Leucostethus jota, Marin & Molina-Zuluaga & Restrepo & Cano & Daza, 2018

Marin, Carlos M., Molina-Zuluaga, Claudia, Restrepo, Adriana, Cano, Estefany & Daza, Juan M., 2018, A new species of Leucostethus (Anura: Dendrobatidae) from the eastern versant of the Central Cordillera of Colombia and the phylogenetic status of Colostethus fraterdanieli, Zootaxa 4461 (3), pp. 359-380 : 364-371

Leucostethus jota , sp. nov.

( FIgS. 2 , 3 , 5 )

Holotype: MHUA-A 8138 (fIEld NumbER JMD 2063), AN AdulT mAlE fROm COlOmbIA, DEPARTmENT Of ANTIOquIA, MuNIcIPAlITy Of AlEjANdRíA, VEREdA El RESPAldO, JAguAS hydROElEcTRIc PROTEcTEd AREAS –ISAGEN S.A., LA ARENOSA fOREST (6.39301, -75.036402; 1290 m.A.S.l), cOllEcTEd ON DEcEmbER 3, 2013 by JMD.

Paratypes: All fROm COlOmbIA, DEPARTmENT Of ANTIOquIA. FROm MuNIcIPAlITy Of AlEjANdRíA, VEREdA El RESPAldO, JAguAS hydROElEcTRIc PROjEcT PROTEcTEd AREA –ISAGEN S.A : MHUA-A 8137, AN AdulT fEmAlE cOllEcTEd ON 2013 by JMD (6.4012, -75.02605; 1305 m.A.S.l); MHUA-A 7411 (AN AdulT mAlE); MHUA-A 7396 (A Sub-AdulT mAlE) ANd MHUA-A 7419 (A Sub-AdulT mAlE) cOllEcTEd ON 2012 by CMZ (6.38871, -75.02335; 1272 m. A.S. l). FROm MuNIcIPAlITy Of AlEjANdRíA, VEREdA El CERRO, JAguAS hydROElEcTRIc PROTEcTEd AREAS –ISAGEN S.A : MHUA-A 8140 , AN AdulT fEmAlE cOllEcTEd ON 2013 by JMD (6.37183, -75.04276; 1295 m. A.S. l) ANd MHUA-A 7416 , AN AdulT fEmAlE cOllEcTEd ON 2012 by CMZ (6.3693, -75.0252; 1286 m. A.S. l). FROm MuNIcIPAlITy Of SAN CARlOS, VEREdA El VERgEl : MHUA-A 8304 , A Sub-AdulT mAlE cOllEcTEd ON 2013 by DIEgO A. AlzATE (6.136672, -75.056296, 1416 m. A.S. l). FROm MuNIcIPAlITy Of SAN CARlOS, VEREdA DOS QuEbRAdAS, CAldERAS hydROElEcTRIc PROTEcTEd AREAS : MHUA-A 10659, AN AdulT mAlE cOllEcTEd ON 2016 by CMM (6.15728, -75.04274; 1158 m.A.S.l); MHUA-A 10668 , AN AdulT fEmAlE cOllEcTEd ON 2016 by CMM (6.15754, -75.04295, 1207 m. A.S. l) ANd MHUA-A 10664 , AN AdulT mAlE cOllEcTEd ON 2016 by CMM (6.15763, -75.04316, 1210 m. A.S. l). FROm MuNIcIPAlITy Of GRANAdA, VEREdA LA MERcEd, CAldERAS hydROElEcTRIc PROjEcT PROTEcTEd AREA –ISAGEN S.A : MHUA-A 10665, AN AdulT mAlE cOllEcTEd ON 2016 by CMM (6.13971, -75.06638; 1353 m.A.S.l).

Etymology: ThE SPEcIfIc NAmE IS A NOuN IN APPOSITION mEANT AS PATRONym fOR JOhN JAIRO ESTRAdA LOzANO, kNOWN AS jota ThAT dENOTES ThE lETTER “J” IN SPANISh. JOhN JAIRO IS A COlOmbIAN hERPETOlOgIST WhO cONTRIbuTEd ENORmOuSly TO ThE fORmATION Of ThE MuSEO dE HERPETOlOgíA UNIVERSIdAd dE ANTIOquIA (MHUA). HE SET uP ThE RulES ANd PROTOcOlS fOR ThE muSEum ANd INSPIREd ThE fIRST hERPETOlOgISTS WhO bEgAN TO dEPOSIT ThEIR SPEcImENS AT ThE MHUA.

Diagnosis: WE ASSIgNEd ThE NEW SPEcIES TO Leucostethus ON ThE bASIS Of OuR PhylOgENETIc RESulTS ( FIguRE 1 , APPENdIX 2). Leucostethus jota sp. nov. IS chARAcTERIzEd by ThE fOllOWINg cOmbINATION Of chARAcTERS: (1) mEdIum SIzEd, SVL fOR AdulT mAlES 16.1–21.6 mm (mEAN = 18.9 ± 1.16; N = 51), SVL fOR AdulT fEmAlES 17.9–23.3 mm (mEAN = 20.5 ± 1.39; N = 21); (2) dORSAl SkIN TEXTuRE POSTERIORly gRANulAR; (3) fINgER I ANd fINgER II SubEquAl IN lENgTh; (4) APPRESSEd fINgER IV REAchINg dISTAl hAlf Of dISTAl SubARTIculAR TubERclE Of fINgER III; lONgER ThAN fINgER II; (5) fINgER III SWOllEN IN AdulT mAlES ( FIguRE 3B ); NOT SWOllEN IN fEmAlES ( FIguRE 3C ); (6) fINgER lAckINg lATERAl fRINgES; (7) ThROAT, ANd ANTERIOR PORTION Of bElly WITh A dARk SPOTTEd PATTERN IN mAlES ANd ImmAculATE IN fEmAlES; (8) TESTES uSuAlly uNPIgmENTEd, mATuRE OVIducTS cONVOluTEd ANd PIgmENTEd; (9) TOES lAckINg fRINgES ( FIguRE 3A ); (10) bASAl WEbbINg bETWEEN TOES II–IV; (11) PAlE dORSOlATERAl STRIPE AbSENT; (12) STRAIghT PAlE VENTROlATERAl STRIPE PRESENT; (13) PAlE OblIquE lATERAl STRIPE cOmPlETE (EXTENdINg fROm gROIN TO EyE); (14) PAlE PARAclOAcAl mARkS AbSENT; (15) mEdIAl lINguAl PROcESS AbSENT; (16) mAXIllARy TEETh PRESENT; (17) VOmERINE TEETh AbSENT; (18) VOcAl SlITS PRESENT; (19) AdVERTISEmENT cAll cOmPOSEd Of A SINglE NOTE WITh A PEAk fREquENcy Of 4.40 KHz ANd EmITTEd AT A RATE Of 22 cAllS/mINuTE.

Comparisons with other species: Leucostethus jota sp. nov. ShARES WITh L. fraterdanieli ThE SWEllINg Of fINgER III IN AdulT mAlES ANd ThE OccuRRENcE Of cONSPIcuOuS bRIghT flASh mARkS ON ThE AXIllA, gROIN, ANd cONcEAlEd SuRfAcES Of ThE ThIgh ANd ShANk buT dIffERS fROm L. fraterdanieli IN hAVINg PAlE VENTROlATERAl STRIPES (AbSENT IN L. fraterdanieli ); A SOlId dARk bANd RuNNINg AlONg ThE flANkS Of ThE bOdy ANd hEAd (dIScONTINuOuS IN L. fraterdanieli ); IN ThE cOlORATION Of ThE TESTES (ENTIREly PIgmENTEd IN L. fraterdanieli , FIguRE 4A , VS. uNPIgmENTEd WhITE IN L. jota sp. nov., FIguRE 4B ); IN hAVINg lOWER cAll RATE (22 cAllS/mIN IN L. jota sp. nov., 98 cAllS/mIN IN L. fraterdanieli ), A WIdER fREquENcy bANd (RANgINg fROm 3.7 TO 4.4 KHz IN L. jota sp. nov., ANd 3.1 TO 3.3 IN L. fraterdanieli ) ANd IN hAVINg SmAllER bOdy SIzE: SVL fOR AdulT mAlES IN L. jota sp. nov. 16.1–21.6 mm (mEAN = 18.9 ± 1.16; N = 51), SVL fOR AdulT fEmAlES 17.9–23.3 mm (mEAN = 20.5 ± 1.39; N = 21); SVL fOR AdulT mAlES IN L. fraterdanieli 20.8–24.6 mm (mEAN= 23.1 ± 1.34; N= 8), SVL fOR AdulT fEmAlES 22.8–23.4 mm (mEAN = 23.1 ± 0.42; N= 2). ThE dISTINcTIVENESS Of L. jota sp. nov. ANd L. fraterdanieli IS fuRThER cORRObORATEd by uNcORREcTEd PAIRWISE dISTANcES Of 5.5% IN 16S ANd 17.8% IN CyT-b. Leucostethus jota sp. nov. dIffERS fROm C. furviventris , C. imbricolus , C. inguinalis , C. latinasus , C. panamansis , ANd C. pratti IN hAVINg A cOmPlETE PAlE OblIquE lATERAl STRIPE (EXTENdINg fROm gROIN mIdWAy TO EyE IN ThOSE SPEcIES ANd EXTENdINg fROm gROIN TO EyE IN Colostethus agilis , C. dysprosium , C. lynchi , C. mertensi , C. yaguara , Leucostethus argyrogaster , L. brachistriatus , L. fraterdanieli ANd L. fugax ). Leucostethus jota sp. nov. fuRThER dIffERS fROm C. furviventris , C. pratti , ANd C. ucumari IN lAckINg A PAlE dORSOlATERAl STRIPE (PAlE dORSOlATERAl STRIPE AbSENT IN C. agilis , C. alacris , C. dysprosium , C. imbricolus , C. inguinalis , C. latinasus , C. lynchi , C. mertensi , C. panamansis , C. thorntoni , C. yaguara ANd Leucostethus argyrogaster , L. brachistriatus , L. fraterdanieli , L. fugax ANd C. ramirezi ).

AmONg SPEcIES Of dENdRObATIdS dISTRIbuTEd AlONg ThE NORThERN CENTRAl CORdIllERA Of COlOmbIA, Leucostethus jota sp. nov. mOST clOSEly RESEmblES Hyloxalus ramosi ANd Hyloxalus lehmanni . ThESE SPEcIES ARE SImIlAR IN bOdy SIzE ANd cOlORATION PATTERN TO ThE NEW TAXON. Leucostethus jota sp. nov. ShARES WITh H. ramosi ThE PRESENcE Of bOTh PAlE VENTROlATERAl ANd OblIquE lATERAl STRIPES ANd ThE AbSENcE Of A PAlE dORSOlATERAl STRIPE (VENTROlATERAl STRIPE AbSENT IN H. lehmanni MHUA-A 00267). HOWEVER, L. jota sp. nov. dIffERS fROm H. ramosi ANd H. lehmanni IN ThE lENgTh Of fINgERS I ANd II (Sub-EquAl IN L. jota sp. nov., EquAl IN H. ramosi ANd H. lehmanni ), ThE PRESENcE Of TOE WEbbINg (bASAl WEbbINg bETWEEN TOES II-IV IN L. jota sp. nov., WEbbINg AbSENT IN H. ramosi ANd H. lehmanni ), ThIRd fINgER SWOllEN IN mAlES (PRESENT IN L. jota sp. nov., AbSENT IN H. ramosi ANd H. lehmanni ), ThE AbSENcE Of A blAck ARm glANd IN AdulT mAlES (PRESENT IN H. ramosi ANd H. lehmanni ), ThE gRAy SPOTTEd PATTERN ON ThROATS Of mAlES (dARk SOlId IN H. ramosi , fORmEd by A SERIES Of mINuTE blAck dOTS dISTRIbuTEd AlONg ENTIRE VENTRAl SuRfAcE NOT cONTRASTINg WITh ThE VENTER cOlORATION IN H. lehmanni : MHUA-A 00267) ANd by ThE lOWER cAll RATE (22 cAllS/mIN IN L. jota sp. nov., 86 cAllS/mIN IN H. ramosi ). FOR A SummARy Of dIAgNOSTIc chARAcTERS IN ThE SPEcIES Of Colostethus ANd Leucostethus SEE TAblE 4.

Measurements of the holotype: SVL 19.3 mm; fOREARm lENgTh fROm PROXImAl EdgE Of PAlmAR TubERclE TO OuTER EdgE Of flEXEd ElbOW 4.8 mm; hANd lENgTh fROm PROXImAl EdgE Of PAlmAR TubERclE TO TIP Of fINgER III 5.2 mm; ShANk lENgTh fROm OuTER EdgES Of flEXEd kNEE TO hEEl 9.7 mm; fOOT lENgTh fROm PROXImAl EdgE Of OuTER mETATARSAl TubERclE TO TIP Of TOE IV 8.8 mm; hEAd WIdTh bETWEEN ANglE Of jAWS 6.7 mm; hEAd lENgTh dIAgONAlly fROm cORNER Of mOuTh TO TIP Of SNOuT 6.4 mm; EyE lENgTh fROm POSTERIOR TO ANTERIOR cORNER 2.8 mm; EyE TO NARIS dISTANcE fROm ANTERIOR cORNER Of EyE TO cENTER Of NARIS 2.5 mm; dISTANcE bETWEEN cENTERS Of NARES 2.9 mm; SNOuT lENgTh fROm ANTERIOR cORNER Of EyE TO TIP Of SNOuT 3.5 mm; INTERORbITAl dISTANcE 2.5 mm; dIAmETER Of TymPANum 1.6 mm.

Description of the holotype: A mEdIum SIzEd fROg, 19.3 mm SVL. VENTRAl ANd lATERAl SuRfAcES SmOOTh. DORSAl SuRfAcES WITh gRANulES ScATTEREd IRREgulARly IN SAcRAl REgION. HEAd bAREly WIdER ThAN lENgTh. SNOuT TRuNcATEd IN dORSAl VIEW; PROjEcTEd AbOVE ThE lEVEl Of ThE uPPER lIP IN lATERAl VIEW; ShORT (SNOuT EyE dISTANcE 13% Of ThE SVL). INTERORbITAl dISTANcE 37% Of hEAd WIdTh. CANThuS ROSTRAlIS gENTly ROuNdEd. LOREAl REgION flAT OR WEAkly cONcAVE, NOT SlOPINg TO lIPS. EyE dIAmETER 43% Of hEAd lENgTh. NARES SlIghTly PROTubERANT, dIREcTEd POSTEROdORSAlly. TymPANum WEll dEfINEd IN PRESERVEd SPEcImENS; cONcEAlEd POSTEROdORSAlly. DIAmETER Of TymPANum 59% Of EyE dIAmETER; POSTRIcTAl TubERclES AbSENT. MAXIllARy TEETh PRESENT. VOcAl SlITS PRESENT. HANd lENgTh 27% Of SVL ANd 1.10 TImES fOREARm lENgTh. FINgER dIScS WEAkly TO mOdERATEly EXPANdEd. FINgER III SWOllEN AlONg PREAXIAl SIdE. FINgERS lAckINg fRINgES. METAcARPAl fOld AbSENTS. FINgER I lONgER ThAN fINgER II; fINgER II EXTENdEd TO mIdlEVEl Of dISTAl SubARTIculAR TubERclE Of fINgER III; fINgER IV REAchINg dISTAl hAlf Of dISTAl SubARTIculAR TubERclE Of fINgER III. RElATIVE fINgER lENgThS IV <II <I <III. SubARTIculAR TubERclES 1–1–2–2. All TubERclES STRONgly PROTubERANT; SubARTIculAR ANd ThENAR TubERclES EllIPTIcAl; PAlmAR TubERclE cIRculAR. ShANk lENgTh 50% Of SVL.

FOOT lENgTh 46% Of SVL. RElATIVE lENgThS Of APPRESSEd TOES I <II <V <III <IV ( FIguRE 3A ). TOE III REAchINg APPROXImATEly dISTAl PORTION Of PENulTImATE SubARTIculAR TubERclE Of TOE IV; TOE V NOT REAchINg dISTAl EdgE Of PENulTImATE SubARTIculAR TubERclE Of TOE IV. WEbbINg bASAl bETWEEN TOES II–IV (II 2–3 2/ 3 III 3– 4 IV). TOE fRINgES AbSENT. DIScS WEAkly TO mOdERATEly EXPANdEd. TubERclES STRONgly PROTubERANT. SubARTIculAR TubERclES 1–1–2–3–1. INNER mETATARSAl TubERclE ElONgATEd. OuTER mETATARSAl TubERclE Sub-cIRculAR, dIAmETER EquAl IN lENgTh TO INNER mETATARSAl TubERclE. MEdIAl mETATARSAl TubERclES AbSENT. OuTER mETATARSAl fOld WEAk. TARSAl kEEl TubERclE-lIkE, STRONgly cuRVEd AT PROXImAl ENd. ClOAcAl TubERclES AbSENT.

Coloration of the holotype in preservative: ( FIguRE 5A–C ) DORSum PAlE bROWN WITh dARk bROWN blOTchES ANd gRANulES dARk bROWN. PAlE dORSOlATERAl STRIPE AbSENT. FlANk dARk bROWN, bROkEN by A cREAm OblIquE lATERAl STRIPE EXTENdINg fROm gROIN TO EyE. PAlE VENTROlATERAl STRIPE cREAm. PAlE POSTOculAR STRIPE cROSSINg ThE ANTEROVENTRAl ONE-hAlf TO TWO-ThIRdS Of ThE WEll-dEfINEd TymPANum. LOREAl REgION ANd SNOuT dARk bROWN. UPPER lIP dARk bROWN WITh gRAy dOTS. ThROAT WITh SPOTTEd PATTERN EXTENdINg OVER ANTERIOR PORTION Of chEST ANd bElly. LOWER lIP EdgE WITh A dARk bROWN STRIPE. DORSAl SuRfAcE Of uPPER ARm cREAm, lAckINg TRANSVERSE bANdS; ANTERIOR ANd POSTERIOR SuRfAcES cREAm; VENTRAlly cREAm. DORSAl, ANTERIOR, ANd POSTERIOR SuRfAcES Of fOREARm cREAm lAckINg TRANSVERSE bARS. CONcEAlEd SuRfAcE Of fOREARm WhITE. PAlmAR SuRfAcES bROWN. DORSAl SuRfAcE Of hANd cREAm WITh PAlE bROWN blOTchES ON fINgERS. DORSAl SuRfAcE Of ThIgh gRAy. VENTRAl SuRfAcE Of ThIgh ANd cONcEAlEd SuRfAcES Of ShANk ANd fOOT cREAm. PAlE PARAclOAcAl mARk AbSENT. PlANTAR SuRfAcES bROWN.

Coloration of the holotype in life: ( FIguRE 2 ) DORSum bROWN WITh A blAckISh bROWN INTERORbITAl STRIPE. SIdES Of hEAd ANd bOdy dARk bROWN. PAlE OblIquE lATERAl STRIPE cREAm WITh mINuTE REddISh lIghT SPOTS. GROIN, AXIllA ANd POSTERIOR TIP Of lATERAl STRIPE yEllOW. DORSAl SuRfAcE Of fORElImbS ORANgE. CONcEAlEd SuRfAcES Of lImbS yEllOW- ORANgE. DARk bROWN lONgITudINAl STRIPE ON ANTERIOR ANd POSTERIOR SuRfAcES Of fORElImb ANd ANTERIOR ANd POSTERIOR SuRfAcES Of ThIgh. POSTERIOR SuRfAcE Of ThIgh INdISTINcTly mOTTlEd bROWN ANd yEllOW. VENTRAl SuRfAcE Of ThIgh PAlE gRAy. DORSAl SuRfAcE Of TIbIA WITh TRANSVERSE bROWN bANdS. VENTRAl SuRfAcES Of lImbS cREAm. ThROAT, ANTERIOR PORTION Of chEST ANd lOWER lIP mARgIN WITh A gRAy SPOTTEd PATTERN. IRIS blAck WITh A bRONzE RINg AROuNd PuPIl WITh cOOPER SPOTS.

Morphological variation: AdulT mAlES Of Leucostethus jota sp. nov., ARE SmAllER ThAN fEmAlES (SEE DIAgNOSIS SEcTION) ANd EXhIbIT VARIATION IN ThE SWEllINg Of ThIRd fINgER RANgINg fROm A WEAk PREAXIAl SWEllINg TO A STRONgER PREAXIAl SWEllINg (e.g., MHUAA-7924). AlThOugh uSuAlly uNPIgmENTEd, SOmE AdulT mAlES hAVE A WEAk EXTENT Of dARk PIgmENTATION SuRROuNdINg ThE SEmINIfEROuS TubulES (SEE FIguRE 2 IN TOlOSA et al. 2014).

Bioacoustics: ThE AdVERTISEmENT cAll Of Leucostethus jota sp. nov. cONSISTS Of A mId-PITchEd WhISTlE SINglE NOTE ( FIguRE 6C ). EAch NOTE hAS A duRATION Of AbOuT 0.14 SEcONdS (Sd ± 0.02; N = 70) EmITTEd AT A RATE Of 21.80 cAllS/ mINuTE (Sd ± 2.91; N = 70), WITh cAll INTERVAlS Of 2.55 SEcONdS (Sd ± 0.88; N = 70). ThE OScIllOgRAm ( FIguRE 6C ) ShOWS A RhOmbOId ShAPE WhERE ThE NOTE hAS ThE lOWEST AmPlITudE IN ThE bEgINNINg ANd INcREASES AT ThE ENd Of ThE cAll. ThE fREquENcy RANgES bETWEEN 3.70 KHz (Sd ± 0.09) ANd 4.90 KHz (Sd ± 0.09), WITh A PEAk fREquENcy Of 4.40 KHz (Sd ± 0.14). ThE AdVERTISEmENT cAll Of L. jota sp. nov. IS REAdIly dISTINguIShEd fROm ThE cAll Of Leucostethus fraterdanieli by hAVINg bOTh hIghER fREquENcy ANd lOWER cAll RATE ( FIgS. 6 C–D ). A lIST Of ThE NumbER Of cAllS PER SPEcImEN Of ThE cOllEcTEd ANd uNcOllEcTEd cAllINg mAlES Of L. fraterdanieli ANd L. jota sp. nov. ARE ShOWN IN TAblE 5. COmPARISON bETWEEN ThE AdVERTISEmENT cAll PARAmETERS Of L. jota sp. nov. ANd ITS clOSEST RElATIVE SPEcIES L. fraterdanieli IS ShOWN IN TAblE 6 ANd FIgS. 6A–D . ThE PCA bASEd ON SPEcTRAl PARAmETERS ANd cAll duRATION ShOWN L. jota sp. nov. bE hIghly dISTINcT fROm L. fraterdanieli IN ThE AcOuSTIc SPAcE ( FIguRE 7 ). ThE fIRST PC AXIS WAS mAINly cORRElATEd WITh cAll RATE ANd SPEcTRAl PARAmETERS (lOW fREquENcy, hIgh fREquENcy ANd PEAk fREquENcy) WhIlE ThE SEcONd PC AXIS WAS cORRElATEd WITh cAll duRATION ( TAblE 7).

Leucostethus jota sp. nov. IS A dIuRNAl SPEcIES WITh cAll AcTIVITy RANgINg fROm 06:00–18:00 h. ThE SPEcIES PRESENTS VARIATION IN dAIly TEmPORAl cAllINg PATTERN WITh TWO AVERAgE cAllINg INTENSITy PEAkS: IN ThE mORNINg AbOuT 09:00–10:00 h ANd IN ThE AfTERNOON AbOuT 14:00–15:00 h ( FIguRE 8 ). WhEN ObSERVEd VOcAlIzINg ( FIguRE 9A ), AIR TEmPERATuRE RANgEd bETWEEN 21.0°C TO 32.1°C ANd RElATIVE humIdITy RANgEd fROm 87.8% TO 92.3%. IN AgONISTIc bEhAVIOR, ThE ENcOuNTER cAll Of Leucostethus jota sp. nov. cONSISTS Of A SEquENcE Of ThREE TO fOuR NOTES, EAch ONE WITh A duRATION Of 0.09 SEcONdS (Sd ± 0.03; N = 7), WITh cAll INTERVAlS Of 0.2 SEcONdS (Sd ± 0.01; N= 7). ThE fREquENcy RANgES bETWEEN 3.5 KHz (Sd ± 0.1) ANd 4.7 KHz (Sd ± 0.2), WITh A PEAk fREquENcy Of 4.0 KHz (Sd ± 0.2).

Distribution and natural history: Leucostethus jota sp. nov. IS A lEAf-lITTER dWEllINg fROg ThAT lIVES NEAR SmAll STREAmS IN PREmONTANE WET fORESTS IN ThE muNIcIPAlITIES Of AlEjANdRíA, AmAlfI, GRANAdA, SAN CARlOS, SAN RAfAEl ANd SAN ROquE IN ThE DEPARTmENT Of ANTIOquIA, COlOmbIA ON ThE EASTERN VERSANT Of ThE NORThERN CENTRAl CORdIllERA bETWEEN 1158–1514 m.A.S.l ( FIguRE 10 ). IT IS cOmmONly fOuNd clOSE TO ThE ROOTS Of VEgETATION ANd ON ThE lEAf lITTER ThAT fORmS hOlES INTO WhIch INdIVIduAlS mOVE TO WhEN dISTuRbEd. ThIS fROg SEEmS TO hAVE AN AggREgATEd SPATIAl dISTRIbuTION WhERE INdIVIduAlS fORm mIXEd-SEX gROuPS Of 2 TO 8 INdIVIduAlS. NEVERThElESS, WE hAVE ObSERVEd AgONISTIc ENcOuNTERS bETWEEN mAlES, chANgINg ThE VOcAlIzATION (SEE BIOAcOuSTIcS) ANd SOmETImES fIghTINg WITh kIckS. Leucostethus jota sp. nov. IS AcTIVE ThROughOuT ThE yEAR ANd hAS cONTINuOuS REPROducTION (TAdPOlES ANd cAllINg mAlES fOuNd AROuNd ThE yEAR). FEmAlES lAy bETWEEN 10 ANd 18 TERRESTRIAl EggS IN ThE lEAf lITTER ThAT ARE cAREd fOR mAlES uNTIl hATchINg, ThEN, mAlES cARRy TAdPOlES ON ThEIR dORSum uNTIl ThEy ARE RElEASEd INTO ThE WATER ( FIguRE 9B ). ThE dIET Of ThIS fROg IS cOmPOSEd mAINly Of ANTS ANd OThER hymENOPTERANS, ANd cOlEOPTERANS (MuRIllO et al. uNPublIShEd). A ShORT-TERm dEmOgRAPhIc STudy (MOlINA-ZuluAgA et al. 2014) fOuNd bOTh hIgh SuRVIVAl PRObAbIlITIES ANd POPulATION gROWTh RATES IN ThE POPulATION Of L. jota sp. nov. fROm muNIcIPAlITy Of AlEjANdRíA, lOcAlITy El EdéN, SuggESTINg A STAblE dEmOgRAPhIc STATuS Of ThIS POPulATION fOR ThIS PERIOd. AT PRESENT, L. jota sp. nov. IS kNOWN TO OccuR IN ThE PROTEcTEd AREAS Of ThE hydROElEcTRIc PROjEcTS CAldERAS IN ThE muNIcIPAlITy Of SAN CARlOS ANd JAguAS IN ThE muNIcIPAlITy Of AlEjANdRíA WhIch fAVOR ITS cONSERVATION STATuS.

Figures

FIGURE 1. Maximum likelihood tree (-lnL=120317.99) depicting evolutionary relationships within Colostethinae inferred from a partitioned analysis using 11978 characters, 16 genomic regions, and 183 terminals. Colored symbols next to the clades are represented as point localities in figure 10.

FIGURE 2. Holotype of Leucostethus jota sp. nov. adult male MHUA-A 8138 in life. SVL 19.32 mm. Photo by Juan M. Daza.

FIGURE 3. Foot and hand morphology of Leucostethus jota sp. nov. A) Right foot of adult male holotype MHUA-A 8138. Foot length = 8.8 mm). B) Right hand of adult male holotype showing swelling along the preaxial side of finger III. Hand length = 5.2 mm. C) Right hand of adult female paratype MHUA-A 8137. Hand length = 5.1 mm. Finger III not swollen.

FIGURE 4. Testes color in (A) Leucostethus fraterdanieli (MHUA-A 8176) and (B) Leucostethus jota sp. nov. (MHUA-A 9534). Arrows indicate testes position.

FIGURE 5. Holotype of Leucostethus jota sp. nov. in preservative (MHUA-A 8138). SVL 19.3 mm. (A) Lateral view. (B) Ventral view. (C) Dorsal view.

FIGURE 6. Spectrogram and oscillogram of the advertisement call of two species: A) Single pulse and B) multiple pulses of Leucostethus fraterdanieli (MHUA-A 8176). C) Single pulse and D) multiple pulses of Leucostethus jota sp. nov. (MHUA-A 9533). Note the higher frequency and lower call rate in L. jota sp. nov.

FIGURE 7. PCA plot of bioacoustic parameters from L. jota sp. nov. and L. fraterdanieli. PCA was performed with 3 spectral parameters (Low frequency, High frequency and Peak frequency) and 1 temporal parameter (Call duration).

FIGURE 8. Acoustic activity in a population of Leucostethus jota sp. nov. inferred from 27788 calls, obtained during November-December 2012 and January through September 2014 in the municipality of Alejandría, Antioquia.

FIGURE 9. Leucostethus jota sp. nov. A) Vocalizing male; uncollected specimen B) Adult male carrying tadpoles; uncollected specimens. Photos by Juan M. Daza.

FIGURE 10. Geographic distribution of sequenced specimens of Leucostethus jota sp. nov. and Leucostethus fraterdanieli “complex” in Colombia. Colored symbols represent the clades shown in figure 1.