Frogs of the genus Eleutherodactylus (Leptodactylidae) in western Ecuador : systematics, ecology, and biogeography

252 Pages·1997·19.8 MB·English

by  Lynch, John D

rogs of the
Genus
Eleutherodactylus
Western Ecuador
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Sj^stematics, Ecology,
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John D. Lynch and William E.
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THEUl Museum of Comparative Zoology { MUSEUM
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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