Leptolalax rowleyae, Nguyen & Dzung Trung Le & Vo & Phan & Duong & Murphy & Nguyen, 2018

Nguyen, Luan Thanh, Dzung Trung Le, Nikolay A. Poyarkov Jr., Vo, Ba Dinh, Phan, Hoa Thi, Duong, Tang Van, Murphy, Robert W. & Nguyen, Sang Ngoc, 2018, A new species of Leptolalax (Anura: Megophryidae) from Son Tra Peninsula, central Vietnam, Zootaxa 4388 (1), pp. 1-21 : 7-17

Leptolalax rowleyae sp. nov.

FIgS. 2–5 ; TAbLeS 3–5.

Holotype. ITBCZ 2783, ADULT MALe, COLLeCTeD WHILe CALLINg fROM A SMALL HOLe UNDeR A ROCK NeAR THe HeADWATeR Of A SMALL STReAM WITH 1.5– 2.0 M IN WIDTH IN eVeRgReeN TROPICAL fOReST IN SON TRA NR, DA NANg CITY, VIeTNAM (16.12754°N, 108.24634°E, CA. 380 M A.S.L.); COLLeCTeD ON 0 8 JANUARY 2013 bY LUAN T. NgUYeN.

Paratypes. FOURTeeN SPeCIMeNS, ALL COLLeCTeD fROM SON TRA NR, DA NANg CITY, VIeTNAM: ITBCZ 4502 , A gRAVID feMALe, ReCORDeD ON fOReST fLOOR, AbOUT 5 M IN DISTANCe fROM A SMALL STReAM (16.12248°N, 108.29821°E, CA. 450 M A.S.L.), COLLeCTeD ON 10 AUgUST 2013 bY LUAN T. NgUYeN ; ITBCZ 2779–81, THRee ADULT MALeS (ALL COLLeCTeD WHILe CALLINg), ReCORDeD NeAR A CASCADe STReAM IN TROPICAL eVeRgReeN fOReST (16.13538°N, 108.25198°E, CA. 440 M A.S.L.), COLLeCTeD ON 0 3 JANUARY 2012 bY LUAN T. NgUYeN AND HOA T. PHAN; ITBCZ 2782, ITBCZ 2784–9 , SeVeN ADULT MALeS (ALL ObSeRVeD AND COLLeCTeD WHILe CALLINg), COLLeCTeD ON 0 9 JANUARY 2013 AT THe SAMe LOCATION AS THe HOLOTYPe bY LUAN T. NgUYeN ; ITBCZ 4471, A gRAVID feMALe AND ITBCZ 4472–3, TWO jUVeNILeS, ReCORDeD ON fOReST fLOOR NeAR A SMALL STReAM (CA. IN 20 M fROM THe STReAM) (16.13534°N, 108.25222°E, CA. 640 M A.S.L.), COLLeCTeD ON 11 SePTeMbeR 2013 bY LUAN T. NgUYeN AND MANH V. Le.

Diagnosis. Leptolalax rowleyae sp. nov. IS DISTINgUISHeD fROM ALL Of ITS CONgeNeRS bY A COMbINATION Of THe fOLLOWINg MORPHOLOgICAL CHARACTeRS: (1) ADULT SVL 23.4–25.4 MM IN MALeS AND 27.0–27.8 MM IN feMALeS; (2) PReSeNCe Of DISTINCT DARK/bROWN DORSOLATeRAL MARKINgS WITH bLACK SPOTS ON fLANKS; (3) PINKISH MILK-WHITe TO LIgHTLY bROWN CHeST AND beLLY WITH NUMeROUS WHITe SPeCKLeS; (4) TYMPANUM eXTeRNALLY DISTINCT; (5) NO WebbINg AND LATeRAL DeRMAL fRINgeS ON fINgeRS AND TOeS; (6) PeCTORAL gLANDS COMPARATIVeLY SMALL, COMPRISINg 2.6–3.7% Of SVL IN ADULT MALeS; (7) VeNTROLATeRAL gLANDS INDISTINCT; (8) IRIS bICOLOReD WITH COPPeR-ORANge UPPeR HALf, fADINg TO gOLDeN IN ITS LOWeR THIRD. THe ADVeRTISeMeNT CALL Of THe NeW SPeCIeS, CONSISTINg Of 4–6 NOTeS, LACKINg A DISTINCT INTRODUCTORY NOTe, WITH AN AVeRAge DOMINANT fReqUeNCY Of 3.2–3.5 KHZ, ALSO DISTINgUISHeS THe NeW SPeCIeS fROM Leptolalax SPeCIeS fOR WHICH CALLS ARe KNOWN. THe NeW SPeCIeS IS ALSO MARKeDLY DISTINCT fROM ALL CONgeNeRS fOR WHICH COMPARAbLe SeqUeNCeS ARe AVAILAbLe (MITOCHONDRIAL geNe 16S RRNA; UNCORReCTeD geNeTIC DISTANCe ±7.4%).

Description of holotype. SMALL-SIZeD SPeCIMeN IN gOOD STATe Of PReSeRVATION; HAbITUS STOCKY ( FIg. 2 ). VeNTRAL SURfACe Of HOLOTYPe DISSeCTeD ON RIgHT SIDe beLLY, DISSeCTION-LeNgTH CA. 2.0 MM ( FIg. 2B ). Head. HeAD WIDTH SLIgHTLY SHORTeR THAN HeAD LeNgTH (HDW/HDL 0.91), fLATTeNeD; SNOUT SLIgHTLY TRUNCATe IN DORSAL VIeW AND ROUNDeD IN PROfILe, PROjeCTINg SLIgHTLY beYOND MARgIN Of LOWeR jAW; NOSTRIL OVAL-SHAPeD, SLIgHTLY ROUNDeD, LOCATeD MUCH CLOSeR TO SNOUT THAN TO eYe; CANTHUS ROSTRALIS INDISTINCT, geNTLY ROUNDeD; LOReAL RegION CONCAVe, geNTLY SLOPINg; PUPIL VeRTICAL; eYe DIAMeTeR NOTAbLY SMALLeR THAN SNOUT LeNgTH (EYE/SNT 0.66); TYMPANUM DISTINCT, ROUNDeD, TYMPANUM DIAMeTeR SMALLeR THAN THAT Of eYe; TYMPANIC RIM eLeVATeD ReLATIVe TO SKIN Of TeMPORAL RegION; VOMeRINe TeeTH AbSeNT; PINeAL OCeLLUS AbSeNT; VOCAL SAC OPeNINgS, OVAL, LOCATeD LATeRALLY ON fLOOR Of MOUTH; TONgUe WIDe WITH bROAD, SHALLOW NOTCH AT POSTeRIOR TIP; SUPRATYMPANIC fOLD fORMINg A DISTINCT WIDe RIDge, RUNNINg fROM POSTeRIOR CORNeR Of eYe POSTeRIORLY TOWARDS DORSAL eDge Of TYMPANUM, geNTLY CURVINg DOWNWARDS TOWARDS AXILLA; SUPRATYMPANIC RIDge COMPARATIVeLY SMOOTH WITH feW fLAT TUbeRCLeS ( FIg. 2A–C ). Forelimbs. FOReLIMbS THIN, SLeNDeR; fINgeR TIPS ROUNDeD, VeRY SLIgHTLY SWOLLeN; ReLATIVe fINgeR LeNgTHS: I <II <IV <III; NUPTIAL PAD AbSeNT; SUbARTICULAR TUbeRCLeS AbSeNT; A LARge, ROUND INNeR PALMAR TUbeRCLe SePARATeD fROM SMALL, LATeRALLY COMPReSSeD OUTeR PALMAR TUbeRCLe bY A THIN MeDIAL gROOVe; fINgeR WebbINg AND DeRMAL fRINgeS AbSeNT ( FIg. 2E ). Hindlimbs. HINDLIMbS SLeNDeR, TIbIA NeARLY HALf Of SNOUT-VeNT LeNgTH (TIB/SVL 0.49). TOe TIPS SIMILAR THOSe Of fINgeRS; ReLATIVe TOe LeNgTHS: I <II <V <III <IV; SUbARTICULAR TUbeRCLeS AbSeNT, RePLACeD bY DISTINCT DeRMAL RIDgeS; LARge, OVAL INNeR MeTATARSAL TUbeRCLe PReSeNT, OUTeR MeTATARSAL TUbeRCLe AbSeNT; bASAL WebbINg PReSeNT beTWeeN ALL fIVe TOeS, Web SLIgHTLY DISTINCT beTWeeN TOeS II–III, AND gReATLY ReDUCeD beTWeeN TOeS I–II, II–IV, AND IV–V; NO LATeRAL fRINgeS SeeN ON ANY TOe ( FIg. 2F ). Skin texture and skin glands. SKIN ON DORSUM MOSTLY SMOOTH WITH NUMeROUS TINY TUbeRCLeS AND PUSTULeS fINeLY AND ReLATIVeLY eVeNLY SCATTeReD ON DORSAL AND LATeRAL SURfACeS Of TRUNK, HeAD AND LIMbS; UPPeR eYeLID WITH NUMeROUS SMALL TUbeRCLeS; SKIN ON VeNTRAL SURfACeS Of TRUNK, HeAD AND LIMbS COMPLeTeLY SMOOTH; PeCTORAL gLANDS LOCATeD AT fOReLIMb bASIS ON VeNTRAL SURfACe Of AXILLARY RegION, ROUNDeD, 0.8 MM IN DIAMeTeR, DISTINCT IN LIfe AND IN PReSeRVATIVe; feMORAL gLANDS SMALL, OVAL, APPROXIMATeLY 0.9 MM IN DIAMeTeR, LOCATeD ON POSTeROVeNTRAL SURfACeS Of THIgHS, CLOSeR TO KNee THAN TO VeNT, DISTINCT IN LIfe AND IN PReSeRVATIVe; SUPRA-AXILLARY gLAND LOCATeD IN AXILLARY RegION DORSALLY fROM INSeRTION Of fOReLIMb, OVAL, fLATTeD, 1.1 MM IN DIAMeTeR. VeNTROLATeRAL gLANDS INDISTINCT, ROUNDeD, fLAT, fORMINg AN INDISTINCT LINe ( FIg. 2A–C ).

Color of holotype in life. COLORATION Of HOLOTYPe ReCORDeD AT NIgHT jUST AfTeR CAPTURe ( FIg. 2F,G ). DORSAL SURfACe Of TRUNK LIgHT bROWN; ONe V-SHAPeD INTeRORbITAL bAR MARKINg WITH PALe CReAM eDgINg ANTeRIORLY; SNOUT DORSALLY beIge; DARK-bROWN W-SHAPeD MARKINg AT SCAPULAR RegION WITH INDISTINCT LIgHT-bROWN eDgINg. DARK bLACKISH-bROWN LINe WITH UNCLeAR bORDeRS LATeRALLY ALONg CANTHUS ROSTRALIS, eNCOMPASSINg NAReS RUNNINg TOWARDS ANTeRIOR CORNeR Of eYe AND A LIgHTeR bROWN LINe MeDIALLY ALONg CANTHUS ROSTRALIS RUNNINg TOWARDS UPPeR eYeLID; DISTINCT bLACKISH SPOT IN ANTeRIOR CORNeR Of eYe; DARK bLACKISH STRIPe beLOW SUPRATYMPANIC RIDge RUNNINg fROM POSTeRIOR CORNeR Of eYe TOWARDS TYMPANUM, TeRMINATINg AbOVe AXILLA, DARK LINe eNCOMPASSINg MOST Of TYMPANUM fORMINg A CHARACTeRISTIC DIAMOND-SHAPeD DARK MARKINg IN TeMPORAL RegION; UPPeR eYeLID LIgHT bROWN WITH SMALL WHITISH SPOTS ON CILIARY eDge; CANTHUS ROSTRALIS ReDDISH-bROWN; DORSAL SURfACeS Of fOReLIMbS (LOWeR ARMS) AND HINDLIMbS LIgHTeR bROWNISH, fOReARMS, AS WeLL AS DORSAL SURfACeS Of fINgeRS AND TOeS MUCH LIgHTeR WITH ORANge bACKgROUND COLOR AND SMALL ReDDISH-bROWN TUbeRCLeS; TRANSVeRSe DARK-bROWN bARS ON DORSAL SURfACe Of THIgHS, TIbIA, TARSUS, LOWeR ARMS, fINgeRS AND TOeS; WHeN HINDLIMb IS fLeXeD, TRANSVeRSe DARK bARS ON DORSAL SURfACeS Of THIgHS CONTINUe ON DORSAL SURfACeS Of SHANKS AND TARSUS ( FIg. 2F ); eLbOWS AND UPPeR ARMS ORANge WITH LIgHT ReDDISH-bROWN SPOTS ON TUbeRCLeS; LATeRAL SURfACeS Of TRUNK LIgHT PURPLISH-bROWN; VeNTRAL SURfACeS LIgHTLY COLOReD, PINKISH MILK-WHITe WITH DeNSe WHITISH SPeCKLINg eVeNLY SCATTeReD ON eNTIRe VeNTRAL SURfACe INCLUDINg THROAT, CHeST, ARMS, LegS, AND jAWS; WHITISH fLeCKS geTTINg LARgeR AND eSPeCIALLY NUMeROUS ON LATeRAL SIDeS Of bODY. TIP Of fINgeRS AND TOeS ARe WHITISH-PURPLe. LATeRAL SIDeS Of bODY WITH 7 DISTINCT bLACKISH-bROWN bLOTCHeS ON eACH SIDe (0.3–1.4 MM IN DIAMeTeR) LOCATeD fROM AXILLA TO gROIN, MOST POSTeRIOR bLACK SPOT LOCATeD IN gROIN AReA IS AbOUT 4.5 TIMeS LARgeR THAN OTHeR SPOTS. THRee DISTINCT bLACKISH-gReY SPOTS ON LIgHT bROWN bACKgROUND COLOR ON eDge Of UPPeR jAW LOCATeD fROM SNOUT TO AReA UNDeR eYe; MIDDLe ONe LASTINg DORSALLY TOWARDS NARIS eNCOMPASSINg MOST PART Of LOReAL AReA ( FIg. 2F ). SUPRA-AXILLARY gLAND MILK-WHITe; feMORAL, PeCTORAL, AND VeNTROLATeRAL gLANDS WHITe. IRIS bICOLOReD: COPPeR-ORANge IN UPPeR HALf, fADINg TO gOLDeN IN LOWeR THIRD; fINe bLACK ReTICULATIONS THROUgHOUT IRIS.

Color of holotype in preservative. IN PReSeRVATION COLORATION fADeS TO DARK gReY-bROWN ON DORSUM AND fLANKS ( FIg. 2A–E ); IRIS COLORATION fADeS TO COMPLeTeLY DARK bROWN. BANDINg ON LIMbS fADe TO bROWNISH COLOR AND TURN LeSS DISTINCT; WHITe fLeCKS ON bODY fLANKS AND VeNTRAL SIDeS ReMAIN PRONOUNCeD IN PReSeRVATIVe. VeNTRAL SURfACe Of CHeST, beLLY, THROAT, VeNTRAL PORTIONS Of ARMS AND THIgHS fADe TO WHITISH-bROWN; MACROgRANDS TURN WHITISH.

Variation. MeASUReMeNTS Of THe TYPe SeRIeS ARe SHOWN IN TAbLe 2 AND RePReSeNTATIVe PHOTOgRAPHS Of PARATYPeS IN LIfe ARe SHOWN IN FIg. 3 . SPeCIMeNS VARY IN bODY SIZe, NUMbeR AND SIZe Of bLACK VeNTROLATeRAL bLOTCHeS AND SLIgHTLY IN COLOR AND PATTeRN IN LIfe. MALeS ITBCZ 2779–80 HAVe DARKeR bROWN COLORATION Of DORSAL SURfACeS AND LIgHT-bROWN VeNTRAL SURfACe WITH DeNSe WHITISH SPeCKLINg COMPAReD WITH THe HOLOTYPe . MALe ITBCZ 2787 HAS INDISTINCT DARK V-SHAPeD INTeRORbITAL bAR; MALe ITBCZ 2782 HAS LeSS DISTINCT DARK bARS ON LIMbS, WHILe MALe ITBCZ 2788 HAS VeRY THIN ALMOST INDISTINCT DARK bARS ON LIMbS. MALe ITBCZ 2784 HAS LIgHT-bROWNISH DORSAL SURfACe WITH TWO bIg PALe MARKINgS beTWeeN AXILLAe AND ReDDISH-bROWN TINT ON bODY fLANKS. MALeS ITBCZ 2780–83 HAVe WHITISH-bROWN COLORATION ON VeNTRAL SURfACeS. FeMALeS TeND TO HAVe LARgeR bODY SIZe (SVL 27.0–27.8; N =2) THAN MALeS (SVL 23.4–25.4; N =11). JUVeNILe ITBCZ 4473 HAS LIgHTeR DORSAL COLORATION COMPAReD TO HOLOTYPe. AS IN MANY OTHeR SPeCIeS Of Leptolalax , COLORATION Of DORSAL SURfACeS IS LIgHTeR AT NIgHT AND geTS DARKeR IN DAYTIMe (IN ALL SPeCIMeNS), WHILe VeNTRAL COLORATION Of ALL SPeCIMeNS beCOMeS DARKeR AND CHANgeS COLOR fROM WHITISH TO ReDDISHbROWN OR bROWN AfTeR CAPTURe IN DAYTIMe OR AfTeR eUTHANIZATION. SKIN TeXTURe APPeARS TO be LeSS TUbeRCULATe IN PReSeRVATIVe THAN IT IS IN LIfe.

MALeS Of L. rowleyae sp. nov. WHeN CALLINg INfLATe A COMPARATIVeLY LARge SINgLe SUbgULAR VOCAL SAC ( FIg. 3C ).

Advertisement call. AS IN OTHeR MeMbeRS Of L. applebyi SPeCIeS gROUP, THe ADVeRTISeMeNT CALL Of THe NeW SPeCIeS RePReSeNTS A SeRIeS Of fAINT, qUITe RASPINg CLICKINg SIgNALS, TO HUMAN eAR SUPeRfICIALLY ReSeMbLINg A CALL Of AN ORTHOPTeRAN. CALL DeSCRIPTION IS bASeD ON THe ReCORDINg Of THe ADVeRTISeMeNT CALL Of THe HOLOTYPe, TAKeN AT 21.5 °C AMbIeNT TeMPeRATURe. THe NeW SPeCIeS HAS SINgLe TYPe Of ADVeRTISeMeNT CALLS (N =10) CONSISTeD Of 4–6 NOTeS (MeAN 5.1 NOTeS PeR CALL; N =51), OR CLICKS, Of VARIAbLe (276–435 MS, MeAN 363 MS) DURATION, RePeATeD AT A RATe Of 8.6–11.3 (MeAN 11.2) NOTeS PeR SeCOND ( FIg. 4 ). A DISTINCT INTRODUCTORY NOTe WAS NOT DISCeRNIbLe. NOTe DURATION VARIeD fROM 4–96 MS (MeAN 20 MS; N =51) AND CONSISTeD Of 1–7 DISTINCT PLUSeS (MeAN 2.1 PULSeS PeR NOTe; N =51). THe DOMINANT fReqUeNCY WAS 3.3 KHZ, HARMONICS WeRe NOT CLeAR bUT CAN be DeTeCTeD AT APPROXIMATeLY 10.5 AND 13.8 KHZ, AND THe fUNDAMeNTAL fReqUeNCY WAS NOT eVIDeNT. CALLS HAD AN AVeRAge INTeRCALL INTeRVAL Of 179 MS (68–428 MS; N =10). NOTe INTeRVAL AND THe NUMbeR Of NOTeS PeR CALL VARIeD SLIgHTLY WITHIN CALLINg bOUTS AND AMONg INDIVIDUALS. TAbLe 4 SHOWS THe VARIATION Of OTHeR CALL CHARACTeRISTICS AMONg eXAMINeD INDIVIDUALS Of THe NeW SPeCIeS.

Tadpole description. DeSCRIPTION Of LARVAL MORPHOLOgY IS bASeD ON fOUR TADPOLeS ITBCZ 2790, ITBCZ 3579– 81; GOSNeR (1960) STAgeS 27–38 ( FIg. 5 ). TADPOLeS WeRe COLLeCTeD IN A MOUNTAIN STReAM, WHILe HIDINg beTWeeN PebbLeS ON THe STReAMbeD. TADPOLeS Of THe NeW SPeCIeS WeRe ACTIVe AT NIgHT AND AVOIDeD LIgHT bY HIDINg THeMSeLVeS beTWeeN ROCKS OR DeAD LeAVeS. Dorsal view. BODY Of TADPOLe ReACHeS A LeNgTH Of 13.8 MM IN THe STAge 31; bODY eLLIPTICAL, eLONgATeD (BW/BL 0.55–0.87, MeAN 0.69; N =4), WITH A ROUNDeD SNOUT. NAReS ANTeRODORSALLY POSITIONeD, NeAReR TO TIP Of SNOUT THAN TO PUPIL (RND/NPD 0.32–0.53; IND/IP 0.73–0.93). EYeS DORSOLATeRAL, LOCATeD AT THe POSTeRIOR eDge Of THe ANTeRIOR ONe fOURTH Of bODY LeNgTH. EYeS COMPARATIVeLY SMALL AND DISTINCT (ED/BL 0.06–0.12, MeAN 0.08; IP/BW 0.46–0.51, MeAN 0.48; N =4). THe MAXIMUM WIDTH Of TAIL MUSCULATURe COMPRISeS LeSS THAN A HALf Of bODY WIDTH (TMW/BW 0.42–0.46, MeAN 0.44; N =4) ( FIg. 5B ). NARIAL APeRTUReS WITH CHARACTeRISTIC geNTLY ROUNDeD RIMS, SLIgHTLY LIfTeD AbOVe bODY SURfACe. Lateral view. BODY LATeRALLY DePReSSeD (BH/BW 0.76–0.83, MeAN 0.79; N =4). CONe-SHAPeD SPIRACLe SINISTRAL, DIReCTeD POSTeRODORSALLY, POSITIONeD LATeRALLY AT POSTeRIOR HALf Of bODY (SS/BL 0.53–0.71, MeAN 0.61; N =4); CONICAL TUbe Of SPIRACLe IS fUSeD TO bODY WITH A fRee SHORT DISTAL PORTION. VeNT TUbe DeXTRAL, SePARATeD fROM bODY AND ATTACHeD TO LOWeR fIN ( FIg. 5A ). Tail. TAIL MUSCULAR, LONg, TAIL LeNgTH eXCeeDINg TWO TIMeS bODY LeNgTH (TAL/BL 2.21–2.95, MeAN 2.60; N =4), WITH A POINTeD TAIL TIP. TAIL MUSCULATURe DISTINCT (MTH/BH 0.52–0.71, MeAN 0.62; N =4), RUNNINg PARALLeL fROM TAIL bASe TO ANTeRIOR TWO THIRDS Of TAIL LeNgTH, THeN gRADUALLY TAPeRINg TO THe TIP Of TAIL. TAIL fINS ReLATIVeLY LOW; UPPeR TAIL fIN STARTS POSTeRIORLY Of TAIL bASe, UPPeR TAIL fIN eDge SLIgHTLY CONVeX; bOTH UPPeR AND LOWeR TAIL fINS ReACH MAXIMUM HeIgHT IN POSTeRIOR THIRD. Oral disc. CUP-LIKe ORAL DISC ANTeROVeNTRALLY POSITIONeD (ODW/BW 0.32–0.37, MeAN 0.34; N =4), fRINgeD WITH SHORT CONICAL PAPILLAe; KeRATODONT ROW fORMULA: 4(2–4)/3(1–2), WITH AbOUT TWO THIN SPIKe-LIKe KeRATODONTS PeR 0.1 MM IN THe STAge 38 ( FIg. 5E ); jAW SHeATHS bLACK, RObUST; UPPeR jAW AND LOWeR jAW SHeATHS ReMARKAbLY DeVeLOPeD WITH DISTINCTLY SeRRATeD eDgeS ( FIg. 5D ). Coloration in life. BODY OLIVe-bROWN TO gReY WITH DISTINCT WHITISH-gOLDeN SPeCKLINg ON DORSAL AND LATeRAL SURfACeS. IRIS DARK, MARbLeD, WITH fOUR bLACK STRIPeS AT THe eXTeRNAL MARgIN. VeNTRAL SIDe Of bODY TRANSLUCeNT, WITH INTeSTINAL SPIRAL CLeARLY VISIbLe. INTeRNAL gILLS DISTINCTLY ReDDISH. TAIL fIN TRANSLUCeNT, gReYISH-bROWN ( FIg. 5A–C ). Coloration in preservative. WHeN PReSeRVeD IN 5% SOLUTION Of fORMALIN, COLORATION fADeS SIgNIfICANTLY, bODY TURNS SeMI-TRANSLUCeNT, DORSALLY AND DORSOLATeRALLY LIgHT bROWN, VeNTRALLY TRANSLUCeNT, TURNINg WHITISH ON THROAT AND CHeST RegION. TAIL MUSCULATURe fADeS TO gReY, UPPeR AND LOWeR TAIL fIN ALMOST TRANSLUCeNT WITH SLIgHT INDISTINCT gReY PIgMeNTATION. DORSOLATeRAL PARTS Of HINDLIMbS WITH WHITISH TO gReYISH PIgMeNTATION. Measurements. THe VARIATION Of MORPHOLOgICAL CHARACTeRS Of THe TADPOLeS IS SHOWN IN TAbLe 5.

Comparisons. Leptolalax rowleyae sp. nov. bOTH MORPHOLOgICALLY AND MOLeCULARLY IS MOST SIMILAR TO THe MeMbeRS Of THe SMALL-bODIeD L. applebyi SPeCIeS gROUP INHAbITINg IN THe CeNTRAL HIgHLANDS Of CeNTRAL AND SOUTHeRN VIeTNAM AND THe NORTHeASTeRN PART Of CAMbODIA INCLUDINg L. applebyi , L. ardens , L. bidoupensis , L. kalonensis , L. maculosus , L. melicus , L. pallidus , L. pyrrhops , AND L. tadungensis . COMPARISONS Of THe NeW SPeCIeS WITH THe MeMbeRS Of L. applebyi SPeCIeS gROUP APPeAR TO be THe MOST PeRTINeNT.

Leptolalax rowleyae sp. nov. CAN be DISTINgUISHeD fROM ALL OTHeR SPeCIeS Of Leptolalax (NON-MeMbeRS Of L. applebyi SPeCIeS gROUP) RePORTeD fOR INDOCHINA AND SURROUNDINg AReAS IN ITS OVeRALL MORPHOLOgY AS fOLLOWS: IN HAVINg SMALL bODY SIZe (SVL 23.4–25.4 MM IN ADULT MALeS, 27.0–27.8 MM IN ADULT feMALeS), Leptolalax rowleyae sp. nov. CAN be DISTINgUISHeD fROM THe LARgeR CONgeNeRS INCLUDINg L. bourreti (MALeS 28.0–36.2 MM, feMALeS 42.0– 45.0 MM), L. botsfordi (MALeS 29.1–32.6 MM, feMALeS 30.0–31.8 MM), L. eos (MALeS 33.1–34.7 MM, feMALe 40.7 MM), L. firthi (MALeS 26.4–29.2 MM, feMALeS 25.7–36.9 MM), L. fuliginosus (MALeS 28.2–30.0 MM), L. minimus (MALeS 25.7–31.4 MM, feMALeS 31.6–37.3 MM), L. nahangensis (MALe 40.8 MM), L. nyx (MALeS 26–30 MM, feMALeS 36–42 MM), L. pelodytoides (MALeS 27.5–32.3 MM, feMALeS 35.5–37.8 MM), L. petrops (MALeS 23.6–27.6 MM, feMALeS 30.3–47.0 MM); L. platycephalus (MALe 35.1 MM, feMALe 46.0 MM), L. puhoatensis (MALeS 24.2–28.1 MM, feMALeS 27.3–31.5 MM), L. solus (MALe 27.6 MM), AND L. sungi (MALeS 48.3–52.7 MM, feMALeS 56.7–58.9 MM), AND fROM THe SMALLeR SPeCIeS INCLUDINg L. kecil (MALeS 19.3–20.5 MM, feMALe 25 MM) AND L. pluvialis (MALeS 21.3–22.3 MM).

IN HAVINg PINKISH MILK-WHITe TO LIgHTLY bROWN CHeST AND beLLY WITH NUMeROUS WHITe SPeCKLeS, THe NeW SPeCIeS ALSO DIffeRS fROM L. aereus , L. bourreti , L. firthi , L. fuliginosus , L. mininus , L. nahangensis , L. nyx , L. pelodytoides , L. solus , L. sungi , L. tuberosus , AND L. zhangyapingi (ALL Of WHICH HAVe MOSTLY WHITe, CReAMY WHITe OR PALe gReY VeNTeRS, WITHOUT WHITe SPeCKLINg AND WITH OR WITHOUT DARK SPOTS OR MOTTLINg); fROM L. botsfordi (VS. ReDDISH-bROWN VeNTRAL SURfACeS WITH WHITe SPeCKLINg); fROM L. croceus (VS. ORANge beLLY); fROM L. eos (VS. CReAMY WHITe beLLY WITH bROWN SPeCKLINg ALONg ITS MARgINS); fROM L. petrops (VS. PALe PINK AND SLIgHTLY TRANSLUCeNT beLLY, PARTICULARLY AT eITHeR SIDe Of THROAT; eDgeS Of THROAT PALe bROWN WITH WHITe SPeCKLINg, CONCeNTRATeD TOWARDS SNOUT; VeNTRAL SURfACe Of CHeST AND AbDOMeN IMMACULATe WHITe); fROM L. pluvialis (VS. gReY VeNTeR WITH DARK gReY MARbLINg, UNIfORM PALe gReY THROAT WITH SPeCKLINg AROUND ITS MARgINS); fROM L. puhoatensis (VS. ReDDISH-bROWN, OfTeN WITH fAINT WHITe DUSTINg IN MALeS AND A PALe PINK IN feMALeS); fROM L. melanoleucus AND L. ventripunctatus (bOTH DISPLAY LARge PATCHeS Of DISTINCT bROWN AND WHITe MARbLINg); fROM L. heteropus (VS. gReY VeNTeR, SPeCKLeD WITH bLACK); AND fROM L. kecil (VS. A UNIfORMLY DARK VeNTeR WITH LARge, DARK ORANge PeCTORAL gLANDS). BY HAVINg bLACK MARKINgS (SPOTS OR bLOTCHeS) ON THe fLANKS, THe NeW SPeCIeS CAN be fURTHeR DIffeReNTIATeD fROM L. aereus , L. croceus , L. eos , L. firthi , L. isos , AND L. tuberosus , ALL Of WHICH DO NOT HAVe THe MARKINgS.

IN HAVINg A DISTINCT TYMPANUM, THe NeW SPeCIeS CAN be eASILY DIAgNOSeD fROM L. croceus AND L. tuberosus (THeSe TWO SPeCIeS TYMPANUM IS HIDDeN) AND fROM L. sungi (WHICH IS RePORTeD TO LACK eXTeRNAL TYMPANUM, HOWeVeR IN SOMe POPULATIONS AN INDISTINCT TYMPANUM IS STILL DISCeRNIbLe). IN HAVINg TOeS WITH bASAL WebbINg AND NO LATeRAL fRINgINg, Leptolalax rowleyae sp. nov. CAN be DISTINgUISHAbLe fROM L. eos , L. firthi , AND L. isos , ALL Of WHICH HAVe MORe eXTeNSIVe TOe WebbINg AND DISTINCT LATeRAL fRINgeS ON TOeS.

IN HAVINg DORSAL SKIN MOSTLY SMOOTH WITH NUMeROUS TINY TUbeRCLeS, THe NeW SPeCIeS IS ALSO DISTINgUISHAbLe fROM L. croceus , L. isos , L. minimus , L. petrops , L. solus , L. tuberosus , AND L. ventripunctatus (ALL THeSe SPeCIeS HAVe ROUgHLY gRANULAR OR TUbeRCULATe SKIN TeXTURe WITH DISTINCT SKIN RIDgeS). BY HAVINg bICOLOReD IRIS, THe NeW SPeCIeS CAN be DISTINgUISHeD fROM SPeCIeS WITH UNIfORM IRIS COLORATION, SUCH AS: L. aereus (VS. UNIfORMLY bRONZe IRIS), L. botsfordi (VS. UNIfORMLY bROWNISH-gOLDeN IRIS), L. firthi (VS. bRIgHT gOLDeN IRIS), L. kecil (VS. DARK ReD IRIS), L. nahangensis (VS. UNIfORMLY gOLDeN IRIS), L. platycephalus (VS. ORANge ReD OR ReDDISH bROWN IRIS), L. pluvialis (VS. UNIfORMLY DARK gOLDeN IRIS), L. sungi (VS. gOLDeN-gReeN IRIS), AND L. tuberosus (VS. gOLDeN IRIS).

FROM MeMbeRS Of L. applebyi SPeCIeS gROUP, Leptolalax rowleyae sp. nov. CAN be DISTINgUISHeD fROM ALL OTHeR SPeCIeS Of THe gROUP NOT ONLY bY MTDNA SeqUeNCe DIffeReNCeS bUT ALSO bY bOTH MORPHOLOgICAL ATTRIbUTeS (ALL fOLLOWINg MORPHOMeTRIC DIffeReNCeS RefeR TO MALeS ONLY) AND bIOACOUSTICS (See beLOW).

THe NeW SPeCIeS DIffeRS fROM L. applebyi , L. ardens , AND L. melicus bY HAVINg A LARgeR SIZe (23.4–25.4, MeAN 24.4 MM, N= 13 IN THe NeW SPeCIeS VS. 19.6–22.3 MM, MeAN 20.5 MM, N=9 [ROWLeY et al. 2016]; 17.7–20.8 MM, MeAN 19.4 MM, N=10 [OUR DATA] IN L. applebyi ; 21.3–24.7 MM, MeAN 22.8 MM, N=16 IN L. ardens ; AND 19.5–22.7 MM, MeAN 20.7 MM, N= 8 IN L. melicus ); COMPARATIVeLY LARgeR HeAD WIDTH (HDW/SVL 0.35–0.38, MeAN 0.36 [N= 13] IN THe NeW SPeCIeS VS. 0.34–0.35, MeAN 0.35 IN L. applebyi [N= 9]); AND CAN be fURTHeR DIAgNOSeD fROM L. kalonensis AND L. pyrrhops bY HAVINg SMALLeR bODY SIZe (23.4–25.4 MM IN THe NeW SPeCIeS VS. 25.8–30.6 IN L. kalonensis AND 30.8–34.3 MM, MeAN 33.2 MM, N= 7 IN L. pyrrhops ).

IN LACKINg LATeRAL fRINgeS ON TOeS, Leptolalax rowleyae sp. nov. IS DISTINgUISHeD fROM L. bidoupensis , L. maculosus , AND L. tadungensis (THeSe THRee SPeCIeS HAVe WeAK, SLIgHT OR DISTINCT LATeRAL fRINgeS ON TOeS). THe NeW SPeCIeS ALSO DIffeRS fROM L. pallidus AND L. pyrrhops bY HAVINg MOSTLY SMOOTH SKIN (VS. COARSeLY SHAgReeNeD IN L. pallidus AND fINeLY SHAgReeNeD TO TUbeRCULATe SKIN IN L. pyrrhops ).

IN HAVINg bICOLOReD IRIS, Leptolalax rowleyae sp. nov. CAN be fURTHeR DISTINgUISHeD fROM L. applebyi , L. ardens , L. melicus , AND L. tadungensis , ALL Of WHICH HAVe UNIfORM COLORATION Of IRIS. IN HAVINg IRIS COLORATION WITH COPPeR UPPeR HALf AND gOLDeN-bROWN IN LOWeR HALf, THe NeW SPeCIeS CAN be DIAgNOSeD fROM L. bidoupensis (VS. ReDDISH IN THe UPPeR HALf, AND SILVeR IN THe LOWeR HALf Of IRIS IN L. bidoupensis ). MOReOVeR, Leptolalax rowleyae sp. nov. IS CLeARLY DISTINCT fROM ALL OTHeR MeMbeRS Of L. applebyi SPeCIeS gROUP eXCePT fOR L. melicus bY HAVINg IN LIfe LIgHT PINKISH, MILK-WHITe TO LIgHT-bROWN COLORATION Of VeNTRAL SURfACeS (VS. DARK bROWNISH-ReD, ReDDISH-bROWN OR PINKISHgReY TO DARK bROWNISH-VIOLeT VeNTRAL COLORATION IN THeSe SPeCIeS).

BY HAVINg A MILK-WHITe SUPRA-AXILLARY gLAND, THe NeW SPeCIeS DIffeRS fROM ALL SPeCIeS Of L. applebyi SPeCIeS gROUP WITH eXCePTION Of L. applebyi AND L. bidoupensis , ALL Of WHICH HAVe COPPeR TO ORANge COLORATION Of THe SUPRA- AXILLARY gLAND. Leptolalax rowleyae sp. nov. IS ALSO DISTINgUISHeD fROM L. ardens AND L. bidoupensis bY HAVINg TINY SMALL WHITe SPOTS ON THe LATeRAL MARgIN Of THROAT (VS. bROKeN ROW Of SLIgHTLY eNLARgeD WHITe SPOTS ON LATeRAL MARgIN Of THROAT IN L. ardens AND L. bidoupensis ); AND fURTHeR DISTINgUISHeD fROM L. bidoupensis bY HAVINg A DISTINCT bLACK SUPRATYMPANIC LINe (VS. INDISTINCT bLACK SUPRATYMPANIC LINe IN THe LATTeR SPeCIeS); AND fROM L. pyrrhops bY HAVINg A LARge POSTeRIOR-MOST bLACK LATeRAL bLOTCH LOCATeD IN THe gROIN AReA, WHICH IS AbOUT 4.5 TIMeS LARgeR THAN THe OTHeR SPOTS IN THe NeW SPeCIeS (VS. gROIN LATeRAL bLOTCH CA. 1.5 TIMeS LARgeR THAN OTHeR LATeRAL SPOTS IN L. pyrrhops ).

THe ADVeRTISeMeNT CALL Of Leptolalax rowleyae sp. nov. IS UNIqUe AMONg CONgeNeRS WITH KNOWN CALLS IN INDOCHINA AND SURROUNDINg AReAS. FROM Leptolalax SPeCIeS OUT Of THe L. applebyi SPeCIeS gROUP, L. rowleyae sp. nov. CAN be DISTINgUISHeD fROM L. croceus , L. fuliginosus , L. heteropus , AND L. tuberosus bY HAVINg HIgHeR DOMINANT fReqUeNCY (3.2–3.5 KHZ, 20.8–21.5 °C IN THe NeW SPeCIeS VS. 2.6–3.0 KHZ, 21.6–25.1 °C IN L. croceus ; 2.3–2.4 KHZ, 19.3–19.6 °C IN L. fuliginosus ; 2.8 KHZ, 21 °C IN L. heteropus ; AND 2.6– 2.8 KHZ, 22.5–24.5 °C IN L. tuberosus ). THe CALL Of Leptolalax rowleyae sp. nov. APPeARS TO HAVe LOWeR fReqUeNCY (See AbOVe) COMPAReD TO THe CALLS ATTRIbUTeD TO L. aereus (6.2–7.9 KHZ, 22.4–25.7 °C), L. firthi (5.4–6.6 KHZ, 18.3–21.2 °C), L. isos (5.8–6.2 KHZ, 22.4–22.8 °C), L. pelodytoides (6.4–6.6 KHZ, 22.7 °C), L. petrops (5.6–6.4 KHZ, 24.5–25.3 °C), AND L. puhoatensis (4.9–5.6 KHZ, 22.3–25.8° C). Leptolalax rowleyae sp. nov. HAS SIMILAR DOMINANT fReqUeNCY WITH L. botsfordi , L. kecil , L. melanoleucus , AND L. solus , bUT THe NeW SPeCIeS SHOWS ONLY A SINgLe TYPe Of RASPINg SIgNAL COMPAReD TO THe TWO TYPeS Of CALLS IN L. botsfordi (HIgHeR AMPLITUDe CLICKINg CALLS AND LOW AMPLITUDe RASPINg CALLS RePORTeD IN L. botsfordi ). Leptolalax rowleyae sp. nov. CAN be fURTHeR DIffeReNTIATeD fROM L. kecil , L. melanoleucus , AND L. solus bY HIgHeR CALL DURATION (276–517 MS IN THe NeW SPeCIeS VS. 110–130 MS IN L. kecil ; 40–63 MS IN L. melanoleucus ; AND 130–260 IN L. solus ); AND HIgHeR NOTe RePeTITION RATeS (8.6–11.3 NOTe/S) COMPAReD TO L. solus (2.1–2.3 NOTe/S).

THe MALe ADVeRTISeMeNT CALL Of L. rowleyae sp. nov. DIffeRS fROM SIgNALS Of ALL OTHeR MeMbeRS Of THe L. appleybi gROUP. BY LACKINg A DISTINCT INTRODUCTORY NOTe, THe CALL Of L. rowleyae sp. nov. CAN be DISTINgUISHeD fROM THAT Of ALL OTHeR MeMbeRS Of L. applebyi gROUP eXCePT fOR L. bidoupensis AND L. maculosus (INTRODUCTORY NOTe RePORTeD fOR CALLS Of L. applebyi , L. ardens , L. kalonensis , L. melicus , L. pallidus , L. pyrrhops , AND L. tadungensis ). THe CALL Of L. rowleyae sp. nov. CAN be fURTHeR DIffeReNTIATeD fROM THe CALLS Of L. kalonensis , L. maculosus , L. pallidus , L. pyrrhops , AND L. tadungensis bY HAVINg HIgHeR DOMINANT fReqUeNCY (3.2–3.5 KHZ, MeAN 3.3 KHZ, 20.8– 21.5 °C IN THe NeW SPeCIeS VS. 2.8 KHZ, 26.4 °C IN L. kalonensis ; 2.7–2.8 KHZ, 23.3–24.1 °C IN L. maculosus ; 2.4– 2.7 KHZ, 14.0–21.4 °C IN L. pallidus ; 1.91–2.23 KHZ, 25 °C IN L. pyrrhops ; AND 2.6–3.1 KHZ, MeAN 2.8 KHZ, 12.9– 22.3 °C IN L. tadungensis ). THe CALL Of L. rowleyae sp. nov. HAS A LOWeR DOMINANT fReqUeNCY COMPAReD TO THAT Of L. applebyi (4.0–4.3 KHZ, 21.5 °C). THe CALL Of Leptolalax rowleyae sp. nov. HAS DOMINANT fReqUeNCY SIMILAR TO THAT Of L. ardens AND L. melicus bUT CAN be DISTINgUISHeD fROM CALLS Of THe LATTeR SPeCIeS bY HAVINg LOWeR NOTe RePeTITION RATe (8.6–11.3 NOTe/S IN THe NeW SPeCIeS VS. 15.1–29.9 NOTe/S IN L. ardens AND 15.9–21.1 NOTe/S IN L. melicus ); THe CALL Of THe NeW SPeCIeS CAN be fURTHeR DIffeReNTIATeD fROM CALLS Of L. bidoupensis bY HAVINg LOWeR INTeRCALL INTeRVAL (68–576 MS VS. 3590–13106 MS IN L. bidoupensis ); LOWeR NOTe RePeTITION RATe (8.6–11.3 NOTe/S VS. 14–21 NOTe/S IN L. bidoupensis ), AND HIgHeR NUMbeR Of PLUSeS PeR NOTe (1–7 PLUSeS PeR NOTe IN THe NeW SPeCIeS VS. SINgLe PLUSeD- NOTe IN L. bidoupensis ). MOReOVeR, THe CALL Of L. rowleyae sp. nov. CAN be fURTHeR DISTINgUISHeD fROM THe CALLS Of L. ardens , L. melicus , L. pallidus , AND L. tadungensis bY HAVINg NO DISTINCT HARMONICS AND fUNDAMeNTAL fReqUeNCY (ALL Of THe LATTeR SPeCIeS HAVe CALLS WITH DISTINCT OR WeAK HARMONICS PReSeNT).

FINALLY, THe NeW SPeCIeS IS MARKeDLY DISTINCT fROM ALL OTHeR CONgeNeRS fOR WHICH COMPARAbLe SeqUeNCeS ARe AVAILAbLe bY ReLATIVeLY LARge geNeTIC DISTANCeS IN 16S RRNA MTDNA geNe fRAgMeNT (p ±7.4%).

Etymology. THe SPeCIfIC ePITHeT IS A PATRONYM HONOURINg JODI J. L. ROWLeY, AN AMPHIbIAN bIOLOgIST IN AUSTRALIAN MUSeUM (SYDNeY), WHO gReATLY CONTRIbUTeD TO THe KNOWLeDge ON INDOCHINeSe AMPHIbIANS AND THe geNUS Leptolalax IN PARTICULAR, DeSCRIbINg TWO-THIRDS Of INDOCHINeSe SPeCIeS Of Leptolalax . We ReCOMMeND “ Rowley’s litter frog ” AS THe COMMON ENgLISH NAMe Of THe NeW SPeCIeS AND THe COMMON NAMe IN VIeTNAMeSe AS “ cóc mày rôly ”.

Distribution and ecology. Leptolalax rowleyae sp. nov. IS CURReNTLY KNOWN ONLY fROM SON TRA PeNINSULA, DA NANg CITY, VIeTNAM. ALL SPeCIMeNS Of THe NeW SPeCIeS WeRe fOUND IN eVeRgReeN TROPICAL fOReSTS Of SON TRA NR ALONg OR IN CLOSe VICINITY fROM CASCADe STReAMS ON eLeVATIONS beTWeeN 380–645 M A.S.L. ( FIg. 6 ). Leptolalax rowleyae sp. nov. OCCURS SYMPATRICALLY WITH Leptobrachium banae LATHROP, MURPHY, ORLOV & HO, Limnonectes poilani (BOURReT), Occidozyga martensii (PeTeRS), Hylarana attigua (INgeR, ORLOV & DAReVSKY), Polypedates SP., Kurixalus banaensis (BOURReT), AND Theloderma vietnamense POYARKOV, ORLOV, MOISeeVA, PAWANgKHANANT, RUANgSUWAN, VASSILIeVA, GALOYAN, NgUYeN & GOgOLeVA. MALeS WeRe fOUND ON THe fOReST fLOOR, HIDINg UNDeR DeAD LeAVeS, USUALLY CALLINg fROM SMALL HOLeS IN THe gROUND OR fROM CReVICeS IN THe ROCKS IN THe HeAD Of DRY STReAM beD OR ALONg CASCADe STReAMS WITH SLOW CURReNT. BOTH THe gRAVID feMALeS WeRe fOUND NeARbY THe SMALL STReAMS WHeRe MALeS WeRe CALLINg, IN AbOUT 10–30 M fROM THe STReAM. CALLINg MALeS WeRe ObSeRVeD fROM AUgUST TO FebRUARY.

TADPOLeS WeRe fOUND IN A SMALL POOL IN THe STReAM beD CONNeCTeD WITH SLOW-CURReNT STReAM IN SON TRA NR, AT AN eLeVATION Of CA. 440 M A.S.L. LARVAe Of Limnontetes poilani AND Leptobrachium banae WeRe fOUND IN THe SAMe STReAM WHeRe TADPOLeS Of THe NeW SPeCIeS WeRe ObSeRVeD.

Figures

FIGURE 2. Holotype of Leptolalax rowleyae sp. nov. in preservative (A–E) and in life (F,G), respectively. A–C, dorsal, ventral, and lateral views of body; D, volar view of right hand; E, thenar view of right foot; F, dorsolateral view; G, ventral view. Note: the semicircle seen in the eye of frog in F is from a ring flash and not a natural coloration. Photos by Luan T. Nguyen.

FIGURE 3. Paratypes of Leptolalax rowleyae sp. nov. in life. A, male ITBCZ 2782 in situ; B, male ITBCZ 2798; C, calling male ITBCZ 2786 in situ (shows comparatively large single subgular vocal sac inflated); D, juvenile IBTCZ 4472; E and F, dorsolateraland ventral views of female ITBCZ 4471. Note:the semicircle seen in the eye of frog in D is from a ring flash and not a natural coloration.Photos by Luan T.Nguyen.

FIGURE 4. Advertisement call of Leptolalax rowleyae sp. nov. A, B, a 15 s waveform of relative amplitude (Rel. amp.) of holotype ITBCZ 2783 (recorded at ambient air temperature of 20.8 o C) and of an un-vouchered individual (recorded at ambient air temperature 21°C); power spectrum (relative amplitude vs. frequency) of a call (C); expanded 0.4 s waveform of relative amplitude (D) and corresponding spectrogram (E) of a call of holotype ITBCZ 2783; expanded 0.4 s waveform of relative amplitude (F) and corresponding spectrogram (G)of a call of an un-vouchered individual (same as inB).

FIGURE 5. Tadpole of Leptolalax rowleyae sp. nov. (ITBCZ 2790, stage 38). A–C, lateral, dorsal and ventral views, in life (photos by Luan T. Nguyen); D, schematic drawing of the oral disc in preservative; E, keratodonts of first posterior tooth row (P1) (drawn and photo by Dung T.Le).

FIGURE 6. Typical breeding habitat of Leptolalax rowleyae sp. nov. in the type locality: Son Tra Nature Reserve, Son Tra Peninsula, Da Nang City, Vietnam.Frogs were recorded along the cascade stream. Photo by Luan T.Nguyen.