A new species of the genus Megophrys (Anura: Megophryidae) from Yunnan Province, China Author Yang, Jian-Huan Author Wang, Jian Author Wang, Ying-Yong text Zootaxa 2018 2018-04-23 4413 2 325 338 journal article 30210 10.11646/zootaxa.4413.2.5 d9d49c4c-4662-44c9-8da4-63b9e67e32f2 1175-5326 1226983 9F0E14D1-6B87-4DD3-92DC-2A62E04641F3 Megophrys feii sp. nov. Fig.1–3 ; Table 1 Holotype . SYS a006524, adult male, cOllected frOm XiaOlangsu Village , TOngbiguan TOwn, Yingjiang COunty , Yunnan PrOvince , China (24˚30'3.23"N, 97˚34'16.75"E, 700 m a.s.l. ), On 5 May 2016 by Jian-Huan Yang (JHY, hereafter). Paratypes . SYS a006525–6526, twO adult females, data identical tO hOlOtype . SYS a006527–6529, three adult males, cOllected frOm adjacent fOrest stream in TOngbiguan TOwn at higher elevatiOn ( 1200 m a.s.l.), On 22 April 2017 by JHY. Etymology. The specific epithet “ feii ” is a patrOnymic nOun in the genitive singular; derived frOm the name Of PrOfessOr Liang Fei Of the Chengdu Institute Of BiOlOgy, Chinese Academy Of Sciences, China , in recOgnitiOn Of his lOng-term and significant cOntributiOn tO the advancement Of amphibian research in China , particularly his research On the MegOphryidae family. Diagnosis. The new species is assigned tO the genus Megophrys s. l. by having a snOut Obtusely prOtruding in lateral view, snOut tip prOjecting far beyOnd the lOwer lip, canthus rOstralis sharp, and small hOrned-like tubercles present On the upper eyelid (Fei et al. 2009; Fei & Ye 2016). Megophrys feii sp.nov. can be distinguished frOm Other cOngeners by a cOmbinatiOn Of fOllOwing characters: (1) bOdy slender and small (SVL 24.3–25.1 mm in fOur males, 28.2–28.9 mm in twO females); (2) head length abOut equal tO head width (HDL/HDW 96.9–99.0%); (3) tympanum circular and distinct; (4) maXillary teeth present; (5) vOmerine ridges and vOmerine teeth absent; (6) tOngue rOund, slightly nOtched pOsteriOrly; (7) tibia lOng (TIB/SVL 48.2–52.0% in fOur males, 54.3–55.4% in twO females), shanks Overlapping when thighs are held at right angles tO the bOdy; (8) dOrsal skin distinctly granular and densely cOvered with small tubercles; (9) lOwer flanks and lateral sides Of belly scattered with small but prOminent, white, tubercles; (10) webbing between tOes rudimentary; (11) lateral fringes On tOes mOderate tO wide; (12) breeding males withOut nuptial pads and spines On fingers; (13) prOtruding prOjectiOn pOsteriOr tO clOaca present in bOth seXes; (14) grOin and ventral thigh cOlOratiOn in life nOt cOntrasting with surrOunding regiOns; (15) the presence Of an indistinct and small hOrn-like tubercle at the edge Of the eyelid; and (16) an advertisement call with a dOminant frequency Of 4.74–4.91 kHz (at 18 ˚C). Description of Holotype . SYS a006524, adult male, bOdy size small and slender (SVL 25.0 mm) cOmpared tO Other cOngeners in the genus. Head small, head length abOut equal tO head width; snOut rOunded in dOrsal view, Obtusely prOtruding in lateral view, withOut rOstral prOjectiOn; lOreal regiOn vertical and cOncave; canthus rOstralis sharp; dOrsal regiOn Of snOut slightly cOncave; eye diameter (EYE 3.1 mm ) greatly larger than tympanum diameter (TMP 1.8 mm ), and shOrter than snOut length (SNT 3.5 mm ); eye-tympanum distance (TEY 1.2 mm ) shOrter than tympanum diameter; tympanum circular and distinct, with upper margin slightly cOncealed by supratympanic ridge; pupil in life slightly hOrizOntal; nOstril Oriented laterally, situated slightly clOser tO the eye than tO the tip Of the snOut (EN 1.6 mm , NS 1.9 mm ); internarial distance (IND 3.2 mm ) slightly greater than upper eyelid width (UEW 2.9 mm ), and abOut equal tO interOrbital distance (IUE 3.1 mm ); pineal Ocellus nOt visible eXternally; maXillary teeth present; vOmerine ridges and vOmerine teeth absent; tOngue rOund, slightly nOtched pOsteriOrly, withOut medial lingual prOcess. FOrelimbs thin and lOng, lOwer arm slightly thicker relative tO upper fOrelimb; length Of lOwer arm and hand lOnger than half Of snOut-vent length (LAHL/SVL 54.5%); fingers shOrt, finger length fOrmula I<II<IV<III; mOderate lateral fringes present On Outer mOst three fingers while absent On the first finger; interdigital webbing, subarticular and supernumerary tubercles absent; thenar and palmar tubercles barely distinguishable; finger tips slightly dilated and rOunded with Oval pads; terminal grOOves absent; nuptial pads and spines absent ( Fig. 1 ). TABLE 2. Morphological comparison of Megophrys feii s.. with all 43 congeners reporteđ from Yunnan anđ the borđering provinces of southwest China (i.e., Guangxi, Guizhou, Sichuan Tibet), anđ from neighbouring countries in Vietnam, Laos, Myanmar, anđ northeast Inđia east of Brahmaputra river. Grey shađing inđicates a đistinct đifference when compaređ to the new species. Male SVL Female SVL Maxillary Tympanum Nuptial Toes friđge Toes webbing Vomerine Vomerine Dorsal skin References teeth pađs/spines teeth riđge texture . feii s.. 2. 2. 2.2 2. 2 s s s! " # $ s s s %&$ % "& This stuđy . ancrae 39.1̅45.0, n=8 48.9, n=1 present đistinct present absent ruđimentary present present weakly granular Mahony et al . 2013. binchuanensis 32.0̅36.0, n=4 40.2̅42.5, n=2 present đistinct present wiđe ruđimentary absent absent smooth Fei et al . 2009, 2012. binlingensis 45.1̅51.0, n=3 -- present đistinct present narrow ruđimentary absent present smooth Fei et al . 2009, 2012. brachykolos 33.7̅39.3, n=5 33.9̅45.9, n=2 present đistinct present absent ruđimentary absent? smooth Inger & Romer 1961 ; Fei et al . 2009, 2012. carinense 91.6̅122.5, n=15 137.0, n=1 present hiđđen present wiđe mođerate/wiđe present present smooth Fei et al . 2009, 2012. chuannanensis 91.4̅109.4, n=12 -- present hiđđen present wiđe ruđimentary present present smooth Fei et al . 2009, 2012. daweimontis 34.0̅37.0, n=18 40.0̅46.0, n=3 present đistinct? absent absent present present smooth Rao & Yang 1997; Fei et al . 2009, 2012. elfina 26.9̅33.9, n=29 35.1̅36.5, n=6 absent đistinct present absent ruđimentary absent absent shagreeneđ Poryakov et al. 2017. feae 78.0̅101.8, n=22 91.0̅113.5, n=8 present hiđđen present wiđe ruđimentary present present smooth Fei et al . 2009, 2012. gerti 31.7̅42.2, n=15 43.1̅47.4, n=3 absent đistinct present absent ruđimentary absent absent shagreeneđ Ohler 2003 ; Poryakov et al. 2017 . gigantica 80.5̅107.0, n=6 110.4̅115.4, n=2 present hiđđen present wiđe ruđimentary absent absent smooth Fei et al . 2009, 2012. glandulosa 76.3̅81.0, n=10 76.5̅99.5, n=3 present đistinct present wiđe ruđimentary present present smooth Fei et al . 2009, 2012. hansi 33.4̅43.1, n=12 45.1̅53.9, n=5 absent đistinct present absent absent absent absent shagreeneđ Ohler 2003 ; Poryakov et al. 2017 . intermedia 86.0̅103.0 92.0, n=1 present hiđđen present wiđe ruđimentary present present smooth Smith 1921 . jingdongensis 53.0̅56.5, n=3 63.5, n=1 present đistinct present wiđe half webbeđ present present smooth Fei et al . 2009, 2012. koui 28.0̅30.0, n=7 25.0̅26.0, n=2 absent đistinct present absent absent/basal absent absent shagreeneđ Fei et al . 2009, 2012; Poryakov et al. 2017. latidactyla 38.9, n=1 -- present đistinct present wiđe half webbeđ present present tuberous Orlov et al . 2015. liboensis 34.7̅67.7 60.8̅70.6 present đistinct? mođerate ruđimentary present present smooth @hang et al . 2017. major 77, n=1 94, n=1 present đistinct present wiđe small present present smooth Boulenger 1908. maosonensis 61.9̅83.4, n=8 -- present đistinct present narrow? present present smooth Chen et al . 2017; examineđ specimens. medogensis 57.2̅68.0, n=16 -- present đistinct present absent absent/narrow present present smooth Fei et al . 2009, 2012 ……continued on the next page Male SVL Female SVL Maxillary Tympanum Nuptial Toes friđge Toes webbing Vomerine Vomerine Dorsal skin TABLE 2. (Continueđ) teeth pađs/spines teeth riđge texture megacephala 45.9̅53.4, n=8 64.4, n=1 present đistinct present absent ruđimentary present present smooth Mahony et al . 2011 microstoma 34.3̅44.4, n=10 39.4̅57.0, n=7 absent đistinct present absent absent/basal absent absent shagreeneđ Fei et al . 2009, 2012; Poryakov et al. 2017 minor 32.3̅40.5, n=15 42.0̅48.2, n=4 present đistinct present absent ruđimentary absent weak smooth Fei et al . 2009, 2012 nankiangensis 39.1, n=1 44.0̅52.9, n=10 present hiđđen? narrow ruđimentary absent absent smooth Fei et al . 2009, 2012 omeimontis 56.0̅59.5, n=10 68.0̅72.5, n=3 present đistinct present narrow ruđimentary present present smooth Fei et al . 2009, 2012 oropedion 32.8̅39.2, n=7 44.1̅48.7, n=2 present đistinct present absent absent present present weakly granular Mahony et al . 2011 pachyproctus 35.3̅36.2, n=2 35.8, n=1 present đistinct present? absent present present granular Fei et al . 2009, 2012 palpebralespinosa 36.2̅38.0, n=2 -- present đistinct present wiđe half webbeđ present present rough Fei et al . 2009, 2012 parva 41.5̅44.0, n=6 -- present đistinct present absent absent/narrow present? smooth Fei et al . 2009, 2012 robusta 114.0, n=1 present đistinct?? ruđimentary present present smooth Boulenger 1908 rubrimera 26.7̅30.5, n=8 present đistinct present narrow absent present present weakly granular Tapley et al . 2017 serchhipii 37.1, n=1 (sex unknown) present đistinct?? ruđimentary present? smooth Mathew & Sen 2007 shapingensis 65.9̅84.2, n=10 76.6̅104.0, n=10 present absent absent wiđe half webbeđ absent absent smooth Fei et al . 2009, 2012 shuichengensis 102.0̅118.3, n=7 99.8̅115.6, n=6 present đistinct absent wiđe half webbeđ absent present smooth Tian et al . 2000; Fei et al . 2009, 2012 spinata 47.2̅54.4, n=18 54.0̅55.0, n=2 present đistinct present wiđe half webbeđ absent weak smooth Fei et al . 2009, 2012 synoria 38.2̅53.7, n=14 51.4̅70.7, n=3 absent đistinct present? absent/basal absent absent shagreeneđ Stuart et al . 2006 vegrandis 27.5̅30.6, n=4 present đistinct absent wiđe ruđimentary absent absent smooth Mahony et al . 2013 wawuensis 34.4̅42.8, n=4 47.0̅49.8, n=2 present đistinct present absent ruđimentary absent absent smooth Fei et al . 2009, 2012 wuliangshanensis 27.3̅31.6, n=10 41.0̅41.5, n=2 present đistinct present absent absent absent absent đensely granulate Fei et al . 2009, 2012 wushanensis 30.4̅35.5, n=10 38.4, n=1 present đistinct present wiđe/absent absent/narrow absent absent smooth Fei et al . 2009, 2012 zhangi 32.5̅37.2, n=3 present đistinct present narrow absent present present smooth Fei et al . 2009, 2012 zunhebotoensis 30.0, n=1 39.0, n=1 present đistinct??? present? smooth Mathew & Sen 2007 Hind limbs relatively lOng and thin, shanks Overlap when thighs are held at right angles tO the bOdy, thigh length (ML 12.8 mm ) abOut equal tO shank length (TIB 13.0 mm), and lOnger than fOOt length (FL 10.5 mm ); tOes shOrt and flattened with brOad lateral fringes that eXtend tO the tips Of all digits; relative tOe lengths I<II<V<III<IV; tips nOt dilated, but with distinct pads; terminal grOves absent; base Of all tOes with rudimentary webbing; Outer metatarsal tubercle, subarticular and supernumerary tubercles absent; inner metatarsal tubercle present but barely distinguishable, 1.3 mm in length; ridge Of callOus tissue present On ventral surface Of all digits. PrOtruding prOjectiOn pOsteriOr tO clOaca present and distinct in life, while nearly invisible in preservative. FIGURE 1. HOlOtypE Of Megophrys feii sp. nov. SYS A006524 In lIfE: (A) dOrsAl vIEW; (B) vEntrAl vIEW; (C) vEntrAl vIEW Of rIght hAnd; (D) vEntrAl vIEW Of rIght fOOt; (E) dOrsOlAtErAl vIEW; (F) lAtErAl vIEW Of lEft hEAd, nOtEd thE IrIs cOlOrAtIOn. PhOtOs by JHY. The skin Of dOrsal surfaces Of the head, bOdy and limbs are rather granular and densely cOvered with small tubercles; all tubercles On the back terminate in asperities; tympanum smOOth, bOrders slightly raised, surrOunding area pOsteriOr tO eye granular; upper eyelid densely cOvered with small tubercles; an indistinct and small hOrn-like tubercle present at the edge Of the eyelid; supratympanic fOld distinct and narrOw, withOut widening pOsteriOrly, eXtends frOm Orbit and curves dOwn thrOugh upper bOrder Of tympanum terminating abOve shOulder; narrOw dOrsOlateral skin fOld distinct and interrupted, eXtending frOm behind supratympanic fOld tO apprOXimately One third distance tO grOin; twO OppOsing v-shaped skin fOlds present On dOrsum; pectOral glands small and raised weakly ( 0.7 mm in diameter), slightly smaller than the tips Of fingers, and abOut equal tO small white tubercles On lateral sides Of belly in size; femOral glands small and weakly raised ( 0.8 mm in diameter), abOut equal tO the tips Of tOes in size, pOsitiOned sub-equally distant frOm knee and clOaca; lOwer flanks scattered with small distinctly raised white tubercles, which eXtending tO lateral sides Of belly; ventral surfaces primarily smOOth, withOut asperities ( Fig. 1 ). Color of the holotype in life. DOrsal and lateral surfaces Of head, bOdy and limbs primarily light grey-brOwn; dark brOwn triangular marking present On tOp Of the snOut; dark brOwn inverted triangular marking withOut light central blOtch present between eyes; dark brOwn dOrsOlateral and rOughly “X”-shaped markings present On dOrsal surface Of bOdy; flanks with irregular dark brOwn spOts; dark brOwn vertical bars present On upper lips, the One belOw eye largest and much distinct; tympanum entirely dark brOwn; dOrsal surface Of upper arm paler brOwn; distinct and wide dark brOwn transverse bars present On dOrsal surfaces Of lOwer arms and hindlimbs; fingers and tOes with dark brOwn bars On dOrsal surfaces. ThrOat and chest dark grey and mOttled with white; an indistinct lOngitudinal dark brOwn stripe present On the thrOat; margin Of lOwer mandibles dark brOwn and scattered with white spOts; ventral surfaces Of bOdy and limbs primarily white, and scattered with distinct irregular small dark brOwn spOts and flecks On belly; distinct, small and white tubercles present On lOwer flanks and lateral sides Of belly; pectOral and femOral glands white; ventral surfaces Of tarsus and feet dark brOwn; area surrOunding clOaca and pOsteriOr surfaces Of thighs dark brOwn. Iris bicOlOred, cOpper-brOwn in the upper twO-thirds and yellOwish- Orange in the lOwer One-third ( Fig. 1 ). Color of the holotype in preservative. Dark brOwn markings On the dOrsum and bars On the limbs remain visible. Ventral surface dull white and scattered with irregular and distinct dark brOwn spOts; small and white tubercles On lateral sides Of belly and lOwer flanks still distinct; pectOral glands becOme less distinct. Variation. All five paratypes match the Overall characters Of the hOlOtype (fOr measurements Of the type series see Table 1 ). Nuptial pads and spines absent in all fOur male types . SYS a06527 and a006529 have a “X”-shaped skin fOld On dOrsum, rather than twO OppOsing v-shaped skin fOlds. Lateral fringes present On fingers in hOlOtype and three paratypes SYS a006527–6529, but absent in twO paratypes SYS a6525–6526. Lateral fringes On tOes mOderate in hOlOtype and paratypes SYS a006525–6526, while distinctly wider in three paratypes SYS a006527– 6529. SYS a006529 has a relatively darker venter and its thigh mOttled with dark brOwn and white ( Fig. 2–3 ). Sexual dimorphism. NO significant seXual dimOrphism Observed, hOwever females are slightly larger than males in bOdy size, have a relatively darker thrOat, distinctly smaller dark brOwn spOts On the venter, and relatively lOnger hind limbs than males (HLL/SVL 172.7–177.2% in twO females, 157.8–166.8% in fOur males); the prOtruding prOjectiOn pOsteriOr tO the clOaca is distinctly lOnger in twO female types than fOur male types . Advertisement call. The advertisement call Of Megophrys feii sp. nov. was recOrded frOm the male paratype SYS a006528 at 00:20 h On 20 April 2017 . Ambient air temperature during the recOrding was ca. 18˚C. The call series cOnsists Of a single call Of 115–150 ms duratiOn (132.6±6.3 ms, N=44), cOntaining 14–20 pulses per call (16.8±1.38, N=44). Pulse interval gradually decreases frOm the beginning tO the end Of the call. Calls were repeated in series at a rate Of apprOXimately 2.0 calls per secOnd, with an irregular intercall-interval Of 134–736 ms duratiOn (371.5±106.7 ms, N=42). The duratiOn Of the call series was nOt recOrded, but calls were cOntinuOusly emitted thrOughOut the lOngest recOrding made Of 21 s. The dOminant frequency is 4.74–4.91 kHz and a harmOnic is present at 5.94–6.29 kHz ( Fig. 4 ). Distribution and ecology. Megophrys feii sp. nov. is currently knOwn frOm Yingjiang COunty, Yunnan PrOvince , China ( Fig. 5 ). The type lOcality Of the new species is less than One kilOmeter frOm the bOundary Of TOngbiguan PrOvincial Nature Reserve, and less than twO kilOmeters frOm the internatiOnal bOrder with Myanmar’s Kachin State; thus we are cOnfident that the new species Occurs in the in adjOining, well-preserved, mOntane fOrests Of TOngbiguan Nature Reserve and Kachin State in Myanmar . The new species was fOund in small, shallOw streams and nearby seepages in well-preserved mOntane evergreen brOadleaf fOrest at elevatiOns between 700 and 1200 m . During survey On 5 May 2016 , male hOlOtype (SYS a006524) and female paratype (SYS a006526) were fOund in ampleXus On leaf litter On the fOrest flOOr ( Fig. 3A ). All calling adult males were fOund perched On shrub leaves by the stream during surveys ( Fig. 3 , B & C). FIGURE 2. DOrsAl (A) And vEntrAl (B) vIEWs Of typE spEcImEns Of Megophrys feii sp. nov. In prEsErvAtIvE: (1) mAlE hOlOtypE SYS A006524; (2) mAlE pArAtypE SYS A006527; (3) mAlE pArAtypE SYS A006528; (4) mAlE pArAtypE SYS A006529; (5) fEmAlE pArAtypE SYS A006525 And (6) fEmAlE pArAtypE SYS A006526. PhOtOs by JHY. FIGURE 3. Megophrys feii sp. nov. (A) A pAIr In AmplEXUs (mAlE hOlOtypE SYS A006524 And fEmAlE pArAtypE SYS A006526); (B) mAlE pArAtypE SYS A006528 vOcAlIzIng; (C) mAlE pArAtypE SYS A006529. PhOtOs by JHY. Comparisons. The new species can be distinguished frOm the 43 knOwn cOngeners repOrted frOm Yunnan and bOrdering prOvinces Of sOuthwest China , and frOm the neighbOuring cOuntries Of Vietnam , LaOs , Myanmar and nOrtheast India east Of the Brahmaputra river by its small bOdy size (SVL 24.3–25.1 mm in fOur males, 28.2–28.9 mm in twO females; and it is One Of the smallest species in the genus Megophrys described tO date), presence Of maXillary teeth, distinct tympanum, absence Of nuptial pads and spines in breeding males, absence Of vOmerine teeth and ridge, degree Of webbing and fringing On the tOes, and granular skin teXture On the dOrsum (see Table 2 ). In additiOn, Megophrys feii sp. nov. further differs M. carinense , M. feae , M. intermedia and M. popei (all Of which have been allOcated tO the subgenus Brachytarsophrys On the basis Of mOlecular data [MahOny et al. 2017]) by the absence Of a transverse fOld at the base Of the head (vs. present), slender bOdy shape (vs. fleshy and rObust bOdy shape) and having a head that is nOt depressed and is apprOXimately as wide as lOng (vs. head distinctly depressed and much wider than lOng). Megophrys feii sp. nov. further differs M. carinense , M. feae , M. gerti , M. hansi , M. intermedia , M. koui , M. latidactyla , M. liboensis , M. microstoma , M. palpebralespinosa , M. popei , M. shuichengensis and M. synoria by the presence Of a small and indistinct hOrn-like tubercle at the edge Of the upper eyelid (vs. a relatively prOminent and elOngated tubercle), and frOm M. binchuanensis , M. binlingensis , M. daweimontis , M. minor , M. nankiangensis , M. omeimontis , M. parva , M. spinata , M. wawuensis and M. wuliangshanensis by having grOin and ventral thigh cOlOratiOn in life nOt cOntrasting with the surrOunding regiOns in males (vs. grOin and/Or ventral thigh cOlOured red and distinctly cOntrasting with the surrOunding regiOn in males in life). TABLE 1. Measurements of type specimens of Megophrys feii sp. nov. . Abbreviations defined in text.

SYS A06524	SYS A06527	SYS A06528	SYS A06529	mEAn ±SD (N=4)	SYS A06525	SYS A06526	mEAn ±SD (N=2)
hOlOtypE	pArAtypE	pArAtypE	pArAtypE	pArAtypE	pArAtypE	pArAtypE
SEX	m	m	m	m	f	f
SVL	25.0	24.4	24.3	25.1	24.7±0.41	28.2	28.9	28.6±0.49
HDL	9.5	8.8	9.3	9.6	9.3±0.36	10.7	10.6	10.7±0.07
HDW	9.6	8.9	9.6	9.8	9.5±0.39	10.9	10.9	10.9±0.00
SNT	3.5	3.4	3.7	3.8	3.6±0.18	3.9	4.1	4.0±0.14
EYE	3.1	3.1	3.5	3.6	3.3±0.26	4.1	3.8	4.0±0.21
IOD	3.1	2.8	2.8	2.9	2.9±0.14	3.2	3.4	3.3±0.14
IND	3.2	3.0	3.2	3.0	3.1±0.12	3.2	3.5	3.4±0.21
UEW	2.9	2.7	2.8	3.0	2.9±0.13	3.3	3.5	3.4±.14
EN	1.6	1.5	1.7	1.8	1.7±0.13	1.7	2.1	1.9±0.28
TMP	1.8	1.8	1.8	1.9	1.8±0.05	2.1	2.2	2.2±0.07
TEY	1.2	1.2	1.2	1.2	1.2±.00	1.5	1.6	1.6±0.07
HND	7.4	7.1	7.4	7.2	7.3±0.15	8.1	8.2	8.2±0.07
LAHL	13.6	13.0	13.5	13.2	13.3±0.28	15.6	15.1	15.4±0.35
TIB	13.0	12.4	12.5	12.1	12.5±0.37	15.3	16.0	15.7±0.49
ML	12.8	11.3	12.1	12.1	12.1±0.61	13.9	14.4	14.2±0.35
FL	10.5	11.1	10.4	10.9	10.7±0.33	12.9	12.6	12.8±0.21
LFT	16.9	17.1	16.4	16.4	16.7±0.36	20.1	21.1	20.6±0.71
HLL	41.7	39.0	40.3	39.6	40.2±1.16	48.7	51.2	50.0±1.77
HDL/HDW	0.99	0.99	0.97	0.98	0.98±0.01	0.98	0.97	0.98±0.01
HDL/SVL	0.38	0.36	0.38	0.38	0.38±0.01	0.38	0.37	0.37±0.01
TMP/EYE	0.58	0.58	0.51	0.53	0.55±.03	0.51	0.58	0.55±0.05
LAHL/SVL	0.54	0.53	0.56	0.53	0.54±0.01	0.55	0.52	0.54±0.02
HLL/SVL	1.67	1.60	1.66	1.58	1.63±0.04	1.73	1.77	1.75±0.03
TIB/SVL	0.52	0.51	0.51	0.48	0.51±0.02	0.54	0.55	0.55±0.01
LFT/SVL	0.68	0.70	0.67	0.65	0.68±0.02	0.71	0.73	0.72±0.01

Megophrys feii sp. nov. differs frOm the mOst mOrphOlOgically similar species, M. vegrandis frOm nOrtheast India , by the presence Of a prOtruding prOjectiOn pOsteriOr tO the clOaca in bOth seXes (vs. absence in male M. vegrandis ), tOngue nOt nOtched pOsteriOrly (vs. weakly nOtched pOsteriOrly), abdOmen scattered with small dark brOwn spOts (vs. large dark brOwn blOtches present On ventrOlateral sides Of abdOmen), presence Of distinct skin asperities present On dOrsal surfaces Of head, bOdy and limbs (vs. absence Of asperities On all surfaces), cOmparatively lOng head in males (HDL/SVL 36.1–38.3% vs. 33.3–36.0%, MahOny et al . 2013), and cOmparatively shOrt hand in males (HAL/SVL 28.7–30.5% vs. 30.7–32.7%, MahOny et al . 2013). Megophrys feii sp. nov. differs frOm the small-sized M. wuliangshanensis by having the ventral surfaces Of hands and feet withOut Orange-red pigmentatiOn in life (vs. ventral surfaces Of the tips Of fingers and tOes, thenar and palmar tubercles On hand, and metatarsal tubercles On feet distinctly cOlOred Orange-red in life, Fei & Ye 2016), and cOmparatively lOng fOrelimbs (mean LAHL/SVL 54% in bOth seXes vs. 48.2% in males and 45.7% in females, Fei et al . 2009). FIGURE 4. AdvErtIsEmEnt cAll Of thE mAlE pArAtypE SYS A006528 Of Megophrys feii sp. nov. : (A) WAvEfOrm And spEctrOgrAm shOWIng tEn cAlls; (B) WAvEfOrm And spEctrOgrAm shOWIng A sInglE cAll. Megophrys feii sp. nov. differs frOm the small-sized M. rubrimera by having the dOrsal surfaces Of the head, bOdy and limbs strOngly granular (vs. weakly granular), dark brOwn marking On the dOrsum nOt edged with light cOlOr (vs. bOrder Of marking beige), and thenar and metatarsal tubercles nOt cOlOred in Orange-red in life (vs. distinctly cOlOred Orange-red). Bioacoustic comparison. The male advertisement call Of Megophrys feii sp. nov. differs frOm M. elfina by having a call duratiOn Of 115–150 ms at 18˚C (vs. 25–112 ms at 11.3–17.5˚C); frOm M. minor by having a duratiOn Of 115–150 ms (vs. 75–110 ms), a call repetitiOn rate Of 2.0 calls/s (vs. average 4 calls/s), 14–20 pulses per call (vs. 8–16), a dOminant frequency Of 4.74–4.91 kHz at 18˚C (vs. 3.4–3.5 kHz at 14˚C); frOm M. oropedion by having a duratiOn Of 115–150 ms (vs. 30–55 ms), a call repetitiOn rate Of 2.0 calls/s (vs. 3–4 calls/s), 14–20 pulses per call (vs. 6–10), and a dOminant frequency Of 4.74–4.91 kHz at 18˚C (vs. 3.1–4.3 kHz at 18˚C); frOm M. rubrimera by having a duratiOn Of 115–150 ms (vs. 56–85 ms), a call repetitiOn rate Of 2.0 calls/s (vs. 3.1–3.4 calls/s), and a dOminant frequency Of 4.74–4.91 kHz at 18˚C (vs. 3.2–3.4 kHz at 21–22.9˚C); frOm M. synoria by having a duratiOn Of 115–150 ms (vs. 37–85 ms), a call repetitiOn rate Of 2.0 calls/s (vs. 4.4–6.6 calls/s), and a dOminant frequency Of 4.74–4.91 kHz at 18˚C (vs. 3.6–3.9 kHz at 21˚C).