer), inner metatarsal tubercle indistinct (vs.
distinct), dark blotch on the snout (vs. absent); X. robusta
(in parentheses) in having smaller male SVL (vs. larger),
eye diameter twice the tympanic diameter (vs. less the
twice), eye-tympanum distance more than 1.5 times of the
tympanum diameter (vs. slightly larger), internarial distance
greater than inter-orbital distance (vs. equal), relative nger
length I<II=IV<III (vs. IV<I=II<III), dark blotch on the
snout (vs. absent); from X. monticola (in parentheses) in
having larger male SVL (vs. smaller), head broader (vs. as
broad as long), snout pointed (vs. rounded), eye diameter less
the snout length (vs. eye diameter longer than snout length),
eye-tympanum distance more than tympanum diameter
(vs. less), nostril near to eye (vs. equidistant), relative nger
length I<II=IV<III (vs. I<II<IV<III), dark blotch on the
snout (vs. absent); X. avipunctata (in parentheses) in having
smaller male SVL (vs. larger), snout pointed (vs. bluntly
pointed), eye diameter twice the tympanum diameter (vs.
2.5 times), inter-orbital distance larger then upper eyelid
width (vs. smaller), relative nger length I<II=IV<III (vs.
IV<I=II<III), femur length shorter than foot length (vs.
longer); X. oreocrypta (in parentheses) in having smaller
SVL (vs. larger), snout pointed (vs. bluntly pointed), relative
nger length I<II=IV<III (vs. IV<II<I<III), femur length
shorter than foot length (vs. longer), dark blotch on the snout
(vs. absent); from X. awuh (in parentheses) in having larger
male SVL (vs. smaller), head broader (vs. as broad as long),
snout pointed (vs. bluntly pointed), snout length larger than
eye diameter (vs. equal), eye-tympanum distance larger than
tympanum diameter (vs. shorter), upper border of tympanum
concealed by supra-tympanic fold (vs. not concealed),
internarial distance larger than inter-orbital distance (vs.
equal), relative nger length I<II=IV<III (vs. I=II<IV<III),
dark blotch on the snout (vs. absent); from X. dzukou (in
parentheses) in having larger male SVL (vs. smaller), head
broader (vs. longer), snout pointed (vs. bluntly pointed),
eye diameter twice the tympanum diameter (vs. more than
twice), tympanum obliquely elongated (vs. circular), upper
border of tympanum concealed by supra-tympanic fold
(vs. not concealed), eye-tympanum distance larger than
tympanum diameter (vs. equal), inter-orbital space greater
than upper eyelid width (vs. lesser), relative nger length
I<II=IV<III (vs. I=II<IV<III), a single dark blotch on the
snout (vs. multiple small dots); from X. numhbumaeng (in
parentheses) in having larger male SVL (vs. smaller), head
broader (vs. longer), snout pointed (vs. bluntly pointed),
eye diameter twice the tympanic diameter (vs. more than
twice), eye diameter less than snout length (vs. longer), eye-
tympanum distance larger than tympanum diameter (vs.
less), tympanum obliquely elongated (vs. circular), relative
nger length I<II=IV<III (vs. I=II<IV<III), femur length
shorter than foot length (vs. longer), dark blotch on the
snout (vs. absent); from X. serchhipii (in parentheses) in
having larger male SVL (vs. smaller), head broader (vs. as
broad as long), snout pointed (vs. rounded), eye-tympanum
distance larger than tympanum diameter (vs. equal), relative
nger length I<II=IV<III (vs. IV<I=II<III), femur length
less than tibia length (vs. equal), femur length shorter
than foot length (vs. longer); from X. zunhebotoensis (in
parentheses) in having larger male SVL (vs. smaller), head
broader (vs. as broad as long), snout pointed (vs. rounded),
eye diameter twice the tympanic diameter (vs. less than
On a New Species of Xenophrys (Anura: Megophryidae) from Tale Wildlife Sanctuary, Arunachal Pradesh, India
with Comments on the Earlier Erroneous Report as X. maosonensis (Bourret, 1937)
25
Zoological Survey of India
twice), eye diameter less than snout length (vs. longer), eye-
tympanum distance larger than tympanum diameter (vs. half
of tympanum diameter), tympanum obliquely elongated (vs.
circular), upper border of tympanum concealed by supra-
tympanic fold (vs. not concealed), relative nger length
I<II=IV<III (vs. I=II<IV<III), femur length less than tibia
length (vs. equal), femur length shorter than foot length (vs.
longer), dark blotch on the snout (vs. absent); from X. ancrae
(in parentheses) in having larger male SVL (vs. smaller),
head broader (vs. longer), snout pointed (vs. rounded), eye-
tympanum distance larger than tympanum diameter (vs.
equal), internarial distance larger than upper eyelid width (vs.
equal), relative nger length I<II=IV<III (vs. I<II<IV<III),
femur length shorter than foot length (vs. longer); from
X. oropedion (in parentheses) in having larger male SVL
(vs. smaller), head broader (vs. as broad as long), snout
pointed (vs. rounded), eye diameter twice the tympanum
diameter (vs. less than twice), eye-tympanum distance larger
than tympanum diameter (vs. less), tympanum obliquely
elongated (vs. slightly oval), upper border of tympanum
concealed by supra-tympanic fold (vs. not concealed),
nostril nearer to eye (vs. equidistant), internarial distance
larger than upper eyelid width (vs. equal), relative nger
length I<II=IV<III (vs. I=II=IV<III); from X. megacephala
(in parentheses) in having pointed snout (vs. rounded),
eye-tympanum distance distinctly longer than tympanum
diameter (vs. slightly longer), tympanum obliquely elongated
(vs. distinctly oval), nostril nearer to eye (vs. equidistant),
internarial distance larger than inter-orbital distance (vs.
less), upper eyelid width less than inter-orbital distance (vs.
equal), relative nger length I<II=IV<III (vs. IV<II<I<III),
femur length shorter than tibia length and foot length (vs.
longer).
Etymology: e specic name is named aer the Apatani
Tribe of Arunachal Pradesh, which predominantly reside
in the Lower Subansiri Valley and in recognition of their
ingenuity in conservation of wild ora and fauna.
Suggested Common Name: We are suggesting Apatani
Horned Frog as the common name for the proposed new
species.
Discussion:
Until the revisions of the Xenophrys s.l. population of India
by Mahony et al. (2018, 2020), the erroneous reports of
many Chinese species into India persisted over the years
(such as Boulenophrys boettgeri, X. glandulosa, etc.). Also,
the past reports of small-sized Xenophrys s.l. from India
were mostly attributed to X. parva, before they were split
into multiple new species, and the subsequent removal of X.
parva from the fauna of India. With this paper, we are also
correcting the erroneous report of X. maosonensis s.s. from
India by elevating the population attributed to this species
from Arunachal Pradesh, India (Saikia et al., 2019) as a new
species, X. apatani sp. nov.
Prior to the erection of Jingophrys genus, the Indian
population of Xenophrys s.l. was represented by 18 species.
Post the division of the erstwhile Xenophrys s.l. into 10
genera, the present Indian Xenophrys s.s. is represented by 16
species (including the erroneous report of X. maosonensis),
and Jingophrys by two species. With the description of this
new species, the total diversity of Xenophrys s.s. in India
remains the same.
Northeast India is located at the intersection of the Eastern
Himalayas and the Indo-Burma biodiversity hotspots and
the geographic distribution of all the Indian extant species of
Xenophrys s.s. clearly delineate between these two hotspots.
e diversity of Xenophrys s.s. in the Himalayan region is
relatively limited, encompassing six species, namely, X.
himalayana, X. periosa, X. robusta, X. monticola, X. ancrae
and Xenophrys apatani sp. nov. In contrast, the Indo-Burma
hotspot portion of India hosts a more extensive diversity
of Xenophrys s.s. including X. avipunctata, X. oreocrypta,
X. major, X. dzukou, X. numhbumaeng, X. serchhipii, X.
zunhebotoensis, X. oropedion, X. megacephala and X. awuh.
Notably, both the hotspots harbour three larger-sized
species of Xenophrys s.s. (SVL > 60 mm): X. himalayana,
X. periosa and X. robusta in the Eastern Himalayas, and X.
avipunctata, X. oreocrypta and X. major in the Indo-Burma
region. However, it is intriguing that most of the smaller-
sized Xenophrys s.s. species in India are primarily found in
the Indo-Burma biodiversity hotspot, with seven species;
while the Eastern Himalayas hotspot has only three species,
including the newly described species in this study.
Arunachal Pradesh, situated in the easternmost part of India
within the Eastern Himalayas, has seen a notable increase
in the discovery of new species of amphibians since the
beginning of the new century (Mathew and Sen, 2009; Sondhi
and Ohler, 2011; Kamei et al., 2013; Mahony et al., 2013,
2018; Saikia et al., 2017, 2022a and b; Saikia and Sinha, 2019;
Patel et al., 2021; Boruah et al., 2023a, b and c). erefore, it is
highly likely that many more unnamed species, particularly
smaller-sized Xenophrys s.s., are awaiting discovery in this
bio-diverse state of India.
Bhaskar Saikia, Bikramjit Sinha, A. Shabnam, Ilona Jacinta Kharkongor and K. P. Dinesh
26 Vol 124(1S) | iS 2024 | www.recordsofzsi.com
is research paper presents the rst comprehensive
phylogenetic tree incorporating all Indian species of
Xenophrys (16 species) and Jingophrys (2 species), thereby
validating the reliability of genetic data for future integrative
taxonomic approach. Furthermore, it claries that the
earlier erroneous report of Xenophrys maosonensis from
India (Saikia et al., 2019) is rather an unnamed species of
Xenophrys which is now formally named as Xenophrys
apatani sp. nov.
Zoobank Registration
Article: urn:lsid:zoobank.org:pub:B4C7BEBF-1FD1-4445-
B815-8B590BDF9479
Species: urn:lsid:zoobank.org:act:E9B4A8D7-399D-4053-
B1BF-0B534B10FD86
TABLE:
Tabl e 1 : Comparative morphmetrics of Xenophrys apatani sp. nov., X. maosonensis and X. mangshanensis vide Saikia et al.
(2019), Lyu et al. (2023) and Huang et al. (1998)
Characters Xenophrys apatani
sp. nov.
Xenophrys
maosonensis
Xenophrys
mangshanensis
Sex ♂ ♂ ♀ ♀ ♂
SVL 54.0 66.2 76.9 73.0 62.5
HL 16.8 26.1 29.7 27.3 22.1
HW 22.6 24.0 29.4 28.4 22.0
MN 14.1 - - - -
MFE 11.7 - - - -
MBE 6.3 - - - -
IFE 11.1 - - - -
IBE 16.7 - - - -
IN 7.2 7.5 9.5 - -
EN 3.0 - - - -
EL (horizontal) 6.0 8.3 9.0 9.6 8.0
SN 4.3 - - - -
SL 6.9 8.9 9.9 10.0 7.5
TYD 3.0 3.9 4.2 - -
TYE 4.8 4.6 5.4 - -
IUE 5.5 5.7 7.3 8.9 7.7
UEW 5.2 - - 7.2 5.8
HAL 13.7 17.0 18.6 18.9 16.3
FLL 12.7 13.1 16.3 - -
FL 28.0 - - - -
TL 32.5 31.5 42.7 39.7 32.4
TFOL 44.1 44.4 54.9 53.2 43.0
FOL 28.3 - - 33.6 27.7
On a New Species of Xenophrys (Anura: Megophryidae) from Tale Wildlife Sanctuary, Arunachal Pradesh, India
with Comments on the Earlier Erroneous Report as X. maosonensis (Bourret, 1937)
27
Zoological Survey of India
Fig. 1: Live photograph of the holotype of Xenophrys apatani sp. nov.
FIGURES:
Fig. 2: In-situ photograph of the holotype of Xenophrys apatani sp. nov. from the type locality
Bhaskar Saikia, Bikramjit Sinha, A. Shabnam, Ilona Jacinta Kharkongor and K. P. Dinesh
28 Vol 124(1S) | iS 2024 | www.recordsofzsi.com
Fig. 3: A map showing the type locality of Xenophrys apatani sp. nov. (red circle) in
India and that of X. maosonensis (green circle) in Vietnam and distribution range
(yellow shaded) of the later in South East Asia.
Fig. 4: Bayesian tree for the members of Megophryinae including all Indian Xenophrys
based on 542 bp of mt 16s rRNA. Node values represent Bayesian Posterior Probabilities
(BPP).
On a New Species of Xenophrys (Anura: Megophryidae) from Tale Wildlife Sanctuary, Arunachal Pradesh, India
with Comments on the Earlier Erroneous Report as X. maosonensis (Bourret, 1937)
29
Zoological Survey of India
Fig. 5: Holotype image of Xenophrys apatani sp. nov. in preserved condition (A: dorsal view; B: ventral
view).
Fig. 6: Habitat photograph of Pange, Tale WLS, the type locality of Xenophrys apatani sp. nov.
Bhaskar Saikia, Bikramjit Sinha, A. Shabnam, Ilona Jacinta Kharkongor and K. P. Dinesh
30 Vol 124(1S) | iS 2024 | www.recordsofzsi.com
SUPPLEMENTARY TABLE:
S1: Sequence details for mt 16S gene (542bps) used in Bayesian phylogenetic analysis. Sequence names are followed aer Lyu et al. (2023)
Sr.
No Accession nu. Names used in this study Species name as per NCBI Voucher ID Locality References as per NCBI
1 MH423746.1 Xenophrys apatani sp. nov. X. maosonensis NE 3 India Saika et al. (2019)
2 KY022213.1 Xenophrys sps. X. maosonensis AMNH 169335 Viet Nam: ua Tien Hue, A Luoi District Mahony et al. (2017)
3 KJ560372 Xenophrys mangshanensis Xenophrys mangshanensis SYS a002025 China, Guangdong, Mt Nankun/Mt.
Nanling Wan g et al. (2014)
4 OQ181002.1 X. maosonensis X. maosonensis SYS a008766 China Lyu et al. (2023)
5 OQ180999.1 X. maosonensis X. maosonensis SYS a008747 China Lyu et al. (2023)
6 OQ181001.1 X. maosonensis X. maosonensis SYS a008749 China Lyu et al. (2023)
7 OQ181000.1 X. maosonensis X. maosonensis SYS_a008748 China Lyu et al. (2023)
8 KX811782.1 X. maosonensis X. maosonensis ROM35558 China Chen et al. (2017)
9 KX811780.1 X. maosonensis X. maosonensis KIZ016045 China Chen et al. (2017)
10 ON146205.1 X. maosonensis X. maosonensis IEBRA4969 Veitnam: Truong Son Range Luong et al. (2022)
11 KX811785.1 X. maosonensis X. maosonensis KIZ048765 China Chen et al. (2017)
12 ON146206.1 X. maosonensis X. maosonensis IEBRA4968 Veitnam: Truong Son Range Luong et al. (2022)
13 KX811783.1 X. maosonensis X. maosonensis ROM6930 China Chen et al. (2017)
14 KX811781.1 X. maosonensis X. maosonensis ROM26311 China Chen et al. (2017)
15 KX811787.1 X. maosonensis X. maosonensis KIZ048900 China Chen et al. (2017)
16 OQ180991.1 X. maosonensis X. maosonensis SYS_a004527 China Lyu et al. (2023)
17 ON146204.1 X. maosonensis X. maosonensis IEBRA4970 Veitnam: Truong Son Range Luong et al. (2022)
18 ON146203.1 X. maosonensis X. maosonensis IEBRA4971 Veitnam: Truong Son Range Luong et al. (2022)
19 KX811784.1 X. maosonensis X. maosonensis ROM16679 China Chen et al. (2017)
20 KX811786.1 X. maosonensis X. maosonensis SDBDU
2000.2935 China Chen et al. (2017)
21 KR828086.1 X. maosonensis X. maosonensis ROM30830 Viet Nam: Vinh Phuc, Tam Dao Grosjean et al. (2015)
22 MT412390.1 X. maosonensis X. maosonensis VNMN010904 Veitnam: Hoang Lien Range Tapley et al. (2020)
On a New Species of Xenophrys (Anura: Megophryidae) from Tale Wildlife Sanctuary, Arunachal Pradesh, India
with Comments on the Earlier Erroneous Report as X. maosonensis (Bourret, 1937)
31
Zoological Survey of India
Sr.
No Accession nu. Names used in this study Species name as per NCBI Voucher ID Locality References as per NCBI
23 KY022307.1 Xenophrys avipunctata Megophrys major SDBDU
2009.297 India, Meghalaya, East Khasi Hills Mahony et al. (2018)
24 MH406754.1 Xenophrys glandulosa Megophrys glandulosa SYS a003757 China, Yunnan, Mt Gaoligong Liu et al. (2018)
25 MH406725.1 Xenophrys medogensis Megophrys medogensis SYS a002932 China, Xizang, Medog Liu et al. (2018)
26 KY022314.1 Xenophrys robusta Megophrys robusta SDBDU
2011.1057 India, West Bengal, Darjeeling Mahony et al. (2018)
27 KY022309.1 Xenophrys periosa Megophrys major SDBDU
2009.1243 India: Arunachal Pradesh Mahony et al. (2018)
28 KY022311.1 Xenophrys himalayana Megophrys major SDBDU
2009.750 India: Arunachal Pradesh Mahony et al. (2017)
29 KY022308.1 Xenophrys major Megophrys major CES 18903 India, Manipur,Tamenglong Mahony et al. (2017)
30 MH647510 .1 Xenophrys monticola Megophrys montana SDBDU
2011.420 India, West Bengal, Darjeeling Mahony et al. (2018)
31 KX811765.1 Xenophrys zhangi Megophrys zhangi KIZ 014278 China, Xizang, Nyalam Chen et al. (2017)
32 KY022306.1 Xenophrys oreocrypta Megophrys major SDBDU
2009.1104 India, Meghalaya, West Garo Hills Mahony et al. (2017)
33 KY022212.1 Xenophrys sps. Megophrys maosonensis AMNH 168679 Viet Nam: Lao Cai, Van Ban, Nam a
Commune Mahony et al. (2017)
34 MH406728.1 Xenophrys lancangica Megophrys major SYS a002961 China, Yunnan, Mengla; Vietnam, Lao
Cai, Van Ban Liu et al. (2018)
35 KY022318.1 Xenophrys ancrae Megophrys ancrae SDBDU
2009.727 India, Arunachal, Changlang Mahony et al. (2018)
36 KY022320.1 Xenophrys awuh Megophrys sp. 3 SDBDU
2007.111 India, Nagaland, Kohima Mahony et al. (2017)
37 KY022322.1 Xenophrys zunhebotoensis Xenophrys zunhebotoensis CES 19909 India, Nagaland, Zunheboto Mahony et al. (2017)
38 KY022323.1 Xenophrys serchhipii Xenophrys serchhipii SDBDU
2009.612 India, Tripura, North Tripura Mahony et al. (2018)
39 KY022316 .1 Xenophrys numhbumaeng Megophrys oropedion SDBDU
2007.041 India, Manipur, Tamenglong Mahony et al. (2017)
Bhaskar Saikia, Bikramjit Sinha, A. Shabnam, Ilona Jacinta Kharkongor and K. P. Dinesh
32 Vol 124(1S) | iS 2024 | www.recordsofzsi.com
Sr.
No Accession nu. Names used in this study Species name as per NCBI Voucher ID Locality References as per NCBI
40 KY022317.1 Xenophrys oropedion Megophrys oropedion SDBDU
2009.299 India, Meghalaya, East Khasi Hills Mahony et al. (2018)
41 KY022315.1 Xenophrys megacephala Megophrys megacephala ZSI A 11213 India, Meghalaya, Ri Bhoi Mahony et al. (2018)
42 KY022324.1 Xenophrys dzukou Megophrys sp. 2 SM-2017 SDBDU
2007.106 India, Nagaland, Kohima Mahony et al. (2017)
43 KX811807.1 Xenophrys auralensis Megophrys auralensis NCSM 79599 Cambodia, Kampong Speu, Aural Chen et al. (2017)
44 KY022214.1 Xenophrys lekaguli Xenophrys legkaguli FMNH 265955 ailand, Sa Kaeo, Mueang Sa Kaeo Mahony et al. (2017)
45 KY022215 .1 Xenophrys takensis Megophrys takensis FMNH 261711 ailand, Kampaeng, Khlong Lan Mahony et al. (2017)
46 MH406746.1 Xenophrys dehongensis Megophrys sp. 43 SYS a003443 China, Yunnan, Yingjiang Liu et al. (2018)
47 MH406737.1 Xenophrys parva Megophrys parva SYS a003042 China, Yunnan, Mengla/Jinghong Liu et al. (2018)
48 KX811925.1 Grillitschia aceras Megophrys aceras
KIZ
025467; KU
KUH332489
ailand, Nakhon Si ammarat, Khao
Nan National Park Chen et al. (2017)
49 AB530656.1 Grillitschia longipes Megophrys longipes IABHU 21101 Malaysia, Genting highland Hasan et al. (2014)
50 MT710708.1 Megophrys acehensis Megophrys sp.
MZBAmph26098
MZB Amph
26098 Indonesia, Sumatra, Aceh Munir et al. (2021)
51 KX773567.1 Megophrys lancip Megophrys sp. ENS 7577 ENS 7577 Indonesia, Sumatra, Bengkulu Mahony et al. (2017)
52 KX811927.1 Megophrys montana Megophrys montana LSUMZ 81916 Indonesia, Java, Jawa Barat Chen et al. (2017
53 KY679898.1 Megophrys parallela Megophrys parallela RMAS 022 Indonesia, Sumatra, West Sumatra Munir et al. (2018)
54 MT710704.1 Megophrys selatanensis Megophrys sp. MZB Amph
22411
MZB Amph
22411 Indonesia, Sumatra, Lampung Munir et al. (2021)
55 KY425389.1 Ophryophryne elna Ophryophryne elna ZMMU NAP-
02658 Vietnam, Dak Lak, Chu Yang Sin Poyarkov et al. 2017
56 KY425365.1 Ophryophryne gerti Ophryophryne gerti KIZ 013663 Vietnam, Lam Dong, Mt Langbian Poyarkov et al. 2017
57 KY022199 .1 Ophryophryne microstoma Megophrys microstoma AMNH 168682 Vietnam, Lao Cai, Van Ban Mahony et al. (2017)
58 KY022198 .1 Ophryophryne synoria Ophryophryne synoria FMNH 262778 Cambodia, Mondolkiri, O’Reang Mahony et al. (2017)
59 KY022202.1 Ophryophryne poilani Megophrys microstoma AMNH 169287 Vietnam, ua Tien Hue, A Luoi Mahony et al. (2017)
60 KY022204.1 Ophryphryne hansi Ophryphryne hansi AMNH 169285 Vietnam, ua ien Hue Mahony et al. (2017)
On a New Species of Xenophrys (Anura: Megophryidae) from Tale Wildlife Sanctuary, Arunachal Pradesh, India
with Comments on the Earlier Erroneous Report as X. maosonensis (Bourret, 1937)
33
Zoological Survey of India
Sr.
No Accession nu. Names used in this study Species name as per NCBI Voucher ID Locality References as per NCBI
61 KJ579119.1 Boulenophrys acuta Megophrys acuta SYS a002266 China, Guangdong, Fengkai Li et al. (2014)
62 MH406645.1 Boulenophrys nanlingensis Megophrys sp. 7 ZL-2018 SYS a001962 China, Guangdong, Mt Nanling Liu et al. (20180)
63 KJ560403.1 Boulenophrys brachykolos Megophrys brachykolos SYS a002258 China, Hong Kong Wang et al. (2014)
64 KX811813.1 Boulenophrys baolongensis Megophrys baolongensis KIZ 019216 China, Chongqing, Wushan Chen et al. (2017)
65 KX811849.1 Boulenophrys
binchuanensis Megophrys binchuanensis KIZ 019441 China, Yunnan, Mt Jizu Chen et al. (2017)
66 MH406892.1 Boulenophrys binlingensis Megophrys binglingensis SYS a005313 China, Sichuan, Mt Wawu/Hongya Liu et al. (2018)
67 MF667878.1 Boulenophrys boettgeri Boulenophrys boettgeri SYS a004149 China, Fujian, Mt Wuyi Wang et al. (2017)
68 MH406796.1 Boulenophrys caudoprocta Megophrys caudoprocta SYS a004293 China, Hunan, Sangzhi Liu et al. (2018)
69 MF667873.1 Boulenophrys cheni Megophrys cheni SYS a004050 China, Jiangxi, Mt Jinggang Wa n g et al. (2017)
70 MH406888.1 Boulenophrys chishuiensis Megophrys sp. 41 ZL-2018 SYS a005307 China, Sichuan, Mt Emei Liu et al. (2018)
71 MH406880.1 Boulenophrys
congjiangensis Megophrys sp. 33 ZL-2018 SYS a005134 China, Guangxi, Mt Jiuwan Liu et al. (2018)
72 MH406643.1 Boulenophrys daiyunensis Megophrys sp. 18 ZL-2018 SYS a001733 China, Fujian, Mt Daiyun Liu et al. (2018)
73 MH406783.1 Boulenophrys daoji Megophrys sp. 20 ZL-2018 SYS a004089 China, Zhejiang, Fenghua Liu et al. (2018)
74 KX811867.1 Boulenophrys daweimontis Boulenophrys daweimontis KIZ 048997 China, Yunnan, Mt Dawei Chen et al. (2017)
75 MH406648.1 Boulenophrys
dongguanensis Megophrys sp. 11 ZL-2018 SYS a001974 China, Guangdong, Dongguan Liu et al. (2018)
76 MH406808.1 Boulenophrys
fanjingmontis Megophrys sp. 37 ZL-2018 SYS a004350 China, Guizhou, Mt Fanjing Liu et al. (2018)
77 MH514886.1 Boulenophrys fansipanensis Megophrys fansipanensis VNMN 2018.01 Vietnam, Lao Cai, Sapa Tapley et al. (2018)
78 MH406848.1 Boulenophrys
fengshunensis Megophrys sp. 14 ZL-2018 SYS a004724 China, Guangdong, Mt Tongguzhang Liu et al. (2018)
79 MT364279.1 Boulenophrys frigida Boulenophrys frigida AMS R186131 Vietnam, Lao Cai, Bat Xat Unpublished
80 MH514890.1 Boulenophrys
hoanglienensis Megophrys hoanglienensis VNMN 07034 Vietnam, Lao Cai, Sapa Tapley et al. (2018)
81 MF667882.1 Boulenophrys
huangshanensis Megophrys huangshanensis SYS a002702 China, Anhui, Mt Huangshan Wang et al. (2017)
Bhaskar Saikia, Bikramjit Sinha, A. Shabnam, Ilona Jacinta Kharkongor and K. P. Dinesh
34 Vol 124(1S) | iS 2024 | www.recordsofzsi.com
Sr.
No Accession nu. Names used in this study Species name as per NCBI Voucher ID Locality References as per NCBI
82 OL635597.1 Boulenophrys hungtai Boulenophrys hungtai SYS a008576 China, Guangdong, Jiedong Unpublished
83 MH406662.1 Boulenophrys insularis Megophrys sp. 13 ZL-2018 SYS a002168 China, Guangdong, Nan’ao Liu et al. (2018)
84 KX811911.1 Boulenophrys jiangi Megophrys sp. 11 JMC-
2016 KIZ 03076 China, Chongqing, Youyang Chen et al. (2017)
85 MH406773.1 Boulenophrys jingdongensis Boulenophrys jingdongensis SYS a003928 China, Yunnan, Mt Wuliang Liu et al. (2018)
86 MH406857.1 Boulenophrys jinggangensis Boulenophrys jinggangensis SYS a004824 China, Hunan, Mt Sifang Liu et al. (2018)
87 MH406778.1 Boulenophrys jiulianensis Megophrys sp. 30 ZL-2018 SYS a004011 China, Guangdong, Mt Nankun Liu et al. (2018)
88 MF667881.1 Boulenophrys kuatunensis Boulenophrys kuatunensis SYS a003449 China, Jiangxi, Mt Wuyi Wang et al. (2017)
89 MH406674.1 Boulenophrys leishanensis Megophrys sp. 34 ZL-2018 SYS a002214 China, Guizhou, Mt Leigong Liu et al. (2018)
90 KJ560414.1 Boulenophrys lini Xenophrys lini SYS a002382 China, Jiangxi, Mt Jinggang Liu et al. (2018)
91 MH406756.1 Boulenophrys lushuiensis Megophrys sp. 40 ZL-2018 SYS a003785 China, Yunnan, Tengchong Liu et al. (2018)
92 MF667865.1 Boulenophrys minor Xenophrys minor SYS a003212 China, Sichuan, Dujiangyan Liu et al. (2018)
93 MH406681.1 Boulenophrys mirabilis Megophrys sp. 25 ZL-2018 SYS a002289 China, Guangxi, Lingui Liu et al. (2018)
94 MK524107.1 Boulenophrys
mufumontana
Boulenophrys
mufumontana SYS a006419 China, Hunan, Mt Mufu Wang et al. (2019)
95 MH406824.1 Boulenophrys nankunensis Megophrys sp. 10 ZL-2018 SYS a004503 China, Guangdong, Mt Nankun Liu et al. (2018)
96 KJ579123.1 Boulenophrys obesa Megophrys obesa SYS a002275 China, Guangdong, Fengkai Li et al. (2014)
97 MH406650.1 Boulenophrys ombrophila Megophrys sp. 9 ZL-2018 SYS a001976 China, Guangdong, Renhua Liu et al. (2018)
98 MH406887.1 Boulenophrys omeimontis Megophrys omeimontis SYS a005301 China, Sichuan, Mt Emei Liu et al. (2018)
99 KY022209 .1 Boulenophrys
palpebralespinosa
Megophrys
palpebralespinosa FMNH 258098 Laos, Phongsaly, Phou Dendin; Vietnam,
anh Hoa, Pu Hu Mahony et al. (2017
100 KX811855.1 Boulenophrys qianbeiensis Megophrys spinata YPX28756 China, Guizhou, Kuankuoshui Chen et al. (2017)
101 KX811871.1 Boulenophrys rubrimera Megophrys sp. 3 JMC-2016 KIZ 020425 China, Yunnan, Jinping Chen et al. (2017)
102 MH406802.1 Boulenophrys sangzhiensis Boulenophrys sangzhiensis SYS a004313 China, Hunan, Sangzhi Liu et al. (2018)
103 MH406699.1 Boulenophrys sanmingensis Megophrys sp. 15 ZL-2018 SYS a002499 China, Fujian, Jiangle Liu et al. (2018)
104 MH406655.1 Boulenophrys shimentaina Megophrys sp. 29 ZL-2018 SYS a002077 China, Guangdong, Yingde Liu et al. (2018)
On a New Species of Xenophrys (Anura: Megophryidae) from Tale Wildlife Sanctuary, Arunachal Pradesh, India
with Comments on the Earlier Erroneous Report as X. maosonensis (Bourret, 1937)
35
Zoological Survey of India
Sr.
No Accession nu. Names used in this study Species name as per NCBI Voucher ID Locality References as per NCBI
105 MH406684.1 Boulenophrys
shunhuangensis Megophrys sp. 24 ZL-2018 SYS a002295 China, Guangxi, Mt Mao’er Liu et al. (2018)
106 MH406675.1 Boulenophrys spinata Megophrys sp. inata ZL-
2018 SYS a002226 China, Guizhou, Mt Leigong Liu et al. (2018)
107 MH406744.1 Boulenophrys tongboensis Panophrys tongboensis SYS a003227 China, Jiangxi, Mt Tongbo Liu et al. (2018)
108 MH406801.1 Boulenophrys
tuberogranulata Panophrys tuberogranulata SYS a004310 China, Hunan, Sangzhi Liu et al. (2018)
109 MH406854.1 Boulenophrys wugongensis Megophrys sp. 12 ZL-2018 SYS a004801 China, Jiangxi, Mt Wugong Liu et al. (2018)
110 MH406771.1 Boulenophrys
wuliangshanensis
Megophrys
wuliangshanensis SYS a003924 China, Yunnan, Mt Wuliang Liu et al. (2018)
111 MH406733.1 Boulenophrys wushanensis Megophrys wushanensis SYS a003009 China, Hubei, Shennongjia; China,
Sichuan, Mt Guangwu Liu et al. (2018)
112 MH406713.1 Boulenophrys
xiangnanensis Megophrys sp. 2 ZL-2018 SYS a002874 China, Hunan, Shuangpai Liu et al. (2018)
113 MH406704.1 Boulenophrys xianjuensis Megophrys sp. 19 ZL-2018 SYS a002668 China, Zhejiang, Mt Dapan Liu et al. (2018)
114 MH406814.1 Boulenophrys
xuefengmontis Megophrys sp. 26 ZL-2018 SYS a004365 China, Hunan, Wugang Liu et al. (2018)
115 MH406719.1 Boulenophrys
yangmingensis Megophrys sp. 28 ZL-2018 SYS a002888 China, Hunan, Shuangpai Liu et al. (2018)
116 MH406863.1 Boulenophrys yaoshanensis Megophrys sp. 31 ZL-2018 SYS a004878 China, Guangxi, Mt Dayao Liu et al. (2018)
117 MH406898.1 Boulenophrys yingdeensis Megophrys sp. 4 ZL-2018 SYS a005447 China, Guangdong, Yingde Liu et al. (2018)
118 MH406844.1 Boulenophrys yunkaiensis Boulenophrys yunkaiensis SYS a004638 China, Guangdong, Mt Yunkai Liu et al. (2018)
119 OQ181007.1 Jingophrys feii Jingophrys feii SYS a003876 China, Yunnan, Yingjiang Lyu et al. (2023)
120 MN963206.1 Jingophrys pachyproctus Megophrys sp. h SS-2020 CIB MT171054 China, Xizang, Medog Shi et al. (2020)
121 KY022305.1 Jingophrys vegrandis Megophrys vegrandis ZSIC A 11605 India: Arunachal Pradesh, West Kameng Mahony et al. (2018)
122 KX811908.1 Jingophrys yeae Megophrys pachyproctus KIZ 010978 China, Xizang, Medog Chen et al. (2017)
123 MN963207.1 Jingophrys zhoui Megophrys zhoui CIB MT171053 China, Renqinbeng, Medog Shi et al. (2020)
124 DQ642121.1 Pelobatrachus baluensis Xenophrys baluensis IRSNB 15926 Malaysia: Sabah, Mt. Kinabalu Lyu et al. (2023);
Mahony et al. (2017)
Bhaskar Saikia, Bikramjit Sinha, A. Shabnam, Ilona Jacinta Kharkongor and K. P. Dinesh
36 Vol 124(1S) | iS 2024 | www.recordsofzsi.com
Sr.
No Accession nu. Names used in this study Species name as per NCBI Voucher ID Locality References as per NCBI
125 KX811918.1 Pelobatrachus edwardinae Megophrys edwardinae FMNH 273694 Malaysia, Sarawak, Bintulu Chen et al. (2017)
126 AB719248.1 Pelobatrachus
kalimantanensis Megophrys nasuta KUHE 53577 Malaysia, Sabah, Crocker, Tenom Hamidy et al. (2012)
127 KJ831313.1 Pelobatrachus kobayashii Megophrys kobayashii UNIMAS 8148 Malaysia, Sabah, Gunung Kinabalu
National Park
Oberhummer et al.
(2014)
128 KY022192.1 Pelobatrachus ligayae Megophrys ligayae KU
KUH309095 Philippines, Palawan Mahony et al. (2017)
129 KY022186.1 Pelobatrachus nasutus Megophrys nasuta FMNH 231281 Malaysia, Sabah Mahony et al. (2017)
130 KY022190.1 Pelobatrachus stejnegeri Megophrys stejnegeri FMNH 250842 Philippines, Mindanao Mahony et al. (2017)
131 KJ831316.1 Sarawakiphrys dringi Xenophrys dringi UNIMAS 8948 Malaysia, Sarawak, Gunung Mulu
National Park
Oberhummer et al.
(2014)
132 MH406902.1 Brachytarsophrys
chuannanensis Megophrys chuannanensis SYS a004927 China, Sichuan, Hejiang Liu et al. (2018)
133 MH406900.1 Brachytarsophrys feae Megophrys feae SYS a003913 China, Yunnan, Jingdong Liu et al. (2018)
134 OQ180989.1 Brachytarsophrys orientalis Brachytarsophrys orientalis SYS a004225 China, Jiangxi, Mt Jiulian Lyu et al. (2023)
135 OQ180990.1 Brachytarsophrys
platyparietus Megophrys platyparietus SYS a005919 China, Yunnan, Dayao Lyu et al. (2023)
136 KM504256.1 Brachytarsophrys popei Megophrys popei SYS a001864 China, Hunan, Yanling Zhao et al. (2014)
137 MH406775.1 Atympanophrys gigantica Megophrys gigantica SYS a003933 China, Yunnan, Jingdong Liu et al. (2018)
138 EF397243.1 Atympanophrys
nankiangensis Megophrys nankiangensis CIB XM835 China, Sichuan, Nanjiang Fu et al. (2007)
139 MH406890.1 Atympanophrys
shapingensis Megophrys shapingensis SYS a005310 China, Sichuan, Mt Wawu Liu et al. (2018)
140 MH406891.1 Atympanophrys wawuensis Megophrys wawuensis SYS a005311 China, Sichuan, Mt Wawu Liu et al. (2018)
141 MH406905.1 Leptobrachella alpina Leptobrachella alpina SYS a003927 China, Yunnan, Jingdong Liu et al. (2018)
142 MH406903.1 Leptobrachella laui Leptobrachella laui SYS a003471 China, Guangdong, Mt Wutong Liu et al. (2018)
On a New Species of Xenophrys (Anura: Megophryidae) from Tale Wildlife Sanctuary, Arunachal Pradesh, India
with Comments on the Earlier Erroneous Report as X. maosonensis (Bourret, 1937)
37
Zoological Survey of India
Acknowledgement:
e authors are grateful to the Director, Zoological Survey
of India, Kolkata for providing facilities. e authors
also acknowledge the Department of Environment and
Forests, Arunachal Pradesh for the permission vide letter
No.CWL/G/13 (95)/2011-12/Pt./1048-49 dt. 22/06/2015 to
survey and collect in Tale Wildlife Sanctuary. e authors are
also grateful to the respective Ocer-in-Charge of NERC,
ZSI, Shillong; WRC, ZSI, Pune and APRC, ZSI, Itanagar
for the support, and special gratitude to Sahil of WRC, ZSI,
Pune and Yomto Mayi of APRC, ZSI, Itanagar for their help
during the preparation of the manuscript.
References:
Boruah, B., Deepak, V., Patel, N. G., Jithin, V., Yomcha, T. and Das. A. 2023a. A new species of green tree frog of the genus
Gracixalus (Anura: Rhacophoridae) from the evergreen forest of Northeast India. Vertebrate Zoology, 73: 557–574
Boruah, B., Narayanan, S., Gerard, J. D., Das, A. and Deepak. V. 2023b. Discovery of a new species of dwarf frog (Anura:
Ceratobatrachidae: Alcalus) extends the northwestern distributional limits of the genus to Northeast India. Systematics
and Biodiversity, 21(1: 2249891): 1–15.
Boruah, B., Deepak, V. and Das, A. 2023c. Musicians in the marsh: A new species of music frog (Anura: Ranidae: Nidirana)
from Arunachal Pradesh, India. Zootaxa, 5374: 51–73.
Bourret, R. 1937. Notes herpétologiques sur l’Indochine française. XIV. Les batraciens de la collection du Laboratoire des
Sciences Naturelles de l’Université. Descriptions de quinze especes ou variétés nouvelles. Annexe au Bulletin Général de
l’Instruction Publique, 1937: 5–56.
Chen, J.M., Zhou, W.W., Poyarkov Jr, N.A., Stuart, B.L., Brown, R.M., Lathrop, A., Wang, Y.Y., Yuan, Z.Y., Jiang, K., Hou, M.
and Chen, H.M., 2017. A novel multilocus phylogenetic estimation reveals unrecognized diversity in Asian horned toads,
genus Megophrys sensu lato (Anura: Megophryidae). Molecular Phylogenetics and Evolution, 106:28-43.
Dinesh, K.P., Radhakrishnan, C., Deepak, P. and Kulkarni, N.U. 2023. A Checklist of Indian Amphibians with Common names
for the country and their IUCN Conservation Status. pp 1–19. Version 5.0 (updated till January 2023) available at http://
zsi.gov.in (online only).
Frost, D. R. 2024. Amphibian Species of the World: an Online Reference. Version 6.2 (05 March, 2024). Electronic Database
accessible at https://amphibiansoheworld.amnh.org/index.php. American Museum of Natural History, New York, USA.
doi.org/10.5531/db.vz.0001
Grosjean, S., Ohler, A., Chuaynkern, Y., Cruaud, C. and Hassanin, A. 2015. Improving biodiversity assessment of anuran
amphibians using DNA barcoding of tadpoles. Case studies from Southeast Asia. Comptes Rendus. Biologies, 338(5):
351-361.
Fu, J.Z., Weadick, C.J. and Bi, K., 2007. A phylogeny of the high‐elevation Tibetan megophryid frogs and evidence for the
multiple origins of reversed sexual size dimorphism. Journal of Zoology, 273(3):315-325.
Hasan, M., Islam, M., Khan, M., Igawa, T., Alam, M.S., Djong, H.T., Kurniawan, N., Joshy, H., Sen, Y.H., Belabut, D.M. and
Kurabayashi, A., 2014. Genetic divergences of South and Southeast Asian frogs: a case study of several taxa based on 16S
ribosomal RNA gene data with notes on the generic name Fejervarya. Turkish Journal of Zoology, 38(4): 389-411.
Huang, C.-M., Lathrop, A. and Murphy, R.W. 1998. Translations of “Two new species of amphibian in Tibet” Huang & Fei
1981 and “Description of two new species of the genus Megophrys” Fei, Ye, & Huang 1995. Smithsonian Herpetological
Information Service, 118: 1–20.
Kamei, R. G., Gower, D. J., Wilkinson, M. and Biju, S. D. 2013. Systematics of the caecilian family Chikilidae (Amphibia:
Gymnophiona) with the description of three new species of Chikila from Northeast India. Zootaxa, 3666: 401–435.
Bhaskar Saikia, Bikramjit Sinha, A. Shabnam, Ilona Jacinta Kharkongor and K. P. Dinesh
38 Vol 124(1S) | iS 2024 | www.recordsofzsi.com
Kurabayashi, A., Matsui, M., Belabut, D.M., Yong, H.S., Ahmad, N., Sudin, A., Kuramoto, M., Hamidy, A. and Sumida, M.,
2011. From Antarctica or Asia? New colonization scenario for Australian-New Guinean narrow mouth toads suggested
from the ndings on a mysterious genus Gastrophrynoides. BMC Evolutionary Biology, 11: 1-12.
Lanfear, R., Calcott, B., Ho, S. Y. W. and Guindon, S. 2012. PartitionFinder: combined selection of partitioning schemes and
substitution models for phylogenetic analyses. Molecular Biology and Evolution, 29(6): 1695–1701.
Li, Y.-l., M.-j. Jin, Z.-y., Zhao, Liu, Z.-y., Wang, Y. and Pang, H. 2014. Description of two new species of the genus Megophrys
(Amphibia: Anura: Megophryidae) from Heishiding Nature Reserve, Fengkai, Guangdong, China, based on molecular
and morphological data. Zootaxa, 3795: 449–471.
Liu, Z., Chen, G., Zhu, T., Zeng, Z., Lyu, Z., Wang, J., Messenger, K., Greenberg, A.J., Guo, Z., Yang, Z. and Shi, S., 2018.
Prevalence of cryptic species in morphologically uniform taxa–Fast speciation and evolutionary radiation in Asian frogs.
Molecular Phylogenetics and Evolution, 127: 723-731.
Luong, A.M., Pham, C.T., Nguyen, T.T., Orlov, N., Ziegler, T. and Nguyen, T.Q. 2022. A new species of Xenophrys (Amphibia:
Anura: Megophryidae) from Truong Son Range, Vietnam. Zootaxa, 5150(3): 333-356.
Lyu, Z.-T., Qi, S., Wang, J., Zhang, S.-Y., Zhao, J., Zeng, Z.-C., Wan, H., Yang, J.-h., Mo, Y.-m. and Wang, Y.-y. 2023. Generic
classication of Asian Horned Toads (Anura: Megophryidae: Megophryinae) and monograph of Chinese species.
Zoological Research, 44: 380–450.
Mahony, S., Sengupta, S., Kamei, R. G. and Biju, S. D. 2011. A new low altitude species of Megophrys Kuhl and van Hasselt
(Amphibia: Megophryidae), from Assam, Northeast India. Zootaxa, 3059: 36–46.
Mahony, S., Teeling, E. C. and Biju, S. D. 2013. ree new species of horned frogs, Megophrys (Amphibia: Megophryidae), from
northeast India, with a resolution to the identity of Megophrys boettgeri populations reported from the region. Zootaxa,
3722: 143–169.
Mahony, S., Foley, N.M., Biju, S.D. and Teeling, E.C., 2017. Evolutionary history of the Asian Horned Frogs (Megophryinae):
integrative approaches to timetree dating in the absence of a fossil record. Molecular Biology and Evolution, 34(3):744-
771.
Mahony, S., Kamei, R.G., Teeling, E.C. and Biju, S.D. 2018. Cryptic diversity within the Megophrys major species group
(Amphibia: Megophryidae) of the Asian Horned Frogs: Phylogenetic perspectives and a taxonomic revision of South
Asian taxa, with descriptions of four new species. Zootaxa, 4523(1):1-96.
Mahony, S., Kamei, R.G., Teeling, E.C. and Biju, S.D. 2020. Taxonomic review of the Asian Horned Frogs (Amphibia:
Megophrys Kuhl & Van Hasselt) of Northeast India and Bangladesh previously misidentied as M. parva (Boulenger),
with descriptions of three new species. Journal of Natural History, 54(1-4):119-194.
Mathew, R., and Sen, N. 2009. Studies on little known amphibians of Northeast India. Records of the Zoological Survey of India.
Occasional Papers, 293: 1–64 + 23 plates.
Munir, M., Hamidy, A., Farajallah, A. and Smith, E.N. 2018. A new Megophrys Kuhl and Van Hasselt (Amphibia: Megophryidae)
from southwestern Sumatra, Indonesia. Zootaxa, 4442(3):389–412.
Munir, M., Nishikawa, K., Hamidy, A. and Smith, E.N. 2021. Two new species of Megophrys Kuhl and Van Hasselt (Amphibia:
Megophryidae) from Sumatra, Indonesia. Zootaxa, 5057(4):503-529.
Nguyen, T.Q., Pham, C.T., Nguyen, T.T., Luong, A.M. and Ziegler, T. 2020. A new species of Megophrys (Amphibia: Anura:
Megophryidae) from Vietnam. Zootaxa, 4722(5): 401–422.
Oberhummer, E., Barten, C., Schweizer, M., Das, I., Haas, A. and Hertwig, S.T. 2014. Description of the tadpoles of three
rare species of megophryid frogs (Amphibia: Anura: Megophryidae) from Gunung Mulu, Sarawak, Malaysia. Zootaxa,
3835(1): 59–79.
On a New Species of Xenophrys (Anura: Megophryidae) from Tale Wildlife Sanctuary, Arunachal Pradesh, India
with Comments on the Earlier Erroneous Report as X. maosonensis (Bourret, 1937)
39
Zoological Survey of India
Patel, N. G., Garg, S., Das, A., Stuart, B. L. and Biju, S. D. 2021. Phylogenetic position of the poorly known montane cascade
frog Amolops monticola (Ranidae) and description of a new closely related species from Northeast India. Journal of
Natural History, 55: 1403-1440.
Poyarkov Jr, N.A., Van Duong, T., Orlov, N.L., Gogoleva, S.S., Vassilieva, A.B., Nguyen, L.T., Nguyen, V.D.H., Nguyen, S.N.,
Che, J. and Mahony, S. 2017. Molecular, morphological and acoustic assessment of the genus Ophryophryne (Anura,
Megophryidae) from Langbian Plateau, southern Vietnam, with description of a new species. ZooKeys, 672: 49-120.
Saikia B, Sinha, B. and Kharkongor, I. J. 2017. Odorrana arunachalensis: A New Species of Cascade Frog (Anura: Ranidae) from
Talle Valley Wildlife Sanctuary, Arunachal Pradesh, India. Journal of Bioresources, 4(2): 30-41.
Saikia, B., Sinha, B., akur, M. and Kharkongor, I. J. 2019. Megophrys maosonensis Bourret, 1937 (Anura: Megophyridae)—A
new India record from Tale Valley Wildlife Sanctuary, Arunachal Pradesh, India. Records of the Zoological Survey of India,
119: 456-462.
Saikia, B., and Sinha, B. 2019. On the Liurana (Anura: Ceratobatrachidae) of India with the description of three new species
from Tale Valley Wildlife Sanctuary in Arunachal Pradesh, eastern Himalayas. Records of the Zoological Survey of India,
119: 303–315.
Saikia, B., Laskar, M. A., Dinesh, K. P. Shabnam, A. and Sinha, B. 2022a. Description of two new species of Amolops (Anura:
Ranidae) from Arunachal Pradesh, Northeast India under the morphological ‘Viridimaculatus species group’. Records of
the Zoological Survey of India, 122: 247–266.
Saikia, B., Sinha, B., Laskar, M. A., Shabnam, A. and Dinesh, K. P. 2022b. A new species of Amolops (Anura: Ranidae)
representing the morphological ‘Marmoratus species group’ from Sessa Orchid Sanctuary, Arunachal Pradesh, Northeast
India. Records of the Zoological Survey of India, 122: 303–322.
Shi, S., Zhang, M., Xie, F., Jiang, J., Liu, W., Ding, L., Luan, L. and Wang, B., 2020. Multiple data revealed two new species
of the Asian horned toad Megophrys Kuhl & Van Hasselt, 1822 (Anura, Megophryidae) from the eastern corner of the
Himalayas. ZooKeys, 977: 101-161.
Sondhi, S. and Ohler, A. 2011. A blue eyed Leptobrachium (Anura: Megophrydiae) from Arunachal Pradesh. Zootaxa, 2912:
28-36.
Tamura, K., G. Stecher, and S. Kumar. 2021. MEGA11: Molecular Evolutionary Genetics Analysis version 11. Molecular Biology
and Evolution, 38: 3022–3027.
Tapley, B., Cutajar, T.P., Mahony, S., Nguyen, C.T., Dau, V.Q., Luong, A.M., Le, D.T., Nguyen, T.T., Nguyen, T.Q., Portway, C.
and Luong, H.V. 2018. Two new and potentially highly threatened Megophrys Horned frogs (Amphibia: Megophryidae)
from Indochina’s highest mountains. Zootaxa, 4508(3): 301-333.
Tapley, B., Nguyen, L.T., Cutajar, T., Nguyen, C.T., Portway, C., Luong, H.V. and Rowley, J.J. 2020. e tadpoles of ve Megophrys
Horned frogs (Amphibia: Megophryidae) from the Hoang Lien Range, Vietnam. Zootaxa, 4845(1): 35-52.
Wang, J., Liu, Z.Y., Lyu, Z.T., Zeng, Z.C. and Wang, Y.Y. 2017. A new species of the genus Xenophrys (Amphibia: Anura:
Megophryidae) from an oshore island in Guangdong Province, southeastern China. Zootaxa, 4324(3): 541-556.
Wang, J., Lyu, Z.T., Liu, Z.Y., Liao, C.K., Zeng, Z.C., Zhao, J., Li, Y.L. and Wang, Y.Y. 2019. Description of six new species of the
subgenus Panophrys within the genus Megophrys (Anura, Megophryidae) from southeastern China based on molecular
and morphological data. ZooKeys, 851: 113-164.
Wang, Y., Zhao, J., Yang, J., Zhou, Z., Chen, G. and Liu, Y., 2014. Morphology, molecular genetics, and bioacoustics support two
new sympatric Xenophrys toads (Amphibia: Anura: Megophryidae) in Southeast China. PLoS One, 9(4): e93075.
Bhaskar Saikia, Bikramjit Sinha, A. Shabnam, Ilona Jacinta Kharkongor and K. P. Dinesh
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