TABLE OF CONTENTS IRCF REPTILES & AMPHIBIANS • 19(4):254 ...€¦ · IRCF REPTILES &...

IRCFREPTILES&AMPHIBIANS•19(4):254–262•DEC2012

etal.2007).Attheepicenterofthisdiversityliesoneoftheworld’smostprominentfacilitiesforecotourismandconser-vation,ReservaAmazónica,formerlyCuscoAmazónico. Since1975,Inkaterra(www.inkaterra.com),theorga-nizationthatoperatesReservaAmazónicaandseveralotherPeruviantourismlodges,haspromotedecologicalresearch

TheTambopataregionofPerùyieldsthehighestbiodiver-sityofreptilesandamphibiansintheworldandincludes

morethan210amphibianandreptilianspecies(DoanandArizábal2002).Previoussurveysoftheregion’sbiodiversityhavebeenusedtocategorizetheIUCNandCITESstatusofspeciesandcontributedtofuturereservedesign(vonMay

Herpetological Ecotourism and Conservation: Reserva Amazónica, Perù

SarahA.Orlofske1,HaydenD.Hedman1,JoséE.Koechlin2,andRobertC.Jadin1

1DepartmentofEcologyandEvolutionaryBiology,UniversityofColoradoBoulder,Boulder,Colorado80309,USA(rcjadin@gmail.com)2Inkaterra,Lima,Peru

PhotographsbyRobertC.Jadin,exceptwherenoted.

254

IRCF REPTILES & AMPHIBIANS • VOL15, NO 4 • DEC 2008 189TABLE OF CONTENTS

T A B L E O F C O N T E N T S

F E A T U R E A R T I C L E S

Chasing Bullsnakes (Pituophis catenifer sayi) in Wisconsin: On the Road to Understanding the Ecology and Conservation of the Midwest’s Giant Serpent ...................... Joshua M. Kapfer 190

The Shared History of Treeboas (Corallus grenadensis) and Humans on Grenada: A Hypothetical Excursion ............................................................................................................................Robert W. Henderson 198

R E S E A R C H A R T I C L E S

The Texas Horned Lizard in Central and Western Texas ....................... Emily Henry, Jason Brewer, Krista Mougey, and Gad Perry 204 The Knight Anole (Anolis equestris) in Florida

.............................................Brian J. Camposano, Kenneth L. Krysko, Kevin M. Enge, Ellen M. Donlan, and Michael Granatosky 212

C O N S E R V A T I O N A L E R T

World’s Mammals in Crisis ............................................................................................................................................................. 220 More Than Mammals ...................................................................................................................................................................... 223 The “Dow Jones Index” of Biodiversity ........................................................................................................................................... 225

H U S B A N D R Y

Captive Care of the Central Netted Dragon ....................................................................................................... Shannon Plummer 226

P R O F I L E

Kraig Adler: A Lifetime Promoting Herpetology ................................................................................................ Michael L. Treglia 234

C O M M E N T A R Y

The Turtles Have Been Watching Me ........................................................................................................................ Eric Gangloff 238

B O O K R E V I E W

Threatened Amphibians of the World edited by S.N. Stuart, M. Hoffmann, J.S. Chanson, N.A. Cox, R. Berridge, P. Ramani, and B.E. Young .............................................................................................................. Robert Powell 243

CONSERVATION RESEARCH REPORTS: Summaries of Published Conservation Research Reports ................................. 245 NATURAL HISTORY RESEARCH REPORTS: Summaries of Published Reports on Natural History ................................. 247 NEWBRIEFS ...................................................................................................................................................................................... 248 EDITORIAL INFORMATION ..................................................................................................................................................... 251 FOCUS ON CONSERVATION: A Project You Can Support ............................................................................................... 252

Front Cover. Shannon Plummer.Totat et velleseque audant mo estibus inveliquo velique rerchil erspienimus, quos accullabo. Ilibus aut dolor apicto invere pe dolum fugiatis maionsequat eumque moditia erere nonsedis ma sectiatur ma derrovitae voluptam, as quos accullabo.

Back Cover. Michael KernTotat et velleseque audant mo

estibus inveliquo velique rerchil erspienimus, quos accullabo. Ilibus

aut dolor apicto invere pe dolum fugiatis maionsequat eumque

moditia erere nonsedis ma sectia-tur ma derrovitae voluptam, as

IRC

F

REPTILES & AMPHIBIANSC O N S E R V AT I O N A N D N AT U R A L H I S T O R Y

Copyright©2012.SarahOrlofske.Allrightsreserved.

WWW.IRCF.ORG/REPTILESANDAMPHIBIANSJOURNAL

Figure 1.ThecanopywalkatReservaAmazónicaismorethan30mhighand344mlongandhastwotowers,eightplatforms,andsevenbridges.Recently,National Geographic AdventurerecognizedReservaAmazónicaaspossessingthe“MostExtraordinaryCanopyWalkway”(Davis2009),whichevenincludesaprivatecabinforadventurousguests.Touriststakeearlymorninghikesinsearchofthemanyspeciesofimpressivebirdsthathavebeenrecorded—butherpetologistscanseeotherspeciessuchasGreenThornytailIguanas(Uracentron azureum,insert).

255

IRCFREPTILES&AMPHIBIANS•19(4):254–262•DEC2012

andconservationsupportedbytourism.Themissionstate-mentofInkaterrais“conservingtheenvironment,preserv-ingthenativecultures,anddevelopingsustainabletourisminPerù.”Inkaterra’sstrategyforimplementingthismissionistoconservetheregion’sbiodiversitywhilealsoeducatingtouristsaboutthenaturalenvironment.ReservaAmazónicahasreceivedawardsfromorganizationsandpraiseinmanyinternational travelmagazines for its conservationeffortsandecotourism(Fig.1).Inkaterra’smissionforconservationremainsevidentthroughvariouscollaborationswithresearchscientists.Inparticular,collaborationswithherpetologistsattheUniversityofKansashaveresultedinReservaAmazónicabeingoneofthemostwellstudiedareasinSouthAmericaforamphibiansandreptiles.Inparticular,Duellman’sbookCuscoAmazónico(2005)extensivelydocumentstheregion’sbiogeographyandherpetofaunaldiversity. WithinfourhoursbyboattoBolivia,ReservaAmazónicasitsalongtheAmazonBasininTambopata,Perù.Theexclu-siveresortresideswithinaone-hourboatridefromthenear-est city,PuertoMaldonado (Fig.2).ReservaAmazónicaallowseachguestpersonalizationoftheirrainforestexcursionsregardlessofweatherconditionsandlengthofstay.Bilingualnatureinterpreters,specializedinthefloraandfaunaofthe

Tambopataregion,facilitatetheseexcursions.Visitorscanwalkalongwellestablishedtrailsystems,includingawet-landsboardwalk;theycantakecanoeexcursionsaroundlakesValenciaandSandoval,aswellasGamitanaCreek;ortheycantakeatourtoariverisland,alargebutterflyhouse,orchidandmedicinalplantgardens,andmore.Thepackagesreso-natewithalllevelsofbiologicalandtravelenthusiasts.Beyondthenaturalmarvels,theresortencompassesfullbarservices,exquisitethree-coursecuisine(vegetarianoptionsincluded),andriverfrontmassagetherapy. Ourpersonalexpeditionsdescribedinthisarticlewerefocusedonamphibiansandreptiles(Fig.3).NotonlywerewevisitingReservaAmazónicatoobservethesespecies,buttostudytheirdiversity,abundance,habitatuse,andspeciesinteractions.Throughourmonitoringstudyincollabora-tionwithReservaAmazónica,wehelpfillacriticalneedforlong-termpopulationdataontropicalspecies(Fig.4),whilesimultaneouslyequippingtheinterpretersandtouristswithinformationabouttheseimportantorganisms.

Up Close and PersonalAround the bungalow area during the day, AmazonRacerunners(Ameiva ameiva)commonlysprintfrombeneath

ORLOFSKEETAL.

Figure 2.TypicalriverboatontheMadredeDiosRiverduringsunsettakenfromtheboatdockatReservaAmazónica.

256

ORLOFSKEETAL. IRCFREPTILES&AMPHIBIANS•19(4):254–262•DEC2012

bungalowstepstonearbybushesandpaceacrossthefoot-paths.Atnight,thebungalowporchesareilluminatedwithindividualkerosenelanterns.Thelightinghelpsavoidstep-pingontheabundantCaneToads(Rhinellamarina)forag-ingalongthepaths.Numerous treefrogs (e.g.,Gunther’sBandedTreefrog,Hypsiboas fasciatus,andMilkyTreefrog,Trachycephalus typhonius [formerly Phrynohyas venulosa];Fig.5)commonlyperchonthevegetationaroundthecab-ins.FromwithinthecabinsyoucantypicallyhearcallsofBolivian Sheepfrogs (Hamptophryne boliviana), Amazon

Rainfrogs(Leptodactylus[Adenomera] spp.),andseveralspe-ciesoflargeLeafFrogsinthegenusPhyllomedusa.ThecallsoftheBolivianSheepfrogsareunmistakable,soundingremark-ablylikesheep. Alargebridgeelevatedaboveasmallswampconnectsthebungalowareatothemainlodge.Thisswampyareaishometonumerousspeciesoffrogsthatcallintheeveningandcaneasilybeobservedwithaflashlight.Onmorethanoneoccasion,aswestoppedtoinspectthecolorful leavesoftheHeliconia, a groupoftouristsandinterpreterswould

Figure 4.TheendemicKoechlin’sTreefrog(Dendropsophus koechlini;a.adultmale,b.amplectantpair)wasdescribedfromReservaAmazónica(DuellmanandTrueb1989).Althoughconsideredoneofthemostabundantspeciesinthearea,theactivityperiodofD. koechliniisshortandcorrelatesstronglywithitsbreedingseasonthatstartswiththefirstheavyshowersoftherainyseason.

Figure 3.TheBlack-spottedSkink(Mabuya nigropunctata)istheonlyskinkintheregion;itisbronzeandblackwithsmoothandshinyscales.ThisspeciesisfrequentlyseenontrailsatReservaAmazónicabut,likethisindividual,canalsobefoundonthewalktoLagoSandoval.ThecanoetouraroundLagoSandovalisverypopularasvisitorsoftenencountermonkeys,macaws,riverotters,andevenBlackCaiman(Melanosuchus niger).

ORLOFSKEETAL. IRCFREPTILES&AMPHIBIANS•19(4):254–262•DEC2012

257

gatheraroundusforacloserlook,givingusanopportunitytodescribethespeciesandourresearchinmoredetail.Everyoneleavesthebridgealittlereluctantlyaftersuchcloseandunex-pectedencounterswithbeautifulBasinTreefrogs(Hypsiboas lanciformis;Fig.6)andotherspecies.

Exploring the Trails — Day and NightThetrailsystematReservaAmazónicaiswellmarkedandincludestrailsaslongasseveralkilometers.Interpretersareavailabletoleadprivatetoursbothbydayandnight.Bootsarehighlyrecommendedwhilewalkingonthetrailsforbothsafety

Figure 5. One night at about 2 am, the calls of Milky Treefrogs(Trachycephalus typhonius) weresoloudthatweawoketofindthemalecallingfromourcabinshower!Wehadtobecarefulduringhandlingtonotalarmthefrogintoreleasingthedefensiveglue-likemilkysecretionsfromglandsinitsskin,fromwhichitgetsitscommonname.Apreviousencounterwiththisspeciestaughtusthatthesecretionsarenearlyimpos-sibletoremove.

Figure 7.Fivespeciesofturtlesoccurintheforests,ponds,andriversinandaroundReservaAmazónica.TheseincludetheYellow-footedTortoise(Chelonoidis denticulata).Adultticks(Amblyomma humerale)parasitizedbothtortoisesobservedbytheauthorsduringseparatetripstoReservaAmazónica.PhotographbySarahA.Orlofske.

Figure 6.AlthoughrecordedfortheTambopataregion(vonMayetal.2007),thisphotographisthefirstrecordoftheBasinTreefrog(Hypsiboas lanci-formis)atReservaAmazónica.Thisindividual,alargefemale,wasfoundjustoutsidethemainlodge,apparentlyattractedtothecallsofmalesinchorusthroughoutthemonthofDecember.

ORLOFSKEETAL.

258

IRCFREPTILES&AMPHIBIANS•19(4):254–262•DEC2012

andsupportthroughthemuddyterrain.Thelodgehasrubberbootsinallsizesavailableforguests.Onceequippedwithboots,youwillonlyhavetotakeafewstepsfromyourcabanatoseesomeinterestingreptilesandamphibians(Fig.7). Manytimesduringthedayandatnightwewouldgatherournotebooksandcameras,putonourbootsandheadoutonthetrails.OnTrail-Awesawplentyofactivityintheleaflitterandonthetrees.AfewminutesintoourtrailwalkwewouldlikelyencounteracolorfulmaleBridledForestGecko(Gonatodes humeralis;Fig.8)perchedonatreealongthetrail.Themorecrypticfemalesareslightlylessnoticeable.Cocha(Kentropyx altamazonica)andForest(K. pelviceps) Whiptailsarelarger,crypticlizards,butactive,soyouwillmostlikelynoticethemshufflingaroundinareasnearrecenttreefallswhenyoudisturbtheirbasking.Theextraordinarydiversityoflizardsisundoubtedlysupportedbythediversityofinverte-brates.Walkingevenashortsectionoftrailandnotobservingornatelycoloredgrasshoppers,walkingsticks,beetles,spiders,andbutterfliesisimpossible. Moresecretivethanthelizards,snakesarediversebutmoredifficulttoobserveintherainforest.Atleast50spe-ciesofsnakeshavebeendocumentedatReservaAmazónica.Walkingthetrailsystemyoumayfindfrog-eatingspecialistslikeBlack-headedCalicoSnakes (Oxyrhopus melanogenys;Fig.9)orWucherer’sGroundSnakes(Xenopholis scalaris),snail-eatersincludingseveralspeciesinthegenusDipsas,orlizard-eatingspecialistssuchasBlunt-headedTreesnakes(Imantodes cenchoa;Fig.10A).Additionally,severalboidsoccasionallycanbefoundonthetrails.TheRainbowBoa(Epicrates cen-chria) ismost frequently encountered.Several venomousspeciesofsnakes,includingtheSouthAmericanBushmaster

Figure 8.FemalesoftheBridledForestGecko(Gonatodes humeralis)aredrabbrownorgrayandblendwellwiththeirenvironment,whereasmales(picturedabove)aremuchmorebrightlycolored.

Temporary Wetlands – Enduring Interactions

Although observing numerous species is pos-sibleatanytimeofyear,byfarthebesttime

to observe amphibians is during the wet season(November–March). The evening after the firstheavyraineruptsinfrogcalls.Followingthesametrailthatwasdrythedaybeforeleadsnowtoatem-porary wetland flooding the trail. When closelyinspectingleavesandbranches,silhouettesoftree-frogsontheothersidesof leavesbecomevisible.Thesearemaleswehaveinterceptedduringtheirmigrationstothechorusesaroundthenewlyformedpond.Several speciesarepresentatone locationandallarecalling.ThesetypicallyincludeTarauacaSnoutedTreefrogs(Scarthyla goinorum),SanCarlosTreefrogs(Dendropsophus leali),AmazonSnoutedFrogs(Scinax ictericus), andthemostabundantofall,Koechlin’sTreefrogs(Dendropsophus koechlini).Thesoundisdeafening,soloudthathearinganotherpersontalkingtoyoufromevenafewfeetawayisdifficult.Toourdismay,gettingaclearrecordingofjustasinglespeciesatatimewasalmostimpossible.Bythenextevening, thewater level inthepondhasrisenevenhigherandfemaleshavearrived.Themaleslooklikebackpacksclingingtofemalesastheyseekalocationtolaytheireggs.Bythefollowingday,eggmassesbegintoappearunderleavesandfloatingonthesurfacealongthepondedges.Withinafewdays,tinytadpoleswillhatchandbeginthejourneytometamorphosis,whichtheywillcompleteafewweekslater.Almostasrapidlyasthefrogsarrivedsohasathrivingaquaticinvertebratecommunitythatincludessnails,waterscorpions,predaceousdivingbeetles,anddragonflies.Here,Sarahisconductingonecomponentofourresearch,assessingthediver-sityofaquaticinvertebratesintemporarywetlands,particularlythosethatpreyontadpolesandmayplayaroleinstructuringtheamphibiancommunity.

ORLOFSKEETAL.

259

IRCFREPTILES&AMPHIBIANS•19(4):254–262•DEC2012

(Lachesis muta),arenativetotheregion;however,encounterswithvenomousspeciesarerareatReservaAmazónica. TheAnacondaBoardWalk, a200-meterboardwalkelevated1mabovetheAguajalesswampsisnamedforananacondasightingduringitsconstructionandliesapproxi-mately1.5kmintothetrail system.Duringtheevening,thevegetationsurrounding theboardwalkresonateswithcacophonous calls of various anurans including Beireis’Treefrogs(Dendropsophus leucophyllatus; Fig.11),ParadoxicalFrogs(Pseudisparadoxa),Henle’sSnoutedTreefrogs(Scinax pedromedinae; Fig. 12), and Orinoco Lime Treefrogs(Sphaenorhynchus lacteus).Whilesearchingforfrogsintheleavesandbranchesoverhangingtheboardwalk,weoccasion-allystartledasleepingSlenderAnole(Anolisfuscoauratus),perchedprecariouslyattheveryedgeoftheleaf,preparedatthefirstmovementcausedbyapredatortoleaptosafetyonanotherbranch(Figure10B).Inornearthewater,wehaveobservedScorpionMudTurtles(Kinosternon scorpioides)andRoyalGroundSnakes(Liophis reginae),andnoticedtheeyeglareofSpectacledCaiman(Caiman crocodilus).

Research and ConservationInkaterraencouragesecologicalresearchoftheregionandtheInkaterraAssociation,anaffiliatedNGO,runsanadjacentresearchstationavailablefornumerousbiologicalstudiesofplantsandanimals.Thattheregionisahotspotforamphib-ian and reptilian diversity allows extensive tropical fieldexperimentsandnaturalhistoryresearch.Asapartnership,Inkaterraoftenfacilitatestheeducationofitsinterpreterson

Figure 9.AjuvenileBlack-headedCalicoSnake(Oxyrhopus melanogenys)isoneofseveralcoral-snakemimicsintherainforest.

Figure 10.SpeciesinthegenusImantodesprobablyhavethebestcan-tileveringabilitiesamongNeotropicalsnakes(Ray2012).Thisisaveryimportantadaptationgiventhatmanyofthesespeciesarelizard-eatingspe-cialists.A.ABluntheadedTreesnake(Imantodes cenchoa)searchesthetreesatnightforitsprey.B.SlenderAnoles(Anolis fuscoauratus)sleepatnightontheedgeofleaves.Thisdefenseservesthemwellastheselizardswillfalltotheforestflooriftheydetectvibrationsoftheirperches.Consequently,thesnakemustcantilevertothelizardwithoutdisturbingthebranchonwhichthelizardissleepinginordertoobtainameal.

ORLOFSKEETAL.

260

IRCFREPTILES&AMPHIBIANS•19(4):254–262•DEC2012

Antspatrolthetrailsbydayandnight.Careisneededtoavoidunintentionally intercepting their trails,

particularlyfortheLeafCutterAnts.TheabundanceofantsintheTambopataregionisgreaterthanthatofanyothertypeofinsectandincludesdiversitynumbersupwardof43speciesinasingletree(Wilson1987).Thisdiversityandabundanceofantsprovidesasignificantpreyresourceforamphibians,andhasresultedintheevolutionofalargenumberofant-eatingspecialists. Duringthedaytwocrypticallycoloredfrogscanbeheardcallingfromtheleaflitter.Althoughbothspecieshavevarieddiets,bothincludeamajorityofants.ThefirstistheThree-stripedRocketFrog(Allobates trilineatus),whichfrequentlycallsfromrootbuttresses,signalinghissuperiormatingqualitiesordefendinghisterritoryfromintruders.Althoughnotascolorfulassomeofitsrelatives,themoretoxicspeciesofPoisonFrogs,itshuesofgoldandcoppermakeitastrikingsightbeforeitretreatsintotheleaflitter.Thesecondspecies,thePale-stripedPoisonFrog(Ameerega hahneli), isslightlymorecolorful,withpaleyellowstripesonabrownish-blackbody,andlikewisebolderinitschoiceofcallinglocations.

Atnight,onecanobserveuptofivemicrohylidspe-ciesthatfollowanttrailsforeasymeals.Whenfollow-ingananttrailforevenashortdistance,oneislikelytoencounterAmazonSheepfrogs(Hamptophryne bolivi-ana) orTwo-coloredOvalFrogs(Elachistocleis bicolor,pictured) partaking of a “buffet.” The ant moundsthemselves represent a highly diverse microhabitat.Approachinganantmoundatnightofferedusaglimpseofotheruniquespeciesrarelyseenelsewhereinthefor-est,includinganothermicrohylid,theBrownEggFrog(Ctenophrynegeayi), andthepoison-frogmimicGold-stripedFrog(Leptodactyluslineatus).Tarantulaburrowsareanotheruniquemicrohabitat,perhapsonewhereyouwouldnotexpecttofindanamphibian.However,theDottedHummingFrog(Chiasmocleis ventrimaculata),thesmallestmicrohylidinthearea,canoccasionallybefoundinsidetheseburrows,whereitfeedsontheantsthatpreyonjuveniletarantulas.Thismutualisticrela-tionshipkeepsthesmallfrogssafefromtheadultfemaletarantulas,whooftenfeedonotherfrogs,asthefemaleapparently“knows”thatthesefrogshelpprotectherspi-derlingsfromants.

Ants, More Ants, and Frogs that Eat Ants

ORLOFSKEETAL.

261

IRCFREPTILES&AMPHIBIANS•19(4):254–262•DEC2012

regionalplantandanimaldiversitybycollaboratingwithvis-itingbiologists.Theinterpretersandstaffareeagertolearn,aswellastoteachus,aboutthelocalbiodiversity.Evenafterlonghoursalreadyintheforest,theywouldjoinusonoureveningexcursionslookingforfrogs,snakes,andmanyotherorganisms.Theywouldhelpustakephotographs,recordcalls,andcollectdata.Forseveralyearsnow,InkaterraandReservaAmazónicahavesupportedthefieldworkofSAOandRCJmakingitpossibleforustocontinueourresearchaswellasmentorthenextgenerationoftropicalherpetologists.Manystudentsofherpetologyhavebeguntheirtropicalresearchatthisverysite,atrendthatislikelytocontinuewellintothefuture.However,theexcellentaccommodationsandnaturalatmospherearejustasinvitingforthosejustlongingtowit-nessamazingbiodiversityandlearnaboutitsconservation.

AcknowledgmentsWethanktheInkaterrastaffforhelpinthefieldandassistingwithourstay.FundingfortraveltoReservaAmazónicawasprovidedtoRCJthroughaSocietyofSystematicBiologists’AwardforGraduateStudentResearch.

Literature CitedDavis,W.2009.Thebig trip:UltimatePeru.National Geographic Adventure

December2008/January2009:44.

Doan,T.M.andW.ArizábalA.2002.Microgeographicvariationinspeciescom-positionof theherpetofaunalcommunitiesofTambopataRegion,Peru.Biotropica34:101–117.

Duellman,W.E.2005.Cusco Amazónico: The Lives of Amphibians and Reptiles in an Amazonian Rainforest.CornellUniversityPress,Ithaca,NewYork.

Duellman,W.E.andJ.E.Koechlin.1991.TheReservaCuzcoAmazonico,Peru:Biologicalinvestigations,conservation,andecotourism. Occasional Papers of

Figure 11.Morethan40speciesoftreefrogsoccuratReservaAmazónica.Oneoftheflagshipspecies,Beireis’Treefrog(Dendropsophus leucophyllatus;a.“clown”phase,b.“giraffe”phase),breedsinseveralplacesbutiseasilyseenfromtheAnacondaBoardwalkduringtherainyseason.

Figure 12.Henle’sSnoutedTreefrog(Scinax pedromedinae) demonstrat-ingcrypsisonalichen-coveredtree.

ORLOFSKEETAL.

262

IRCFREPTILES&AMPHIBIANS•19(4):254–262•DEC2012

the Natural History Museum, The University of Kansas142:l–38.

Duellman,W.E.andL.Trueb.1989.TwonewtreefrogsoftheHyla parvicepsgroupfromtheAmazonBasininsouthernPeru.Herpetologica 45:1–10.

Ray,J.M.2012.Bridgingthegap:Interspecificdifferencesincantileveringabil-ityinaNeotropicalarborealsnakeassemblage.South American Journal of Herpetology7:35–40.

vonMay,R., J.M.Jacobs,R.D.Jennings,A.Catenazzi,andL.O.Rodríguez.2007.Anfibios de Los Amigos, Manu y Tambopata, Perú.AsociaciónparalaConservacióndelaCuencaAmazónica,Lima,Perú.

Wilson,E.O.1987.ThearborealantfaunaofPeruvianAmazonforests:Afirstassessment.Biotropica 19:245–251.