517
Re ision o bush ogs, Rao ches es and Philau us
(Amphibia: Anu a: Rhacopho idae) om he no heas
Indian biodi e si y ho spo wi h desc ip ion o hi een
new species
Bi upan Bo uah1, V. Deepak2,3,4, Abhiji Das1
1 Wildli e Ins i u e o India, Chand abani, Deh adun, U a akhand 248001, India
2 Museum o Zoology, Senckenbe g Na u al His o y Collec ions, Königsb ücke Lands aße 159, 01109 D esden, Ge many
3 The Na u al His o y Museum, C omwell Rd, Sou h Kensing on, SW7 5BD London, Uni ed Kingdom
4 Newcas le Uni e si y, Newcas le NE1 7RU, Uni ed Kingdom
h ps://zoobank.o g/7C8226BF-FEA3-4EE2-9012-C0B859797028
Co esponding au ho : Abhiji Das ([email p o ec ed].in)
Academic edi o Uwe F i z | Recei ed 28 Janua y 2025 | Accep ed 21 Oc obe 2025 | Published 20 No embe 2025
Ci a ion: Bo uah B, Deepak V, Das A (2025) Re ision o bush ogs, Rao ches es and Philau us (Amphibia: Anu a: Rhacopho idae) om he
no heas Indian biodi e si y ho spo wi h desc ip ion o hi een new species. Ve eb a e Zoology 75: 517–625. h ps://doi.o g/10.3897/ z.75.
e148133
Abs ac
Bush ogs cu en ly in he gene a Rao ches es and Philau us a e poo ly documen ed om no heas India when compa ed o he
Wes e n Gha s o sou hwes e n India. They a e mo phologically c yp ic, bu a iable in hei acous ic epe oi e and gene ic di e -
gence. We p esen a long-o e due e ision o he bush ogs o no heas India by sampling 81 locali ies in eigh Indian S a es (in-
cluding nine ype locali ies o known species) and by compa ing his o ical (especially, ype) ma e ial. Using an in eg a i e app oach
ha combines le els o di e gence in mi ochond ial DNA, a nuclea -encoded gene and compa ing mo phological and bioacous ic
da a, we demons a e he exis ence o 13 new species, which a e o mally desc ibed he e. In addi ion, based on he e idence om
molecula and mo phological da a we synonymise ou p e iously desc ibed species om his egion. We esol e he axonomic
iden i ies o h ee o he og species ound in no heas India. A e ex ensi e su ey ac oss no heas India, we did no ind a single
specimen o he genus Philau us. The only species in his genus wi h ambiguous gene ic iden i y is P. dubius which equi es u he
axonomic in es iga ion.
Keywo ds
Bioacous ics, c yp ic species, Indo-Bu ma, Indochina, mo phology, phylogeny, edesc ip ion, axonomy
In oduc ion
The anu an amily Rhacopho idae is ep esen ed by 473
species in 23 gene a (F os 2025). Membe s o his am-
ily a e dis ibu ed h oughou sub-Saha an A ica, Sou h,
Sou heas and Eas Asia (Ellepola and Meegaskumbu a
2023; F os 2025). In Asia he e a e h ee gene a o bush
ogs/sh ub ogs namely: Philau us Gis el, 1848, Pseu-
dophilau us Lau en , 1943, and Rao ches es Biju e al.,
2010. Wi hin Rhacopho idae, hese h ee gene a can be
Ve eb a e Zoology 75, 2025, 517–625 | DOI 10.3897/ z.75.e148133
Copy igh Bi upan Bo uah e al. This is an open access a icle dis ibu ed unde he e ms o he C ea i e Commons A ibu ion License (CC BY 4.0), which pe mi s un es ic ed use,
dis ibu ion, and ep oduc ion in any medium, p o ided he o iginal au ho and sou ce a e c edi ed.
Bo uah B e al.: Re ision o bush ogs om no heas India
518
dis inguished om o he gene a by hei di ec de el-
opmen o leci ho ophic adpole and small o medium
body size (maximum snou - en leng h 65 mm) in adul s
(Bossuy and Dubois 2001; Biju e al. 2010; E e e al.
2021; Ga g e al. 2021). The e a e cu en ly 212 species o
Asian bush/sh ub ogs, 121 o which ha e been desc ibed
in he las wo decades. Species in his polyphyle ic g oup
we e o iginally desc ibed unde many hacopho id and
non- hacopho id gene a un il Bossuy and Dubois (2001)
placed hem in he genus Philau us. Subsequen ly, Li e
al. (2009) esu ec ed he genus Pseudophilau us which
cu en ly has 78 species, dis ibu ed mos ly in S i Lanka,
and an addi ional h ee species ound in he Wes e n Gha s
and one species in he Eas e n Gha s o India (Bha a h e
al. 2024). Biju e al. (2010) e ec ed Rao ches es o ea
a la ge adia ion (82 species) o Asian bush ogs dis ib-
u ed in he Wes e n Gha s, cen al, eas e n and no heas
India, Nepal, sou he n China, Myanma , Thailand, Laos,
Vie nam, Cambodia and Malaysia (Du e al. 2024; F os
2025). Recen obus phylogene ic s udies ha e no only
desc ibed new species wi hin he genus, bu ha e also
ans e ed species ha we e o me ly alloca ed o o he
gene a, such as Thelode ma Tschudi, 1838 (Chen e al.
2020; Ga g e al. 2021). Based on Biju e al. (2010), a ew
species om no heas India ha e been ans e ed o he
genus Rao ches es, and a ew doub ul species con inue
o be placed in he genus Philau us (Ga g e al. 2021).
A p esen , 15 species o bush ogs a e known om
no heas India unde he genus Rao ches es [R. ande -
soni (Ahl, 1927), R. annandalii (Boulenge , 1906), R.
asakg ensis Na een, Chand amouli & Babu, 2024, R.
ga o (Boulenge , 1919), R. jadoh Wa j i, Pu kayas ha,
Lal emsanga & Das, 2025, R. jakoid Wa j i, Pu kayas ha,
Lal emsanga & Das, 2025, R. kempiae (Boulenge , 1919),
R. manipu ensis (Ma hew & Sen, 2009), R. ezakhani Al-
Razi e al., 2020, R. sahai (Sa ka & Ray, 2006) and R.
shillongensis (Pillai & Chanda, 1973)] and Philau us [P.
dubius (Boulenge , 1882), P. kempii (Annandale, 1912),
P. mic odiscus (Annandale, 1912), P. namdaphaensis
(Sa ka & Sanyal, 1985)]. O hese, 13 species ha e been
desc ibed om no heas India (Fig. 1A; Table S1A). Ou
o hese 13 species, 10 (R. asakg ensis, R. ga o, R. jadoh,
R. jakoid, R. manipu ensis, R. sahai, R. shillongensis, P.
dubius, P. mic odiscus and P. namdaphaensis) a e endem-
ic o he egion. Bo uah e al. (2018) iden i ied he phy-
logene ic posi ion o R. shillongensis and edesc ibed he
species and Na een e al. (2024) edesc ibed R. ga o and
R. kempiae and gene a ed some gene ic da a and added
o he phylogene ic in o ma ion. Mos ecen ly, Wa j i e
al. (2025) desc ibed wo new species om Meghalaya.
Howe e , hese s udies elied p ima ily on single-locus
da ase s and geog aphically limi ed sampling. In con-
as , comp ehensi e mul i-locus phylogene ic app oach-
es ha e been shown o be mo e e ec i e in esol ing
axonomic unce ain ies (Chan e al. 2018, 2020). Addi-
ionally, mos desc ip ions o he emaining species om
no heas India a e o e a cen u y old and based on a sin-
gle specimen. Since he o iginal desc ip ions, less e o
was made o e isi he ype locali ies o hese species.
Subsequen ly, many au ho s (Chanda 1994; Choudhu y
e al. 2002; Ao e al. 2003; Sa ka and Ray 2006; Sen and
Ma hew 2008; Ma hew and Sen 2009, 2010; Sengup a
e al. 2010; Sen e al. 2013; Saikia e al. 2016; Ohle e
al. 2018; Pu kayas ha e al. 2020) epo ed o lis ed hese
species ollowing he o iginal desc ip ion, al hough hei
axonomic iden i ies we e no eassessed. The e o e, a
comp ehensi e and sys ema ic s udy o he bush ogs o
no heas India is a long o e due. Thus, ou p esen s udy
aims o ill his gap by u ilising mo phological, molecula
and acous ic da a.
Unlike he Rao ches es om he Wes e n Gha s which
a e di e se in body size (16–50.5 mm), colou a ion and
ecomo phology (Biju and Bossuy 2009; Zacha iah e al.
2011; Vijayakuma e al. 2014; Ga g e al. 2021), he spe-
cies om no heas India a e mo e conse a i e in mo -
phology, wi h a less di e ence in ange o body size (13–28
mm) and colou a ion, and occupy simila ecological nich-
es. Howe e , he e a e mul iple geog aphic ba ie s and
a ious habi a ypes in no heas India whe e se e al small
amphibian and ep ile species ha e di e si ied (Aga wal e
al. 2014; Mahony e al. 2020; Bo uah e al. 2024).
The e ec i eness o using bioacous ic e idence in sys-
ema ic s udies e ealing c yp ic anu an species has been
highligh ed o o he ogs (P asad e al. 2020; T o ime s
e al. 2024; Vences e al. 2024). Rao ches es species a e
highly ocal, wi h males emi ing dis inc i e ad e ise-
men calls ha ha e been p o en use ul o axonomic
classi ica ion (Ga g e al. 2021). Howe e , in o ma ion
a ailable on he bioacous ics o bush ogs om no heas
India is limi ed.
Be ween he yea s 2016 and 2024, we sampled ac oss
no heas India (see Fig. 1B), collec ing mul iple popu-
la ions o bush ogs wi h pa icula ocus on opo ypic
ma e ial. In his e ision we eassess he phylogene ic
ela ionships o he bush ogs o no heas India, esol e
p e ious ambiguous species iden i ica ions and desc ibe
13 new species based on molecula da a, mo phological
cha ac e s, and acous ics.
Ma e ials and me hods
Field su ey and sample collec ion
We collec ed 204 specimens om 81 locali ies ac oss
eigh Indian s a es, namely A unachal P adesh, Assam,
Manipu , Meghalaya, Mizo am, Nagaland, Sikkim and
Wes Bengal du ing 2016–2024 (Fig. 1B; Table S1B).
Specimens we e loca ed isually a nigh using a hand-
held o chligh o by ollowing he calls o male ogs.
Collec ed specimens we e pho og aphed in li e and hen
eu hanized using icaine me hanesul ona e (MS-222).
Specimens we e ixed in 4% o malin and s o ed in 70%
e hanol a e insing in wa e . P io o ixa ion, issue
samples we e collec ed om high muscle o li e and
s o ed in absolu e e hanol o DNA ex ac ion. Collec ed
specimens we e deposi ed in he eposi o y a he Wildli e
Ins i u e o India, Deh adun.
Ve eb a e Zoology 75, 2025, 517–625 519
Figu e 1. Map o no heas India showing dis ibu ion. A Type locali ies o p e iously desc ibed bush og species om no heas
India, Indochina and Indo-Bu ma; B sampling locali ies o bush ogs du ing his s udy be ween 2016 and 2024 (see Table S1 o
locali y de ails). Poli ical bounda ies o neighbou ing coun ies a e ma ked wi h black lines and s a e bounda ies wi hin he poli ical
bounda y o India a e ma ked wi h g ey lines.
Bo uah B e al.: Re ision o bush ogs om no heas India
520
DNA ex ac ion and ampli ica ion
We ex ac ed genomic DNA om issue samples using
Qiagen DNeasy Blood and Tissue Ki ollowing he man-
u ac u e ’s ins uc ions. We ampli ied pa ial sequences
o h ee mi ochond ial genes, 16S RNA (16S; 560 base
pai s (bp)), cy och ome b (cy b; 610 base pai s) and cy-
och ome c oxidase subuni I (COI; 680 base pai s), and
one nuclea gene, hodopsin (RHO; 320 base pai s) using
p e iously published p ime s and p o ocols (Table S2).
Ampli ied PCR p oduc s we e un on a 2% aga ose gel
and iewed unde UV ansillumina o . The pu i ied PCR
p oduc s we e sequenced in bo h di ec ions on an Applied
Biosys ems Gene ic Analyze 3500 XL using BigDye 3.1
ki s. The newly gene a ed sequences in his s udy we e de-
posi ed in DNA Da a Bank o Japan (DDBJ) wi h he eg-
is a ion numbe s LC896952, LC896953, and LC898195–
LC898646 (Table S3).
Phylogene ic analyses
We conduc ed wo phylogene ic analyses, one using con-
ca ena ed da ase o mi ochond ial and nuclea genes
consis ing o a o al o 2034 bp (16S 556 bp, cy b 571
bp, COI 617 bp and RHO 290 bp) and one using conca -
ena ed da ase o only mi ochond ial genes. Bidi ec ional
sequences we e manually checked using he CHROMAS
2.6.6 so wa e (h p:// echnelysium.com.au/wp/ch omas)
and aligned using Clus alW (Thompson e al. 1994) wi h
de aul p io se ings implemen ed in MEGA 7.1 (Kuma
e al. 2016). Fo he p o ein-coding genes, we checked o
unexpec ed s op codons by ansla ing he sequence in o
amino acids in MEGA 7.1. We aligned he newly gene a -
ed sequences wi h sequences downloaded om GenBank
(Benson e al. 2007) (Table S3). We used six Rao ches es
species, h ee each om he no he n and sou he n clades
om he Wes e n Gha s and wo species o Pseudophilau-
us as ou g oups ollowing Ga g e al. (2021).
We pe o med maximum likelihood analyses using he
GUI e sion o he IQTREE (Nguyen e al. 2015), imple-
men ed in PHYLOSUITE (Zhang e al. 2020) one on he
conca ena ed da ase o mi ochond ial and nuclea genes
and ano he one on conca ena ed da ase o only mi ochon-
d ial genes. We pa i ioned he wo da ase s by gene and
codon posi ion using MODELFINDER (Kalyaanamoo -
hy e al. 2017) o ind he bes - i pa i ions and models o
sequence e olu ion o each pa i ion (Table S4).
We also pe o med a Bayesian in e ence (BI) analy-
sis o he conca ena ed da ase on mi ochond ial and
nuclea genes using he p og am M Bayes 3.2 (Ronquis
e al. 2012), wi h de aul p io se ings. The same da a
se used o he maximum likelihood (ML) analysis was
pa i ioned by codon posi ions using Pa i ionFinde
1.1.1 (Lan ea e al. 2012) wi h de aul se ings o ind
he bes - i model o sequence e olu ion (Table S4A).
Fou sepa a e uns we e se up wi h eigh Ma ko chains
each ini ia ed om andom ees and allowed o un o
10 million gene a ions, sampling e e y 100 gene a ions.
Analyses we e e mina ed when he s anda d de ia ion o
he spli equencies was less han 0.001, he i s 25% o
ees we e disca ded as bu n-in, and ees we e cons uc -
ed using a 50% majo i y consensus ule. We ob ained he
ESS alues using TRACER 1.6 (Rambau e al. 2018)
and con i med g ea e han 200 o he p io s. Suppo o
he in e nal b anches o he ML and BI was quan i ied
using 10,000 pseudo eplica es (ul a as boo s ap UFB)
and pos e io p obabili y (PP), espec i ely. B anch sup-
po s (UFB and PP) highe han 95 (UFB: see Minh e al.
2013) and >0.95 we e conside ed as s ong suppo . The
esul ing ee was edi ed in FigT ee .1.4.4 (h p:// ee.
bio.ed.ac.uk/so wa e/ ig ee).
Mo phological examina ion o he ype se ies o
Rao ches es sahai sugges ed ha hey did no belong o
he genus Rao ches es (see below in axonomic conclu-
sions sec ion). In o de o con i m he iden i y o his spe-
cies, we pe o med a BLAST sea ch (Al schul e al. 1990)
on he NCBI BLAST websi e (h p://blas .ncbi.nlm.nih.
go ) o he sequences (16S) gene a ed om samples ha
we e collec ed om he ype locali y and a e mo phologi-
cally simila o he ype se ies. These opo ypic sequences
showed simila i y o ano he Rhacopho id genus a he
han Rao ches es and Philau us. To u he con i m he
iden i y o he species, he newly gene a ed sequences
we e aligned wi h 34 samples om GenBank ep esen -
ing 15 hacopho id gene a and Bue ge ia oxycephala
(Schlegel, 1858) was used as an ou g oup (Benson e al.
2007; Table S5). Fo his da a se o 526 base pai s (16S),
we pe o med a hi d ML analysis using he GUI e sion
o IQTREE (Nguyen e al. 2015) implemen ed in PHY-
LOSUITE (Zhang e al. 2020).
Species delimi a ion
To delinea e pu a i e species bounda ies o no heas
Indian, Indo-Bu mese and Indochinese Rao ches es, we
pe o med wo di e en species delimi a ion me hods
namely: Mul i-Ra e Poisson T ee P ocesses (mPTP)
(Kapli e al. 2017) and Assemble Species by Au oma ic
Pa i ioning (ASAP) (Puilland e e al. 2021). ASAP is a
dis ance-based species delimi a ion me hod ha iden i ies
pu a i e species bounda ies based on pai wise gene ic dis-
ances. Whe eas mPTP accoun s o a ying coalescen
a es among clades, enabling i o e lec di e en le els
o in aspeci ic gene ic di e si y ac oss lineages. These
me hods ha e been ecen ly used in amphibian axono-
my (Oswald e al. 2023; Ca ne and Viei es 2024; A ias e
al. 2025). The conca ena ed mi ochond ial da ase (16S +
cy b + COI) was used o bo h he delimi a ion analyses.
The ML ee in e ed om conca ena ed mi ochond ial
da ase was u he used as he inpu ee o mPTP in
a web-se e accessible a h p://mp p.h-i s.o g and he
ollowing unc ions we e used: MCMC = 10,000,000,
minimum b anch leng h = minb _au o. The same con-
ca ena ed mi ochond ial da ase was used o ASAP in a
web-se e accessible a h ps://bioin o.mnhn. /abi/pub-
lic/asap. Fo he ASAP analysis, he subs i u ion model o
p dis ances was used o compu e he dis ances along wi h
he de aul alues o o he pa ame e s.
Ve eb a e Zoology 75, 2025, 517–625 521
Unco ec ed pai wise dis ances (p dis ances) we e cal-
cula ed in MEGA 7.1 wi h pai wise dele ion o missing
da a and gaps o each mi ochond ial gene (16S, cy b and
COI) sepa a ely. The pai wise gene ic dis ances o he
16S da ase we e used as a guide o lineage delimi a ion.
The gene ic di e gences (16S) we e conside ed as low
(1% o 2%), mode a e (>2% o <4%) and high (4 % o
6%) ollowing Vijayakuma e al. (2014).
Mo phological examina ion
Mo phome ic measu emen s we e aken o he nea -
es 0.1 mm using a digi al slide callipe (Mi u oyo CD-
6˝CSX). Me ic and me is ic cha ac e s we e examined
using a s e eomic oscope (Olympus SZX10). Sex and ma-
u i y we e de e mined based on ei he p esence o ocal
sac and nup ial pad in males, o by examina ion o go-
nads h ough a en ola e al incision. Measu emen s and
associa ed e minology ollowed Bo uah e al. (2023a),
unless o he wise s a ed. The ollowing measu emen s
we e aken: SVL (snou - en leng h), HW (head wid h
a angle o jaw), HL (head leng h om ea o mandi-
ble o ip o snou ), SL (snou leng h om ip o snou o
an e io o bi al ma gin), EN (dis ance om on o eye
o nos il), NS (dis ance om nos il o ip o snou ), IN
(in e na ial dis ance), EL (ho izon al eye leng h be ween
o bi al ma gins), IUE (sho es in e -uppe eyelid wid h
be ween uppe eyelids), UEW (maximum uppe eye-
lid wid h), IFE (in e nal on o eyes, sho es dis ance
be ween an e io o bi al ma gins), IBE (in e nal back o
eyes, sho es dis ance be ween pos e io o bi al ma gins),
HTYD (maximum ho izon al ympanum diame e ), MN
(dis ance be ween ea o mandible and pos e io bo de
o nos il), MAE (dis ance be ween ea o mandible and
an e io bo de o o bi (eye)), MPE (dis ance be ween
ea o mandible and pos e io bo de o o bi (eye)), FAL
( o ea m leng h om lexed elbow o base o ou e palma
ube cle), HAL (hand leng h om base o ou e palma u-
be cle o ip o hi d inge ), AG (dis ance be ween axilla
and g oin), TL ( high leng h om en o knee), TBL ( ib-
ia leng h om knee o heel), Ta L ( a sus leng h om i-
bio- a sal a icula ion o base o inne me a a sal ube cle),
FOL ( oo leng h om base o inne me a a sal ube cle o
ip o ou h oe), FL ( inge leng h om ip o inge o i s
base whe e i joins adjacen inge ), TOL ( oe leng h om
ip o oe o i s base whe e i joins adjacen oe), digi ( in-
ge and oe) numbe s a e ep esen ed by Roman nume als
I–V (inside o ou side), FD (disc wid h o inge s), TD
(disc wid h o oes). All measu emen s in he ex a e gi -
en in millime es. Measu emen s we e aken om he igh
side o he specimen. We also examined he skin ex u e o
he collec ed specimens b oadly ca ego ised as: ube cles
(small de mal p ojec ion wi h blun ip), spinules (small
de mal p ojec ion wi h sha ply poin ed ip) and g anules
on en al side o head, abdomen and limbs.
We colla ed body size (SVL) da a o o he congene s
om no heas India and Indochina and Indo-Bu ma (Table
S6). The da a we e plo ed using ggplo 2 (Wickham 2016)
in he p og am R .4.4 (R Co e De elopmen Team 2021).
We compa ed newly desc ibed species wi h he con-
gene s known om no heas India and Sou heas Asia
and he edesc ibed species using o iginal desc ip ions.
Ou mo phological compa isons we e based on measu e-
men s, skin ex u e, snou shape and colou pa e ns o he
geno yped specimens included in he phylogene ic ee
( Table 1A, B). Commonly used a ios o species diag-
nosis and compa isons in his s udy we e 1) head leng h
s. head wid h, 2) snou leng h s. eye leng h, 3) snou
leng h s. in e -uppe eyelid wid h, 4) in e -uppe eyelid
wid h s. eye leng h, 5) in e na ial dis ance s. uppe eye-
lid wid h, and 6) high leng h s. ibia leng h.
Mul i a ia e analyses
We used P incipal Componen Analysis (PCA) o assess
he o e all mo phological a ia ion be ween di e en
popula ions in he so wa e R (R Co e Team 2021) using
he R package MASS (Venables and Ripley 2002). Fo
his analysis, we used 38 mo phome ic a iables o 176
indi iduals collec ed om 81 locali ies ac oss no heas
India. We excluded emales, sub-adul s, and indi idu-
als o which cha ac e s we e damaged o no a ailable.
We used he G oupS uc R package (Chan and G isme
2022) o calcula e and accoun o allome ic e ec s in he
mo phome ic da ase , ollowing he allome ic equa ions
desc ibed by Tho pe (1975, 1983). This is o a oid size
dependen in e co ela ion e ec s, p io o conduc ing
PCA be ween he pu a i e axa wi hou making a p io i
assump ions abou g oupings. Mo phome ic da a we e
analysed in he so wa e R, and he package ggplo 2 was
used o p epa e plo s (Wickham 2009). The esul s we e
plo ed g oup wise excep o R. annandalii, R. minda
Dos e al., 2025 and wo pu a i e species wi h low nodal
suppo . These ou species we e plo ed along wi h o he
species om hei closes g oup.
We an ano he PCA using eigh mo phome ic mea-
su emen s (SVL, IUE, UEW, EL, HL, HW, TL, and TBL)
ou o he 38 mo phome ic measu emen s which we e
also a ailable o R. ga o (ZSI19187) and R. kempiae
(ZSI18859) ypes, o de e mine he mo phospace o hese
ypes in ela ion o he newly collec ed ma e ial o he
wo species.
Bioacous ic eco ding and analysis
We eco ded ocalisa ions in he ield using a digi al audio
eco de (Ma an z PMD 620 MK–II) wi h a unidi ec ion-
al handheld mic ophone (Sennheise MKH 416). Calls
we e eco ded a a dis ance o app oxima ely 30–50 cm
om he calling male. These calling indi iduals we e col-
lec ed and DNA sequences o he same indi iduals we e
used o subsequen phylogene ic analyses. We used RA-
VEN PRO .1.5 (h p://www.bi ds.co nell.edu/ a en) o
isualise and analyse he acous ic p ope ies ( as Fou ie
ans o m, wid h 1024 samples, Hann window wi h 50%
o e lap, 49.1 Hz esolu ion and 3db il e bandwid h).
The acous ic p ope ies and e minology ollow Köhle
Bo uah B e al.: Re ision o bush ogs om no heas India
522
Table 1A. Mo phological compa ison o Rao ches es species included in his s udy gi en as mean ± SD ( ange). NA=da a no a ailable.
Species SVL in male
(mm)
SVL in emale
(mm)
Snou Shape
(do sal iew)
HL/HW SL/EL SL/IUE IUE/EL Nos il posi ion
ela i e o eye and
snou
UEW/IN EL/HL HTYD/EL
R. ande soni 18.75±7.42
(13.5–24.0)
NA NA HL<HW NA NA IUE<EL close o snou ip NA NA 0.25
R. annandalii 18.58±0.85
(17.3–19.6)
20.34±1.12
(18.9–21.6)
ounded o sub
o oid
1.0±0.01
(0.98–1.01)
(1.05±0.07
(0.96–1.15)
1.1±0.1
(0.97–1.23)
0.95±0.06
(0.81–1.0)
close o eye o
equidis an
0.76±0.06
(0.65–0.84)
0.38±0.02
(0.35–0.42)
0.22±0.15
(0.25–0.36)
R. cine ascens
no . comb.
16 NA ounded NA SL<EL SL=IUE NA NA NA NA NA
R. dulongensis 16.7±2.0
(15.0–19.0)
NA ob usely poin ed 1.06±0.01
(1.05–1.07)
1.0±0.04
(0.97–1.04)
1.07±0.08
(1.0–1.15)
0.94±0.09
(0.87–1.04)
close o snou ip 0.76±0.05
(0.71–0.8)
0.43±0.04
(0.4–0.47)
0.49±0.07
(0.43–0.57)
R. ga o 24.54±1.8
(19.9–26.9)
26.9 ounded o sub
o oid
0.95±0.04
(0.91–1.06)
0.93±0.06
(0.81–1.03)
1.11±0.07
(0.96–1.28)
0.84±0.06
(0.74–0.94)
close o snou ip o
equidis an
0.86±0.07
(0.72–1.0)
0.41±0.02
(0.36–0.46)
0.24±0.07
(0.21–0.34)
R. hekouensis 17.0±0.7
(16.1–17.5)
21.1 ounded 0.9±0.05
(0.82–0.95)
0.77±0.19
(0.55–0.96)
0.85±0.19
(0.61–1.04)
0.9±0.04
(0.85–0.96)
NA 0.79±0.11
(0.65–0.9)
0.45±0.05
(0.41–0.53)
0.5±0.09
(0.35–0.6)
R. hillisi 16.4±1.0
(15.9–17.7)
17.5 ounded 1.13±0.08
(1.03–1.25)
1.21±0.08
(1.09–1.33)
1.21±0.06
(1.14–1.35)
1.0±0.07
(0.91–1.1)
close o snou ip 0.78±0.06
(0.69–0.88)
0.31±0.02
(0.69–0.88)
0.37±0.04
(0.32–0.43)
R. huanglianshan 18.2± 0.8
(17.0–19.6)
21.5 ounded 0.98±0.01
(0.95–1.0)
1.18±0.09
(1.0–1.33)
1.08±0.08
(0.96–1.22)
1.09±0.11
(0.95–1.28)
close o snou ip 0.73±0.05
(0.65–0.84)
0.35±0.02
(0.30–0.38)
0.40±0.05
(0.33–0.5)
R. jadoh 13.8± 0.23
(14.0–13.6)
NA ounded 0.55±0.02
(0.53–0.57)
1.7 1.37 1.24 close o snou ip 0.80±0.05
(0.77–0.83)
0.43 —
R. jakoid 20.0± 2.21
(18.4–21.6)
NA ounded 0.78±0.05
(0.74–0.81)
1.35±0.1
(1.34–1.36)
1.27±0.24
(1.1–1.4)
1.08±0.2
(0.94–1.2)
close o snou ip 0.78±0.02
(0.77–0.79)
0.32±0.04
(0.29–0.34)
—
R. kempiae 21.13±1.57
(17.8–22.9)
23.15±0.21
(23.0–23.3)
ounded o sub
o oid o poin ed
0.97±0.03
(0.91–1.01)
0.97±0.06
(0.82–1.04)
1.18±0.04
(1.12–1.28)
0.82±0.06
(0.71–0.93)
close o snou ip o
equidis an
0.82±0.09
(0.67–0.92)
0.4±0.02
(0.36–0.42)
0.27±0.03
(0.22–0.34)
R. leik ho 15.77±0.06
(15.7–15.8)
NA ounded 1.00±0.08
(0.91–1.08)
1.07±0.2
(0.89–1.29)
1.19±0.09
(1.14–1.29)
0.9±0.11
(0.79–1.0)
close o snou ip 0.83±0.22
(0.67–1.08)
0.44±0.05
(0.4–0.5)
0.32±0.07
(0.25–0.38)
R. longchuanensis 17.8–23.9 NA ounded HL=HW NA NA IUE>EL NA NA NA NA
R. malipoensis 16.8±1.0
(14.6–17.7)
18.8±0.4
(18.3–19.3)
ob usely poin ed 0.87±0.08
(0.74–1.0)
1.0±0.14
(0.83–1.23)
0.84±0.14
(0.62–1.0)
1.20±0.12
(1.0–1.38)
close o snou ip 0.69±0.11
(0.54–0.86)
0.38±0.06
(0.30–0.5)
0.59±0.08
(0.5–0.71)
R. menglaensis 18.87±1.27
(16.6–21.6)
19.7±1.13
(18.9–20.5)
ounded 1.0±0.24
(1–1.18)
1.04±0.26
(1.03–1.34)
1.1±0.27
(1.0–1.28)
0.9±0.22
(0.84–1.08)
close o snou ip 0.78±0.19
(0.68–0.95)
0.33±0.8
(0.30–0.38)
0.29±0.08
(0.24–0.39)
R. pa ulus 17.88±0.69
(17.0–18.6)
23.0 ounded 1.01±0.03
(0.98–1.05)
0.95±0.16
(0.70–1.08)
1.1±0.15
(0.88–1.24)
0.86±0.03
(0.8–0.88)
close o snou ip o
equidis an
0.94±0.13
(0.7–1.0)
0.41±0.4
(0.36–0.45)
0.29±0.09
(0.23–0.25)
R. ezakhani 20.25±0.86
(18.8–21.0)
NA ounded 0.71±0.14
(0.64–0.95)
0.9±0.07
(0.83–1.0)
1.04±0.14
(0.94–1.25)
0.86±0.05
(0.8–0.9)
close o snou ip o
equidis an
0.9±0.04
(0.85–0.95)
0.54±0.07
(0.41–0.59)
0.43±0.06
(0.33–0.48)
R. shillongensis 17.02±1.51
(13.9–20.4)
18.52±1.56
(15.1–21)
ounded o sub
o oid o nea ly
poin ed
0.95±0.04
(0.92–1.0)
0.99±0.05
(0.93–1.05)
1.13±0.04
(1.08–1.16)
0.88±0.05
(0.81–0.93)
close o snou ip o
equidis an
0.83±0.06
(0.76–0.89)
0.4±0.03
(0.36–0.44)
0.29±0.05
(0.22–0.33)
R. y hus no . comb. NA 20.9–21.2 ounded HL=HW SL>EL SL<IUE IUE>EL close o snou ip IN>UEW NA NA
R. yadongensis 20.3±2.7
(17.8–24.1)
NA ob usely poin ed 0.71±0.2
(0.51–0.9)
1.01±0.02
(1.0–1.03)
0.74±0.09
(0.61–0.82)
1.38±0.17
(1.27–1.63)
close o snou ip 1.04±0.08
(1.0–1.17)
0.57±0.08
(0.44–0.81)
0.32±0.04
(0.26–0.35)
R. a unachalensis
sp. no .
23.57±1.87
(19.6–26.3)
25.0±3.82
(22.3–27.7)
ounded o sub
o oid
0.98±0.03
(0.92–1.03)
0.9±0.06
(0.8–1.05)
1.13±0.07
(1.03–1.25)
0.8±0.05
(0.67–0.89)
close o snou ip 0.93±0.09
(0.71–1.04)
0.41±0.02
(0.38–0.45)
0.27±0.04
(0.21–0.34)
R. ba akensis sp. no . 18.92±1.17
(16.7–21.0)
NA ounded o sub
o oid
0.98±0.04
(0.89–1.02)
0.95±0.05
(0.82–1.04)
1.12±0.09
(0.96–1.27)
0.85±0.05
(0.72–0.96)
close o snou ip o
equidis an
0.8±0.05
(0.7–0.91)
0.41±0.02
(0.38–0.45)
0.29±0.05
(0.19–0.38)
Ve eb a e Zoology 75, 2025, 517–625 523
Species SVL in male
(mm)
SVL in emale
(mm)
Snou Shape
(do sal iew)
HL/HW SL/EL SL/IUE IUE/EL Nos il posi ion
ela i e o eye and
snou
UEW/IN EL/HL HTYD/EL
R. boulenge i sp. no . 18.18±0.68
(17.5–19.1)
NA ounded 1.0±0.02
(0.98–1.02)
0.97±0.06
(0.89–1.04)
1.15±0.14
(1.04–1.35)
0.85±0.05
(0.77–0.88)
equidis an 0.8±0.04
(0.75–0.85)
0.41±0.03
(0.38–0.45)
0.31±0.03
(0.28–0.35)
R. dibangensis
sp. no .
20.7±1.56
(19.6–21.8)
NA ounded 1.0±0.02
(0.96–1.03)
0.93±0.04
(0.86–1.0)
1.12±0.06
(1.03–1.24)
0.83±0.04
(0.77–0.91)
close o snou ip 0.96±0.07
(0.81–1.04)
0.37±0.03
(0.35–0.39)
0.24±0.05
(0.21–0.28)
R. eaglenes ensis sp.
no .
21.96±1.24
(18.8–23.6)
22.35±0.64
(21.9–22.8)
ounded o sub
o oid
0.99±0.04
(0.94–1.07)
0.93±0.06
(0.86–1.07)
1.02±0.04
(0.97–1.1)
0.91±0.05
(0.84–0.97)
close o snou ip o
equidis an
0.84±0.12
(0.5–0.92)
0.41±0.02
(0.37–0.43)
0.28±0.04
(0.19–0.33)
R. minda 17.7±0.61
(16.7–18.3)
NA ounded 0.97±0.07
(0.88–1.07)
0.98±0.09
(0.92–1.18)
1.16±0.35
(0.92–1.9)
0.89±0.19
(0.5–1.05)
close o snou ip o
equidis an
0.73±0.09
(0.6–0.84)
0.4±0.03
(0.35–0.43)
0.32±0.07
(0.27–0.46)
R. khonoma
sp. no .
19.23±2.43
(16.9–22.0)
NA ounded 0.98±0.03
(0.94–1.0)
0.97±0.04
(0.93–1.0)
0.99±0.05
(0.94–1.04)
0.98±0.02
(0.96–1.0)
equidis an 0.94±0.08
(0.75–0.91)
0.4±0.01
(0.39–0.40)
0.33±0.02
(0.31–0.35)
R. lawng laiensis sp.
no .
18.9±1.13
(18.1–19.7)
NA ounded 0.96±0.01
(0.95–0.97)
0.83 1.07±0.03
(1.04–1.09)
0.78±0.03
(0.76–0.8)
equidis an 0.95 0.44±0.03
(0.42–0.46)
0.22±0.02
(0.21–0.23)
R. magnus
sp. no .
24.37±1.71
(22.5–27.4)
26.95±2.05
(25.5–28.4)
ounded o sub
o oid
1.0±0.02
(0.96–1.03)
0.93±0.04
(0.86–0.97)
1.12±0.06
(1.03–1.24)
0.83±0.03
(0.77–0.91)
close o snou ip 0.96±0.07
(0.81–1.04)
0.4±0.02
(0.38–0.43)
0.28±0.04
(0.21–0.36)
R. mawsyn amensis
sp. no .
17.26±0.99
(16.0–18.7)
20 ounded 0.97±0.03
(0.92–1)
0.93±0.06
(0.87–1)
1.07±0.06
(0.96–1.15)
0.87±0.06
(0.8–0.96)
equidis an 0.77±0.09
(0.62–0.89)
0.41±0.01
(0.38–0.43)
0.29±0.04
(0.24–0.37)
R. monoli hus sp. no . 19.57±1.07
(17.9–20.6)
NA ounded 0.94±0.03
(0.89–0.97)
0.97±0.05
(0.89–1.0)
0.99±0.03
(0.96–1.04)
0.98±0.03
(0.93–1.0)
equidis an 0.77±0.08
(0.67–0.89)
0.4±0.02
(0.38–0.43)
0.24±0.12
(0.22–0.32)
R. na puhensis
sp. no .
19.81±0.87
(18.0–21.1)
23.1 sub o oid o
nea ly acu e
0.99±0.03
(0.94–1.01)
0.94±0.07
(0.81–1.03)
1.09±0.07
(0.96–1.21)
0.86±0.04
(0.8–0.93)
close o snou ip 0.82±0.1
(0.68–0.95)
0.41±0.02
(0.39–0.46)
0.16±0.12
(0.19–0.31)
R. nasu a
sp. no .
18.6±1.16
(17.0–19.9)
NA acu e 1.06±0.05
(1.03–1.11)
1.02±0.02
(1–1.04)
1.09±0.05
(1.04–1.17)
0.93±0.03
(0.88–0.96)
equidis an 0.81±0.03
(0.76–0.86)
0.38±0.01
(0.37–0.39)
0.31±0.02
(0.29–0.31)
R. o ien alis
sp. no .
18.67±1.49
(14.6–20.8)
20.85±1.48
(19.8–21.9)
ounded o sub
o oid
0.99±0.04
(0.9–1.07)
0.99±0.05
(0.92–1.07)
1.14±0.1
(0.96–1.32)
0.88±0.08
(0.73–1.04)
close o snou ip o
equidis an
0.81±0.1
(0.64–0.95)
0.39±0.02
(0.35–0.42)
0.28±0.06
(0.21–0.41)
Table 1B. Mo phological compa ison o Rao ches es species included in his s udy gi en as mean ± SD ( ange). NA=da a no a ailable.
Species Nup ial pad TBL/TL TL/SVL Skin on
do sum
Conca e s i-
pes on back
Pa e n on
g oin
Mid do sal line Bands on
hindlimbs
Pa e n on
abdomen
Whi e band
on in e o bi al
space
Re e ences
R. ande soni NA NA NA sca e ed
ube cles
absen (?) la ge da k
i egula spo
wi h wo ligh
spo s
NA single band b own punc-
ua ions wi h
yellow g anules
pale yellow
bands may
be p esen o
absen
Ande son (1978),
Wu e al. (2021)
R. annandalii F1 0.96±0.02
(0.93–1.0)
0.5±0.02
(0.48–0.54)
sca e ed
ube cles
p esen absen absen one o h ee
bands (ou e
wo bands may
be ain )
whi ish spo s p esen o
absen
This s udy
R. cine ascens
no . comb.
NA TL>TBL NA sca e ed
ube cles
p esen a da k spo pa -
ially enci cled
by ligh inge
absen h ee bands no pa e n absen S oliczka (1870),
his s udy
Bo uah B e al.: Re ision o bush ogs om no heas India
524
Species Nup ial pad TBL/TL TL/SVL Skin on
do sum
Conca e s i-
pes on back
Pa e n on
g oin
Mid do sal line Bands on
hindlimbs
Pa e n on
abdomen
Whi e band
on in e o bi al
space
Re e ences
R. dulongensis absen 0.96±0.08
(0.88–1.03)
0.49±0.03
(0.46–0.52)
nume ous
ube cles
p esen da k pa ches
be ween wo
ligh pa ches
p esen o
absen
h ee bands da k-b own and
whi e spo s
absen Wu e al. (2021)
R. ga o F1 0.96±0.03
(0.91–1.01)
0.51±0.02
(0.46–0.54)
mode a e o
hea y spinules;
may be in e mi-
xed wi h blun
ube cles
absen (hou -
glass shaped
ma king may
be p esen )
da k pa ches
wi h o wi hou
ligh spo s
p esen o
absen
h ee bands pale-c eam
colou ed blo -
ches p esen o
absen
p esen o
absen
Boulenge (1919),
his s udy
R. hekouensis F1 and F2 0.81 (da a
a ailable only
o holo ype)
0.53 (da a
a ailable only
o holo ype)
nume ous
ube cles
p esen da k pa ches
be ween wo
ligh pa ches
p esen o
absen
h ee bands small black
spo s and whi e
ube cles
absen Du e al. (2024)
R. hillisi F1 1.03±0.05
(0.98–1.12)
0.46±0.03
(0.42–0.5)
sca e ed
ube cles
p esen a da k blo ch p esen o
absen
h ee bands whi ish spo s absen Jiang e al. (2020)
R. huanglianshan F1 0.98±0.03
(0.93–1.0)
0.44±0.02
(0.41–0.50)
sca e ed
ube cles
p esen a da k blo ch p esen o
absen
single band whi ish spo s absen Jiang e al. (2020)
R. jadoh F1 1.13±0.1
(1.06–1.2)
0.49±0.01
(0.48–0.49)
small wa s
sca e ed
p esen — absen 1 (?) no pa e n absen Wa j i e al. (2025)
R. jakoid F1 1.19±0.15
(1.08–1.3)
0.40±0.02
(0.38–0.42)
small wa s
sca e ed
p esen — absen 3 (?) no pa e n absen Wa j i e al. (2025)
R. kempiae F1 0.98±0.03
(0.95–1.07)
0.51±0.02
(0.48–0.55)
spinules
in e mixed wi h
ube cles
absen o ain a sho da k
s ipe
p esen o
absen
h ee bands bu
excep middle
ba , ou e wo
bands ba ely
isible
no pa e n absen Boulenge (1919),
his s udy
R. leik ho F1 1.03±0.06
(0.98–1.09)
0.51±0.04
(0.47–0.55)
nume ous
spinules
Absen (a pai
o an e io ly
me ged s ipes
p esen )
a sho b ow-
nish oli e ba
absen one o h ee
bands
no pa e n p esen o
absen
Köhle e al. (2025)
R. longchuanensis F1 NA NA NA absen o indis-
inc
NA NA NA NA NA Jiang e al. (2020),
Wu e al. (2021),
Huang e al. (2023)
R. malipoensis F1 1.08±0.09
(0.94–1.21)
0.46±0.04
(0.42–0.54)
small ube cles
p esen
p esen da k pa ches absen h ee bands whi e and black
lecks
absen Huang e al. (2023)
R. menglaensis F1 0.98±0.24
(1.0–1.09)
0.45±0.11
(0.40–0.51)
sca e ed
ube cles
p esen a da k blo ch p esen o
absen
h ee bands whi ish spo s absen Jiang e al. (2020)
R. pa ulus NA 1.04±0.04
(0.99–1.1)
0.45±0.01
(0.43–0.46)
sca e ed
spinules
p esen da k ma bling
enclosing a
ligh blo ch
absen one on high
and one o wo
on ibia
b own and whi-
e mo ling
absen Boulenge (1893),
Köhle e al. (2025)
R. ezakhani F1 0.94±0.06
(0.85–0.99)
0.52±0.04
(0.48–0.59)
sca e ed
ube cles
p esen sho da k s ipe absen h ee bands black o whi e
spo s
absen Al-Razi e al. (2020),
his s udy
Ve eb a e Zoology 75, 2025, 517–625 525
Species Nup ial pad TBL/TL TL/SVL Skin on
do sum
Conca e s i-
pes on back
Pa e n on
g oin
Mid do sal line Bands on
hindlimbs
Pa e n on
abdomen
Whi e band
on in e o bi al
space
Re e ences
R. shillongensis F1 0.92±0.02
(0.89–0.94)
0.48±0.01
(0.47–0.50)
ube cles mode-
a e o hea y
p esen ( a ely
hou -glass
shaped pa ch)
da k pa ch wi h
ligh c own-
shaped ma k o
di used da k
pa ch
absen one b oad band
o h ee bu
ou e wo may
be di used/
indis inc
whi e and black
blo ches p esen
o absen
absen This s udy
R. y hus
no . comb.
NA TL>TBL NA sca e ed
ube cles
p esen enla ged ligh
pa ch
absen single band i egula da k
and ligh spo s
absen Smi h (1940), his
s udy
R. yadongensis F1 NA NA small ube cles
p esen
p esen ( ain ) NA absen single indis inc
band
whi e pa ches absen Zhang e al. (2022)
R. a unachalensis
sp. no .
F1 0.96±0.02
(0.92–1.0)
0.54±0.02
(0.51–0.58)
mode a e o
hea ily ube -
cula ed; may be
in e mixed wi h
spinules
p esen da k and ligh
pa ches
absen h ee bands whi e blo ches
p esen o
absen
p esen o
absen
This s udy
R. ba akensis
sp. no .
F1 0.95±0.03
(0.9–0.99)
0.52±0.03
(0.47–0.57)
dense spinules p esen absen absen h ee bands no pa e n absen This s udy
R. boulenge i
sp. no .
F1 0.93±0.03
(0.9–0.97)
0.51±0.01
(0.49–0.51)
dense spinules p esen da k and ligh
pa ches
absen h ee bands
( ain )
whi ish spo s absen This s udy
R. dibangensis
sp. no .
F1 0.94±0.01
(0.93–0.94)
0.49±0.01
(0.48–0.50)
dense spinules
on middle o
back
p esen (indis-
inc )
absen absen wo bands
( ain )
whi ish spo s p esen o
absen
This s udy
R. eaglenes ensis
sp. no .
F1 0.96±0.02
(0.92–0.99)
0.51±0.03
(0.48–0.57)
dis inc u-
be cles
p esen absen absen h ee bands whi e blo ch
p esen o
absen
absen This s udy
R. minda F1 1.03±0.12
(0.91–1.25)
0.46±0.04
(0.41–0.52)
dense spinules p esen (Indis-
inc )
enla ged da k
and ligh
pa ches
absen one o h ee
bands
whi ish spo s absen Köhle e al. (2025),
his s udy
R. khonoma
sp. no .
F1 0.94±0.02
(0.92–0.96)
0.49±0.02
(0.47–0.51)
sca e ed
ube cles
absen o ain da k and ligh
pa ches
absen single band whi e spo s
wi h o wi hou
black spo s
absen This s udy
R. lawng laiensis
sp. no .
F1 0.93±0.01
(0.92–0.94)
0.5±0.01
(0.49–0.51)
mode a ely
dense spinules
p esen da k band wi h
ligh edge
absen single band whi ish spo s absen This s udy
R. magnus
sp. no .
F1 0.95±0.03
(0.92–1.01)
0.52±0.02
(0.48–0.55)
spinules
in e mixed wi h
ube cles
p esen enla ged da k
pa ch
p esen o
absen
h ee bands da k b own
blo ches p esen
o absen
p esen o
absen
This s udy
R. mawsyn amensis
sp. no .
F1 0.94±0.02
(0.9–0.96)
0.49±0.02
(0.48–0.52)
dense spinules p esen da k pa ches absen h ee bands
( ain )
whi ish spo s absen This s udy
R. monoli hus
sp. no .
F1 0.89±0.01
(0.87–0.9)
0.49±0.02
(0.47–0.52)
sca e ed
ube cles
ba ely isible enla ged da k
pa ch
absen single band whi ish spo s absen This s udy
Bo uah B e al.: Re ision o bush ogs om no heas India
532
Figu e 4. A, B Maximum Likelihood ee in e ed om conca ena ed mi ochond ial da ase . Values a he in e nal b anches a e
Ul a-Fas Boo s ap (UFB) suppo . Ou g oups a e no shown in his igu e. Colou ed ba s on he igh a e lineages delimi ed by
ASAP and mPTP me hods. Scale ba : subs i u ion pe si e. Values abo e ASAP ba s a e he anks gi en by ASAP analysis.
Ve eb a e Zoology 75, 2025, 517–625 533
Figu e 4 (con inued).
Bo uah B e al.: Re ision o bush ogs om no heas India
534
dalii, R. ba akensis sp. no ., R. boulenge i sp. no ., R.
minda , R. hekouensis, R. hillisi, huanglianshan, lawng -
laiensis sp. no ., R. mawsyn amensis sp. no . and R. ma-
lipoensis a e smalle in body-size compa ed o he o he
species (Fig. 5). Whe eas R. magnus sp. no ., R. ga o and
R. a unachalensis sp. no . a e ela i ely la ge (Fig. 5).
Al hough R. ga o and R. cangyuanensis a e gene ically
simila wi h e y low di e gences (0.0–0.8% in he 16S
gene) (Figs 2, 3) he popula ion om no heas India is
la ge in body size han he ones om China (Fig. 5). This
could be a esul o geog aphic sepa a ion and egional
a ia ion be ween hese wo popula ions. The e o e, body
size alone canno be a eliable cha ac e o dis inguish
some o he widely dis ibu ed species in his complex.
Mul i a ia e analyses
P incipal Componen Analysis (PCA) compu ed o all
38 mo phome ic cha ac e s o males e ealed he sp ead
o mo phological a ia ion in he desc ibed and pu a i e
species (gene ic lineages) om no heas India. Nin y i e
pe cen o he a ia ion in he PCA was explained by 15
PCA axes. PCA axes one (PC1) and wo (PC2) explained
mos o he a ia ion (73.9%) in he da ase (Table S8).
On he i s PC axis (PC1), inge leng hs (FLI, FLII and
FLIV) and i h oe leng h had highe ac o loadings
(Table S9). The a ia ion in PC axis (PC2) wo was less
a iable compa ed o PC1 o he 18 pu a i e lineages in-
cluded in his analysis (Fig. 6). In he R. ga o g oup, he
o dina ion plo showed clea mo phological sepa a ion
be ween R. ga o and o he h ee lineages wi h no o e -
lap (Fig. 6A). The o he h ee lineages in his g oup ha e
pa ial o e lap among hem (Fig. 6A). Among he pu a-
i e lineages o R. ezakhani g oup we s udied, all he
membe s ha e pa ial o e lap in mo phospace. Rao ches-
es mawsyn amensis sp. no . is di e en among he ou
species and has mino o e lap wi h R. ba akensis sp.
no . (Fig. 6B). Among he pu a i e lineages o he R.
kempiae g oup, R. shillongensis is dis inc om all he
o he lineages (R. kempiae, R. dibangensis sp. no . and
R. monoli hus sp. no .) o he g oup while R. kempiae
has pa ial o e lap wi h he o he wo lineages. In he R.
annandalii g oup, he lineages ha e no o e lap excep he
o e lap be ween R. a unachalensis sp. no . and R. mag-
nus sp.no . (Fig. 6D). Sample size o R. boulenge i sp.
no ., R. dibangensis sp. no ., R. khonoma sp. no ., and
R. lawng laiensis sp. no . species is low (2–4) he e o e
hese esul s may a y in u u e wi h addi ion o mo e
samples.
In he PCA esul s o R. ga o and R. kempiae, 95% o
he a ia ion is explained by PC1 o PC5. Axes one (PC1)
Figu e 5. Visualisa ion ( iolin plo supe imposed on a box plo ; indi idual da a poin s a e o e laid as do s) o body size di e ences
o he male Rao ches es species desc ibed in his s udy and hei congene s om Indo-Bu ma and Indochina a e a anged alphabe -
ically. “Rao ches es cangyuanensis” and “R. kempiae” in Na een e al. (2024) a e synonyms o R. ga o. “Rao ches es asakg ensis ”
is a synonym o R. kempiae . Rao ches es sp1 is an undesc ibed species misiden i ied as R. ga o by Na een e al. (2024). The shape
o each iolin plo ep esen s he dis ibu ion densi y o he da a. Wide sec ions indica e a highe concen a ion o obse a ions,
while na owe sec ions e lec ewe da a poin s.
Ve eb a e Zoology 75, 2025, 517–625 535
and wo (PC2) explained mos o he a ia ion (78.4%)
in he da ase (Table S8B). Despi e one o he ype spec-
imens (ZSI19187) being a subadul , he e is a clea sepa-
a ion be ween he R. ga o and R. kempiae samples (Fig.
6E). Type specimens o bo h he species a e well clus e ed
wi h hei espec i e opo ypes. This u he emphasizes
he ac ha he newly collec ed ma e ials a e co ec ly
alloca ed o he espec i e species.
Bioacous ics
The ad e isemen calls o he 12 species included in his
s udy a e o a single ype. Calls can be b oadly di ided
in o pulsa ile and non-pulsa ile calls. Th ee species (R.
ba akensis sp. no ., R. dibangensis sp. no . and R. kempi-
ae) ha e pulsa ile calls while he emaining species ha e
non-pulsa ile calls (Table 2). Rao ches es eaglenes ensis
Figu e 6. P incipal Componen Analysis (PCA) o 176 specimens o Rao ches es using 38 mo phome ic cha ac e s (A–D). A Ra-
o ches es ga o g oup, B Rao ches es ezakhani g oup, C Rao ches es kempiae g oup + R. khonoma sp. no . + R. eaglenes ensis sp.
no ., D Rao ches es annandalii g oup + R. minda + R. a unachalensis sp. no . + R. magnus sp. no . Sis e species plo ed wi h
same symbols bu wi h di e en colou s. E PCA o R. ga o and R. kempiae specimens using eigh mo phome ic cha ac e s.
Bo uah B e al.: Re ision o bush ogs om no heas India
536
sp. no . and R. mawsyn amensis sp. no . ha e he
sho es call du a ion (9–20 ms) and R. ba akensis
sp. no . has he longes call du a ion (91–4464 ms).
Among he species wi h pulsa ile calls, R. ba ak-
ensis sp. no . has 2–59 pulses/call while R. diban-
gensis sp. no . and R. kempiae has 2–4 pulses/calls.
Rao ches es mawsyn amensis sp. no . and R. o ien-
alis sp. no . ha e he highes dominan equency
calls while R. a unachalensis sp. no ., R. ga o and
R. eaglenes ensis sp. no . ha e he lowes dominan
equency calls (Table 2). Ad e isemen calls o he
species s udied a e in suppo o phylogene ically
iden i ied lineages. This makes i a use ul ool o
ield iden i ica ion o bush ogs in no heas India.
Taxonomic conclusions
Based on he phylogene ic esul s wi h suppo o
mo phology, we e e all he collec ed samples o
bush ogs in his s udy o he genus Rao ches es ex-
cep o “R. sahai” which we ans e ed o he ge-
nus Nasu ixalus and synonymise his species wi h N.
je donii (see ollowing sec ion). Addi ionally, based
on ou axonomic app oach we desc ibe hi een new
species o Rao ches es om no heas India. In he
ollowing sec ion, we i s discuss p e iously de-
sc ibed species wi h ambiguous iden i y, ollowed by
new species desc ip ions.
Nasu ixalus je donii (Gün he , 1876)
Figu es 7, 8; Tables S10, S11
Synonymy and ch esonymy.
Polypeda es je donii Gün he , 1876: 571.
Rhacopho us je donii — Boulenge (1882): 80; Das and Du a
(1998): 67.
Rhacopho us (Rhacopho us) je donii — Ahl (1931): 114; Du a
(1997): 101.
Rhacopho us (Rhacopho us) bue ge i je donii — Wol (1936):
172.
Philau us (Ki ixalus) je donii — Dubois (1987): 73.
Philau us (Philau us) je donii — Bossuy and Dubois (2001): 25.
Philau us sahai Sa ka & Ray, 2006: 303.
Pseudophilau us sahai — Li e al. (2009): 519.
Philau us sahai — Ahmed e al. (2009): 16.
Rao ches es sahai — Biju e al. (2010): 1120.
F ankixalus je donii — Biju e al. (2016): 6–13.
Nasu ixalus je donii — Si ongxay e al. (2016): 439–440.
Commen s on axonomic s a us. Rao ches es sahai
(Fig. 7) was o iginally desc ibed as Philau us sahai
by Sa ka and Ray (2006) based on wo specimens
collec ed by S. S. Saha on 29 Feb ua y 1988 om
10 km no h o Gandhig am, Changlang Dis ic ,
A unachal P adesh, India. A e Biju e al. (2010),
Table 2. Compa ison o ad e isemen calls o Rao ches es species o no heas India desc ibed in his s udy and R. ezakhani om Bangladesh.
Species Call deli e y pa e n Tempo al
s uc u e
No. o calls
pe g oup
Call du a ion
(ms)
Call ise ime
(ms)
Call all ime
(ms)
In e call
in e al (ms)
Pulses/call Pulse du a-
ion
Pulse a e
(pulses/sec)
Dominan
equency
(Hz)
Re e ence
R. a unachalensis
sp. no .
deli e ed in g oup non-pulsa ile 2–33 15–25 1–2 14–24 125–158 — — — 2411–2842 This s udy
R. ba akensis
sp. no .
no deli e ed in g oup pulsa ile — 91–4464 72–2327 19–2137 534–2281 2–59 10–34 11.73–14.08 3445–3832 This s udy
R. boulenge i
sp. no .
deli e ed in g oup non-pulsa ile 2 20–31 1 20–30 173–216 — — — 3703–3789 This s udy
R. dibangensis sp. no . no deli e ed in g oup pulsa ile — 58–127 1–50 11–126 48–514 2 o 3 10–15 16.95–22.22 3143–3316 This s udy
R. eaglenes ensis
sp. no .
deli e ed in g oup non-pulsa ile 2–4 9–20 1 12–18 224–270 ———2971–3100 This s udy
R. ga o deli e ed in g oup non-pulsa ile 2–13 20–32 1 19–31 144–170 ———2799–2971 This s udy
R. kempiae no deli e ed in g oup pulsa ile — 67–200 1 64–199 398–1695 2–4 9–37 18.52–21.74 3057–3488 This s udy
R. mawsyn amensis
sp. no .
no deli e ed in g oup non-pulsa ile — 13–20 1 12–19 1127–2133 — — — 4263–4608 This s udy
R. monoli hus
sp. no .
deli e ed in g oup non-pulsa ile 5–13 12–29 1 11–28 118–285 — — — 3316–3531 This s udy
R. na puhensis sp. no . no deli e ed in g oup non-pulsa ile — 13–25 1 12–24 544–823 — — — 3574–4134 This s udy
R. o ien alis
sp. no .
deli e ed in g oup non-pulsa ile 2–6 15–28 1–3 13–27 162–295 — — — 3617–4177 This s udy
R. ezakhani NA pulsa ile NA NA NA NA NA 5–11 3–29 10–19 4320–4770 Al-Razi e al. (2020)
R. shillongensis deli e ed in g oup non-pulsa ile 2–59 12–22 1 11–21 215–281 — — — 3660–3746 This s udy
Ve eb a e Zoology 75, 2025, 517–625 537
his species was placed unde Rao ches es by implica-
ion (F os 2025). Since i s desc ip ion, his species has
no been epo ed om he ype locali y and no a emp
has been made o s udy he ypes o compa e hem wi h
o he species in Rao ches es o o he gene a o small ha-
copho ids. We examined he ype se ies (ZSIA8500 and
ZSIA8501) a ailable a he Zoological Su ey o India,
Kolka a. Ou examina ion o he ypes e ealed some
cha ac e is ics ha con adic he o iginal desc ip ion o
he species by Sa ka and Ray (2006), such as ome ine
ee h p esen in bo h ypes al hough unde de eloped ( s.
ome ine ee h absen in o iginal desc ip ion) (Fig. 7);
snou unca ed in do sal and la e al iew, ounded in
en al iew ( s. snou ounded in o iginal desc ip ion);
udimen a y webbing p esen on hand ( s. no men ioned
in o iginal desc ip ion). Thus, p esence o ome ine ee h
in he ype specimens sugges ha his species does no
belong o he nominal genus Rao ches es. Be ween he
yea s 2022 and 2023, we collec ed h ee hacopho id
specimens om Gandhig am ha mo phologically co -
espond o he ype specimens o R. sahai based on he
ollowing se o cha ac e s such as unca ed snou , p o-
uding eyes, ome ine ee h p esen , ympanum dis inc ,
udimen a y webbing p esen on hand, webbing well de-
eloped on oo , disc on inge s and oes, abdomen and
high g anula (Figs 7 and 8).
Fu he mo e, in ou phylogene ic analysis (ML) based
on he 16S mi ochond ial gene, he newly collec ed sam-
ples (o bo h subadul and adul ) o med a clade wi h Na-
su ixalus je donii (Gün he , 1876) om no heas India
(Fig. 8A). The unco ec ed p dis ance be ween he newly
collec ed samples and a ailable sequences o N. je donii
om India included in his s udy a e 0.0–0.2% in he 16S
gene (Table S10).
We also examined he ype specimens o N. je donii
a he Na u al His o y Museum, London collec ion
(BMNH1947.2.7.84 and BMNH1947.2.7.85). Ou opo-
ypes ag eed wi h he N. je donii ypes based on he ol-
lowing se o mo phological cha ac e s: 1) snou ounded
o almos unca e in do sal iew and nea ly e ical in la -
e al iew; 2) ympanum dis inc ; 3) small webbing p es-
en on hand; 4) ci cum-ma ginal g oo e p esen on inge
and oe disc; 5) oe webbing mode a e; 6) g anula belly
and en al aspec o high; 7) da k pa ch co e ing pos e-
io pa o head on do sal aspec , con inuing o do sum
and pos e io ly bi u ca e. The ollowing excep ions o
a ia ions we e obse ed in he newly collec ed ma e ial
and p e iously published desc ip ions. Biju e al. (2016)
men ioned ha ome ine idges p esen in wo oblique
se ies wi hou ee h. Howe e , he o iginal desc ip ion
men ioned ha ome ine ee h a e p esen . Based on ou
specimens collec ed om di e en pa s o no heas In-
dia (Meghalaya, Mizo am, and A unachal P adesh), mos
o he specimens o N. je donii ha e blun ome ine ee h
on wo oblique se ies. In some o he specimens ( o ex-
ample WII-ADA3240) which is gene ically simila , om-
e ine idge is p esen wi hou ee h. Addi ionally, Jiang e
al. (2016) men ioned ha ome ine ee h a e p esen in N.
medogensis Jiang, Wang, Yan & Che, 2016, while Yang
and Chan (2018) men ioned ome ine idge p esen wi h-
ou ee h in N. yingjiangensis Yang & Chan, 2018. Thus,
his cha ac e is no consis en wi hin he genus and spe-
cies. Biju e al. (2016) men ioned ha me aca pal ube -
cles a e absen . Howe e , me aca pal ube cles a e isible
in he ypes as well as in ou newly collec ed specimens.
Based on he mo phological simila i y o he opo ypes o
he holo ype and pa a ype and he phylogene ically close
posi ion o he opo ypes o N. je donii, we conside “R.
sahai” o be a junio subjec i e synonym o N. je donii.
Howe e , none o he a ailable DNA sequences o N. je -
donii by Biju e al. (2016) and Muansanga e al. (2022)
a e om he ype locali y in Da jeeling. Those a ailable
sequences a e om he hill anges o he sou he n slope
o he B ahmapu a Valley. The e o e, opo ypic samples
a e needed o de e mine whe he N. je donii sensu s ic o
om Da jeeling and he popula ions om sou h o B ah-
mapu a Valley cu en ly e e ed o as N. je donii belong
o he same species o ep esen di e en species.
Sa ka and Ray (2006) placed his species in he genus
Philau us p obably because o he small body size o he
collec ed ma e ial. Al hough Sa ka and Ray (2006) e-
e ed o he ype se ies as adul s, we con i med ha he
ypes a e subadul s based on he newly collec ed subadul
and adul specimens om he ype locali y a Gandhi-
g am. Addi ionally, his species was ne e compa ed wi h
N. je donii in he pas p obably because bo h he species
we e placed in di e en gene a. Fo axonomic s abili y,
we p o ide he e an expanded desc ip ion o N. je donii
based on he newly collec ed ma e ial om he ype lo-
cali y o “R. sahai”.
Ma e ials examined. A subadul emale (WII-
ADA1770) and an adul male (WII-ADA1773) collec ed
by BB, AD and NGP on 16 Sep embe 2022 a Gand-
hig am (27.265139°N, 96.937041°E, ele a ion 1120 m
a.s.l.), Changlang Dis ic , A unachal P adesh; an adul
male (WII-ADA3240) collec ed om same locali y by
AD, SD, RNV and JDG on 23 May 2023.
Desc ip ion o newly collec ed male specimen (WII-
ADA1773; Fig. 8B–H). Medium sized hacopho id og,
SVL 41 mm; head sligh ly wide han long (HW/HL =
0.98); snou ounded in do sal and semici cula in en al
iew, e ical in la e al iew; snou leng h smalle han
eye leng h (SL/EL = 0.9); can hus os alis indis inc ,
oblique; lo eal conca e; nos ils o al, la e ally posi ioned
and obliquely o ien ed; nos il close o snou ip han eye;
snou an e io ly dep essed a in e na ial space; in e na ial
space equal o in e -uppe eyelid wid h and uppe eyelid
wid h; eyes p o uding, mode a e in size, less han hal o
head leng h (EL/HL = 0.42); ympanum dis inc , ound,
nea ly hal o eye leng h (HTYD/EL = 0.47); ongue pos-
e io ly no ched; choanae ound; ome ine ee h p esen
be ween choanae and angula o body axis; a pai o in-
e nal ocal sac openings on lowe jaw; habi us s ou , i s
leng h nea ly hal o snou - en leng h (AG/SVL = 0.47).
Fo elimbs slende ; o ea m leng h smalle han hand
leng h (FAL/HAL = 0.71); hi d inge longes ; inge s
wi h ounded disc; disc on hi d and ou h inge nea -
ly equal o ympanic diame e ; ci cum-ma ginal g oo e
Bo uah B e al.: Re ision o bush ogs om no heas India
538
p esen on disc; suba icula ube cles ounded and dis-
inc ; supe nume a y ube cles p esen ; palma ube cles
dis inc , o al shaped; udimen a y webbing p esen be-
ween inge s; nup ial pad no isible.
Hindlimbs compa a i ely s ou ; high leng h and ib-
ia leng h equal, less han hal o snou - en leng h (TL/
SVL = 0.43); oo leng h g ea e han high and ibia
leng h (TL/FOL = 0.96); ou h oe longes ; oes wi h
ounded discs; oe discs wid h sligh ly smalle han ha
o inge s; ci cum-ma ginal g oo e p esen on discs; sub-
a icula ube cles dis inc and ounded; supe nume a y
ube cles absen ; inne me a a sal ube cle p esen ; ou e
me a a sal ube cle absen ; webbing mode a e, eaching
i s suba icula ube cle on ou h oe.
Skin on do sal aspec o snou and head smoo h wi h nu-
me ous enla ged, dis inc spinules including on uppe eye-
lids; simila spinules on lo eal egion, below eyes, and on
mandibula egion; skin on do sum smoo h wi h nume ous
spinules, which dec ease in numbe pos e io ly; lank wi h
la g anula ube cles; en um o head and ches smoo h;
abdomen g anula ; g anules on high indis inc ; la ube -
cles below en ; o elimbs, ibia, and a sus smoo h.
Figu e 7. “Rao ches es sahai” (= Nasu ixalus je donii). A–C Holo ype o “R. sahai” (ZSIA8500). A Do sal iew, B en al iew,
C la e al iew o head; D–I pa a ype o “R. sahai” (ZSIA8501). D Do sal iew, E en al iew, F la e al iew o head, G en al
iew o le hand, H en al iew o le oo ; I–J ome ine ee h in pa a ype and holo ype espec i ely. Scale ba = 10 mm.
Ve eb a e Zoology 75, 2025, 517–625 539
Figu e 8. A ML ee in e ed om 16S mi ochond ial gene. UFB alues below 50 a e no shown on ee. Scale ba ep esen s
subs i u ion pe si e. Newly collec ed specimens o N. je donii om ype locali y o “R. sahai” Gandhig am, A unachal P adesh;
B–E opo ypic collec ion WII-ADA1773 in p ese ed condi ion; F WII-ADA1773 in li e; G–H WII-ADA1770 in p ese ed con-
di ion. Scale ba = 10 mm.
Bo uah B e al.: Re ision o bush ogs om no heas India
540
Colou a ion in li e (based on specimen WII-ADA1773;
Fig. 8F). Top and la e al sides o head b own; ip o snou
wi h enla ged oli e-b own pa ch o below nos ils and e -
ical pale yellow line a ip o snou ; a simila ly colou ed
spo in on o in e o bi al space; oli e-b own s ipe on
lo eal egion below can hus os alis; a simila ly colou ed
pa ch below eyes, bo de ed wi h pale yellow; da k b own
i is wi h i egula golden pa ches; oli e-b own s eak
along an e io pa o sup a ympanic old and a ew i eg-
ula spo s on ympanum; do sum pale yellowish b own
wi h enla ged oli e b own hou glass shaped pa ch, s a -
ing om in e o bi al space and co e ing pos e io hal o
uppe eyelids, pos e io ly bi u ca ing in middle o do -
sum, di ec ing owa ds g oin as g adually di using; wo
pale yellowish b own spo s on da k pa ch, one on back o
head and ano he elonga ed one on an e io pa o back;
lank pale b own wi h some i egula small oli e-b own
spo s; do sum o o elimb pale yellowish b own; b oad
oli e-b own band on o ea m; simila pa ch on base o
hands and do sal aspec o inge s; hindlimbs pale yel-
lowish b own on op wi h h ee b oad oli e-b own, i eg-
ula bands on each high and ibia; small pa ch o simila
colou on knees; simila bands on a sus, base o oo , and
on do sal aspec o oes; en um o head, abdomen, and
limbs uni o m pale c eam colou ed.
Colou a ion in p ese a i e. Do sal aspec o head, do -
sum, and limbs b own; ma kings on head, do sum, and
limbs isible as in li e; en um o head, back and limbs
uni o m pale b own.
Mo phological a ia ion. O e all, he newly collec ed
ma e ial om he ype locali y o “R. sahai” simila o
he ypes o N. je donii. Howe e , ollowing a ia ions
we e obse ed among he indi iduals. Head sligh ly wid-
e han leng h in WII-ADA1773 which is simila o ha
o holo ype o “R. sahai” (ZSIA8500). Howe e , in he
pa a ype (ZSIA8501) and wo newly collec ed specimens
(WII-ADA3240 and WII-ADA1770), head leng h is equal
o head wid h. Snou leng h is equal o eye leng h in he
ype specimens o “R. sahai” (ZSIA8500, ZSIA8501), bu
sligh ly sho e in he newly collec ed specimens. Snou
leng h g ea e han eye leng h in he ypes o N. je donii
(BMNH 1947.2.7.84, BMNH 1947.2.7.85). In e -uppe
eyelid wid h g ea e han uppe eyelid wid h in he ypes
o “R. sahai” (ZSIA8500, ZSIA8501) and ypes o N.
je donii (BMNH 1947.2.7.84, BMNH 1947.2.7.85). In-
e -uppe eyelid wid h is equal o he uppe eyelid wid h
in WII-ADA1773 and in wo specimens (WII-ADA3240
and WII-ADA1770), in e -uppe eyelid wid h is small-
e han uppe eyelid wid h. In e na ial dis ance equal
o in e -uppe eyelid wid h in he ypes o “R. sahai”
(ZSIA8500, ZSIA8501) and WII-ADA1773. In e na ial
dis ance equal o in e -uppe eyelid wid h in he ypes o
“R. sahai” (ZSIA8500, ZSIA8501) and WII-ADA1773
while in e na ial dis ance g ea e han in e -uppe eyelid
wid h in WII-ADA3240 and WII-ADA1770. In e na ial
dis ance smalle han in e -uppe eyelid wid h in he ypes
o N. je donii (BMNH 1947.2.7.84, BMNH 1947.2.7.85).
Tibia leng h smalle han high leng h in ypes o “R.
sahai” (ZSIA8500, ZSIA8501) and he wo opo ypes
(WII-ADA3240 and WII-ADA1770) bu high leng h and
ibia leng h equal in WII-ADA1773. Tibia leng h g ea -
e han high leng h in he ypes o N. je donii (BMNH
1947.2.7.84, BMNH 1947.2.7.85). A de ailed mo phome -
ic a ia ion o he newly collec ed ma e ial is p o ided in
Table S11. Do sal ma kings on head, do sum and limbs o
he ypes a e ba ely isible unlike he opo ypes. In addi-
ion, WII-ADA3240 has a dis inc nup ial pad on he i s
inge ; i egula shaped and sized b own pa ches p esen on
en um o head, ches , and o elimbs o WII-ADA1770.
Dis ibu ion and na u al his o y. Nasu ixalus je donii is
widely epo ed om no heas India (Biju e al. 2016). I
was p e iously epo ed om Da jeeling (Gün he 1876);
Mawphlang Sac ed G o e, Wahlynkien (Ma ai Kaphon),
Che apunji in Meghalaya; Za aeng ung, Raenghzaeng
Village in Manipu ; Sechüma illage, Zubza, Me iema
Village, Seukwehii, Tseminyu Village in Nagaland (Biju
e al. 2016); Hmui ang Communi y Rese e Fo es , Mu -
len Na ional Pa k in Mizo am (Muansanga e al. 2022).
Rahman e al. (2020) epo ed his species om Zi adum,
Myanma . Zug (2022) epo ed his species om no h-
e n Myanma (Kachin). Du ing ou s udy, we eco ded
he species om Gandhig am and nea Glaw Lake a
Kamlang Tige Rese e in A unachal P adesh, and in
Hmui ang, Mizo am. A Gandhig am, subadul indi idu-
als we e eco ded on lea es a a heigh o ~20 cm abo e
g ound nea a small s eam ~300 cm wide a a ound 20:00
h s. Adul indi iduals we e eco ded on lea es a a heigh
o ~100 cm abo e g ound a a ound 22:00 h s in Sep em-
be 2022. Calling males we e obse ed ac i e among ee
e ns a h ee me e s abo e g ound in la e May.
Nasu ixalus mic odiscus (Annandale, 1912)
no . comb.
Ch esonymy.
Rhacopho us mic odiscus Annandale, 1912: 114.
Rhacopho us (Rhacopho us) mic odiscus — Ahl (1931): 57, 121.
Philau us (Ki ixalus) mic odiscus — Dubois (1987): 73.
Philau us (Philau us) mic odiscus — Bossuy and Dubois (2001): 40.
Philau us mic odiscus — Ahmed e al. (2009): 16.
Commen s on axonomic s a us. Wha is cu en ly
known unde Philau us mic odiscus (Fig. 9) was o iginal-
ly desc ibed by Annandale (1912), based on a single speci-
men (ZSI16924), as Rhacopho us mic odiscus om Kobo
in he oo hills o Abo (now in Assam; Fig. 1A). This spe-
cies has ne e been epo ed since i s desc ip ion. Go ham
(1974) synonymized his species wi h Rhacopho us je -
donii (now Nasu ixalus je donii) bu his was no ollowed
by Dubois (1987) and Bossuy and Dubois (2001). Go -
ham (1974) was only ollowed by Sa ka and Ray (2006).
Dubois (1987) placed his species in he genus Philau us
unde he subgenus Ki ixalus. La e , Bossuy and Du-
bois (2001) ans e ed i o he nomino ypical subgenus
Philau us. Howe e , Ga g e al. (2021) ques ioned he
placemen o his species in he genus Philau us. Based on
Ve eb a e Zoology 75, 2025, 517–625 541
molecula da a o bush ogs collec ed om ac oss no h-
eas India, we a gue ha he e a e no species o he genus
Philau us (sensu s ic o) p esen in his egion.
A e a e iew o Annandale’s o iginal desc ip ion and
examina ion o he holo ype (ZSI16924) we p o ide he
ollowing cha ac e s o he holo ype: small body size
(28.7 mm); p esence o ome ine ee h; p esence o u-
dimen a y webbing on hand; p esence o discs on digi s.
Annandale (1912) men ioned ha he disc on he hi d
inge is less han hal o he ympanic diame e , bu his
may be because he specimen was in a dehyd a ed s a e as
seen on he ske ch p o ided in he o iginal desc ip ion);
g anula abdomen and highs; sub iangula ma king on
head; conca e s ipes on do sum; c ossba s on do sal
su ace o limbs (Fig. 9); la e al sides o highs eddish
(based on o iginal desc ip ion).
The holo ype being a small bodied og (28.7 mm) wi h
a compa a i ely sho ounded snou and discs on he in-
ge s and oes (as wi h species in Nasu ixalus, Philau us,
Pseudophilau us, and Rao ches es), om no heas India
(as we a gue only wi h Nasu ixalus and Rao ches es) wi h
ome ine ee h (absen in Rao ches es) s ongly sugges s
ha “P. mic odiscus” belongs o he genus Nasu ixalus Ji-
ang e al., 2016. Species o he genus Nasu ixalus (e.g., N.
medogensis) a e known o occu in he Adi (= Abo ) Hills
o A unachal P adesh (Biju e al. 2016; Yang e al. 2018).
Du ing ou ield su eys o e he pas i e yea s we eco d-
ed se e al popula ions o Nasu ixalus in he uppe each-
es o he Adi Hills and h oughou he S a e o A unachal
P adesh. These eco ded specimens pa ially ma ch An-
nandale’s desc ip ion by he ollowing se o cha ac e s:
do sally ounded and la e ally nea ly unca e snou shape,
dis inc can hus os alis, p esence o ome ine ee h, dis-
inc ympanum, p esence o do sal s ipes, p esence o
da k ba on in e -uppe eyelid space, c ossba s on limbs,
g anula belly, discs on inge s, webbing on oo . Thus,
we o mally ans e Philau us mic odiscus o Nasu ixalus
mic odiscus no . comb. bu acknowledge ha u he s ud-
ies in ol ing collec ing specimens om he ype locali y
o nea he ype locali y o N. mic odiscus and including
Figu e 9. Holo ype o Nasu ixalus mic odiscus no . comb. (ZSI16924). A Do sal iew (pho og aphed in 2012) showing conca e
da k s ipes on do sum (wi h black a ow), B do sal iew, C en al iew, D la e al iew o head, E en al iew o head showing
ome ine ee h indica ed wi h black a ow ( aken in 2024), Ske ch o he same specimen om Annandale’s (1912) o iginal desc ip-
ion o “Rhacopho us mic odiscus”. Scale ba = 10 mm.
Bo uah B e al.: Re ision o bush ogs om no heas India
548
based on he newly collec ed ma e ial om he ype lo-
cali y is necessa y.
Desc ip ion o lec o ype. Small sized Rao ches es, SVL
16 mm (Fig. 14E, F); lowe jaw b oken an e io ly; bo h
he o elimbs b oken a elbow; le hind limb b oken a
g oin and ibio a sal a icula ion; igh hindlimb b oken a
knee and oes comple ely damaged; longi udinal incision
on belly. Snou ounded in do sal and en al aspec ; can-
hus os alis indis inc ; ome ine idge o ee h absen ;
ympanum indis inc ; sup a ympanic old dis inc ; in-
ge disc ounded, ci cumma ginal g oo e p esen ; high
leng h sligh g ea e han ibia leng h (TBL/TL = 0.98);
oe disc ounded, wi h ci cumma ginal g oo e; skin on
do sal aspec o head, do sum and limbs smoo h; a ew
indis inc ube cles on uppe eyelids and pos e io pa o
head behind uppe eyelids; on en al aspec , gula egion
and ches smoo h; belly and high g anula ; ibia smoo h
on en al aspec .
Colou a ion in p ese a i e. Do sal aspec o head,
do sum and limbs b own, sligh ly pale owa ds lank
and base o he hindlimb; en al aspec nea ly uni o m
b own bu sligh ly pale han he do sal colou . Excep
sligh ly da k pa ch on g oin and sligh ly da k and single
b oad c oss ba on high and ibia, o he colou pa e n
such as da k ba on in e -uppe eyelids space, s ipes on
do sum; bands on o ea ms, sil e y inge enci cling da k
pa ch on g oin as men ioned in he o iginal desc ip ion
a e no isible now.
Rao ches es ga o (Boulenge , 1919)
Figu es 15–17; Tables 1, 2, S6, S7, S12
Synonymy and ch esonymy.
Ixalus ga o Boulenge , 1919: 207.
Rhacopho us (Philau us) ga o — Ahl (1931): 70.
Philau us ga o — Bou e (1942): 450–451; Inge (1985): 528; Ahmed
e al. (2009): 15.
Philau us (Philau us) ga o — Bossuy and Dubois (2001): 41.
Philau us namdaphaensis Sa ka and Sanyal, 1985: 287–289.
Rao ches es manipu ensis Ma hew & Sen, 2009: 43, 44, pla e XVI.
Philau us namdaphaensis — Ahmed e al. (2009): 16, 147.
Rao ches es cangyuanensis Wu e al., 2019: 558–561.
Rao ches es kempiae — Na een e al. (2024): 362, 365–367.
Rao ches es ga o — Na een e al. (2024): 362.
Commen s on axonomic s a us. Boulenge (1919) de-
sc ibed “Ixalus” ga o based on a single specimen (Fig.
15 A–D) collec ed by S anley Wells Kemp om “abo e
Tu a”, Ga o Hills. I was subsequen ly placed in o he
hacopho id gene a including Rhacopho us Kuhl & an
Hassel , 1822 and Philau us (Ahl 1931; Bou e 1942;
Inge 1985; Bossuy and Dubois 2001). This species has
been epo ed om Assam (Choudhu y e al. 2001), Naga-
land (Ao e al. 2003) and Shillong in Meghalaya (Ma hew
and Sen 2009; Kha kongo e al. 2016). Howe e , i has
ne e been epo ed om i s ype locali y (abo e Tu a in
Ga o Hills, Meghalaya) since i was i s desc ibed. Re-
cen ly, Na een e al. (2024) epo ed his species om
Da ibokg e Communi y Rese e in he Eas Ga o Hills
Dis ic o Meghalaya, claiming ha hei collec ed spec-
imen is a “ opo ype” (see discussion below). Du ing ou
ield su ey on 27 May 2022, we encoun e ed an indi-
idual o he bush og (Fig. 16A–H) a he ype locali y
Tu a Peak (25.50302°N, 90.23853°E, ele a ion 1030 m
a.sl.), Meghalaya. These indi iduals esemble he cha -
ac e s men ioned in he o iginal desc ip ion o R. ga o
by Boulenge (1919) as ollows: 1) da k-b own hou glass
shaped ma k co e ing head om in e o bi al space and
back, 2) lo eal and empo al egion da k-b own, 3) da k-
b own c ossba s on limbs and 4) dis inc ympanum, 5)
can hus os alis dis inc . Al hough he o iginal desc ip-
ion s a ed he snou shape is unca ed, i is damaged in
he holo ype (squeezed, see Fig. 15). Ou examina ion o
he newly collec ed ma e ial ound ha he snou shape is
ounded o sub-o oid.
Ma e ial examined. Holo ype: ZSI 19187, subadul (sex
could no be de e mined); collec ed by S.W. Kemp abo e
Tu a, Wes Ga o Hills Dis ic , Meghalaya, India.
Newly collec ed ma e ial. Topo ypes: i e adul males
(WII-ADA1493–1496, WII-ADA1499) collec ed by
BB, VJ, and AD on 27 May 2022 om Tu a Peak Re-
se ed Fo es (25.50302°N, 90.23853°E, ele a ion 1030
m a.s.l.), Wes Ga o Hills Dis ic , Meghalaya. Re e ed
ma e ials: one adul male (WII-ADA1479) collec ed
by BB, VJ, and AD on 20 May 2022 om Da ibokg e
(25.48973°N, 90.32373°E, ele a ion 1140 m a.s.l.),
Wes Ga o Hills Dis ic , Meghalaya; wo adul males
(WII-ADA858, WII-ADA859) collec ed by BB and AD
on 1 Augus 2021 om nea Kiphi e Di isional Fo es
Complex (25.89940°N, 94.76947°E, ele a ion 1300
m a.s.l.), Kiphi e Dis ic , Nagaland; one adul male
(WII-ADA861) collec ed by BB and AD on 2 Augus
2021 om A amasang am, (25.83128°N, 94.87319°E,
ele a ion 1480 m a.s.l.), Kiphi e Dis ic , Nagaland;
se en adul males (WII-ADA601–WII-ADA605, WII-
ADA607, WII-ADA608) collec ed by BB on 18 Ap il
2019 om Lakhiche a (24.97502°N, 92.77305°E, ele a-
ion 50 m a.sl.), Cacha Dis ic , Assam; wo adul males
(WII-ADA609, WII-ADA610) collec ed by BB on 20
Ap il 2019 om Subhang (24.97613°N, 92.79206°E, el-
e a ion 170 m a.s.l.), Cacha Dis ic , Assam; wo adul
males (WII-ADA1035 and WII-ADA1036) collec ed by
BB, NGP, and AD on 9 Sep embe 2021 om Ngengpui
Wildli e Sanc ua y (22.48696°N, 92.77307°E, ele a ion
180 m a.s.l.), Lawng lai Dis ic , Mizo am; one adul e-
male (WII-ADA1142) collec ed by BB, NGP, and AD on
16 Sep embe 2021 om Tei ei (23.694°N, 92.45147°E,
ele a ion 270 m a.s.l.), Dampa Tige Rese e, Mami Dis-
ic , Mizo am; one adul male (WII-ADA1645) collec -
ed by BB on 25 July 2022 om Leima ak (24.59375°N,
93.66256°E, ele a ion 480 m a.sl.), Chu achandpu
Dis ic , Manipu ; wo adul males (WII-ADA1648 and
WII-ADA1649) collec ed by BB on 26 July 2022 om
Cha oikhullen (24.6071°N, 93.71986°E, ele a ion 1180
Ve eb a e Zoology 75, 2025, 517–625 549
Figu e 15. Rao ches es ga o. A–D Holo ype o Rao ches es ga o (ZSI19187), A do sal iew, B en al iew, C, D la e al iew
o head; E–N “Philau us namdaphaensis”, E–G holo ype (ZSIA 7177); H–J pa a ype (ZSIA 7179); K–N pa a ype (ZSIA 7178),
N g anula belly skin. Scale ba = 10 mm.
Bo uah B e al.: Re ision o bush ogs om no heas India
550
m a.s.l.), Chu achandpu Dis ic , Manipu ; ou adul
males (WII-ADA3211, WII-ADA3213, WII-ADA3217,
and WII-ADA3219) collec ed by RNV and SD on 9 May
2023 om Haldiba i (27.52453°N, 96.39913°E, ele a-
ion 500 m a.s.l.), Namdapha Tige Rese e, Changlang
Dis ic , A unachal P adesh.
Diagnosis. Small o medium sized Rao ches es, SVL
19.9–26.9 mm in adul males; head leng h equal o i s
wid h o sligh ly wide han long (HL/HW = 0.9–1.01);
snou shape ounded o sub-o oid in do sal iew, snou
leng h sligh ly less han o equal o eye leng h (SL/EL =
0.81–1.03); snou leng h g ea e han o equal o in e -up-
pe eyelid wid h (0.96–1.28); nos ils close o snou ip
han eye o equidis an be ween he wo; in e na ial dis-
ance smalle han in e -uppe eyelid wid h (IN/IUE =
0.71–0.94) and g ea e o equal o uppe eyelid wid h
(UE/IN = 0.73–1.0), in e -uppe eyelid wid h smalle
han eye leng h (0.74–0.94); spinules o ube cles on do -
sum; b own pa ches on g oin and high; h ee da k bands
on high and ibia.
Redesc ip ion o holo ype (ZSIC 19187; Fig. 15A–D).
Specimen comple ely dehyd a ed; snou squeezed; lowe
jaw b oken a igh side; ou h inge disc on le side
damaged; igh hind limb is dislodged om body; a punc-
u e on an e io pa o abdomen. Small sized og (SVL
= 12.3 mm); head leng h equal o wid h; eyes mode a e,
less han hal o head leng h (EL/HL = 0.38); in e o -
bi al space equal o eye leng h and g ea e han uppe
eyelid wid h (UEW/IUE = 0.43); ympanum dis inc ,
ound; ongue pos e io ly no ched; sup a ympanic old
isible; o elimbs slende , o ea m leng h smalle han
hand leng h (FAL/HAL = 0.81); each inge wi h ounded
discs; ci cum-ma ginal g oo e p esen on discs; suba ic-
ula ube cles dis inc , ounded; palma ube cles indis-
inc ; hindlimbs slende , high leng h mo e han hal o
snou - en leng h (TL/SVL = 0.57) and sligh ly longe
han ibia (TBL/TL = 0.97); each oes wi h ounded disc;
discs wi h ci cum-ma ginal g oo e, as wide as hose on
inge s; suba icula ube cles dis inc , ounded; inne
me a a sal ube cle p esen and ou e me a a sal ube cle
absen ; do sal aspec o head and do sum smoo h; do sal
su ace o limbs smoo h; lank smoo h; h oa , ches and
en al side o limbs smoo h; abdomen g anula .
Colou a ion o holo ype (ZSIC 19187) in p ese a-
i e. Head do sally b own and pale la e ally; do sum
b own and pale owa ds he lank; enla ged, sligh ly
da ke hou glass shaped ma k isible on do sum s a -
ing a in e o bi al space, pos e io ends much da ke nea
g oin; limbs b own do sally; pale b own en ally; b own
specks on h oa , ches , and an e io pa o abdomen and
lowe a ms.
Expanded desc ip ion based on he newly collec ed o-
po ype (WII-ADA1495: Fig. 16A–H). A medium sized
Rao ches es, SVL 25.3 mm; head wide han long (HL/
HW = 0.93); snou ounded in do sal iew, sligh ly p o-
uding beyond lowe jaw in en al iew; snou leng h
sligh ly smalle han eye leng h (SL/EL = 0.92) and equal
o in e -uppe eyelid wid h; a sligh dep ession on in e -
na ial space; nos ils o al, obliqued, close o snou ip
han eyes (NS/EN = 0.88); na ial egion p o uding; can-
hus os alis dis inc , obliqued; lo eal egion conca e;
eyes p o uding, mode a e in size (EL/HL = 0.38), g ea e
han in e -uppe eyelid wid h (IUE/EL = 0.89); in e na ial
dis ance equals o uppe eyelid wid h; ympanum ound,
dis inc , nea ly one ou h o eye leng h; sup a ympanic
old dis inc ; ome ine ee h absen ; choanae ound; unk
less han hal snou - en leng h (TRL/SVL = 0.48).
Fo elimbs slende ; o ea m sho e han hand leng h
(FAL/HAL = 0.86); hi d inge longes ; inge s wi h
ounded discs; ci cum-ma ginal g oo e p esen on discs;
disc on inge II, III and IV g ea e han ympanic diam-
e e ; palma ube cles indis inc ; suba icula ube cles
dis inc , ound; p oximal suba icula ube cle on inge
III and IV smalle han dis al suba icula ube cles; ine
g anula nup ial pad on i s inge ; webbing absen .
Hindlimbs slende ; high leng h hal o snou - en
leng h (TL/SVL = 0.52) and sligh ly g ea e han ibia
leng h (TBL/TL = 0.98); ou h oe longes ; ci cum-ma -
ginal g oo e p esen on oe discs; suba icula ube cles
dis inc , ounded; p oximal suba icula ube cle on ou h
oe indis inc and small; supe nume a y ube cles absen ;
oe disc wid h equal o ha o inge s; inne me a a sal
ube cle p esen , i s leng h equal o disc wid h o second
oe; ou e me a a sal ube cle absen ; webbing small,
eaching second suba icula ube cle on ou h oe.
Skin on do sal aspec o snou and head smoo h wi h
indis inc ube cles on uppe eyelids; side o head smoo h,
wi h ew ube cles behind angle o jaw; spinules s a ing
behind uppe eyelids and abo e sup a ympanic olds uns
along do sola e al side o unk and sca e on middle o
unk, pos e io pa o do sum smoo h; lank g anula ;
o elimbs and hindlimbs smoo h do sally; head on en-
al side smoo h; ches , lowe a ms, abdomen and highs
g anula ; ibia smoo h.
Colou a ion in li e. Do sal aspec o head yellowish
b own om snou o an e io one hi d o uppe eyelids
wi h i egula b own spo s; do sum yellowish b own wi h
da k b own spo s; an hou glass shaped b own ma k s a -
ing om in e o bi al space co e ing pos e io wo hi d
o uppe eyelids o an e io o en , pos e io ly di used;
da k b own spo s along he edges o hou glass shaped
ma k; lo eal egion and a ea an e io o nasals b own,
a ea below eyes, lips, ympanic egion and mandibula
egion pale b own wi h sligh ly da k b own specks and
spo s; indis inc b own s eak below sup a ympanic old
s a ing om pos e io co ne o eyes o ea o mandi-
ble; lowe a m o o elimbs yellowish b own do sally
and o ea m g eyish b own do sally; a b own c ossba on
o ea ms; hands g eyish b own wi h b own specks; disc
o he i s and second inge s pale yellow; highs and
ibia yellowish b own do sally; g eyish b own wi h ain
eddish inge owa ds ou e la e al side o ibia and a sus;
indis inc c ossba s on ibia; c ossba s on highs in he
o m o in e connec ed spo s; la ge da k b own pa ch on
ou e la e al side o highs and inne la e al side o ibia;
Ve eb a e Zoology 75, 2025, 517–625 551
i egula shaped da k b own enla ged spo s on g oin and
a sligh ly pale elonga ed pa ch en al o i ; ain b own
pa ch on inne la e al side o highs; a ew da k b own
spo s abo e en ; h oa , ches , abdomen, en al aspec
o o elimbs, and base o highs lesh colou ed; b own
mo ling and c eamy-whi e spo s on an e io pa o low-
Figu e 16. Rao ches es ga o. A–H WII-ADA1495 om he ype locali y, Tu a Peak. A Do sal iew (hou glass shaped da k pa ch
on do sum is shown wi h a black a ow), B en al iew, C la e al iew o head in p ese ed condi ion, D do sal iew, E do sola -
e al iew, F en al iew, G g oin, H highs. I–M WII-ADA608 om Lakhiche a, Assam. I Do sal iew in p ese ed condi ion
(Inse image showing spinules on do sum), J do sola e al iew, K en al iew, L g oin, M. high; N–Q WII-ADA1499 om he
ype locali y, N do sola e al iew, O do sal iew, P g oin, Q high. R–T WII-ADA1142 om Te ei, Mizo am. R Do sola e al iew,
S g oin, T high; U WII-ADA3217 om Haldiba i, A unachal P adesh; V, W WII-ADA607 and WII-ADA603 espec i ely om
Lakhiche a, Assam; X. WII-ADA1035 om Ngengpui, Mizo am. Scale ba = 10 mm.
Bo uah B e al.: Re ision o bush ogs om no heas India
552
e jaw and i egula ly shaped small b own pa ches along
i s edge; gula egion sligh ly pale yellow; b own specks
on an e io pa o abdomen, en al aspec o o elimbs,
and palm, hea ily speckled on o ea ms; en al aspec
o oo , ibia, and o dis al end o highs hea ily speckled
wi h b own.
Colou a ion in p ese a i e. Do sal ma kings on head,
do sum and lank same as when ali e; do sum o snou
pale c eam colou ed up o in e o bi al space including
an e io pa o uppe eyelids, a b own pa ch in he mid-
dle; side o head da k b own; do sum owa ds lank pale-
c eam colou ed wi h da k b own spo s; he da k b own
hou glass shaped ma k co e ing head, and do sum isible
as in li e; lowe a ms c eam colou ed and o ea ms da k
b own; highs and ibia c eam colou ed; ou e do sola e -
al side o ibia da k b own; a sus and oo da k b own;
dis inc da k b own c ossba s on high and ibia; da k
b own ma bling on en al side mo e dis inc han in li e,
dense ma bling on chin, along lowe jaw, o ea ms, ches ,
ibia, a sus and oo .
Sexual dimo phism and mo phological a ia ion.
Adul males ha e ex e nal ocal sac, in e nal ocal sli s
on lowe jaw, and nup ial pad on i s inge . De ails o
a ia ion in mo phome ic cha ac e s among indi iduals
a e gi en in Table S12. In addi ion, ma king pa e ns on
head, do sum, g oin and high a y among indi iduals
o he species (Figs 16, 17). Do sal colou a ies as pale
yellowish b own, pale eddish b own and g eyish b own.
Hou glass shaped ma king da k g ey o da k b own (WII-
ADA1495). This ma king is b oken in some indi iduals
(WII-ADA607; Figs 16V, 17B) o may be eplaced by a
pai o b oad conca e s ipes (WII-ADA1035), o com-
ple ely absen (WII-ADA603; Figs 16W, 17F). Some in-
di iduals ha e a whi e ba on o ehead co e ing he an-
e io pa o he uppe eyelids (WII-ADA608; Figs 16I,
J, 17A); g oin and highs may ha e i egula da k-b own
pa ches o da k-b own pa ch wi h b igh i egula sized
and shaped yellow spo s; pale yellow mid-do sal line
may be p esen and a simila line on hindlimbs o igina -
ing om he mid-do sal line abo e en and unning do -
sally along highs and ibia and en o-la e ally on a sus
(WII-ADA1499; Fig. 16N, O, U); da k b own spo s may
be p esen o absen on lank and limbs; one indi idual
om Ngengpui Wildli e Sanc ua y ha e pa ches o yel-
low lecks on head, back, hind limbs and lowe a ms we e
pale-yellow on op (WII-ADA1035; Figs 16X, 17G); in-
ensi y on he spinules and ube cles on head, back and
limbs a ies among indi iduals as dense o sca e ed.
Mo phological compa ison. Rao ches es ga o di e s
om R. annandalii, R. dulongensis, R. hekouensis, R. hil-
lisi, R. huanglianshan, R. jadoh, R. leik ho, R. malipoen-
sis, R. minda and R. pa ulus by la ge body size, SVL
19.9–26.9 mm in adul males ( s. SVL in adul males,
17.3–19.6 mm in R. annandalii, 15.0–19.0 mm in R. du-
longensis, 16.1–17.5 mm in R. hekouensis, 15.9–17.7
mm in R. hillisi, 17.0–19.6 mm in R. huanglianshan,
13.6–14.0 mm in R. jadoh, 15.7–15.8 mm in R. leik-
ho,14.6–17.7 mm in R. malipoensis, 16.7–18.3 mm in
R. minda , 17.0–18.6 mm in R. pa ulus); i di e s om
R. annandalii, R. cine ascens no . comb., R. pa ulus,
and R. shillongensis by he absence o conca e s ipes on
do sum ( s. p esen ); i di e s om R. ande soni by he
p esence o spinules on do sum ( s. sca e ed ube cles
on do sum) and h ee da k bands on high and ibia ( s.
single band on high and ibia); i di e s om R. kempiae
by dis inc can hus os alis ( s. indis inc can hus os a-
lis), enla ged da k b own pa ches wi h o wi hou ligh
pa ches p esen on g oin and la e al aspec o high ( s. a
small b own s eak p esen on g oin and no da k pa ches
on la e al aspec o high), a whi e ba on in e -uppe eye-
lid space may be p esen o absen ( s. a dis inc o indis-
inc da k ba may be p esen o absen ), hou glass pa e n
may be p esen co e ing head and do sum o absen ( s.
hou glass pa e n absen , only “)(“ ma k may be p esen
on do sum); i di e s om R. hillisi, R. huanglianshan
and R. menglaensis by i s snou leng h being smalle han
o equal o eye leng h ( s. snou leng h longe han eye
leng h); di e s om R. jakoid by p esence o enla ged
da k b own pa ches wi h o wi hou ligh pa ches on g oin
and la e al aspec o high ( s. absen ); i di e s om R.
longchuanensis by in e -uppe eyelid wid h smalle han
eye leng h ( s. in e -uppe eyelid wid h g ea e han eye
leng h); i di e s om R. malipoensis by in e -uppe eye-
lid wid h smalle han eye leng h ( s. in e -uppe eyelid
wid h g ea e han eye leng h); i di e s om R. meng-
laensis by i s snou leng h being smalle han o equal o
eye leng h ( s. snou leng h longe han eye leng h); i
om R. ezakhani by p esence o spinules on do sum ( s.
sca e ed ube cles on do sum), conca e s ipes on do -
sum absen ( s. p esen ), enla ged b own pa ches wi hou
o wi h yellow spo s on g oin p esen ( s. sho b own
s eak on g oin); i di e s om R. y hus no . comb. by
i s snou leng h being smalle han o equal o eye leng h
( s. snou leng h g ea e han eye leng h), in e -uppe eye-
lid wid h smalle han eye leng h ( s. in e -uppe eyelid
wid h g ea e han eye leng h), in e -uppe eyelid wid h
smalle han o equal o snou leng h ( s. in e -uppe
eyelid wid h g ea e han snou leng h); i di e s om R.
yadongensis by snou leng h being g ea e han o equal
o in e -uppe eyelid wid h ( s. in e -uppe eyelid wid h
g ea e han eye leng h), in e -uppe eyelid wid h smalle
han eye leng h ( s. in e -uppe eyelid wid h g ea e han
eye leng h). De ailed mo phological compa ison wi h
o he congene s included in his s udy is gi en in Table 1.
Acous ics. The calls o R. ga o we e eco ded a Tu a
Peak, Meghalaya on 27 May 2022 a an ambien em-
pe a u e o 28.2°C and a Lakhiche a, Assam on 18
Ap il 2019 a an ambien empe a u e o 28.8°C be ween
18:30 h s and 19:30 h s. The call desc ip ion is based on
42 calls om wo indi iduals (WII-ADA1495 and WII-
ADA601). The calls a e single ype, non-pulsa ile (Fig.
18), ini ially emi ed as a single call wi hou a g oup, and
g adually emi ed in g oups (4.72 ± 2.22 calls pe call
g oup; 2–13 calls/call g oup). The mean call du a ion is
28.57 ± 2.74 ms (20–32 ms) wi h a call ise ime o 1 ms
and call all ime o 27.55 ± 2.71 ms (19–31 ms). The
Ve eb a e Zoology 75, 2025, 517–625 553
in e al be ween calls is 154.31 ± 7.02 ms (144–170 ms)
when emi ed in g oups. The mean dominan equency
o he call is 2860.84 ± 49.55 Hz (2799.3–2971.6 Hz). A
de ailed compa ison o he ad e isemen calls wi h hose
o o he congene s is p esen ed in Table 2.
Figu e 17. Di e en mo phs o Rao ches es ga o in hei na u al habi a . A–F Ba ail hills, Assam; G Ngengpui Wildli e Sanc ua y,
Mizo am; H Leima ak, Manipu ( opo ype o R. “manipu ensis” WII-ADA1645).
Bo uah B e al.: Re ision o bush ogs om no heas India
554
Phylogene ic ela ionship and gene ic di e gence.
Rao ches es ga o is sis e o R. leik ho (UFB 99 and PP 1;
Figs 2, 3). The gene ic di e gence o R. ga o wi h o he
congene s included in his s udy a e 4.2–9.2% in he 16S,
13.7–21.2% in he cy b and 9.5–16.4% in he COI gene
(Table S7A–C).
Dis ibu ion and na u al his o y. This species is dis ib-
u ed in no heas India om wes e n pa o Meghalaya
o he eas e n pa o A unachal P adesh in he Namda-
pha Tige Rese e. This species is also dis ibu ed in he
Ba ail Hills o Assam and Nagaland, in Manipu , and in
Mizo am. I is mos ly associa ed wi h lowland e e g een
o es s (Fig. 19A). This species was so a known om
50–1480 m a.s.l. ac oss hills on he sou he n slopes o
B ahmapu a Valley, India. Al-Razi e al. (2020a) epo ed
his species as R. longchuanensis om Sa cha i Na ional
Pa k in Bangladesh. Wu e al. (2019) epo ed his species
as R. cangyuanensis om Cangyuan, China. Lal onunga
e al. (2021) epo ed om Mizo am as R. cangyuanensis.
Choudhu y e al. (2001) epo ed his species om Ga -
bhanga Rese e Fo es and Kulsi Rese e Fo es in Assam,
and Ao e al. (2003) epo ed i om Dzulake, Nagaland
which needs u he examina ion o con i m he eco ds.
Calling males we e eco ded du ing Ap il–June om
pe ches 1–2 me es abo e he g ound and om oadside
o es a eas domina ed by bamboo hicke s (Fig. 20A, B).
We also eco ded he species calling om he lea es o a
small ee abou i e me es abo e he g ound and om a
bamboo hicke a a heigh o six me es abo e he g ound
in Cha oikhullen, Manipu . We obse ed a single emale
indi idual o his species on oadside ege a ion in Tei ei,
Dampa Tige Rese e, Mizo am in Sep embe 2021.
Rema ks. Na een e al. (2024) epo ed edisco e y o
“R. ga o” based on i e specimens (SACON VA 809, 163,
305, 306, 308) collec ed om an unnamed locali y wi h
geog aphical coo dina e gi en as “25.51°N, 90.38°E,
895 m” in Ga o Hills, Meghalaya and one specimen
(SACON VA 129) om Da ibokg e Communi y Rese e
(25.47297°N, 90.3148°E, 1200 m), ha hey claimed as
a “ opo ype”. Howe e , he 16S gene sequence da a in
Na een e al. (2024) comes om a specimen (SACON
VA 809) collec ed app oxima ely 15 km eas o Tu a.
Mo phological desc ip ion o he alleged “ opo ype”
(SACON VA 809) do no ma ch he o iginal desc ip ion
in e ms o do sal colou pa e n. The o iginal desc ip ion
o R. ga o men ions an hou glass pa e n on do sum, bu
his was absen in SACON VA 809. Al hough Na een e
al. (2024) men ioned ha wo o he specimens such as
VA 305 and VA 308 ha e hou glass pa e ns, hey did no
gene a e molecula da a o hese wo specimens. The ho-
lo ype (ZSIC19187) o R. ga o was o iginally collec ed
by S.W. Kemp om “abo e Tu a” which is he idge/peak
immedia ely abo e Tu a (see Pai a 1919). We he e o e
a gue ha wi hou jus i ica ion, Na een e al. (2024) con-
side ed SACON VA 809 as a opo ype o R. ga o, and
p o ided a misleading phylogene ic s a us o R. ga o.
The DNA sequence (16S) o he “R. ga o” opo ypical
specimen (SACON VA 809) submi ed by Na een e al.
(2024) did no clus e wi h ou opo ypical samples o R.
ga o (WII-ADA1493, WII-ADA1494, WII-ADA1499)
collec ed om Tu a Peak (= abo e Tu a) (25.50302°N;
90.23853°E; ele a ion 1030 m a.s.l.; Figs 2, 3). The
gene ic di e gence o he ue opo ypical samples o
R. ga o and he specimen o “R. ga o” o Na een e al.
(2024) is 6.5–7.0% in 16S gene (Table S7A–C). Na een
Figu e 18. Ad e isemen calls o R. ga o (WII-ADA1495) on le and R. kempiae (WII-ADA1497) on igh . F om op o bo om,
A, B oscillog ams, C spec og am, D powe spec um. The call ma ked wi h an a ow on op in A is expanded in B.
Ve eb a e Zoology 75, 2025, 517–625 555
e al. (2024) s a ed ha R. ga o is phylogene ically allied
o R. shillongensis. Howe e , ou phylogene ic analyses
indica e R. shillongensis ( opo ype; WII-ADA1460) as
dis inc om R. ga o ( opo ype; WII-ADA1493, WII-
ADA1494, WII-ADA1499; Figs 2, 3). Ou phylogene ic
analyses a e s ongly suppo ed by acous ic cha ac e is-
Figu e 19. Dis ibu ion o Rao ches es species in no heas India. A Rao ches es ga o and h ee o he s closely ela ed new species;
B Rao ches es kempiae, R. shillongensis, R. minda and h ee new species. Type locali ies a e deno ed wi h s a s.
Bo uah B e al.: Re ision o bush ogs om no heas India
556
Figu e 20. Habi a a he ype locali ies o Rao ches es species o no heas India. A Fo es ail a Tu a Peak, he ype locali y o
R. ga o and R. kempiae; B o es co e and adjacen lowlands o Ga o Hills, om Da ibokg e; C a landscape iew o o es co e s
and D a o es ail a Namdapha TR, ype locali ies o R. o ien alis sp. no . and R. nasu a sp. no .; E hills a Mawsyn am, nea he
ype locali y o R. mawsyn amensis sp. no .; F Nohkalikai Wa e all and o es co e a he ype locali y o R. boulenge i sp. no .
in Che apunji; G d y s eam bed and ege a ion in Ma uwache a, nea Ba ail WLS, he ype locali y o R. ba akensis sp. no . and
H o es a ound Jain ia Hills in Na puh WLS, a he ype locali y o R. na puhensis sp. no .
Ve eb a e Zoology 75, 2025, 517–625 557
ics o he wo species (see unde espec i e desc ip ion
and compa ison). This sugges s ha SACON VA 809
epo ed by Na een e al. (2024) is a misiden i ica ion
o R. ga o and i ep esen s an undesc ibed lineage o
Rao ches es. Thus, we conside SACON VA 809 as an
in alid opo ype o R. ga o based on accu a e ype locali-
y and he phylogene ic esul s o his s udy.
“Philau us namdaphaensis” was o iginally desc ibed
by Sa ka and Sanyal (1985) om Fa mbase Camp (a 350
m a.s.l.), Namdapha Na ional Pa k, A unachal P adesh
based on h ee adul males collec ed by S. Biswas. La e ,
Sengup a e al. (2000) epo ed his species om Ga bhan-
ga Rese e Fo es , Assam, and Ma hew and Sen (2009),
and Sen e al. (2013) epo ed i om Nok ek Biosphe e
Rese e, Meghalaya. On 9 May 2023 we collec ed wo
specimens o bush ogs om Haldiba i (27.52453°N;
96.39913°E; ele a ion 500 m a.s.l.), Namdapha Tige Re-
se e, A unachal P adesh (Fig. 15U) on he no h bank
o he Noa-Dihing Ri e . These wo specimens con o m
o he o iginal desc ip ion o “Philau us namdaphaensis”
based on he ollowing cha ac e s: 1) mid do sal line om
snou o en , 2) limbs wi h c ossba s and a line o igi-
na ing a en eaching o heel h ough do sal aspec o
high and ibia, 3) whi e ba on in e o bi al space, 4) da k
b own blo ches on g oin and la e al aspec s o highs.
“Rao ches es manipu ensis” was desc ibed by Ma hew
and Sen (2009) om Leima ak, Manipu based on a sin-
gle specimen. So a , his species is only known om
he ype locali y. On 25 July 2022 we collec ed a opo-
ypical specimen (WII-ADA1645; Fig. 17H) om Lei-
ma ak (24.59375°N; 93.66256°E; ele a ion 480 m a.sl.)
ha con o ms o he o iginal desc ip ion. This indi idual
ma ches he o iginal desc ip ion by he ollowing se o
cha ac e s: 1) head wide han leng h, 2) dis inc can hus
os alis, 3) ome ine ee h absen , 4) a ain hou glass
shaped pa ch co e ing head om in e -uppe eyelid space
and back (da k in holo ype), pos e io ends o his pa ch
di ec ing owa ds g oin; 5) b own and pale whi e pa ches
on lank nea g oin.
Ou phylogene ic analyses in ol ing opo ypical sam-
ples o “P. namdaphaensis” and “R. manipu ensis” e-
sul ed wi h hem nes ed wi h he opo ypical samples o
R. ga o (Figs 2, 3). The opo ypical samples o R. ga o
ha e gene ic di e gences o 0.0–0.6% wi h opo ypi-
cal samples o “P. namdaphaensis” and 0.4–1.1% wi h
he opo ypical sample o “R. manipu ensis” in he 16S
gene (Table S7A–C). Mo phological examina ions o
he opo ypes o he h ee species (R. ga o, “P. namda-
phaensis” and “R. manipu ensis”) and examina ions o
he ype se ies o “P. namdaphaensis” did no e eal any
s ong di e en ia ing cha ac e s o sepa a e hese h ee
species.
Mo phological and molecula s udies o he newly col-
lec ed ma e ial om di e en locali ies in no heas India
sugges s R. ga o is a polymo phic species (see unde e-
desc ip ion sec ion abo e). We ha e eco ded indi iduals
o R. ga o om Tu a peak ha esemble he ypical mo -
pho ype o “P. namdaphaensis” (Fig. 16A–H). Ma hew
and Sen (2008) also epo ed “P. namdaphaensis” om
Nok ek Biosphe e Rese e, Meghalaya p obably based
on his cha ac e . The colou and pa e n o “R. mani-
pu ensis” p o ided in he o iginal desc ip ion and he
pho og aph o he holo ype by Ma hew and Sen (2009)
esemble he ypical mo pho ype o R. ga o om he
ype locali y. The e o e, hese a e un eliable and canno
be used as diagnos ic cha ac e s. Based on molecula
da a and mo phological cha ac e s, we p opose o ea
“Philau us namdaphaensis” and “R. manipu ensis” as ju-
nio subjec i e synonyms o R. ga o.
“Rao ches es cangyuanensis” was desc ibed om
Cangyuan, China (Wu e al. 2019). Subsequen ly, Lal o-
nunga e al. (2021) epo ed his species om Hmui ang,
Mizo am and e e ed o a p e ious eco d o R. longch-
uanensis om Sa cha i Na ional Pa k, Bangladesh (Al-
Razi e al. 2020a) as “R. cangyuanensis”. Rahman e al.
(2022) epo ed he species om no heas Bangladesh.
Based on he esul s o hei phylogene ic s udy,
Na een e al. (2024) sugges ed “R. cangyuanensis” as a
junio subjec i e synonym o R. kempiae. Howe e , in
ou cu en phylogene ic analyses, “R. cangyuanensis” is
nes ed wi h he opo ypical samples o R. ga o (Fig. 3)
collec ed in his s udy (see commen s unde R. kempiae).
The gene ic di e gence be ween R. ga o collec ed o his
s udy and “R. cangyuanensis” samples is 0.0–0.9% in he
16S gene (Table S7A–C). Thus, we conside “R. cangyu-
anensis” as a junio subjec i e synonym o R. ga o and
s a e ha he p e ious eco ds o “R. cangyuanensis”
om Mizo am should be conside ed as R. ga o.
Rao ches es kempiae (Boulenge , 1919)
Figu es 21, 22; Tables 1, 2, S6, S12
Synonymy and ch esonymy.
Ixalus kempiae Boulenge , 1919: 208.
Rhacopho us (Philau us) kempiae — Ahl (1931): 53, 69.
Philau us kempiae — Bou e (1942): 450–451; Inge (1985): 91;
Ahmed e al. 2009: 16.
Philau us (Philau us) kempiae — Bossuy and Dubois (2001): 41.
Rao ches es asakg ensis Na een e al., 2024: 362, 365–370.
Rao ches es kempiae — Na een e al. (2024): 365–367.
Commen s on axonomic s a us. Boulenge (1919) de-
sc ibed Rao ches es kempiae based on a single specimen
collec ed by A. Kemp om “abo e Tu a”, Ga o Hills.
The ype locali y e e s o he idge/peak immedia e-
ly abo e Tu a (see Pai a 1919). I is possible ha his
specimen was collec ed in 1917, which is e iden om
Pai a (1919) and B une i (1918) ha A. Kemp was pa
o hei expedi ion. Howe e , he e was no alloca ed
ouche numbe a ailable o he ype in he o iginal de-
sc ip ion. Chanda (1994) men ioned a possible loca ion
o he ype specimen as he “B i ish Museum, (Na u al
His o y), London, U. K.” which was ollowed by Bossuy
and Dubois (2001) and epea ed in F os (2025). Chan-
da e al. (2000) did no p o ide he p esence o he ype
specimen o “Ixalus kempiae” o “Philau us kempiae”
in he collec ion o he Na ional Zoological Collec ion
o Zoological Su ey o India (ZSI), Kolka a (Calcu a).
Bo uah B e al.: Re ision o bush ogs om no heas India
564
Sexual dimo phism and mo phological a ia ion.
Males ha e a pai o in e nal ocal sac openings, subgu-
la ex e nal ocal sac and nup ial pad on he i s inge ;
head longe han wide in emales, bu in males head wide
han long o wid h equal o leng h. Do sal colou a ies
om pale g eyish b own o pale yellowish b own; con-
ca e s ipes on do sum may be b oken o ain in some
indi iduals; numbe o c ossba s on high and ibia a -
Figu e 23. Rao ches es o ien alis sp. no . A Do sal iew, B en al iew o holo ype (WII-ADA1309) in p ese ed condi ion;
C do sal iew o WII-ADA1389 in p ese ed condi ion; D do sola e al iew and E en al iew o holo ype in li e (inse image
showing g anula skin o abdomen); F WII-ADA3423 in li e; G-J WII-ADA1387 in li e, G do sola e al iew (inse image showing
enla ged blun ly conical ube cles on uppe eyelids), H en al iew, I g oin, J pos e io iew o highs; K an uncollec ed calling
male om Kamala Valley, Namdapha TR; L WII-ADA1631 in li e. Scale ba = 10 mm.
Ve eb a e Zoology 75, 2025, 517–625 565
ies be ween 2–4; axilla and g oin may be pale yellow in
some indi iduals; disc on inne wo inge s may be pale
yellow. De ailed mo phome ic a ia ion be ween he in-
di iduals o his species is gi en in Table S12.
Mo phological compa ison. Rao ches es o ien alis sp.
no . di e s om R. ande soni by sho s ipe p esen on
g oin ( s. la ge black spo wi h wo yellow spo s p esen
on g oin); i di e s om R. annandalii by skin on do -
sum wi h spinules ( s. skin on do sum smoo h); i di e s
om R. cine ascens no . comb. by sho s ipe p esen
on g oin ( s. a spo pa ially enci cled by yellow inge
p esen on g oin), dense spinules on do sum ( s. sca e ed
ube cles on do sum); i di e s om R. dulongensis by
p esence o nup ial pad on i s inge in males ( s. nup ial
pad absen ); i di e s om R. hekouensis by nup ial pad
p esen only on i s inge ( s. nup ial pad p esen on i s
and second inge ); i di e s om R. ga o by sho s ipe
p esen on g oin ( s. enla ged da k b own pa ches wi h
o wi hou whi e o yellow pa ches p esen on g oin); i
di e s om R. hillisi, R. huanglianshan, R. menglaensis,
and R. y hus no . comb. by snou leng h being less han
Figu e 24. Compa a i e spec og ams and oscillog ams illus a ing ep esen a i e calls o ou species o Rao ches es desc ibed
in his s udy. A, B Oscillog ams, C spec og am, D powe spec um. The call ma ked wi h an a ow on op in A is expanded in B.
Bo uah B e al.: Re ision o bush ogs om no heas India
566
o equal o eye leng h ( s. snou longe han eye leng h);
i di e s om R. jadoh and R. jakoid by snou leng h
being less han o equal o eye leng h ( s. snou leng h
g ea e han eye leng h), and high leng h being g ea e
han o equal o ibia leng h ( s. high leng h less han ib-
ia leng h); i di e s om R. kempiae by p esence o dis-
Figu e 25. Dis ibu ion o Rao ches es species in no heas India. A Rao ches es ezakhani and ou o he closely ela ed species;
B Rao ches es annandalii and h ee new species. Type locali ies a e deno ed by s a s, and o he locali ies a e deno ed by ci cles.
Ve eb a e Zoology 75, 2025, 517–625 567
inc “)-(“ on do sum ( s. compa a i ely ain “)(“ma k
on do sum), 2–4 c oss ba s on ibia and high ( s. h ee
ba s on ibia and high); i di e s om R. leik ho by p es-
ence o a pai o conca e s ipes on do sum ( s. pai o
an e io ly connec ed s ipes on do sum); i di e s om
R. longchuanensis by in e -uppe eyelid wid h being less
han o equal o eye leng h ( s. in e -uppe eyelid wid h
g ea e han eye leng h); i di e s om R. malipoensis by
sho s ipe p esen on g oin ( s. black pa ches p esen on
g oin); i di e s om R. minda by sho s ipe p esen on
g oin ( s. enla ged black and whi e pa ches p esen on
g oin); i di e s om R. pa ulus by sho s ipe p esen
on he g oin ( s. da k b own ma bling enclosing a whi -
ish blo ch p esen on g oin); i di e s om R. ezakhani
by p esence o dense spinules on do sum ( s. sca e ed
ube cles on do sum); i di e s om R. shillongensis by
sho s ipe p esen on g oin ( s. da k b own pa ch wi h
whi e c own shaped ma k p esen on g oin); i di e s
om R. yadongensis in e -uppe eyelid wid h being less
han o equal o eye leng h ( s. in e -uppe eyelid wid h
g ea e han eye leng h). Mo phological compa isons
wi h o he new species desc ibed in his s udy a e gi en
in espec i e sec ions and Table 1.
Acous ics. The calls o R. o ien alis sp. no . we e eco d-
ed in Deban, Namdapha Tige Rese e on 10 May, 2022
and in Kamala Valley, Namdapha Tige Rese e on 16
May 2022 a 19:25 h s and 19:30 h s a ambien empe a-
u es o 25.8°C and 22.7°C espec i ely. The desc ip ion
o calls is based on 60 calls including hose o he wo
indi iduals (WII-ADA1309 and WII-ADA1367). The
calls a e single ype, non-pulsa ile, emi ed as a single call
a s a , g adually emi ing calls in g oups (2–6 calls pe
g oup) (Fig. 24). The mean call du a ion is 24.6 ± 2.79 ms
(15–28 ms) wi h a ise ime o 1.13 ± 0.39 ms (1–3 ms)
and a all ime o 23.43 ± 2.85 ms (13–27 ms). The mean
in e al be ween calls is 220.92 ± 23.13 ms (162–295 ms)
when calls emi ed in g oups. The dominan equency
is 3958.52 ± 202.2 Hz (3617.6–4177.4 Hz). A de ailed
compa ison o he ad e isemen calls wi h hose o he
congene s is p esen ed in Table 2.
Phylogene ic ela ionship and gene ic di e gence.
Rao ches es o ien alis sp. no . is sis e o wo o he new
species wi h s ong node suppo (UFB 100, PP 1) (Figs
2, 3). The gene ic di e gence wi h i s congene s is 3.2–
9.1% in he 16S, 9.3–18.7% in he cy b and 6.0–14.9%
in he COI genes (Table S7A–C).
E ymology. The speci ic epi he is de i ed om he wo d
“o ien al” meaning eas e n, in e e ence o i s dis ibu ion
in he eas e nmos pa o India.
Sugges ed common name. Eas e n bush og.
Dis ibu ion and na u al his o y. Rao ches es o ien a-
lis sp. no . is cu en ly known om Namdapha Tige Re-
se e, Kamlang Tige Rese e and Gandhig am Village in
A unachal P adesh, and Liyai Khunou in Manipu (Fig.
25A). Calling males we e eco ded among unde s o ey o
e e g een o es a app oxima ely one me e heigh abo e
g ound (Fig. 20C, D). In Gandhig am his species was
eco ded among sh ubs nea small s eams in he o es s
and nea human se lemen s.
Rao ches es mawsyn amensis sp. no .
h ps://zoobank.o g/D770C5B1-65BD-4BAC-8BCF-9653DF-
C6F9C5
Figu e 26; Tables 1, 2, S12
Holo ype. An adul male (WII-ADA1531) collec ed
by BB on 1 June 2022 nea Maw apa (25.26144°N,
91.53468°E, ele a ion 1140 m a.s.l.), Eas Khasi Hills,
Meghalaya, India.
Pa a ypes. Six adul males (WII-ADA1533–WII-
ADA1538), and an adul emale (WII-ADA1530). Col-
lec ion de ails a e he same as o he holo ype.
Diagnosis. Small sized Rao ches es, SVL 16–18.7 mm
in adul male, SVL in adul emale a leas 20.0 mm;
head leng h equal o sligh ly less han wid h (HL/HW =
0.92–1.0); ome ine ee h absen ; snou ounded on do -
sal iew; snou leng h equal o o sligh ly less han eye
leng h (SL/EL = 0.87–1.0); nos il equidis an be ween
eye and snou ip; dense spinules on do sal aspec o head,
do sum and limb, in e mixed wi h sca e ed blun ly con-
ical ube cles; do sally pale b own o da k b own; da k
b own “)-(“ ma k on do sum; indis inc o dis inc da k
b own c ossba s on o elimb and hindlimb; yellow ba on
in e -uppe eyelid space may be p esen ; ips o blun ly
conical ube cles whi e; i egula sized whi e lecks on
lank, la e al aspec o head, and on abdomen; sup a ym-
panic old and lowe a m pale yellow.
Desc ip ion o he holo ype. Holo ype in good condi ion
excep o an incision on en al side o igh high; SVL
17.8 mm, do so en ally la ened; head sligh ly wide
han long (HL/HW = 0.92); snou ounded in do sal iew
and nea ly unca ed in la e al iew, sligh ly p o uding
in en al iew; snou leng h sligh ly less han eye leng h
(SL/EL = 0.88); can hus os alis dis inc , ounded and
nea ly e ical; lo eal egion conca e; ympanum indis-
inc , ounded, less han one hi d o eye leng h (HTYD/
ED = 0.28); sup a ympanic old dis inc ; in e na ial dis-
ance smalle han in e -uppe eyelid wid h (INS/IUE =
0.9) and sligh ly g ea e han uppe eyelid wid h (UEW/
IUE = 0.89); nos il o al, obliquely o ien ed, equidis an
be ween eye and snou ip, na ial egion sligh ly p o ud-
ing, in e na ial space sligh ly conca e on do sal aspec ;
in e -uppe eyelid space la ; ome ine ee h absen ; cho-
anae ound; ongue no ched pos e io ly; unk do so en-
ally la ened, less han hal o snou - en leng h (AG/
SVL = 0.47).
Fo elimbs slende , o ea m sho e han hand leng h
(FAL/HAL = 0.75); digi s wi h ounded disc, ci -
cum-ma ginal g oo e p esen on each disc; me aca pal
Bo uah B e al.: Re ision o bush ogs om no heas India
568
ube cles indis inc ; disc wid hs o hi d and ou h inge
g ea e han ympanic diame e ; dis inc , la ge and inely
g anula nup ial pad p esen on i s inge , co e ing en-
ola e al and do sola e al aspec ; suba icula ube cles
ound, p oximal ones on hi d and ou h inge s small-
e and indis inc han dis al ones and indis inc , SAT =
1:1:2:2; webbing among inge s absen ; ela i e leng h o
inge s = I<II<IV<III.
Hind limbs slende ; high leng h hal o snou - en
leng h (TL/SVL = 0.49), sligh ly g ea e han ibia leng h
(TBL/TL = 0.95), and g ea e han oo leng h (FOL/
TL = 0.77); heels sligh ly o e lapping when ibia old-
ed pe pendicula o body axis; ela i e leng h o oes =
I<II<II<V<IV; oes wi h ounded disc, wid h o oe disc
equal o hose on inge , ou h oe disc la ges , disc on
ou h and i h oes wide han ympanic diame e ; ci -
cum-ma ginal g oo e p esen on each disc; na ow inne
me a a sal ube cle p esen , ou e one absen ; suba icula
ube cles ound, p oximal ones on TIII-TV indis inc ; su-
pe nume a y ube cles absen ; webbing among he oes
sligh , be ween TI and TII udimen a y, webbing no
eaching second suba icula ube cle o ou h oe.
Skin on do sal aspec o snou and head shag een,
spinules no isible as in li e condi ion; indis inc u-
be cles on in e -uppe eyelid space and on uppe eyelid;
blun ly conical ube cles on head and uppe eyelid dis-
inc ly isible in li e; ew dis inc ube cles behind an-
gle o jaw; do sum shag een wi h dense spinules, pos-
e io ly spinules no as dense as an e io pa ; spinules
mo e dis inc do sal o sup a ympanic old and do sum;
lank wi h dense spinules; o elimb and hindlimb smoo h
in p ese ed condi ion, bu spinules and blun ube cles
dis inc ly isible in li e; indis inc la ube cles p esen
a ound en ; h oa and ches smoo h; abdomen and en-
al aspec o high g anula ; ibia smoo h; la ube cle
along ou e do so en al aspec o a sus; small g anules
sca e ed on do sal su ace o ongue.
Colou a ion in li e. Do sal aspec o head and do sum
da k b own; la e al aspec o head whi ish wi h iny
b own speckle, i egula whi e pa ches and iny do s on
uppe and lowe jaw; ip o ube cles and spinules whi e;
sup a ympanic old o ange yellow; lank whi ish wi h
pale b own inge and b own speckle; lowe a m o ange
yellow; uppe a m pale b own wi h wo indis inc and
sligh ly da k b own c ossba s; inge disc wi h dense
b own speckle, o ange yellow pa ch on some o in-
ge discs; high do sally pale b own wi h sligh ly da k
b own speckle on i , knee sligh ly da ke ; la e al aspec
o high pale b own, owa ds en whi ish wi h b own
speckle, enla ge da k b own pa ch a ound en ; ibia pale
b own wi h indis inc sligh ly da ke c ossba and whi -
ish speckle; ibia and oo pale b own wi h da ke b own
and whi e speckles; oe disc wi h dense b own speckle;
ip o ube cles on hindlimb whi e; ube cles along a sus
whi e; h oa , abdomen and en al aspec o limb whi -
ish wi h b own mo ling; small whi e spo s i egula ly
placed on gula egion; whi e spo s o i egula size on
abdomen, en ola e al aspec o lank, uppe a m, high,
and a sus.
Colou a ion in p ese a i e. Head do sally pale b own,
uppe eyelid da k g ey; in e ed iangula da k g ey ma k
on pa ie al egion; lo eal egion sligh ly da ke , la e al as-
pec o head pale yellowish wi h b own speckle; do sum
g eyish b own, pale owa ds la e al side; a sligh ly da k
b own “)-(“ ma k on do sum o which an e io ends s a s
om uppe eyelid and pos e io ends ex ends o g oin; in
li e condi ion his ma king was no isible; o elimb and
hindlimb pale yellowish wi h b own mo ling; indis inc
b own c ossba s isible on o ea m, high, and ibia; en-
ally uni o m pale-yellowish wi h b own mo ling.
Sexual dimo phism and mo phological a ia ion.
Snou - en leng h o he collec ed males (N = 7) smalle
han ha o he single emale. In males, a pai o in e nal
sli s p esen on he lowe jaw; an ex e nal la ge subgula
ocal sac p esen in male; enla ged nup ial pad p esen on
he i s inge o males. Li e colou a ion a ies among
indi iduals, pa a ype (WII-ADA1531) had an o ange-yel-
low ba on in e -uppe eyelid space ollowed by a pai
o sho o ange yellow s ipe which a e join pos e io ly;
in WII-ADA1530 eddish b own hou glass shaped ma k
on da k b own p esen on do sum. In addi ion o hese
a ia ion, in ensi y o do sal spinules and ube cles a ies
among indi iduals, WII-ADA1530 and WII-ADA1537
has compa a i ely ewe sca e ed spinules on do sum
unlike o he indi iduals; ma king on do sum and c oss-
ba s on limbs ain in WII-ADA1537 and WII-ADA1538;
c ossba s on limbs o WII-ADA1536 na ow and mo e
dis inc han ha o o he indi iduals; do sal skin on head
nea ly smoo h in WII-ADA1530, WII-ADA1536 and
WII-ADA1538; o he mo phome ic a ia ions a e gi en
in Table S12.
Mo phological compa ison. Rao ches es mawsyn a-
mensis sp. no . di e s om R. ande soni, R. leik ho and
R. malipoensis by he posi ion o nos il equidis an be-
ween eye and snou ip ( s. nos il close o snou ip han
eye); i di e s om R. annandalii by p esence o dense
spinules on do sum ( s. skin on do sum smoo h); i di e s
om R. ga o by smalle body size, SVL 16.0–18.7 mm in
adul males ( s. SVL 19.9–26.9 mm in adul males); i
di e s om R. cine ascens no . comb. and R. ezakhani
by p esence o dense spinules on do sum ( s. sca e ed
ube cles on do sum); i di e s om R. dulongensis by
head leng h being equal o o less han wid h ( s. head
longe han wide), posi ion o nos il equidis an be ween
eye and snou ip ( s. nos il close o snou ip han eye);
i di e s om R. hekouensis by p esence o nup ial pad
only on i s inge ( s. nup ial pad p esen on i s and
second inge s); i di e s om R. hillisi and R. huanglian-
shan by snou leng h less han eye leng h ( s. snou leng h
snou leng h g ea e han eye leng h), posi ion o nos il
equidis an be ween eye and snou ip ( s. nos il close o
snou ip han eye); i di e s om R. jadoh by la ge body
size in adul males, SVL 16.0–18.7 mm ( s. SVL 13.6–
14.0 mm); i di e s om R. jadoh and R. jakoid by snou
leng h being less han o equal o eye leng h ( s. snou
leng h g ea e han eye leng h), and high leng h being
g ea e han o equal o ibia leng h ( s. high leng h less
Ve eb a e Zoology 75, 2025, 517–625 569
Figu e 26. Rao ches es mawsyn amensis sp. no . A–C Holo ype (WII-ADA1531) in p ese ed condi ion, A do sal iew (inse
image showing do sal spinules on do sum), B en al iew, C la e al iew o head. D–G pa a ype (WII-ADA1530) in p ese ed
condi ion. D Do sal iew, E en al iew (inse image showing g anula abdomen skin), F en al iew o le hand, G en al iew
o le oo . H Holo ype in li e, I pa a ype (WII-ADA1538) in li e; J pa a ype (WII-ADA1534) in li e; K pa a ype (WII-ADA1530)
in li e. Scale ba = 10 mm.
Bo uah B e al.: Re ision o bush ogs om no heas India
570
han ibia leng h); i di e s om R. kempiae by p esence
o da k b own “)-(“ ma k on do sum ( s. conca e s ipes
on do sum absen o ain ); i di e s om R. longchuan-
ensis by in e -uppe eyelid wid h being less han o equal
o eye leng h ( s. in e -uppe eyelid wid h g ea e han eye
leng h); i di e s om R. menglaensis by snou leng h be-
ing smalle han eye leng h ( s. snou leng h snou leng h
longe han eye leng h), head leng h being equal o o less
han wid h ( s. head longe han wide); i di e s om
R. minda by p esence o da k pa ches on g oin ( s. en-
la ged black and whi e pa ches p esen on g oin), absence
o whi e pa ches on la e al aspec o high ( s. enla ged
whi e pa ches on la e al aspec o high); i di e s om
R. pa ulus by p esence o dense spinules on do sum ( s.
sca e ed spinules on do sum); i di e s om R. o ien alis
sp. no . by p esence o compa a i ely dense spinules on
head and do sum ( s. compa a i ely less spinules on head
and do sum), dis inc enla ged ube cles absen on do sal
aspec o head absen ( s. enla ged blun ly conical ube -
cles on do sal aspec o head p esen ), h ee c ossba s on
high and ibia ( s. wo o ou c ossba s on high and
ibia); i di e s om R. shillongensis p esence o spinules
on do sum ( s. small blun ube cles on do sum); i di -
e s om R. y hus no . comb. by snou leng h being less
han o equal o eye leng h ( s. snou leng h g ea e han
eye leng h); i di e s om R. yadongensis by in e -uppe
eyelid wid h being less han o equal o eye leng h ( s.
in e -uppe eyelid wid h g ea e han eye leng h. Mo pho-
logical compa isons o R. mawsyn amensis sp. no . wi h
ou newly desc ibed species in his s udy a e p o ided in
espec i e sec ions and in Table 1.
Acous ics. The calls o R. mawsyn amensis sp. no .
we e eco ded a i s ype locali y on 1 June 2022 be-
ween 18:30–19:15 h s a an ambien empe a u e o
24.4°C–26.0°C. The call desc ip ion is based on 40 calls
including hose om wo indi iduals (WII-ADA1531 and
WII-ADA1534). The calls a e single ype, non-pulsa ile,
emi ed as a single call a egula in e als, no in g oups
(Fig. 24). The mean call du a ion is 16.2 ± 1.99 ms (13–
20 ms) wi h a ise ime o 1 ms and a all ime o 15.48
± 2.09 ms (12–19 ms). The mean in e al be ween calls
is 1418.5 ± 264.94 ms (1127–2133 ms). The dominan
equency is 4403.55 ± 74.74 Hz (4263.6–4608.1 Hz). A
de ailed compa ison o he ad e isemen calls wi h hose
o o he congene s is p esen ed in Table 2.
Phylogene ic ela ionship and gene ic di e gence.
Rao ches es mawsyn amensis sp. no . is sis e o R. bou-
lenge i sp. no . (UFB 100, PP 1.0; Figs 2, 3) wi h gene ic
di e gence o 2.5–2.9% in he 16S, 3.9–4.4% in he cy b
and 3.6–3.9% in he COI genes. The gene ic di e gences
wi h o he congene s included in his s udy a e 3.2–9.8%
in he 16S, 9.3–18.2% in he cy b and 3.6–15.6% in he
COI genes (Table S7A–C).
E ymology. This speci ic epi he is a oponym de i ed
om he name “Mawsyn am” in Meghalaya S a e which
is he we es place on ea h.
Sugges ed common name. Mawsyn am bush og.
Dis ibu ion and na u al his o y. Rao ches es mawsyn-
amensis sp. no . is known only om he ype locali y
a ound Maw apa , app oxima ely 5 km sou hwes o
Mawsyn am, Meghalaya (Fig. 25A). Calling males we e
ound abou 1–1.5 m abo e g ound on a slope in he o -
es ; he a ea was co e ed by mos ly bamboo species and
a ew palm ees, e ns and sh ubs wi h hick lea li e s;
he e was a small s eam a his si e and a oad h ough he
o es connec ing Maw apa and Mawsyn am (Fig. 20E).
We ound he single emale indi idual among lea li e .
O he sympa ic anu an species ound a his locali y in-
clude Rhacopho us bipunc a us Ahl, 1927, Ku ixalus sp.
Xenoph ys sp. and Amolops sp.
Rao ches es boulenge i sp. no .
h ps://zoobank.o g/B794ACB2-00BF-4C16-9434-
EA761972F390
Figu e 27; Tables 1, 2, S12
Holo ype. An adul male (WII-ADA1539) collec ed
by BB on 2 June 2022 om Che apunji (25.27696°N,
91.69745°E, ele a ion 1380 m a.s.l.), app oxima ely one
kilome e eas o Nohkalikai Wa e all, Eas Khasi Hills,
Meghalaya, India.
Pa a ypes. Th ee adul males (WII-ADA1540, WII-
ADA1541 and WII-ADA1542). Collec ion de ails a e he
same as o he holo ype.
Diagnosis. Small sized Rao ches es, adul male, SVL
17.5–19.1 mm; emale unknown; head leng h equal o
wid h; ome ine ee h absen ; snou ounded in do sal
iew; snou leng h less han o equal o eye leng h; snou
leng h g ea e han in e -uppe eyelid wid h; nos il equi-
dis an be ween eye and snou ip; blun ly conical ube -
cles on do sal aspec o head, uppe eyelid, behind eyelid,
do sola e al aspec o unk, on do sal aspec o ibia, and
a sus; spinules on do sum; nup ial pad p esen on i s
inge in males; pai o conca e da k b own s ipes on
do sum; c ossba s on o ea m, high, and ibia; lowe a m
o ange yellowish; sup a ympanic old pale yellowish; in-
ge disc yellow; i egula sized whi e pa ches on axilla,
g oin and abdomen.
Desc ip ion o he holo ype. Holo ype is in good condi-
ion excep o an incision unde igh high; adul male
wi h ocal sacs, SVL 17.9 mm, o e all specimen in good
condi ion, a small po ion o muscle om en al side o
igh high was sampled. Body slende , do so en ally
la ened, head o al in shape, do so en ally la ened,
leng h equal o wid h; snou ounded in do sal iew,
acu e in la e al iew, sligh ly p o uding beyond lowe
jaw; ome ine ee h absen ; choanae ound; ongue pos-
e io ly b oad and sligh ly no ched medially a pos e io
end; pai o sli like openings on lowe jaw; snou leng h
Ve eb a e Zoology 75, 2025, 517–625 571
sligh ly less han eye leng h; can hus os alis dis inc ,
ounded and e ical; lo eal egion conca e; in e -uppe
eyelid space la ; na ial egion sligh ly aised; in e na ial
dis ance sligh ly less han in e -uppe eyelid wid h (INS/
IUE = 0.87) and g ea e han uppe eyelid wid h (UEW/
INS = 0.75); nos il o al, la e ally posi ioned, obliquely
o ien ed, equally posi ioned be ween eye and snou ip;
eye mode a e in size (EL/HL = 0.45), p o uding in li e
condi ion; ympanum ba ely isible, (HTYD/EL = 0.3);
sup a ympanic old dis inc ; unk slende , do so en ally
la ened, less han hal o snou - en leng h (AG/SVL =
0.45).
Fo elimbs slende ; o ea m sligh ly longe han
hand leng h; digi s wi h ounded disc, disc on hi d and
ou h inge sligh ly wide han ympanic diame e ; ci -
cum-ma ginal g oo e on disc p esen ; palma ube cles
ba ely isible; indis inc la g anules p esen on palm;
suba icula ube cles la ge dis inc and ound, p oximal
ones on inge III and IV smalle han dis al suba icula
ube cles; suba icula ube cles on inge I–IV = 1:1:2:2;
enla ged nup ial pad on i s inge ; ela i e leng h o in-
ge s = I<II<IV<III; webbing among inge s absen .
Hindlimbs slende ; high leng h nea ly equal o ibia
leng h (TBL/TL = 0.97) and hal o he snou - en leng h
(TL/SVL = 0.49); ibia longe han oo leng h (FOL/TBL
= 0.89); oes wi h ounded disc; ela i e leng h o oes
= I<II<V<III<IV; ci cum-ma ginal g oo e p esen on oe
disc; disc on ou h oe as wide as disc on inge III and
IV; ellip ical inne me a a sal ube cle p esen , ou e one
absen ; suba icula ube cles isible, ound (1:1:2:3:2);
webbing sligh , no eaching second suba icula ube cle
on ou h oe.
Do sal skin on head nea ly smoo h, uppe eyelid g an-
ula wi h indis inc blun ube cles; ew ube cles on man-
dibula egion pos e io o angle o jaw; ube cles abo e
sup a ympanic old and abo e axilla unning do so-la -
e ally o middle o he unk; lank g anula , becoming
dis inc en o-la e ally; ube cles and spinules on head
and do sum in p ese ed condi ion no dis inc as in li e;
ube cles on o elimb and hindlimb no isible in p e-
se ed condi ion unlike in li e; gula skin loose; skin on
ches , abdomen, and on en al aspec o high g anula ,
bu g anules no as dis inc on abdomen; en al side o
o elimbs wi h ba ely isible la g anules o palm; ibia
and a sus smoo h.
Colou a ion in li e. Head do sally b own; an e io pa
o in e -uppe eyelid space sligh ly eddish b own; in-
dis inc , b own in e ed iangula ma k on head s a ing
om in e -uppe eyelid space and ex ending pos e io ly;
do sum eddish b own, pai o b oad da k b own s ipes
on do sum, inwa dly cu ed and ou e ma gins di used
owa d lank, s a ing om pos e io end o uppe eyelid,
ex ending up o g oin; ip o ube cles on head, uppe eye-
lid, do sum and lank whi e; la e al aspec o head pale
g eyish b own wi h b own mo ling on i , small i egu-
la whi e and ain b own pa ches on lips; sup a ympanic
old c eam colou ed, lowe end b oad, ain b own s ipe
in on o sup a ympanic old s a ing om pos e io co -
ne o he eye o angle o jaw; la e al aspec unk pale
g eyish b own wi h pinkish inge and b own mo ling on
i ; ew la ge whi e pa ches on g oin, en ola e al aspec
o lank and axilla; lowe a m do sally o ange yellow,
o ea ms and hands pale b own; a sligh ly da ke c ossba
on o ea m; inge disc yellow, b own mo ling on ou h
inge disc; hind limb pale b own, sho da k b own c oss-
ba s on high, ibia, and a sus, base o high do sally pale
yellowish, ibio- a sal a icula ion and ou e la e al aspec
o a sus eddish b own; disc o inne h ee oes yellow;
en ally gula egion wi h b own and whi e mo ling, ab-
domen wi h i egula whi e pa ches and b own mo ling.
Colou a ion in p ese a i e. Do sally head pale b own;
uppe eyelid da k g ey; a sligh ly da ke g eyish b own
e e sed iangula ma k on head, s a ing be ween uppe
eyelids ex ending pos e io ly, pos e io end o he ma k-
ing di used; lo eal egion sligh ly da ke , emaining la -
e al aspec o head pale ; do sum an e io ly pale b own,
pai o pale c eam colou ed b oad s ipes, inwa dly
cu ed, bo h s ipes me ge and di used pos e io ly; o e-
limb, hindlimb, and digi al discs do sally pale yellowish
wi h b own mo ling; ain b own c ossba s on o ea m,
high, and ibia. Ven ally pale yellowish wi h b own mo -
ling.
Mo phological a ia ion. De ailed mo phome ic a i-
a ions a e p o ided in Table S12. In addi ion, pai o
s ipes on do sum p ominen in he pa a ypes han ho-
lo ype; ube cles behind uppe eyelid and spinules on
do sum p ominen in WII-ADA1541 han ha o o he
pa a ypes and holo ype; c ossba on o ea m and ibia
p ominen in WII-ADA1542 han ha o o he ype spec-
imens; skin on do sal aspec o snou and head shag een
in WII-ADA1540 and WII-ADA1541; snou ip sligh ly
mo e p onounced in WII-ADA1541 han holo ype and
o he pa a ypes.
Mo phological compa ison. Rao ches es boulenge i sp.
no . di e s om R. ande soni by head leng h being equal
o wid h ( s. head leng h less han wid h), posi ion nos il
equidis an be ween eye and snou ip ( s. nos il close
o snou ip han eye); i di e s om R. annandalii by
p esence o da k b own and whi e pa ches on g oin ( s.
no pa e n on g oin), p esence o dense spinules on do -
sum ( s. sca e ed ube cles on do sum); i di e s om
R. cine ascens no . comb. by snou leng h being g ea e
han in e -uppe eyelid wid h ( s. snou leng h equal o
in e -uppe eyelid wid h), posi ion o nos il equidis an
be ween eye and snou ip ( s. nos il close o snou ip
han eye); i di e s om R. dulongensis by head leng h
being equal o head wid h ( s. head longe han i s wid h),
nup ial pad p esen on i s inge in males ( s. nup ial pad
absen ); i di e s om R. ga o by smalle body size, SVL
17.5–19.1 mm in adul males ( s. SVL 19.9–26.9 mm in
adul males); i di e s om R. hekouensis by p esence o
nup ial pad only on i s inge ( s. nup ial pad p esen
on i s and second inge ); i di e s om R. hillisi and
R. menglaensis by head leng h being equal o head wid h
( s. head longe han wide), snou leng h being less han
o equal o eye leng h ( s. snou leng h g ea e han eye
Bo uah B e al.: Re ision o bush ogs om no heas India
572
leng h); i di e s om R. huanglianshan by in e -uppe
eyelid wid h being less han eye leng h ( s. in e -uppe
eyelid wid h g ea e han o equal o eye leng h); i di e s
om R. jadoh by la ge body size in adul males, SVL
17.5–19.1 mm ( s. SVL 13.6–14.0 mm), and in e -uppe
eyelid wid h being less han eye leng h ( s. in e -uppe
eyelid wid h g ea e han eye leng h); i di e s om R. ja-
koid by snou leng h being less han o equal o eye leng h
( s. snou leng h g ea e han eye leng h), and high leng h
being g ea e han ibia leng h ( s. high leng h less han
ibia leng h); i di e s om R. leik ho by la ge body size
in adul males, SVL 17.5–19.1 mm ( s. SVL 15.7–15.8
mm), posi ion o nos il equidis an om eye and snou
ip ( s. nos il close o snou ip han eye); i di e s om
R. longchuanensis and R. yadongensis by in e -uppe
eyelid wid h being smalle han eye leng h ( s. in e -up-
pe eyelid wid h g ea e han eye leng h); i di e s om
R. malipoensis by in e -uppe eyelid wid h being smalle
han eye leng h ( s. in e -uppe eyelid wid h g ea e han
o equal o eye leng h); i di e s om R. minda by p es-
ence o compa a i ely smalle whi e pa ches on g oin ( s.
enla ged black and whi e pa ches p esen on g oin), ab-
sence o whi e pa ches on la e al aspec o high ( s. en-
la ged whi e pa ches on la e al aspec o high); i di e s
om R. pa ulus by high leng h g ea e han ibia leng h
( s. high leng h smalle han o equal o ibia leng h),
h ee c ossba s p esen on high and ibia ( s. one on
high and one o wo on ibia); i di e s om R. ezakhani
by head leng h being equal o wid h ( s. head wide han
long), snou leng h being g ea e han in e -uppe eyelid
wid h ( s. snou leng h less han o equal o in e -uppe
eyelid wid h); i di e s om R. shillongensis by p esence
o dense spinules on back ( s. skin on do sum mode -
a ely o hea ily ube cula ed); i di e s om R. y hus
no . comb. by snou leng h being less han o equal o
eye leng h ( s. snou longe han eye leng h), posi ion o
nos il equidis an be ween eye and snou ip ( s. nos il
close o snou ip han eye); mo phologically R. bou-
lenge i sp. no . is close o R. mawsyn amensis sp. no .
A de ailed mo phological compa ison o his species wi h
i s congene s is summa ized in Table 1.
Acous ics. The calls o R. boulenge i sp. no . eco ded in
i s ype locali y on 2 June 2022 a 18:15 h s a an ambien
empe a u e o 25.9°C. The call desc ip ion is based on
40 calls om he holo ype (WII-ADA1539). The calls a e
single ype, non-pulsa ile (Fig. 24), emi ed as a single call
a he beginning and g adually modi ied in g oups (2 calls
pe g oup). The mean call du a ion is 25.03 ± 3.48 ms
(20–31 ms) wi h a ise ime o 1 ms and all ime o 24.1
± 3.47 ms (20–30 ms). The mean in e al be ween calls is
187.06 ± 13.18 ms (173–216 ms) when calls we e emi ed
in g oups. The dominan equency is 3740.33 ± 20.81 Hz
(3703.7–3789.8 Hz). The call o R. boulenge i sp. no .
di e s om he call o i s sis e species R. mawsyn amen-
sis sp. no . by he ollowing cha ac e s: call du a ion o
R. boulenge i sp. no . is gene ally longe , 20–31 ms ( s.
call du a ion 13–20 ms), in e -call in e al is lowe , 173–
216 ms ( s. in e -call in e al 1127–2133 ms), and peak
equency ange is lowe , 3703–3789 Hz ( s. 4263–4608
Hz). A de ailed compa ison o he ad e isemen calls
wi h hose o he congene s is p esen ed in Table 2.
Figu e 27. Rao ches es boulenge i sp. no . A–C Holo ype (WII-ADA1539), A do sal iew and B en al iew in p ese ed con-
di ion, C holo ype in li e. D–F Pa a ype (WII-ADA1542) in p ese ed condi ion, D do sal iew, E en al iew, F la e al iew o
head. Scale ba = 10 mm.
Ve eb a e Zoology 75, 2025, 517–625 573
Phylogene ic ela ionship and gene ic di e gence.
Rao ches es boulenge i sp. no . is sis e o R. mawsyn-
amensis sp. no . (Figs 2, 3). The gene ic di e gence be-
ween hese wo species is 2.5–2.9% in he 16S, 3.9–4.4
in he cy b, and 3.6–3.9% in he COI genes. The gene ic
di e gence o R. boulenge i sp. no . wi h o he conge-
ne s a ied be ween 3.4–9.8% in he 16S, 9.3–18.0% in
he cy b and 6.0–14.6% in he COI genes (Table S7A–C).
E ymology. This species is named a e Geo ge Albe
Boulenge o his con ibu ion o Indian he pe ology, pa -
icula ly bush ogs.
Sugges ed common name. Boulenge ’s bush og.
Dis ibu ion and na u al his o y. Rao ches es bou-
lenge i sp. no . is cu en ly known only om he ype
locali y (Fig. 25A). Calling indi iduals o his species
we e eco ded on sh ubs and bamboo hicke s in a small
isola ed o es pa ch. The o es pa ch was su ounded
by g asslands. A small s eam was p esen app oxima e-
ly 300 m wes o he o es . O he sympa ic anu an spe-
cies eco ded in he a ea we e Amolops sp. and Mine -
a ya sp.
Rao ches es ba akensis sp. no .
h ps://zoobank.o g/52117744-1919-4F83-A854-
0A72C30F7935
Figu e 28; Tables 1, 2, S12
Holo ype. An adul male (WII-ADA612) collec ed by
BB on 18 Ap il 2019 om Ma uwache a (24.973744°N,
92.768593°E, ele a ion 50 m a.s.l.), Cacha Dis ic , As-
sam, India.
Pa a ypes. Six adul males (WII-ADA611, WII-
ADA613– WII-ADA617). Collec ion de ails a e same as
o he holo ype.
Re e ed ma e ial. Th ee adul males (WII-ADA862,
WII-ADA863 and WII-ADA865) collec ed by BB and
AD on 2 Augus 2021 om A amsang am (25.8312°N,
94.8731°E, ele a ion 1460 m a.s.l.), app oxima ely 1.6
km (ae ial dis ance) no heas o Pung o, Kiphi e Dis ic ,
Nagaland; one adul male (WII-ADA942) collec ed by
BB, NGP and AD on 4 Sep embe 2021 om Hmui ang
(23.45357°N, 92.75236°E, ele a ion 1500 m a.s.l.), Aiza-
wl Dis ic , Mizo am; one adul male (WII-ADA1050)
collec ed by BB, NGP and AD on 13 Sep embe 2021 om
Phuldungsei (23.53537°N, 92.41584°E, ele a ion 780 m
a.s.l.), Dampa Tige Rese e, Mami Dis ic , Mizo am;
se en adul males (WII-ADA1335, WII-ADA1340,
WII-ADA1346–WII-ADA1350) collec ed by BB and
AD on 13 May 2022 om Mo ijheel T ail (27.49963°N,
96.33256°E, ele a ion 370 m a.s.l.), Namdapha Tige
Rese e, Changlang Dis ic , A unachal P adesh; h ee
adul males (WII-ADA1451–WII-ADA1453) collec -
ed by BB, VJ and AD on 20 May 2022 om Gibbons
Land (27.50168°N, 96.33169°E, ele a ion 350 m a.s.l.),
Namdapha Tige Rese e, Changlang Dis ic , A unachal
P adesh; wo adul males (WII-ADA1643 and WII-
ADA1644) collec ed by BB on 25 July 2022 om Lam-
dan (24.5954°N, 93.70851°E, ele a ion 1270 m a.s.l.),
Chu achandpu Dis ic , Manipu ; one adul male (WII-
ADA3207) collec ed by RNV and SD on 5 May 2023
nea M’Pen illage (27.50166°N, 96.32147°E, ele a ion
370 m a.s.l.), Changlang Dis ic , A unachal P adesh.
Diagnosis. Small sized Rao ches es, SVL 16.7–21.0 mm
in adul males; head leng h equal o o less han wid h
(HL/HW = 0.89–1.01); snou leng h less han o equal o
eye leng h (SL/EL = 0.82–1.03); nos il equidis an be-
ween eye and snou ip o sligh ly close o snou ip; in-
e na ial dis ance less han o equal o in e -uppe eyelid
wid h (IN/IUE = 0.74–1.0) and g ea e han uppe eyelid
wid h (UEW/IN = 0.7–0.91); in e -uppe eyelid wid h
smalle han eye leng h (IUE/EL = 0.72–0.96); ome ine
ee h absen ; nup ial pad p esen on i s inge in males;
blun ube cles on head, uppe eyelid, behind uppe eye-
lid, o ea m, high, iba, and a sus; spinules on do sum
in e mixed wi h blun ube cles; a b own “)-(“ ma king
on do sum; b own c ossba s on o ea m, high and ibia;
disc on inne wo inge s yellow; bony p ojec ion on hu-
me us on en al aspec .
Desc ip ion o he holo ype. Holo ype in good condi ion
excep o an incision on abdomen owa ds le side and
ano he incision on en al side o igh high. Adul male
wi h ocal sac, 20.4 mm in SVL; head leng h equal o
wid h; snou ounded in do sal iew, unca ed in la e al
iew and sligh ly p o uding beyond lowe jaw; can hus
os alis smoo h, obliqued; lo eal egion conca e; snou
leng h equal o eye leng h; nos il o al, la e ally posi-
ioned, obliquely o ien ed, sligh ly close o snou ip han
eye (NS/EN = 0.86); in e na ial dis ance sligh ly smalle
han in e -uppe eyelid wid h (IN/IUE = 0.88) and g ea -
e han uppe eyelid wid h (UEW/IN = 0.86); in e na ial
egion sligh ly conca e; a ea be ween nos il and snou
ip sligh ly conca e; ympanum dis inc , ound, i s diam-
e e less han one hi d o eye leng h; ongue pos e io ly
wide and deeply no ched; ome ine ee h absen ; choanae
ound; pai o in e nal sli like openings p esen on low-
e jaw owa ds edges; symphysial knob p esen on lowe
jaw; habi us s ou , hal o he snou - en leng h.
Fo elimbs slende , hand leng h sligh ly less han o e-
a m leng h (HAL/FAL = 0.94), digi wi h ounded disc;
ci cum-ma ginal g oo e p esen on each disc; hi d inge
longes , ela i e leng h o inge s = I<II<IV<III; disc on
hi d inge la ges and sligh ly wide han ympanic di-
ame e (HTYD/FIIID = 0.73); webbing on hand absen ;
suba icula ube cles dis inc , enla ged and ound excep
p oximal ones on hi d and ou h inge s which a e indis-
inc and smalle ; suba icula ube cle o mula on inge s
I–IV = 1:1:2:2; palma ube cles indis inc ; bony p ojec-
ion on hume us di ec ed en ally (Fig. 25)
Hindlimbs slende ; high leng h hal o snou - en
leng h (TL/SVL = 0.51); ibia leng h nea ly equal o high
Bo uah B e al.: Re ision o bush ogs om no heas India
580
la ged yellow spo s on high); di e s om R. hekouensis
by la ge body size, SVL 18.0–21.1 mm in adul males
( s. SVL 16.1–17.5 mm in adul males), nup ial pad p es-
en only on i s inge ( s. nup ial pad p esen on i s
and second inge ); i di e s om R. hillisi by la ge body
size, SVL 18.7–21.2 mm in adul males and SVL a leas
up o 23.1 mm in adul emale ( s. SVL 14.5–17.7 mm in
adul males and 17.5 in adul emale), head leng h equal
o wid h o sligh ly wide han long ( s. head longe han
wide), da k b own pa ch on g oin absen ( s. p esen ); i
di e s om R. huanglianshan by da k b own pa ch on
g oin absen ( s. p esen ), snou leng h being equal o o
less han eye leng h ( s. snou leng h g ea e han eye
leng h); i di e s om R. jadoh by la ge body size in
adul males, SVL 18.0–21.1 mm ( s. 13.6–14.0 mm), and
in e -uppe eyelid wid h being less han eye leng h ( s. in-
e -uppe eyelid wid h g ea e han eye leng h); i di e s
om R. jakoid by snou leng h being less han o equal o
eye leng h ( s. snou leng h g ea e han eye leng h), in-
e -uppe eyelid wid h less han eye leng h ( s. in e -up-
pe eyelid wid h g ea e han o equal o eye leng h), and
high leng h being g ea e han o equal o ibia leng h
( s. high leng h less han ibia leng h); i di e s om R.
kempiae by do sal skin, sca e ed ube cles p esen do -
sum ( s. do sal skin o do sum wi h spinules and blun
ube cles), absence o ma king on g oin ( s. sho b own
s ip p esen on g oin); i di e s om R. longchuanen-
sis by inne -uppe eyelid being smalle han eye leng h
( s. in e -uppe eyelid g ea e han eye leng h); i di e s
om R. leik ho by la ge body size in adul males, SVL
Figu e 31. Compa a i e spec og ams and oscillog ams illus a ing ep esen a i e calls o h ee species o Rao ches es. A, B Oscil-
log ams, C spec og am, D powe spec um. The call ma ked wi h an a ow on op in A is expanded in B.
Ve eb a e Zoology 75, 2025, 517–625 581
18–21.1 mm ( s. SVL 15.7–15.8 mm); i di e s om
R. malipoensis by i s la ge body size, SVL 18.7–21.2
mm in adul males and SVL a leas up o 23.1 mm in
adul emale ( s. SVL 14.6–17.7 mm in adul males and
18.3–19.3 in adul emales), black pa ches on g oin and
la e al aspec o high absen ( s. p esen ); i di e s om
R. menglaensis by head leng h being equal o wid h o
sligh ly wide han long ( s. head longe han wide), da k
b own o black blo ch on g oin absen ( s. p esen ); i di -
e s om R. minda by absence o da k o whi e pa ches
on g oin ( s. enla ged black and whi e pa ches p esen on
g oin), no whi e pa ches on la e al aspec o high ( s. en-
la ged whi e pa ches on la e al aspec o high); i di e s
om R. boulenge i sp. no . and R. mawsyn amensis sp.
no . in ha ing ela i e smoo h skin wi h sca e ed ube -
cles on do sum ( s. dense spinules on do sal aspec o
head, do sum, and limbs), being nos il close o snou
ip han eyes ( s. nos il equidis an om eyes and snou
ip in R. boulenge i sp. no . and R. mawsyn amensis sp.
no .); di e s om R. o ien alis sp. no . by p esence o
sca e ed ube cles on do sum ( s. dense spinules on do -
sum), p esence o b oad da k b own c ossba on high
and ibia ( s. h ee c ossba s on high and ibia), p esence
o pai o conca e s ipes on do sum ( s. a “)-(“ ma k
on do sum); i di e s om R. pa ulus snou sub o oid
o nea ly acu e ( s. snou ounded), absence o ma king
on g oin ( s. da k b own ma bling enclosing a whi ish
blo ch on g oin); i di e s om R. ezakhani by p esence
o single b oad c ossba on each high and ibia ( s. a
leas h ee na ow da k b own c ossba s on each high
and ibia), absence o b own pa ch on g oin ( s. a sho
b own s eak p esen on g oin); i di e s om R. shillon-
gensis by absence o ma king on g oin ( s. da k b own
pa ch wi h whi e c own shaped ma k o di used b own
pa ch on g oin); i di e s om R. y hus no . comb. by
snou leng h being equal o o less han eye leng h ( s.
snou leng h g ea e han eye leng h), absence o spo o
ma king on g oin ( s. a la ge whi e spo on g oin); i di -
e s om R. yadongensis by absence o whi e pa ches on
ches and abdomen ( s. p esen ). A de ailed mo phologi-
cal compa ison is p o ided in Table 1.
Acous ics. The calls o R. na puhensis sp. no . we e
eco ded in Na puh Wildli e Sanc ua y, Meghalaya on
29 May 2022 be ween 19:00–21:00 h s a an ambien
empe a u e o 27.5°C. The call desc ip ion is based
on 40 calls om wo indi iduals (WII-ADA1509 and
WII-ADA1510). Calls a e o a single ype, non-pulsa-
ile, emi ed a egula in e als and no in g oups (Fig.
31). The mean call du a ion is 18.7 ± 4.02 ms (13–25
ms) wi h a ise ime o 1 ms and a all ime o 17.68 ±
3.83 ms (12–24 ms). The mean call in e al be ween calls
is 670.17 ± 85.57 ms (544–823 ms). The dominan e-
quency is 3954.57 ± 165.48 Hz (3574.5–4134.4 Hz). A
de ailed compa ison o he ad e isemen calls wi h hose
o he congene s is p esen ed in Table 2.
Phylogene ic ela ionship. Rao ches es na puhensis sp.
no . is sis e o R. lawng laiensis sp. no . (UFB 100, PP
1.0; Figs 2, 3) wi h gene ic di e gence o 2.3–2.7% in
he 16S, 8.1–8.2% in he cy b and 8.7–8.9% in he COI
genes. The closes ae ial dis ance be ween ype locali ies
o hese wo species is 293 km. Rao ches es na puhensis
sp. no . has gene ic di e gence o 5.1–10.0% in he 16S,
13.7–20.8% in he cy b and 8.7–15.7% in he COI genes
wi h he o he membe s o he genus (Table S7A–C).
E ymology. The speci ic epi he is a oponym de i ed
om he name “Na puh Wildli e Sanc ua y” in Megha-
laya S a e whe e he ype se ies we e collec ed.
Sugges ed common name. Na puh bush og.
Dis ibu ion and na u al his o y. Cu en ly Roa ches-
es na puhensis sp. no . is known om i s ype locali y
in and a ound Na puh Wildli e Sanc ua y, and in Riwai
o Meghalaya (Fig. 19A). We encoun e ed calling males
and a single emale a ~1–2 m abo e g ound pe ching on
sh ubs and on small bamboo lea es, Zingibe sp. on hill
slopes (Fig. 20H). O he sympa ic species ound wi h he
new species we e Chi ixalus sp. and Polypeda es c . hi-
malayensis.
Rao ches es lawng laiensis sp. no .
h ps://zoobank.o g/CD636615-6E5F-46AC-A69A-F1D656D-
EF592
Figu e 32; Tables 1, 2, S12
Holo ype. WII-ADA947, an adul male collec ed by
AD, BB and NGP on 6 Sep embe 2021 om Ngengpui
Wildli e Sanc ua y (22.48498°N, 92.75653°E, ele a ion
210 m a.s.l.), Lawng lai Dis ic , Mizo am, India.
Pa a ype. An adul male (WII-ADA946) collec ed by
AD, BB and NGP on 6 Sep embe 2021 om Ngengpui
Wildli e Sanc ua y (22.48906°N, 92.75678°E, ele a ion
210 m a.s.l.), Lawng lai Dis ic , Mizo am.
Diagnosis. A small sized Rao ches es, SVL = 18.1–19.7
mm in males, emale unknown; snou ounded, sligh -
ly less han eye leng h; nos il close o snou ip han
eye; head sligh ly wide han long o equal p opo ions;
in e -uppe eyelid wid h less han eye leng h; ome ine
ee h absen ; nup ial pad p esen on i s inge in males;
do sal aspec pale yellowish b own; “)(“ shaped b oad
da k b own ma king on do sum, pos e io ends o his
ma king bo de ed by i egula whi e spo s; an oblique
da k b own s ipe bo de ed wi h whi e spo s on lank;
da k c ossba s bo de ed wi h whi e spo s on o elimb and
on hindlimb; i egula small whi e spo s on do sal aspec
o head; do sum and limb; ympanic egion da k b own;
a da k b own s eak en al o eye bo de ed wi h whi e
s eak; g oin, axilla, hind a m, la e al side o high, inge
and oe discs yellow; i egula whi e lecks on ches and
abdomen; o elimb and hindlimb da k b own; gula e-
gion yellowish b own.
Bo uah B e al.: Re ision o bush ogs om no heas India
582
Desc ip ion o he holo ype. Adul male, SVL 18.1 mm;
head wide han long (HL/HW = 0.95); snou ounded
wi h a blun ube cula p ojec ion a ip, unca ed in
la e al iew, snou leng h less han eye leng h (SL/EL =
0.83); nos il o al, equidis an be ween eyes and snou
ip; in e -uppe eyelid wid h g ea e han in e na ial dis-
ance (IN/IUE = 0.76); in e na ial dis ance equal o uppe
eyelid wid h; eye leng h nea ly hal o head leng h (EL/
Figu e 32. Rao ches es lawng laiensis sp. no . A–E Holo ype (WII–ADA947) in p ese a i e and F, G holo ype in li e; A do sal
iew, B en al iew, C la e al iew o head, D en al iew o igh hand, E en al iew o igh oo ; F do sola e al iew, G en al
iew. H, I Pa a ype (WII-ADA946) in p ese a i e and J–M in li e; H do sal iew, I en al iew, J, K do sola e al iew, L en al
iew, M g oin. Scale ba = 10 mm.
Ve eb a e Zoology 75, 2025, 517–625 583
HL = 0.46); can hal idge dis inc , oblique; lo eal sligh ly
conca e; ympanum ounded, dis inc ; ympanic diame-
e one i h o eye leng h; sup a ympanic old dis inc ;
ome ine ee h absen ; choanae ound; ongue no ched
medially a pos e io end and wi h spa sely dis ibu ed
la g anules.
Fo elimbs slende , o ea m sho e han hand leng h
(FAL/HAL = 0.86); hi d inge longes , ela i e leng h o
inge s = I<II<IV<III; inge wi h ounded disc, disc wi h
ci cum-ma ginal g oo e; disc on second, hi d and ou h
inge wide han ho izon al ympanic diame e ; suba ic-
ula ube cles ound and dis inc excep p oximal ones on
hi d and ou h inge s; palma ube cles indis inc ; nup-
ial pad p esen on i s inge .
Hindlimbs slende , high leng h longe han shank
leng h (TBL/TL = 0.92) and oo leng h (FOL/TL =
0.85); shank longe han oo leng h (FOL/TBL = 0.92);
ou h inge longes oes, ela i e leng h o oes =
I<II<III<V<IV; oe wi h ounded disc, disc wi h ci -
cum-ma ginal g oo e, disc wid h o ou h oe sligh ly
less han ha o hi d oe; suba icula ube cles ound
and dis inc excep p oximal ones on ou h and i h oes;
inne me a a sal ube cle p esen , longe han ho izon al
ympanic diame e , ou e me a a sal ube cle absen .
Skin on do sal aspec o head smoo h wi h sca e ed
spinules; dis inc spinules on uppe eyelid; ew g anula
ube cles behind angle o jaw; nume ous spinules on do -
sum, abo e sup a ympanic old and lank; limb smoo h
do sally; h oa , ches , abdomen and en al aspec o
high g anula ; ibia and a sus smoo h.
Colou a ion in li e. Do sal aspec o head and do sum
pale yellowish b own; sligh ly da k b own pa ch on
snou and pos e io pa o head; ube cles on snou , up-
pe eyelid and pos e io pa o head whi e; la e al aspec
o head pale yellowish b own; na ial egion da k b own;
al e na i e da k b own and whi e bands on uppe jaw;
whi e speckles on angle o jaw and mandibula egion;
ympanic egion da k b own; ew i egula sized whi e
spo s on do sum; a pai o conca e da k b own s ipes
on do sum, pos e io ly p ominen and edged wi h whi e
speckles; ano he simila sho s ipe on lank; whi e
lecks on axilla; i egula whi e spo s on o elimb; inge
discs yellow; whi e spo s o i egula size and shape on
hindlimb; an indis inc b oad c ossba on high and ibia;
g oin and la e al aspec o high pale yellow; en al as-
pec o head yellowish wi h dense b own and whi e ma -
bling, ew small whi e pa ches along lowe jaw; i egula
whi e lecks on ches , abdomen, o elimb and hind limb;
inge and disc pale yellow; yellow inge on oe, i s oe
on le side yellow.
Colou a ion in p ese a i e. Do sal and en al co-
lou a ion simila wi h ha o colou a ion in li e excep
ha whi e spo s isible in li e a e no isible in p ese ed
condi ion; b own e icula ion on pale-c eam colou ed
backg ound on ches and abdomen.
Mo phological compa ison. Rao ches es lawng laien-
sis sp. no . di e s om R. ande soni by posi ion o
nos il equidis an be ween eye and snou ip ( s. nos il
close o snou ip han eye); i di e s om R. annandalii
by in e na ial dis ance being equal o uppe eyelid wid h
( s. in e na ial dis ance g ea e han uppe eyelid wid h),
snou leng h being less han eye leng h ( s. snou leng h
equal o o g ea e han eye leng h); i di e s om R. ba -
akensis sp. no . by absence o bony p ojec ion on hu-
me us ( s. bony p ojec ion on hume us p esen ); i di e s
om R. boulenge i, R. kempiae, R. o ien alis sp. no .,
and R. shillongensis sp. no . by in e na ial dis ance being
equal o uppe eyelid wid h ( s. in e na ial dis ance being
g ea e han uppe eyelid wid h); i di e s om R. cine -
ascens no . comb. by p esence o a single b oad c ossba
on high and ibia ( s. h ee c ossba s on high and ibia);
i di e s om R. dulongensis by snou leng h being less
han eye leng h ( s. snou leng h equal o eye leng h); i
di e s om R. ga o by smalle body size, SVL 18.1–19.7
mm in adul males ( s. SVL 19.9–26.9 in adul males); i
di e s om R. hekouensis, R. hillisi and R. malipoensis
by la ge body size, SVL 18.1–19.7 mm in adul males
( s. SVL 16.1–17.5 mm in R. hekouensis, 15.9–17.7 mm
in R. hillisi and 14.6–17.7 in R. malipoensis); i di e s
om R. hillisi by snou leng h being less han eye leng h
( s. snou leng h g ea e han eye leng h); i di e s om
R. huanglianshan by snou leng h being less han eye
leng h ( s. snou leng h g ea e han eye leng h); i di e s
om R. jadoh by la ge body size in adul males, SVL
18.1–19.7 mm ( s. 13.6–14.0 mm), and in e -uppe eye-
lid wid h being less han eye leng h ( s. in e -uppe eye-
lid wid h g ea e han eye leng h); i di e s om R. ja-
koid by snou leng h being less han eye leng h ( s. snou
leng h g ea e han eye leng h), in e -uppe eyelid wid h
being less han eye leng h ( s. in e -uppe eyelid wid h
g ea e han o equal o eye leng h), and high leng h be-
ing g ea e han o equal o ibia leng h ( s. high leng h
less han ibia leng h); i di e s om R. leik ho by la ge
body size in adul males, SVL 18.1–19.7 mm ( s. SVL
15.7–15.8 mm); i di e s om R. longchuanensis by in-
e -uppe eyelid wid h being less han eye leng h ( s. in-
e -uppe eyelid wid h g ea e han eye leng h); i di e s
om R. malipoensis by in e -uppe eyelid wid h being
less han eye leng h ( s. in e -uppe eyelid wid h g ea e
han o equal o eye leng h); i di e s om R. mawsyn-
amensis sp. no . by p esence o mode a ely dense
spinules on do sum ( s. compa a i ely dense spinules
on do sum), p esence o da k b own s ipe edged wi h
whi e spo s on lank ( s. absen ); i di e s om R. meng-
laensis by snou leng h being smalle han eye leng h ( s.
snou leng h longe han eye leng h), head leng h being
equal o o sligh ly smalle han i s wid h ( s. head longe
han i s wid h); i di e s om R. minda by p esence o
da k b own band bo de ed wi h whi e spo s on g oin ( s.
enla ged black and whi e pa ches p esen on g oin), by
absence o whi e pa ches on la e al aspec o high ( s.
enla ged whi e pa ches on he la e al aspec o high); i
di e s om R. na puhensis sp. no . by in e na ial dis-
ance being equal o uppe eyelid wid h ( s. in e na ial
dis ance g ea e han uppe eyelid wid h), p esence o
da k b own s ipe edged wi h whi e spo s on g oin ( s.
no pa e n on g oin); i di e s om R. pa ulus by high
Bo uah B e al.: Re ision o bush ogs om no heas India
584
Figu e 33. Habi a a he ype locali ies o Rao ches es species in no heas India. A Lowland o es s eam a he ype locali y
o R. lawng laiensis sp. no . in Ngengpui WLS; B landscape iew o Mehao WLS om Abango, ype locali y o R. dibangensis
sp. no . in Abango; C. hill s eam and ipa ian habi a a he ype locali y o R. shillongensis in Malki, Shillong; D landscape iew
o he hills a Pumdunlong, he ype locali y o R. monoli hus sp. no . a Willong Khullen Village; E landscape iew o hills and
lowland a eas a Khonoma, he ype locali y o R. khonoma sp. no .; F o es co e a Bompu, Eaglenes WLS a he ype locali y
o R. eaglenes ensis sp. no .; G hill s eam and o es co e a Abo hills, habi a o R. magnus sp. no .; H landscape iew o Abo
hills and adjacen lowland a eas, habi a o R. a unachalensis sp. no .
Ve eb a e Zoology 75, 2025, 517–625 585
being longe han ibia ( s. high sho e han o equal o
ibia leng h); i di e s om R. ezakhani by p esence o
da k b own s ipe edged wi h whi e spo s on lank ( s.
no b own s ipe on lank); i di e s om R. y hus no .
comb. by snou leng h being less han eye leng h ( s.
snou leng h g ea e han eye leng h); i di e s om R.
yadongensis by in e -uppe eyelid wid h being less han
eye leng h ( s. in e -uppe eyelid wid h g ea e han eye
leng h), by snou leng h being less han eye leng h ( s.
snou leng h equal o eye leng h). A de ailed mo pholog-
ical compa ison is gi en in Table 1.
Phylogene ic ela ionship and gene ic di e gence.
Rao ches es lawng laiensis sp. no . is sis e o R. na -
puhensis sp. no . (Figs 2, 3). The gene ic di e gence wi h
o he congene s included in his s udy is 2.3– 9.3% in he
16S, 13.9– 21.0% in he cy b and 13.9– 21.0% in he COI
genes (Table S7A–C).
E ymology. The speci ic epi he is a oponym de i ed
om he name “Lawng lai” Dis ic in Mizo am S a e
whe e he ype locali y o his species lies.
Sugges ed common name. Lawng lai bush og.
Dis ibu ion and na u al his o y. Rao ches es lawng -
laiensis sp. no . is cu en ly known om i s ype locali-
y in Ngengpui Wildli e Sanc ua y, Lawng lai, Mizo am
(Fig. 19A). The holo ype was ound calling on a lea ap-
p oxima ely a one me e heigh abo e g ound along a
o es ail a a ound 23:00 h s. The pa a ype was eco d-
ed while i was calling om a bamboo b anch app ox-
ima ely 1.5 m abo e g ound a a ound 20:15 h s a he
edge o a small s eam o app oxima ely wo me e s in
wid h (Fig. 33A).
Rao ches es nasu a sp. no .
h ps://zoobank.o g/F55DCD22-E950-431A-B542-F145DC-
C252C9
Figu e 34; Tables 1, 2, S12
Holo ype. An adul male (WII-ADA3388) collec ed by
RNV on 12 June 2023 om Ranijheel (27.53903°N,
96.4888°E, ele a ion 910 m a.s.l.), Namdapha Tige Re-
se e, Changlang Dis ic , A unachal P adesh, India.
Pa a ypes. Two adul males (WII-ADA3398 and WII-
ADA3399) collec ed by RNV on 13 June 2023 om
Bulbulia (27.53872N, 96.4763E, ele a ion 880 m a.s.l.),
Namdapha Tige Rese e, Changlang Dis ic , A unachal
P adesh.
Re e ed Ma e ial. Two adul males (WII-ADA3216
and WII-ADA3218) collec ed by RNV and SD on 10
May 2023 om Ho nbill (27.53810°N, 96.44037°E, el-
e a ion 660 m a.s.l.), Namdapha Tige Rese e, Chang-
lang Dis ic , A unachal P adesh.
Diagnosis. A small sized Rao ches es, SVL 17.0–19.9
mm in adul males, emale unknown; head longe han
wide; snou acu e wi h de mal p ojec ion on snou ip;
snou leng h equal o eye leng h; in e -uppe eyelid wid h
less han o equal o eye leng h (IUE/EL = 0.88–0.96);
nos il equidis an be ween eye and snou ip; ho n like
de mal p ojec ion on edge o uppe eyelid; an hou -glass
shaped da k b own ma k s a ing on in e -uppe eyelid
space and ex ending o do sum; b igh yellow pa ches on
g oin and high; inge and oe disc yellow.
Desc ip ion o he holo ype. Holo ype well p ese ed
excep o an incision on he en al side o igh high.
Adul male, SVL 17.9 mm; head longe han wide (HW/
HL = 0.9); snou acu e in do sal and la e al iew as a de -
mal p ojec ion p esen on snou ip, p o uding beyond
lowe jaw in en al iew; snou leng h equal o eye leng h
and sligh ly g ea e han in e -uppe eyelid wid h (IUE/
SL = 0.85); symphysial knob p esen on lowe jaw; na ial
egion p o uding; nos il o al, la e ally posi ioned and
obliquely o ien ed, equidis an be ween eye and snou ip;
sligh ly conca e on do sal aspec o snou ; can hus os-
alis ounded, e ical; lo eal conca e; eye mode a e in
size (EL/HL = 0.37); in e na ial dis ance nea ly equal o
in e -uppe eyelid wid h (IN/IUE = 0.91) and g ea e han
uppe eyelid wid h (UEW/IN = 0.86); ympanum indis-
inc , ound, one hi d o eye leng h (HTYD/EL = 0.31);
sup a ympanic old dis inc ; ome ine ee h absen ; cho-
anae ound; ongue pos e io ly deeply no ched and he
wo lobes a e acu e; pai o in e nal ocal sac openings
on lowe jaw; habi us less han hal o snou - en leng h
(AG/SVL = 0.46).
Fo elimbs slende ; o ea m nea ly equal o hand
leng h (FAL/HL = 0.96); ela i e leng h o inge s =
FI<FII<FIV<FIII; ounded disc on inge , disc on inge
III and IV wide han ympanic diame e ; ci cum-ma -
ginal g oo e on each disc; udimen a y webbing p esen
be ween inge s; palma ube cles indis inc ; suba icula
ube cles ound, lowe suba icula ube cle on inge III
and IV small and indis inc ; enla ged, ine g anula nup-
ial pad on i s inge .
Hindlimbs slende ; high longe han ibia (TBL/TL
= 0.89) and hal o snou - en leng h (TL/SVL = 0.5);
ela i e leng h o oes = TI<TII<TIII<TV<TIV; ounded
disc on oe, smalle han hose o inge s; ci cum-ma -
ginal g oo e p esen each disc; suba icula ube cles
dis inc in li e condi ion, p oximal suba icula on ou h
and i h oes indis inc ; inne me a a sal ube cle p esen ,
i s leng h equal o wid h o discs o oes II–V; e y hin
de mal inge along i h oe; webbing mode a e, eaching
beyond second suba icula ube cle on ou h oe.
Skin on do sal aspec o snou and head smoo h (sha-
g een in li e condi ion); indis inc ube cles on uppe
eyelid, a ho n like ube cle on edge o each uppe eye-
lid; ube cles on angle o jaw; skin on do sum (shag een
in li e condi ion) wi h spinules, sca e ed on middle and
dense owa ds do sola e al aspec ; a weak mid-do sal
de mal idge isible in li e condi ion s a ing om snou
and becoming di used pos e io ly; lank smoo h; do sal
aspec o o elimb and hindlimb smoo h; h oa and ches
Bo uah B e al.: Re ision o bush ogs om no heas India
586
smoo h; abdomen and high g anula ; ibia and a sus
smoo h.
Colou a ion in li e. Do sal aspec o snou and op o
head, do sum b own; an hou glass shaped da k b own
ma king s a ing a in e -uppe eyelid space and co e ing
pos e io hal o uppe eyelid ex ending o do sum, be-
coming di used pos e io ly; a di used da k b own b oad
s ipe along sup a ympanic old; ano he simila s ipe
on lo eal egion eaching snou ip; i egula pale c eam
colou ed whi e spo s on uppe jaw; wo yellow spo s on
g oin on each side and inne la e al aspec o high; on
ou e la e al aspec o high yellow pa ches ba ely isible;
a b oad c oss ba on each high and ibia ba ely isible;
disc on digi s yellow; en al aspec o head, ches , ab-
domen and high b own, pale han do sal aspec ; dense
da k b own mo ling on chin, h oa wi h ew whi e lecks
on an e io pa o he jaw; b own mo ling on en al
aspec o o elimb ches , abdomen and high no dense;
i egula whi e lecks on o ea m, high, a sus and oo ;
g anules on an e io o middle o he abdomen wi h whi e
lecks; b own mo ling on oo dense;
Colou a ion in p ese a i e. Colou a ion da ke han
in li e; ma king on head and do sum isible; mandibula
egion da k b own; do sal aspec o dis al hal o o e-
a m and hand da k b own; knee da k b own; c ossba s on
high and ibia dis inc ly isible; one b oad c oss ba on
each high and wo on each ibia; disc on digi s pale c eam
colou ed wi h b own mo ling; da k b own pa ch a ound
en ; chin and h oa b own; ches , abdomen and en al
aspec o high pale c eam colou ed wi h b own lecks;
o ea m, ibia, a sus, and ee b own; yellow spo s on
g oin and high and whi e lecks on abdomen no isible.
Mo phological a ia ion. De ailed mo phome ic a i-
a ions a e gi en in Table S12. Addi ionally, yellow spo s
on g oin and highs a y among indi iduals (Fig. 34).
Fo example, WII-ADA3216 has yellow pa ches on base
o lowe a m, en al aspec o high and ibia; dis inc
enla ged yellow pa ches on ou e la e al aspec o high;
yellow pa ches on g oin much enla ged unlike holo ype;
a ube cle on in e -uppe eyelid space. Do sal ma king on
head and do sum in WII-ADA3216 dis inc in p ese ed
condi ion and ba ely isible in WII-ADA3218, WII-
ADA3398 and WII-ADA3399. B own mo ling on he
en al side in pa a ypes is no as dense as in holo ype.
Mo phological compa ison. Rao ches es nasu a sp.
no . di e s om i s congene s by i s acu e snou shape;
u he i di e s om R. ande soni by posi ion o nos il
equidis an be ween eye and snou ip ( s. nos il close
o snou ip han eye); i di e s om R. annandalii, R.
boulenge i sp. no ., and R. kempiae by p esence o en-
la ged yellow pa ches on g oin and high ( s. absen ); i
di e s om R. ba akensis sp. no . by head being longe
han wide ( s. head leng h less han o equal o wid h),
enla ged yellow pa ches p esen on g oin ( s. enla ged
b own pa ch p esen on g oin), by p esence o da k b own
hou glass shaped pa e n on do sum ( s. pai o conca e
da k b own s ipes p esen on do sum); i di e s om R.
cine ascens no . comb. by snou leng h being equal o
eye leng h and longe han in e -uppe eyelid wid h ( s.
snou leng h less han eye leng h and equal han in e -up-
pe eyelid wid h); i di e s om R. dulongensis by posi-
ion o nos il equidis an be ween eye and snou ip ( s.
nos il close o snou ip han eye), enla ged yellow spo s
p esen on g oin and high ( s. absen ); i di e s om R.
ga o by p esence o enla ged yellow pa ches on g oin
and high ( s. da k b own pa ches wi h o wi hou yellow
spo s p esen ); i di e s om R. hekouensis by head being
longe han wide ( s. head wide han long), snou being
longe han in e -uppe eyelid wid h ( s. snou leng h less
han o equal o in e -uppe eyelid wid h), nup ial pad
p esen only on i s inge ( s. nup ial pad p esen on i s
and second inge ); i di e s om R. hillisi and R. meng-
laensis by posi ion o nos il equidis an be ween eye and
snou ip ( s. nos il close o snou ip han eye), snou
leng h being equal o eye leng h ( s. snou leng h g ea e
han eye leng h); i di e s om R. huanglianshan by head
being longe han wide ( s. head leng h less han o equal
o wid h), by snou leng h being equal o eye leng h ( s.
snou leng h g ea e han eye leng h); i di e s om R.
jadoh by la ge body size in adul males, SVL 17.0–19.9
( s. SVL 13.6–14.0 mm); i di e s om R. jadoh and
R. jakoid snou acu e ( s. snou ounded), by posi ion o
nos il equidis an be ween eye and snou ip ( s. nos il
close o snou ip han eye), and enla ged yellow pa ches
p esen on g oin and high ( s. absen ); i di e s om R.
lawng laiensis sp. no . by head being longe han wide
( s. head leng h equal o o smalle han wid h), by snou
leng h being equal o eye leng h ( s. snou leng h less han
eye leng h); i di e s om R. leik ho by la ge body size
in adul males, SVL 17.0–19.9 mm ( s. SVL 15.7–15.8
mm), an hou glass shaped pa e n p esen on do sum ( s.
absen ); i di e s om R. longchuanensis by in e -uppe
eyelid wid h being less han o equal o eye leng h ( s.
in e -uppe eyelid wid h g ea e han eye leng h); i di -
e s om R. malipoensis by head being longe han wide
( s. head wide han long), by snou being longe han in-
e -uppe eyelid wid h ( s. snou leng h less han o equal
o in e -uppe eyelid wid h); i di e s om R. mawsyn a-
mensis sp. no . and R. shillongensis by head being longe
han wide ( s. head leng h less han o equal o wid h),
enla ged yellow pa ches p esen on g oin and high ( s.
absen ); i di e s om R. minda by p esence o yellow
pa ches on g oin and high ( s. enla ged black and whi e
pa ches p esen on g oin and high), an hou glass shaped
pa e n p esen on do sum( s. absen ); i di e s om R.
na puhensis sp. no . by head being longe han wide ( s.
head leng h less han o equal o wid h), by posi ion o
nos il equidis an be ween eye and snou ip ( s. nos il
close o snou ip han eye); i di e s om R. pa ulus
by high being longe han ibia ( s. high leng h smalle
han o equal o ibia leng h); i di e s om R. ezakhani
by snou leng h being equal o eye leng h ( s. snou leng h
g ea e han eye leng h), by head being longe han wide
( s. head wide han long); i di e s om R. y hus no .
comb. by head being longe han wide ( s. head leng h
equal o wid h), snou leng h being equal o eye leng h
Ve eb a e Zoology 75, 2025, 517–625 587
( s. snou leng h g ea e han eye leng h); i di e s om
R. yadongensis by head being longe han wide ( s. head
wide han long), in e -uppe eyelid wid h being less han
o equal o eye leng h ( s. in e -uppe eyelid wid h g ea -
e han eye leng h). A de ailed mo phological compa ison
is p o ided in Table 1.
Figu e 34. Rao ches es nasu a sp. no . A–D Holo ype (WII-ADA3388) in p ese a i e, A Do sal iew, B en al iew, C la e al
iew, D en al iew o igh hand. E, F Pa a ype (WII-ADA3216) in p ese a i e. G–J Holo ype in li e, G do sola e al iew,
H en al iew, I do sal iew, J g oin. K–O pa a ype (WII-ADA3216) in li e, K do sola e al iew, L do sal iew o head, M en al
iew, N g oin, O pos e io iew o highs. Scale ba = 10 mm. Pho os (G–O) by Raji N.V.
Bo uah B e al.: Re ision o bush ogs om no heas India
588
Phylogene ic ela ionship and gene ic di e gence.
Rao ches es nasu a sp. no . is sis e o an undesc ibed
species om Tibe and Adi hills, A unachal P adesh
(UFB 100, PP 1.0; Figs 2, 3). These wo lineages ha e a
shallow gene ic di e gence o 1.5–2.0% in he 16S gene.
The gene ic di e gence o R. nasu a sp. no . wi h o he
congene s included in his s udy a e 3.1–8.9% in he 16S,
9.1–19.2% in he cy b and 3.2–14.8% in he COI genes
(Table S7A–C).
E ymology. The speci ic epi he is de i ed om he La in
wo d nose “Nasu a”, in e e ence o he p ominen acu e
snou o his species.
Sugges ed common name. Poin y-nosed bush og.
Dis ibu ion and na u al his o y. Rao ches es nasu a sp.
no . is cu en ly dis ibu ed wi hin Namdapha Tige Re-
se e in Ranijheel, Bulbulia and Ho nbill on he no he n
bank o he Noa-Dihing Ri e , A unachal P adesh, India
be ween 600–1000 m a.s.l. ele a ion (Fig. 20A). Calling
agg ega ion o he species was eco ded be ween 18:00–
22:00 h s on ege a ion along s eams and o es ails.
Rao ches es dibangensis sp. no .
h ps://zoobank.o g/03F4255C-EBDB-4925-92A6-AF-
654C29C440
Figu e 35; Tables 1, 2, S12
Holo ype. An adul male (WII-ADA1677) collec ed
by BB on 7 Augus 2022 om Abango (28.09974°N,
95.89392°E, ele a ion 410 m a.s.l.), Lowe Dibang Val-
ley Dis ic , A unachal P adesh, India.
Pa a ype. An adul male (WII-ADA1671) collec ed
along wi h holo ype om he same locali y.
Diagnosis. A small sized Rao ches es, SVL 19.6–21.8
mm in adul males, emale unknown; head leng h equal
o wid h; snou ounded, i s leng h equal o o less han
eye leng h; ympanum indis inc ; small blun ube cles
on uppe eyelid; spinules on middle o do sum, abo e
sup a ympanic old, lank and spa sely p esen on limb;
wo indis inc c ossba s on high and ibia; do sum pale
b own wi h a pai o ain conca e s ipes, pos e io ends
o hese s ipes dis inc nea g oin; whi e ba p esen on
in e -uppe eyelid space in some indi iduals; small whi e
spo s isible on abdomen in li e.
Desc ip ion o he holo ype. Holo ype well p ese ed
excep o an incision on unde side o igh high; small
sized. Adul male wi h ocal sac, SVL 21.8 mm; head
leng h equal o wid h; snou ounded in do sal iew and
nea ly unca ed in la e al iew, i s leng h (2.9 mm) equal
o eye leng h (2.9 mm); nos il obliquely o al in shape,
la e ally posi ioned and oblique, close o snou ip han
eye (NS/EN = 0.69); na ial egion aised; can hus os-
alis smoo h, obliqued; lo eal egion conca e; in e na i-
al dis ance equal o in e -uppe eyelid wid h and g ea e
han uppe eyelid wid h (UEW/IN = 0.68); ympanum in-
dis inc , ound, less han one hi d o eye leng h (HTYD/
EL = 0.28); sup a ympanic old dis inc ; ongue pos e-
io ly wide and deeply no ched; small la ube cles on
ongue; choanae o al; ome ine ee h absen ; a pai o sli
like openings on lowe jaw nea angle o jaw; symphysial
knob p esen on lowe jaw.
Habi us slende , do so- en ally la ened, less han
hal o snou - en leng h (AG/SVL = 0.44); o elimb
slende ; hand leng h g ea e han o ea m leng h (FAL/
HAL = 0.86); digi wi h ounded disc; ci cum-ma ginal
g oo e p esen on each disc; hi d inge longes , ela i e
leng h o inge s = I<II<IV<III; disc on hi d inge wide
han ympanic diame e (HTYD/TIIID = 0.73); palma
ube cles ba ely isible; suba icula ube cles ounded,
p oximal ones on hi d and ou h inge smalle han dis-
al ones and indis inc ; no webbing among he inge s;
inely g anula nup ial pad on i s inge ; hindlimb slen-
de ; high longe han ibia (TBL/TL = 0.94) and nea ly
hal snou - en leng h (TL/SVL = 0.48); ou h oe lon-
ges ; oe wi h ounded disc, disc wid h equal o hose o
inge s; ela i e leng h o oes = I<II<III=V<IV; no de -
mal inge along i h oe; webbing sligh , no eaching
he second suba icula ube cle on ou h oe; small inne
me a a sal ube cle p esen , ou e one absen ; suba icula
ube cles ound, p oximal suba icula ube cles on oe
III–V ba ely isible; no supe nume a y ube cles.
Skin on do sal aspec o head smoo h; snou on do sal
aspec and uppe eyelids shag een; indis inc ube cles on
uppe eyelid; a leas wo blun ube cles on mandibula
egion, behind angle o jaw; spinules on do sola e al as-
pec o unk and middle o do sum, s a ing abo e pos e-
io pa o sup a ympanic old, ew sca e ed on an e io
and pos e io pa o do sum; lank, o elimb and hind-
limb smoo h on do sal aspec ; h oa and ches smoo h;
gula skin loose; abdomen g anula ; high g anula bu
no as dis inc as on abdomen; ibia smoo h on en al
aspec .
Colou a ion in p ese a i e. Do sal aspec o head
b own; pale whi e ba on in e -uppe eyelid space ol-
lowed by a sligh ly da ke in e ed iangula ma k on
head; uppe eyelid da k g ey; lo eal egion da ke han
do sal aspec o head; sup a ympanic old da k b own;
do sum da ke han head, pale on lank; o elimb pale
b own on do sal aspec ; a sligh da k b own c ossba on
o ea m; a simila sho ba on hand below ou h inge ;
hindlimbs pale b own do sally, sligh ly da ke owa ds
knee; single da k b own c oss ba on each high; a single
c oss ba on le ibia and wo on igh ibia; wo c ossba s
on each a sus, ain ; a sho da k b own ba on oo be-
low i h oe; a ea a ound en da ke ; chin, h oa , ches ,
abdomen, en al aspec o limb, hand, and ee c eamy
whi e wi h b own mo ling, chin owa ds he edge o low-
e jaw hea ily mo led.
Colou a ion in li e. Head and body and limbs on op pale
g eyish b own; a b oken pale whi e ba on in e -uppe
Ve eb a e Zoology 75, 2025, 517–625 589
eyelid space; i is whi ish wi h b own spo s and i egula
wa y b own lines, golden mo ling on uppe hal o i ;
a pai o ba ely isible, ain b own conca e s ipes on
do sum; single ain b own c ossba on o ea m; simila
wo c ossba s on each high and ibia and single c oss ba
on a sus owa ds dis al end; disc on inne wo inge s and
inne wo oes yellow; en al aspec o head, abdomen
and limbs pale lesh colou ed wi h b own mo ling; ocal
sac pale yellow and pale semi- anspa en ,; whi e spo s
o i egula size on g anules o abdomen; i egula whi e
spo s on high, ibia, a sus, and o ea m.
Mo phological a ia ion. Mo phome ic de ails a e gi -
en in Table S12. The pa a ype does no ha e a whi e ba
on in e -uppe eyelid space; c ossba s on limbs no isi-
ble in p ese ed condi ion unlike holo ype; lowe jaw and
ches hea ily mo led wi h b own unlike holo ype.
Mo phological compa ison. Rao ches es dibangensis
sp. no . di e s om R. ande soni by head leng h being
equal o wid h ( s. head wide han leng h); R. annan-
dalii by p esence o dense spinules on do sum ( s. skin
on do sum smoo h); i di e s om R. ba akensis sp. no .
by absence o pa e n on g oin ( s. enla ged b own pa ch
on g oin), wo indis inc c ossba s p esen on high and
ibia ( s. h ee na ow bands p esen on high and ib-
ia), conca e s ipes on do sum indis inc o no isible
( s. conca e s ipes on do sum dis inc ); i di e s om
Figu e 35. Rao ches es dibangensis sp. no . A Do sal and B en al iew o holo ype (WII-ADA1677) in p ese ed condi ion.
C–E pa a ype (WII-ADA1671) in p ese ed condi ion, C do sal iew, D en al iew, E la e al iew o head. F Do sola e al and
G en al iew o holo ype in li e. Scale ba = 10 mm.
Bo uah B e al.: Re ision o bush ogs om no heas India
596
do sum); i di e s om R. hillisi by la ge body size in
males, SVL 17.9–20.6 mm ( s. SVL 15.9–17.7 mm in
males), by head being wide han long ( s. head longe
han wide), by snou leng h being equal o o sligh ly less
han eye leng h ( s. snou leng h longe han eye leng h),
ympanum indis inc ( s. ympanum dis inc ), absence o
da k s ipes on do sum ( s. a da k “)(“ shaped ma king on
do sum); i di e s om R. huanglianshan by snou leng h
equal o o sligh ly less han eye leng h ( s. snou leng h
g ea e han eye leng h), ympanum indis inc ( s. ym-
panum dis inc ), absence o da k ma king on in e -uppe
eyelid space ( s. a b own iangula ma king on in e -up-
pe eyelid space), and by absence o “X” ma k on do sum
( s. a da k “X” ma k on do sum); i di e s om R. jadoh
and R. jakoid by snou leng h equal o in e -uppe eye-
lid wid h ( s. snou leng h g ea e han in e -uppe eyelid
wid h), by nos il being equidis an be ween snou ip and
eye ( s. nos il close o snou ip han eye), and high
leng h being g ea e han ibia leng h ( s. high leng h
less han ibia leng h); i di e s om R. kempiae by snou
leng h being equal o in e uppe eyelid wid h ( s. snou
leng h g ea e han in e -uppe eyelid wid h), in e -uppe
eyelid wid h being equal o eye leng h ( s. in e uppe
eyelid wid h smalle han eye leng h); i di e s om R.
lawng laiensis sp. no . by in e uppe eyelid wid h being
equal o eye leng h ( s. in e uppe eyelid wid h g ea e
han eye leng h), in e na ial dis ance being g ea e han
uppe eyelid wid h ( s. in e na ial dis ance equal o uppe
eyelid wid h); i di e s om R. leik ho by la ge body size
in adul males, SVL 17.9–20.6 mm ( s. SVL 15.7–15.8
mm); i di e s om R. longchuanensis by smalle body
size in males, SVL 17.9–20.6 mm ( s. SVL 21.4–23.9
mm in males), by in e uppe eyelid wid h being equal o
eye leng h ( s. in e uppe eyelid wid h g ea e han eye
leng h), ympanum indis inc ( s. dis inc ); i di e s R.
mawsyn amensis sp. no . by do sal skin smoo h wi h ew
sca e ed spinules on i ( s. dense spinules on do sum),
absence o “)-(“ on do sum ( s. p esen ), ma king absen
on in e -uppe eyelid space absen ( s. p esen ); i di e s
om R. menglaensis by head being wide han long ( s.
head longe han wide), snou leng h being equal o o
sligh ly less han eye leng h ( s. snou leng h longe han
eye leng h),, no “X” shaped ma king on do sum ( s. a
“X” ma king on do sum p esen ), ympanum indis inc
( s. ympanum dis inc ); i di e s om R. minda by ab-
sence o black and whi e pa ches on g oin ( s. enla ged
black and whi e pa ches p esen on g oin), by absence
o whi e pa ches on la e al aspec o highs ( s. enla ged
whi e pa ches on la e al aspec o highs); i di e s om
R. na puhensis sp. no . by p esence o b own pa ch wi h
whi e c own shaped pa ch below i on g oin ( s. g oin
yellowish wi hou da k o ligh pa ches), and by posi ion
o nos il equidis an om eye and snou ip ( s. nos il
close o snou ip han eye); i di e s om R. nasu a sp.
no . by ounded snou ( s. acu e snou ), by head being
wide han long ( s. head longe han i s wid h), and
snou leng h being equal o in e -uppe eyelid wid h ( s.
snou leng h being g ea e han in e uppe eyelid wid h);
i di e s om R. o ien alis sp. no . by sca e ed ube cles
on do sum ( s. dense spinules on do sum), da ke s ipes
on do sum ba ely isible o absen ( s. conca e do sal
s ipes dis inc ); i di e s om R. pa ulus by absence
o in e -uppe eyelid ba and s ipes on do sum ( s. ba
o iangula ma k be ween uppe eyelids and “)(“ ma k
on do sum p esen ), ympanum indis inc ( s. ympanum
dis inc ); i di e s om R. ezhakhani by absence o con-
ca e s ipes on do sum ( s. a “)(“ o “)-(“ ma k on do -
sum), by p esence o single c ossband on high and ibia
( s. h ee c ossbands on high and ibia); i di e s om
R. shillongensis by absence o s ipe o blo ch on do sum
( s. do sal colou a iable, dis inc o indis inc “)(“ ma k
o da k hou glass shaped ma k may be p esen ); i di e s
om R. y hus no . comb. by posi ion o nos il equi-
dis an om eye and snou ip ( s. nos il close o snou
ip han eye), by snou leng h being equal o eye leng h
( s. snou leng h longe han eye leng h), absence o “)(“
ma king on do sum ( s. a “)(“ ma k on do sum); i di e s
om R. yadongensis by in e -uppe eyelid wid h being
sligh ly less han o equal o eye leng h ( s. in e -uppe
eyelid wid h g ea e han eye leng h), absence o iangu-
la ma k be ween uppe eyelids ( s. dis inc da k iangu-
la ma king be ween uppe eyelids), ympanum indis inc
( s. ympanum dis inc ). De ailed mo phological com-
pa ison wi h o he congene ic species included in his
s udy is p o ided in Table 1.
Acous ics. The calls o R. monoli hus sp. no . we e e-
co ded in Pumdunlong, Manipu on 21 July 2022 a 18:50
h s a an ambien empe a u e o 21.6°C. The call de-
sc ip ion is based on 70 calls om wo indi iduals (WII-
ADA1634 and WII-ADA1635). The calls a e single ype,
non-pulsa ile (Fig. 31) and a e emi ed in g oups (5–13
calls pe call g oup). In each call g oup, he ampli ude
peaks a e he ini ial 1–3 calls. The mean call du a ion
is 22.01 ± 16.07 ms (12–29 ms) wi h a call ise ime o
1 ms and call all ime o 18.52 ± 4.62 (11–28 ms). The
in e call in e al is 164.78 ± 25.14 ms (118–285 ms).
The mean dominan equency is 3470.51 ± 59.77 Hz
(3316.1–3531.4 Hz). A de ailed compa ison o ad e -
isemen calls wi h hose o he congene s is p esen ed
in Table 2.
Phylogene ic ela ionship and gene ic di e gence.
Rao ches es monoli hus sp. no . is sis e o R. shillon-
gensis and R. jadoh wi h s ong nodal suppo (UFB 100,
PP 1.0; Figs 2, 3). The gene ic di e gence wi h o he
congene s included in his s udy is 3.9–8.6% in he 16S,
11.8–18.2% in he cy b and 6.5–13.8% in he COI genes
(Table S7A–C).
E ymology. This species is named a e he “monoli hs”
loca ed in Willong Khullen illage in Manipu S a e, 4
km sou hwes om he ype locali y o his species.
Sugges ed common name. Willong-Khullen bush og.
Dis ibu ion and na u al his o y. Rao ches es mono-
li hus sp. no . is known only om i s ype locali y and
a ound Willong Khullen Village in Manipu (Fig. 19B).
Calling indi iduals we e eco ded be ween 18:00–20:00
Ve eb a e Zoology 75, 2025, 517–625 597
h s on sh ubs and small woody ege a ion a pe ch heigh s
o ~2 m. Calling agg ega ions we e eco ded in second-
a y o es s a a ious deg ada ion s a e (Fig. 33D). We ob-
se ed b eeding agg ega ion o Hyla annec ans (Je don,
1870) in he same habi a in empo a y wa e puddles.
Rao ches es khonoma sp. no .
h ps://zoobank.o g/F8DFBFD3-C19D-4B52-AC34-
BA08D2237356
Figu e 38; Tables 1, 2, S12
Holo ype. An adul male (WII-ADA897) collec ed by BB
and AD on 8 Augus 2021 om 2.7 km wes o Khono-
ma illage (25.64145°N, 93.99801°E, ele a ion 2040 m
a.s.l.), Kohima Dis ic , Nagaland, India.
Pa a ypes. Th ee adul males (WII-ADA894–896) col-
lec ed om he same locali y as holo ype.
Diagnosis. Small sized Rao ches es, SVL 16.9–22.0
mm in adul males; head wid h equal o o sligh ly wid-
e han leng h (HL/HW = 0.94–1.0); snou ounded, i s
leng h equal o o sligh ly less han eye leng h (SL/EL =
0.93–1.0); snou leng h less han o equal o in e -uppe
eyelid wid h (0.94–1.04); nos il equidis an o snou ip
and eye; in e -uppe eyelid wid h equal o eye leng h; in-
e na ial dis ance g ea e han uppe eyelid wid h (UEW/
IN = 0.75–0.91); do sal colou pale yellowish b own wi h
golden inge; sligh ly da ke b own and whi e pa ches
on g oin and inne la e al aspec o high; single b oad,
b own c ossba on high and ibia.
Desc ip ion o he holo ype. Adul male wi h ocal sac,
SVL 22 mm; head leng h equals o wid h; snou ounded
in bo h do sal and la e al iews, i s leng h sligh ly less
han eye leng h (SL/EL = 0.94) and in e -uppe eyelid
wid h (SL/IUE = 0.94); snou sha ply sloped an e io ly
om in e na ial egion; lo eal egion sligh ly conca e;
can hus os alis ounded and oblique; in e o bi al space
la ; nos ils o al and obliquely o ien ed, equidis an be-
ween snou ip and eye; eye mode a e in size, p o uding
(EL/HL = 3.4); pineal ocellus absen ; ympanum indis-
inc , ounded; sup a ympanic old dis inc ; in e na ial
dis ance smalle han in e uppe eyelid wid h (IN/IUE
= 0.71) and equal o uppe eyelid wid h; ongue pos e i-
o ly b oad and bilobed; ome ine ee h absen ; choanae
ounded; pai o sli like openings on lowe jaw; sym-
physial knob on lowe jaw; maxilla y ee h p esen ; ans-
lucen ex e nal subgula ocal sac p esen .
Habi us slende (AG/SVL = 0.5), do so en ally la -
ened; o elimbs slende , o ea m leng h nea ly equal
o hand leng h (FAL/HAL = 0.97); hi d inge longes ,
ela i e leng h o inge s = I<II<IV<III; all digi s wi h
ounded discs; la ges disc on hi d and ou h inge , disc
wid h o he inge IV g ea e han ympanic diame e ;
ci cum-ma ginal g oo e p esen on each disc p esen ;
palma ube cles ba ely isible on le hand; suba icu-
la ube cles la ge and ound, p oximal ones on hi d and
ou h inge small and indis inc ; suba icula ube cles
on inge = 1:1:2:2; on la g anules p esen on palm, a
la ge ine g anula nup ial pad on i s inge s.
Hindlimbs slende ; high sligh ly longe han ibia
(TBL/TL = 0.93) and hal o snou - en leng h (TL/SVL
= 0.5); ibia longe han oo leng h (FOL/TBL = 0.86);
ela i e leng h o oes = I<II<V<III<IV; oe wi h ounded
disc, as wide as on inge s; ci cum-ma ginal g oo e p es-
en on each oe; inne me a a sal ube cle p esen , ou e
one absen ; suba icula ube cles dis inc , ound, p ox-
imal suba icula ube cles on ou e h ee oes smalle ;
suba icula ube cles on oes = 1:1:2:3:2; webbing sligh ,
no eaching he second suba icula ube cle o ou h
inge .
Skin on do sal aspec o head smoo h o shag een on
do sum; ew small ube cles sca e ed on uppe eyelid;
ew ube cles abo e sup a ympanic old; ube cles do -
sola e al aspec o do sum and on ibia spa sely p esen ;
ube cles pos e io o angle jaw ba ely isible; o elimb
smoo h; gula egion, ches g anula bu no as dis inc on
abdomen; abdomen g anula ; lank owa ds en ola e al
aspec g anula ; en al aspec o high g anula bu o
ibia smoo h.
Colou a ion in li e. Do sally head, back and limbs
pale-yellowish b own wi h golden inge; a e y ain ba
on in e -uppe eyelid space; ain c ossba s on high and
ibia; en ally, an e io pa o head pale g eyish b own,
pos e io ly pale yellowish b own wi h dense whi e mo -
ling on and spa se b own mo ling; abdomen whi ish
wi h b own mo ling, ew black i egula spo s on an e i-
o pa o he abdomen; o elimb en ally pale yellowish
wi h b own mo ling and b own spo s on o ea ms; highs
en ally pale yellow b own wi h b own mo ling and i -
egula whi e spo s; la ge c eamy whi e pa ches on ibia;
la ge b own pa ch on g oin and sligh ly smalle whi e
pa ch on ei he side o i ; ano he whi e pa ch on inne
la e al aspec o high owa ds base.
Colou a ion in p ese a i e. Do sally head and back
g ey, uppe eyelid sligh ly da ke , an indis inc sligh ly
da k ba be ween uppe eyelids; lo eal egion da ke , wo
indis inc da k s eaks on uppe jaw adia ing om he
eye; o elimbs and hindlimbs do sally g eyish b own,
sligh ly da ke b own c oss ba on o ea m, high, ibia
and a sus; lank pale g eyish b own; on en al aspec ,
head c eam colou ed wi h b own mo ling, dense on an-
e io pa o lowe jaw; abdomen c eam colou ed wi h
small indis inc whi e spo s and b own mo ling, i egu-
la small b own pa ches on abdomen and owa ds lank;
o elimb en ally wi h dense b own mo ling; hindlimb
c eam colou ed wi h dense b own mo ling, b own c oss-
ba s on high and ibia isible, pale-whi e spo s on high
and ibia; no ma king on ou e la e al aspec o high and
a ound en .
Mo phological a ia ion. De ailed mo phological a i-
a ions a e p o ided in Table S12. In addi ion, black spo s
on he abdomen a e absen in he pa a ypes; a pai o in-
Bo uah B e al.: Re ision o bush ogs om no heas India
598
dis inc conca e and sligh ly da k s ipes on he do sum
isible on WII-ADA896 in p ese ed condi ion.
Mo phological compa ison. Rao ches es khonoma sp.
no . di e s om R. ande soni and R. ezakhani by in-
e -uppe eyelid wid h being equal o eye leng h ( s. in-
e -uppe eyelid wid h smalle han eye leng h); i di e s
om R. annandalii by snou leng h being equal o o less
han in e -uppe eyelid wid h ( s. snou leng h g ea e
han in e -uppe eyelid wid h); i di e s om R. ba a-
kensis sp. no . by absence o bony p ojec ion on hume -
us ( s. bony p ojec ion on hume us p esen ), in e -uppe
Figu e 38. Rao ches es khonoma sp. no . A–C and F, G holo ype (WII-ADA897) in p ese ed condi ion. A Do sal iew, B en al
iew, C la e al iew o head, F en al iew o le hand, G en al iew o le ee . D Do sal and E en al iew o pa a ype (WII-
ADA896) in p ese ed condi ion. H–L Holo ype in li e, H do sola e al iew, I en al iew, J g oin, K pos e io iew o high,
L g anules on abdomen. Scale ba = 10mm.
Ve eb a e Zoology 75, 2025, 517–625 599
eyelid wid h being equal o eye leng h ( s. in e -uppe
eyelid wid h smalle han eye leng h); i di e s om R.
boulenge i sp. no . by in e -uppe eyelid wid h being
equal o eye leng h ( s. in e -uppe eyelid wid h smalle
han eye leng h), by absence o spinules on do sum, ew
blun sca e ed ube cles p esen ( s. dense spinules on
do sum), and by absence o conca e s ipes on do sum,
may be ba ely isible in p ese ed condi ion ( s. dis inc
pai o da k b own conca e s ipes p esen on do sum); i
di e s om R. cine ascens no . comb. by p esence o a
single b oad b own c ossba on high and ibia ( s. h ee
c ossba s on high and ibia); i di e s om R. dibangen-
sis sp. no . and R. na puhensis sp. no . by in e -uppe
eyelid wid h being equal o eye leng h ( s. in e -uppe
eyelid wid h less han eye leng h), by nos il being equi-
dis an be ween eye and snou ip ( s. nos il close o
snou ip han eye); i di e s om R. dulongensis by head
leng h being equal o o less han head wid h ( s. head
longe han wide), by nos il being equidis an be ween
eye and snou ip ( s. nos il close o ip o snou ); i di -
e s om R. ga o by absence o spinules on do sum ( s.
spinules p esen on do sum); i di e s om R. hekouensis
by p esence o nup ial pad only on i s inge ( s. nup ial
pad p esen on i s and second inge s), p esence o ew
sca e ed ube cles on do sum ( s. nume ous ube cles on
do sum); i di e s om R. hillisi and R. menglaensis by
head leng h being less han o equal o head wid h ( s.
head longe han wide), snou leng h being equal o o
less han eye leng h ( s. snou leng h g ea e han eye
leng h); i di e s om R. huanglianshan by snou leng h
being equal o o less han eye leng h ( s. snou leng h
longe han eye leng h), nos il being equidis an be ween
eye and snou ip ( s. nos il close o snou ip han eye);
i di e s om R. jadoh by la ge body size in adul males,
SVL 16.9–22.0 mm ( s. SVL 13.6–14.0), and in e -uppe
eyelid wid h being equal o eye leng h ( s. in e -uppe
eyelid wid h g ea e han eye leng h); i di e s om R.
jakoid by snou leng h being less han o equal o eye
leng h ( s. snou leng h g ea e han eye leng h), nos il
being equidis an be ween snou ip and eye ( s. nos il
close o snou ip han eye); i di e s om R. kempiae
by in e -uppe eyelid wid h being equal o eye leng h
( s. in e -uppe eyelid wid h smalle han eye leng h), by
snou leng h being smalle han o equal o in e -uppe
eyelid wid h ( s. snou leng h g ea e han in e -uppe
eyelid wid h); i di e s om R. lawng laiensis sp. no .
by in e -uppe eyelid wid h being equal o eye leng h ( s.
in e -uppe eyelid wid h less han eye leng h), in e na ial
dis ance being g ea e han uppe eyelid wid h ( s. in-
e na ial dis ance equal o uppe eyelid wid h); i di e s
om R. leik ho by la ge body size in adul males, SVL
16.9–22.0 mm ( s. SVL 15.7–15.8 mm), nos ils being
equidis an be ween eye and snou ip ( s. nos ils close
o snou ip), p esence o sca e ed ube cles on do sum
( s. nume ous spinules on do sum), b own and whi e
pa ches p esen on g oin ( s. a sho b own ba p esen on
he g oin); i di e s om R. longchuanensis by in e -up-
pe eyelid wid h being equal o eye leng h ( s. in e -uppe
eyelid wid h g ea e han eye leng h); i di e s om R.
malipoensis by nos il being equidis an be ween eye and
snou ip ( s. nos il close o snou ip han eye); di e s
om R. mawsyn amensis sp. no . by absence o spinules
on head and do sum, ew blun sca e ed ube cles p esen
( s. dense spinules on op o head and do sum), by ab-
sence o conca e s ipes on do sum, may be ba ely isi-
ble in p ese ed condi ion ( s. dis inc pai o da k b own
conca e s ipes p esen on do sum); i di e s om R.
minda by p esence o b own and whi e pa ches on g oin
( s. enla ged black and whi e pa ches p esen on g oin),
by p esence o sca e ed ube cles on do sum ( s. dense
spinules on do sum); i di e s om R. nasu a sp. no .
by head leng h being less han o equal o i s wid h ( s.
head longe han wide), ounded snou ( s. snou acu e),
in e -uppe eyelid wid h being g ea e han o equal o
snou leng h ( s. snou leng h g ea e han in e -uppe
eyelid wid h), p esence o sligh ly da k b own and whi e
pa ch on g oin and high ( s. b igh yellow pa ches on
g oin and high); i di e s om R. o ien alis sp. no . by
p esence a ew blun sca e ed ube cles on do sum ( s.
spinules on do sum), a single b oad c oss ba isible on
high and ibia ( s. h ee o ou na ow c ossba s on high
and ibia); i di e s om R. pa ulus by high leng h be-
ing g ea e han ibia leng h ( s. high leng h less han o
equal o ibia leng h); i di e s om R. shillongensis by
absence o “)(“ ma k on do sum ( s. a da k b own “)(“
ma k p esen on do sum); i di e s om R. y hus no .
comb. by snou leng h being equal o o less han eye
leng h ( s. snou leng h longe han eye leng h); i di e s
om R. yadongensis by in e -uppe eyelid wid h being
equal o eye leng h ( s. in e -uppe eyelid wid h g ea e
han eye leng h), by posi ion o nos ils equidis an be-
ween eye and snou ip ( s. nos ils close o snou ip
han eye). Mo phologically R. khonoma sp. no . close o
R. monoli hus sp. no . De ailed mo phological compa i-
son wi h o he congene ic species is p o ided in Table 1.
Phylogene ic ela ionship and gene ic di e gence.
Rao ches es khonoma sp. no . is sis e o he clade which
includes R. dulongensis, R. hillisi and R. yadongensis
wi h weak suppo (UFB 71, PP <50; Figs 2, 3). The ge-
ne ic di e gence o R. khonoma sp. no . wi h he con-
gene s included in his s udy a e 3.1–6.7% in he 16S,
11.2–17.0% in he cy b and 7.8–14.3% in he COI genes
(Table S7A–C).
E ymology. The speci ic epi he is a oponym de i ed
om he name o he illage “Khonoma” in Nagaland
S a e whe e he ype se ies was collec ed.
Sugges ed common name. Khonoma bush og.
Dis ibu ion and na u al his o y. Rao ches es khono-
ma sp. no . is cu en ly known om i s ype locali y
Khonoma and Dzuleke illage in he Kohima Dis ic o
Nagaland om an ele a ion o 2000–2040 m a.s.l. (Fig.
19B). This species has also been eco ded om low ele-
a ion a eas a Khonoma Na u e Conse a ion and T a-
gopan Sanc ua y (KNCTS). Indi iduals o he species
we e mos commonly ound along he sub opical o es
ma gins and hilly e ace c oplands (Fig. 33E). G a id
Bo uah B e al.: Re ision o bush ogs om no heas India
600
emales we e pho og aphed in June and calling males
we e eco ded om May o Sep embe . We obse ed
males calling on sh ubs a a pe ch heigh o app oxima e-
lyone me e abo e g ound le el. Many indi iduals we e
obse ed calling om Eupa o ium hicke s. The a ea a
he ype locali y was mos ly co e ed by jhum and e ace
cul i a ion.
Rao ches es eaglenes ensis sp. no .
h ps://zoobank.o g/7035E94D-EA2B-4018-8D95-46EA5D-
A9309C
Figu e 39; Tables 1, 2, S12
Holo ype. Adul male (WII-ADA1619) collec ed by
BB, KB and DSG on 7 July 2022 nea Bompu camp
(27.06641°N, 92.40599°E, ele a ion 1970 m a.s.l.),
Eaglenes Wildli e Sanc ua y, Wes Kameng Dis ic ,
A unachal P adesh, India.
Pa a ypes. Eigh adul males (WII-ADA1612–WII-
ADA1618, WII-ADA1626) collec ed om he same lo-
cali y and same da e as he holo ype by BB, KB and DSG.
Re e ed ma e ial. Th ee adul males (WII-ADA1556–
WII-ADA1558) collec ed by BB and KB on 30 June
2022 om Glow Ba i (27.17112°N, 92.46007°E, el-
e a ion 2160 m a.s.l.), app oxima ely 5 km no h o
Eaglenes Wildli e Sanc ua y, Wes Kameng Dis ic ,
A unachal P adesh; wo adul emales (WII-ADA2513
and WII-ADA2514) and one adul male (WII-ADA2518)
collec ed by KB on 23 Augus 2022 nea Bompu camp
(27.07193°N, 92.4005°E, ele a ion 2000 m a.s.l.), Ea-
glenes Wildli e Sanc ua y, Wes Kameng Dis ic ,
A unachal P adesh.
Diagnosis. Small sized Rao ches es, SVL 18.8–23.6 mm
in adul males and SVL 21.9–22.8 mm in adul emales;
head leng h equal o wid h o sligh ly less han wid h in
males (HL/HW = 0.93–1.01) and head longe han wide
in emales (HL/HW = 1.07); ome ine ee h absen ; snou
ounded o sub-o al, i s leng h sligh ly less han o equal
o eye leng h (SL/EL = 0.85–1.0); nup ial pad p esen on
i s inge ; blun ube cles on op o head, uppe eyelid,
do sum and do sal su ace o limb; pai o oli e b own,
conca e s ipes on do sum; a b oad b own c ossba on
o ea m; h ee c ossba s on high and ibia.
Desc ip ion o he holo ype. Holo ype is well p ese ed
excep o an incision on unde side o igh high. Snou -
en leng h 22.1 mm; head sligh ly wide han long; de-
p ession on do sal aspec o snou be ween in e nasal
egion and snou ip; snou ounded in do sal iew and
nea ly acu e in la e al iew, sligh ly p o uding beyond
lowe jaw, blun ube cula p ojec ion on snou ip; i s
leng h less han eye leng h (SL/EL = 0.9); can hus os a-
lis ounded, obliqued; lo eal egion conca e; nos il o al,
la e ally posi ioned and obliquely o ien ed; nos ils equal-
ly posi ioned be ween eye and snou ip; eye mode a e in
size, leng h less han hal o head leng h (EL/HL = 0.41);
ympanum ound, dis inc , nea ly one hi d o he eye
leng h (HTYD/EL = 0.29); sup a ympanic old dis inc ;
in e nasal dis ance less han in e -uppe eyelid wid h (IN/
IUE = 0.85) and g ea e han uppe eyelid wid h (UEW/
IN = 0.7); choanae o al; ome ine ee h absen ; ongue
pos e io ly no ched and pos e io lobe on igh side sho -
e han lobe on le side; pai o sli like openings p esen
on lowe jaw; unk do so- en ally la ened, slende ,
hal o snou - en leng h (AG/SVL = 0.51).
Fo elimbs slende , o ea m sho e han hand leng h
(FAL/HAL = 0.72); ela i e leng h inge s = I<II<IV<III;
inge wi h ounded disc; ci cum-ma ginal g oo e on disc
p esen ; disc o he inge s II–IV wide han ympanic di-
ame e ; webbing absen ; suba icula ube cles ounded
and enla ged excep he p oximal suba icula ube cle on
hi d and ou h inge which smalle and indis inc ; sub-
a icula ube cles on inge s I:II:III:IV = 1:1:2:2; palma
ube cles no isible; inge wi hou la e al de mal inge;
ine g anula nup ial pad p esen on base o i s inge
co e ing inne la e al and do sal su ace, supe nume a y
ube cles absen .
Hindlimbs slende , high leng h hal o he snou -
en leng h (TL/SVL = 0.5); ela i e leng h oes =
I<II<III<V<IV; ibia leng h equal o high leng h and lon-
ge han oo leng h (FOL/TBL = 0.84); oe wi h ounded
disc; disc wid h equal o hose o inge s; ci cum-ma ginal
g oo e p esen ; suba icula ube cles ounded, p oximal
suba icula ube cles on oe III–V smalle and indis inc ;
indis inc inne me a a sal ube cle p esen ; ou e me a-
a sal ube cle absen ; supe nume a y ube cles absen ;
no de mal inge along oe V; webbing sligh , eaching
below second suba icula ube cle o ou h oe.
Smoo h skin on do sal aspec o snou , uppe eyelids
and head; h ee ube cles in a longi udinal s aigh line on
middle o head; indis inc ube cles on uppe eyelid; ew
p ominen ube cles on mandibula egion behind angle
o jaw; ube cles sca e ed do sal o sup a ympanic olds,
on do sum and sp eading o lank; ube cles absen on
pos e io pa o do sum; o elimb and hindlimb smoo h
do sally; ube cles on limb no isible as in li e condi ion;
h oa , ches , en al aspec o o elimb and ibia smoo h;
abdomen g anula , indis inc ; g anules on high ba ely
isible.
Colou a ion in li e. Do sal su ace o head, do sum and
limbs pale eddish b own; uppe eyelids pale g ey; la -
e al aspec o head pale-g eyish wi h eddish inge; i is
da k b own, speckled wi h golden, hea ily speckled do -
sally; in e ed iangula pa ch on do sal su ace o head
co e ing in e -uppe eyelid space and pos e io pa o
head; a b oad “X” shaped, oli e-b own colou ed pa e n
on do sum; indis inc c oss ba on o ea ms; simila ba s
on do sal su ace o hand and ou e wo inge s; h ee
b oad c ossba s on high and ibia; simila ba s on do sal
su ace o oo and ou e h ee oes; disc on inne wo
inge s and inne h ee oes yellow, es o he discs ed-
dish wi h b own mo ling; on en um, head, abdomen
and limbs lesh-colou ed wi h b own mo ling; enla ged
Ve eb a e Zoology 75, 2025, 517–625 601
whi e blo ches on ches and abdomen; la e al side o high
pale- eddish.
Colou a ion in p ese a i e. Do sal su ace o snou
and la e al aspec o head pale b own; sligh ly da k b own
s ipe on lo eal egion; uppe eyelids da k g ey; la ge da k
b own pa ch on head; da k b own spo s on uppe jaw, be-
low eyes o mandibula egion; pos e io pa o head o
an e io pa o do sum pale yellowish b own, pos e io ly
do sum sligh ly da ke ; a “X” shaped da k b own ma k on
do sum, i s an e io ends s a s om pos e io co ne o
uppe eyelid, pos e io ends b oade and di ec ed owa ds
Figu e 39. Rao ches es eaglenes ensis sp. no . A Do sal and B en al iew o holo ype (WII-ADA1619) in p ese ed condi ion.
C Do sal iew o WII-ADA2514 in p ese ed condi ion. D–G Holo ype in li e, D do sola e al iew, E en al iew, F g oin,
G pos e io iew o high. H WII-ADA1556 in li e; I pa a ype (WII-ADA1615) in li e; J, K wo uncollec ed males om Bompu,
Eaglenes Wildli e Sanc ua y. Scale ba = 10mm.
Bo uah B e al.: Re ision o bush ogs om no heas India
602
g oin; o elimbs pale yellowish b own, a b oad b own
c oss ba on o ea m; i egula sized, b own pa ches on
op o hands and wo ou e inge s; hindlimb pale yellow-
ish b own; h ee b oad c ossba s on each high and ibia;
simila ba s on a sus and oo and ou e h ee oes; an
enla ged da k b own pa ch on en a ea; h oa , ches , ab-
domen, en al aspec o elimb and hindlimb pale c eam
colou ed wi h b own mo ling.
Sexual dimo phism and mo phological a ia ion. In
males head leng h equals o wid h o sligh ly smalle han
wid h s. head longe han head wid h in emale; a subgu-
la ocal sac p esen in males; a pai o in e nal ocal sac
openings p esen on lowe jaw and a nup ial pad p esen
in males. Snou shape a ies among he indi iduals om
ounded o sub-o al; in some indi iduals, a whi e ba on
head connec ing he uppe eyelids was obse ed; i egu-
la shaped and sized yellow spo s p esen on head, back
and limbs p esen in some indi iduals. Mo phome ic
a ia ions a e p o ided in Table S12.
Mo phological compa ison. Rao ches es eaglenes ensis
sp. no . di e s om R. annandalii by snou leng h be-
ing smalle han o equal o eye leng h ( s. snou leng h
g ea e han o equal o eye leng h); i di e s om R. ba-
a kensis sp. no . by absence o ex a bone on hume us
( s. an ex a bone p esen on hume us); i di e s om
R. boulenge i sp. no ., R. mawsyn amensis sp. no ., and
R. o ien alis sp. no . by p esence o dis inc ube cles on
do sum ( s. dense spinules on do sum); i di e s om
R. cine ascens no . comb. by absence o enla ged da k
and ligh pa ches on g oin ( s. spo pa ially enci cled
by yellow inge on g oin); i di e s om R. dibangensis
sp. no . by p esence o dis inc bun ube cles on do sum
( s. dense spinules on do sum), p esence o h ee b oad
c ossba s on high and ibia ( s. wo indis inc c ossba s
on high and ibia); i di e s om R. dulongensis by p es-
ence o nup ial pad on i s inge ( s. nup ial pad absen );
i di e s om R. ga o by absence o da k b own pa ch-
es on g oin and high ( s. p esen ), do sum wi h blun
ube cles ( s. do sum wi h spinules); i di e s om R.
hekouensis, R. hillisi and R. malipoensis by la ge body
size, SVL 18.8–23.6 mm in adul males ( s. 16.1–17.5
mm in R. hekouensis, 15.9–17.7 mm in R. hillisi and
14.6–17.7 mm in R. malipoensis); i di e s om R. huan-
glianshan, R. menglaensis, and R. y hus no . comb. by
snou leng h being equal o o less han eye leng h ( s.
snou leng h longe han eye leng h); i di e s om R.
jadoh by la ge body size in adul males, SVL 18.8–23.6
mm ( s. SVL 13.6–14.0 mm), and by in e -uppe eyelid
wid h being less han o equal o eye leng h ( s. in e -up-
pe eyelid wid h g ea e han eye leng h); i di e s om
R. jakoid by snou leng h being less han o equal o eye
leng h ( s. snou leng h g ea e han eye leng h); i di e s
om R. khonoma sp. no . by p esence o h ee c ossba s
on high and ibia ( s. single c ossba on high and ibia),
p esence o dis inc “X” ma k o conca e s ipes on do -
sum ( s. s ipes on do sum ba ely isible), p esence o u-
be cles p onounced on head and do sum ( s. ube cles on
head and do sum compa a i ely small and mo e spa sely
dis ibu ed), and absence o b own and whi e pa ch on
g oin ( s. b own and whi e pa ches on g oin p esen ); i
di e s om R. lawng laiensis sp. no . by in e na ial dis-
ance being g ea e han uppe eyelid wid h ( s. in e na i-
al dis ance equal o uppe eyelid wid h); i di e s om R.
leik ho by la ge body size in adul males, SVL 18.8–23.6
mm ( s. SVL 15.7–15.8 mm); i di e s om R. longc-
huanensis, R. yadongensis by in e uppe eyelid wid h
being equal o eye leng h ( s. in e uppe eyelid wid h
g ea e han eye leng h); i di e s om R. menglaensis
Figu e 40. Compa a i e spec og ams and oscillog ams illus a ing ep esen a i e calls o wo new species o Rao ches es. A, B Os-
cillog ams, C spec og am, D powe spec um. The call ma ked wi h an a ow on op in A is expanded in B.
Ve eb a e Zoology 75, 2025, 517–625 603
by snou leng h being equal o o smalle han eye leng h
( s. snou leng h longe han eye leng h); i di e s om
R. minda by absence o black o whi e pa ches on g oin
( s. enla ged black and whi e pa ches p esen on g oin),
absence o whi e pa ches on he la e al aspec o high
( s. enla ged whi e pa ches on la e al aspec o high); i
di e s om R. monoli hus sp. no . by he dis inc “X”
ma k o conca e s ipes on do sum ( s. s ipes on do sum
ba ely isible), ube cles p onounced on head and do sum
( s. ube cles on head and do sum compa a i ely small
and mo e spa sely dis ibu ed), and absence o b own and
whi e pa ch on g oin ( s. ligh b own and whi e pa ch-
es on g oin p esen ); i di e s om R. nasu a sp. no .
by ounded o sub-o al shaped snou ( s. snou acu e);
i di e s om R. na puhensis sp. no . by p esence o
h ee c ossba s on high and ibia ( s. single c oss ba on
high and ibia); i di e s om R. pa ulus by p esence o
h ee c ossba s on highs ( s. single c oss ba on high);
i di e s om R. ezakhani by absence o da k pa e n on
g oin ( s. a sho b own s eak p esen on g oin); i di e s
om R. shillongensis by p esence o h ee b oad c oss-
ba s on high and ibia ( s. single b oad c ossba on high
and ibia). A de ailed mo phological compa ison wi h i s
congene ic species is p o ided in Table 1. Mo phological
cha ac e s o R. eaglenes ensis sp. no . and R. kempiae
a e simila bu hey ha e mino o e lap in mo phospace
(Fig. 6C). Rao ches es kempiae is es ic ed o sou h o
B ahmapu a Valley while R. eaglenes ensis sp. no . is
es ic ed o no h o B ahmapu a Valley.
Acous ics. The calls o R. eaglenes ensis sp. no . we e
eco ded in Glowba i, nea Eaglenes Wildli e Sanc ua y
on 30 June 2022 a 20:10 h s and a an ambien empe a-
u e o 21.4°C; in Bompu, Eaglenes Wildli e Sanc ua y
on 7 July 2022 a 20:00 h s and a ambien empe a u e o
21.1°C. The call desc ip ion is based on 60 calls includ-
ing wo indi iduals (WII-ADA1558 and WII-ADA1619).
The calls a e single ype, non-pulsa ile, emi ed in g oups
(2–4 calls pe g oup) in egula in e als (Fig. 40). The
mean call du a ion is 16.3 ± 1.74 ms (9–20 ms) wi h call
ise ime o 1 ms and call all ime o 15.45 ± 1.7 ms (12–
18 ms). The mean in e -call in e al is 236.56 ± 8.89 ms
(224–270 ms). The mean dominan equency is 3029.73
± 61.33 Hz (2971.6–3100.8 Hz). A de ailed compa ison
o ad e isemen calls wi h hose o congene s is sum-
ma ised in Table 2.
Phylogene ic ela ionships and gene ic di e gence.
Phylogene ically R. eaglenes ensis sp. no . was eco -
e ed as a basal lineage o a la ge clade con aining R. hil-
lisi g oup, R. kempiae g oup, R. ezakhani g oup, and R.
ga o g oup in ML analysis wi h weak suppo (UFB <50)
and in BI analysis i nes ed wi h hillisi g oup wi h weak
suppo (PP <0.5; Figs 2, 3). The gene ic di e gence o
R. eaglenes ensis sp. no . wi h he o he membe s o he
genus a ies be ween 2.9–8.0% in he 16S, 9.1–19.1% in
he cy b and 7.3–15.4% in he COI genes (Table S7A–C).
E ymology. The speci ic epi he is a oponym de i ed
om he name o he “Eaglenes Wildli e Sanc ua y” in
he S a e o A unachal P adesh whe e he ype specimens
we e collec ed.
Sugges ed common name. Eaglenes Bush og.
Dis ibu ion and na u al his o y. Rao ches es eagle-
nes ensis sp. no . is cu en ly known only om he Ea-
glenes Wildli e Sanc ua y and adjacen a eas such as
Glow Ba i wi hin an ele a ion ange o 1950–2200 m
a.s.l. (Figs 25B, 33F) We eco ded he indi iduals on
hill slopes along o es ails; in he mon hs o June and
July. Calling males we e obse ed on e ns, bamboo and
sh ubs a a heigh o 0.5–2 m abo e g ound. O he am-
phibian species eco ded a his locali y include Nasu i-
xalus sp., Xenoph ys sp. Amolops wangyali Mahony e
al., 2022, Amolops sp., Rhacopho us bu manus (Ande s-
son, 1939), Lep ob achium bompu, Du aph ynus hima-
layanus (Gün he , 1864), Nano ana liebigii (Gün he ,
1860) and Nano ana sp. A h eya (2006) epo ed se e al
mo phs o uniden i ied Rao ches es sp. (as Philau us sp.)
om Eaglenes Wildli e Sanc ua y and sugges ed ha i
had a leas ou di e en calls. A h eya (2006) did no
p o ide de ails o he loca ions and acous ic pa ame e s
o compa ison. Howe e , some o he pho og aphs o
Philau us sp. p o ided in A h eya (2006) alls wi hin he
mo phological a ia ion o R. eaglenes ensis sp. no .
Rao ches es a unachalensis sp. no .
h ps://zoobank.o g/CE8875FB-F369-4B4E-B5B8-A2EC-
99C4A6E5
Figu e 41; Tables 1, 2, S12
Ch esonymy.
Ixalus ube cula us — Annandale (1912): 7, 17
Ixalus annandalei — Annandale (1912): 7, 16 (spelling e o )
Rao ches es annandalii — Ohle e al. (2018): 282, 286
Holo ype. An adul male (WII-ADA1553) collec ed by
BB on 17 June 2022 om Rengging illage (28.14136°N,
95.27059°E, ele a ion 470 m a.s.l.), Eas Siang Dis ic ,
A unachal P adesh, India.
Pa a ype. An adul male (WII-ADA1554) collec ed
along wi h holo ype om he same locali y.
Re e ed ma e ial. Two adul emales (WII-ADA676,
WII-ADA677) and ou subadul males (WII-ADA491,
WII-ADA678, WII-ADA680 and WII-ADA681) col-
lec ed by BB and AD on 17 Oc obe 2019 om Po in
(27.34786°N, 93.84970°E, ele a ion 580 m a.s.l.), Low-
e Subansi i Dis ic , A unachal P adesh; one subadul
(WII-ADA1712) collec ed by BB on 29 Oc obe 2021
om Balek illage (28.06245°N, 95.27216°E, ele a ion
520 m a.s.l.), Eas Siang Dis ic , A unachal P adesh;
wo adul males (WII-ADA1308 and WII-ADA1327)
collec ed by BB and AD on 12 May 2022 nea Deban
(27.49662°N, 96.38949°E, ele a ion 380 m a.s.l.), Nam-
Bo uah B e al.: Re ision o bush ogs om no heas India
604
dapha Tige Rese e, A unachal P adesh; wo adul males
(WII-ADA1336, WII-ADA1345) collec ed by BB VJ and
AD on 13 May 2022 om Mo ijheel ail (27.48690°N,
96.33134°E, ele a ion 660 m a.s.l.), Namdapha Tige
Rese e, A unachal P adesh; wo adul males (WII-
ADA1551 and WII-ADA1552) collec ed by BB on 16
June 2022 om Ro ung (28.14113°N, 95.15619°E, el-
e a ion 390 m a.s.l.), Eas Siang Dis ic , A unachal
P adesh; wo adul males (WII-ADA1549 and WII-
ADA1550) collec ed by BB, KB and DSG on 29 June
2022 nea Sessa (27.10079°N, 92.53770°E, ele a ion
1030 m a.s.l.), Wes Kameng Dis ic , A unachal P adesh;
ou adul males (WII-ADA1704–WII-ADA1707) col-
lec ed by BB on 8 Augus 2022 nea Mehao nallah
(28.15436°N, 95.90596°E, ele a ion 930 m a.s.l.), Me-
hao Wildli e Sanc ua y, Lowe Dibang Valley Dis ic ,
A unachal P adesh; one adul male (WII-ADA3203)
collec ed by RNV and SD on 2 May 2023 om Deban
(27.49284°N, 96.38569°E, ele a ion 490 m a.s.l.), Nam-
dapha Tige Rese e, A unachal P adesh; one adul male
(WII-ADA3215) collec ed by RNV and SD on 10 May
2023 om Ho nbill (27.53810°N, 96.44037°E, ele a-
ion 660 m a.s.l.), Namdapha Tige Rese e, A unachal
P adesh; one adul male (WII-ADA3250) collec ed by
AD, RNV, SD and JDG on 25 May 2023 om 70 mile
(27.38734°N, 96.78273°E, ele a ion 870 m a.s.l.), Nam-
dapha Tige Rese e, A unachal P adesh; one adul male
(WII-ADA3265) collec ed by AD, RNV, SD and JDG
on 29 May 2023 om Kamala Valley (27.46148°N,
96.42758°E, ele a ion 650 m a.s.l.), Namdapha Tige Re-
se e, A unachal P adesh; one subadul (WII-ADA2537)
collec ed by KB on 27 Augus 2022 om Sessni
(27.04747°N, 92.41797°E, ele a ion 1260 m a.s.l.), Wes
Kameng Dis ic , A unachal P adesh; one subadul (WII-
ADA2548) collec ed by KB on 28 Augus 2022 om
Khellong (27.01314°N, 92.41408°E, 800 m a.s.l.), Wes
Kameng Dis ic , A unachal P adesh.
Diagnosis. A medium sized Rao ches es, SVL 19.6–26.3
mm in adul males and SVL 22.3–27.7 mm in adul e-
males; ome ine ee h absen ; snou ounded o subo a e
in do sal iew; nos il close o snou ip han eye; in-
e -uppe eyelid wid h less han eye leng h; nup ial pad
p esen on i s inge in males; nume ous ube cles on
head, do sum and limb in e mixed wi h spinules; p om-
inen blun ly conical ube cles on uppe eyelid; a b oad
da k b own “X” ma king on do sum ex ending o g oin;
b oad da k b own c ossba s on o elimb and hindlimb.
Desc ip ion o he holo ype. Holo ype well p ese ed
excep o an incision on unde side o igh high. Me-
dium sized Rao ches es, adul male wi h ocal sac, 24.8
mm in SVL; head leng h nea ly equal o wid h (HL/HW
= 0.97); snou sub-o al in do sal iew, unca ed in la e -
al iew, sligh ly p o uding beyond lowe jaw in en al
iew; snou sligh ly dep essed do sally a in e na ial e-
gion; snou leng h less han eye leng h (SL/EL = 0.81);
can hus os alis dis inc , obliqued; lo eal egion sligh ly
conca e; na ial egion p o uding; nos il o al, la e al-
ly posi ioned and obliquely o ien ed; nos il equidis an
be ween eye and snou ip; in e na ial dis ance less han
in e -uppe eyelid wid h (IN/IUE = 0.85) and equal o up-
pe eyelid wid h; eye mode a e in size, less han hal o
head leng h (EL/HL = 0.45); ympanum dis inc , ound-
ed less han one ou h o eye leng h; sup a ympanic old
dis inc ; ome ine ee h absen ; choanae ounded; ongue
pos e io ly no ched; unk do so- en ally la ened, i s
leng h nea ly hal o snou - en leng h (AG/SVL = 0.49).
Fo elimbs slende , o ea m leng h nea ly equal o
hand leng h (FAL/HAL = 0.94); ela i e leng h o inge s
= FI<FII<FIV<FIII; inge s wi h ounded disc; disc on
ou h inge wides and wide han ympanic diame e
(HTYD/FIVD = 0.63); ci cum-ma ginal g oo e p esen
on discs; palma ube cles indis inc ; suba icula ube -
cles dis inc and ounded, p oximal suba icula ube cle
on hi d and ou h inge s smalle han dis al ones; en-
la ged inely g anula nup ial pad p esen on i s inge ;
webbing be ween inge s absen .
Hindlimbs slende , high sligh ly longe han ib-
ia (TBL/TL = 0.96) and mo e han hal o he snou -
en leng h (TL/SVL = 0.54); ela i e leng h o oes =
TI<TII<TIII<TV<TIV; each oe wi h ounded disc; disc
as wide as inge disc; ci cum-ma ginal g oo e p esen
on each disc; elonga ed me a a sal ube cle p esen ; ou e
me a a sal ube cle absen ; webbing mode a e, eaching
second suba icula ube cle o ou h oe.
Skin on do sal aspec o snou and an e io pa o
head smoo h wi h sca e ed ube cles; p ominen ube -
cles on uppe eyelid; pos e io pa o he head wi h dense
spinules in e mixed wi h blun sligh ly la ge ube cles;
sca e ed ube cles below lo eal egion and ew p omi-
nen ube cles on mandibula egion; nume ous spinules
on do sum and lank; blun ube cles sca e ed among
spinules on lank; o elimbs smoo h do sally; hindlimbs
smoo h do sally wi h sca e ed blun ube cles; on en-
al aspec o h oa and ches smoo h; abdomen and high
g anula ; ibia smoo h; ube cles on en o-la e al aspec
o a sus and o ea m.
Colou a ion in li e. Do sal aspec o head, do sum, limbs
g eyish b own; indis inc ligh ba wi h da ke edge p es-
en on in e -uppe eyelid space a b oad da k b own “X”
ma k on do sum, pos e io ends o his ma king ex ending
o g oin whe e i is da ke ; da k b own s ipe along su-
p a ympanic old co e ing ympanum; b oad da k s ipe
on uppe jaw en al o eye; na ial egion da k b own; a
b oad da k b own band on o ea m, one on base o he
hand and one on each hi d and ou h inge ; h ee b oad
da k b own c ossba s on high and ibia; one simila ba
on each a sus and base o oo ; i egula da k b own
pa ches on op o oo and oes; da k b own pa ches on
la e al aspec o high; enla ged da k b own pa ch a ound
en ; on en al aspec o lowe jaw, ches , abdomen and
limbs whi ish; b own mo ling along lowe jaw; pale yel-
lowish pa ches on gula a ea and base o lowe a m.
Colou a ion in p ese a i e. Do sal aspec o head and
do sum b own; uppe eyelid da k g ey; ma kings on head,
do sum and limbs isible as in li e; o ea m, high, and
ibia pale han do sum; hand, a sus, and oo pale c eam
Ve eb a e Zoology 75, 2025, 517–625 605
Figu e 41. Rao ches es a unachalensis sp. no . A–C Holo ype (WII-ADA1553) in p ese ed condi ion, A do sal iew, B en al
iew, C la e al iew o head. D–G WII-ADA3203 in li e om Namdapha TR, D la e al iew, E en al iew, F g oin, G pos e io
iew o high. H Do sola e al and I en al iew o WII-ADA678 in li e om Po in. J Do sola e al iew o holo ype in li e. K Pa a-
ype (WII-ADA1554) in li e. L Do sola e al iew o WII-ADA1549 om Sessa in li e. M Do sola e al iew o WII-ADA1705 om
Mehao WLS in li e. Inse image showing dense ube cles on do sum in A, dis inc blun ube cles in D, g anula skin on abdomen
in I. Scale ba = 10 mm.
Bo uah B e al.: Re ision o bush ogs om no heas India
612
shape, oblique and la e ally posi ioned, equidis an om
snou ip and eyes; in e na ial dis ance smalle han in-
e -uppe eyelid wid h (IN/IUE = 0.79) and g ea e han
uppe eyelid wid h (UEW/IN = 0.84); in e uppe eyelid
space sligh ly con ex; can hus os alis smoo h, sligh ly
oblique; lo eal egion conca e; ympanum ound, indis-
inc , less han one hi d o eye leng h (HTYD/EL = 0.28);
eye mode a e in size, less han hal o head leng h (EL/
HL = 0.41).
Fo elimbs slende , o ea m leng h equal o hand
leng h, nea ly one hi d o snou - en leng h (FAL/SVL
= 0.27); hi d inge longes , ela i e leng h o inge s =
I<II<IV<III; inge s wi h ounded disc; ci cum-ma ginal
g oo e p esen on each disc; disc on hi d inge wides
and g ea e han he ympanic diame e (HTYD/FID =
0.78); webbing absen be ween inge s; suba icula u-
be cles dis inc , p oximal one on inge III and IV smalle
in size han ha o uppe one; supe nume a y ube cles
absen ; palma ube cles indis inc .
Hindlimbs slende ; high longe han ibia (TBL/TL
= 0.94) and oo (FOL/TL = 0.8); ibia longe han oo
(FOL/TBL = 0.85); ou h oe longes , ela i e leng h
o oes = I<II<III<V<IV, oes wi h ounded disc; ci -
cum-ma ginal g oo e p esen on each disc; disc on ou h
oe wide han ympanic diame e and sligh ly small-
e han ha o inge III; suba icula ube cles dis inc ,
ounded, p oximal one on oe IV e y small; supe nume -
a y ube cles absen ; inne me a a sal ube cle p esen ,
ou e me a a sal ube cle absen .
Skin on do sal aspec o head and do sum smoo h; dis-
inc and blun ly conical ube cles on uppe eyelid, pos-
e io pa o head, pos e io o uppe eyelid and an e io
pa o do sum; la e al aspec o head smoo h, ew ound-
ed ube cles behind angle o jaw; sup a ympanic olds
dis inc ; do sum wi h dense spinules, sp eading owa ds
lank; o elimbs and hindlimbs smoo h wi h indis inc u-
be cula p ojec ions; on en um, head smoo h, abdomen,
en al aspec o lowe a m, and high dis inc ly g anula ;
nume ous la g anules on palm and en al aspec o oo .
Colou a ion in li e. Do sally head, do sum and limb da k
b own, lank sligh ly pale ; enla ged black pa ch be ween
wo whi e pa ches p esen on g oin, co e ing pos e io
hal o he lank; simila black and whi e pa ches o i eg-
ula size and shape p esen on la e al and en al aspec s
o high, ibia, and la e al aspec o a sus; one b oad
black o da k b own c ossba p esen on do sal side o
each o ea m, high and ibia; i egula small pale bluish
whi e spo s on uppe jaw p esen ; en ally da k b own;
small i egula pale-bluish whi e spo s p esen on gula
egion, o ea m and hind a m, ches , abdomen and en al
side o a sus; digi al disc yellowish on do sal aspec .
Colou a ion in p ese a i e. Do sal aspec o head and
do sum simila o ha o li e colou a ion; do sal aspec
o limbs sligh ly pale b own han in li e; en al aspec
pale b own, all whi e and black ma kings isible as when
i was ali e.
Mo phological a ia ion. De ails o mo phome ic a i-
a ions a e p o ided in Table S12. Shape and size o he
black and whi e pa ches on g oin and hindlimbs a ies
among indi iduals; in ensi y o he small whi e spo s on
he en al aspec o body and on chin a ies among indi-
iduals o his species.
Phylogene ic ela ionship. Rao ches es minda is sis e
o a g oup con aining R. huanglianshan and R. meng-
laensis wi h weak suppo in ML analysis (Fig. 2). In BI
analysis, R. minda eco e ed as a sis e o R. annandalii
wi h weak suppo (Fig. 3). The gene ic di e gence o
R. minda wi h i s congene s included in his s udy a e
2.4–7.4% in he 16S, 11.7–17.7% in he cy b genes (Ta-
ble S7A–B).
Dis ibu ion and na u al his o y. Rao ches es minda
was known only om he ype locali y, be ween Hline Th-
oa illage and Gyin Dwe illage, O a ma aung Na ional
Pa k o Minda Dis ic , Myanma . The p esen s udy e-
po s his species om India o he i s ime (Fig. 19B)
and i ex ends he no heas dis ibu ion bounda y o he
species by 548 km om he ype locali y in Myanma . We
eco ded calling males in a deg aded ag icul u al a m on
he edge o Fakim Wildli e Sanc ua y (Fig. 44A). Calling
indi iduals we e ound on he hill slopes o an abandoned
jhum ield be ween 18:00–19:00 h s on lea es o Eupa-
Figu e 44. Habi a s o Rao ches es minda and R. annandalii A Landscape iew o he hills a Fakim WLS and B landscape iew
a Da jeeling.
Ve eb a e Zoology 75, 2025, 517–625 613
o ium sp., one me e abo e g ound. The a ea was mos ly
co e ed in maize and soybean as well as in asi e plan s
such as Eupa o ium sp. and Pa henium sp., on a slope
wi h an incline o app oxima ely 60 deg ees.
Rao ches es annandalii (Boulenge , 1906)
Figu es 45, 46; Tables 1, 2, S12
Ch esonymy.
Ixalus annandalii Boulenge , 1906: 385.
Ixalus annandalei — Annandale (1908): 305; inco ec spelling.
Rhacopho us (Philau us) annandalii — Ahl (1931): 71.
Philau us annandalii — Bou e (1942): 451; Go ham (1974): 166;
Sa ka e al. (1992): 90; Du a (1997): 74; Ahmed e al. (2009): 15,
147.
Philau us (Philau us) annandalii — Bossuy and Dubois (2001): 38.
Pseudophilau us annandalii — Li e al. (2009): 519.
Rao ches es annandalii — Subba e al. (2017): 4, 11; F os (2025).
Commen s on axonomic s a us. Boulenge (1906) de-
sc ibed Rao ches es annandalii om Ku seong, Da jeel-
ing (= Da jiling), Wes Bengal, India. Bossuy and Dubois
(2001) edesc ibed he lec o ype (BMNH 1947.2.26.58;
emale). We e-examined he lec o ype a NHM, London
and we collec ed opo ypes om Da jeeling. We ound he
ollowing excep ion in ou collec ed specimens agains
he lec o ype desc ip ion by Bossuy and Dubois (2001):
1) head longe han wide ( s. head leng h equal o wid h),
snou leng h equal o eye leng h ( s. snou leng h g ea e
han o equal o eye leng h); ibia longe han high ( s.
ibia sho e han o equal o high). Based on new in-
o ma ion, we p o ide a e ised diagnosis and expanded
desc ip ion o his species.
Ma e ial examined. Lec o ype (BMNH 1947.2.26.58;
Fig. 45) collec ed by N. Annandale om Ku seong, Da -
jeeling (= Da jiling), Wes Bengal.
Newly collec ed ma e ial. Fou adul males (WII-
ADA521, WII-ADA537, WII-ADA548, WII-ADA551)
and h ee adul emales (WII-ADA520, WII-ADA530
and WII-ADA533) collec ed by NGP on 23 Ap il 2018
om Ra angla (27.2879°N, 88.34717°E, ele a ion 1780
m a.s.l.), Namchi Dis ic , Sikkim; one adul male (WII-
ADA2721) and wo adul emales (WII-ADA2716 and
WII-ADA2722) collec ed by NGP on 8 Augus 2022 nea
Rock Ga den wa e all (27.02778°N, 88.22889°E, ele a-
ion 1500 m a.s.l.), Da jeeling Dis ic , Wes Bengal.
Diagnosis. A small sized Rao ches es (Fig. 46), SVL
17.3–19.6 mm in adul males and SVL 18.9–21.6 mm
in adul emales; head leng h equal o head leng h; om-
e ine ee h absen ; snou ounded o sub o oid in do sal
iew; snou sligh ly longe han eye leng h o equal o eye
leng h (EL/SL = 0.82–1.03); snou leng h g ea e han o
equal o in e -uppe eyelid wid h (SL/IUE = 0.97–1.23);
snou leng h g ea e han o equal o eye leng h (SL/EL =
0.96–1.15); nos ils equidis an be ween eye and snou ip
o close o eye han snou ip; nup ial pad p esen on i s
inge in males; do sal skin on head and do sum smoo h
o shag eened; sca e ed ube cles owa ds do sum and
lank; da k b own band on in e -uppe eyelid space; a pai
o conca e da k b own s ipes on do sum; b own mo -
ling and whi e lecks on en um.
Desc ip ion o a newly collec ed ma e ial (WII-
ADA2721). An adul male, small size, SVL 17.3 mm;
head as long as wide (HL/HW = 0.98); snou ounded in
do sal iew, p o uding beyond lowe jaw in la e al and
en al iews; snou leng h sligh ly g ea e han eye leng h
(EL/SL = 0.92); can hus os alis ounded, oblique; lo e-
al conca e; na ial egion p o uding; in e na ial dis ance
sligh ly less han in e uppe eyelid wid h (IN/IUE =
0.91) and g ea e han uppe eyelid wid h (UEW/IN =
0.75); in e -uppe eyelid space sligh ly con ex; nos il
o al, la e ally posi ioned and obliquely o ien ed, equidis-
an be ween eye and snou ip; eye in mode a e size, less
han hal o head leng h (EL/HL = 0.4), sligh ly g ea e
han in e uppe eyelid wid h (IUE/EL = 0.92); ympanum
ba ely isible, ounded, one ou h o eye leng h (HTYD/
EL = 0.25); sup a ympanic old dis inc ; ome ine ee h
absen ; choanae ound; ongue pos e io ly wide and me-
dially no ched, pos e io wo lobes widely sepa a ed; pai
o in e nal ocal sac opening on lowe jaw; habi us do -
so- en ally la ened, leng h mo e han hal o snou - en
leng h (AG/SVL = 0.52).
Fo elimbs slende ; hand longe han o ea m (FAL/
HAL = 0.83); ela i e leng h inge s = I<II<IV<III; in-
ge s wi h ounded disc; disc on hi d inge sligh ly wid-
e han ympanic diame e (HTYD/FIIID = 0.75); ci -
cum-ma ginal g oo e p esen on each disc; suba icula
ube cles dis inc and ounded excep he p oximal sub-
a icula ube cles on hi d and ou h inge which small
and indis inc ; palma ube cles indis inc ; webbing on
inge s absen ; a ine g anula nup ial pad on i s inge .
Hindlimbs slende ; high longe han ibia (TBL/TL =
0.94) and less han hal o snou - en leng h (TL/SVL =
0.48); ela i e leng h oes = I<II<III<V<IV; ounded disc
on oe; disc wid h equal o ha o inge s; suba icula
ube cles dis inc , ounded excep he lowe suba icula
ube cles on oe III–V which a e small and indis inc ; in-
dis inc inne me a a sal ube cle p esen and ou e one
absen ; webbing sligh , eaching second suba icula u-
be cle on ou h oe.
Snou and do sal aspec o head smoo h; an indis inc
longi udinal idge on middle o head; la e al aspec o
head smoo h, a ube cle p esen on mandibula egion;
do sum smoo h; lank g anula owa ds en ola e al as-
pec , indis inc ; o elimbs and hindlimbs smoo h on op;
on en um, h oa smoo h; pos e io ly gula skin g an-
ula ; ches , and abdomen g anula ; g anules on en al
aspec o high indis inc ; ibia smoo h on en al aspec .
Colou a ion in li e. Head, do sum and limbs b own on
do sal aspec ; an e io pa o head o in e o bi al space
sligh ly pale han pos e io pa o head and do sum; lo-
eal egion sligh ly da ke ; indis inc da k b own pa ch
en al o eye; a da k-b own s ipe below he sup a ym-
Bo uah B e al.: Re ision o bush ogs om no heas India
614
panic old; da k b own conca e ba on in e -uppe eyelid
space and da k b own pa ch pos e io o i ; pai o con-
ca e da k b own s ipes on do sum, s a ing behind uppe
eyelid and pos e io ends di ec ing owa ds g oin; a b oad
da k b own c ossba on o ea m; simila band on each
high, ibia, and a sus; disc on inge and oe yellow;
g oin, la e al aspec o high, inne la e al side o ibia
and a sus eddish-b own; da k b own pa ch a ound en ;
b own ma bling and i egula whi e lecks on en al side
o head, abdomen and limbs.
Sexual dimo phism and mo phological a ia ion.
Males ha e a pai o in e nal ocal sac openings on low-
e jaw; ex e nal subgula ocal sac, and nup ial pad on
i s inge . Do sal conca e s ipes and c oss ba s may be
indis inc in some indi iduals o his species; ube cle o-
wa ds lank may be comple ely absen ; De ailed mo pho-
me ic a ia ions a e gi en in Table S12.
Mo phological compa ison. Rao ches es annandalii
di e om R. ande soni, R. dulongensis, R. hillisi, R. hu-
an gli anshan, R. menglaensis, and R. y hus no . comb.
by posi ion o nos il equidis an be ween eye and snou
ip o close o eye han snou ip ( s. nos il close o
snou ip han eye); i di e s om R. cine ascens no .
comb. by snou leng h being g ea e han o equal o eye
leng h ( s. snou leng h smalle han eye leng h); i di e s
om R. dulongensis by nos il being equidis an be ween
eye and snou ip o close o eye han snou ip ( s. nos il
close o snou ip han eye); i di e s om R. hekouensis
by p esence o nup ial pad only on i s inge ( s. nup ial
pad p esen on i s and second inge s); i di e s om R.
hillisi and R. huanglianshan by nos il being equidis an
be ween eye and snou ip o close o eye han snou ip
( s. nos il close o snou ip han eye); i di e s om R.
jadoh by la ge body size in adul males, SVL 17.3–19.6
mm ( s. 13.6–14.0 mm); i u he di e s om R. jadoh
and R. jakoid by head leng h being g ea e han o equal
o head wid h ( s. head wide han long), and high leng h
being g ea e han o equal o ibia leng h ( s. high leng h
less han ibia leng h); i di e s om R. leik ho by la ge
body size in adul males, SVL 17.3–19.6 mm ( s. SVL
15.7–15.8 mm); i di e s om R. longchuanensis by in e
uppe eyelid wid h being less han o equal o eye leng h
( s. in e -uppe eyelid wid h g ea e han eye leng h); i
di e s om R. malipoensis by absence o black pa ch-
es on g oin and inne la e al aspec o high ( s. black
pa ches p esen on g oin and inne la e al side o high);
Figu e 45. Rao ches es annandalii (lec o ype; BMNH 1947.2.26.58). A Do sal and B en al iew, C en al aspec o hand, D
en al aspec o oo , E g anula skin on abdomen. scale ba = 10 mm.
Ve eb a e Zoology 75, 2025, 517–625 615
i di e s R. menglaensis by nos il being equidis an be-
ween eye and snou ip o close o eye han snou ip
( s. nos il close o snou ip han eye); i di e s om
R. minda by he absence o black o whi e pa ches on
g oin ( s. enla ged black and whi e pa ches p esen on
g oin), by absence o whi e pa ches on he la e al side o
high ( s. enla ged whi e pa ches on he la e al side o
high); i di e s om R. pa ulus absence o da k b own
and whi ish blo ch on g oin ( s. da k b own ma bling en-
closing a whi ish blo ch p esen on g oin); i di e s om
R. ezakhani by absence o da k o ligh pa e n on he
g oin ( s. a sho b own s eak p esen on g oin), and by
snou leng h being g ea e han in e -uppe eyelid wid h
( s. snou leng h less han o equal o in e uppe eyelid
wid h); i di e s om R. y hus no . comb. by posi ion
nos il equidis an be ween eye and snou ip o close o
eye han snou ip ( s. nos ils close o snou ip han
eye); i di e s om R. yadongensis by in e -uppe eyelid
wid h being less han o equal o eye leng h ( s. in e -up-
pe eyelid wid h g ea e han eye leng h). Mo phological
compa isons o R. annandalii wi h newly desc ibed spe-
cies in his s udy a e p o ided in he mo phological com-
pa ison sec ion o each espec i e species and Table 1.
Phylogene ic ela ionship and gene ic di e gence.
Phylogene ically R. annandalii eco e ed as basal lin-
eage o h ee species including R. a unachalensis sp.
no ., R. magnus sp. no . and R. longchuanensis in ML
analysis wi h weak suppo (Fig. 2). In BI analysis, i is
sis e o R. minda wi h weak suppo (Fig. 3). The gene -
ic di e gence wi h i s congene s included in his s udy
a e 3.8–8.2% in he 16S, 13.7–19.7% in he cy b and
10.0–16.8% in he COI genes (Table S7A–C).
Dis ibu ion and na u al his o y. Sa ka e al. (1992)
epo ed his species om Kalimpong, Goom i, Mahana-
di, Ghum Bhanjan in No he n Wes Bengal; Subba e al.
(2017) epo ed i om Sikkim; S es ha (2001), Ande s
(2002), Kha iwada e al. (2021) and Shah and Tiwa i
(2004) epo ed i om Nepal. Calling agg ega ions we e
obse ed on sh ubs a e he dusk du ing Ap il–July (Fig.
44B).
Rema ks. Annandale (1912) epo ed his species om
Mangaldai, Assam; Chanda (1992, 1994) epo ed his
species om Goalpa a in Assam and Namdapha in
A unachal P adesh ollowed by Sa ka and Ray (2006).
Ao e al. (2003) epo ed his species om Nagaland.
Howe e , ou sampling ac oss he egion shows ha R.
annandalii is es ic ed o Sikkim (Ra angla), Da jeeling
(Rock Ga den Wa e all) and eas e n Nepal. Fu he mo e,
he p esen s udy sugges s ha he p e ious eco ds o his
Figu e 46. Newly collec ed ma e ial o R. annandalii. A Do sola e al and B en al iew o opo ype (WII-ADA2721) in li e (inse
image showing g anula skin on abdomen). C WII-ADA534 in li e. D–G WII-ADA520, D do sal iew immedia ely a e eu haniza-
ion (inse image showing ine g anula skin on do sum), E do sal iew, F la e al iew o head, G en al iew. Scale ba = 10 mm.
Pho os A–D by Nai ik G. Pa el.
Bo uah B e al.: Re ision o bush ogs om no heas India
616
species om Assam (Annandale 1912), Nagaland (Ao e
al. 2003) and A unachal P adesh (Sa ka and Ray 2006;
Ma hew and Sen 2010) a e e oneous. Wangyal e al.
(2020) epo ed his species om Bhu an, which equi es
u he e i ica ion o con i m he iden i y o his species.
Discussion
The desc ip ion o hi een new species and one new e-
co d (R. minda ), as well as he e ision o he axonomy
o p e iously desc ibed species, has inc eased he o al
numbe o species in he genus Rao ches es om no h-
eas India o 21. In his e ision we esol ed he axonom-
ic iden i y and phylogene ic s a us o eigh his o ically
known species om he egion (R. ga o, R. kempiae, “R.
asakg ensis”, “R. manipu ensis”, “R. namdaphaensis”,
and “R. sahai”, Liu ana kempii no . comb., and Nasu ix-
alus mic odiscus no . comb.). The cu en gene ic place-
men o P. dubius s ill emains un esol ed and equi es
u he s udy. A ew s udies in ol ing la ge scale egion-
al sampling ha e been conduc ed in he Wes e n Gha s
and in no heas India (Biju and Bossuy 2009; Biju e al.
2011; Biju e al. 2014a, 2014b; Vijayakuma e al. 2014;
Ga g e al. 2021; Kamei e al. 2013; Mahony e al. 2018).
Howe e , his is he i s s udy o i s kind o include he
genus Rao ches es om he sub-Himalayan egion which
was p e iously he subjec o anecdo al s udies by Pil-
lai and Chanda (1979), Chanda (1992, 1994), Sengup a
e al. (2000), Choudhu y e al. (2001), Ao e al. (2003);
Sa ka and Ray (2006), Sen and Ma hew (2008), Ma hew
and Sen (2010), Sengup a e al. (2010), Sen e al. (2013),
Saikia e al. (2016), Ohle e al. (2018), Pu kayas ha e
al. (2020a) o he iden i y o he species was no de e -
mined (Das e al. 2009; Bo uah e al. 2018; Pu kayas ha
e al. 2020b). One o he ea u es o he no heas clade
o Rao ches es is he o e lapping mo phological cha ac-
e s o he species in he clade, unlike hei congene s in
he Wes e n Gha s which a e dis inc i e in hei mo pho-
logical cha ac e s, colou a ion and body size (Biju and
Bossuy 2005; Gu u aja e al. 2007; Biju and Bossuy
2009; Biju e al. 2010; Zacha iah e al. 2011; Seshad i e
al. 2012; Vijayakuma e al. 2014; Rajkuma e al. 2016;
Zacha iah e al. 2016; Ga g e al. 2021). All he species
desc ibed in his s udy a e endemic o no heas India.
Among he new species, R. boulenge i sp. no ., R. diban-
gensis sp. no ., R. khonoma sp. no ., R. lawng laiensis
sp. no ., R. mawsyn amensis sp. no . and R. monoli hus
sp. no . a e poin endemics cu en ly known only om
he ype locali ies. Fu he esea ch may ex end he dis i-
bu ion ange o neighbou ing egions. Ou s udy ex ended
he dis ibu ion o R. minda o India which was ecen -
ly desc ibed om Minda in Myanma by Köhle e al.
(2025). O he wo widesp ead species o he egion, R.
a unachalensis sp. no . and R. ga o, he la e is dis ib-
u ed in he sou he n pa o he B ahmapu a Valley name-
ly Meghalaya, Assam, Mizo am, Manipu , Nagaland and
sou heas e n pa o A unachal P adesh, Bangladesh o
Yunnan, China. The eco ds o R. pa ulus om he Ga-
ampani-Nambo -Doigu ung landscape in Assam (Sen-
gup a e al. 2010) and Bangladesh (Rahman e al. 2022)
need u he e i ica ion. The pho og aph o R. ande soni
(as Thelode ma ande soni) om Bangladesh (Ahmad
and Mim 2020) seems o be one o he mo phs o R. ga o.
Simila ly, he epo o R. ande soni om Bhu an (Wang-
yal and Jam sho 2022) equi es u he e i ica ion.
Recen s udies ha e highligh ed he impo ance o in e-
g a i e axonomic app oach especially o iden i y and de-
sc ibe c yp ic amphibians (Ga g e al. 2018, 2021; Go in
e al. 2024; Vences e al. 2024). Using mul iple lines o
e idence, we we e able o esol e he mis aken axonom-
ic iden i ies o he R. asakg ensis, R. cangyuanensis, R.
manipu ensis, R. sahai, and “P. namdaphaensis”. In addi-
ion, he p esen s udy demons a es he impo ance o in-
cluding he opo ypic ma e ial in de e mining he co ec
iden i y o c yp ic species wi h o e lapping dis ibu ions
such as in he cases o R. ga o and R. kempiae. Re-ex-
amina ion o he ype specimens enabled us o ans e
Philau us mic odiscus and Philau us kempii o Nasu ixa-
lus mic odiscus no . comb. and Liu ana kempii no .
comb. espec i ely. Addi ionally, we ecommend gene ic
ealloca ion o wo species o Philau us om Myanma o
he genus Rao ches es (R. cine ascens no . comb. and R.
y hus no . comb.).
In ou mul i a ia e analysis (PCA), R. annandalii, R.
ga o, R. minda , and R. shillongensis we e dis inc in
mo phospace wi hou o e lap wi h he o he membe s
o hei espec i e g oups (Fig. 6). Be ween he phylo-
gene ically sis e species such as R. shillongensis and R.
monoli hus sp. no ., R. mawsyn amensis sp. no . and
R. boulenge i sp. no . he e was no o e lap in mo pho-
space, while R. a unachalensis sp. no . and R. magnus
sp. no . ha e pa ial o e lap. Sample sizes we e low o
R. dibangensis sp. no . and R. lawng laiensis sp. no .,
addi ional sampling is equi ed o unde s and he ex en
o mo phological a ia ion o hese wo species. In ad-
di ion o his, phylogene ically dis an ly ela ed species
such as R. ba akensis sp. no ., R. boulenge i sp. no ., R.
eaglenes ensis sp. no ., R. kempiae, R. khonoma sp. no .,
R. mawsyn amensis sp. no ., R. monoli hus sp. no ., R.
na puhensis sp. no ., R. nasu a sp. no . and R. o ien a-
lis sp. no . showed pa ial o e lap in mo phospace. The
PCA analysis showed ha he small-bodied species a e
mo phologically c yp ic, while he e is clea mo phologi-
cal sepa a ion o la ge-bodied species (R. a unachalensis
sp. no ., R. eaglenes ensis sp. no ., R. ga o, and R. mag-
nus sp. no . All Rao ches es species ound in mainland
Asia, excep in no heas India, a e small-bodied. The e-
o e, he e is a high likelihood o disco e ing addi ional
c yp ic species om p e iously unsampled loca ions in
mainland Asia.
Phylogene ically he no heas Indian lineages o
Rao ches es adia ed om he “bombayensis g oup” (Vi-
jayakuma e al. 2016; Ga g e al. 2021). The phyloge-
ne ic ela ionships o many o he subclades wi hin his
sis e clade ound in no heas India and he Indochina
egion emain un esol ed wi h low nodal suppo (Figs
2, 3). Addi ional axon sampling ac oss his egion and
Ve eb a e Zoology 75, 2025, 517–625 617
nex -gene a ion sequencing may help esol e hese weak-
ly suppo ed nodes. Gene ic di e gences be ween sis e
species ound in no heas India a e mos ly mode a e
(>2% o <4% in he 16S gene) excep o a single spe-
cies (R. ga o) ha has high (≥4%) di e gence in he 16S
gene (Table S7A–C). Addi ionally, R. nasu a sp. no . has
he lowes (1.5–2%) di e gence in he 16S gene (Table
S7A–C) om an undesc ibed lineage ound in Tibe and
Adi Hills, A unachal P adesh (Figs 2, 3). Ou esul s also
shows ha widesp ead species such as R. a unachalen-
sis sp. no . ha e no able in a-species gene ic di e gence
(0.0–2.2%) in he 16S gene.
The gene ic di e gence among he species o his ge-
nus in no heas India is unlike ha o Wes e n Gha s. In
Wes e n Gha s, Rao ches es species a e mos ly highly
di e gen (Vijayakuma e al. 2014). This high di e -
gence can be a ibu ed o: i) he age o he lineages in he
Wes e n Gha s being olde (ca. 31 million yea s (my)),
and ii) geog aphical ba ie s including isola ed massi s,
and deep alleys such as he Palgha Gap and ele a ion-
al g adien s wi hin he Wes e n Gha s Moun ain anges.
Whe eas, he lineages in no heas India a e ecen adia-
ion (ca. 13 my; Vijayakuma e al. 2013) and he e ain
in he no heas Indian hills a e less a iable compa ed
o he Wes e n Gha s. Al hough he B ahmapu a Valley
has been iden i ied as a geog aphical ba ie o ce ain
e eb a e species (Mani 1974; Ripley and Beehle 1990;
Pawa e al. 2007; Deepak and Ka an h 2018) he ole o
hill anges and alleys in he specia ion o smalle e e-
b a es in no heas India emains poo ly unde s ood.
The di e si y o Rao ches es species ound in he
no heas hills (Sensu Mani 1974) sou h o he B ah-
mapu a alley is highe (16 species) compa ed o hose
ound no h o he alley ( i e species). A simila pa e n
is obse ed in he di e si y o ben - oed geckos in no h-
eas India (Aga wal e al. 2014; Bo uah e al. 2024). In
addi ion o his, he eas e nmos pa o he egion mo e
speci ically he Namdapha-Kamlang egion appea s o be
a con ac zone o he widely dis ibu ed species om he
no h o he alley (R. a unachalensis sp. no .) and spe-
cies om sou h o he alley (R. ga o).
Al i udinal g adien s in he Wes e n Gha s a e known
o ha e played a majo ole in he di e si ica ion o bush
ogs (Vijayakuma e al. 2016). Howe e , i is poo ly un-
de s ood in no heas India. Based on he samples we col-
lec ed om 82 locali ies ac oss no heas India, we ound
ha mos o he species in no heas India a e dis ibu ed
ac oss low o mid o high ele a ions (<100 m o >1500
m), excep o wo species (R. dibangensis sp. no . and
R. lawng laiensis sp. no .) ha a e es ic ed o low el-
e a ions (<500 m) and h ee species (R. eaglenes ensis
sp. no ., R. minda and R. khonoma sp. no .) ha a e e-
s ic ed o highe ele a ions (>1500 m). Addi ional sam-
pling in his egion and biogeog aphic analyses will ei he
con i m o e ise hese esul s and e eal di e si ica ion
pa e ns in bush ogs o no heas India and he ac o s
in luencing i .
Bioacous ic s udies ha e p o en o be use ul in am-
phibian axonomy especially in di e en ia ing c yp ic an-
u an species (P asad e al. 2020; Ga g e al. 2021; Vences
e al. 2024; Go in e al. 2024). The acous ic epe oi e o
Rao ches es om no heas India can be b oadly classi-
ied in o pulsa ile and non-pulsa ile calls. Compa a i ely
small-bodied species such as R. mawsyn amensis sp. no .
and R. na puhensis sp. no . ha e he highes dominan
equency calls, while la ge-bodied species such as R.
a unachalensis sp. no ., R. ga o, and R. eaglenes ensis
sp. no . ha e he lowes dominan equency calls. Some
o he Wes e n Gha s congene s ha e mo e han one ype
o call (1–3 ypes) and some o hem emi bo h pulsa-
ile and non-pulsa ile calls (e.g., R. esplendens Biju e
al., 2010; see Ga g e al. 2021). Howe e , species om
no heas India ha e a single ype o call and emi ei he
pulsa ile o non-pulsa ile calls. The p esen s udy is he
i s o i s kind in no heas India o desc ibe acous ics o
12 species o bush ogs. This will aid u u e esea ch and
acous ic moni o ing o anu ans in his egion.
Among he 13 new species o Rao ches es desc ibed
in his s udy, se en a e om p o ec ed a eas (PAs) o in
he icini y o PAs namely: R. ba akensis sp. no . (nea
Ba ail WLS, Assam), R. dibangensis sp. no . (nea Me-
hao WLS, A unachal P adesh), R. eaglenes ensis sp. no .
(Eaglenes WLS, A unachal P adesh), R. lawng laiensis
sp. no . (Ngengpui WLS, Mizo am), R. nasu a sp. no .
(Namdapha TR, A unachal P adesh), R. na puhensis sp.
no . (Na puh WLS, Meghalaya) and R. o ien alis sp. no .
(Namdapha TR, A unachal P adesh). The ype locali ies
o R. a unachalensis sp. no . and R. magnus sp. no . do
no all wi hin PAs bu hese species a e dis ibu ed in PAs
(Kamlang TR, Mehao WLS, Mouling NP, Namdapha TR,
A unachal P adesh). Among he PAs men ioned abo e,
Namdapha TR now has he iches di e si y o bush ogs
ep esen ed by i e species. In addi ion, ecen ly h ee
new amphibian species (G acixalus pa kaiensis, Alcalus
on inalis, Nidi ana noadihing Bo uah e al., 2023) ha e
been desc ibed om Namdapha TR, which ep esen s he
no he n limi o opical ain o es dis ibu ion (P oc o
e al. 1998). Ou s udy ei e a es he ac ha ex ensi e
sampling ac oss mul iple physiog aphic o ma ions is
c ucial o delinea ing c yp ic di e si y ha is h ea ened
om an h opogenic ac i i ies (A unachalam e al. 2004).
Du ing ou ieldsu eys we obse ed ongoing apid de-
o es a ion in he egion due o jhum cul i a ion, ca da-
mom cul i a ion, and linea in as uc u e and mega dam
p ojec s. P o ec ed a eas o he no heas Indian egion
hus a e co ne s ones o he conse a ion o endemic am-
phibian di e si y.
Acknowledgemen s
We hank he Na ional Geog aphic Socie y o he awa d o Na-
ional Geog aphic Explo e G an (NGS-74044R-20), SERB-DST
(CRG/2018/000790) and Meghalaya Biodi e si y Boa d (SBB.84/Re-
sea ch S udies/CAMPA/2021 da ed Ma ch 2022) o inancial suppo .
Fieldwo k was made possible wi h app op ia e pe missions and logis-
ical suppo by he S a e Fo es Depa men s o A unachal P adesh,
Assam, Meghalaya, Mizo am, Manipu , Nagaland, T ipu a, Sikkim
and Wes Bengal ( ide le e nos. FWC/G/173/P -111/3897-908 da ed
9 Feb ua y, 2016; CWL/GEN/13(95)/11-12/P .V/438-40 da ed 2 May
Bo uah B e al.: Re ision o bush ogs om no heas India
618
2018; CWL/G/173/2018-19/P .VII/1100-07 da ed 22 Augus 2019;
WL/FG.31/Technical Commi ee/2019, da ed 18 July, 2019; CWL/
GEN/96(Vol-III)/550 da ed 8 July, 2019; B.19060/1/2020-CWLW/112
da ed 2 Feb ua y 2021; CWL/GEN/355/2021/3178 da ed 28 Sep em-
be 2021; 866/WL/4R-28/2021 da ed 7 Ap il, 2021; No.F.8(163)/Fo -
WL-2020/MoU/1846-51 da ed 19 Ma ch, 2021; FOR.77/2019/66 da ed
27 June, 2022; CWL/173/2018-19/P .VII(A)/296-303 da ed 20 Ap il
2023). We since ely acknowledge he suppo ecei ed om o icials o
A unachal P adesh Fo es Depa men namely Sh. Milo Tase , Sh. Aduk
Pa on, Sh. Tajum Yomcha, Sh. Dhawan Kuma Rawa , Sh. Ha sh aj
Wa ho e, Sh. Taluk Rime, Sh. Bun y Tao, Sh. B. Da ang, Sh. Kabuk
Lego, Sh. Taluk Rime, Sh. Rupi Boli, Kashok Solung, Sh. Kamin Dai,
Sh. Tashi Mize, Basan Taye, Sh. Mayu Va iya, Sh. Adi ya Das o hei
suppo du ing ield wo k. We hank Sh. Aochuba (Field di ec o , In-
anki NP, Nagaland), Bendang Temsu (Range O ice , Kiphi e, Naga-
land), Lim hu e Yimchunge (Nagaland). We hank B. Lal inmuanpuia
(Mizo am) and Pa sanga (Ngengpui) o hei help. Ou since e hanks
o Sh. A piyush Sangma (Meghalaya Fo es dep .) o aluable suppo .
We hank Gobind Saga Bha dwaj (Di ec o ), Ruchi Badola (Dean) and
Sh. Ma ia Ch is u Raja D. (Regis a ) o Wildli e Ins i u e o India, Deh-
adun o hei cons an suppo . We a e g a e ul o he Di ec o , Zoolog-
ical Su ey o India (ZSI), Kolka a o gi ing us pe mission o examine
ype specimens. We hank Kaushik Deu i, ZSI, Kolka a o his suppo
du ing ype specimen examina ion. We hank Je S eiche o his cu a-
o ial suppo a NHM, London and Ma k Wilkinson o checking om-
e ine ee h o some o he ype specimens a NHM. We hank Ind aneil
Das and We ne Con adie o hei aluable inpu s on he i s d a o
his manusc ip . We hank San anu Dey, Nai ik G. Pa el, Vijayan Ji hin,
Jason D. Ge a d, Samuel Lal onunga, Sou a Du a, Raji N.V., K ish-
nendu Bane jee, Deb Shankha Goswami, Gau a Rai Pa iyal, P anoy K.
Bo ah, Vignesh, Mi za G. Ghazi, Shahid Da , Neelam Du a, Malsawm-
dawngliana, Isaac Zosangliana and K. Lalhmangaiha, o ield suppo .
We hank A. Madhan aj, Su ya P asad Sha ma, Swa i Nawani, Su ya
Na ayanan, Kumudani Bala Gau am, Bhim Singh o hei help du ing
lab wo k. BB hanks Shahid Da , Vinee Rana and Hem Rana o hei
suppo du ing he museum isi in Kolka a. We hank Ami ha Balan
and S abani Bose o hei help in s a is ical analyses. We hank S.K.
Du a, Ind aneil Das, U am Saikia and Sam a Mondol o hei suppo .
We hank ield assis an s p o ided by La e Teibo Ma wein, Lishi Gunia,
Aphu Yoha Yobin, Akhida, John Tayeng, Punya Chada, B in on Wa j i,
Chip a Mekola, Nga e Mega, Hachi Molo, Dochi Molo, Bhai i, Awaaz
and Saga . We hank he edi o and e iewe s o hei aluable sugges-
ions on p e ious e sions o his manusc ip .
Re e ences
Aga wal I, Baue AM, Jackman TR, Ka an h KP (2014) Insigh s in o
Himalayan biogeog aphy om geckos: A molecula phylogeny
o Cy odac ylus (Squama a: Gekkonidae). Molecula Phyloge-
ne ics and E olu ion 80: 145–155. h ps://doi.o g/10.1016/j.ym-
pe .2014.07.018
Ahl E (1927) Zu Sys ema ik de asia ischen A en de F oschga ung
Rhacopho us. Si zungsbe ich e de Gesellscha na u o schende
F eunde zu Be lin 1927: 35–47.
Ahl E (1931) Amphibia: Anu a III (Polypeda idae). Das Tie eich 55:
1–477.
Ahmad F, Mim EZ (2020) Rapid assessmen o amphibians a Milon-
cha i a ea in Banda ban Dis ic , Bangladesh. Rep iles & Amphibi-
ans 27: 36–41. h ps://doi.o g/10.17161/ anda. 27i1.14444
Ahmed MF, Das A, Du a SK (2009) Amphibians and Rep iles o No h-
eas India: A Pho og aphic Guide. Aa anyak, Guwaha i, 170 pp.
Al-Razi H, Ma ia M, Hasan S, Muza a SB (2020a) Fi s eco d o
Rao ches es longchuanensis Yang and Li, 1978 (Anu a: Rhacoph-
o idae) om no heas e n Bangladesh sugges s wide habi a ole -
ance. Amphibian and Rep ile Conse a ion 14: 119–131.
Al-Razi H, Ma ia M, Muza a SB (2020b) A new species o c yp ic
bush og (Anu a, Rhacopho idae, Rao ches es) om no heas e n
Bangladesh. ZooKeys 927: 127–151. h ps://doi.o g/10.3897/zook-
eys.927.48733
Al schul SF, Gish W, Mille W, Mye s EW, Lipman DJ (1990) Basic local
alignmen sea ch ool. Jou nal o Molecula Biology 215: 403–410.
Ande s CC (2002) Class Amphibia (Amphibians). In: Schleich HH,
Käs le W (Eds) Amphibians and Rep iles o Nepal: Biology, Sys-
ema ics, Field Guide. A.R.G. Gan ne Ve lag, Ruggell, 133–340.
Ande son J (1871) A lis o he ep ilian accession o he Indian Muse-
um, Calcu a, om 1865 o 1870, wi h a desc ip ion o some new
species. Jou nal o Asia ic Socie y o Bengal 40: 12–39.
Ande son J (1879 “1878”) Ana omical and Zoological Resea ches:
Comp ising an Accoun o he Zoological Resul s o he Two Expe-
di ions o Wes e n Yunnan in 1868 and 1875; and a Monog aph o
he Two Ce acean Gene a, Pla anis a. B. Qua i ch, London, 984 pp.
h ps://doi.o g/10.5962/bhl. i le.50434
Annandale N (1908) Ba achia – No es on some Ba achia ecen ly add-
ed o he collec ion o he Indian Museum. Reco ds o he Zoologi-
cal Su ey o India 2: 304–304.
Annandale N (1912) Zoological esul s o he Abo expedi ion, 1911–
1912. I. Ba achia. Reco ds o he Indian Museum 8: 7–36.
Annandale N (1913) Some new and in e es ing Ba achia and liza ds
om India, Ceylon and Bo neo. Reco ds o he Indian Museum 9:
301–307.
Ao JM, Bo doloi S, Ohle A (2003) Amphibian auna o Nagaland wi h
nine een new eco ds om he s a e including i e new eco ds o
India. Zoos’ P in Jou nal 18: 1117–1125. h ps://doi.o g/10.11609/
JoTT.ZPJ.18.6.1117-25
A unachalam A, Sa mah R, Adhika i D, Majumde M, Khan ML (2004)
An h opogenic h ea s and biodi e si y conse a ion in Namdapha
na u e ese e in he Indian Eas e n Himalayas. Cu en Science 87:
447–454.
A h eya R (2006) Eaglenes Biodi e si y P ojec (2003–2006): Con-
se a ion Resou ces o Eaglenes Wildli e Sanc ua y. Kaa i T us ,
Pune, 196 pp.
Benson DA, Ka sch-Miz achi I, Lipman DJ, Os ell J, Wheele DL
(2007) GenBank. Nucleic Acids Resea ch 35: D21–D25.
Bha a h B, Dinesh KP, Ka u hapandi M, Kun e KB, Shabnam A,
Jaiswal D, Jadha SS (2024) New eco ds o he p ima ily S i Lank-
an Pseudophilau us egius (Manamend a-A achchi & Pe hiyagoda,
2005) (Amphibia; Rhacopho idae) om he Eas e n Gha s o India,
c ossing he Palk S ai sal wa e ba ie in he Indian Ocean. Zoo -
axa 5477: 219–236. h ps://doi.o g/10.11646/zoo axa.5477.2.6
Biju SD, Bossuy F (2005) Two new Philau us (Anu a: Ranidae: Rhaco-
pho inae) om Ponmudi Hill in he Wes e n Gha s o India. Copeia
2005: 29–37. h ps://doi.o g/10.1643/CH-04-194R1
Biju SD, Bossuy F (2009) Sys ema ics and phylogeny o Philau-
us Gis el, 1848 (Anu a, Rhacopho idae) in he Wes e n Gha s o
India, wi h desc ip ions o 12 new species. Zoological Jou nal o
he Linnean Socie y 155: 374–444. h ps://doi.o g/10.1111/j.1096-
3642.2008.00466.x
Biju SD, Ga g S, Gu u aja KV, Shouche Y, Walujka SA (2014a) DNA
ba coding e eals unp eceden ed di e si y in dancing ogs o India
Ve eb a e Zoology 75, 2025, 517–625 619
(Mic ixalidae, Mic ixalus): A axonomic e ision wi h desc ip ion
o 14 new species. Ceylon Jou nal o Science (Biological Sciences)
43: 37–123. h ps://doi.o g/10.4038/cjsbs. 43i1.6850
Biju SD, Ga g S, Mahony S, Wijaya hilaka N, Sene i a hne G, Meegas-
kumbu a M (2014b) DNA ba coding, phylogeny and sys ema ics o
golden-backed ogs (Hyla ana, Ranidae) o he Wes e n Gha s-S i
Lanka biodi e si y ho spo , wi h he desc ip ion o se en new spe-
cies. Con ibu ions o Zoology 83: 269–335.
Biju SD, Sene i a hne G, Ga g S, Mahony S, Kamei RG, Thomas A,
Shouche Y, Raxwo hy CJ, Meegaskumbu a M, Bocxlae IV (2016)
F ankixalus, a new hacopho id genus o ee hole b eeding ogs
wi h oophagous adpoles. PloS One 11: e0145727. h ps://doi.
o g/10.1371/jou nal.pone.0145727
Biju SD, Shouche Y, Dubois A, Du a SK, Bossuy F (2010) A g ound-
dwelling hacopho id og om he highes moun ain peak o he
Wes e n Gha s o India. Cu en Science 98: 1119–1125.
Biju SD, Van Bocxlae I, Mahony S, Dinesh KP, Radhak ishnan C, Zach-
a iah A, Gi i V, Bossuy F (2011) A axonomic e iew o he nigh
og genus Nyc iba achus Boulenge , 1882 in he Wes e n Gha s,
India (Anu a: Nyc iba achidae) wi h desc ip ion o wel e new spe-
cies. Zoo axa 3029: 1–96. h ps://doi.o g/10.11646/zoo axa.3029.1.1
Bo uah B, Das A (2020) W es ling in he bushes: Agg essi e beha -
io o Rao ches es shillongensis (Pillai e Chanda, 1973). Russian
Jou nal o He pe ology 27: 296–298. h ps://doi.o g/10.30906/1026-
2296-2020-27-5-296-298
Bo uah B, Deepak V, Pa el NG, Ji hin V, Yomcha T, Das A (2023a) A
new species o g een ee og o he genus G acixalus (Anu a: Rha-
copho idae) om he e e g een o es o No heas India. Ve eb a e
Zoology 73: 557–574. h ps://doi.o g/10.3897/ z.73.e98444
Bo uah B, Na ayanan S, A a ind NA, Lal onunga S, Deepak V, Das A
(2024) Desc ip ion o six new species o Cy odac ylus G ay (Squa-
ma a: Gekkonidae) om no heas e n India. Ve eb a e Zoology 74:
453–486. h ps://doi.o g/10.3897/ z.74.e124752
Bo uah B, Raj P, Du a SK, Das A (2018) Redesc ip ion and geog aphic
dis ibu ion o Rao ches es shillongensis (Anu a: Rhacopho idae)
om Meghalaya, no heas India. Phyllomedusa 17: 3–20. h ps://
doi.o g/10.11606/issn.2316-9079. 17i1p3-20
Bo uah B, Saikia U, Das, A (2017) Rep oduc i e beha iou o R. shill-
ongensis om Meghalaya, No heas India. In: Das A (Ed.) Di e si y
and Ecology o Amphibians o India. ENVIS Bulle in: Wildli e and
P o ec ed A eas 19. Wildli e Ins i u e o India, Deh adun, 217–228.
Bossuy F, Dubois A (2001) A e iew o he og genus Philau us Gis-
el, 1848 (Amphibia, Anu a, Ranidae, Rhacopho inae). Zeylanica 6:
1–112.
Boulenge GA (1882) Ca alogue o he Ba achia Salien ia s. Ecauda a
in he collec ion o he B i ish Museum. Na u e 25: 601. h ps://doi.
o g/10.1038/025601a0
Boulenge GA (1893) Concluding epo on he ep iles and ba achians
ob ained in Bu ma by Signo L. Fea dealing wi h he collec ion
made in Pegu and he Ka in Hills in 1887–88. Annali del Museo
Ci ico di S o ia Na u ale di Geno a 13: 304–337.
Boulenge GA (1906) Desc ip ion o wo new Indian ogs. Jou nal o
he Asia ic Socie y o Bengal 2: 385–386.
Boulenge GA (1919) XII Desc ip ions o h ee new ba achians om
he Ga o Hills, Assam. Reco ds o he Zoological Su ey o India
16: 207–208.
Bou e R (1942) Les ba aciens de l’Indochine. Gou e nemen Géné al
de l’Indochine, Hanoi, 547 pp.
B oadley DG (1971) The Rep iles and Amphibians o Zambia. Kenya
Li ho, Nai obi, 143 pp.
B une i E (1918) Re ision o he O ien al Tipulidae wi h desc ip ions
o new species. Pa II. Reco ds o he Zoological Su ey o India
15: 255–340.
Chanda SK (1992) Fu he no e on he dis ibu ion pa e n o Amphibia
in no h-eas India. Jou nal o Bengal Na u al His o y Socie y, New
Se ies 11: 25–33.
Chanda SK (1994) Anu a (Amphibia) o no h eas e n India. Memo ies
o Zoological Su ey o India 18: 1–143.
Chanda SK, Das I, Dubois A (2000) Ca alogue o amphibian ypes in
he collec ion o he Zoological Su ey o India. Hamad yad 25:
100–128.
Chan KO, G isme LL (2022) G oupS uc : An R package o allome ic
size co ec ion. Zoo axa 5124: 471–482. h ps://doi.o g/10.11646/
zoo axa.5124.4.4
Chan KO, G isme LL, B own RM (2018) Comp ehensi e mul i-locus
phylogeny o old wo ld ee ogs (Anu a: Rhacopho idae) e eals
axonomic unce ain ies and po en ial cases o o e -and unde es i-
ma ion o species di e si y. Molecula Phylogene ics and E olu ion
127: 1010–1019. h ps://doi.o g/10.1016/j.ympe .2018.07.005
Chan KO, Hu e CR, Wood PLJ, G isme LL, B own RM (2020) Ta -
ge -cap u e phylogenomics p o ide insigh s on gene and species
ee disco dances in old wo ld ee ogs (Anu a: Rhacopho idae).
P oceedings o he Royal Socie y B 287: 20202102. h ps://doi.
o g/10.1098/ spb.2020.2102
Che J, Jiang K, Yan F, Zhang Y (2020) Amphibians and Rep iles in Ti-
be – Di e si y and E olu ion. Science P ess, Beijing, xxiii, 803 pp.
[in Chinese].
Chen JM, P endini E, Wu YH, Zhang BL, Suwannapoom C, Chen HM,
Jin JQ, Lemmon EM, Lemmon AR, S ua BL, Raxwo hy CJ (2020)
An in eg a i e phylogenomic app oach illumina es he e olu iona y
his o y o old wo ld ee ogs (Anu a: Rhacopho idae). Molecula
Phylogene ics and E olu ion 145: 106724. h ps://doi.o g/10.1016/j.
ympe .2019.106724
Choudhu y NK, Hussain B, Ba uah M, Saikia S, Sengup a S (2001)
Amphibian auna o Kam up Dis ic , Assam, wi h no es on hei
na u al his o y. Hamad yad 26: 276–282.
Das A, Saikia U, Mu hy BHCK, Dey S, Du a SK (2009) A he pe o-
aunal in en o y o Ba ail Wildli e Sanc ua y and adjacen egions,
Assam, no h-eas e n India. Hamad yad 34: 117–134.
Das I, Du a SK (1998) Checklis o he amphibians o India, wi h En-
glish common names. Hamad yad 23: 63–68.
Decemson H, Gouda S, Lalbiakzuala, Lalmuansanga, Hma GZ,
Vabei yu eilai M, Lal emsanga HT (2021) An anno a ed check-
lis o amphibians in and a ound Dampa Tige Rese e, Mizo am,
India. Jou nal o Th ea ened Taxa 13: 17918–17929. h ps://doi.
o g/10.11609/jo .6319.13.3.17918-17929
Deepak V, Ka an h P (2018) A idi ica ion d i en di e si ica ion o
an- h oa ed liza ds om he Indian subcon inen . Molecula Phy-
logene ics and E olu ion 120: 53–62. h ps://doi.o g/10.1016/j.
ympe .2017.11.016
Delo me M, Dubois A, G osjean S, Ohle A (2006) Une nou elle e go-
axinomie des Megoph yidae (Amphibia, Anu a). Aly es 24: 6–21.
De e JA, Fui en AM, Konu Ö, Wilkinson JA (2012) C yp ic o en
ogs o Myanma : An examina ion o he Amolops ma mo a us
species complex wi h he esu ec ion o Amolops a ghanus and
he iden i ica ion o a new species. Copeia 2012: 57–76. h ps://doi.
o g/10.1643/CH-10-180
Dubois A (1987) Miscellanea axinomica ba achologica (II). Aly es 6:
1–9.
Bo uah B e al.: Re ision o bush ogs om no heas India
620
Dubois A, Ohle A (1998) A new species o Lep ob achium (Vib issa-
pho a) om no he n Vie nam, wi h a e iew o he axonomy o
he genus Lep ob achium (Peloba idae, Megoph yinae). Dume ilia
4: 1–32.
Du L, Xu Y, Liu S, Yu G (2024) A new species o Rao ches es (Anu-
a, Rhacopho idae) om Yunnan P o ince, China. ZooKeys 1192:
213–235. h ps://doi.o g/10.3897/zookeys.1192.106013
Du a SK (1997) Amphibians o India and S i Lanka (Checklis and
Bibliog aphy). Odyssey Publishing House, Bhubaneswa , 342 pp.
Ellepola G, Meegaskumbu a M (2023) Di e si ica ion and bioge-
og aphy o Rhacopho idae – A model es ing app oach. F on ie s
in Ecology and E olu ion 11: 1195689. h ps://doi.o g/10.3389/
e o.2023.1195689
E e L, Haas A, Lee CC, Min PY, Das I, He wig ST (2021) Ou o
he ap: A new phy o helm-b eeding species o Philau us and an
upda ed phylogeny o Bo nean bush ogs (Anu a: Rhacopho idae).
Jou nal o Zoological Sys ema ics and E olu iona y Resea ch 59:
1064–1096. h ps://doi.o g/10.1111/jzs.12465
Fei L (Ed.) (1999) A las o Amphibians o China. Henan Science &
Technology P ess, Zhengzhou, 432 pp.
Fei L, Hu SQ, Ye CY, Huang YZ (2009) Fauna Sinica. Amphibia. Vol-
ume 2. Anu a. Chinese Academy o Science, Science P ess, Beijing,
957 pp.
Fei L, Ye CY, Jiang JP (2012) Colo ed A las o Chinese Amphibians
and Thei Dis ibu ions. Sichuan Publishing House o Science and
Technology, Chengdu, 619 pp.
F os DR (2025) Amphibian Species o he Wo ld: An Online Re e -
ence. Ve sion 6.1. Ame ican Museum o Na u al His o y, New Yo k,
NY, h p:// esea ch.amnh.o g/he pe ology/amphibia/index.h ml [ac-
cessed 31 July 2025]
Ga g S, Sene i a hne G, Wijaya hilaka N, Phuge S, Deu i K, Manamen-
d a-A achchi K, Meegaskumbu a M, Biju SD (2018) An in eg a i e
axonomic e iew o he Sou h Asian mic ohylid genus Upe odon.
Zoo axa 4384: 1–88. h ps://doi.o g/10.11646/zoo axa.4384.1.1
Ga g S, Suyesh R, Das S, Bee MA, Biju SD (2021) An in eg a i e ap-
p oach o in e sys ema ic ela ionships and de ine species g oups
in he sh ub og genus Rao ches es, wi h desc ip ion o i e new
species om he Wes e n Gha s, India. Pee J 9: e10791. h ps://doi.
o g/10.7717/pee j.10791
Go ham SW (1974) Checklis o Wo ld Amphibians up o Janua y 1,
1970. New B unswick Museum, Sain John, 173 pp.
Go in VA, O lo NL, B agin AM, Pawangkhanan P, Mil o KD, Le
DX, Van Nguyen T, Du esnes C, Suwannapoom C, Poya ko NA
(2024) Phylogeog aphic pa e n and axonomic e ision o he Ka-
loula balea a species complex (Amphibia, Anu a, Mic ohylidae)
wi h desc ip ion o wo new species om Indochina. He pe ozoa 37:
391–420. h ps://doi10.3897/he pe ozoa.37.e137394
Gün he A (1875) Second epo on Indian ep iles ob ained by he
B i ish Museum. P oceedings o he Zoological Socie y o London
1875: 224–234.
Gün he A (1876) Thi d epo on collec ions o Indian ep iles ob ained
by he B i ish Museum. P oceedings o he Zoological Socie y o
London 1875: 567–577.
Gu u aja KV, A a ind NA, Ali S, Ramachand a TV, Vela an TP, K ish-
nakuma V, Agga wal RK (2007) A new og species om he cen-
al Wes e n Gha s o India, and i s phylogene ic posi ion. Zoologi-
cal Science 24: 525–534. h ps://doi.o g/10.2108/zsj.24.525
Hakim J, T agese SJ, Ghose A, Rashid SMA, Rahman SC (2020) Am-
phibians and ep iles om Lawacha a Na ional Pa k in Bangladesh.
Check Lis 16: 1239–1268. h ps://doi.o g/10.15560/16.5.1239
Hou M, Yu G-h, Chen H-m, Liao C-l, Zhang L, Chen J, Li P-p, O -
lo NL (2017) The axonomic s a us and dis ibu ion ange o six
Thelode ma species (Anu a: Rhacopho idae) wi h a new eco d
in China. Russian Jou nal o He pe ology 24: 99–127. h ps://doi.
o g/10.30906/1026-2296-2017-24-2-99-127
Huang J, Liu XL, Du L, Be ns ein JM, Liu S, Yang Y, Yu G, Wu Z (2023)
A new species o bush og (Anu a, Rhacopho idae, Rao ches es)
om sou heas e n Yunnan, China. ZooKeys 1151: 47–65. h ps://
doi.o g/10.3897/zookeys.1151.95616
Inge RF (1985) Rhacopho idae. In: F os DR (Ed.) Amphibian Species
o he Wo ld: A Taxonomic and Geog aphical Re e ence. Associ-
a ion o Sys ema ics Collec ions and Allen P ess, Law ence, KS,
525–550.
Je don TC (1870) No es on Indian he pe ology. P oceedings o he Asi-
a ic Socie y o Bengal 1870: 66–85.
Jiang K, Ren J, Wang J, Guo J, Wang Z, Liu Y, Jiang D, Li J (2020)
Taxonomic e ision o Rao ches es menglaensis (Kou, 1990) (Am-
phibia: Anu a), wi h desc ip ions o wo new species om Yunnan,
China. Asian He pe ological Resea ch 11: 263–281. h ps://doi.
o g/10.16373/j.cnki.ah .200018
Jiang K, Yan F, Wang K, Zou DH, Li C, Che J (2016) A new genus and
species o ee og om Medog, sou heas e n Tibe , China (Anu a,
Rhacopho idae). Zoological Resea ch 37: 15–20.
Kalyaanamoo hy S, Minh BQ, Wong TK, on Haesele A, Je miin LS
(2017) ModelFinde : Fas model selec ion o accu a e phylogene -
ic es ima es. Na u e Me hods 14: 587–589. h ps://doi.o g/10.1038/
nme h.4285
Kamei RG, Gowe DJ, Wilkinson M, Biju SD (2013) Sys ema ics o
he caecilian amily Chikilidae (Amphibia: Gymnophiona) wi h he
desc ip ion o h ee new species o Chikila om no heas India.
Zoo axa 3666: 401–435. h ps://doi.o g/10.11646/zoo axa.3666.4.1
Kapli P, Lu e opp S, Zhang J, Kobe K, Pa lidis P, S ama akis A, Flou-
i T (2017) Mul i- a e Poisson ee p ocesses o single-locus species
delimi a ion unde maximum likelihood and Ma ko chain Mon e
Ca lo. Bioin o ma ics 33: 1630–1638. h ps://doi.o g/10.1093/bio-
in o ma ics/b x025
Kha kongo IJ, Saikia B, Deb R (2016) De ailed s udy on Philau us
ga o (Boulenge , 1919) (Amphibia: Anu a: Rhacopho idae) – An
ex emely a e and endemic species o no heas India. Reco ds o
he Zoological Su ey o India 116: 439–444.
Kha iwada JR, Wang B, Zhao T, Xie F, Jiang J (2021) An in eg a i e ax-
onomy o amphibians o Nepal: An upda ed s a us and dis ibu ion.
Asian He pe ological Resea ch 12: 1–35. h ps://doi.o g/10.16373/j.
cnki.ah .200050
Köhle G, Dos O, Than NL, Ohle A, Cha un ochana PT, Chuaynke n
Y, Chuaynke n C, Geiss K (2025) A axonomic e ision o he genus
Rao ches es in Myanma and Thailand wi h he desc ip ion o wo
new species om Myanma (Amphibia, Anu a, Rhacopho idae).
Zoo axa 5613: 47–81. h ps://doi.o g/10.11646/zoo axa.5613.1.2
Köhle J, Jansen M, Rod iguez A, Kok PJ, Toledo LF, Emm ich M,
Glaw F, Haddad CF, Rödel MO, Vences M (2017) The use o bio-
acous ics in anu an axonomy: Theo y, e minology, me hods and
ecommenda ions o bes p ac ice. Zoo axa 4251: 1–124. h ps://
doi.o g/10.11646/zoo axa.4251.1.1
Kuma S, S eche G, Tamu a K (2016) MEGA7: Molecula e olu ion-
a y gene ics analysis e sion 7.0 o bigge da ase s. Molecula Biol-
ogy and E olu ion 33: 1870–1874. h ps://doi.o g/10.1093/molbe /
msw054
Lal onunga S, Van amliana VA, Lal amliana LA, Lalhmingliani E
(2021) A new coun y eco d o Rao ches es cangyuanensis Wu,
Ve eb a e Zoology 75, 2025, 517–625 621
Suwannapoom, Xu, Mu phy & Che 2019 and addi ional eco d o
Ku ixalus yangi Yu, Hui, Rao & Yang 2018 (Anu a: Rhacopho idae:
Rhacopho inae) om India. Zoo axa 4974: 383–390. h ps://doi.
o g/10.11646/zoo axa.4974.2.7
Lan ea R, Calco B, Ho SY, Guindon S (2012) Pa i ionFinde : Com-
bined selec ion o pa i ioning schemes and subs i u ion models o
phylogene ic analyses. Molecula Biology and E olu ion 29: 1695–
1701. h ps://doi.o g/10.1093/molbe /mss020
Li JT, Che J, Mu phy RW, Zhao H, Zhao EM, Rao DQ, Zhang YP
(2009) New insigh s o he molecula phylogene ics and gene ic
assessmen in he Rhacopho idae (Amphibia: Anu a) based on i e
nuclea and h ee mi ochond ial genes, wi h commen s on he e o-
lu ion o ep oduc ion. Molecula Phylogene ics and E olu ion 53:
509–522. h ps://doi.o g/10.1016/j.ympe .2009.06.023
Mahony S, Kamei RG, Teeling EC, Biju SD (2018) C yp ic di e si y
wi hin he Megoph ys majo species g oup (Amphibia: Megoph y-
idae) o he Asian ho ned ogs: Phylogene ic pe spec i es and a
axonomic e ision o Sou h Asian axa, wi h desc ip ions o ou
new species. Zoo axa 4523: 1–96. h ps://doi.o g/10.11646/zoo -
axa.4523.1.1
Mahony S, Kamei, RG, Teeling EC, Biju SD (2020) Taxonomic e iew
o he Asian ho ned ogs (Amphibia: Megoph ys Kuhl & an Has-
sel ) o no heas India and Bangladesh p e iously misiden i ied as
M. pa a (Boulenge ), wi h desc ip ions o h ee new species. Jou -
nal o Na u al His o y 54: 119–194. h ps://doi.o g/10.1080/002229
33.2020.1736679
Mahony S, Teeling EC, Biju SD (2013) Th ee new species o ho ned
ogs, Megoph ys (Amphibia: Megoph yidae), om no heas India,
wi h a esolu ion o he iden i y o Megoph ys boe ge i popula-
ions epo ed om he egion. Zoo axa 3722: 143–169. h ps://doi.
o g/10.11646/zoo axa.3722.2.2
Mani MS (1974) Biogeog aphy o he Himalaya. In: Mani MS (Ed.)
Ecology and Biogeog aphy in India. W. Junk, The Hague, 664–681.
Ma hew R, Sen N (2008) On he occu ence o Philau us namda-
phaensis Sa ka and Sanyal and Rhacopho us naso Annandale (An-
u a: Rhacopho idae) om Meghalaya, no h eas e n India. Cob a 2:
14–16.
Ma hew R, Sen N (2009) S udies on li le known amphibian species o
no heas India. Reco ds o Zoological Su ey o India, Occasional
Pape 293: 1–64, 23 pla es.
Ma hew R, Sen N (2010) Pic o ial Guide o he Amphibians o No h-
eas India. Zoological Su ey o India, Kolka a, 144 pp.
Minh BQ, Nguyen MAT, on Haesele A (2013) Ul a as app oxima-
ion o phylogene ic boo s ap. Molecula Biology and E olu ion
30: 1188–1195. h ps://doi.o g/10.1093/molbe /ms 024
Muansanga L, Malsawmdawngliana F, Vabei yu eilai M, Bualchhuak T,
Lal emsanga HT (2022) Colo polymo phism in Je don’s ee og,
Nasu ixalus je donii (Gün he 1876), om No heas India wi h
no es on die and dis ibu ion in Mizo am, India. Rep iles and Am-
phibians 29: 55–58. h ps://doi.o g/10.1761/ anda. 29i1.16209
Na een RS, Chand amouli SR, Babu S, Ryndongsngi AM, Ka unaka an
PV, Kuma a HN (2024) Redisco e y and edesc ip ion o Ixalus
ga o Boulenge , 1919, and Ixalus kempiae Boulenge , 1919, wi h a
eassessmen o he axonomic s a us o Rao ches es cangyuanensis
Wu, Suwannapoom, Xu, Mu phy & Che, 2019 and he desc ip ion
o a new species om he Ga o Hills o Meghalaya. He pe ozoa 37:
359–372. h ps://doi.o g/10.3897/he pe ozoa.37.e122825
Na een RS, Na h KP, Lal emsanga HT, Deu i K, Decemson H, Muan-
sanga L, Vabei yu eilai M, Malsawmdawngliana F, Wa j i H, Pu -
kayas ha J (in p ess) A bush og wi h mul iple names: New popula-
ions o Rao ches es kempiae (Boulenge , 1919) and eassessmen o
he axonomic s a us o Philau us namdaphaensis (Sa ka and Sany-
al, 1985) and Rao ches es manipu ensis (Ma hew and Sen, 2009).
Jou nal o Asia-Paci ic Biodi e si y 18. h ps://doi.o g/10.1016/j.
japb.2025.05.003
Nguyen LT, Schmid HA, on Haesele A, Minh BQ (2015) IQ-TREE:
A as and e ec i e s ochas ic algo i hm o es ima ing maxi-
mum-likelihood phylogenies. Molecula Biology and E olu ion 32:
268–274. h ps://doi.o g/10.1093/molbe /msu300
Ohle A (2003) Re ision o he genus Oph yoph yne Boulenge , 1903
(Megoph yidae) wi h desc ip ion o wo new species. Aly es 21:
23–44.
Ohle A, Bo ah MM, Das MK, Tesia C, Bo doloi S (2018) A s udy on
amphibian auna o A unachal P adesh (India). Aly es 36: 276–288.
Pai a CA (1919) Rhyncho a om he Ga o Hills, Assam. Reco ds o he
Zoological Su ey o India 16: 349–377.
Pawa S, Koo MS, Kelley C, Ahmed MF, Chaudhu i S, Sa ka S (2007)
Conse a ion assessmen and p io i iza ion o a eas in No heas In-
dia: P io i ies o amphibians and ep iles. Biological Conse a ion
136: 346–361. h ps://doi.o g/10.1016/j.biocon.2006.12.012
Pillai RS, Chanda SK (1973) Philau us shillongensis, a new og (Ra-
nidae) om Meghalaya, India. P oceedings o he Indian Academy
o Sciences 78: 30–36.
Pillai RS, Chanda SK (1979) Amphibian auna o Khasi hills, Megha-
laya. Reco ds o he Zoological Su ey o India 383–395.
Poya ko NA, O lo J NL, Moisee a AV, Pawangkhanan P, Ruang-
suwan T, Vassilie a AB, Galoyan EA, Nguyen TT, Gogole a SS
(2015) So ing ou moss ogs: m DNA da a on axonomic di e -
si y and phylogene ic ela ionships o he Indochinese species o
he genus Thelode ma (Anu a, Rhacopho idae). Russian Jou nal
o He pe ology 22: 241–280. h ps://doi.o g/10.30906/1026-2296-
2015-22-4-241-280
P asad VK, Gau am KB, Gup a SK (2020) Iden i ica ion o anu an
species di e si y o he Panna Tige Rese e, Cen al India, us-
ing an in eg a ed app oach. Zoo axa 4851: 450–476. h ps://doi.
o g/10.11646/zoo axa.4851.3.2
P oc o J, Ha idasan K, Smi h (1998) How a no h does lowland e -
e g een opical ain o es go? Global Ecology and Biogeog aphy
Le e s 7: 141–146. h ps://doi.o g/10.2307/2997817
Puilland e N, B ouille S, Achaz G (2021) ASAP: Assemble species by
au oma ic pa i ioning. Molecula Ecology Resou ces 21: 609–620.
h ps://doi.o g/10.1111/1755-0998.13281
Pu kayas ha J, Khan N, Roychoudhu y S (2020a) A p elimina y check-
lis o he pe o auna occu ing in Rowa Wildli e Sanc ua y, T ipu a,
India. In: Roy N, Roychoudhu i S, Nau iyal S, Aga wal SK, Baksi S
(Eds) Socio-economic and Eco-biological Dimensions in Resou ce
Use and Conse a ion: S a egies o Sus ainabili y. Sp inge , Cham,
225–233. h ps://doi.o g/10.1007/978-3-030-32463-6_10
Pu kayas ha J, Roychoudhu y S, Biswa BB, Das M, Sengup a S (2020b)
He pe o aunal di e si y and conse a ion s a us in Amchang Wild-
li e Sanc ua y o Assam, India. In: Roy N, Roychoudhu i S, Nau iyal
S, Aga wal SK, Baksi S (Eds) Socio-economic and Eco-biological
Dimensions in Resou ce Use and Conse a ion: S a egies o Sus-
ainabili y. Sp inge , Cham, 217–223. h ps://doi.o g/10.1007/978-
3-030-32463-6_9
R Co e Team (2021) R: A language and en i onmen o s a is ical
compu ing. R Founda ion o S a is ical Compu ing, Vienna. h ps://
www.R-p ojec .o g
Rahman MM, Chen JM, Wu YH, Chen HM, Lwin YH, Mu phy RW,
Li GG, Che J (2020) New coun y eco ds o h ee species o
