A taxonomic revision of the Ingerophrynus parvus species complex (Amphibia, Anura, Bufonidae) with the description of a new species from Thailand

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) he pe ozoa.penso .ne 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) he pe ozoa.penso .ne 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 . 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