I Herp I QL I 668 I .E2 I R53 I 1994 IqQuitos Region, Amazonian Peru ReticulatedPoison Frog Dendrohates reticiilanis i Lily O. Rodriguez | AND I William E. Duellman \ MaseuBJ of Comparative Zoology Llbr&r? Harvard University A copy of the Guide to the Frogs ofthe Iquitos Region, AmazonianPerumaybeobtainedbywritingto: PublicationsSecretary NaturalHistoryMuseum TheUniversityofKansas Lawrence,Kansas 66045.USA ThechargeforeachcopyoftheGuideis$19.95U.S.currency, plus $1.50 for postage and handling. (Overseas postage is $3.00.) MasterCard and VISA are accepted. To order by phone,call913-864-4540. AfreecopyofourmostrecentCatalogueofPublications maybeobtainedfromtheaboveaddress. The University of Kansas Natural History Museum SpecialPublication No. 22 December 1994 GUIDE TO THE FROGS OF THE IQUITOS REGION, AMAZONIAN PERU By Lily O. Rodriguez AND William E. Duellman Published by asociaciondeecologiayconservacion, AMAZONcenterFORENVIRONMENTALEDUCATIONAND RESEARCH, AND NATURALHISTORYMUSEUM,THEUNIVERSITYOFKANSAS Lawrence, Kansas 1994 UNIVERSITYOFKANSASPUBLICATIONS,NATURALHISTORY MUSEUM Editor:LindaTrueb SpecialPublicationNo.22 pp.i-ii, 1-80 Plates 1-12 PublishedDecember 1994 ISBN0-89338-047-4 ^O^^^ NaturalHistory Museum \J\^^ TheUniversityofKansas Lawrence,Kansas 66045 ^M ^'' USA Frontcover:Dendrobatesreticulatus. PhotographbyW.W.Lamar CoverandpublicationdesignbyLindaTrueb ©NaturalHistoryMuseum,TheUniversityofKansas Printedonacid-free,recycledpaper BY UniversityofKansasPrintingService Lawrence,Kansas Contents BIOGRAPHICALSKETCHES iv PREFACE V INTRODUCTION I BiologyofFrogs 1 HowtoUsethisGuide 2 Names 4 Terminology 5 GUIDETOTHEFROGS 11 Toads (FamilyBufonidae) 11 PoisonArrowFrogs (FamilyDendrobatidae) 15 Treefrogs (FamilyHylidae) 20 Egg-broodingFrogs{GenusHemiphractus) 21 Treefrogs(Genus//y/a) 22 Broad-headedTreefrogs(GeneraOsteocephalusand 39 Phrynohyas) LeafFrogs{GentxdiAgalychnisandPhyllomedusa) 43 Narrow-headedTreefrogs(GeneraScarthylaandScinax) 47 Hatchet-facedTreefrogs{GenusSphaenorhynchus) 50 TerrestrialFrogs (FamilyLeptodactylidae) 52 Eleutherodactylines 53 Leptodactylines 64 Narrow-mouthedFrogs (FamilyMicrohylidae) 73 AquaticFrogs (FamilyPipidae) 77 PondFrogs (FamilyRanidae) 78 SELECTEDREFERENCES 79 PLATES 1-12 Followingp. 80 MAPOFIQUITOS REGION InsideBackCover Biographical Sketches TheauthorsatExploramaLodge,Peru,April 1993. PhotographbyCharlesOlmstead. LilyO.RodriguezreceivedherdoctoraldegreeinecologyattheUniver- sity ofParis in 1991. Since 1979, she has conducted field research in Amazonian Peru on the taxonomy, ecology, and life histories offrogs. SheisaResearchAssociateoftheMuseodeHistoriaNatural,Universidad NacionalMayordeSanMarcosinLima,Peru,andaFieldAssociateof theAmericanMuseumofNaturalHistory,NewYork. Address: Museo deHistoriaNatural,Apartado 14-0434,Lima 14,Peru. WilliamE.DuellmanreceivedhisPh.D.fromTheUniversityofMichi- ganin 1956. Since 1959,hehasbeenassociatedwithTheUniversityof Kansas,whereheisCuratorofHerpetologyintheNaturalHistoryMu- seumandProfessorintheDepartmentofSystematicsandEcology. Most ofhismorethan270publicationsdealwiththesystematics,lifehistory, andecologyoffrogsintropicalAmerica. Address:NaturalHistoryMu- seum,UniversityofKansas,Lawrence,Kansas66045-2454,USA. Preface Within the past two decades devastation oftheAmazon rainforest has attracted the attention ofnotonly biologists and conservationists, butalso thegeneralpublic. Biologistshaveendeavoredtoinventory thebiota,and conservationists have struggled to preserve large tracts of the rainforest. Amateurnaturalistsandlaypersonsfromallwalksoflifehaveventuredto theAmazon Basin towitnessthe wondersofthis, the largestrainforeston earth;asaconsequence,manyofthesepersonshavebecomeeffectiveadvo- catesfortheconservationandstudyoftherainforestandthefantasticwealth oflivingorganismsfoundthere. Iquitos,Peru,isoneofthemostimportantdeparturepointsforbiological investigations,conservationefforts,andecologicallysensitivetourism. (Other majorpointsarePuertoMaldonado,Peru,andManausandBelem,Brazil.) Through the combined efforts ofExploramaTours ofIquitos and Interna- tional Expeditions, Inc., in the USA, a series of International Rainforest Workshopswasinitiatedin 1991. Theseworkshopshavesubsequently in- volveddozensofvolunteerinstructorsandservedasaneducationalexperi- enceforseveralhundredworkshopparticipants. Itwaswhileservingasan instructorinthefirstoftheseworkshopsthatthefirstauthorobservedmany photographsoffrogstakenintheareapreviouslybyJamesH. Marlett,and the ideaforaguidetothe frogs wasconceived. In subsequentworkshops (1992-1994),thetwoofuscombinedoureffortstoproducethisguide. WearegratefultoRichardRyelandRichardMills,InternationalExpedi- tions,Inc.,forinvitingustoparticipateintheworkshops. PeterJensonand hisstaffofExploramaToursprovideduswithcomfortableaccommodations andefficientlogistics. Innumerableworkshopparticipantsfoundmanyfrogs andposedcountlessquestionsthathelpedusformulatethisbook. Forpro- vidingsomeofthephotographsusedherein,wethankLuisA.Coloma(LAC), MichaelJ.Doolittle(MJD),WalterHodl(WH),JohnD.Lynch(JDL),James H.Marlett(JHM),RoyW.McDiarmid(RWM),CharlesW.Myers(CWM), AndrewYoung(AY),andespeciallyWilliamW. Lamar(WWL). Theirre- spectivephotographsareidentifiedbytheforegoinginitialsinthecaptions; photographs by the authors are identified by the initials LOR and WED. Figures 1-3 were drawn byAmyLathrop, whodeservesaccoladesforthe clarityofherillustrations. WeareespeciallygratefultoJoelSoudersofthe IMG Corporation ofLawrence, Kansas, forhis efforts and cooperation in providingimagesscannedfrom35-mmtransparenciesinanelectronicfor- mat; these were composed into the color plates using PageMaker^"^ on a Macintoshcomputer. WeareespeciallyindebtedtoLindaTrueb,whothor- oughly edited the textand skillfully manipulated the textand illustrations electronically, thereby achieving the high quality ofthis publication. The entiretextwasreadbyWilliamW.LamarandErikR.Wild;wethankthem fortheirhelpfulcomments. RichardMillsandB.AnthonyLuscombealso offerconstructivecomments on the text. Publicationwas achievedbythe collaborative efforts of B. Anthony Luscombe, President, Asociacion de Ecologiay Conservacion (ECCO); Dick Mills andRichardRyel,Amazon CenterforEnvironmentalEducationandResearch(ACEER);andPhilipS. Humphrey,Director,NaturalHistoryMuseum,TheUniversityofKansas. Lastly,weareindebtedtothefrogsthemselves. Theirbeautyandfasci- nating behavior have captivated our attention for many years. Although they do not receive the same general recognition as birds, cats, and mon- keys,theydoplayanimportantroleintherainforestecosystem,forwhich theydeservefarmorecreditthantheyusuallyareaccorded. LilyO.Rodriguez WilliamE.DuelIman Explorama,Loreto,Peru March1994 Introduction About4,000speciesoffrogsarerecognizedworldwide;nearly 1,600of these liveinSouthAmerica,wherethegreatestnumberofspeciesoccurin theAmazonrainforestandinthehumidmontaneforestsontheslopesofthe Andes. More than 80 species have been found at single sites in the upper Amazon BasininEcuadorandPeru; nowhereelse inthe world isthe frog faunamorediverse.Forcomparison,thisnumberisaboutequaltoallofthe speciesknown intheUnitedStates andCanadaandmorethanthree times thenumberofspeciesknownfromallofEurope.TheIquitosregionhasa tremendousdiversityoffrogs,andrecentinvestigationshavediscoveredfive speciesnotpreviouslyknownfromthatregion,aswellassomespeciesthat arenewtoscience. Hereinwepresentaccountsof112speciesoffrogsknownfromtheIquitos region; 111 oftheseareillustratedincolor.Thisguid—eisdesignedforuseby amateur naturalists and professional herpetologists anyone interested in identifying adult frogs or larvae from the region and learning something abouttheirnaturalhistory. BIOLOGYOFFROGS Frogsandtoadsareone(Anura)ofthethreeordersoflivingamphibians; the others are Caudataand Gymnophiona. The Caudata includes the sala- manders, which are most diverse in the Northern Hemisphere. Only two generaofsalamandersenterSouthAmerica, andone speciesoccurs inthe Iquitosregion.TheGymnophionaareelongate,limbless,wormlikeamphib- ians.Thesepoorlyknownanimalsusuallyliveunderground(someareaquatic) intropicalregions;severalspeciesareknownfromtheIquitosregion.Frogs differfromotheramphibiansbybeingspecializedforjumping.Thesespe- cializationsincludeashorteningofthebody,absenceofatail,andpresence of long, powerful hind limbs. Like the skin ofother amphibians, that of frogsispermeabletowater,whichinwarm,dryairwillevaporatefromthe body and result in desiccation and death. Consequently, frogs most com- monlyliveinplaceswhereeitherthegroundand/orairaremoistorwhere they can replenish theirwatersupply by entering water. Replenishmentis accomplishedbysoakingupmoisturefromthesubstrateorair,notbydrink- ing. Unlikemostterrestrial animals suchas salamanders, snakes,andmam- mals,frogsdonotleavescenttrailstomarkterritoriesandidentifypotential mates.Instead,frogs,likebirds,advertisetheirpresencebysound.Withfew exceptions,onlymalefrogsvocalize;soundsareproducedwhenairisforced fromthe lungsoverthevocalcordsintothemouthandintoanexpandable vocalsacthatactsasaresonatingchamber.Inagivenregion,eachspecies offrog has a distinctive call that is recognized by other members ofthe 1 2 RODRIGUEZANDDUELLMAN species(andprobablybysomepredators).Somefrogsproducedifferentkinds ofnotes;themostcommonisthecourtshipcalltoattractfemales,whereas territorialcallsarerecognizedbyothermalesandencountercallsarepro- ducedduringclose-rangeinteractionsbetweenmales. MostNorthAmericansandEuropeansreceivedastereotypedconceptof amphibianreproductioninhighschoolorcollegebiologycourses,inwhich thelifehistory of''thefrog" waspresentedas asimple, annual, three-step — process (1) clumps ofeggs are laid in ponds and fertilized by the male whilehegraspsthefemale; (2)eggshatch intoaquaticlarvae (tadpolesor poUywogs)thatfeedonalgae,growlegs,andmetamorphose intofroglets; and (3) youngfrogsmove ontoland, where theyfeedon insects andgrow into adult frogs that return to the pond the following spring to repeat the reproductiveprocess.AlthoughmostfrogsinNorthAmericaandEuropedo follow this pattern, there are many otherdiverse andfascinating modes of anuranreproductionanddevelopment,especiallyinthetropics.IntheIquitos region,manyfrogsbreedthroughouttheyear,andonefemalemaydeposit manyclutchesofeggsannually;otherspecieshaveexplosivebreedingperi- odsfollowingheavyrains.Manytreefrogsmateinthetreesanddeposittheir eggs onvegetationoverponds; the hatchlingtadpoles drop intothe water. Anothergroup(Leptodactylines)deposittheireggsinfoamlikenestsonthe surfaceofpondsoronlandadjacenttoponds;theeggsdevelopinthenest, which reduces chances ofdesiccation and predation. Eleutherodactylines anddendrobatidsdepositeggsonland;intheformer,thelargeeggsundergo direct development into froglets without having an aquatic tadpole stage, whereasindendrobatidshatchlingtadpoleswriggleontothebackofanat- tending adult who carries the tadpoles to water. There are other kinds of parental care that can be observed in frogs in the Iquitos region. Forex- ample,femaleegg-broodingtreefrogscarrytheireggsontheirback where they develop directly into froglets. In females ofthe aquatic frogs ofthe genusPipa,eggsareimbeddedintheskinonthebackwheretheyundergo directdevelopment intofroglets. In some dendrobatids and treefrogs, tad- polesdevelopinwaterinarborealbromeliads,andthefemalesperiodically returntothebromeliadstodepositunfertilizedeggsforthetadpolestoeat. Despiteoursteadilyincreasingknowledgeaboutthenumbersandkinds offrogsandtheirbehaviorandlifehistories,littleinformationexistsabout manyofthespeciesintheIquitosregion.Excitingnewinformationcanbe gainedbyanyobservantpersonwhohappenstobeintherightplaceatthe right time when frogs are doing something never observed and recorded before. HOWTOUSETHISGUIDE Thisguideisdesignedtohelpyouidentifyfrogs.Relatedfrogsaregrouped undertheheadingsgiveninthetableofcontents; followingtheaccountof