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^^CZ Hern ^^''^' LIBRARY 668 NOV 5 2001 Papers ^Scientific u^w^^e^^ . Museum Natural History The University of Kansas crt iL> QJ 21 September 2001 Number 22:1-10 •o=- —C 1c^a A New Species of Marsupial Frog a/a?>).<3N^^cC«fl (Anura: Hylidae: Gastrotheca) z B ° from the Cordillera Azul in Peru 2U g 3 " C By William E. Duellman^ Edgar Lehr^ and Cesar Aguilar^ I ^Natural HistoryMuseum and BiodiversityResearch Center, and ; I DepartmentofEcologyand Evolutionary Biology. \ The UniversityofKansas, Lawrence, Kansas 66045-2454, USA. u d -StaatlicheNaturhistorischeSarnmlungen DresdenMuseumfiirTierkunde, i- %x A.-B.-Meyer,-Bau KoenigsbrueckerLandstrasse 159, D-01109 Dresden, Germany X 2 ^DepartamentodeHerpetologia, MuseodeHistoria Naturalde San Marcos, Apartado 14-0434, Lima 14, Peru Z 3 = s S-^ CONTENTS ABSTRACT 1 RESUMEN 2 IINTRODUCTION 2 Acknowledgments 2 MATERIALSAND METHODS 2 DESCRIPTIONOF NEWSPECIES 2 DISCUSSION 9 LITERATURE CITED 9 APPENDIX; SPECIMENS EXAMINED 9 ABSTRACT AnewspeciesofGastrothecafromthesouthernendoftheCordilleraAzulinPeruseems to be most closely related to G. lateonota known only from the Cordillera Huancabamba in northern Peru. Both of these species and G. marsupiata, monticola and peruana produce tadpoles, whereas the otherspecies ofGastrotheca in PeruvianAndeshave directdevelopment. KeyWords: Anura; Hylidae; Gastrotheca;new species;Peru;Andes. )NaturalHistoryMuseum,TheUniversityofKansas ISSNNo.1094-0782 Scientific Papers, Natural History Museum, The University of Kansas RESUMEN Una nueva especies de Gnstwtlieca del extreme surde la CordilleraAzul aparentemente esta realcionada a G. lateonota quesolo esta registrada para laCordillera de Huancabamba alnorte de Peru.AmbasespeciesjuntoconG. niarsiipiata, inonticola, yperuanaproducenrenacuajos,mientras que las otras especies de losAndes peruanos tienen desarrollo directo. PalabrasClaves: Anura, Hylidae, Gastwtliecn; especienueva; Peru; losAndes. INTRODUCTION Marsupial frogs ofthe genus Gnstwtlieca are highlydi- Theothersevenspecieshaverestricteddistributionsin verse in theAndes, especially in Colombia, Ecuador, and theAndes. Gastrotheca abdita is known only from the iso- Peru, where 31 species have been recorded; 14 of these lated Cordillera Colan, and G. galeata and G. lateonota are species have been reported from Peru (Duellman, 1987; knownonlyfromtheCordilleradeHuancabambainnorth- Duellman and Fritts, 1972; Duellman and Trueb, 1988; ern Peru. Four other species (G. excubitor, oclioai, Duellman and Wild, 1993;Trueb andDuellman, 1978). Of pacchamama, and rebeccae)have restricted allopatricdistri- these species, G. longipes occurs in the upper Amazon butionsathighelevationsintheCordilleraOrientalincen- Basin, and G. testudinea and G. weinlandii have exten- traland southern Peru. sive distributions in cloud forests on the easternslopes Field work in 1998 by Lehr and Aguilar in the Cordil- of the Andes from Colombia to Bolivia. The other spe- leraAzul,apartoftheCordilleraOriental incentralPeru, cies in Peru are restricted to the Andes, mostly at el- resultedinthediscoveryofseveralnewspeciesofanurans evations above 2000 m. (Lehr,2001; Lehretal.2001,2002).Amongtheseisaprevi- Ofthese high Andean species, four have rather exten- ously unknown species ofGastrotheca. sivedistributions. Gastrotheca mnrsupiatn rangesfromcen- Acknowledgments tralPeruintoBoliviaandG.griswoldioccursincentralPeru; the range of G. peruana extends from central Peru north- ThefieldworkwassupportedbytheForschungsinstitut ward in the Cordillera Occidental, whereas that of G. und Naturmuseum Senckenberg. Lehr is grateful to per- monticolaincludesthenorthernpartsoftheCordilleraOc- somiel at the Instituto Nacional de Recursos Naturales of cidental and Cordillera Central, the Cordillera de theMinisteriodeAgriculturainLima,Peru,forproviding Huancabamba, the Huancabamba Depression in Peru collectingandexportpermits.JavierIcocheaandGunther and Ecuador, and the southern part of the Cordillera Kohler made helpful suggestions. The manuscript ben- Occidental in Ecuador. The nominal species G. lojana efitedfromcriticalreviewsbyJosephR.MendelsonIIIand was placed in the synonymy of G. monticola by ErikR.Wild,and fromcarefuleditingbyLindaTrueb.The Duellman and HilHs (1987). accuraterenderingsofthefrogsarefromthetalentedhand ofChristopherA. Shell (Kansas). MATERIALSAND METHODS The 16 morphological measurements and 25 external Historia Natural Universidad San Marcos, Lima, Peru descriptive characters are those used by Duellman and (MHNSM). The distribution map is based on the Mapa Pyles (1980), Duellman and Hillis (1987), and Duellman FisicoPolitico(InstitutoGeograficoMilitardelPeru,1973). andTrueb(1988).Allmeasurementsareinmm;snout-vent StatisticalanalyseswereperformedwithStatView4.5(Aba- length is abbreviated SVL. Museum codes are those of cusConcepts,Inc., 1992-1995). Numberedcolorsrefertocor- Leviton et al. (1985) with the addition of the Museo de respondingcolors in the colorguides ofSmithe (1975, 1981). DESCRIPTION OF NEW SPECIES Gastrotheca stictopleura new species — A MHNSM Diagnosis. moderate-sizedspecies(femalesto68.3 Holotype.— 20319, an adult female, from mm)having (1) tibialength49.9%SVL,noticeablyshorter TrancaGrandeatChagUa (09°51'08"S,75°54'37"W,eleva- thanfoot;(2)mterorbitaldistancenoticeablygreater(160%) tion 3090 m), Provincia de Pachitea, Departamento de than width of upper eyelid; (3) skin on dorsum finely Huanuco,Pena,obtainedon23August1998byEdgarLehr. shagreen, not co-ossified with skull, lacking transverse — Referred specimens. SMF 80328-29, juveniles, from ridges; (4) supraciliary processes absent; (5) heel lacking Cochacalla at Chaglla, Provincia de Pachitea, calcar or pronounced tubercles; (6) tympanic annulus Departamento de Huanuco, Peru, obtained on23Avigust smooth; (7) Finger I shorter than FingerII; width ofdisks 1998by Edgar Lehr. greater than thatofdigits; (8) fingers unwebbed; (9) web- New Species ofMarsupialFrog Table 1. MeasurementsofadultfemalesofthreespeciesofGastrotlwca. Meansand 1 SDgivenbelowranges Scientific Papers, Natural History Museum, The University of Kansas Fig.2.VentralcolorationinadultfemalesofthreespeciesofGastrotheca. A. G. lateonota,KU181736,63.3mmSVL. B.G. montkola,KU181743, .1 mmSVL. C. G.stictoplewa,MHNSM20319,68.3mmSVL. Thesearesmallerspecies(SVL<57mm) thathavesmaller turing of the underlying bone, and the skin is moveable discsonthefingersandlesswebbing,whichextendsonly onthetop ofthe head. — to the penultimate subarticular tubercle on Toe V. Description of holotype. Brooding female having a Gastrothecaperuana differs furtherbyhavingFingersIand SVLof68.3 mm; head slightly wider than long, narrower II equal inlength, a granular tympanic annulus, and pus- thanbody;headlength29.7%SVL;headwidth33.8%SVL; tular skin on the dorsum. Furthermore, these three spe- snout-moderately long, acutely rounded in dorsal view, cies differ from G. stictopleura in dorsal coloration. inclined anteroventrallyinprofile; nostrilsslightlyprotu- Gastrotheca griswoldi and G. niarsupiata have a dark inter- berant, directed anteroventrally, situated at a point 54.2% orbitalbar; in G. grisiuoldi, thebar usually is connected to distancefromanteriorleveloforbittotipofsnout,atlevel diagonal or paravertebral marks on the body, whereas in well behind margin of lower jaw; eye-nostril distance G. marsupiata the bar is separate from the dark flecks or 86.2% length of eye. Canthus rostralis straight, elevated, spotsonthebody. Gastrothecaperuanalacksadarkinteror- rounded in profile; loreal region noticeably concave; lips bitalbarandhaselongateparavertebralmarksonthebody. rounded; internarial region slightly concave; top ofhead Other species of Gastrotheca inhabiting the Cordillera depressed; interorbital distance 34.6% head width, 160% Oriental in Peru (G. excubitor, ochoai, paccharnama, and width ofuppereyelid. Tympanum vertically ovoid, sepa- rebeccae) are smaller than G. stictopleura (maximumSVL < ratedfromeyebydistance1.4xlengthoftympanum;tym- 46mm) and havedirectdevelopment. Two speciesinhab- panic araiulus distinct anteriorly and ventrally, smooth; iting cloud forests on the Andean slopes ofthe cordillera supratympanicfoldmoderatelyweak,barelyoverlapping (G. testudmea and G. weinlandii) differ from G. stictopleura upper edge of tympanum, extending from posterior cor- in several characters. Both are much larger, with females ner oforbit topoint above insertion offorearm. attainingaSVLinexcessof80mm,andhaveFingerIlonger Armmoderatelyrobust;handmoderatelylarge;fingers than Finger II, discs on digits much wider than digit at unwebbed; distinct lateral fringes on Fingers II-IV; discs base of the disc, granular tympanic annulus, and a tan moderately large with median longitudinal groove inan- dorsumwithtransversebarkbrownbars(G. testudinea) or terior part of each pad; width of disc on third finger no- chevrons(G. lueinlandii).Thelatterhastheskinonthehead ticeablygreaterthanlengthoftympanum; relativelength co-ossified with the skull and has large calcars, whereas of fingers 1 < 2 < 4 < 3; subarticular tubercles large, theskinisnotco-ossifiediiiG. testudinea,whichhasasmall rounded,nonebifid;supernumerarytuberclesabsent;pal- tubercle on theheel. On cursoryexamination, the skin on martuberclelow,elliptical;prepoUicaltubercleelongately the dorsal surface of the skull appears to be co-ossified elliptical (Fig. 3A). Hind limb moderately robust; tibia with the underlying cranial elements in some preserved length 49.9% SVL; foot length 44.7% SVL; heels of specimens of G. lateonota, uionticola, stictopleura, and adpressed limbs overlapping by about one-fourth length testudhiea;however,thetextureoftheskinreflectstosculp- ofshank;calcar,heeltubercle,andoutertarsalfoldabsent; New Species ofMarsupial Frog stripeonmarginofupperlip,followedposteriorlybytwo white spots above insertion of forelimb. Axilla and me- dianandposterodorsalsurfacesofflanksbluishgraywith small white spots; groin pale grayish tan without markings.Narrow,transversewhitestripeabovevent;crestsof vertical folds and tubercles lateral to vent white; narrow whitestripeonventrolateraledgeofforearm;narrowwhite bar on heel. Dorsal surfaces of fingers and Toes I-lII tan; othertoesandalldiscsbluishgray.Tliroatpalebluishgray with creamy white granules; other ventral surfaces and anterior and posterior surfaces ofthighs dull creamy tan; somegranulesonthebellywithasmallamountofbrown pigmentation (Fig. 3C). Color in life: From color photographs and field notes by Lehr. Dorsalsurfacesofhead,body,and limbs, and loreal and tympanic regions bright dark green (162B) with diffusesalmon(6) onbodyandlimbs;labialand dorsolateral stripes cream (54); dorsolateral stripe bordered ventrally by brown (38) stripe; spots on flanks, above insertion of forearm, and lateral to vent white. Throat, chest, andventralsurfacesofarmscream(54)toyellow(56)with diffusegreen(162B)andsalmon(6)spots;bellypalebrown (223D); ventral surfaces of thighs dark brown (119A); Fig.3. Hand(A)andfoot(B)ofholotypeofCastrothecastictopleum, shanks green (162B); hands and feet dark gray (79). Iris MHNSM20319. metallicorange (17)withfineblackreticulations (Fig. 4E). Measurements ofholotype: SeeTable 1. inner tarsal fold distinct, extending full length of tarsus; Cranialosteology: Byliftingtheskincoveringtheright outermetatarsaltubercleabsent;innermetatarsaltubercle posterolateral part of the skull, it was possible to deter- ovoid,barelyvisiblefrom above. Toeslong, slender,bear- mine the association ofthe frontoparietaland squamosal. ing distinct lateral fringes; relative length oftoes a < 2 < 3 The squamosal lacks a medial flange and is notin contact <5 <4;toeslessthanone-halfwebbed;webbingformulaI withtheoticflangeofthefrontoparietal. Thiscondition is 2—2II1—3III2—3IV3—2-V;discsslightlysmallerthan like that described for Gnstrotliecn lateonota by Duellman thoseonfingers;subarticulartuberclesprominent,round; and Trueb (1988) and differs from that in G. montkola, Ln supernumerarytuberclesprominent,presentonproximal which a median flange of the squamosal is inbroad con- segments ofall toes (Fig. 3B). tactwith the otic flange ofthe frontoparietal (Fig. 5). — Skin on dorsum shagreen; skin on ventral surfaces of Variation. Two juveniles have SVLs of 28.5 and 32.7 forearms and shanks smooth; skin on other ventral sur- mm. The coloration of the larger of these (SMF 80328) is facesgranular;verticalcloacalfoldspresent;transverserow like that of the holotype, except that there are faint tan — offourroundedtuberclesonposteriorsurfaceofeachthigh markings elongate, X-shaped mark on the dorsum be- lateral to cloaca, decreasing in size distally; opening of ginningontheuppereyelidsandterminatingpostsacrally brood pouch V-shaped, with the apex just posterior to withtheintersectioninthescapularregion;a short,longi- sacrum. Vomerine odontophores slightly oblique hidinal, middorsal mark posterior to the sacrum; irregu- posteromedially,narrowlyseparatedmedially,atposterior lar marks on dorsal surfaces oflimbs. The posterior part margins ofchoanae,bearing 5 and 6 teeth; choanae mod- oftheflanksiswhitewiththreeverticalbrownmarks.The erately large, nearly round; tongue bluntly ovoid, shal- dorsal coloration of the smaller specimen (SMF 80329) is lowly notched posteriorly, free posteriorly for about one likethatoftheholotype,buttheventerhasmoregranules third ofits length. withbrownpigmentation; theflanksposteriortotheaxil- Colorinpreservative: Dorsumofhead,body,andlimbs lary region are white withbrown spots. bluishgraywithnarrowcreamstripeoriginatingasverti- Field notes on coloration in life of these twojuveniles cal line on snout, passing along canthus rostralis, margin are: SMF80328: Dorsum green (159) with metallicsalmon ofupper eyelid, and supratympanic fold, terminating on (6) spots onhead, body, and extremities. SMF80329: Dor- upper flank at about midlength of body; narrow cream sum green (6), hands and feet cream (54); cloacal region Scientific Papers, Natural History Museum, The University of Kansas A. Gastmtliccamoiiticohi,female,KU212022,51.5mmSVL. B. GastwtJiccnttioiiticoh New Species of Marsupial Frog 200 " 180 160 140 CO 120 <0 100 E 80 - Fig.5. SkullsofGastrotheca monlicola (left)and G. lateonota(right) showingrelationshipoffrontoparietalandsquamosal. Theconditionin G.stktoplcuraislikethatinG. lateonota.Scalebar=5mm.Adaptedfrom 60 DuellmanandTrueb(1988). 40 - darkbrown (119B)borderedbywhite;canthalanddorso- lateral stripes cream (54) bordered below by darkbrown (223B)stripe;lipscream(54);flankswhitewithdarkbrown 20 (119) spots; throat pale brown (92); chest and belly dark brown (119B)withpalebrown (92) spots;ventralsurfaces of extremities dark brown (119B) with horn-colored (92) andsalmon(106) spots; iris salmon (132D) (Fig. 4F). — Life history and development. The holotype is a brooding female. Opening thebrood pouch revealed ova mm about 5.3 in diameter. Eggs were not removed from the pouch, but an estimated 82 eggs are present. At least some of these eggs appear tobe in developmental Stages 17-20 (Gosner, 1960);thesestagesareequivalenttoStages 16-18 described for Gastrotheca riobambaeby del Pino and Escobar(1981). Thenumberand sizeofeggsindicate that theeggsofGastrothecastictopleurahatchintotadpolesthat complete theirdevelopmentinponds (Fig. 6). Gastrotheca lateonota and G. monticola also produce eggs thathatch as tadpoles.ElevenbroodingfemalesofG.lateonotacontained 66-152(x= 103.1 ±27.6)eggsindevelopmentalStages31- 35 (Gosner, 1960) having diameters of 5.3-7.3 (x = 6.4 ± 0.66)mm.FourbroodingfemalesofG. moiiticolacontained 66-186 (x= 129.5 ±57.3) eggshaving diameters of3.6^.9 (x = 4.4 ± 0.57) rmn. The number of eggs brooded by females of species having direct development is much smaller (Fig. 6). — Distribution and ecology. Gastrotheca stictopleura is known only from the immediate vicinity of Chaglla, m Departamento Huanuco, Peru (Fig. 7). This site is the southernend ofthe CordilleraAzul; this name is applied to that part of the Cordillera Oriental lying to the east of Scientific Papers, Natural History Museum, The University of Kansas G. lateonota G. monticola G. stictopleura Zl Area above 3000 m. 100 200 Kilometers Fig.7. DistributionsofthreespeciesofGastvotliecainsoutliernEcuadorandnortliernPeru. — Etymology. ThespecificnameisaLatinizedfeminine orspotted, and theGreekpleura, meaning side. Thename adjectivederivedfromtheGreekstiktos,meaningdappled refers to the spotted pattern on the flanks. New Species ofMarsupial Frog DISCUSSION In size, external morphology, nature of the frontopari- The co-occurrence of Gastrotheca griswoldi, in which etalsandsquamosals,andmodeoflifehistory,Gnstrothccn eggs hatch as froglets in the brood pouch, and G. stictoplewa is most like G. lateonota, known only from the stictoplenra, inwhicheggshatchastadpolesthatcomplete CordilleradeHuancabambainnorthernPeru(Fig.6).With their development in ponds, reflects a consistent pattern the exception of the nature of the frontoparietals and inGastrotheca in thehighAndes.Allcases ofsympatryin- squamosals, these two species are more like G. moiiticola clude one species that produces tadpoles and another in thananyothermembersofthegenus.Gastrothecamonticola whichthe eggs undergo directed development. Thus, the rangesfromsouthernEcuador throughoutnorthern Peru direct-developing G.griszmldioccurssympatricallyindif- (Fig. 7). ferentpartsofitsrangewithG.peruana and G.stictoplenra, Like many other species of Gastrotheca in the high bothofwhichproducetadpoles.Likewise,thewidespread Andes, G. lateonota, monticola, and stictopleiira exhibitcon- G. marsnpiata, which produces tadpoles, occurs sympatri- siderablevariationindorsalcoloration. InG. lateonota,the cally with slightly smaller species that exhibit direct de- dorsumisgreenorbrownwithorwithoutdarkerparaver- velopment(G. excubitor,oclioai, andpacchamama) through- tebral marks, and in G. monticola, the dorsum is green or out their separate ranges in the Cordillera Oriental. On brown,usuallywithadarkermiddorsalblotchorparaver- the other hand, no two species that produce tadpoles tebral marks (Fig. 4A-D). The three known specimens of are known to occur sympatrically. The same situation G.,stictoplenraaregreendorsally,butonespecimen(SNM prevails in the highAndes ofBolivia, Ecuador, and Co- 80328) also has tan markings on the dorsum. lombia. LITERATURE CITED Del Pino, E. M., and B. Escobar. 1981. Embryonic stages ofGastrotheca INRENA(InstitutoNacionaldeRecursosNaturales).1995.MapaForestal riobambae (Fowler) during maternal incubation and comparison of delPeru. Lima:DireccionGeneralForestal. developmentwiththatofotheregg-broodinghylidfrogs.Journalof INRENA(InstituteNacionaldeRecursosNaturales).1996.GiiiaExplkativa Morphology167:277-295. delMapaForestal. Lima:DireccionGeneralForestal. Duellman, W. E. 1987. Two new species of marsupial frogs (Anura: Lehr,E.2001.AnewspeciesofP/in/uofn.^(Anura:Leptodactylidae)from Hylidae)fromPeru.Copeia 1987:903-909. theeasternAndeanslopesofcentralPeru.Salamandra37:11-20. Duellman,W.E.,andT.H.Fritts.1972.Ataxonomicreviewofthesouth- Lehr,E.G.Kohler,andE.Ponce.2001.AnewspeciesofPhn/nopiis(Anura: ernAndean marsupial frogs (Hylidae: Gastrotheca). Occasional Pa- Leptodactylidae)fromPeru. SenckenbergianaBiologie80:205-212. persMuseumNaturalHistoryUniversityofKansas9:1-37. Lehr. E.. C. Aguilar. and G. Kohler. 2002. Two sympatric new species of Duellman,W.E.,andD.M.Hillis.1987.Marsupialfrogs(Anura:Hylidae: Phrxnopus (Anura: Leptodactylidae) from a cloud forest in the Peruvian Gastrotheca)oftheEcuadorianAndes:resolutionoftaxonomicprob- Andes.Journal ofHerpetology(inpress). lemsandphylogeneticrelationships.Herpetologica43:141-173. Leviton, A. E., R. H. Gibbs, Jr., E. Heal, and C. E. Dawson. 1985. Stan- Duellman,W. E.,and R.A.Pyles. 1980.Anewmarsupialfrog(Hylidae: dardsinherpetologyandichthyology:Part1.Standardsymboliccodes Gastrotheca)fromtheAndesofEcuador.OccasionalPapersMuseum forinstitutionalresourcecollectionsinherpetologyandichthyology. NaturalHistoryUniversityofKansas84:1-13. Copeia 1985:802-832. Duellman,W.E.,andL.Trueb. 1988.Crypticspeciesofhylid marsupial Smithe,EB.1975. Naturalist'sColorGuide.Part1.ColorGuide.NewYork: frogsinPeru.JournalofHerpetology22:159-179. AmericanMuseumofNaturalHistory. Duellman,W.E.,andE.R.Wild.1993.AnuranamphibiansfromtheCor- Smithe,F. B. 1981. Naturalist'sColorGuide. Part11.ColorGuideSupple- dillera de Huancabamba, northern Peru: systematics, ecology, and ment.ColorSwatches.NewYork:AmericanMuseumofNaturalHis- biogeography.OccasionalPapersMuseumNaturalHistory Univer- tory. sityofKansas157:1-53. Trueb,L.,andW.E.Duellman.1978.Anextraordinarynewcasque-headed Gosner, K. L. 1960. Asimplified table for staging anuran embryos and marsupialfrog(Hylidae:Gastrotheca).Copeia 1978:498-503. larvaewithnotesonidentification.Herpetologica16:183-190. APPENDIX SpecimensExamined Gastrothecalateonota:PERU: Piura: ElTambo,31.5km ECanchaque, 138410 (skeleton), 148568. Zamora-Chinchipe: Zamora (?), BMNH 2180m,KU 181729,181730(holotype),181731-39, 181836-37(skeleton), 1933.6.24.45. MHNSM 1635. PERU: Amazonas: N slope Abra Barro Negro, 27 km WSW Gastrotheca monticola:ECUADOR:Azuay:Giron,2240-2500m, KU Leimebamba,3440m,KU212078;Chachapoyas,2340m,KU138238-41, 138401-03.Loja:Celica,2130m,BMNH1931.11.3.3-4;Loja,2150m,BMNH 215627 (tadpoles), MCZ 88897-901, MHNSM 6277 (tadpoles); 20.5 km 1931.2.12.10-13, 1933.6.3.18-44, 1935.11.3.26-32, 1947.2.31.6-12, WSWLeimebamba,3130m,KU181741;22KMWSWLeimebamba,3220 1931.2.31.13(holotypeofG./o/i7)M),1931.2.31.14-18,KU120673-74,USNM m, KU 212495 (tadpoles), MHNSM 6294 (tadpoles); 24 km WSW 258851-58; 2 km N Loja, 2100 m, KU 142846 (tadpoles); 5 km N Loja, Leimebamba,3370m,FSM30080;5kmNLevanto,2850m,KU212021;6 2150m,KU138235-36,138237(skeleton);2kmELoja,2200m,KU120675, kmNWMendoza,2200m,KU209421;Molinopampa,2400m,KU212022- USNM 258849-50; 6.8 km E Loja, 2640 m, KU 217511-12; 9 km E Loja, 31,212493(young),212494(tadpoles),MHNSM6116-21,6287(tadpoles), 2660m, KU 121387(tadpoles); 10km ELoja,2600m, KU142855 (tads), 6292(young); Pomacochas(Florida),2180m,KU 181742-70, 181838-39 k17m84W70-L7o6ja;,22k31m0SmL,ojKaU,C2A02S68983,89280;3354k7m(tWadpLoojlae,s)2;155.05mk,mKWU1Lo3j8a2,332;3350.,2 (losckaelliettyo,n)M,2H1N20S32M-362,04M;HBNelSlaMvis1t0a4,01(59)4,76.122.22-23.14.7-C4a8j,ama1r9c4a7:.2N.o25s.p7e7c-i7f8i;c KU142603-08,148549-51;7.9kmWLoja,2440m,KU203548(tadpoles); Cutervo,2620m,KU212055-66,NMW6483;8kmNWCutervo,2560m, 10 km W Loja, 2500 m, KU 138234; Saraguro, 2500 m, KU 138404-09, KU 212067, 212492 (lads); Querocotillo, MCZ 5328-30. Piura:Ayabaca, 10 Scientific Papers, Natural History Museum, The University of Kansas 2S7a0n0Img,naMciHoNtrSailM, 2760220(2m),;WKUsl1o9p6e8C19e;rrHouCahnicnagbuaemlbaao,n1H9u6a0nmc,abAaMmbNaH- mm,,KKUU210891786794-7(0t,adMpoHleNs)S.M15420-21;18.5kmWSWHuancabamba,2740 7551,KU219771,MCZ5290(holotype),5291-93,5296-97,5299-300,5302, Gastrothecasttctopleura:PERU:Huamico:ChagUa,MHNSM20319(ho- 5304-07, 5309, 5312-15, 5317, 5319, 5328-30; 1.8 km N Huancabamba, lotype),SMF80328-29. 1980m,KU219767-68,MHNSM15418-19;4kmNHuancabamba,1900