Description of a new and highly distinctive genus and species of Euptychiina (Lepidoptera: Nymphalidae: Satyrinae) from the Brazilian southern Amazon

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REVISTA

BRASILEIRA

DE

Entomologia

AJournalonInsectDiversityandEvolution

w w w . r b e n t o m o l o g i a . c o m

Systematics,

Morphology

and

Biogeography

Description

of

a

new

and

highly

distinctive

genus

and

species

of

Euptychiina

(Lepidoptera:

Nymphalidae:

Satyrinae)

from

the

Brazilian

southern

Amazon

André

V.L.

Freitas

∗

,

Luísa

L. Mota,

Thamara

Zacca

,

Eduardo

P. Barbosa

UniversidadeEstadualdeCampinas,InstitutodeBiologia,DepartamentodeBiologiaAnimal,MuseudeZoologia,Campinas,SP,Brazil

a

r

t

i

c

l

e

i

n

f

o

Articlehistory:

Received18December2018 Accepted6May2019 Availableonline31May2019 AssociateEditor:LiviaPinheiro Keywords: Amazonia Immaturestages Morphology Molecularphylogeny Taxonomy

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b

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t

AnewgenusandspeciesofEuptychiina(Satyrinae),CristalinaiavitoriaMota,Zacca&Freitasgen.et

sp.nov.,isdescribedbasedonthreespecimenscollectedintheregionoftheCristalinoRiver, Alta

Floresta,MatoGrosso,Brazil.Thisrarespeciesisknownonlyfromthisregion,whereitﬂiesinside

thedense bamboopatchestypical ofthat area.Thelast instarlarvaand thepupa aredescribed;

thelarva was observedfeeding on matureleaves of thecommon bambooGuadua aff. paniculata

Munro.urn:lsid:zoobank.org:pub:D61EDE8D-CAE9-41C6-B24D-BB789873566E

Introduction

Inthelast decade,agoodnumberofgenerainthe Neotrop-ical subtribe Euptychiina (Nymphalidae: Satyrinae) have been more or less intensively studied, especially since the start of a large collaborative international effort that began in 2013 and has resulted in 50 publications on the ecology, natu-ral history, taxonomy and evolution of this group (see details in http://www.flmnh.ufl.edu/museum-voices/euptychiina/). Dur-ing the process, the immature stages have been described for severalspecies(e.g.,Freitasetal.,2016a,b;Freitas,2017),new phy-logenetichypothesishavebeenproposedbasedonmorphologyand moleculardata(Marín etal.,2017;Espelandetal.,2019), many smallandlargegenerahavebeenreorganized (e.g.Zaccaetal., 2017,2018),andnewgeneraandspecieshavebeendescribedfrom throughouttheNeotropics(e.g.Barbosaetal.,2015,2016;Freitas etal.,2015,2018;Huertasetal.,2016;Nakaharaetal.,2015,2018, 2019;Zaccaetal., 2014;Benmesbah etal.,2018).Anumber of newspecieshavebeenidentifiedbasedonintegrativetaxonomic approachestargetingcrypticspeciescomplexesinlargegenera,as inthegenusYphthimoidesForster,1964(Barbosaetal.,2015),while othersareclearlydistinctbasedonexternalmorphology,aswasthe casewiththegenusMoneuptychiaForster,1964,whichincreased

∗ Correspondingauthor.

E-mail:baku@unicamp.br(A.V.Freitas).

from2to8speciesinadecade(Freitasetal.,2015andreferences therein).

However, in the great majority of the above cases, newly describedspecieswerepartofcrypticcomplexesthatremained unnoticedinmuseumdrawers,ascaneasilybeobservedfromlists ofexaminedmaterial,includingthetypes.Bycontrast,thepresent studyisoutstandingbydescribingataxonapparentlynotpresent inanyofthemajorexaminedcollections.Moreover,thisnewtaxon wasinitiallydiscoveredfromasinglefield-collectedlarvathatwas rearedtoadult,resultinginahighlydistinctivesatyrinenotclosely resemblinganyotherknown.Subsequentintensivefieldworkin thesameregionresultedinonlytwoadditionalspecimens,which composethetypeseriesofthisnewspecies.Molecularand mor-phologicalcharactersconfirmthatthisnewEuptychiinaspeciesis seeminglynotcloselyrelatedtoanyoftheknowndescribedgenera. Accordingly,thepresentpaperdescribesthissouthernAmazonian taxonasanewgenusandspecies,anddiscussestherelevanceof thepresentfindingforEuptychiinasystematicsandconservation oftheregion.

Materialandmethods Studysite

Allthreeknownindividualswerecollectedintheregionofthe “CristalinoLodge”(9◦3551S,55◦5552W),locatednearthewest banksoftheCristalinoRiver,inthemunicipalityofAltaFloresta, https://doi.org/10.1016/j.rbe.2019.05.004

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Fig.1.HabitatandholotypeofCristalinaiavitoriagen.etsp.nov.inthetypelocalityatCristalinoLodge,AltaFloresta,MatoGrosso,Brazil.(A)Ageneralviewofthearea;(B) acloseviewofthebamboopatchwherethelarvawasfound;(C)maleholotypeofCristalinaiavitoriasp.nov.(dorsalabove,ventralbelow).

northernMatoGrossoState(near theborder withParáStatein northernBrazil) (Fig.1A,B).The climateof theregioniswarm and wet,withaverageannualtemperatureof 24◦C and annual rainfallabove 2400mm,with a dry seasonlasting 3–5months (Nimer, 1989).Thevegetationintheregionistransitionalfrom ombrophiloustosemidecidual(IBGE,2004).

Morphologicalstudies

Thelast instarcollectedin theﬁeld wasrearedin a 500mL plasticcontainerwithfreshleavesofitsbamboohostplantoffered adlibitum.Photographsofthelarvaandpupaweretakenwitha NikonD3100,andofthelastinstarcephaliccapsuleandmale gen-italiausingaZeissSteREODiscoveryV.20Stereomicroscope(Zeiss, Germany)inassociationwiththeAxioVisionRel.4.8softwarefor focus-stackingimages.

Oneadultspecimenhaditsabdomendetachedandsoakedina heatedtesttubewith10%potassiumhydroxidesolution(KOH)for about3–5mininbainmariebeforedissectionofthegenitalia.Wing sizewasmeasuredfrombasetoapexontheforewingandbaseto outermarginonthehindwingusingaruler.Theterminologyfor wingvenation,wingpatternelementsandgenitaliafollowsZacca etal.(2018).Thefollowingabbreviationsareusedthroughoutthe text:FW–forewing;HW–hindwing;DW–dorsalwings;VW– ven-tralwings;DFW–dorsalforewing;VFW–ventralforewing;DHW– dorsalhindwing;VHW–ventralhindwing;*–specimendissected. ThetaxonomichigherclassificationfollowsLamas(2004)modified afterPe ñaetal.(2006,2010)andWahlbergetal.(2009).

SpecimensofdifferentgroupsofSatyrinihavebeen morpho-logicallyexamined,photographedand/ordissectedbyallauthors inmorethan30publicandprivatecollections(seee.g.Zaccaetal., 2018),inadditiontotheoriginaldescriptionsofEuptychiinagenera (e.g.Forster,1964).Inaddition,photographsofNeotropical butter-ﬂytypespecimenstakenbyGerardoLamasandavailableinWarren etal.(2013)werealsoconsultedandcomparedwiththenewtaxon heredescribed.

DNAsamplingandanalysis

TotalDNAwasextractedfromtwolegsofadults(YPH-0835, BLU883andBLU1055)usingtheDNeasyBlood&TissueKit proto-col(QIAGEN,Düsseldorf,Germany).TotalDNA(Deoxyribonucleic acid)wasstoredinTEbufferat₋₂₀◦C.Themitochondrialgene cytochrome coxidaseI (COI– 1498bp),and thenuclear genes Glyceraldehyde-3-phosphate dehydrogenase (GAPDH - 691bp) and Ribosomal ProteinS5(RpS5 -610bp) for mostspecimens, wereamplified,purifiedandsequencedusingstandardtechniques (seeBarbosaetal.,2015;Silva-Brandãoetal.,2005;Wahlbergand Wheat, 2008). Theobtained sequenceswere deposited in Gen-Bank (Benson et al., 2013)(see Table 1 for voucher numbers). AllsequenceswereexaminedwiththeprogramFinchTVv.1.4.0 (Geospiza,PerkinElmerInc.,Waltham,MA),andposteriorlyaligned manually withsequences obtainedpreviously and available on GenBankbyusingBioEditv.7.2.4(Hall,2013).Thefinalmatrix com-prised81specimensfrom58speciesof34genera(includingthree specimensfromthenewgenusCristalinaiagen.nov.).The phy-logeneticrelationshipsofthenewspecieswereestimatedusing a Maximum Likelihood (ML)analysis. Analyses were runusing RAxML(Stamatakisetal.,2008)with250thoroughsearchesforthe maximumlikelihoodtopology,followedby1000rapidbootstrap replicateswiththedatamodeledaccordingtotheGTR+CATmodel (Stamatakiset al.,2008)andpartitionedbygene ontheCIPRES portal(Milleretal.,2010).

Results

CristalinaiaFreitas,Barbosa&Zaccagen.nov.

Type species. Cristalinaia vitoria Mota, Zacca & Freitas sp.n. urn:lsid:zoobank.org:act:C917EF80-C62A-4E41-8AA8- 1CCD6637EB5Eurn:lsid:zoobank.org:act:0A7CC6EA-95F5-4179-AAF8-6220389C4B6D

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SpeciesofEuptychiinawithcode,samplingsitedataandGenBankaccessionnumbersforsequencedgenes.Cristalinaiagen.nov.isinbold.Acronymsareexplainedinthe Methodssection.

Species DNAvoucher Locality COI GAPDH RPS5

Oressinomasorata CP06-89 Oxapampa,Peru GQ357209 GQ357440 GQ357570 Ypthimabaldus NW98-5 CentralSulawesi,PaloloValley,SEBerdikari,Indonesia DQ338875 EU528416 EU528469 Melanargialachesis NW149-3 Languedoc,Aude,Villegly,France GQ357222 GQ357463 GQ357593 Karanasabolorica NW166-10 EPamir,Karatekedistrict,Russia GQ357246 GQ357504 GQ357630 Erebiaoeme EW24-7 Languedoc,Ariege09,Ustou,BoisduPiéjau,France DQ338780 EU141479 EU141375 Maniolajurtina EW4-5 SantCliment,NSpain,Spain AY090214 EU141481 EU141376 Pronophilathelebe CP03-70 QuebradaSieteJeringas,Junín,Peru DQ338859 EU528410 EU528461 Paramaceraxicaque CP15-08 DistritoFederal,Mexico GQ357210 GQ357442 GQ357571 Cyllopsispertepida NW165-3 Guanajuato,Mexico GQ357204 GQ357428 GQ357557 Atlanteuptychiaernestina YPH-0191 SerradoJapi,Jundiaí,SãoPaulo,Brazil KP994863 KU340904 KU340937 Atlanteuptychiaernestina EE001 SerradoJapi,Jundiaí,SãoPaulo,Brazil MG209749 MG546307 — Atlanteuptychiaernestina EE002 SerradoJapi,Jundiaí,SãoPaulo,Brazil MG209750 MG546308 — Atlanteuptychiaernestina NW136-14 SãoLuísdoParaitinga,SãoPaulo,Brazil DQ338793 GU205947 GU206007

Cristalinaiavitoria YPH-0835 CristalinoLodge,AltaFloresta,MatoGrosso,Brazil MK575530 — —

Cristalinaiavitoria BLU883 CristalinoLodge,AltaFloresta,MatoGrosso,Brazil MK575531 — —

Cristalinaiavitoria BLU1055 CristalinoLodge,AltaFloresta,MatoGrosso,Brazil MK575532 MK575543 MK575548

Euptychiaatlantica MGCL-LOAN-216 ReservaSerraBonita,Camacan,Bahia,Brazil KX833117 — — Euptychiaenyo CP06-73 CordilleradelCóndor,Peru GQ357205 GQ357430 GQ357559 Euptychiamollina YPH-0154 Tapajós,Pará,Brazil KP890210 KU340902 KU340935 Inbiohilara LEP-14997 c.6kmELita,CachacoIM,Ecuador KR818704 — —

Inbiohilara LEP-19587 —— MK575533 — MK575549

Euptychiasp. SN-2015 —— KR818703 — —

Chloreuptychiacatharina CP01-68 MadredeDios,Peru GQ864749 GQ864942 GQ865413 Hermeuptychiaharmonia CP06-93 Oxapampa,Peru GU205842 GU205955 GU206014 Hermeuptychiapimpla CP04-10 QuebradaSieteJeringas,Junín,Peru GU205843 GU205956 GU206015 Hermeuptychiamaimoune YPH-0239 RioMadeira,PortoVelho,Rondônia,Brazil KR349479/KT880199 KR349475 KR349472 Hermeuptychiafallax YPH-0368 SantaTeresinha,Bahia,Brazil KU340868 — KU340950 Pindissquaminstriga NW165-5 Guanajuato,Mexico GQ357211 GQ357445 GQ357574 Godartianaarmilla PM04-02 Brasília,DistritoFederal,Brazil JQ392582 JQ392816 JQ392921 Godartianaluederwaldti YPH-0200 Goiás,Goiás,Brazil KT880201 KT880210 — Godartianamuscosa YPH-0319 FozdoIguac¸u,Paraná,Brazil KT880203 KT880212 — Godartianamuscosa NW127-8 SerradoJapi,Jundiaí,SãoPaulo,Brazil DQ338582 GQ864974 GQ865443

Amphidectacalliomma YPH-0355 —— MK575534 MK575544 MK575550

Amphidectacalliomma YPH-0356 —— MK575535 MK575545 MK575551

Amphidectapignerator YPH-0308 FozdoIguac¸u,Paraná,Brazil MK575536 MK575546 MK575552

Zischkaiapacarus CP14-02 —— GQ864819 GQ865049 GQ865512

Splendeuptychiaitonis CP02-44 MadredeDios,Peru DQ338811 GQ357447 GQ357576 Splendeuptychiahygina YPH-0317 FozdoIguaçu,Paraná,Brazil MK575537 — — Splendeuptychiaboliviensis CP02-48 MadredeDios,Peru GU205866 GU205980 GU206041 Splendeuptychialibitina YPH-0318 FozdoIguaçu,Paraná,Brazil MK575538 MK575547 — Forsterinariaboliviana CP04-88 QuebradaSieteJeringas,Junín,Peru DQ338799 GQ357435 GQ357564 Forsterinarianecys NW126-10 RibeirãodasPedras,SãoPaulo,Brazil GU205837 GU205950 — Forsterinariaquantius PM10-05 SerradoRolaMoça,NovaLima,MinasGerais,Brazil JQ392596 JQ392829 JQ392934 Forsterinariaitatiaia YPH-0453 SerradoCaparaó,MinasGerais,Brazil MK575539 — — Forsterinariaitatiaia YPH-0454 SerradoCaparaó,MinasGerais,Brazil MK575540 — — Seponapunctata YPH-0240 Abunã,PortoVelho,Rondônia,Brazil KR349480 KR349476 — Seponapunctata YPH-0494 ConceiçãodoMatoDentro,MinasGerais,Brazil KR349482 — — Seponapunctata YPH-0503 ReservaTrabijú,Pindamonhangaba,SãoPaulo,Brazil KR349484 — — Taygetistripunctata YPH-0579 Aquidauana,MatoGrossodoSul,Brazil MK575541 — MK575553 Taygetinabanghaasi LEP-00435 FilodeChumbiriatza,Zamora-Chinchipe,Ecuador JQ392633 JQ392854 JQ392964 Harjesiaobscura YPH-0238 RioMadeira,PortoVelho,Rondônia,Brazil KU340858 KU340905 KU340938 Erichthodesantonina YPH-0325 Alagoas,Brazil KU340864 KU340913 KU340946 Prendaclarissa AVLF-1 SãoFranciscodePaula,RioGrandedoSul,Brazil HQ444284 — — Taydebispeculiaris NW149-11 CamposdoJordão,SãoPaulo,Brazil GQ864811 GQ865036 GQ865499 Taydebispeculiaris YPH-0375 CamposdoJordão,SãoPaulo,Brazil MK575542 — — Nhambikuaramima MGCL-LOAN-550 PortoVelho,Rondônia,Brazil MF489994 MF490009 MF490025 Nhambikuaramima MGCL-LOAN-175 Ibateguara,Alagoas,Brazil MF489993 — — Nhambikuaracerradensis YPH-0572 SerradoCipó,SantanadoRiacho,MinasGerais,Brazil MF489984 MF490004 MF490017 Nhambikuaracerradensis YPH-0573 SerradoCipó,SantanadoRiacho,MinasGerais,Brazil MF489985 MF490005 MF490018 Nhambikuaracerradensis YPH-0574 SerradoCipó,SantanadoRiacho,MinasGerais,Brazil MF489986 MF490006 MF490019 Nhambikuaracerradensis YPH-0575 Aquidauana,MatoGrossodoSul,Brazil MF489987 — MF490020 Nhambikuaracerradensis YPH-0576 Aquidauana,MatoGrossodoSul,Brazil MF489988 MF490007 MF490021 Nhambikuaracerradensis MGCL-LOAN-134 Itirapina,SãoPaulo,Brazil MF489995 — — Splendeuptychiafurina CP02-39 MadredeDios,Peru GU205868 GU205982 GU206043 Splendeuptychiatoynei LEP-10657 LosEncuentros-Zarza,Zamora-Chinchipe,Ecuador MF489990 — MF490023 Splendeuptychiadoxes MGCL-LOAN-386 SerradoJapi,Jundiaí,SãoPaulo,Brazil MF489992 — — Splendeuptychiadoxes BC-DZWill-092 Pq.EstadualLagoAzul,CampoMourão,Paraná,Brazil MF489991 — MF490024 Splendeuptychiadoxes NW126-8 Atibaia,SãoPaulo,Brazil GU205867 GU205981 GU206042 Splendeuptychiadoxes YPH-0172 SerradoJapi,Jundiaí,SãoPaulo,Brazil KR349478 KR349474 — Splendeuptychialatia MGCL-LOAN-500 ReservaBiológicadeUna,Una,Bahia,Brazil MF489996 — — Paryphthimoidesgrimon CP10-01 Saibadela,SeteBarras,SãoPaulo,Brazil DQ338806 GQ865015 GQ865483 Cissiamyncea CP01-58 TambopataResearchCenter,MadredeDios,Peru GU205832 GU205944 GU206004 Cissiamyncea NW108-6 Picinguaba,Ubatuba,SãoPaulo,Brazil DQ338581 GQ357427 GQ357556 Magneuptychiamoderata CP01-36 MadredeDios,Peru GU205847 GU205960 GU206019

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Table1(Continued)

Species DNAvoucher Locality COI GAPDH RPS5

Splendeuptychiaashna CP01-19 MadredeDios,Peru GU205865 GU205979 GU206040 Cissiasimilis DNA97-014 OrangeWalkDistrict,Belize AY508529 — — Cissiasimilis 07-SRNP-14171 AreadeConservacionGuanacaste,CostaRica JQ538434 — — Magneuptychiaocypete CP01-32 TambopataResearchCenter,MadredeDios,Peru GU205848 GU205961 GU206020 Magneuptychiapallema CP02-41 AlberguePosadaAmazonas,MadredeDios,Peru GU205849 GU205962 GU206021 Capronnieriagalesus NW167-5 Pq.Nac.SerradoItajai,Apiúna,SantaCatarina,Brazil GU205826 GU205938 GU205998 Archeuptychiacluena NW149-9 Est.Biol.Boracéia,Salesópolis,SãoPaulo,Brazil GQ864736 GQ864926 GQ865392

Fig.2.LateralviewoftheheadofCristalinaiavitoriagen.etsp.nov.

Diagnosis.Cristalinaiagen.nov.differsfromallothergeneraof Eup-tychiinabytheVHWocelliwithverybroadorangeocellarrings, reducedblackocelarspotswithnopupils(Fig.1C)andthemale8th abdominaltergitestronglysclerotized,exceptbytheunsclerotized antero-dorsalregion(Fig.4B,C).Thefemaleisunknown.

Etymology.ThegenericnameisderivedfromtheCristalinoRiver, ariverofdarktranslucentwatersnearwhichthespecimenswere collected(theword “cristalino”is Portuguesefor“crystalclear”, alludingtothetranslucentwatersoftheriver).Italsoalludesto theReservaParticulardo PatrimônioNaturalCristalino (Cristal-ino PrivateReserve) and tothe“Fundac¸ãoEcológicaCristalino” (CristalinoEcologicalFoundation),bothfoundedbyVitoriadaRiva Carvalho.Thegenderofthenameshouldbeconsideredfeminine. CristalinaiavitoriaMota,Zacca&Freitassp.nov.(Figs.1–6). Typematerial.Holotypemale,depositedintheMuseude Zoolo-giadaUniversidadeEstadualdeCampinas(ZUEC),Campinas,São Paulo,Brazil (Fig. 1C). Labelson the holotype (ﬁvelabels sep-arated by transverse bars): /HOLOTYPUS/BRAZIL, Mato Grosso, AltaFloresta, CristalinoLodge,9◦3541S55◦5552W,2.II.2016, 240–260m,LLM280,LuisaL.Motaleg.,exlarva/DNAvoucher– BLU883/ZUECLEP10648/CristalinaiavitoriaMota,Zacca&Freitas det.2018/.

Paratypes.2males(1dissected).BRAZIL–MatoGrosso:Alta Flo-resta,CristalinoLodge,9◦3541S55◦5552W,1male,20.IV.2016, LLM 509 (DNA voucher BLU 1055), ZUEC LEP 10650, 1 male, 3.VIII.2018, LLM 52.2018 (DNA voucher YPH 0835), ZUEC LEP 10649*,LuisaL.Motaleg,ZUEC.

Diagnosis. Cristalinaia vitoria sp. nov. superﬁcially resembles several other species of Euptychiina (e.g. Pareuptychia species, Pseudeuptychia species, Splendeuptychia toynei Willmott & Hall, 1995,S.aurigera(Weymer,[1911]),S.triangula(Aurivillius,1929)) bythewhitegroundcoloronthewings,butitcanbeeasily dis-tinguishedfromthembytheincompletemedianlineandfourtiny ocellifromM1toCuA2ontheVFW,bytheﬁveVHWocellibetween

M1to2Athathaveaverybroadorangeocellarringandblackocellar

Sc R1 R2 R3 R4 R5 M1 M1 CuA1 CuA1 CuA2 CuA2 2A 2A 3A 1 cm hu Sc+R1 Rs M2 M2 M3 M3

Fig.3. WingvenationofCristalinaiavitoriagen.etsp.nov.Sc,subcostalvein;R, radialvein;M,medianvein;CuA,cubitalvein;A,analvein;h,humeralvein;+, fusionbetweenveins.

spotinthedistalhalfofringplusaverysmallocelliinRs-M1,andthe

broadenedVHWreddishbrownmarginallineformingcrescentsin eachcell.

Description.Head:Brownandglabrous eyes;antennaewith38 flagellomeres(n=1),whitishshortscalesatbaseofeach flagellom-ere;labialpalpicoveredbydarkbrownelongatedscalesmixedwith creamyscales;thirdsegmentwithshortbrownscalesandcreamy scalesinlateralview,firstandthirdlabialpalpisegmentsalmost samelength,secondsegment1½lengthofthirdsegment(Fig.2). Thorax:darkbrowncoveredbywhitishshortscalesandelongated filiformscales;coxaeandfemurcoveredbywhitishshortscales, tibiaandtarsuscoveredbydarkbrownshortscalesandreduced spines;pairoftibialspursatdistalendoftibia.Wings:FWlength: 16.5mm(HT)–17–19mm(PT).Venation(Fig.3):FWwith sub-costalandmedia-cubitalveinsdilatedatmid-basalregion(more than10timeswiderthanmid-apicalregionofvein)and2Aslightly dilatedat mid-basalregion(about twicewiderthan mid-apical regionofvein);discalcellhalf-lengthofwinglength.HW with

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t1 t2 t3 _t8 s8 s7 s6 s5 s4 s3 s1+2 tg c u g v sa sp 2 mm

B

D

E

F

G

* *

H

C

Fig.4. AbdomenandmalegenitaliaofCristalinaiavitoriagen.etsp.nov.:(A) Abdomen(lateral);(B–C)8thabdominaltergite:(B)lateral,(C)dorsal;(D–H)Male genitalia:(D)lateral,(E)dorsal,(F)ventral,(G)Phallus,lateral;(H)Phallus,dorsal.t, tergite;s,sternite;tg,tegumen;g,gnathos;u,uncus;v,valvae;c,costae;sa,saccus; sp,spine.ThesmallwhiteasteriskinEandFindicatestheinsectpinusedforplacing thegenitaliainposition.

developedhumeralveincurvedindirectiontocostalmargin;discal cell2/3lengthofentirelengthofwing.DFW:groundcolorbrown, whitishareabetweenM3andinnermarginfrombasetomedian

region.DHW:groundcolorbrown,whitishareabetweencostaland innermarginfrombasetomedianregion.Elementsofventralwings canbeseenthroughouttransparency.VFW:similartoDFW;thin submedianlinerestrictedtodiscalcell,irregularthinmedianline fromcostalmarginto2A,thincrenulatedsubmarginallinefrom

tornus;fourtinyocellifromM1 toCuA2.VHW:thinsubmedian

linefromcostaltoinnermargin,thinmedianlinerestrictedtoCuA2

andinnermargin,submarginallinecrenulatedfromcostaltoinner margin,broadmarginallineformingcrescentsineachcell(Fig.1C); sixocellifromRsto2A,tinyocellusincellRs-M1,ocellifromM1

to2Awithabroadorangeocellarringsurrounding,blackocellar spotdistallyandwithnopupils.Abdomen:darkbrowncovered bywhitishshortscale;8thtergiterectangularandstrongly sclero-tized,exceptedbytheantero-dorsalunsclerotizedregion(Fig.4B, C);8thsternitereduced.Malegenitalia(Fig.4A,D–H):Tegumen subtriangularlaterally;appendixangularisabsent;ventral projec-tionoftegumenanddorsalprojectionofsaccusfusedandangulated atmid-region;anteriorprojectionofsaccuswell-developedand ventrallyopen;uncusrobust,taperingatapexlaterally;gnathos smallerthanuncusandrobust;valvaetrapezoid,2/3lengthof sac-cuswithtworeduced spines atapex,covered bysmallﬁliform scales,costaereduced;phallus2/3lengthofanteriorprojectionof saccus,vesicawithoutcornuti.Female:Unknown.

Distribution.Cristalinaiavitoriasp.nov.isknownonlyfromits typelocalityintheregionofCristalinoLodge,AltaFloresta, north-ernMatoGrosso.Thesiteisaprivateprotectedarea(the“Reserva ParticulardoPatrimônioNaturalCristalino”).

Naturalhistory.Thisspeciesisextremelyscarce;onlythree indi-vidualshavebeenobserved(andcollected)overthecourseofabout 600hof field work,distributedover more than250days inall months,fromSeptember2015toAugust2018.Inaddition, pre-viouscollectingtripstothetypelocalitybyKeithS.BrownJr.and AVLF(inFebruaryandJune2000)andseveralbutterflywatching groupsthatvisitedtheareadidnotdetectthisspecies(LLM&AVLF inprep.).Basedontheveryfewavailabledatafromthethreeknown individuals,C.vitoriasp.nov.isrestrictedtothedensebamboo patchesthatarecommonwithintheforestsoftheregion.Thetwo adultswerefoundwithin5mfromeachother,althoughmorethan twoyearsapart.Theywerebothobservedintheshadyunderstory ofthebamboopatchataroundmiddaywithtemperaturesabove 30◦C,whicharenotunusualinthearea(LLMpers.obs.).Theirflight waslow(between0.5and1.0m)anderratic,andtheyrestedonthe

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CP06-89 Oressinoma sorata

NW98-5 Ypthima baldus NW166-10 Karanasa bolorica

NW149-3 Melanargia lachesis EW24-7 Erebia oeme

EW4-5 Maniola jurtina

CP03-70 Pronophila thelebe

MGCL-LOAN-216 Euptychia atlantica

YPH-0174 Euptychia mollina CP06-73 Euptychia enyo EE001 Atlanteuptychia ernestina

YPH-0191 Atlanteuptychia ernestina

EE002 Atlanteuptychia ernestina

NW136-14 Atlanteuptychia ernestina NW165-3 Cyllopsis pertepida CP15-08 Paramacera xicaque

LEP-19587 Inbio hilara SN-2015 Inbio hilara LEP-14997 Inbio hilara CP01-68 Chloreuptychia catherina

BLU1055 Cristalinaia vitoria YPH-0835 Cristalinaia vitoria BLU883 Cristalinaia vitoria CP04-10 Hermeuptychia pimpla CP06-10 Hermeuptychia harmonia

YPH-0368 Hermeuptychia fallax YPH-0239 Hermeuptychia maimoune

NW165-5 Pindis squamistriga YPH-0308 Amphidecta pignerator

YPH-0356 Amphidecta calliomma

YPH-0355 Amphidecta calliomma

YPH-0319 Godartiana muscosa YPH-0200 Godartiana luederwaldti

YPH-0317 Splendeuptychia hygina YPH-0318 Splendeuptychia libitina YPH-0240 Sepona punctata

YPH-0503 Sepona punctata YPH-0494 Sepona punctata

YPH-0579 Taygetis tripunctata YPH-0238 Harjesia obscura PM10-05 Forsterinaria quantius

CP04-88 Forsterinaria boliviana NW126-10 Forsterinaria necys YPH-0454 Forsterinaria itatiaia YPH-0453 Forsterinaria itatiaia

CP10-01 Paryphythimoides grimon

NW108-6 Cissia myncea CP01-58 Cissia myncea NW167-5 Capronnieria galesus

CP02-41 Magneuptychia pallema CP01-32 Magneuptychia ocypete

DNA97-01 Cissia similis

07-SRNP-14171 Cissia similis CP01-19 Splendeuptychia ashna

CP01-36 Magneuptychia moderata NW149-9 Archeuptychia cluena

YPH-0325 Erichthodes antonina BR-AVLF-1 Prenda clarissa

NW149-11 Taydebis peculiares YPH-0375 Taydebis peculiares

MGCL-LOAN-175 Nhambikuara mima MGCL-LOAN-550 Nhambikuara mima

YPH-0572 Nhambikuara cerradensis YPH-0574 Nhambikuara cerradensis YPH-0573 Nhambikuara cerradensis YPH-0575 Nhambikuara cerradensis MGCL-LOAN_134 Nhambikuara cerradensis YPH-0576 Nhambikuara cerradensis LEP-10657 Splendeuptychia toynei CP02-39 Splendeuptychia furina

MGCL-LOAN-500 Splendeuptychia latia NW126-8 Splendeuptychia doxes BC-DZWill-092 Splendeuptychia doxes

MGCL-LOAN-386 Splendeuptychia doxes YPH-0172 Splendeuptychia doxes 100 100 0.04 99 100 100 100 100 95 74 93 85 100 100 100 100 100 100 100 98 90 94 100 100 100 100 100 100 100 100 100 93 99 99 85 75 83 100 100 100 100 96 84 86 96 98 85

LEP-00435 Taygetina banghaasi PM04-02 Godartiana armilla

CP14-02 Zischkaia pacarus CP02-44 Splendeuptychia itonis

CP02-48 Splendeuptychia boliviensis NW127-8 Godartiana muscosa

Fig.6.MaximumLikelihoodconsensustreeshowingthephylogeneticrelationshipsbetweenCristalinaiavitoriagen.et.sp.nov.andotherEuptychiinagenera.Numbers abovebranchesarebootstrapvalues.Valuesbelow75arenotshown.

uppersideofleaveswithwingsclosed.Asinglelastinstar(Fig.5A, B)wasfoundintheﬁeld,inasunnyareaabout100mawayfrom wheretheadultswerecollected,feedingonmatureleavesofthe commonbambooGuaduaaff.paniculata.Thelarvaisentirelygreen withnumerouslongitudinalthinlightstripesonthebodyandwith apairofcaudalﬁlamentsaboutfourtimesthelengthofthe10th abdominalsegments.Theheadisgreenwithapairofshortscoli, oneoneachsideoftheepicranial/verticalnotch(Fig.5C).Thetotal larvallengthwas27mm,theheadcapsulewidthwas2.34mmand theheadscoliwereoflength1.1mm.Thepupa(Fig.5D,E)is elon-gatedandsmooth,entirelygreenwiththinsubtledorsaldarkgreen stripesandawhitelinedorsallyborderingthewingcases.The ocu-larcapsareveryshortandrounded,thecremasterisgreenandthe dorsalabdomenissmooth,withoutprojections.Thetotal pupal lengthwas13mm.

Etymology.The specificnameis afterVitoriadaRivaCarvalho, inrecognitionofherpioneeringworkontheconservationofthe southern Amazon, and founder of the Cristalino group (which includesthe“FundaçãoEcológicaCristalino”,CristalinoLodgeand CristalinoPrivateNaturalHeritageReserves).Thespecificepithet shouldbeconsideredfeminineandindeclinableinaccordanceto theArticle31.2.3(ICZN,1999).

Phylogeneticanalyses

MaximumLikelihoodanalysisusingDNAsequencedatafrom threegenes(Fig.6)supportsthedescriptionofthisnewgenusand newspecies,althoughmoresequencedataand/ortaxaareneeded tomoreconfidentlyrefinetherelationshipsofCristalinaiagen.nov. withinEuptychiina.Basedonthecurrentanalysis,thenewgenus appearsasanisolatedlongbranchandcannotbeassignedtoanyof theknowngeneraoranyoftheEuptychiinacladespreviously pro-posedbyPe ñaetal.(2010),Marínetal.(2017)andEspelandetal. (2019)orinongoingstudies(EPBunpublisheddata),asituation similartothatreportedforotherstudiesdescribingnew monoba-sicgenera(Freitasetal.,2016c;Kaminskietal.,2017;Nakahara etal.,2019).

Discussion

ThewingcolorpatternofCristalinaiavitoriagen.etsp.nov.is atﬁrstglancesomewhatsimilartothatofseveralother Amazo-niansatyrines,especiallysomespeciesofSplendeuptychiaForster, 1964.ThepatternofthewhitishcolorationontheDFWandDHWof Cristalinaiavitoriagen.etsp.nov.resemblesthatofPseudeuptychia

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Hall,1995,S.aurigera(Weymer,[1911]),S.triangula(Aurivillius, 1929)andmostspeciesofPareuptychiaForster,1964.However,C. vitoriasp.nov.canbeeasilydistinguishedfromallofthesespecies bythecurvedmedianlinefromtheRadiusto2AontheVFW,one tinyocellusinRs-M1andﬁveocelliwithbroadorangeocellarring

fromM1to2AonVHW.

ThemalegenitaliaofC.vitoriasp.nov.resemblesthatfoundin speciesofStevenariaViloria,Costa,Neild&Nakahara,2016(see Costa et al., 2016, Figs 40–41), mainly by the long saccus and phallusoccupyingmorethana halfoftheabdomenlength,but therearesigniﬁcantdifferencesinotherstructuresofC.vitoriasp. nov.,assuchtherobustgnathos,phalluswithsmallspineatthe dorso-apicalregion(Fig.4G),apexofthevalvaewithtworeduced spines(Fig.4D)andvesicawithoutcornuti. Similarlongsaccus andphallusareknowntooccurinsomeclearwingbutterﬂiesin thesubtribe Godyridina(Danainae:Ithomiini;seeWillmottand Freitas,2006),agroupnotcloselyrelatedtoSatyrinae(Wahlberg etal.,2009).AnotherinterestingfeatureofC.vitoriasp.nov.is thepatternofsclerotizationofthe8thabdominaltergite(Fig.4B, C); in otherspecies of Euptychiina, the8th tergite is membra-nouswithasclerotizedhalf-ringattheanteriorregion(Nakahara etal.,2016;Willmottetal.,2018;TZpers.obs.).Theabovecited characteristicsmakeC.vitoriagen.etsp.nov.uniqueamongthe Euptychiina.Othercharactersofthemalegenitaliaandwingcolor patternwerealsoinconclusivetoassignC.vitoriagen.etsp.nov. to any known Euptychiina clade, agreeing with the molecular data.

Concerningtheimmaturestages,thelastinstarisatfirstglance similartothoseofseveralEuptychiinawithgreen-stripedlarva, especiallythoseofZischkaiapacarus(Godart,[1824])and Godar-tianamuscosa (A.Butler,1870) (Freitasand Pe ña,2006; Freitas etal., 2016a,2016b; Zaccaetal., 2017;Seeet al.,2018). How-ever,theshapeoflastinstarheadscoliandthepupalprofileare quitedistinctfromallthesespecies.Infact,thelarvalcolor pat-ternisverysimilartothegroundcolorpresentintheelongated leavesofbamboosandgrasses,suggestingthatcamouflagecouldbe involved(seealsoFreitas,2017forasimilarcase),explainingwhy severalunrelatedspeciesofEuptychiinaareconvergentinlarval coloration.

Finally, it is important to draw attention to the conserva-tion status of the entire region of the Cristalino River in the AltaFloresta municipality.The forestsof the regionhave been highlyimpactedinrecentdecades,especiallyinthewestern fron-tierof the ParqueEstadual Cristalino(Cristalino StatePark (de Paulo et al., 2015)). The regional landscape is now dominated by forest fragments embedded in a matrix of pastures, result-inginobviouslossofhabitatqualityandmajorchangesinlocal climate (Dubreuil et al., 2012). However, in the “Reserva Par-ticulardo PatrimônioNaturalCristalino”, thetype localityof C. vitoria gen. et sp. nov., the forests are still preserved; more-over, the area is contiguous to the large Cristalino State Park and,hopefully,thetotalpreservedarea(summing184,000ha)is largeenoughtoprotectthenewtaxonheredescribed,aswellas theentireregionalbiota.Thisworkthus underlinesthe impor-tance of conservation efforts in the Southern Amazon, where otherremarkable,phylogeneticallydistinctivespeciessurelyawait discovery.

Authorcontributions

Manuscriptidealization:AVLFandLLM;collectingand imma-turestagesrearing:LLM;morphology:TZ;phylogeny:EPB;writing, reviewandﬁgures:Allauthors.

Theauthorsdeclarenoconﬂictsofinterest. Acknowledgements

The authors thank Cristalino Lodge and Fundação Ecológica Cristalinoforlogistic support.Keith Willmottcarefullyreadthe lastversionofthemanuscriptandgavevaluablesuggestions.The ICMBioprovidedthecollectingpermits(10438-1,7462-1, 8969-1).AVLFacknowledgessupportfromFAPESP(Biota-Fapespgrants 2011/50225-3 and 2013/50297-0), from the Brazilian Research Council – CNPq(fellowship 303834/2015-3),from theNational Science Foundation (DEB-1256742) and from USAID and the U.S. National Academy of Sciences (NAS) under the PEER pro-gram(SponsorGrantAwardNumber:AID-OAA-A-11-00012).EPB and TZ thank FAPESP for Pos-Doc fellowships (2016/15873-8, 2017/02264-6,respectively).LLMthanksCAPESforadoctoral fel-lowship.ThisstudywasfinancedinpartbytheCoordenaçãode AperfeiçoamentodePessoaldeNívelSuperior–Brasil(CAPES)– FinanceCode001. ThispublicationispartoftheRedeLep‘Rede NacionaldePesquisaeConservaçãodeLepidópteros’ SISBIOTA-Brasil/CNPq(563332/2010-7).Butterfly speciesareregisteredin theSISGEN(ACBC599).

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