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rogs of the Genus Eleutherodactylus Western Ecuador in ;?v j^"--* *),' 'l»tv ^^.^^^v.^*' \ l^a. -T'»«*'?^S!tr>f>: and Sj^stematics, Ecology, Biogeography Dpd John D. Lynch and William E. Ernst Msyr Lfbrary HlMMtwTt r?f nnmrjarstn/e Zoology THEUl Museum of Comparative Zoology { MUSEUM HERPETOLOGY LIBRARY The Universi ling withVolume 1 in 1946, was disco nerly published in the above series wt Museum Occasional Papers until Number 180 in December 1996. The University ofKansas Natural History Museum Miscellaneous Publications began with Number 1 in 1946 and ended with Number 88 in February 1996. Monographs oftheThe University ofKansas Natural History Museum were initiated in 1970 and discontinued with Number 8 in December 1992. The foregoing publication series now are combined in a new series entitled The University of Kansas Natural History Museum Scientific Papers, begun in January 1997. 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Include expiration date. Cover design by LindaTrueb. Photograph ofEleutherodactylus labiosus by the late Kenneth Miyata. The University of Kansas Natural History Museum Special Publication No. 23 21 February 1997 Frogs of the Genus Eleutherodactylus (Leptodactylidae) in Western Ecuador: Systematics, Ecology, and Biogeography John D. Lynch Professor, SchoolofLife Sciences The UniversityofNebraska, Lincohi, Nebraska 68588 USA, Associate, Division ofHerpetology NaturalHistoryMuseum, The University ofKansas Lawrence, Kansas 66045, USA William E. Duellman Curator, Division ofHerpetology NaturalHistoryMuseum Professor, DepartmentofSystematicsandEcology The University ofKansas Lawrence, Kansas 66045, USA Natural History Museum The UniversityofKansas Lawrence, Kansas SPECIAL PUBLICATIONS Editor: Linda Trueb Managing Editor: Joseph T. Collins Special Publication No. 23 pp. i-iv, 1-236 Plates 1-8 Published 21 February 1997 ISBN 0-89338-054-7 © 1997 BY Natural History Museum Dyche Hall The University of Kansas Lawrence, Kansas 66045-2454 USA Printedby UniversityofKansasPrintingService Lawrence.Kansas CONTENTS ABSTRACT 1 RESUMEN 1 INTRODUCTION 1 Acknowledgments 3 MATERIALS AND METHODS 4 WESTERN ECUADOR 4 Physiography 8 River Systems 9 Climate 9 BiocLiMATic Regimes 12 Vegetation 17 Geologic and Climatic History 18 Human Environmental ModiAcations 23 SYSTEMATICS 24 Description ofCharacters 24 Subgeneraand Species Groups 41 KeytoSpecies 53 Clave de las Especies 58 Accounts of Species 63 Eleutherodactylusachatinus (Boulenger) 63 Eleuthewdactylusactites Lynch 66 Eleutherodactylus anatipes Lynch and Myers 68 Eleutherodactylus anonudus (Boulenger) 69 Eleutherodactylus apiculatus Lynch and Burrowes 70 Eleutherodactylusappendiculatus (Werner) 71 Eleutherodactylus babaxLynch 72 Eleutherodactylus cajamarcensis Barbourand Noble 73 Eleutherodactylus calcarulatus Lynch 74 Eleutherodactylus capriferLynch 76 = Eleutherodactylus celatorLynch 76 Eleutherodactylus cerastes Lynch 78 Eleutherodactylus chalceus (Peters) 79 Eleutherodactyluscolomai new species 81 Eleutherodactylus crenungiiis Lynch 83 Eleutherodactylus crucifer(Boulenger) 85 Eleutherodactylusdegenernew species 86 Eleutherodactylusdissimulatus new species 88 Eleutherodactylus duelhnani Lynch 90 Eleutherodactylus eremitus Lynch 91 Eleutherodactyluseugeniae new species 93 , Eleutherodactylusfloridus new species 94 Eleutherodactylusgentryi new species 97 Eleutherodactylus gularis (Boulenger) 99 Eleutherodactylus hamiotae Flores 102 Eleutherodactylus hectus Lynch and Burrowes 103 Eleutherodactylus helonotus (Lynch) 104 Eleutherodactylusillotus new species 104 Eleutherodactylus lahiosus Lynch, Rui'z, and Ardila 106 Eleutherodactylus laticlavius Lynch and Burrowes 107 Eleutherodactylus latidiscus (Boulenger) 108 Eleutherodactylus leoni Lynch 10 1 1 Eleuthewdactyliis longirostris (Boulenger) 11 Eleutherodactylus loustes Lynch 112 Eleuthewdactylus luteolatemlis Lynch 113 Eleutherodaciylus lyinani Barbour and Noble 114 Eleuthewdactylusmuricatus Lynch and Miyata 115 Eleutherodactylusnecerus Lynch 116 EleutherodactylusnyctophylaxLynch 118 Eleutherodactylus ocellatus Lynch and Burrowes 119 Eleutherodactylus ornatissimus (Despax) 120 Eleutherodactylusparvillus Lynch 121 Eleutherodactylusphoxocephalus Lynch 122 Eleutherodactyluspteridophilus new species 124 Eleutherodactyluspyrrhomerus Lynch 126 Eleutherodactylus quincpiagesimus Lynch andTrueb 127 Eleutherodactylus rosadoi Flores 129 Eleutherodactylus ruidus Lynch 130 Eleutherodactylusscolodiscus Lynch and Burrowes 131 Eleutherodactylus simonbolivariWiens and Coloma 132 Eleutherodactylus siopelus Lynch and Burrowes 133 Eleutherodactylus sobetes Lynch 133 Eleutherodactylus subsigillatus (Boulenger) 134 Eleutherodactylus sulculus Lynch and Burrowes 135 Eleutherodactylus surdus (Boulenger) 136 Eleutherodactylus taeniatus (Boulenger) 138 Eleutherodactylus teuebrionis Lynch and Miyata 138 Eleutherodactylus thymalopsoides Lynch 139 Eleutherodactylustruebae new species 140 Eleutherodactylus unistrigatus (Giinther) 142 Eleutherodactylus verecundus Lynch and Burrowes 143 Eleutherodactylus vertebralis (Boulenger) 144 Eleutherodact\'lus w-nigrum (Boettger) 145 Eleutherodactylus walkeri Lynch 148 ECOLOGY 150 TheEleutherodactylusWay ofLife 150 Eleutherodactyluscommunities 150 Discussion 159 BIOGEOGRAPHY 165 Patterns ofDistribution 165 Patterns ofSpeciation 176 HistoricalBiogeography 182 Comparisons with Other Regions 185 Comparisons with OtherTaxa 188 FUTURE RESEARCH 190 LITERATURE CITED 191 APPENDICES 197 I. Specimens Examined 197 II. Gazetteer 208 III. Species ofEleutherodactylus 219 IV. Distributionaldata 235 PLATES 1-8 Following page 76 IV ABSTRACT Western Ecuador encompasses the Pacific lowlands, a series of low coastal mountains, and the Andes, which rise to heights ofmore than 6000 m. Within this region, bioclimatic regimes range from humid tropical rainforest to desert on the lowlands, from subtropical dry and subtropical humid environments on the lower slopes of the Andes to humid temperate and subtemperate forests, subparamo, and paramo at high elevationsintheAndes.Thisregion is inhabitedby 61 speciesofEleiitherodactylus,each ofwhich istreated in anaccountthatincludesadiagnosis,descriptionorreferencetoadescription,coloration in life, natural history, anddistribution.Ninenewspeciesaredescribed,andE.alberchiisplacedinthesynonymyof£. surdus. Records forE. taeniatitsin Ecuadorarebasedon misidentifications.All morphological characters used in the identifica- tionandsystematicsof£'/£^//r//eraJ«crv/t<5aredefinedandillustrated,andakeytotheidentificationofthespecies in western Ecuador is provided. NinecommunitiescontainingsixtofifteenspeciesofEleiitherodactyluswereanalyzedwithrespecttospecies diversity,rankandrelativeabundanceofspecies,body size,andmicrohabitatuntilization. Fidelityofspeciesto bioclimatic regimes is rather low; the highest degree of endemism is 26% in the humid subtropical regime. Among the Eleutherodactyhis in western Ecuador, 32 species are endemic to the region, and 24 other species reachthesouthernlimitsoftheirdistributionsthere.Altitudinally,thegreatestnumberofspeciesisbetween 1200 m and 2200 m. Sizes ofdistributions are large at low elevations compared to smallerranges ofspecies in the Andes.Patternsofspeciationincludeuplandvicariantsoflowlandrelatives,latitudinalreplacementofsistertaxa in the Andes, and trans-Andean relatives. Key words: Ecuador; Pacific versant; Leptodactylidae; Eleiitherodactylus; Taxonomy; New species; Commu- nity ecology; Biogeography. RESUMEN LaparteoestedeEcuadorabarcalastierrasbajasdelPacifico,unaseriedemontafiasbajascostefias, y losAndesque suben aelevaciones mayoresde los6000m. Dentrodeestaregion losregimenesbioclimaticos variandesdeselvahiimedatropicalhastadesiertoenlastierrasbajas,desdeambientessecossubtropicaleshasta humedos subtropicales en las pendientes bajas de los Andes, y hasta bosques templados y subtemplados, subparamo, y paramo en las elevaciones altas de los Andes. Esta region es habitada por 61 especies de Eleiitherodactylus, cada una de estas se cubre en un sumario que incluye diagnosis, descripcion o referendaa unadescripcion,coloracionenvida,historianatural,ydistribuciongeografica.Sedescribenueveespeciesnuevas y secolocaE. alberchien lasinonimiadeE. surdus. Los registrosdeE. taeniatusenEcuadorhan sidobasados enidentificacioneserroneas.Todos loscaracteresmorfologicosutilizadosen laidentificaciony sistematicason definidos e ilustrados, y se provee unaclave para la identificacion de las especies en el oeste del Ecuador. Nueve comunidades que contienen de seis a quince especies de Eleiitherodactylus fueron analizadas con respecto adiversidad, gradode abundanciay abundancia relativadeespecies, tamaiiodel cuerpo, y utilizacion demicrohabitat.Lafidelidaddeespeciesaregimenesbioclimaticosesalgobajo;elgradomasaltodeendemismo es 26% y se encuentraen el regimen hiimedo subtropical. De las especiesde Eleutherodactyhisen el oeste del Ecuador, 32 son endemicas a la region, y las otras 24 especies llegan a su Ifmite sur en esta region. Altitudinalmente el mayor niimero de especies se encuentran entre 1100 m y 2200 m. El tamaiio de las distribuciones es grande en las tierras bajas en comparacion con las extensiones mas pequehas de las especies enlosAndes.Lospatronesdeespeciacionincluyenvicariantesenlastierrasaltasquetieneparientesenlastierras bajas, reemplazo latitudinal de especies hermanas en los Andes, y parientes transandinos. Palabrasclaves: Ecuador; regionAndina-Pacffica; Leptodactylidae; Eleiitherodactylus; Taxonomia; Especies nuevas; Ecologia de comunidades; Biogeografia. INTRODUCTION Anyone whohas ventured into the humid low- tion and rocks along streams with a headlight land rainforests of northwestern Ecuador on into immediatelyrevealstothevisitoravarietyoffrogs thecloudforestsonthePacificslopesoftheAndes ofthe genus Eleiitherodactylus, the most speciose inEcuadoratnighthasbeengreetedbyavarietyof group of vertebrates in the neotropics, with more "chirps"and"peeps."Inspectionofthelowvegeta- than 500species recognized (Duellman, 1993). In UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 manyplacesintheneotropics,especiallyinAndean ecology ofeleutherodactyline frogs in northwest- cloud forests and adjacent lowland rainforests, ern SouthAmerica. these are the most diverse and abundant anurans. Aseminal paperon Eleuthewdactyhis in west- Assuch, they shouldbe ideal subjectsforecologi- ern Ecuador was that in which Boulenger (1898) cal studies and monitoring of populations. How- described six new species (as Hylodes and ever, such has not been the case, principally be- Syrrhophus) collected by W. F. H. Rosenberg for causethese frogs havebeen anotoriously difficult the British Museum. Ofthe61 speciesnowknown group taxonomically. We anticipate that this di- from western Ecuador, only 16 ofthese were de- lemma can be remedied by thorough and precise scribedpriortoourworkintheregion. Lynch(and taxonomic treatments ofmembers ofthe genus in coworkers) described 32 species, and nine are various geographic regions. named herein; three other species {E. hamiotae Webegan ourinvestigations ofthe Ecuadorian Flores. E. wsadoi Flores, and E. sunonholivari herpetofaunainthe mid-1960s. Duellman worked Wiens and Coloma) have been named in recent sporadically inwesternEcuadorfrom 1967-1984. years. Eightofthe 11 speciesofEleuthewdactylus Lynch's field work there in 1967-1979 focused nowknownfromcloudforestatTandapi,Ecuador, almost exclusively on Eleuthewdactyhis, and al- weretreatedbyLynch(1976a).LynchandBuirowes though his interests shifted to the rich Colombian (1990) reported on the Eleuthew-dactylus in the faunaabout 15yearsago,heretainedanobligation cloudforestatLaPlanadainsouthwesternColom- tosummarizetheEleiitherodactylusfaunaofEcua- bia;oftheeightspeciesdescribedasnew(andthen dor. Four summaries ofthe Eleuthewdactylus of knownonlyfromColombia),wehavefoundsixin geographical regions of the country have ap- collections from Ecuador. Some ofthe most com- — peared the Andes of southern Ecuador (Lynch, monspeciesonthePacificlowlandsofEcuadorare 1979a), the Amazon Basin (Lynch, 1980a), the members of the paraphyletic Eleuthewdactylus AndesofnorthernEcuador(Lynch. 1981a),andthe "fitzingeri"groupofLynch(1976)andLynchand AmazonianslopesoftheAndesofEcuador(Lynch Myers (1983), the Chocoan members of which and Duellman, 1980). Most recently, we (Lynch weretreatedmonographicallybyLynchandMyers and Duellman, 1995) described E. philipi from (1983). paramos in southern Ecuador. Mostofthe speciesknownfromwesternEcua- The present work is a summary of the dor have been described, and we do not repeat Eleiitherodactylus ofthe Pacific lowlands and the descriptionsintheaccountsofthosespecies;rather, westernslopesoftheAndesofEcuador.Herein,the we provide references to adequate descriptions. upperaltitudinal limit isarbitrary; itisdefinednot Complete descriptions are given for the nine spe- by elevation but, rather, by assemblages of cies named herein. We provide a redescription of Eleuthewdactylus.Somespecies(e.g.,E.curtipes) Eleuthewdactylus gularis (Boulenger) because fromtheparamosdoencroachintotheuppercloud existing descriptions (Boulenger, 1898; Cochran forests, but we do not include these species; they andCoin, 1970) are inadequate andbecause more weretreatedbyLynch(1981a).Ourintenthereisto than one species has been confounded under that treat the species confined to forested regions of name. For all species, we provide lengthy, num- western Ecuador. This small fragment ofthe total bereddiagnoses, includingcomparisonsandcom- range ofthe genus contains more than 12% ofthe ments,evenwhendoingsoresultsinduplicationof speciesoiEleuthewdactylus. Inadditiontoathor- materialpublishedelsewhere;wehavedonethisto oughtaxonomictreatment,weexaminepatternsof facilitatecomparisonsandidentifications.Accounts distribution with respect to climate, vegetation, oftwospecies,E. siopelusandE. sulculusthatare topography, and historical biogeography. We also known from extreme southwestern Colombia but focus on several eleuthero-dactyline communities yet to be recorded from Ecuador, are included in western Ecuador and compare these with com- becauseweareconfidentthattheywillbefoundin munities elsewhere. In so doing, we hope to pro- Ecuador.Ourotherdepartureistheinclusionofthe vide the ground work for future studies on the onlyothereleutherodactylines(Barycholospulcher ELEUTHERODACTYLUS IN WESTERNECUADOR and Phyllonastes sp.) known from western Ecua- ResearchCouncil.Fundingforthecolorplateswas dor in the key and in comparisons in appropriate provided by the School of Biological Sciences, diagnoses. University ofNebraska, Lincoln. Thisisthe lastinaseriesofpapersonEcuador- For the loan of specimens and/or providing ian eleutherodactylines, and we take thisopportu- working space in their respective institutions, we nity to review, and expand on, the characters we are grateful to Darrel R. Frost, Charles W. Myers, find to be most useful in the identification ofspe- and Richard G. Zweifel, American Museum of cies. This concern for well-defined characters re- Natural History: Alice G. C. Grandison, British flectsLynch'sinterestinthephylogeneticrelation- Museum (Natural History);Alan E. Leviton, Cali- shipswithinthegenusandthedesiretohavegroups forniaAcademyofSciences;RobertF Inger.Field of species defined by shared derived characters Museum of Natural History; Jose R Rosado and ratherthanbypheneticsimilarity.Theinitialunder- ErnestE.Williams,MuseumofComparativeZool- standingofrelationshipswithinEleiithewdactylus ogy, Harvard University; Josef Eiselt, Natur- came with the recognition that the subgenus historisches Museum, Wien; Dorothy Smith, Uni- Craugastorcould be diagnosed by aderived con- versity of Illinois Museum of Natural History; ditionofthem.adductormandibulae(Lynch, 1986). Arnold G. Kluge, Ronald A. Nussbaum. and the That discovery caused JDL to question (and ulti- late Charles F. Walker, University of Michigan mately to reject) many ofhis earlier groupings of MuseumofZoology;thelateDorisM.Cochran.W. Eleutherodoctylus and to continue to search for Ronald Heyer, RoyW. McDiarmid, the lateJames synapomorphiesforvariousunitswithinthegenus. A.Peters,andGeorgeR.Zug,NationalMuseumof Natural History. We acknowledgethecooperation Acknowledgments ofAnaAlmendariz(EscuelaPolitecnicaNacional), Luis A. Coloma (Pontificia Universidad Catolica More than a quarter of a century has passed del Ecuador), and Alexandra Quiguango (Eco- sinceweinitiatedourworkinEcuador.Duringthat ciencia) for loaning specimens from Ecuadorian time we have built up a debt ofgratitude to many collections and providing dataon localities in Ec- persons who contributed directly or indirectly to uador.ColomaandMarthaL.Crump,whoactedas the completion of this work. We are especially liaison with Ecociencia, provided color slides, as grateful to our field companions whose efforts didPatriciaA. Burrowes and RoyW. McDiarmid. greatly enhanced the collection ofspecimens and Credits forphotographs in Plates 1-8 are noted in fielddata. Atvarioustimes,JDLwasaccompanied the legends as: JDL = John D. Lynch, KM = byThomasJ.Berger,DavidC.Cannatella,Thomas Kenneth Miyata, LAC = LuisA. Coloma, MLC = H.Fritts,RobertW.Hen-derson,MarshaC.Lynch, Martha L. Crump, PAB = Patricia A. Burrowes, and Gerald R. Smith. Duellman variously was RWM = Roy W. McDiarmid. WED = William E. accompanied by Patricia A. Burrowes, Dana T. Duellman.WethankPatriciaA. BurrowesandErik Duellman,DavidM.Hillis,BruceMacBryde,Alan R. Wild forhelpful suggestions on the analysis of H. Savitzky, John E. Simmons, and Linda Trueb. the ecological data, Joseph R. Mendelson III for Duellman'sfieldworkwasaccomplishedaspartof checking specimens against our list ofspecimens research supportedby the National Science Foun- examined, Santiago F Burneo and Ignacio de la dation (Grants DEB 74-02998, 76-09986, and 82- Riva forproviding the Spanish version ofthe key. 19388)andtheNationalGeographicSociety(Grant and Rafael Joglar for assistance with the Spanish 1304);thebiogeographic analysisherein is partof Resumen. We also thank Eugenia del Pino and a project on patterns of anuran speciation and FernandoOrtizC,whogenerouslyprovidedfacili- biogeography supported by the National Science ties for our use at the Universidad Catolica in Foundation (Grant BSR 8805920). Lynch's field Quito, and to Sergio Figueroa andAbel TovarV., workwasmadepossiblebygrantsfromtheSociety MinisteriodeAgriculturay Ganaderfa. forissuing of the Sigma Xi, Watkins Fund of the Natural permits for collection and exportation of speci- History Museum, and the University ofNebraska mens. — UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Anearlierversionofthis manuscriptbenefitted withtheelectronictransferofillustrationsresulted from thoughful comments by Martha L. Crump, in the improvement of our otherwise benign ef- throrough review by LuisA. Coloma. and critical forts. Lastly, we dedicate this paperto the memo- evaluationbyJayM.Savage,whoencouragedusto ries of two deceased friends and colleagues eschewcertainhectoringtermsinourdiscussionof Kenneth Miyata and James A. Peters; they loved anatomy. To these three persons in particular we Ecuador and collected many ofthe specimens on extend our profound thanks. We especially thank which this work is based. LindaTrueb. whosecareful editingandassistance MATERIALS AND METHODS Specimens are identified by standardized mu- taining structural features are defined in a follow- seum codes designated by Leviton et al. (1985), ing section. Description of Characters. Measure- exceptfortheadditionoffourcollectionshousedin ments were taken with dial calipers to the nearest Quito, Ecuador: ECO =Ecociencia (specimens to 0.1 mm; measurements,andproportionsaregiven be deposited in a museum), EPN = Escuela foreach sex when their means differ significantly Politecnica Nacional, MECN = Museo Ecuatori- (P<0.05);otherwisethesedataarecombined.The anodeCienciasNaturales,andQCAZ=Museode following abbreviations are used: E-N = eye-nos- Zoologfa, Pontificia Universidad Catolicadel Ec- tril distance; HL=head length, HW=headwidth, uador. Some specimens collected recently byAna lOD = interorbital distance, SVL = snout-vent Almendariz are identified by field numbers (e.g., length. Areas ofdistribution were measured from EPN-AA). References are made to some speci- mapsusingaMicro-PlanIIimageanalysis system mens in Colombian collections: ICNMHN = (Laboratory ComputerSystems, Inc., Cambridge, InstitutodeCienciasNaturales,MuseodeHistoria MA). Natural, Universidad Nacional de Colombia, Throughoutthetext,wehaveavoidedusingthe Bogota;IND-AN=amphibiancollection,Instituto termsseniorandjuniorauthor,becauseinthiscase Nacional de los Recursos Naturales Renovables y thejunior is seniorchronologically; therefore, the delAmbiente, Bogota; UVC = Universidad Valle authors are noted simply as JDL and WED. Al- de Cauca, Call. though we both assume responsibility forthe ma- All specimens from western Ecuadorthat have terial presented herein, JDL is the primary author WED been studied are listed in Appendix L Localities for the sections on systematics, and is the from which specimens have been examined are primary author for the sections on western Ecua- listedwiththeirgeographiccoordinatesandeleva- dor,ecology,andbiogeography.Allmorphological tions(whenknown)inAppendixILLocalitiesthat illustrationsweredrawnbyJDLusingadissecting havebeenlocatedareplottedondistributionmaps microscope equipped with a camera lucida. All WED intheAccountsofSpecies;becauseofcrowdingof graphs and maps were generated by using symbols, somecloselyapproximatedlocalitiesare Aldus Freehand® on a Macintosh computer. The not shown. colorphotographs werereproducedelectronically Methods for taking measurements and ascer- from colortransparencies. WESTERN ECUADOR Western Ecuador encompasses the area from parts of the provinces of Azuay, Carchi, Cafiar, ColombiatoPeruandfromthePacificOceantothe Chimborazo, Cotopaxi, Imbabura, Loja, and westerncrestoftheAndes; thisregion includesall Pichincha (Fig. 1). In order to provide a working of the provinces of Bolivar, El Oro, Esmeraldas, basisfortheecologicalandbiogeographicanalyses Guayas, Los Rios, and Manabi, and the western oftheEleutherodactylusoftheregion, weprovide