A axonomic e ision o he Inge oph ynus pa us
species complex (Amphibia, Anu a, Bu onidae) wi h he
desc ip ion o a new species om Thailand
Dmi iy V. A khipo 1, Pa inya Pawangkhanan 2,3, Gou am Sa ke 4,5, Vladisla A. Go in1, And ey M. B agin6,
Tan Van Nguyen7,8, Ami Hamidy9, Mohd Abdul Muin10, Nia Ku niawan11, Ra e M. B own12,
Cha mongkon Suwannapoom2, E ic Smi h4, Nikolay A. Poya ko 1
1 Depa men o Ve eb a e Zoology, Lomonoso Moscow S a e Uni e si y, Leninskiye Go y, Moscow 119234, Russia
2 Di ision o Fishe y, School o Ag icul u e and Na u al Resou ces, Uni e si y o Phayao, Phayao 56000, Thailand
3 Rabbi in he Moon Founda ion, Suanphueng, Ra chabu i 70180, Thailand
4 Amphibian and Rep ile Di e si y Resea ch Cen e (ARDRC) and Depa men o Biology, The Uni e si y o Texas a A ling on, A ling on, USA
5 Co ey College, 1000 W Aus in Bl d, Ne ada, MO 64772, USA
6 Join Vie nam - Russia T opical Science and Technology Resea ch Cen e , 63 Nguyen Van Huyen Road, Nghia Do, Cau Giay, Hanoi 122000, Vie nam
7 The School o Medicine & Pha macy, Duy Tan Uni e si y, Da Nang, 550000, Vie nam
8 Cen e o En omology & Pa asi ology Resea ch, Duy Tan Uni e si y, Da Nang, 550000, Vie nam
9 Labo a o y o He pe ology, Museum Zoologicum Bogo iense, Resea ch Cen e o Biology, Indonesian Ins i u e o Sciences-LIPI, Widyasa waloka
Jl. Raya Jaka a Bogo km 46, Cibinong, Wes Ja a, Indonesia
10 Cen e o Global Sus ainabili y S udies, Hamzah Sendu Lib a y 1, Uni e si i Sains Malaysia, 11800 Minden, Penang, Malaysia
11 Depa men o Biology, Uni e si as B awijaya, Jl. Ve e an, Malang, Eas Ja a, Indonesia
12 Biodi e si y Ins i u e and Depa men o Ecology and E olu iona y Biology, Uni e si y o Kansas, Law ence, KS 66045, USA
h ps://zoobank.o g/DFF82964-7476-401C-B0FD-D9722A4CCF21
Co esponding au ho s: Nikolay A. Poya ko ([email p o ec ed]); E ic Smi h ([email p o ec ed]);
Cha mongkon Suwannapoom ([email p o ec ed])
Academic edi o : A hu Tiu enko ♦
Recei ed
15 July 2025 ♦
Accep ed
2 Sep embe 2025 ♦
Published
1 Oc obe 2025
Abs ac
A e iew o he axonomic s a us o he Inge oph ynus pa us species complex, based on mo phological, bioacous ic, and molecula
da a, e ealed a new species om sou he n Thailand, which we desc ibe as Inge oph ynus сh ysolophus sp. no . The new species
is dis inguished om i s congene s by he combina ion o he ollowing mo phological cha ac e s: pa o oid elonga e, na ow, and
sha ply aised; wa s on lanks less ele a ed han hose o do sum; c anial c es s no hickened behind eyes; lo es e ical; ympanum
dis inc , i s diame e sligh ly exceeding wo- hi ds o eye leng h; ibia sho ; i s inge longe han second; ip o hi d oe no each-
ing median suba icula ube cle o ou h oe; suba icula ube cles no enla ged; a sal spine bases small; nup ial pads p esen ;
en e wi h low wa s; g ound colo o lanks and do sum ligh b own; da k b own s ipes along he midline o he back; c anial
idges well-de eloped, b igh o ange. Phylogene ic analysis o he m DNA agmen , including he 12S RNA, 16S RNA, and ND1
gene sequences (up o 4,479 bp), con i ms he placemen o he new species as a sis e o Inge oph ynus pa us s. s . (p-dis ance
5.37%). The gene ic dis inc i eness o Inge oph ynus сh ysolophus sp. no . and I. pa us s. s . is u he con i med by no sha ing
haplo ypes o h ee nuclea genes (POMC, BDNF, and RAG1). Bioacous ic analysis e ealed s able di e ences be ween he male
ad e isemen calls o he wo species. Fu he mo e, ecological modeling shows ha he new species is allopa ic wi h espec o
I. pa us s. s .; hei anges a e likely sepa a ed by a biogeog aphical bounda y o he Kanga -Pa ani Line. The new species is
cu en ly known om low- o mid-ele a ions (ca. 65–1,000 m a.s.l.) in opical o es s o sou he n and wes e n Thailand, sou he n
Myanma , and sou hwes Cambodia. We sugges he new species be conside ed Leas Conce n (LC) acco ding o he IUCN Red Lis
c i e ia. Ou s udy u he unde sco es he u gen need o in ensi ied in eg a i e axonomic esea ch o he genus Inge oph ynus o
cla i y he axonomy o wide- anging species complexes and o elabo a e e ec i e conse a ion measu es.
He pe ozoa 38: 271–297 (2025)
DOI 10.3897/he pe ozoa.38.e165173
Copy igh Dmi iy V. A khipo 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.
he pe ozoa.penso .ne
Dmi iy V. A khipo e al.: A new species o Inge oph ynus om Thailand272
Key Wo ds
Asia, in eg a i e axonomy, Kanga -Pa ani Line, mo phology, sys ema ics
In oduc ion
The oad genus Inge oph ynus F os , G an , Fai o ich,
Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Don-
nellan, Raxwo hy, Campbell, Blo o, Mole , D ewes,
Nussbaum, Lynch, G een & Wheele , 2006 inhabi s op-
ical o es a eas h oughou sou he n mainland China,
Hainan Island, and Indochina h ough Myanma , penin-
sula Thailand, he Thai-Malay Peninsula, Suma a, Bo -
neo, Ja a, Nias, Sulawesi, and he Philippines (F os e
al. 2006; F os 2025). The genus Inge oph ynus is mo -
phologically diagnosed by he ollowing combina ion o
cha ac e s: he p esence o sup ao bi al, pa ie al, and su-
p a ympanic c es s; he absence o a a sal idge; ocal
sacs lacking melanopho es in he su ounding muscle is-
sue; he absence o ibial glands; he absence o supina o
manus hume alis and adduc o longus muscles; he p es-
ence o pai ed c es s on he e eb al column; a ugose
skull; squamosal bones wi h b oad do sal o ic pla es; and
smoo h pala ine bones (Inge 1972; G isme 2007). The
genus Inge oph ynus cu en ly includes 13 ecognized
species (F os , 2025), namely, I. bipo ca us (G a enho s ,
1829), I. celebensis (Gün he , 1859), I. cla ige (Pe e s,
1863), I. di e gens (Pe e s, 1871), I. galea us (Gün he ,
1864), I. gollum G isme , 2007, I. kumqua (Das & Lim,
2011), I. ledongensis (Fei, Ye & Huang, 2009), I. mac o is
(Boulenge , 1887), I. pa us (Boulenge , 1887), I. philip-
pinicus (Boulenge , 1887), I. quad ipo ca us (Boulenge ,
1887), and I. wangyingyongi Liu, Zhang, Ananje a, Liu,
Hou & O lo , 2025. Howe e , he phylogene ic ela ion-
ships among he membe s o he genus Inge oph ynus
ha e no been s udied ex ensi ely, and i is qui e likely
ha i s cu en di e si y is unde es ima ed.
The Malayan dwa oad, Inge oph ynus pa us Bou-
lenge , 1887, was desc ibed by Boulenge (1887) om
he en i ons o Malacca, Malaysia (Fig. 1). This species
has been conside ed widely dis ibu ed ac oss sou he n
Myanma , sou he n, wes e n, and eas e n Thailand, Cam-
bodia, Peninsula Malaysia, and Indonesia (Poya ko e
al. 2021; F os 2025). Recen s udies ha e sugges ed ha
I. pa us may in ac ep esen a complex o species due
o subs an ial mo phological and molecula di e ences
be ween he popula ions in he sou he n and no he n
pa s o he Thai–Malay Peninsula (Mulcahy e al. 2018;
S ion e al. 2018; Chan and G isme 2019).
In his wo k, we p esen an upda ed mi ochond ial
DNA (he ea e m DNA) genealogy o he genus In-
ge oph ynus, combining sequences a ailable in GenBank
wi h newly gene a ed sequences ob ained om an ex en-
si e sampling ac oss Sou heas Asia. We u he analyze
he di e en ia ion wi hin he I. pa us species complex
using h ee nuclea DNA (he ea e nuDNA) loci, bio-
acous ic analysis, and ecological niche modeling. Ou e-
sul s sugges a subs an ial di e en ia ion o he I. pa us
species complex popula ions loca ed on di e en sides o
he Kanga -Pa ani Line, an impo an biogeog aphical
bo de in Sou heas Asia (Poya ko e al. 2021, 2023).
He ein, we p o ide a axonomic e ision o he I. pa us
species complex and desc ibe he popula ions om no h
o he Kanga -Pa ani Line as a new species.
Ma e ials and me hods
Sample collec ion
Fieldwo k was ca ied ou in di e en p o inces o Thai-
land in Ma ch and Decembe 2020, Janua y and Feb-
ua y 2022, No embe 2021, Feb ua y 2024, and July
2024 (Fig. 1). Specimens we e collec ed by hand du ing
excu sions along o es ails o nea b eeding si es, in-
cluding empo a y ain ponds and slow-mo ing s eams.
Geog aphic coo dina es and al i ude da a we e accu a ely
cap u ed using a Ga min GPSMAP 60CSx GPS ecei -
e and eco ded in he WGS 84 da um. The specimens
we e eu hanized using a 20% benzocaine solu ion. P io
o p ese a ion, issue samples we e ex ac ed o gene ic
analysis and s o ed in 96% e hanol; hese samples includ-
ed ei he emo al muscles o a piece o li e . Specimens
we e subsequen ly ixed in 4% bu e ed o malin and la -
e ans e ed o 70% e hanol.
Specimen collec ion and animal use p o ocols in Thai-
land we e app o ed by he Ins i u ional E hical Com-
mi ee o he Ins i u ional E hical Commi ee o Animal
Expe imen a ion o he Uni e si y o Phayao, Phayao,
Thailand (ce i ica e numbe UP-AE64-02-04-005, is-
sued o C. Suwannapoom) and we e s ic ly complian
wi h he ecommenda ions o he Thailand Animal Wel-
a e Ac . Fieldwo k, including he collec ion o animals
in he ield, was au ho ized by he Ins i u e o Animals
o Scien i ic Pu pose De elopmen (IAD), Bangkok,
Thailand (pe mi numbe s U1-01205-2558 and UP-
AE59-01-04-0022, issued o C. Suwannapoom). Re-
sea ch in Indonesia was conduc ed unde esea ch pe mi
149/SIP/FRP/SM/V/2013 (issued o E.N. Smi h).
Addi ional specimens and issue samples we e ob ained
om museum collec ions. Specimens and issues o igina -
ed om he he pe ological collec ions o he Zoological
Museum o Moscow Uni e si y (ZMMU, Moscow, Rus-
sia), Rabbi in he Moon Founda ion (RIM, Suanphueng,
He pe ozoa 38: 271–297 (2025)
he pe ozoa.penso .ne
273
Ra chabu i, Thailand), he School o Ag icul u e and Na -
u al Resou ces, he Uni e si y o Phayao (AUP, Phayao,
Thailand), he Uni e si y o Kansas, Museum o Na u al
His o y (KU, Law ence, Kansas, USA), he Uni e si y o
Texas a A ling on (UTA, A ling on, Texas, USA), and
he Kunming Ins i u e o Zoology, Chinese Academy o
Sciences (KIZ, Kunming, China). The loca ion o he ex-
amined popula ions and he dis ibu ion o he I. pa us
species complex a e shown in Fig. 1.
Labo a o y me hods
DNA was isola ed om he high muscles o li e o
ouche ed specimens. Fo he molecula phylogene -
ic analyses, o al genomic DNA was isola ed using he
s anda d phenol-chlo o o m-p o einase K ex ac ion
p ocedu es wi h consequen isop opanol p ecipi a ion
o a inal concen a ion o abou 1 mg/ml (p o ocols ol-
lowed Hillis e al. 1996; Samb ook and Russell 2001).
Figu e 1. Dis ibu ion o he lineages o he Inge oph ynus pa us species complex as iden i ied in he phylogene ic analyses. S a s
deno e ype locali ies, wi h hei co esponding axon names in colo ed clouds; colo s co espond o hose used in Figs 2–5; symbol
shapes deno e m DNA lineages shown in Fig. 2. Base map c ea ed using simplemapp .ne . Fo locali y in o ma ion, see Suppl. ma-
e ial 1: able S2. Pho og aphs by P. Pawangkhanan and N.A. Poya ko .
he pe ozoa.penso .ne
Dmi iy V. A khipo e al.: A new species o Inge oph ynus om Thailand274
We isualized he isola ed o al genomic DNA using aga-
ose elec opho esis in he p esence o e hidium b omide.
We measu ed he concen a ion o o al DNA in 1 μl using
NanoD op 2000 (The mo Scien i ic) and consequen ly
adjus ed i o ca. 100 ng DNA/μL.
In o al, we ampli ied wo agmen s o m DNA and
h ee nuDNA genes. P ime s used in PCR and sequenc-
ing a e summa ized in Suppl. ma e ial 1: able S1 and
we e aken om Hedges (1994), Goebel e al. (1999),
Wiens e al. (2005), P amuk (2006), Van de Meijden e
al. (2007), Páez-Moscoso and Guayasamin (2012), and
Lied ke e al. (2017). We pe o med DNA ampli ica ion
in 20 μl eac ions using ca. 50 ng genomic DNA, 10 nmol
o each p ime , 15 nmol o each dNTP, 50 nmol addi ion-
al MgCl2, Taq PCR bu e (10 mM T is-HCl, pH 8.3, 50
mM KCl, 1.1 mM MgCl2, and 0.01% gela in), and 1 uni
o Taq DNA polyme ase.
Fi s , we ampli ied a 453 bp long con inuous agmen
o he 16S RNA m DNA gene. This agmen o he 16S
RNA gene is widely used o biodi e si y su eys in am-
phibians (Vences e al. 2005; Viei es e al. 2009) and has
been analyzed in he mos ecen phylogene ic s udies o
he amily Bu onidae (e.g., Sma e al. 2017; Chan and
G isme 2019; Sa ke e al. 2019; Suwannapoom e al.
2021, 2022). The PCR condi ions in ol ed an ini ial de-
na u a ion s ep o 5 min a 94 °C, ollowed by 34 cycles
o dena u a ion o 1 min a 94 °C, p ime annealing o
1 min a 50 °C, ex ension o 1 min a 72 °C, and a inal
ex ension s ep o 5 min a 72 °C.
Addi ionally, o he selec ed samples ep esen ing
di e en lineages wi hin he genus Inge oph ynus, we
ampli ied a 2,100 bp long con inuous m DNA agmen ,
including pa ial sequences o he 12S RNA gene, com-
ple e sequences o RNAVal, 16S RNA, RNALeu, and
pa ial sequences o he NADH dehyd ogenase subuni 1
gene (ND1). The PCR condi ions in ol ed an ini ial de-
na u a ion s ep o 3 min a 94 °C, ollowed by 35 cycles o
dena u a ion o 35 s a 94 °C, p ime annealing o 40 s
a 48 °C, ex ension o 40 s o 12S RNA / 60 s o ND1
a 72 °C, and a inal ex ension s ep o 10 min a 72 °C.
Fo he selec ed samples ep esen ing di e en popu-
la ions wi hin he I. pa us species complex, we also am-
pli ied h ee nuDNA genes: POMC, BDNF, and RAG1.
P ime s used in PCR and sequencing a e summa ized in
Suppl. ma e ial 1: able S1. The PCR condi ions o he
POMC gene we e as ollows: an ini ial dena u a ion s ep
o 3 min a 94 °C, ollowed by 44 cycles o dena u a ion
o 35 s a 94 °C, p ime annealing o 40 s a 54 °C,
ex ension o 40 s a 72 °C, and a inal ex ension s ep o
10 min a 72 °C. The PCR condi ions o he BDNF gene
we e as ollows: an ini ial dena u a ion s ep o 5 min a
94 °C, ollowed by 32 cycles o dena u a ion o 1 min
a 94 °C, p ime annealing o 1 min a 50 °C, ex ension
o 1 min a 72 °C, and a inal ex ension s ep o 10 min
a 72 °C. The PCR condi ions o he RAG1 gene we e
as ollows: an ini ial dena u a ion s ep o 5 min a 94 °C,
ollowed by 10 cycles o dena u a ion o 1 min a 94 °C,
p ime annealing o 1 min wi h ouchdown empe a u e
om 65 °C o 55 °C, wi h empe a u e educing by 1 °C
pe each cycle, ex ension o 1 min a 72 °C, ollowed by
34 cycles o dena u a ion o 1 min a 94 °C, p ime an-
nealing o 1 min a 55 °C, ex ension o 1 min a 72 °C,
and a inal ex ension s ep o 10 min a 72 °C.
We an all ampli ica ions using an iCycle The mal
Cycle (Bio-Rad). We loaded he PCR p oduc s on o 1%
aga ose gels in he p esence o e hidium b omide and
isualized hem by elec opho esis. The success ul a -
ge ed PCR p oduc s we e pu i ied by he Dia om DNA
PCR Clean-Up ki and ou sou ced o E ogen® (Mos-
cow, Russia) o sequencing; sequence da a collec ion
and isualiza ion we e pe o med on an ABI 3730xl Au-
oma ed Sequence (Applied Biosys ems). We deposi ed
he newly ob ained sequences in GenBank unde he ac-
cession numbe s PX209002–PX209038 and PX213461–
PX213508 (Suppl. ma e ial 1: able S2).
Phylogene ic analyses
To econs uc he ma ilineal genealogy o he genus
Inge oph ynus, we used newly ob ained 16S RNA se-
quences o I. pa us om Thailand and he sequences o
he 12S RNA, 16S RNA, and ND1 m DNA agmen s
o he I. pa us species complex membe s om Thai-
land, Myanma , Malaysia, and Indonesia, as well as o h-
e Inge oph ynus species, ob ained om GenBank. Sup-
pl. ma e ial 1: able S2 summa izes he in o ma ion on
GenBank accession numbe s, museum ouche s, and he
locali y o o igin o he sequences used in his s udy. We
used sequences o en Asian Bu onidae ep esen a i es
as ou g oups o oo he ee, namely Ansonia lep opus
(Gün he , 1872), Du aph ynus c . melanos ic us (Schnei-
de , 1799), Lep oph yne bo bonica (Tschudi, 1838),
Ph ynoidis jux aspe (Inge , 1984), Peloph yne mise a
(Mocqua d, 1890), Ren apia hosii (Boulenge , 1892),
Pseudobu o subaspe Tschudi, 1838, Sabahph ynus
macula us (Mocqua d, 1890), Pa apeloph yne scalp a
(Liu & Hu, 1973), and Sigalegaleph ynus ha eyi Sa k-
e , Wos l, Thammacho i, Sidik, Hamidy, Ku niawan &
Smi h, 2019, based on he phylogene ic esul s o F os e
al. (2006), Mulcahy e al. (2018), and Chan and G isme
(2019). In o al, we ob ained m DNA da a o 65 speci-
mens o Inge oph ynus, which included en ou o he 13
cu en ly ecognized Inge oph ynus species and 44 spec-
imens o he I. pa us species complex; he geog aphic
dis ibu ion o he sampled popula ions o he I. pa us
species complex is shown in Fig. 1.
We ini ially aligned nucleo ide sequences using Clus -
alX 1.81 (Thompson e al. 1994) wi h de aul pa ame e s
and hen op imized hem manually in BioEdi 7.0.5.2
(Hall 1999) and MEGA 11.0 (Tamu a e al. 2013). We
u ilized ModelFinde (Kalyaanamoo hy e al. 2017) o
de e mine he mos sui able e olu iona y models o ou
da a se analysis; hey a e p esen ed in Suppl. ma e ial 1:
able S3. We calcula ed pai wise unco ec ed gene ic dis-
ances (p-dis ances) be ween sequences wi h MEGA 11.0.
He pe ozoa 38: 271–297 (2025)
he pe ozoa.penso .ne
275
We in e ed he ma ilineal genealogy using Bayesian
In e ence (BI) and Maximum Likelihood (ML) app oach-
es. We conduc ed BI using M Bayes 3.1.2 (Ronquis and
Huelsenbeck 2003). Me opolis-coupled Ma ko chain
Mon e Ca lo (MCMCMC) analyses we e un wi h one
cold chain and h ee hea ed chains o one million gen-
e a ions and sampled e e y 1,000 gene a ions. We pe -
o med wo independen MCMCMC uns, and he ini ial
100 ees we e disca ded as bu n-in. We assessed con i-
dence in ee opology based on he equency o nodal
esolu ion (pos e io p obabili y; BI PP) (Huelsenbeck
and Ronquis 2001). We used IQ-TREE (Nguyen e al.
2015) o econs uc ML ees. A o al o 10,000 ul a as
boo s ap eplica ions o ML analysis (UFBS) (Minh e
al. 2013) assessed he con idence in ee opology o ML
analysis. In bo h da ase s, we a p io i ega ded ee nodes
wi h BI PP alues o e 0.95 and UF BS alues o e 95%
as s ongly suppo ed. We conside ed BI PP alues be-
ween 0.95 and 0.90 and UF BS alues be ween 95% and
90% as endencies, while lowe alues we e conside ed
o indica e he lack o suppo (Huelsenbeck and Hillis
1993; Minh e al. 2013).
Addi ionally, o he h ee nuclea ma ke s examined
(POMC, BDNF, and RAG1), we cons uc ed allele ne -
wo ks o each gene using he median-joining me hod in
PopA e . 1.5 (Leigh and B yan 2015) wi h a 95% con-
nec ion limi . Fo he pu pose o allele ne wo k cons uc-
ion, sequences wi h mo e han one he e ozygous si e
we e esol ed in PHASE 2.1.1 (S ephens e al. 2001), o
which he inpu da a we e p epa ed in SeqPHASE (Flo
2010). PHASE was un unde de aul se ings, excep o
he p obabili y h eshold, which was se o 0.7.
Mo phological examina ion
Mo phome ic da a we e aken o 44 adul males and 32
emales o he Inge oph ynus pa us species complex
(Table 1). Measu emen s we e aken using a Mi u oyo
digi al calipe o he nea es 0.01 mm and subsequen ly
ounded o 0.1 mm. All measu emen s we e aken on he
igh side o he examined specimen. The mo phome ics
o adul s and cha ac e e minology ollowed G isme
(2007) and included he ollowing cha ac e s: snou -
en leng h, om ip o snou o en (SVL); head leng h
om ip o snou o hind bo de o angle o jaw (HL);
head wid h, wid h o head a i s wides poin (HW); head
heigh in he in e o bi al egion (HD); dis ance om he
ip o he muzzle o he nos ils (S-N); dis ance be ween
nos ils (IND); dis ance om nos il o an e io co ne
o eye (N-E); ho izon al eye diame e (ED); in e o bi -
al dis ance, as he dis ance be ween he inne bo de o
he uppe eyelid (IOD); eyelid wid h a he wides pa
(ELW); dis ance be ween he eye and he ympanic mem-
b ane (ETD); dis ance om eyes o ip o snou (ESD);
ho izon al diame e o ympanum (TD); c anial c es
leng h (CRL); pa o oid Wid h (PGW); pa o oid gland
leng h (PGL); leng h o he o elimbs om he base o
he o elimb o he ip o he longes inge (FLL); leng h
o he o ea m and hand om he elbow o he ip o he
longes inge (FHL); o ea m wid h (FAW); hand leng h
om he p oximal base o he ou e me aca pal ube cle
o he ip o he longes inge (HAL); 1s inge leng h
(F1); 2nd inge leng h (F2); 3 d inge leng h (F3); 4 h
inge leng h (F4); leng h o he in e nal me aca pal u-
be cle (IMC); leng h o he ex e nal me aca pal ube -
cle (OMC); hip leng h om anus o knee (FEL); ibia
leng h, dis ance be ween knee and ibio a sal a icula ion
(TIL); a sus and oo leng h (TFL); oo leng h om he
p oximal base o he in e nal me a a sal om he ube cle
o he ip o he longes oe (FOL); 1s oe leng h (T1); 2nd
oe leng h (T2); 3 d oe leng h (T3); 4 h oe leng h (T4); 5 h
oe leng h (T5); leng h o he in e nal me a a sal ube cle
(IMT); leng h o he ou e me a a sal ube cle (OMT).
Toe webbing and suba icula ube cle o mulas we e de-
sc ibed ollowing Sa age (1975). The sex and ma u i y
o he specimens we e checked by mino dissec ions and
examina ion o gonads and di ec obse a ion o calling
in li ing males p io o collec ion. O he abb e ia ions:
Dis . = Dis ic ; FR = Fo es Rese e; M = Moun ain;
NP = Na ional Pa k; NR = Na u al Rese e; P o . = P o -
ince; asl. = abo e sea le el.
Among he examined popula ions, SVL was compa ed
using a one-way ANOVA wi h he Tukey-K ame es .
The pe cen age a io (R) o each mo phome ic cha ac-
e o SVL was subsequen ly calcula ed; we compa ed
22 cha ac e a ios agains SVL among popula ions using
he K uskal-Wallis es . PCA was conduc ed o examine
o e all mo phological a ia ion among popula ions using
log- ans o med me ic alues ollowing Nishikawa e al.
(2007). When a high co ela ion be ween ce ain pai s o
cha ac e s was ound, we omi ed one o hem om he
analyses o exclude possible o e weigh ing e ec s. S a-
is ical analyses we e pe o med wi h S a is ica 6.0 (S a -
So Inc. 2001). The signi icance le el was se a p < 0.05.
The diagnosis o he genus Inge oph ynus and mo -
phological cha ac e s o compa ison we e aken om
he o iginal desc ip ions and axonomic e iews o he
genus: Boulenge (1887), Inge (1966, 1972), Man hey
and G ossmann (1997), Das and Lim (2001), G isme
(2007), and Fei e al. (2012).
Species dis ibu ion modeling (SDM)
We used he p og am Maxen 3.4.1 (Phillips e al. 2006;
Phillips and Dudik 2008) o model he po en ial dis i-
bu ion o he Inge oph ynus pa us species complex
membe s in Sou heas Asia. A o al o 155 unique geo-
e e enced da a poin s (Suppl. ma e ial 1: able S4) and
19 bioclima ic a iables e lec ing he heigh , aspec ,
and deg ee o inclina ion o he Ea h’s su ace; a idi y
index; ege a ion ypes; he pe cen age o co e age o
he Ea h’s su ace by woody ege a ion; and empe a-
u e and p ecipi a ion da a h oughou he yea (bio1-
19) we e used o gene a e he model a 5 km pixel size.

he pe ozoa.penso .ne
Dmi iy V. A khipo e al.: A new species o Inge oph ynus om Thailand276
Table 1. Mo phome ic cha ac e s (in mm) eco ded om each specimen o Inge oph ynus pa us sensu s ic o and Inge oph ynus
сh ysolophus sp. no . examined in his s udy. No es: H = holo ype, P = pa a ype, R = e e ed ma e ial, M = adul male, F = adul
emale. (Con inued on he nex page).
ZMMU ID Locali y S a us Sex SVL HL HW HD S-N IND N-E ED IOD ELW ETD
Inge oph ynus pa us
ZMMU A-8029 Yala, Thailand – F 48.5 13.8 16.6 6.8 1.7 3.8 3.2 5.3 4.8 4.2 1.1
ZMMU A-8021 Yala, Thailand – F 47.9 13.9 15.9 6.6 1.6 3 2.9 6 4.4 4.2 1.1
ZMMU NAP-09684 Selango , Malaysia – F 44.5 13.2 16.9 8.9 1.6 3.2 2.8 5.3 4.9 3.7 0.5
ZMMU A-8025 Yala, Thailand – M 33.1 9.8 11.6 5.2 0.9 2.5 2.1 4 3.4 3.1 1.2
ZMMU A-8026 Yala, Thailand – M 34.9 9.4 11.2 4.6 1.3 2.3 2.6 4.3 3.6 3.2 0.8
ZMMU A-8024 Yala, Thailand – M 33.8 10.2 11.9 4.4 1.3 2.3 2.4 4.4 3.8 3.1 0.9
ZMMU A-8022 Yala, Thailand – M 33.5 9.9 11.7 5.3 1.1 2.3 2 4.5 3.1 3.2 0.7
ZMMU A-8023 Yala, Thailand – M 36.2 10.1 11.6 4.5 1.2 2.5 2.1 4.9 4 3.1 0.8
ZMMU A-8027 Yala, Thailand – M 35.4 9.8 11.8 5.2 1.4 2.5 2.6 4.4 3.9 3 1.1
ZMMU A-8028 Yala, Thailand – M 36.7 10.8 12.2 5 1.4 2.5 2.5 4.7 3.9 3.4 0.8
Inge oph ynus сh ysolophus sp. no .
ZMMU A-8035 Chumphon, Thailand P F 34.2 10.2 11.8 4.8 1.3 2.4 2.2 4.3 3.3 3.3 0.7
ZMMU A-8030 Chumphon, Thailand H F 34.8 10.6 11 3.7 1.9 2.4 2.8 4.7 3.7 3.2 0.7
ZMMU A-8036 Chumphon, Thailand P F 35 9.6 10.6 5.2 1.2 2.2 2.5 4.3 3.5 3.2 0.7
ZMMU A-8037 Chumphon, Thailand P F 34 9.3 10.6 4.3 1.1 2.2 2.5 4.1 3.7 3.4 0.7
ZMMU NAP-09425 Suan Phueng, Ra chabu i, Thailand R F 38.2 10.7 13.4 5.9 1.2 2.3 2.9 4.8 3.6 3.4 1
ZMMU A-8211 Suan Phueng, Ra chabu i, Thailand R F 35.3 10.2 12 5.1 1.3 2.4 2.3 4.5 4.1 3.3 0.7
ZMMU A-8061 Suan Phueng, Ra chabu i, Thailand R F 36.3 10.5 11.6 5.3 1.5 2.3 2.6 4.2 3.9 3.4 0.8
ZMMU A-8031 Chumphon, Thailand P F 39.5 11.2 12.6 5.2 1.5 2.7 2.8 4.1 3.9 3.5 0.8
ZMMU A-8033 Chumphon, Thailand P F 37.8 10.4 12.9 5.3 1.6 2.8 2.6 4.8 3.7 3.5 0.9
ZMMU A-8032 Chumphon, Thailand P F 42.4 11.3 12.8 5.2 1.5 2.9 2.7 4.7 3.9 3.5 1
ZMMU A-8057 Suan Phueng, Ra chabu i, Thailand R M 30.3 8.7 10.3 3.5 1.2 2 2 3.9 3.2 3.1 0.5
ZMMU A-8055 Suan Phueng, Ra chabu i, Thailand R M 35.7 11 12 4.5 1.4 2.3 2.9 4.3 3.6 3.4 0.8
ZMMU A-8020 Phuke , Thailand R M 32.6 9.9 10.9 5 1.4 2.1 2.2 4 3.5 3.5 0.9
ZMMU A-8049 Suan Phueng, Ra chabu i, Thailand R M 33.3 9.8 10.6 5 1.3 2.2 2.2 3.9 3.7 3.2 0.8
ZMMU A-8060 Suan Phueng, Ra chabu i, Thailand R M 34.4 10.4 12 5.3 1.4 2.2 2.5 4.3 3.8 3.4 0.8
ZMMU A-8210 Suan Phueng, Ra chabu i, Thailand R M 33.9 9.5 11.2 5.2 1 2.4 2.5 4 3.6 3.2 0.7
ZMMU A-8048 Suan Phueng, Ra chabu i, Thailand R M 33.8 10.2 10.8 4.5 1.1 2.1 2.2 4.4 3.6 3 0.8
ZMMU A-8050 Suan Phueng, Ra chabu i, Thailand R M 34.3 10.8 11.3 4.6 1.1 2.5 2.7 4.6 3.4 3 0.9
ZMMU A-8034 Chumphon, Thailand P M 34.1 9.4 11.6 4.2 1.2 2.1 2.2 4.1 3.4 3 0.8
ZMMU A-8039 Chumphon, Thailand P M 30.4 9.7 9.9 4.5 1.1 2.1 2 3.7 3.2 3.4 0.6
ZMMU A-8059 Suan Phueng, Ra chabu i, Thailand R M 32.6 9.8 10.7 4.6 1.2 2.1 2.2 4.4 3.1 3.3 0.7
ZMMU A-8054 Suan Phueng, Ra chabu i, Thailand R M 34 10.1 11.2 4.7 1.1 2.2 2.2 4.3 3.5 3.1 0.7
ZMMU A-8051 Suan Phueng, Ra chabu i, Thailand R M 30.8 8.8 9 3.5 1.1 2 2 3.8 3.1 3 0.5
ZMMU A-8047 Suan Phueng, Ra chabu i, Thailand R M 33.8 9.7 11.1 4.5 1.1 2.3 2.2 4.4 3.6 3.3 0.8
ZMMU A-8056 Suan Phueng, Ra chabu i, Thailand R M 33.4 9.6 10.6 4.9 1.4 2.2 2.4 4.3 3.6 3.2 0.8
ZMMU A-8058 Suan Phueng, Ra chabu i, Thailand R M 33.5 9.8 11.2 4.7 1.5 2.4 2.5 4.5 3.7 3.2 0.8
ZMMU A-8052 Suan Phueng, Ra chabu i, Thailand R M 31.8 9.6 10.7 4.7 1.2 2.2 2.5 4.3 3.7 3.2 0.8
ZMMU A-8053 Suan Phueng, Ra chabu i, Thailand R M 31.2 9.1 10.5 4.7 1.2 2.2 2.5 4.2 3.4 3.1 0.8
ZMMU ID Sex ESD TD CRL1 PGW PGL FLL FHL FAW HAL F1 F2 F3 F4 IMC
Inge oph ynus pa us
ZMMU A-8029 F 5.2 4.3 7.7 3.5 4.6 31.6 24.5 3.6 12.5 5.8 4 7.7 2.5 1.9
ZMMU A-8021 F 4.8 3.2 6.9 3.9 3.8 33.3 24.6 4.3 12.5 7.5 4.1 8.9 3.5 1.3
ZMMU NAP-09684 F 5.2 3.7 7.8 2.9 3.8 31.2 23.5 3.4 11.8 6.6 3.8 7.9 2.7 0.9
ZMMU A-8025 M 4 2 6.1 2.4 3 23.2 17.5 3 8.6 4.8 2.2 5.6 1.8 1.4
ZMMU A-8026 M 4 1.9 6 2.4 2.3 24.2 17.9 3.8 8.7 4.5 2.9 6.3 2.2 0.7
ZMMU A-8024 M 4 2 6 2.4 3 23.3 18.3 3.6 9.5 4.9 3.4 7.2 2 0.9
ZMMU A-8022 M 3.4 2.1 6.3 2.6 2.4 23.7 18.4 3.2 9 4.5 2.8 6.3 1.6 0.8
ZMMU A-8023 M 3.7 2.5 6 2.5 2.9 23.4 18.2 3.1 8.6 4.6 3.5 7 2.3 1
ZMMU A-8027 M 3.8 2.2 6 2.5 2.3 25.5 19.1 3.6 9.5 4.6 3.5 6.5 2.1 1.4
ZMMU A-8028 M 4.2 2.4 6.1 2.4 2.8 26 18.9 3.9 9.9 4.9 3 6.5 2.2 1.2
Inge oph ynus сh ysolophus sp. no .
ZMMU A-8035 F 3.9 2.5 5.9 2.6 2.7 24.2 18.6 2.1 9.2 4.5 3 6.4 2.1 1.3
ZMMU A-8030 F 4.4 2.7 5.7 2.9 2.7 22.1 17.7 2.6 8.7 4.6 2.6 5.8 2.2 1.1
ZMMU A-8036 F 3.8 2.4 4.4 2.5 2.4 22.2 17.4 2.9 8.5 4.8 2.7 5.6 2.1 1.1
ZMMU A-8037 F 3.8 3 4.7 2.6 2.6 22.5 16.8 3 8.3 4.6 2.9 5.6 1.9 1.1
ZMMU NAP-09425 F 3.8 3.7 5.7 2.7 4.4 25.3 19.4 3.5 9.4 3.3 4.7 5.8 2.3 1.3
ZMMU A-8211 F 3.6 2.3 5.2 2.6 5.2 24.9 19.5 2.7 9.3 3.9 3.1 6.3 2.6 1.2
ZMMU A-8061 F 4 2.5 6.5 3.9 3.9 23.3 18.8 2.7 9.8 5.4 3.7 7 2.1 1.5
ZMMU A-8031 F 4.4 2.7 5.9 2.8 2.6 27.9 20.9 2.9 9.9 5.1 4 7.5 2.3 1.4
ZMMU A-8033 F 4.3 2.6 5.6 3.3 3.6 27.5 20.6 3.2 10.1 5.4 3.7 6.6 2.3 1.4
ZMMU A-8032 F 4.5 2.4 5.9 3 3.5 28.7 22.2 2.9 10.4 6 4 7.1 2.6 1.5
ZMMU A-8057 M 3.7 2.3 5.7 2.4 2.6 20.4 16 2.7 7.7 4 2.2 5.6 1.6 0.9
ZMMU A-8055 M 3.9 2.5 6.3 3.7 3.8 21.3 17.4 3.7 8.7 4.1 2.7 5.7 2.1 1
ZMMU A-8020 M 3.8 2.3 5.3 2.9 2.8 20.8 16.8 3.4 8.7 4.8 2.3 5.8 1.9 1.2
ZMMU A-8049 M 3.8 2.5 5.6 2.5 2.9 22.2 16.3 3.2 8.6 4.7 2.6 5.7 2 1
He pe ozoa 38: 271–297 (2025)
he pe ozoa.penso .ne
277
The da a we e aken om he da abases Wo ldclim 1
(h ps://www.wo ldclim.o g), GlobCo e 2009 (h ps://
due.es in.esa.in /page_globco e .php), Global A idi y
and PET (h ps://csido in o.wo dp ess.com), and Pe cen
ee co e age (h ps://gi hub.com/globalmaps/gm_ e_
2). Fo con enience, each laye was abb e ia ed as ol-
lows: BIO1 = annual a e age empe a u e, BIO2 = daily
a e age ange (mon hly a e age (maximum empe a u e
- minimum empe a u e)), BIO3 = iso he mali y (BIO2/
BIO7) (×100), BIO4 = empe a u e seasonali y (s an-
da d de ia ion × 100), BIO5 = maximum empe a u e
o he wa mes mon h, BIO6 = minimum empe a u e
o he coldes mon h, BIO7 = annual empe a u e ange
(BIO5-BIO6), BIO8 = a e age empe a u e o he we es
qua e , BIO9 = a e age empe a u e o he d ies qua -
e , BIO10 = a e age empe a u e o he wa mes qua -
e , BIO11 = a e age empe a u e o he coldes qua e ,
BIO12 = annual p ecipi a ion, BIO13 = p ecipi a ion o
he we es mon h, BIO14 = p ecipi a ion o he d ies
mon h, BIO15 = seasonali y o p ecipi a ion (coe icien
o a ia ion), BIO16 = p ecipi a ion o he we es qua -
e , BIO17 = d ies qua e p ecipi a ion, BIO18 = wa m-
es qua e p ecipi a ion, BIO19 = coldes qua e p e-
cipi a ion, A idIndex = a idiza ion index, Ba en = soil
ZMMU ID Sex ESD TD CRL1 PGW PGL FLL FHL FAW HAL F1 F2 F3 F4 IMC
ZMMU A-8060 M 3.9 2.6 6.1 3.1 3.1 22.7 17.4 3.4 8.7 4.5 2.7 5.9 1.9 1.1
ZMMU A-8210 M 3.6 2.1 5.7 2.5 3.7 21.3 17.7 2.2 8.9 3.9 2.7 5.7 2.1 0.9
ZMMU A-8048 M 3.9 2.4 5.9 2.6 3.6 22 17 3.3 8.9 4.4 2.8 6.5 1.9 1.4
ZMMU A-8050 M 3.6 2.8 5.3 2.6 3.2 22.2 18.2 3.5 9.4 4.3 2.9 5.2 2 1.4
ZMMU A-8034 M 3.8 2.4 5.1 3.2 3.1 23.8 18.3 3.1 8.8 4.1 2.8 6 1.8 1.2
ZMMU A-8039 M 3.5 2.2 5.3 2.2 2.7 22 16 2.8 7.5 3.8 2.7 5.3 2 1.2
ZMMU A-8059 M 3.4 2.4 5.3 2.4 3 23.7 17.6 3.3 8.7 4.3 2.7 6 2 1.1
ZMMU A-8054 M 3.5 2.3 5.6 2.4 3.6 23.9 18.1 2.8 8.6 4.2 2.9 6 2 1
ZMMU A-8051 M 3.6 2.2 5.3 2.5 2.6 20.7 16 2.6 7.8 4.1 2.2 5.2 1.7 0.9
ZMMU A-8047 M 3.9 2.4 5.9 2.7 2.9 22.6 17.4 3.3 8.9 4.4 2.8 6.4 1.8 1.4
ZMMU A-8056 M 3.9 2.6 5.7 3.1 2.8 22.4 17.2 3.1 8.6 4.6 2.7 5.8 1.9 1.2
ZMMU A-8058 M 3.9 2.5 5.8 3.2 3.4 22.3 17.4 3.4 8.7 4.6 2.5 5.9 1.9 1.2
ZMMU A-8052 M 3.8 2.6 5.7 3.2 3.1 22.4 17.5 3.1 8.9 4.7 2.6 5.9 1.9 1
ZMMU A-8053 M 3.7 2.4 5.5 2.7 2.5 20 16.1 3.2 7.7 3.9 2.6 5.2 1 1
ZMMU ID Sex OMC FEL TIL TFL FOL T1 T2 T3 T4 T5 IMT OMT CRL2
Inge oph ynus pa us
ZMMU A-8029 F 3.2 21.2 21.5 28.4 16.5 4.9 2.8 4.4 7.8 2.2 2 1.4 3
ZMMU A-8021 F 2.8 21.3 21.6 29 16.6 4.5 3 4.7 7.8 2 2.1 1.6 2.9
ZMMU NAP-09684 F 2.3 19.8 20.7 29 16.9 4.9 4 6.2 8.9 2.5 1.9 1.7 3
ZMMU A-8025 M 2.2 14.2 14.5 20 10.9 3.1 2 2.9 5.2 1.9 1.1 1.1 2
ZMMU A-8026 M 1.9 15.1 14.5 21.6 11.7 3.1 2.3 3.3 6 1.9 1.4 1.1 2.6
ZMMU A-8024 M 2 16.3 15.2 22.5 12.3 3.4 2.9 3.6 6 1.8 1.2 1.3 2
ZMMU A-8022 M 2.1 14.3 14.8 20.9 11.8 2.4 2.2 4.1 5.4 1.5 1.2 1.1 1.9
ZMMU A-8023 M 2 14.9 15.1 21.2 12.5 3.2 2.7 3.9 5.5 1.7 1.2 1.4 2
ZMMU A-8027 M 2 16.3 15.3 22.4 12.7 3.5 2.1 3.4 6.3 1.8 1.8 1.4 1.9
ZMMU A-8028 M 2.1 17 16.2 22.7 13.3 3.7 2.5 4.1 6.8 2 1.4 1.2 2.1
Inge oph ynus сh ysolophus sp. no .
ZMMU A-8035 F 1.8 14 14 21.8 13.1 3 2.6 3.4 6 1.8 1.1 1.1 2.3
ZMMU A-8030 F 1.4 13.8 14.7 21.5 13 3.4 2.2 4.2 6.7 2 1.3 1.1 2.2
ZMMU A-8036 F 1.9 13.5 13.8 21.1 12.5 3.3 2.2 3.6 6 1.9 1.3 1.3 2.2
ZMMU A-8037 F 2 13.9 13.8 20.7 12 3 2.4 3.6 6.1 1.9 1.2 1.1 2
ZMMU NAP-09425 F 2.2 14.6 15.9 23 15.2 4 2.8 3.9 6.3 2.1 1.4 1.3 2.7
ZMMU A-8211 F 2.2 15.8 15 22.2 12.2 3.8 2 3.4 5.5 1.1 1.5 1.2 2.3
ZMMU A-8061 F 2.1 15.2 14.5 21.5 13.1 4.2 2.4 3.6 6.6 1.9 1.3 1.3 2.5
ZMMU A-8031 F 2.2 17.8 16.6 24.7 14.8 4 2.9 4.4 6.6 1.8 1.3 1.3 3.1
ZMMU A-8033 F 2.1 16.3 16.3 24.3 14.4 3.5 2.6 4.2 7.5 1.8 1.4 1.4 2.4
ZMMU A-8032 F 2.4 17.6 17.6 24.8 14.5 4.3 3 4.6 7.8 2.3 1.6 1.6 2.8
ZMMU A-8057 M 1.7 14.3 12.1 18.6 11 3.2 2 3.3 5.3 1 1.2 1.2 2
ZMMU A-8055 M 1.8 13.6 14.4 21 12.4 3.7 2.1 3.6 6.4 2.2 1.5 1.3 2.3
ZMMU A-8020 M 1.8 13.3 13.5 20.7 12.2 3.4 2.7 3.6 6.5 1.8 1.4 1.4 1.9
ZMMU A-8049 M 1.9 13.4 14.1 20.7 12 3.3 2.1 3.8 5.9 1.6 1.2 1.4 2.1
ZMMU A-8060 M 1.9 13.6 13.7 20.7 12.2 3.5 2.3 3.8 6.3 1.6 1.2 1.1 2.1
ZMMU A-8210 M 2.1 13.4 14 20 12.3 3.4 2.4 3.8 6.6 1.7 1.2 1 2.1
ZMMU A-8048 M 2.2 13.5 14.5 20.8 12.5 3.9 2.3 3.8 5.4 1.6 1.4 1.1 1.9
ZMMU A-8050 M 2 15.2 15.2 21.8 13.2 3.4 2.6 3.7 5.9 1.6 1.3 1.1 2.2
ZMMU A-8034 M 2 14.2 14.5 21.6 13.3 4 2.6 3.3 5.5 1.5 1.3 1.3 2.5
ZMMU A-8039 M 2 11.9 12.8 20.2 11.4 3.5 2.4 3.2 5.3 1.6 1.2 1.1 2
ZMMU A-8059 M 1.9 15.1 14.6 21.8 12.6 3.3 2.4 3.8 5.8 1.6 1.3 1.3 1.9
ZMMU A-8054 M 1.8 15.3 15 20.1 12.1 3.1 2.2 3.7 5.9 1.6 1.3 1.2 1.9
ZMMU A-8051 M 1.8 13.3 12.9 18.8 11 3 2 3.3 5.3 1.5 1.2 1.1 1.8
ZMMU A-8047 M 2 15 14.1 21.1 12.5 3.7 2.4 3.9 5.6 1.6 1.4 1.1 1.9
ZMMU A-8056 M 1.9 14.2 14 20 11.9 3.3 2.1 3.8 6.3 1.7 1.2 1.1 1.9
ZMMU A-8058 M 2 13.8 13.9 20.5 12.2 3.3 2.1 3.7 6.1 1.7 1.4 1.2 2.2
ZMMU A-8052 M 1.9 13.9 13.9 20.5 12.4 3.1 2.2 3.9 6.3 1.6 1.1 1.1 2.1
ZMMU A-8053 M 1.9 12.6 12.8 18.2 11 3.1 1.8 3.2 5 1.2 1.3 1.1 2.1
he pe ozoa.penso .ne
Dmi iy V. A khipo e al.: A new species o Inge oph ynus om Thailand278
e ili y, cul i a ed land cul i a ion, Ele a ion = al i ude,
Exposi ion = slope exposu e, Slope = d ainage angle,
Slope 3 cells = wa e shed angle squa ed 3 × 3 cells,
Mixed = mixed land co e , Homogenei y = homogenei y
o land co e , Sh ub = pe cen age o sh ub co e , He ba-
ceous = pe cen age o g ass co e , Dec-b oad = decidu-
ous b oadlea o es , E e -b oad = e e g een b oadlea
o es s, E e -decid = deciduous-b oadlea o es s. The
p og am ENMTools 1.3 (Wa en e al. 2010) was used o
il e SDM da a and exclude co ela ed a iables.
Models we e assessed by compu ing he a ea unde
he cu e (AUC), which was used o es ima e he ela i e
con ibu ion o a iables o each model. The eliabili y
o he model was e alua ed based on he ecei e ope a -
ing cha ac e is ic cu e o aining and es da a, as well
as hei AUC alues. We used he andom es pe cen age
ool se ings o s a is ical e alua ion o he model, wi h
75% o he andomly selec ed locali ies used o se up
he model and 25% o he poin s used o es i . A model
wi h AUC es alues g ea e han 0.75 was conside ed o
be use ul, and abo e 0.90, e y good (Swe s 1988; Eli h
2002). The inal maps we e designed using he QGIS
Desk op 3.28 so wa e (QGIS De elopmen Team 2021).
The esul ing dis ibu ion image ob ained shows in colo s
he calcula ed p obabili y ha he condi ions a e a o -
able o he species o inhabi ( ed: a high p obabili y o
a o able condi ions; shades o blue: unlikely condi ions).
Bioacous ic analysis
Ad e isemen calls o wo popula ions o he Inge oph y-
nus pa us species complex we e eco ded using a po a-
ble digi al audio eco de , he Zoom H5 (ZOOM Co po a-
ion, Tokyo, Japan), in s e eo mode wi h 48 kHz sampling
equency and 16-bi p ecision. The eco dings we e aken
in Khao K a Jom M ., Suan Phueng, Ra chabu i, Thailand
(locali y 3, Fig. 1; on 10 Feb ua y 2022 a 19.00 h and a 24
°C), and in Gunung Je ai M ., Kedah, Peninsula Malaysia
(on 30 Janua y 2010 a 20.00 h and a 25 °C). The ambien
empe a u e was measu ed a he calling si e immedia e-
ly a e he audio eco ding wi h a digi al he mome e ,
he KTJ TA218A Digi al LCD The mome e -Hyd ome e .
Males we e obse ed calling om he banks o slow- low-
ing o es s eams. Calls we e analyzed using Ra en P o
1.6.5 (K. Lisa Yang Cen e o Conse a ion Bioacous ics
a he Co nell Lab o O ni hology 2024).
Audio spec og ams we e calcula ed wi h a as Fou ie
ans o m (FFT) o 512 poin s, 90% o e lap, and 135 Hz
g id spacing, using he Hanning window. The e minolo-
gy o call analysis and desc ip ion using a call-cen e ed
app oach (de ining unin e up ed uni s as calls whene e
hey a e sepa a ed by long silen in e als) ollows Koe-
hle e al. (2017). In o al, we measu ed eigh eco dings
o calls, including 188 calls om h ee males o he I. pa -
us species complex om wo locali ies (Suppl. ma e ial
1: able S5). Due o he poo quali y o he eco dings, we
we e able o measu e only eigh empo al pa ame e s o
he calls: du a ion o call se ies, in e als be ween call se-
ies, numbe o calls pe se ies, call du a ion, in e als be-
ween successi e calls wi hin se ies, du a ion o call ise
(CRT) and call all (CFT), numbe o pulses pe call, and
one powe pa ame e o calls: peak equency (Fpeak).
Skele al mo phology
To s udy he skele al mo phology o Inge oph ynus c .
pa us om wes e n Thailand, we examined X- ay p ojec-
ions o wo specimens: ZMMU A-8059 (adul male om
Ra chabu i P o ince) and ZMMU A-8032 (adul emale
om Chumphon P o ince) and pe o med a de ailed mi-
c o-CT scanning. The scans we e pe o med on a SkyScan
1272 mic o omog aph (B uke , Bille ica, USA) equipped
wi h a Hamama su L10101-67 sou ce (Hamama su Pho-
onics, Hamama su, Japan) and a Ximea xiRAY16 came a
(Ximea GmbH, Müns e , Ge many) a he Biological Fac-
ul y o Moscow Uni e si y. The specimens we e scanned
a a sou ce ol age o 100 kV and a sou ce cu en o 100
µA wi hou an X- ay il e . The samples we e o a ed 360°
a ound he e ical axis wi h a o a ion s ep o 0.1, wi h 4
ames a e aging. F om he ob ained X- ay p ojec ions,
s acks o i ual c oss sec ions h ough he specimens’
skele al s uc u es we e econs uc ed wi h he so wa e
NRecon® (B uke mic o-CT, Kon ich, Belgium). The e-
sul ing s ack consis ed o 3599 images wi h a 4504x4504
esolu ion and a 5.42 um pixel size. This s ack o imag-
es was impo ed in o he h ee-dimensional isualiza ion
so wa e package A izo 8.1 o subsequen p ocessing and
isualiza ion o os eological ai s. The ob ained scans
we e deposi ed in Mo phoSou ce (h ps://www.mo pho-
sou ce.o g/conce n/media/000771864). Os eological e -
minology ollowed T ueb (1973) and P egil (1981).
Resul s
Phylogene ic in e ence om m DNA
da a
Ou m DNA-based genealogy o he genus Inge oph ynus
(Fig. 2) includes sequences o en ou o he 13 cu en ly
ecognized nominal species o he genus (Suppl. ma e ial
1: able S2) wi h he excep ion o I. kumqua , I. cla ige ,
and I. philippinicus. No ably, we p o ide gene ic da a o
he species I. quad ipo ca us o he i s ime. The esul -
ing ee is based on he analyses o a con inuous m DNA
agmen ha includes he 12S RNA, 16S RNA, and
ND1 gene sequences (wi h a o al leng h o up o 4,479
bp). BI and ML phylogene ic analyses esul ed in almos
iden ical ee opologies, wi h he monophyly o he ge-
nus Inge oph ynus s ongly suppo ed in bo h analyses
(100/1.0; he ea e , alues co espond o he ML UFBS/
BI PP alues o nodal suppo , espec i ely; Fig. 2).
Ou ee shows a opology ha sligh ly di e s om
opologies in p e ious s udies (Chan and G isme 2019;
He pe ozoa 38: 271–297 (2025)
he pe ozoa.penso .ne
279
Liu e al. 2025). Acco ding o ou da a (Fig. 2), he species
o he genus Inge oph ynus o m wo ecip ocally mono-
phyle ic g oups: he Sunda clade (93/0.99) and he Indo-
chinese clade (100/1.0). The Indochinese clade includes
ou species om mainland Sou heas Asia and sou he n
China—I. galea us, I. wangyingyongi, I. ledongensis, and
I. mac o is. No ably, I. galea us was ound o be pa aphyle ic
wi h espec o I. wangyingyongi and I. ledongensis: he la -
e wo species o med a clade (95/0.99) placed wi hin he
I. galea us adia ion wi h high nodal suppo (100/1.0). We
also epo deep di e gence wi hin I. galea us and I. mac-
o is (Fig. 2), wi h unco ec ed gene ic p-dis ance in he
16S RNA gene wi hin hese species eaching up o 2.95%
and 3.44% o subs i u ions, espec i ely (Suppl. ma e ial 1:
Figu e 2. Phylogene ic ela ionships among he Inge oph ynus pa us species complex and o he ela ed Inge oph ynus species
based on he m DNA agmen including he 12S RNA, 16S RNA, and ND1 gene sequences (up o 4,479 bp). Fo ouche speci-
men in o ma ion and GenBank accession numbe s, see Suppl. ma e ial 1: able S2. Numbe s a ee nodes co espond o ML UFBS/
BI PP suppo alues, espec i ely. Pho og aphs by P. Pawangkhanan and N.A. Poya ko .
he pe ozoa.penso .ne
Dmi iy V. A khipo e al.: A new species o Inge oph ynus om Thailand286
idges b igh o ange (Fig. 8C); pa o oids and pos e i-
o ly ex ending do sola e al wa s da k beige, en ally
coun e shaded wi h da k b own (Fig. 9A), o ming a bo -
de be ween he ligh e b ownish do sum and da k b own
lanks; da k b own c ossba s on he limbs excep o he
b achia; he en e ligh b own o g ayish, and en al
wa s ligh beige (Fig. 8B). The pupil is ho izon al and
black, while he i is is also black, ea u ing dense gold-
en e icula ions on he do sal and en al sides and cop-
pe e icula ions medially; he pupil is edged wi h a hin
golden line (Fig. 8C). A e p ese a ion in e hanol o
ou yea s, all aspec s o he colo pa e n emain; pa e ns
Figu e 8. Holo ype o Inge oph ynus ch ysolophus sp. no . om Wa Tham Sanook, Tha Sae, Chumphon, Thailand, in li e—spec-
imen ZMMU A-8030 (adul emale). Do sal aspec (A); en al aspec (B); la e al iew o head (C); do sal iew o head (D); ola
iew o he le hand (E); plan a iew o he igh oo (F). Pho og aphs by N.A. Poya ko . Scale ba s: 5 mm.

He pe ozoa 38: 271–297 (2025)
he pe ozoa.penso .ne
287
o colo a ion o limbs and body a e s ill isible bu no
as conspicuous as in li e; b igh o ange colo s on c anial
c es s and colo a ion o he i is comple ely aded.
Va ia ion. The indi iduals in he ype se ies and he
e e ed specimens a e all e y simila in ex e nal appea -
ance. Indi idual di e ences in size and body p opo ions
a e p esen ed in Table 1. Males a e signi ican ly smalle
han emales (p < 0.05): mean male body leng h 33.0 ± 1.3
mm (SVL = 30.3–35.7 mm, n = 18); mean emale body
leng h 36.8 ± 2.2 mm (SVL = 34.0–42.4 mm; n = 10). Fig.
10 displays he a ia ion in do sal colo a ion o he pa a-
ypes. The e we e no signi ican di e ences in he colo -
a ion o male pa a ypes and emales, excep ha males had
a da ke h oa colo a ion. A emale specimen, ZMMU
A-8031, had a b igh eddish-b own backg ound do sal
colo a ion, ma kedly di e en om he dulle b ownish
colo a ion o all o he specimens examined (Fig. 10).
The e is a ce ain a ia ion in he deg ee o de elopmen
o da k do sal ma kings among he indi iduals: emales
ZMMU A-8032, A-8035, and A-8036 had con as ing
black blo ches in scapula and sac al a eas, while males
ZMMU A-8034 and ZMMU A-8041 and emale ZMMU
A-8037 had almos no da k ma kings on he do sum (Fig.
10). In gene al, males showed dulle do sal pa e ns wi h
ain bo de s and less con as ing da k ma kings.
Tadpole mo phology. A de ailed desc ip ion o
he la al mo phology o Inge oph ynus ch ysolophus
sp. no . om Thailand was p esen ed by Meewa ana
(2022: 30–31) ( e e ed o as I. pa us in his wo k).
Tadpoles o he new species ha e an o al-shaped head-
body, 2/3 longe han wide; eyes wi h do sal o ien a ion;
in e na ial dis ance comp ising 2/3 o in e o bi al dis-
ance; spi acle sinis al; mou h-snou dis ance equal o
snou -eye dis ance; en median; ail b oad wi h weak
ail muscula u e; do sal and en al ail ins beginning a
he ail base, bo h subequal in dep h; and ail ip ounded
(Meewa ana 2022). O al disc sub e minal, wi h en al
o ien a ion, small papillae in mou h co ne , labial oo h
ow o mula 2(2)/3; beaks black wi h se a ed edges.
Tadpoles each 20.0–25.0 mm in leng h; he head-body
is black in do sal and la e al aspec s and anslucen in
en al aspec , wi h in es ines being isible on he pos e-
io hal o he head-body leng h; ail muscles blackish,
ail ins anspa en (Meewa ana 2022).
Figu e 9. Inge oph ynus ch ysolophus sp. no . in li e in si u. A. Holo ype ZMMU A-8030 (adul emale) om Wa Tham Sanook,
Tha Sae, Chumphon, Thailand; B. Pa a ype ZMMU A-8034 (adul male) om Wa Tham Sanook, Tha Sae, Chumphon, Thailand;
C. ZMMU A-8020 (adul male) om Pa Klok, Phuke , Thailand; D. ZMMU A-8059 (adul male) om M . Khao K a Jom, Suan
Phueng, Ra chabu i, Thailand. Pho og aphs by N.A. Poya ko .
he pe ozoa.penso .ne
Dmi iy V. A khipo e al.: A new species o Inge oph ynus om Thailand288
Os eological desc ip ion. The ollowing desc ip ion o
adul skull mo phology is based on he omog aphic da a
ob ained o he adul male (ZMMU A-8059, pa a ype)
and he adul emale (ZMMU A-8032, pa a ype) (Fig. 11).
C anium. O e all, he c anium o Inge oph ynus ch ys-
olophus sp. no . is gene ally well ossi ied; he highly ube -
culous skin appea s o be qui e dense op ically in his spe-
cies and is isible in ou econs uc ions, concealing pa s
o he skull; he denses egions a he ip o he snou and on
he uppe eyelidswe e cu o he scans; some elemen s on
he c anial oo show aces o hype ossi ica ion, while he
o ic egion seems unde ossi ied; he skull shape is almos
iangula in do sal (Fig. 11A, D) and en al iews (Fig.
11B, E) and close o apezoid in la e al iew (Fig. 11C, F).
Snou dis inc ly unca e in do sal (Fig. 11A, D) and en al
(Fig. 11B, E) iews, no p o uding beyond he uppe jaw.
The hyob anchial appa a us is mos ly ca ilaginous and hus
in isible in ou econs uc ion; he only isible elemen is
he well-ossi ied pai ed pos e omedial p ocess o he hyoid.
P emaxilla. The p emaxilla is a pai ed bone, sligh ly
a cua e do sally, oo hless (Fig. 11B, E). The p emaxilla
do sally con ibu es o he in e nasal on anelle (ca um
in e nasale), en ally con ibu es o he an e omedial e-
nes a, and do somedially con ac s he en omedial pa
o he ex e nal na es. The ala y p ocess o he p emaxilla
is o ien ed an e odo sally. The an e io su ace o he p e-
maxilla y bones is sligh ly g anula in emale and smoo h
in male (Fig. 11B, E). The p emaxilla con ac s he maxil-
la y bones pos e ola e ally and con ibu es o he an e io
pa o he nasal ca i y.
Maxilla. The pai ed maxilla is a oo hless bone, un-
ning la e ally om he nasal capsule o he le el o he
o ic egion (Fig. 11B, E). The maxilla con ac s he p e-
maxilla y an e io ly, he nasal do somedially, he pala ine
medially, he p e ygoid pos e omedially, and he quad a-
ojugal pos e io ly.
Nasal. The pai ed nasals o m he do sal pa o he
snou ; hey con ac he maxilla y en ola e ally wi h he
Figu e 10. Va ia ion in do sal colo a ion o he pa a ypes and e e ed ma e ial o Inge oph ynus ch ysolophus sp. no . om Wa
Tham Sanook, Tha Sae, Chumphon, Thailand, in li e. Pa a ypes: adul male, ZMMU A-8034, ZMMU A-8039; adul emale, ZMMU
A-8031–8033, ZMMU A-8035–8037. Re e ed ma e ial: subadul opo ypic males, ZMMU A-8038, RIM NAP-11589; subadul
opo ypic emales, ZMMU A-8040–8044; subadul s (sex no de e mined), ZMMU A-8045, ZMMU A-8046, RIM NAP-11591, RIM
NAP-11593. Pho og aphs by N.A. Poya ko .
He pe ozoa 38: 271–297 (2025)
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289
Figu e 11. C anial mo phology o Inge oph ynus ch ysolophus sp. no .: adul emale, pa a ype ZMMU A-8032 (A–C); adul male,
pa a ype ZMMU A-8059 (D–F), isualized ia mic o-CT scanning. Skulls a e shown in do sal (A, D), en al (B, E), and la e al
(C, F) aspec s. Fo he la e al iews only, he le hal o he c anium is shown. Abb e ia ions: nas – nasal, sphen – sphene hmoid,
p sph – pa asphenoid, p m – p emaxilla, smx – sep omaxilla, pal – pala ine, om – ome , max – maxilla, p – p e ygoid, p
– on opa ie al, qj – quad a ojugal, soc – sup ao bi al c es , sq – squamosal, s c – sup a ympanic c es , p o – p oo ic, exoc –
exoccipi al, s – s apes, mnm – men omeckelian, dn – den a y, ansp – angulosplenial. Visualiza ion by D.V. A khipo and V.A.
Go in. Scale ba : 5 mm; all pic u es a e o scale.
he pe ozoa.penso .ne
Dmi iy V. A khipo e al.: A new species o Inge oph ynus om Thailand290
well-p onounced maxilla y p ocess and he sphene h-
moid pos e io ly, which ills he space be ween he nasals
and on opa ie als (Fig. 11A, C). The nasal con ibu es
en ally o he nasal ca i y. Do sal su aces o nasal
bones a e dep essed medially and o e all a e sligh ly sha-
g eened in male and g anula in emale. The pai ed nasals
a e well sepa a ed om each o he , o ming an in e nasal
on anelle medially (Fig. 11A, D).
F on opa ie al. The la ge pai ed on opa ie als o m
he oo o he skull, sepa a ed by he on opa ie al su-
u e ac oss hei leng h (Fig. 11A, D); hey con ac he
sphene hmoid an e io ly and an e o en ally, he exooc-
cipi als pos e io ly, and he p oo ics pos e odo sola e al-
ly, and a e used wi h he la e pos e ola e ally. The do -
sal su ace o he on opa ie als is smoo h in male and
sligh ly g anula in emale. The do sola e al pa s o he
on opa ie als o m sup ao bi al c es s all ac oss hei
leng h and a e g anula ed in he pos e io hal in male and
along hei en i e leng h in emale.
Sep omaxilla. The pai ed sep omaxilla is a small bone
o a complex shape loca ed in he an e io pa o he
snou (Fig. 11C, F). The sep omaxilla con ibu es o he
ex e nal na es and he an e io wall o he nasal ca i y.
Vome . The ome is a a he small pai ed bone, i a-
dia e in shape, loca ed in he an e io pa o he pala ine
egion, co e ing he c anial base be ween he in e nal na-
es and bea ing no ee h (Fig. 11B, E). A pai o ome s
con ibu es o he medial and do sal walls o he in e nal
na es and o he en al walls o nasal capsules.
Pala ine. The pai ed pala ine is a od-shaped, oo h-
less bone wi h a p ominen en al idge (Fig. 11B, E).
The pala ine is loca ed in he pos e io pa o he pala ine
egion and con ac s he sphene hmoid medially, he max-
illa do sola e ally, and he p e ygoid pos e ola e ally.
Sphene hmoid. The sphene hmoid is a single bone
subcylind ical in shape, o ming he an e io po ion
o he neu oc anium (Fig. 11B, E). I is s uc u ed like
a ypical ca ilaginous bone wi h p onounced inne and
ou e bone laye s. The sphene hmoid con ac s he nasals
an e odo sally, he on opa ie als pos e odo sally, he
pala ines en ola e ally, and he pa asphenoid pos e o-
en ally, and con ibu es o he nasal ca i y an e io ly.
Pa asphenoid. The single pa asphenoid is a la ge
swo d-shaped bone pla e o ming he loo o he c anium
(Fig. 11B, E). The pa asphenoid con ac s he sphene hmoid
an e io ly, he p oo ics pos e odo sally, he p e ygoids pos-
e ola e ally, and he exoccipi als pos e io ly. The pa as-
phenoid is unde -ossi ied la e ally in he emale specimen.
Squamosal. The pai ed hoe-shaped squamosals a e
loca ed a he side o he c anium and a a igh angle
o he jaw a c (Fig. 11C, F). The squamosal con ac s he
p oo ic do somedially, he quad a ojugal en ally, and
app oaches he p e ygoid medially. The squamosal is
densely ube culous do sally, wi h he o ic amus o ming
he sup a ympanic c es on i s do sola e al su ace (Fig.
11A, D). The medial pa o he o ic amus appea s o be
unde ossi ied in he emale specimen. Ven al and zygo-
ma ic ami a e well de eloped, ossi ied, and smoo h.
P e ygoid. The pai ed p e ygoids a e i adia e in
shape, wi h each ha ing h ee b anches ( ami; Fig. 11C,
F). The p ocessus o icum (medial amus) is di ec ed do -
somedially, wi h a p ominen exca a ion in he pos e io
iew, con ac ing he p oo ic. The p ocessus pala inum
(an e io amus) is di ec ed an e io ly, wi h a p ominen
la e al idge, and con ac s he maxilla la e ally and he
pala ine an e io ly (Fig. 11B, E). The p ocessus quad a-
um (pos e io amus) o he p e ygoid is di ec ed pos e i-
o ly, con ac ing he quad a ojugal.
Quad a ojugal. The pai ed quad a ojugals a e a he
small bones, loca ed pos e ola e ally on he skull (Fig.
11B, E). The quad a ojugal con ac s he maxilla an e i-
o ly, he squamosal do sally, app oaching he p e ygoid
pos e omedially, and a icula ing wi h he angulosplenial
bone o he lowe jaw pos e o en ally.
P oo ic. The p oo ic is an incomple ely ossi ied pai ed
bone, seemingly spongious, la gely ca ilaginous pos e-
io ly (Fig. 11B, E). The pos e ola e al pa o he do sal
su ace o he p oo ic bea s an exca a ion. The p oo ic
con ac s he squamosal do sola e ally, he p e ygoid en-
ola e ally, he pa asphenoid en ally, and he on opa-
ie al do somedially and is used wi h he la e an e ome-
dially. The p oo ics la gely con ibu e o he o ic capsules.
S apes. The pai ed s apes (columella) a e la gely mine -
alized, sligh ly a ched, and ex ending medially (Fig. 11C,
F). The s apes is o ien ed la e ally and sligh ly an e io ly and
comes close o he p oo ics, o he a ea o he o al window.
Saccula o oconia. The o ic capsule is pa ially illed
wi h calcium ca bona e in he o m o he saccula o o-
conia. The pai ed saccula o oconia a e well mine alized,
subsphe ical in shape, and loca ed deep inside he inne ea .
Exoccipi al. The pai ed exoccipi al is he pos e io -
mos bone o he skull, o ming occipi al condyles and
he o amen magnum (Fig. 11B, E). The wo exoccipi als
sligh ly ouch each o he pos e odo sally and pos e o en-
ally. The exoccipi als show aces o unde ossi ica ion
la e ally, in he o ic egion. The exoccipi al con ac s he
on opa ie al an e odo sally, he pa asphenoid an e o en-
ally, and con ibu es an e ola e ally o he o ic capsule.
Mandible. The lowe jaw is shaped like a solid bony
a ch and comp ises h ee bone elemen s, namely, pai ed
angulosplenials, den a ies, and men omeckelians, he la -
e wo being comple ely used (Fig. 11C, F). The space
be ween den a ies and angulosplenials is likely illed wi h
Meckel’s ca ilage, in isible in ou econs uc ions.
E ymology. The species name “сh ysolophus” is
a La inized adjec i e in he nomina i e singula , mas-
culine gende , de i ed om he Ancien G eek wo ds
“χρυσός” o “ch ysos,” meaning “gold,” and “λόφος” o
“lophos,” meaning “c es ” o “ idge.” The species name
is gi en in e e ence o he cha ac e is ic golden-o ange
colo a ion o sup a ympanic c es s in he new species.
We sugges he ollowing common names o he new
species: Golden-c es ed Dwa Toad (in English), Khang
kok kh ae hua ong (คางคกแคระหัวทอง, in Thai), and Zla-
og ebnis aya shlemonosnaya zhaba (Златогребнистая
шлемоносная жаба, in Russian).
He pe ozoa 38: 271–297 (2025)
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291
Compa isons. Inge oph ynus ch ysolophus sp. no .
can be dis inguished om I. bipo ca us, I. celebensis,
I. cla ige , I. di e gens, I. galea us, I. ledongensis, I. mac-
o is, I. philippinicus, I. quad ipo ca us, and I. wangy-
ingyongi by ha ing small body size (SVL 30–36 mm in
males, 34–42 mm in emales s. 55–70 mm in males,
60–80 mm in emales o I. bipo ca us; up o 130 mm
o I. celebensis; 33 mm in male, 58–69 mm in emales
o I. cla ige ; 28–45 mm in males, 50–55 in emales o
I. di e gens; up o 50 mm in males, 80 mm in emales
o I. galea us; 47–55 in males, 62–64 mm in emales o
I. ledongensis; up o 50 mm in males, 55 mm in emales
o I. mac o is; 52–78 mm in males, 58–86 mm in emales
o I. philippinicus; 48–50 mm in males, 49–62 in emales
o I. quad ipo ca us; and 44.8–53.3 mm in males, 54.3–
57.9 mm in emales o I. wangyingyongi). Inge oph ynus
сh ysolophus sp. no . u he di e s om I. bipo ca us,
I. di e gens, I. galea us, I. ledongensis, I. quad ipo ca us,
and I. wangyingyongi by ha ing pa o oid no con inuous
wi h an oblique ow o conspicuously enla ged wa s ( s.
con inuous). Inge oph ynus сh ysolophus sp. no . can be
u he dis inguished om I. cla ige and I. philippinicus
by ha ing c anial c es s no hickened behind eyes ( s.
c anial c es s dis inc ly hickened immedia ely behind he
eye le el). Inge oph ynus сh ysolophus sp. no . u he
di e s om I. kumqua by ha ing nup ial pads p esen
( s. absen ) and by ha ing i s inge longe han second
( s. second inge longe han i s ).
Finally, Inge oph ynus сh ysolophus sp. no . supe i-
cially mos closely esembles i s sis e species I. pa us s.
s .; howe e , he new species can be eadily dis inguished
om he la e by ha ing he ollowing sui e o mo pholog-
ical cha ac e s: smalle body size in bo h sexes (SVL 30.3–
35.7 mm [a g. 33.0 mm] in males, 34.0–42.4 mm [a g.
36.8 mm] in emales s. 33.1–36.7 mm [a g. 34.8 mm] in
males, 44.5–48.5 mm [a g. 47.0 mm] in emales); sligh ly
highe HL/HW a io in bo h sexes (0.81–0.98 [a g. 0.90] in
males, 0.80–0.96 [a g. 0.87] in emales s. 0.83–0.89 [a g.
0.85] in males, 0.78–0.87 [a g. 0.83] in emales); highe
a io TD/ED in males (0.53–0.64 [a g. 0.58] s. 0.44–0.51
[a g. 0.48]), bu lowe in emales (0.51–0.77 [a g. 0.60]
s. 0.53–0.81 [a g. 0.68]); lowe a io TIL/SVL in emales
(0.39–0.43 [a g. 0.41] s. 0.44–0.45 [a g. 0.45]); lowe a-
io IMT/T1 in bo h sexes (0.33–0.42 [a g. 0.38] in males;
0.31–0.40 [a g. 0.37] in emales s. 0.35–0.51 [a g. 0.42] in
males, 0.35–0.51 [a g. 0.42] in emales), and by he p esence
o b igh o ange colo a ion o he c anial c es s ( s. b own).
Ad e isemen call. The male ad e isemen call o
Inge oph ynus ch ysolophus sp. no . is desc ibed in de-
ail in he Resul s sec ion (see abo e); he call pa ame e s
a e p esen ed in Suppl. ma e ial 1: able S5; he wa e-
o m and he sonog am o he male ad e isemen call o
he new species a e p esen ed in Fig. 6B. The male ad e -
isemen call o he new species di e s om he call o
I. pa us s. s . by a lowe peak equency (1,292–1,378
Hz s. 2,438–2,625 Hz), ewe sho (0.02–0.05 s s.
0.05–0.07 s) calls in he se ies (8–10 s. 13–15), and by
ha ing ewe pulses in he call (5–6 s. 6–9).
Dis ibu ion. Inge oph ynus ch ysolophus sp. no . is
eliably known om cen al, eas e n, wes e n, and sou h-
e n Thailand (Mae Hong Son, Tak, Kamphaeng Phe ,
U hai Thani, Kanchanabu i, Ra chabu i, Phe chabu i,
P achuap Khi i Khan, Chumpon, Ranong, Su a Thani,
Phang Nga, Phuke , K abi, Nakhon Si Thama a , T ang,
Songkhla, Sa un, T a , and Chan habu i p o inces); he
adjacen pa s o sou he n Myanma (Tanin ha yi Region
and Yangon S a e); and sou hwes Cambodia (Ca damom
Moun ains) (Fig. 1). Acco ding o he esul s o he species
dis ibu ion modeling (Fig. 7B), he occu ence o he new
species is expec ed in he no he nmos Peninsula Ma-
laysia (Pe lis S a e); u he s udies a e equi ed o cla i y
he ex en o i s dis ibu ion in he Thai-Malay Peninsula.
Na u al his o y no es. All indi iduals o he new
species we e collec ed du ing he nigh om swampy
a eas along he slow-mo ing, shallow s eam wi hin a
closed-canopy e e g een mon ane o lowland opical
o es . B eeding and la al de elopmen ake place in
ain pools o side pools along he s eam banks, ypically
wi h sandy o sil y bo oms and nume ous dead lea es
and o he plan s accumula ed on he bo om (Meewa a-
na 2022). In Thailand, he new species occu s in a ious
habi a s, om undis u bed mon ane opical o es s o
hea ily dis u bed bamboo o es s and ubbe plan a ions.
In Suang Phueng (Ra chabu i P o ince), he new species
was eco ded in syn opy wi h h ee o he bu onid species:
Ansonia ka en Suwannapoom, G isme , Pawangkhanan ,
Naiduangchan, Yushchenko, A khipo , Wilkinson & Po-
ya ko , 2021; Ph ynoidis aspe (G a enho s , 1829); and
Du aph ynus c . melanos ic us (Schneide , 1799).
Conse a ion s a us. A p esen , he new species is
known om mul iple loca ions ac oss sou he n, cen al,
wes e n, and eas e n Thailand, sou he n Myanma , and
sou hwes Cambodia (Fig. 7B). The po en ial h ea s o
his species a e habi a loss and deg ada ion due o in-
ensi ied logging and de o es a ion. We p opose he new
species o be classi ied as Leas Conce n (LC) acco ding
o he IUCN’s Red Lis ca ego ies (IUCN 2019).
Discussion
Ou upda ed m DNA-based genealogy is la gely consis-
en wi h p e ious phylogene ic s udies o he genus In-
ge oph ynus (F os e al. 2006; Chan and G isme 2019;
Liu e al. 2025). The genus Inge oph ynus is con i med
as a monophyle ic g oup, including wo majo clades: he
Indochinese clade (I. galea us + I. mac o is) and he Sun-
daland clade (all emaining species) (Fig. 2). In pa icula ,
we o he i s ime p o ide gene ic da a o I. quad ipo -
ca us, which is eco e ed as a sis e species o I. bipo ca-
us (Fig. 2). The I. pa us species complex was ound o
be a membe o he Sundaland clade o Inge oph ynus bu
is only dis an ly ela ed o I. bipo ca us, as hypo hesized
ea lie (Inge 1966; Inge 1972). Fu he mo e, ou phy-
logene ic econs uc ion, coupled wi h gene ic dis ances,
indica es deep in aspeci ic di e si ica ion wi hin such

he pe ozoa.penso .ne
Dmi iy V. A khipo e al.: A new species o Inge oph ynus om Thailand292
wide- anging complexes as I. di e gens and I. galea us
(Fig. 2, Suppl. ma e ial 1: able S6), which equi es u -
he in eg a i e axonomic s udies.
The axonomy o he Inge oph ynus galea us com-
plex equi es a special commen . Recen ly, Liu e al.
(2025) e ised his complex and desc ibed he popula ion
om Guangdong P o ince o China p e iously epo -
ed as I. ledongensis (Gong e al. 2011) as a new species,
I. wangyingyongi. Liu e al. (2025) also demons a ed
ha I. ledongensis s. s . om Hainan Island o China is
placed deeply wi hin he adia ion o he I. galea us com-
plex. In ou m DNA-based genealogy, I. wangyingyongi
and I. ledongensis o m a well-suppo ed clade, ende ing
I. galea us pa aphyle ic (Fig. 2). Howe e , he s udy o
Liu e al. (2025) con ained se e al signi ican laws, and
hei conclusions mus be aken c i ically.
Fi s ly, jus as in ou s udy, Liu e al. (2025) demon-
s a ed ha I. wangyingyongi and I. ledongensis a e
placed wi hin he adia ion o I. galea us; hei posi ion
wi hin he complex is essen ially un esol ed in he anal-
ysis o Liu e al. (2025), and hei ecogni ion makes
I. galea us pa aphyle ic. Ne e heless, Liu e al. (2025)
labeled wo lineages o I. galea us as po en ial new spe-
cies, Inge oph ynus sp. 1 ( om Quang Binh, cen al Vie -
nam) and Inge oph ynus sp. 2 ( om no he n Vie nam),
and e e ed o a lineage om Gia Lai (cen al Vie nam)
as he ue I. galea us s. s . wi hou p o iding any solid
e idence o his decision. As he ull species s a us o
I. wangyingyongi and I. ledongensis is no e iden , he
ac ha hei placemen wi hin I. galea us makes he la -
e species pa aphyle ic canno be used as an a gumen
o spli ing he complex in o se e al species, including
wo unnamed pu a i e “species.” A mo e conse a i e
app oach would sugges ha I. wangyingyongi and I. le-
dongensis should be conside ed junio synonyms o I. ga-
lea us, pending u he molecula and mo phological da a
on he emaining popula ions o he complex.
Secondly, he decision by Liu e al. (2025) o ega d
he Gia Lai popula ion (cen al Vie nam) as nomino ypi-
cal I. galea us s. s . is ques ionable. Liu e al. (2025) e e
o he ype locali y o I. galea us, which was gi en by
Gün he (1864) as “Gamboja” (= Cambodia); he au ho s
show i on he map as a locali y in no he n Cambodia (Liu
e al. 2025: Fig. 5) and conclude ha he popula ion om
he moun ains o Gia Lai P o ince o Vie nam he e o e
belongs o I. galea us s. s . due o i s geog aphic p oxim-
i y. No only did Liu e al. (2025) no examine he ype
specimen I. galea us (holo ype NHMUK 1947.2.21.13),
bu hei conclusion also appea s o be e oneous. S ua e
al. (2006) demons a ed ha Bu o galea us was desc ibed
by Gün he (1864) om M. Hen i Mouho ’s Cambodian
collec ion and ha he ype specimen o his species was
almos ce ainly collec ed du ing Mouho ’s expedi ion o
“B elum.” S ua e al. (2006) showed ha his illage
was likely loca ed on he p esen -day e i o y o Vie nam
nea he bo de wi h Cambodian Modulki i P o ince (ca.
11°58'N, 107°12'E, in Binh Phuoc P o ince o Vie nam,
acco ding o Ashbu on in Mouho 1864). This a ea is
loca ed o e 330 km sou hwa d om Gia Lai P o ince
and belongs o he hilly oo hills o he sou he n Anna-
mi es, an a ea biogeog aphically p o oundly di e en
om he mon ane o es s o cen al Vie nam (Poya ko e
al. 2021, 2023). The e o e, he a ibu ion o he Gia Lai
popula ion o he I. galea us complex o I. galea us s. s .
by Liu e al. (2025) is unwa an ed.
Finally, he le el o gene ic di e gence be ween
I. wangyingyongi and I. ledongensis in he 16S RNA
gene is minimal (p = 2.21%; Suppl. ma e ial 1: able S6),
and hei di e gence om he lineages o I. galea us is
also gene ally lowe han p = 3.0%, he o mal le el o
species-le el di e gence in anu ans (Vences e al. 2005;
Viei es e al. 2009). In he absence o eliable diagnos ic
mo phological cha ac e s, examina ion o he I. galea us
ype specimen and su icien samples o I. galea us, and
he lack o nuclea DNA o bioacous ic da a, we con-
clude ha such low di e gence alues in a single m DNA
ma ke a e insu icien o jus i y he ull species s a us
o I. wangyingyongi and I. ledongensis. Recognizing he
geog aphical and ecological speci ici y o he I. galea-
us complex lineages om sou he n China, we ollow he
ecommenda ions o Du esnes e al. (2023, 2024) and
he e o e p opose o ea hem as subspecies Inge oph y-
nus galea us wangyingyongi s a . no . and Inge oph ynus
galea us ledongensis s a . no . pending u he in eg a i e
axonomic analysis. Fu u e s udies should p o ide a com-
p ehensi e e ision o he I. galea us complex, including
a me iculous analysis o i s a ia ion in nuDNA ma ke s,
as well as in mo phological and bioacous ic cha ac e s.
The axonomic s a us o I. gollum, which was also ound
o be e y closely ela ed o I. di e gens in 16S RNA
gene sequences (p = 2.32%; Suppl. ma e ial 1: able S6),
may also be econside ed in he u u e; he e we e ain
om any axonomic changes pending addi ional gene ic
and mo phological da a on he I. di e gens complex.
I is also no able ha Fei e al. (2012) assigned I. le-
dongensis o a sepa a e genus, Qiangbu o Fei, Ye & Jiang,
2012. Ou s udy, in acco dance wi h he ea lie esul s o
Liu e al. (2025), places I. galea us ledongensis s a . no .
deeply wi hin he adia ion o he genus Inge oph ynus,
as one o he lineages o he I. galea us complex. The e-
o e, ou esul s con i m ha Qiangbu o Fei, Ye & Jiang,
2012 should be ega ded as a junio synonym o Inge o-
ph ynus F os , G an , Fai o ich, Bain, Haas, Haddad, de
Sá, Channing, Wilkinson, Donnellan, Raxwo hy, Camp-
bell, Blo o, Mole , D ewes, Nussbaum, Lynch, G een &
Wheele , 2006, a axonomy ha is al eady widely accep -
ed (F os 2025). The nomen Qiangbu o is hus a ailable
o he mainland Asian clade o Inge oph ynus (including
he I. galea us complex and I. mac o is). Howe e , u -
he mul ilocus s udies o Inge oph ynus a e equi ed o
cla i y he phylogene ic ela ionships o he genus.
The in eg a i e axonomic analysis o he I. pa us
species complex con i med he p esence o wo e olu-
iona ily independen lineages, dis inc in m DNA se-
quences, nuDNA gene alleles, ad e isemen male call
bioacous ic pa ame e s, and ecological specializa ion.
He pe ozoa 38: 271–297 (2025)
he pe ozoa.penso .ne
293
These esul s con i m he conclusions o p e ious s udies,
which demons a ed signi ican di e en ia ion wi hin he
I. pa us complex based on mo phological da a (S ion
e al. 2018) and m DNA sequences (Chan and G isme
2019), and gene ally sugges he exis ence o wo clea ly
de ined sis e species: I. pa us s. s . and Inge oph ynus
ch ysolophus sp. no .
Fu he mo e, ou s udy e ealed ha he dis ibu ion
bounda ies o hese wo species coincide wi h he Kan-
ga -Pa ani Line—an impo an biogeog aphic bounda y
sepa a ing equa o ial o es s om seasonal opical mon-
soon o es s (Mo ley 2000; Poya ko e al. 2021, 2023).
The Kanga -Pa ani Line (KPL) is loca ed a 6–7°N, which
ma ks an impo an shi in bo h lo is ic and clima ic
cha ac e is ics om aseasonal o seasonal e e g een op-
ical o es wi hin he Indo-Sundaic a ea o Sou heas Asia
(Van S eenis 1950; Whi mo e 1984; Mo ley 2000; Wik a-
manayake e al. 2000; Wood u 2003, 2010; Bal ze e al.
2008, 2009; Quah and Anua 2018). I is assumed ha he
shi om Con inen al Asia ic lo a in he no h o Male-
sian lo a in he sou h o he KPL is linked o he p es-
ence o one o mo e mon hs o d ough occu ing no h o
he KPL (Whi mo e 1990). The clima ic o ces ha ha e
shaped he lo a ha e simila ly a ec ed he dis ibu ion o
auna in he egion. Nume ous s udies ha e shown i o be
a signi ican a ea o aunal exchange ac oss a ious axa,
including amphibians (Quah and Anua 2018; Poya ko
e al. 2020; Suwannapoom e al. 2020, 2021, 2022; T o-
ime s e al. 2024), ep iles (G isme 2011; G isme e al.
2014, 2020a, 2020b, 2023; Idiia ullina e al. 2023, 2024),
bi ds (Reddy 2008), and mammals (Wood u and Tu ne
2009; Pa ou e al. 2010). Ou da a u he unde line he
impo ance o his biogeog aphic bounda y in shaping he
di e si y o Sou heas Asian amphibians.
Many new amphibian species a e desc ibed e e y
yea , o en by e ising wide- anging species complex-
es consis ing o lineages wi h supe icially simila ex-
e nal mo phology (T o ime s e al. 2024; Go in e al.
2024). The e is no doub ha in ensi ied su ey e o s,
especially in he unexplo ed emo e moun ain and island
a eas o Sou heas Asia, coupled wi h he in oduc ion
o new echniques, will acili a e he disco e y o o e -
looked di e si y in he genus Inge oph ynus species
(E ans e al. 2003). An in eg a i e axonomic app oach
appea s o be c ucial in analyzing he c yp ic di e si y
o amphibians, and in he case o anu ans, bioacous ic
analysis is pa icula ly p omising (Koehle e al. 2017).
He ein we p o ide b ie desc ip ions o male ad e ise-
men calls o I. pa us s. s . and Inge oph ynus ch yso-
lophus sp. no . and e eal s able di e ences in empo al
and equency pa ame e s among he calls o hese wo
species. I is no able ha , o da e, he exis ing da a on he
bioacous ics o he genus Inge oph ynus a e ex emely
sca ce; o he bes o ou knowledge, only wo pape s
b ie ly desc ibing call pa ame e s o I. quad ipo ca us
and I. di e gens we e published (Sukuma an e al. 2010;
Am am e al. 2018). The e o e, we p esen he i s bio-
acous ic da a o he I. pa us species complex membe s
and encou age u he s udies o male ad e isemen call
a ia ion in he genus Inge oph ynus.
Wi h he desc ip ion o Inge oph ynus ch ysolophus
sp. no . and he p oposed synonymiza ion o I. wangy-
ingyongi and I. ledongensis wi h I. galea us, he o al
numbe o species in he genus eaches 12, h ee o which
a e known o occu in Thailand, namely I. ch ysolophus
sp. no ., I. mac o is, and I. pa us s. s . As a esul o
his e ision, he dis ibu ion o I. pa us s. s . is limi -
ed o Suma a ( he en i e island) and Ja a ( he wes e n-
mos pa o he island), he Malay Peninsula sou h o he
Kanga -Pa ani Line, and se e al o sho e islands (like
he Se ibua A chipelago o Malaysia). Consequen ly,
we emo e I. pa us om he auna o Cambodia and
Myanma . Fu he mo e, wo small-sized Inge oph ynus
species, namely I. bipo ca us and I. di e gens, which a e
supe icially simila o he I. pa us complex, ha e been
p e iously epo ed om Peninsula Thailand wi hou
ouche specimens o de ailed in o ma ion on hese e-
co ds (Chuaynke n and Chuaynke n 2012; Poya ko e
al. 2021). We suspec ha hese a e misiden i ica ions o
species belonging o he I. pa us complex, and he e o e
we p opose o emo e hese wo species om he auna
o Thailand pending u he e idence o hei occu ence.
While i is o en di icul o obse e consis en di -
e ences in ex e nal mo phology among closely ela ed
amphibian species, s udying c anial mo phology shows
g ea p omise and can be pa icula ly in o ma i e o
diagnos ics on di e en le els (e.g., De o el e al. 2021;
Go in e al. 2021; Poya ko e al. 2024; and he e e enc-
es he ein). O e all, he s udies on os eology o he amily
Bu onidae appea o be sca ce, wi h no s udies known o
us dedica ed o he genus Inge oph ynus in pa icula . In
his ega d, we hope ha ou cu en esea ch will p o-
ide a solid amewo k o he subsequen e isions o
he genus Inge oph ynus ha include os eological da a.
Al hough we a e lacking a e e ence wo k o p o ide a
compa ison be ween di e en species o Inge oph ynus,
he ob ained mic o-CT scans p o ided new da a on he
ana omy o he sup ao bi al and sup a ympanic c anial
c es s— he diagnos ic cha ac e o he genus Inge oph y-
nus. Also, ou da a show ha he skull o Inge oph ynus
ch ysolophus sp. no . shows bo h aces o hype ossi ica-
ion and unde ossi ica ion. I seems ha bo h he hype os-
si ica ion o he c anial oo and unde ossi ica ion o he
o ic egion a e mo e p onounced in he emale specimen
han in he male one. Un o una ely, wi hou addi ional
da a, we a e unawa e i hese di e ences can be a ibu ed
o sexual dimo phism o indi idual a iabili y.
Mos species o he genus Inge oph ynus ha e he
conse a ion s a us Leas Conce n (LC), and only such
na ow- anged endemics as I. kumqua and I. gollum a e
lis ed as Endange ed (EN) in he IUCN’s Red Lis ca -
ego ies (IUCN 2019). The desc ip ion o a new species
o his genus emphasizes he uniqueness o he auna o
Thailand, and he occu ence o he species in di e en
locali ies o he coun y gi es eason o belie e ha con-
ce ns abou i s ex inc ion a e minimal.
he pe ozoa.penso .ne
Dmi iy V. A khipo e al.: A new species o Inge oph ynus om Thailand294
Acknowledgmen s
We would like o hank he Labo a o y Animal Re-
sea ch Cen e , Uni e si y o Phayao, and he Ins i u e
o Animal o Scien i ic Pu poses De elopmen (IAD),
Thailand, o hei pe mission o conduc he ieldwo k
he e. We a e deeply g a e ul o T. Ruangsuwan, M.
Naiduangchan, and T. Wo anuch (Thailand), and T.
Ma sukoji (Japan) o hei help du ing he ield su -
eys. We hank he membe s o MSU He pLab, includ-
ing A. V. T o ime s, S. S. Idiia ullina, N. S. Kliukin,
and E. N. Solo ye a, o hei suppo and assis ance.
We exp ess ou since e g a i ude o he academic edi o
A hu Tiu enko and Zeeshan Mi za o nume ous con-
s uc i e commen s and sugges ions, which allowed us
o imp o e he p e ious e sion o he manusc ip .
This wo k was suppo ed by he Russian Science
Founda ion o N.A. Poya ko (G an No. RSF 22-14-
00037-P, specimen collec ion and p ese a ion, molec-
ula and mo phological analyses, and da a analyses),
he Na ional Science Founda ion (G an No. NSF
DEB-1146324 o E.N. Smi h and M.B. Ha ey), and
he Thailand Science Resea ch and Inno a ion Fund
and he Uni e si y o Phayao, Uni o Excellence 2026
on Aqua ic Animals Biodi e si y Assessmen (Phase
II) o C. Suwannapoom. Specimen collec ion and an-
imal use p o ocols in Thailand we e app o ed by he
Ins i u ional E hical Commi ee o Animal Expe imen-
a ion o he Uni e si y o Phayao, Phayao, Thailand
(ce i ica e numbe UP-AE64-02-04-005, issued o
C. Suwannapoom) and we e s ic ly complian wi h
he e hical condi ions o he Thailand Animal Wel a e
Ac . Fieldwo k, including he collec ion o animals in
he ield, was au ho ized by he Ins i u e o Animals
o Scien i ic Pu pose De elopmen (IAD), Bangkok,
Thailand (pe mi numbe s U1-01205-2558 and UP-
AE59-01-04-0022, issued o C. Suwannapoom). Re-
sea ch in Indonesia was conduc ed unde esea ch pe -
mi 149/SIP/FRP/SM/V/2013 (issued o E.N. Smi h).
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