Vol. 1 No. 1 1990 Rondonia butterflies: EMMEL and AUSTIN 1

TROPICAL LEPIDOPTERA, 1(1): 1-12

THE TROPICAL RAIN FORESTBUTTERFLY FAUNA OF RONDONIA, BRAZIL

SPECIES DIVERSITY AND CONSERVATION

THOMAS C. EMMEL and GEORGE T. AUSTINDepartment of Zoology, University of Florida, Gainesville, FL 32611, and

Nevada State Museum and Historical Society, 700 Twin Lakes Drive,Las Vegas, NV 89107, USA

ABSTRACT.— The state of Rondonia in west central Brazil apparently has the highest reported butterfly diversity in the world, with an estimated1,500-1,600 species living within several square kilometers in the central part of that state. A preliminary checklist of over 800 identified speciesis given, and some of the factors contributing to this diversity are described. The tropical rain forest in this area is being rapidly cleared fordevelopment and the creation of one or more inviolate biological preserves is urgently needed in order to save a living sample of the incrediblydiverse fauna and flora for study by future generations.

KEY WORDS: Amazon Basin, butterfly faunas, Hesperiidae, Lycaenidae, Nymphalidae, Papilionidae, Pieridae, rain forest, Riodinidae.

Rondonia, one of the newest states in western central Brazil,occupies some 93,840 square miles (243,044 sq km) in thesouthwestern part of the Amazon Basin of South America. Thisterritory, which borders Bolivia to the south and west, wasformerly a part of the state of Amazonas and until the last twodecades, was primarily important to Brazil's economy only duringthe Amazon rubber boom, which collapsed in 1912. In 1943, thearea was established as Guapore. The territory was renamedRondonia in 1956, after Marshal Camdido Mariano da SilvaRondon, Brazil's famous explorer, and was made a state in 1981.

The rain forest here was sparsely inhabited by Indians, rubbertappers, and former railroad workers until the then-militaryregime of the central Brazilian government began pushing thedevelopment of the Amazon in the 1960's. Thus in 1966,

Fig. 1. Map of the Rondonia area of Brazil, with the inset showing major townsand cities of Rondonia mentioned in the text.

Fig. 2. Aerial view of the Rondonia rain forest (near Porto Velho), showinghighway BR364 across the lower right corner and nearby pastures. (© T. C.Emmel; this photograph and all following photographs)

EMMEL and AUSTIN: Rondonia butterflies TROPICAL LEPIDOPTERA

Fig. 3. Interior view of the lush and diverse lowland tropical rajin forest atFazcnda Rancho Grande, near Caucalandia, Rondonia.

Fig. 4. Nessaea obrinus (Nymphalidae) on rotting papaya fruit.Fig. 5. Morpho achilles (Nymphalidae) sunning itself momentarily on a moss-

covered palm frond.

Fig. 6. Prepona sp. (Nymphalidae) feeding at fermenting sap as it oozes froma wound in tree bark.

SUDAM, the Superintcndcncy for the Development of theAmazon, was created as an agency to oversee the plannedoccupation of the Amazon forest with "civilization." Lucrativeincentives were offered for industrial and agricultural projects,particularly the development of cattle ranches to supply cheapbeef for the Bra/ilian people (relatively little of this beef was toever be exported to the American fast-food industry, unlike thesituation in Central America). Great amounts of valuable timberwere cleared during those years and merely left to be burnedduring the annual dry season, in order to produce large amountsof pasture. Millions of acres of public land in the eastern andnorthern Amazon were occupied along the new highways beingput in, especially along the Bclcm-Brasilia Highway. By 1970most of the settlers had abandoned their developments afterdiscovering that the underlying soils were too poor to sustainagriculture (Revkin, 1990). Wide variations in Amazonian soilquality have consistently defeated attempted settlement. Grassesquickly use up the essential minerals left behind after burning ofthe native forest vegetation (whose biomass formerly held mostof the minerals and other nutrients), and inedible weed speciesand brush then invaded the fields incessantly. Thus pasture thatonce supported one head of cattle for every 2.5 acres (a relativelypoor ratio to begin with) dropped to supporting one head of cattlefor every 10 acres, and the maintenance costs and large land areasrequired for livestock made it unworthwhilc to continue ranching.

The federal government's other major plan to attract people tothe Amazon (aside from encouraging private development) wasto build a vast network of roads. In order to move people awayfrom the overpopulated lands in the southeastern Amazon, thegovernment completed the extension of federal highway BR-364from the capital of Mato Grosso (Cuiaba) to Porto Velho, thecapital of the new state of Rondonia. Channeling immigrants intothe state, this road extension, though not paved completely until1984, created most of the destruction in Rondonia throughout the1970's as the population of the area grew at a tremendous pace(Ellis, 1988). In those ten years, there was an explosion of newarrivals coming from the southwest. Between 1970 and 1985, thepopulation of Rondonia increased from less than 100,000 to more

Vol. 1 No. 1 1990 Rondonia butterflies: EMMEL and AUSTIN 3

Fig. 9

Hamadryas sp. (Nymphalidae) feeding at tree sap.Anaea (ZaretLs) itys (Nymphalidae) drinking human perspiration lhat had

dripped onto dead leaves in the forest floor litter.Heliconius doris (blue and red forms) and other Ileliconius (JNymphalidae)

drinking at a urine-soaked sand bank.

Fig. 10. Panacea divalis (Nymphalidae) attracted to sweat on a net handle.Fig. 11. Diaethria nr. negtecta (Nymphalidae) perched on a leaf.Fig. 12. Callicore sp. (Nymphalidae) feeding at fermenting tree sap.

EMMEL and AUSTIN: Rondonia butterflies TROPICAL LEPIDOPTERA

Fig. 13. Caligo brasiliensis (Nymphalidae: Brassolinae) feeding at fermentingtree sap.

than 730,000 people. Side roads were built and families weregiven tides to 250-acre lots of rain forest as long as they clearedhalf of their plot of land for agriculture. More than 70,000square miles of Amazonian forest were being burned annually,and a sizeable proportion was located in the state of Rondonia.As the settlers cleared and farmed the rolling hills of this area,the little soil on those hillsides quickly washed away. In order to

continue fanning or obtain useful land for cattle, the settlers hadto clear new tracts of land. By 1987, American weather satellitepictures of Rondonia showed Brazilian ecologists that more than100,000 fires were burning at once in the state during the dryseason, and that the rain forest was being cleared at a rate farbeyond what anyone had suspected. An urgent worldwideinterest immediately developed (e.g., Linden, 1989) in preservingthis and other rain forests of the Amazon Basin because of thealready alarming extent of global warming being increased by therelease of large amounts of CO2 from the burning of these forests,the role of the forest in the natural oxygen cycle and hydroliccycles of the western hemisphere itself, and the fact that self-sustainable uses could be made of the rain forests that would bemore profitable to Brazil than the past policy of clearing theforest and planting grass for cattle.

Fig. 14. Hypolhyris euclea (Nymphalidae: Ithomiinae) drinking pyrrolizidinealkaloids from the stems of a decaying Heliolropium plant (Boraginaceae).

Fig. 15. A diurnal Urania moth (L'raniidae) drinking water at a river bank.

During the last several years, our joint interests in tropicalbutterfly diversity have centered on this area of Brazil. Thesenior author has led expeditions of groups of Icpidopterists tothese extraordinarily productive tropical rain forests in March1987, March 1989, and November 1989; the junior author hasaccompanied him on the latter two field trips, with other jointvisits planned for November and December 1990, and March1991. Our sampling in these areas has confirmed that centralRondonia may be the most diverse area in the world for butter-flies. Because of the accelerated human development of thisgeneral region, it is probable that there will be virtually nothingleft of the original rain forest habitat for these butterflies by earlyin the next century — only a decade away — unless one or morelarge preserves are set aside now.

SAMPLING AREA IN RONDONIA

We have concentrated our sampling efforts in the area south ofAriquemes, a town located 199km south of Porto Velho. Some62km south of Ariquemes lies the small village of Caucalandia,and near there is the Fazcnda Rancho Grande, a 750-ha tractowned by the Harald Schmitz family who have been extremelykind in providing accommodations and other logistical support forour surveys. Fazenda Rancho Grande lies 22km west of highwayBR 364, or about 5km west off road B-65 on cross-road C-20.

Vol. 1 No, 1 1990 Rondonia butterflies: EMMEL and AUSTIN

Fig. 16. Euptychia sp. (Nymphalidae) perched on a rain forest leaf.

The area has been sampled for about 10km in all directionsaround the Fazenda, which lies at latitude 10° 32'S and longitude62° 48'W. The terrain is typically low rolling hills or flat plainscovered with wet tropical rain forest, except in areas of humandisturbance.

The general elevation in the area ranges from 160-350m; theFazenda Rancho Grande ranch house is 187m (540ft) above sealevel. East-west "C"-numbered roads (e.g., C-20) have been

Fig. 17. Consul fabius (Nymphalidae) perched on a rain forest leaf near cowmanure, which attracted this butterfly to the ground to feed.

constructed every five kilometers running south from Ariquemes;there arc also north-south "B"-numbered roads (e.g., B-65) every20km that cut perpendicularly across these east-west roads. Thishas created rectangular blocks whose edges arc largely cleared byhomesteaders and ranchers, but whose interiors (perhaps 3.5 x 18or more km) are still largely uncut forest in most places. Thegeneral region is located about 750km from the nearest highmountains to the west. The forest has a relatively open canopyin places, including many palms and bamboos, with some densehill and alluvial forests.

Rainfall and temperature records at Fazenda Rancho Grandehave been kept since 1984 by Harald Schmitz, and he has kindlyshared these data with us. The annual dry season here extendsfrom May to September. At its severe peak, in June, July andAugust, monthly rainfall drops to 10-20mm, and no rain at allmay fall for one or two months annually. At this time the forestdries out enough to allow large-scale burning. The wet seasonstarts in October and continues to late April. Annual rainfallaverages around 2200mm, with the rainiest months being inJanuary (the annual peak, with often around 440mm), February,and March. The recorded temperature extremes range from 9°C(11 Jun 1985) to 43°C (27 Sep and 5 Oct 1987), but normally thetemperatures are more moderate. During the dry season in 1987,for example, the average minimum was 18.7°C and the averagemaximum was 34.1°C in August (coolest month of the year).During the wet season of that same year, the warmest tempera-tures were reached in October (24.9°C average minimum and38.9°C average maximum). During the rainy season, there aremany strongly flowing streams in the area and standing pondsalong roadsides and in depressions in the hilly terrain.

PRELIMINARY LIST OF THE RONDONIABUTTERFLY FAUNA

In visits during 1975 and 1977, Keith S. Brown (1976, 1984)estimated that the species diversity in the area near Jaru (latitude10°26'S, longitude 62°27'W), about 50km to the east of theFazenda Rancho Grande, supported a butterfly fauna of 1,100-1,300 species. In those years, he accumulated normal dailyspecies lists of over 300 species, and had maximum daily lists ofup to 429 species (by one person on 5 Oct 1975) and of 462species (by two persons on 2 Oct 1975). Today, the Jaru area iscleared for agriculture and no sustantial tracts of rain forestremain there. We will never know the total butterfly faunaoriginally present in that rich Jaru site.

In the area near the Fazenda Rancho Grande, vicinity ofCaucalandia, we have now accumulated a list of over 840 species.These are only the species that have been identified at least togenus so far. The fact that we are now at 200 species ofmetalmarks (Riodinidae) alone suggests from comparative faunalsurveys elsewhere in the Neotropics (e.g., 193 riodinids atTambopata, Peru) that the grand total number of species will bein the vicinity of 1,500-1,600 species of butterflies (includingskippers) when the survey is completed. In the meantime, weoffer this preliminary list.

All determinations are very tentative due to time constraintsand the fact that taxonomic revisions are needed among many of

EMMEL and AUSTIN: Rondonia butterflies TROPICAL LEPIDOPTERA

Fig. 18-21. Rondonia Riodinidae: 18. Male Amarynthis meneria sunning and perching on a forest leaf (upper left); 19. Caria sp. drinking animal urine on a foresttrail (lower left); 20. Lasaia sp. drinking animal urine on a forest trail (upper right); 21. Rhetus sp. drinking human urine on a trail (lower right).

the species groups involved. Those names of the Papilionidae,Pieridae, and Nymphalidae arc mostly after D'Abrera (1981;1984; 1987a, b; 1986), those of Hesperiidae are after Evans(1951, 1952, 1953, 1955). The Lycaenidae were determined byRobert K. Robbins (March 1989 sample) and the Riodinidae weredetermined by Donald Harvey (March 1989 sample); the remain-der of the names are mostly from Lewis (1973). Morpho specieswere determined by George E. Martinez. Several riodinids werealso identified by Curtis J. Callaghan from color photographstaken by George O. Krizek. Thus while this list is provisional innature, it will still offer a preliminary idea of the species diversityfound in this tropical rain forest site. It should not be used as asource of data for distributional or taxonomic studies, of particularspecies. Most specimens on which these preliminary determina-tions were made are housed at the University of Florida (Gaines-ville), the Nevada Slate Museum (Las Vegas), or the AllynMuseum of Entomology (Sarasota, Florida) for further study.

To summarize these data by family and subfamily, we have thefollowing species diversity present in the initially identifiedsamples of butterflies from this region of Rondonia:

HESPERIIDAE 231Pyrrhopyginae 3Pyrginae 135Hesperiinae 93

PAPILION1DAE 18PIERIDAE 29RIODINIDAE 203LYCAENIDAE 87

Polyommatinae 2Theclinae 85

NYMPHALIDAE 275Nymphalinae 126Satyrinae 56Bras sol inae 8Morphinae 11Heliconiinae 25Danainae 3Ithomiinae 46

TOTAL species: 843

Vol. 1 No. 1 1990 Rondonia butterflies: EMMEL and AUSTIN 7

HESPERIIDAEHESPERIINAE

Aides aegitaAides epitusAnthoptus epictetusAnthoptus insignisApaustus menesAroma aromaArotis sp. (nr. bryna)Callimormus coradesCallimormus juveniusCallimormus radiola radiolaCallimormus saturnusCarystoides caihaeaCarystoides nosedaCarystoides sicania orbiusCarystus phorcusCobalopsis neroCobalus calvinaCobalus virbius virblusConga chydaeaCorticea corticea noct'tsCorUcea lysias potexCymaenes alumnaCymaenes chela (?)Cymaenes lazaCymaenes uruba groupCymaenes sp. (nr. urubaltabori]Cymaenes sp.Cynea bistrigulaCynea corisanaDalla diraspesDamns clavusDecinea luciferDubiella dubiusEnosis angular is angularisEuphyes peneiaMellana angraMellana barbaraMellana clavusMethionopsis sp.Mnasilus allubitusMnasitheus chrysophrysMacros moerosMorys compta comptaMorys geisa geisaNeoxeniades braesia braesiaNeoxeniades cinciaNiconiades xanthaphesPanoquina sylvicolaPaplas proxlmusPapias subcostulataPapias sp.Paracarystus menestriesParphorus storax storaxPeba striataPerichares philetes philetesPhanes aletesPhanes almodaPhlebodes pertinaxPhlebodes sp.

Pompeius pompelusQuint a cannaeSaliana antoninusSaliana esperiSaturnus tiberlus tiberiusSodalia sodalisStyriodes quadrinotataSynapte infuscoSynapte stilusTalides slnois sinoisThargella caura cauraThespieus dalmanThracides quartaThracides smaragdulusTigasis akurusTuresis basiaTuresis lucasVacerra bonfilius bonfiliusVehilius incaVehilius putusVehilius retusVehilius stictomenes stictomenesVehilius vetulaVehilius vetusVenas evansVettius marcusVettius phyllus phyllusVettius richardi (?)Vettius triangularisVirga virginiusWallengrenia premnasZariaspes mysUnidentified species (3)

PYRGINAE

Ablepsis azinesAchlyodes busirus riojaAchlyodes thrasoAethilla echinaAguna asander asanderAguna claxonAguna coelusAguna hypozoniusAguna metophisAnastrus obscurus narvaAnastrus sempiternus simplidorAnastrus tolimus robigusAnisochoria pedaliodina pedaliodinaAntigonus erosusAntigonus liborius liboriusAntigonus nearchusAstraptes alardus alardusAstraptes anaphus anaphusAstraptes apaslus apastusAstraptes azulAstraptes enotrusAstraptes fulgeratorAutochton longipennisAutochton neisAutochton zarexBolla atahuallpaiBolla cuprelceps

Bungalotis astylosCabirus procas juntaCamptopleura auxoCamptopleura impressusCarrhenes bambaCarrhenes canescens leadaCelaenorrhinus astrigeraCelaenorrhinus jaoCelaenorrhinus shema ochraCelaenorrhinus similis groupCelaenorrhinus sp.Chioides catillus catillusChrysoplectrum pervivaxCogia calchasConognathus platonCycloglypha thrasibulus thrasibulusCycloglypha tisiasDiaeus lacaena variegataDyscophellus euribates euribatesEbrietas evanidusEbrietas infandaEntheus gentiusEntheus pralinaEntheus telemusEpargyreus sp.Eracon paulinusGorgopas trochilusGorgythion begga beggaGorgythion beggina escalophoidesGorgythion plautiaGrais stigmaticus stigmaticusHelias phalaenoides palpalisHeliopetes alanaHeliopetes arsalte arsalteHeliopetes omrinaHeliopetes pelrusflyalothyrus infernalisHyalothyrus leucomelasHyalothyrus neleus neleusMarela tamyrisMarela tamyroidesMictris crispus crispusMilanion hemes hemesMilanion pilumnusMorvina fissimacula remaMorvina morvus cyclopaMylon cajus cajusMy Ion exstincta (?)Myrinia santa monkaNarcosius colossus granadensisNarcosius samsonNisoniades bessus bessusNisoniades borraNisoniades braziaNisoniades castolusNisoniades macariusNisoniades mauraNisoniades mimasOuleus fridericus frider icusOuleus matria matriaPaches exosaParamimus scurra herbertiPellicia meno

8 EMMEL and AUSTIN: Rondonia butterflies TROPICAL LEPIDOPTERA

Pellicia nemaPhanus marshalliiPhanus obscurior prestoniPhareas coelestePlumbago plumbagoPolyctor polyctor polyctorPolygonus manueli manueliPolythrix minvanesPolythrix octomaculataPyrdalus corbulo corbuloPyrgus oileusPythonides grandis assedaPythorddes jovianus fabriciiQuadrus contubernalis contubernalisQuadrus deyrolleiSostrata pusilla pusillaSpathilepla cloniusSpiomades artemidesStaphylus epicaste epicasteStaphylus lizeriStaphylus minorStaphylus putumayo pulumayoTelemiades antiope ioscaTelemlades delalandeTimochreon satyrus satyrusTrina geometrina geometrinaTyphedanus optica goyaTyphedanus undulatusUrbanus albimargo rlcaUrbanus carmelitaUrbanus chalcoUrbanus dorantes dorantesUrbanus doryssus doryssusUrbanus estaUrbanus prontaUrbanus proteusUrbanus reductusUrbanus simpliciusUrbanus lannaUrbanus teleusUrbanus vtrescensXenophanes tryxusUnidentified species (3)

PYRRHOPYGINAE

Aspitha aspitha rufescensOxynetra confusaPyrrhopyge phidlas group

RIODINIDAE

Adelotypa nr. alectorAdelotypa hemileucaAdelotypa nr. huebneriAdelotypa spp. (2)Adelotypa aminias cateniferaAdelotypa aristus (?)Adelotypa leucocyanaAdelotypa leucophaeaAdelotypa sentaAlesa amesis

Alesa nr. lelephaeAmarynthis meneriaAncyluris colubraAnteros nr. bracteataAnteros nr. carausiusAnteros renaldusArgyrogrammana nr. peroneAudre albimaculataCalephelis argyrodinesCalosplla nr. emylianaCalosplla emylius crispinellaCalosplla parthaonCalosplla sp.Calydna catanaCalydna chasebaCalydna thersanderCalydna sp.Caria mantineaCaria nr. chrysameCaria nr. trochilusChalodeta nr. lyperaChamaelimnas sp. (pansa or tircis)Charts nr. aniusCharts jessaCharts sp.Chans caryatisCharts cleonusCharts zamaChorinea amazonChorinea sp. (faunus or batesi)Cremna actorisCremna nr. beltianaCremna meleagrisCrocozona caeciasEmesis nr. emesiaEmesis lucindaEmesis mandanaEmesis ocyporeEmesis temesaEmesis spp. (3)Eunogyra satyrusEurybia nr. dardusEurybia nr. elvinaEurybia nr. franciscanaEurybia halimedeEurybia hyacinthinaEurybia nicaeusEuselasia nr. arbasEuselasia nr. brevicaudaEuselasia calligrammaEuselasia chrysippeEuselasia clithraEuselasia nr. crotopusEuselasia eubagesEuselasia eugeon (?)Euselasia nr. eulioneEuselasia eunaeusEuselasia euodiasEuselasia euryoneEuselasia nr. eustachisEuselasia eutychusEuselasia labdacusEuselasia melaphaea

Euselasia orfitaEuselasia tarintaEuselasia toppiniEuselasia nr. uriaEuselasia spp. (5)Exoplisa cadmeisHyphilaria niciaHyphilaria parthenisIthomeis astreahhomiola nr. floralisJuditha lamisJuditha molpeLasaia agesilas agesilasLasaia arsisLemonias zygiaLepricornis spp. (2)Leucochimona hyphaeaLeucochimona matiscaMelanis nr. andaniaMelanis seleutiaMelanis smithiaeMelanis nr. stenotaeniaMesene epaphus sertataMesene hyaMesene nr. nolaMesene phareusMesophthalma idoteaMesosemia nr. ancepsMesosemia nr. calypsoMesosemia cippusMesosemia eumeneMesosemia furiaMesosemia judicialisMesosemia nr. levisMesosemia nr. maeotisMesosemia marcellaMesosemia melaeneMesosemia nr. minosMesosemia nr. nestiMesosemia olivenciaMesosemia philoclesMesosemia sifia (?)Mesosemia nr. tenebricosusMesosemia spp. (2)Metacharis luciusMetacharis regalisMonethe sp. (albertus or rudolphus)Napaea neposNotheme erotaNymphidium nr. acheroisNymphidium baoetiaNymphidium caricaeNymphidium chioneNymphidium fulminansNymphidium galactinaNymphidium leucosia (?)Nymphidium nr. lisimonNymphidium manicorensisNymphidium mantusNymphidium minutaNymphidium niveumNymphidium nr. baoetiaNymphidium olinda

Vol. 1 No. 1 1990 Rondonia butterflies: EMMEL and AUSTIN

Nymphidium omois (?)NympkidiumlJuditha spp. (5)Nymula gelaNymula ochraOrimba butleriOrimba flammulaOrimba pythionidesOrimba sp.Panara p her eel usParcella amarynthinaParties nycteisParties philotesPhaenochitonia eanesPhaenochltonia sophistesRhetus arciusRhetus dysoniiSarota gyasSarota chrysusSemomesia nr. capaneaSemomesia macariaSemomesia semiairaStalachtis calliopeStalachtis phlegiaSymmachia rubinaSymmachia nr. praxilaSymmachia sp.Synargis orestesSynargis tytiaSynargis sp.Syrmatia dorilasTharops trotschiThemone paisTheope foliorumTheope pediasTheope thestiasTheope sp.This be ireneaThisbe molelaThysanota galenaUnknown species (15)

LYCAENIDAEPOLYOMMATINAE

Hemiargus sp.Leptotes cassius

THECLINAE

Arawacus separataAreas imperialisCalycopis spp. (2)Cyanophrys spp. (2)Cycnus phalerosEvenus gabrielalaspis sp.Lamprospilus geniusMinistrymon phrutusMithras nautesOcaria ocrisiaPanthiades bitias bitiasPanthlades pelion

Pseudolycaena marsyasRekoa melonRekoa palegonStrymon cestriStrymon muluchaThecla celmusThecla cupentusThecla halcionesThecla hisbonThecla malvinaThecla pholeusThecla terentiaThecla spp., etc. (53)Theclopsis lydusTheclopsis nr. epidiusTheritas hemonTheritas mavorsTmolus mutina

NYMPHALIDAEBRASSOLINAE

Caligo brasiliensisCaligo eurilochusCaligo idomeneusCaligo illioneusCatoblepia berecynthiaCatoblepia soranusOpsiphanes invirae (?)Selenophanes cassiope

DANAINAE

Ituna ilioneLycorea cleobaeaLycorea paslnuntia pasinuntia

HELICONIINAE

Agraulis vanillaeDione junoDryadula phaetusaDryas JuliaEueides alipheraEueides IsabellaEueides lampetoEueides lybiaEueides tales pythagorasEueides vibiliaHeliconius aoedeHeliconius burneyiHeliconius demeterHeliconius doris (2 forms)Heliconius elevatusHeliconius eratoHeliconius ethillaHeliconius hecaleHeliconius leucadiaHeliconius melpomeneHeliconius metharmeHeliconius numata (3 forms)Heliconius sara

Heliconius wallaceiPhilaethria dido

ITHOMIINAE

Aeria elaraAeria euritnediaAthyrtis tnechanitis salviniCallithomia leneaCeratinia poecilaCerat'mia sp.Dircenna deroForbestra equicola equicoloidesGodyris zavaleta (?)Heterosais giuliaHypoleria oreasHypoleria mirzaHypoleria proximoHypothyris daphnis daphnisHypothyris euclea bariiHypothyris fluonia (?)Hypothyris leprieuri ninyasHypothyris mamercusHypothyris ninonia neimyiHypothyris semifulva (?)Ithomia agnosiaMechanitis egaensisMechanitis lysimniaMechanitis mazaeusMechanitis polymniaMelinaea maenius maeniusMelinaea marsaeus marsaeusMelinaea menophilusMethona grandiorMethona megistoNapeogenes aethraNapeogenes inachiaNapeogenes pheranthesNapeogenes Stella jamariensisNapeogenes sylphisOleria aquataOleria astraea (?)Oleria nr. assimilisOleria egraOleria nr. manoraPseudoscada genityllusRhodussa cantobricaScada reckiaScada theaphiaThyridia psidiiTithorea harmonia

MORPHINAE

Antirrhea murenaCaerois chorinaeusMorpho achilles (3 forms: Helena,

amazonica, patrochus}Morpho adonisMorpho anaxibiaMorpho dsseis phanodemusMorpho deidamiaMorpho helenor helenor

10 EMMEL and AUSTIN: Rondonia butterflies TROPICAL LEPIDOPTERA

*

Morpho telemachusMorpho menelaus (3 forms: matto-

grossensis, occidentaits,melacheilus

Morpho rhetenor

NYMPHALINAE

Adelpha aethaliaAdelpha boeotiaAdelpha cocalaAdelpha cythereaAdelpha delphicolaAdelpha epioneAdelpha erotiaAdelpha fufia (?)Adelpha fugelaAdelpha iphiclusAdelpha mesentinaAdelpha paraenaAdelpha phylacaAdelpha plesaureAdelpha pollina (?)Adelpha serpaAdelpha sichaeusAdelpha thesprotiaAgrias claudina sardanapalusAnarlia amatheaAntigonis pharsalia felderiArchaeoprepona amphimachus

amphimachusArchaeoprepona demophon

demophonAsterope markiiBaeotus amazonicusBatesia hypochloraBiblis hyperiaCalllcore astarteCallicore cylleneCallicore cynosuraCallicore eunomia (2 forms)Callicore hesperisCallicore hystaspesCastilia angustaCatonephele acontiusCatonephele antinoeCatonephele salaciaColobura dirceConsul fabiusDagonfontus (?)Diaethria clymenaDiaethria nr. neglectaDoxocopa agathina agathinaDynamine agacles coreDynomine areneDynamine artemisiaDynamine athemon ruveataDynamine chryseisDynamine coenus leucotheaDynamine ines inesDynamine mylittaDynamine myrsonDynamine racidula

Dynamine theseusEctima theclaEresia claraEresia eunice (2 forms)Eresia plagiataEunica alpais alpaisEunica concordiaEunica eurota eurotaEunica malvinaEunica margaritaEunica orphiseEunica sophonisbaEunica sydoniaEuptoiela hegisaFountainea ryphea rypheaHaematera pyramus pyramusHamadryas amphinome amphinomeHamadryas arinome arinomeHamadryas chloe chloeHamadryas februa februaHamadryas feronia feroniaHamadryas iphthime iphthimeHamadryas laodamia laodamiaHamadryas velutinaHislorius odius orionHypanartia letheHypna clytemnestra clytemnestraMarpesia beraniaMarpesia chironMarpesia crethonMarpesia eginaMarpesia hermioneMarpesia liviusMarpesia noricaMarpesia orsilochusMarpesia petreusMemphis basiliaMemphis glauceMemphis glauconeMemphis lemnos (?)Memphis leonidaMemphis morvusMemphis oenomaisMemphis philumenaMemphis viciniaMemphis xenocles (?)Memphis sp.Myscelia capenasNapeocles jucundaNessaea obrinusNica flavillaOrtUia gentinaPanacea divalisPaulogramma peristeraPeria lamisPrecis evaretePrepona gnorimaPrepona pheridamusPrepona rothschildiPyrrhogyra amphiroPyrrhogyra crameri nautacaPyrrhogyra otalaisSmyrna blomfildia

Tegosa fragilisTegosa infrequensTegosa similisTelenassa burchelUTemenis laothoeTemenis pulchraTigridia acestaVictorina stelenesVila semistalachlisZaretis itys

SATYRINAE

Bia actorionCithaerias nr. aurorina"Euptychia" anal'ts"Euptychia" antonoe"Euptychia" arnaea"Euptychia" ay ay a"Euptychia" balesii"Euptychia" cephus"Euptychia" confusa"Euptychia" nr. cyanites"Euptychia" erichtho"Euptychia" nr. gracilis"Euptychia" nr. helle"Euptychia" hermes group spp. (2)"Euptychia" herse"Euptychia" hesionides"Euptychia" hewitsoni"Euptychia" nr. lea"Euptychia" libye"Euptychia" libyoidea"Euptychia" nr. myncea"Euptychia" nortia"Euptychia" obscura"Euptychia" ocirrhoe"Euptychia" nr. ocypete"Euptychia" nr. pallerna"Euptychia" penelope"Euptychia" perfuscata"Euptychia" philippa"Euptychia" nr. picea"Euptychia" quadrina"Euptychia" nr. renata"Euptychia" nr. salvini"Euptychia" nr. tenera"Euptychia" terrestris"Euptychia" nr. terrestris"Euptychia" nr. thobiei (?)"Euptychia" lolumnia"Euptychia" tricolor"Euptychia" nr. umbrosa"Euptychia" sp.Haetera pieraPierella astyoche (?)Pierella hyalinusPierella lamiaPierella lenaTaygetis andromedaTaygetis blandaTaygetis celiaTaygetis echo

Vol. 1 No. 1 1990 Rondonia butterflies: EMMEL and AUSTIN 11

Taygetis mermeriaTaygetis peneleaTaygetis virgiliaTaygetis vraziTaygetis xenana

PAPILIONIDAE

Battus belusBattus crassusEurytides ariarathesEurytides dolicaonEurytides pausaniasPapilio anchisiadesPapilio androgeusPapilio garleppiPapilio lycophronPapilio thoasPapilio torquatusParides aeneasParides neophilus consusParides phosphorusParides sesostrisParides steinbachi (?)Parides vertumnusParides zacyrUhus polymetus

PIERIDAE

ArUeos menippeAphrissa statiraAppias drusillaAscia buniae phaloeAscia buniae pharetiaDaptoneura leucadiaDaptoneura lycimniaDismorphia pinthaeus proximaDismorphia theucarilaEnantia liciniaEnaniia meliteEurema agaveEurema albulaEurema deva pseudomorphaEurema elatheaEurema leuceEurema phiale (?)Eurema venustaItaballia demophileItaballia pisonisLeptophobia aripa aripaLeucidia brephosLeucidia elvinaPerrhybris pyrrhaPhoebis argante organicPhoebis phileaPhoebis sennae marcellinaPhoebis tritePseudopieris nehemia (?)

DISCUSSION

Until our recent work near Caucalandia in Rondonia, theworld's richest sites for butterfly diversity were believed to be atJaru in the same Brazilian state (Brown 1984) and at TambopataNature Reserve in the department of Madre de Dios, Peru(Lamas, 1981, 1983). Brown (1984) actually recorded 956species at Jaru, Rondonia, and predicted 1,332 species to haveoccurred there at the time of his studies (prior to deforestation).Lamas (1983, and in Brown 1984) actually recorded 1,032species at Tambopata, Peru, and Lamas predicted 1,624 specieswould eventually be found there. Brown (1984) considered thelatter Tambopata estimate to be rather high. Both Jaru andTambopata have complex microheterogeneity of the physicalenvironment, with diverse soil types and frequent, unpredictableclimatic disturbance by thunderstorms, winds, variably severe dryperiods, and cold fronts that are fatal to many tropical butterflies.Man has invaded both areas with trails, small clearings, and treeremoval by logging (and massive deforestation now in Rondonia).Both climatic and human disturbance maintain the alreadyheterogeneous ecosystems of Tambopata and central Rondonia ina constant state of preclimax vegetation mixed with climax forest,with many secondary succession patches, as pointed out byBrown (1984). These conditions create an ideal mix of moremature and stable forest patches, with the diverse adult and larvalresources and differing microclimates of the more open second-ary-succession areas, just as still exists near Caucalandia today.

Tambopata and Rancho Grande may be approximately equal ineventually-determined levels of butterfly diversity (close to 1,600species in several square kilometers), but the mixture of ecologi-cal and historical causes of species diversity is likely somewhatdifferent at the two sites. Tambopata, for example, is undoubted-ly influenced by the proximity of the Andes whose upliftedmontane habitats start only 35 km to the west, and the potentialfor extensive movement of various sun-loving butterfly popula-tions (especially Papilionidae and Pieridae) along the largerivercourses there, flying in from neighboring cleared andsecondary-succession areas. These and other factors influencingspecies diversity will be addressed in detail in a later paper(Emmel and Austin, in preparation).

CONCLUSIONS

Surveys at Jaru and near Caucalandia in Rondonia, Brazil,show that this central Rondonian area may have the highestbutterfly diversity in the world. In the limited field work to date,over 800 butterfly species at Fazenda Rancho Grande nearCaucalandia have been identified, and that number is expected toreach approximately 1,500-1,600 species with further researchthere. From informal surveys to date, diversity among the birds,mammals, plants, and other organisms seems equally high.Because of the accelerated pace of deforestation and settlementin Rondonia, it seems essential to create as soon as possible oneor more inviolate biological preserves of considerable size to savethis fauna and flora for future generations to visit, study, andenjoy.

12 EMMEL and AUSTIN: Rondonia butterflies TROPICAL LEPIDOPTERA

ACKNOWLEDGMENTS

We thank the Harald Schmitz family at Fazcnda RanchoGrande, Rondonia, for their outstanding hospitality and theirprovision of every possible logistical help in carrying out ourfield surveys in Rondonia. We also wish to express our apprecia-tion to Paul and Sandra Russell (Malibu, California), George E.Martinez (Miami, Florida), and Jim P. Brock (Tucson, Arizona)for contributing their records to this initial survey list. Determi-nations were kindly provided in part by Donald E. Harvey andRobert K. Robbins (Smithsonian Institution), Curtis J. Callaghan(Spain and Brazil), and Steven Steinhauser and Lee D. Miller(Allyn Museum of Entomology, Sarasota, Florida). We also wishto thank Olaf H. H. Mielke (Univcrsidadc Federal do Parana,Curitiba) for assistance in obtaining permits and in first introduc-ing us in March 1987 to the state of Rondonia and its richbutterfly fauna.

LITERATURE CITED

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News Lepid. Soc. (Lawrence), 1984(3): 45-47.D'Abrcra, B.

1981. Butterflies of the Neotropical Region. Parti . Papilionidae &Pieridac. East Melbourne, Aust.: Lansdowne. 172pp.

1984. Butterflies of the Neotropical Region. Part II. Danaidae,Ithomiidae, Heliconidae & Morphidae. Victoria, Aust.: HillHouse. Pp. 174-384.

1987a. Butterflies of the Neotropical Region. Part in. Brassolidae,Acraeidae, & Nymphalidae (partim). Victoria, Aust.: HillHouse. Pp. 386-525.

1987b. Butterflies of the Neotropical Region. Part IV. Nymphalidae(partim). Victoria, Aust.: Hill House. Pp. 528-678.

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