'Species' from two different butterfly genera combined into one: description of a new genus of Euptychiina (Nymphalidae: Satyrinae) with unusually variable wing pattern

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REVISTA

BRASILEIRA

DE

Entomologia

A Journal on Insect Diversity and Evolution w w w . r b e n t o m o l o g i a . c o m

Systematics,

Morphology

and

Biogeography

‘Species’

from

two

different

butterﬂy

genera

combined

into

one:

description

of

a

new

genus

of

Euptychiina

(Nymphalidae:

Satyrinae)

with

unusually

variable

wing

pattern

André

Victor

Lucci

Freitas

a,∗

_,

_Eduardo

_Proenc¸

_a

_Barbosa

a

_,

_Keith

_Richard

_Willmott

b

_,

Niklas

Wahlberg

c,d

_,

_Gerardo

_Lamas

e

a_Department_of_Animal_Biology,_Instituto_de_Biologia,_Universidade_Estadual_de_Campinas,_Campinas,_SP,_Brazil

b_McGuire_Center_for_Lepidoptera_and_{Biodiversity,}_Florida_Museum_of_Natural_History,_University_of_Florida,_Gainesville,_United_States c_Laboratory_of_Genetics,_Department_of_Biology,_University_of_Turku,_Turku,_Finland

d_Department_of_Biology,_Lund_University,_Lund,_Sweden

e_Department_of_Entomology,_Museo_de_Historia_Natural,_Universidad_Nacional_Mayor_de_San_Marcos,_Lima,_Peru

a

r

t

i

c

l

e

i

n

f

o

Articlehistory: Received8May2015 Accepted12January2016 Availableonline4March2016 AssociateEditor:HéctorVargas Keywords: Harjesia Integrativetaxonomy Pseudodebis Satyrini Yphthimoides

a

b

s

t

r

a

c

t

SeponaFreitasandBarbosa,gen.nov.isproposedfortheNeotropicalsatyrinebutterﬂyspecies

Eup-tychiapunctataWeymer,1911anditsjuniorsubjectivesynonymsEuptychiagriseolaWeymer,1911and

TaygetisindecisaRibeiro,1931.Thenewgenushasadistinctivewingpatternandshapeofthevalvaein

themalegenitalia,thelatterbeingauniqueautapomorphywithinthesubtribeEuptychiina.Basedon

moleculardata,thisgenusisnotsistertoanyothersingleeuptychiinegenus,insteadappearingasthe

sistertoallremaininggeneraintheTaygetisclade.Thepresentpaperillustratesthecomplexityofthe

taxonomyofEuptychiina,andtheimportanceofusingdifferentsourcesofevidenceintaxonomicstudies.

accessarticleundertheCCBY-NC-NDlicense(http://creativecommons.org/licenses/by-nc-nd/4.0/).

Introduction

Inrecentyears,thehighlydiversebutterﬂysubfamily

Satyri-naehasbeensubjecttoseveralstudiesattemptingtoclarifyits

internalrelationshipsandtaxonomy(MurrayandProwell,2005;

Pe ˜naet al.,2006,2010; Marínetal.,2011;Matos-Maravi etal., 2013;Siewertetal.,2013;Seraphimetal.,2014).Thesestudies

haverevealedmanynon-monophyleticgenera,andanumberof

complexesofcrypticspecieswaitingtobedisentangled,especially

inthepredominantlylowland,largelyNeotropicalsubtribe

Eupty-chiina(e.g.Pe ˜naetal.,2010;Freitasetal.,2012a,b;Matos-Maravi

etal.,2013;Zaccaetal.,2013;Siewertetal.,2013;Seraphimetal., 2014).

Including10genera,the“Taygetisclade”isoneofﬁvemajor

groups of Euptychiina (Pe ˜na et al., 2010); a preliminary

phy-logenyforthisclade(Matos-Maravietal.,2013)showedthatfour

genera,namelyHarjesiaForster,1964,PseudodebisForster,1964,

∗ Correspondingauthor.

E-mails:baku@unicamp.br(A.V.L.Freitas),niklas.wahlberg@biol.lu.se (N.Wahlberg).

ForsterinariaR.Gray,1973andTaygetisHübner[1819],are

poly-phyletic, requiring somerevised generic combinations and the

description of new genera. The genus Harjesia, as then

con-ceived,included species placed inthree differentclades within

the“Taygetisclade”(Matos-Maravietal.,2013).Inthatphylogeny,

Harjesiagriseola(Weymer,1911)appearedassistertotheentire

“Taygetis clade” (Matos-Maravi et al., 2013), suggesting that it

shouldbeplacedinanewgenus.

Ongoingresearchintothephylogeneticrelationshipsofanother

euptychiinegenus,YphthimoidesForster,1964,showedthatthis

genusisclearlypolyphyletic,withseveralspeciesthatshouldbe

reassignedtoothergenera(Freitasetal.,2012b,Barbosaetal.,2015,

EPBand AVLF,inprep.).Onespecies inparticular, Yphthimoides

punctata(Weymer,1911),isquitedistinctfromallotherdescribed

Yphthimoides,andfurthermorphologicalstudiesrevealedthatY.

punctataandH.griseolaaresimilarenoughtobeconsidered

sub-jectivesynonyms.

Thisstudypresentsevidencebasedonanintegrativetaxonomic

approach(e.g.,Dayrat,2005;Yeatesetal.,2011;Panteetal.,2015)

usingbothmorphologicalandmoleculardataforthesynonymyof

Y.punctataandH.griseola,anddescribesanewgenustoharborthe

resultingsinglespecies.

http://dx.doi.org/10.1016/j.rbe.2016.01.004

0085-5626/© 2016 Sociedade Brasileira de Entomologia. Published by Elsevier Editora Ltda. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).

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Materialandmethods

Adultspecimenswerestudiedina numberof Americanand

Europeancollections,andthefollowingacronymsareusedhere:

AWLW– Allan &Lesley Wolhuter collection, United Kingdom;

DZUP – Departamento de Zoologia, Universidade Federal do

Paraná,Curitiba,Paraná,Brazil;FLMNH–FloridaMuseumof

Nat-uralHistory, Gainesville,FL, USA; KWJH –Keith R.Willmott&

Jason P. W. Hall collection, Gainesville, FL, USA; LBCB: L. & C.

Brévignoncollection,FrenchGuiana;MNHN–MuséumNational

d ´HistoireNaturelle,Paris,France;MNRJ–MuseuNacionaldoRio

deJaneiro,RiodeJaneiro,Brazil;MOBE:MohamedBenmesbah

col-lection,France;MUSM–MuseodeHistoriaNatural,Universidad

NacionalMayordeSanMarcos,Lima,Peru;MZUJ–Muzeum

Zoo-logiczneUniwersytetuJagiellónskiego,Kraków,Poland;NHMUK

–TheNaturalHistoryMuseum,London,UnitedKingdom;YUGA

– Yuvinka Gareca colletion, Santa Cruz, Bolivia; ZSM –

Zool-ogische Staatssammlung München, München, Germany; ZUEC

– Museu de Zoologia da Universidade Estadual de Campinas,

Campinas,SãoPaulo,Brazil;ZUEC-AVLF–AndréV.L.Freitas

Col-lection,UniversidadeEstadualdeCampinas,Campinas,SãoPaulo,

Brazil. Morphology

Dissectionsweremadeusingstandardtechniques.Legs,palpi

andabdomensweresoakedinhot10%potassiumhydroxidefor

nearly 10minutes before dissection, and dissected parts were

storedinglycerol.Inordertoseethevenation,wingswere

diaph-anizedbysoakingtheminalcohol andNaClOsolution(bleach).

Taxonomicnomenclature followsLamas (2004a,b),modiﬁed by

Pe ˜naetal.(2006)andWahlbergetal.(2009).Drawingsand

mea-surements of wings, legsand palpiwere made using a Leica®

MZ7.5stereomicroscopeequippedwithamicrometricscaleand

adrawingtube.Photographsofthemaleandfemalegenitaliawere

takenusingaZeissDiscoveryV20Stereomicroscope.Thefollowing

abbreviationsareused:(FW)forewing,(HW)hindwing,(D)dorsal,

(V)ventral.

Phylogeneticinference

GenomicDNAwasextractedfromtwolegsofadultsbyusing

theDNeasyBlood&TissueKitprotocol(QIAGEN,Düsseldorf,

Ger-many).DNAwasstoredinTEbufferat−20◦_C._The_{mitochondrial}

genecytochromecoxidaseI(Cox1,ca.658bp,correspondingto

the‘DNAbarcode’region)forallspecimensandthenucleargenes

GAPDHforonespecimen(YPH-0240)andRpS5fortheoutgroups

wereampliﬁed,puriﬁedandsequencedusingstandardtechniques

(seeSilva-Brandãoetal.,2005;WahlbergandWheat,2008).The

sequencesofnucleargeneRpS5forHarjesiagriseolawereobtained

fromGenBank.

All the sequences were aligned by eye with sequences

obtainedpreviouslyandavailableonGenBankbyusingBioEditv.

7.2.4(Hall,2013,availableathttp://www.mbio.ncsu.edu/bioedit/

bioedit.html#downloads).The ﬁnal matrix comprised 32

speci-mensfromspeciesof10genera(including11specimensfromthe

newgenusSepona)andthreespeciesusedasoutgroups,namely

Hermeuptychia maimoune (A. Butler, 1870), Paryphthimoides

grimon(Godart[1824])andSplendeuptychiadoxes(Godart[1824])

(seeTable1forthesequencecodes).

Thephylogeneticrelationshipsofthenewspecieswere

esti-matedusingmaximumlikelihood.AnalyseswererunusingRAXML

(Stamatakisetal.,2008)with1000rapidbootstrapreplicatesand

asearchforthemaximumlikelihoodtopologyontheCIPRES

por-tal(Milleretal.,2010).Thedataweremodeledaccordingtothe

GTR+Gmodelforeachpartitionindependently.

Table1

SpeciesofEuptychiinawithcode,samplingsitedata,andGenBankaccessionnumbersforsequencedgenes.

Speciesname Code Locality COI GAPDH RpS5

Seponapunctata YPH-0240 Abunã,PortoVelho,RO,Brazil KR349480 KR349476 – Seponapunctata YPH-0346 Est.Eco.AltoAcre,AssisBrasil,AC,Brazil KR349481 – – Seponapunctata YPH-0494 Conceic¸ãodoMatoDentro,MG,Brazil KR349482 – – Seponapunctata YPH-0502 Est.Eco.AltoAcre,AssisBrasil,AC,Brazil KR349483 – – Seponapunctata YPH-0503 ReservaTrabijú,Pindamonhangaba,SP,Brazil KR349484 – –

Seponapunctata YPH-0506 Bonito,MS,Brazil KR349485 – –

Seponapunctata YPH-0507 Bonito,MS,Brazil KR349486 – –

Seponapunctata YPH-0511 PortoVelho,RO,Brazil KR349487 – –

Seponapunctata CP23-21 ProvinciaLaConvención,Peru JQ392607 JQ392838 JQ392943

Seponapunctata CP24-01 MadredeDios,Peru JQ392608 – JQ392944

Seponapunctata CP24-02 MadredeDios,Peru JQ392609 – JQ392945

Hermeuptychiamaimoune YPH-0239 RioMadeira,PortoVelho,RO,Brazil KR349479 KR349475 KR349472 Paryphthimoidesgrimon YPH-0095 TrêsLagoas,MS,Brazil KR349477 KR349473 KR349471 Splendeuptychiadoxes YPH-0172 SerradoJapi,Jundiaí,SP,Brazil KR349478 KR349474 – Taygetinaoreba CP02-13 TambopataResearchCenter,MadredeDios,Peru JQ392613 JQ392842 JQ392949

Taygetinaoreba CP-CI107 CICRA,Peru GU205839 GU205952 GU206011

Taygetinabanghaasi LEP00435 FilodeChumbiriatza,Zamora-Chinchipe,Ecuador JQ392633 JQ392854 JQ392964 Taygetisweymeri PM03-03 Magistral,Sinaloa,Mexico JQ392708 JQ392918 JQ393027

Taygetislarua PM14-24 – JQ392669 JQ392885 JQ392995

Taygetisvirgilia PM02-03 Amalfí,Porcé,Antioquia,Colombia JQ392700 JQ392912 JQ393021

Taygetislaches PM04-13 Xingu,PA,Brazil JQ392659 JQ392878 JQ392988

Forsterinariaquantius PM10-05 Pq.E.SerradoRolaMoc¸a,NovaLima,MG,Brazil JQ392596 JQ392829 JQ392934 Parataygetislineata NN58 PuentePuntayacu-Qbda.LaSolitaria,Junín,Peru JQ392618 JQ392846 JQ392953 Parataygetisalbinotata CP07-43 Chanchamayo,Junín,Peru JQ392616 JQ392844 JQ392951 Posttaygetispenelea CP01-06 TambopataResearchCenter,MadredeDios,Peru JQ392620 – JQ392955

Posttaygetispenelea UN0403 TeodoroSampaio,SP,Brazil JQ392623 – –

Pseudodebisvalentina CP01-85 TambopataResearchCenter,MadredeDios,Peru JQ392629 – –

Pseudodebisvalentina CP-CI25 CICRA,Peru JQ392631 JQ392852 JQ392962

Harjesiablanda CP01-13 TambopataResearchCenter,MadredeDios,Peru DQ338800 GQ357436 GQ357565

Harjesiablanda CP-CI57 CICRA,Peru JQ392606 JQ392837 JQ392942

Harjesiaobscura CP23-22 Peru JQ392610 JQ392839 JQ392946

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SeponaFreitas&Barbosa,gen.nov.

Typespecies:EuptychiapunctataWeymer, 1911,here desig-nated.

Diagnosis

Moleculardata(Pe ˜naetal.,2010)placethisgenuswithinthe

“Taygetisclade”ofthesatyrinesubtribeEuptychiina.Intermsof

wingshapeandpattern,adultsaresimilartothoseofsomespecies

ofHarjesia,PseudodebisandTaygetinaForster,1964,whichallhave

undulateratherthanstraightdarkdiscalandpostdiscallineson

theventralsurface,butthisgenuscanbedistinguishedfromall

othereuptychiinesbytheuniqueshapeofthevalvaeinthemale

genitalia(Fig.3A).Thevalvaebearalong,thin,inwardlycurved

pro-jectionarisingabruptlyfromtheotherwiseroundedmainbodyof

thevalva.SeeTable2forcomparisonsofadditionalmorphological

charactersofSeponapunctatawithrepresentativesofothergenera

ofthe“Taygetisclade”.TherelationshipsofSeponatoother

eupty-chiinagenera,andjustiﬁcationforitsrecognitionasamonotypic

genus,areaddressedfurtherbelowunder‘Discussion’.

Etymology

Seponaisanarbitrarycombinationofletters,derivedfromthe

Latintransitive verb “sepono”, meaning toput aside, separate,

orremove, inreferencetotheisolatedpositionofthegenusin

comparisonwithothermembersofthe“Taygetisclade”.Itshould

betreatedasafemininenoun.

Seponapunctata(Weymer,1911)comb.nov.

EuptychiapunctataWeymer,20April1911:205.TypeLocality:Brazil,

MinasGerais.Syntype(s),notlocated.

=EuptychiagriseolaWeymer,20July1911:211,n.syn.TypeLocality:

Bolivia[LaPaz],Mapiri.Lectotypefemale(heredesignated):‘Original!/nun

aberheissengriseola//Mapiri//Collection/v.Rosen//grisea/Weym.//Typus

Nr./EuptychiagriseolaStgr.i.l./Weymer/Zoologische/Staatssammlung/

München’,depositedinZSM(examined).

=TaygetisindecisaRibeiro,1931:33,n.syn.TypeLocality:Brazil

[Rondô-nia,RioJamari].Holotypefemale:‘Comm.Rondon/No.43.19.6.14/Coll.

Stolle//HOLOTIPO//N053/645//HolótipoTaygetis/indecisa/Ribeiro,1931

/Mielke&Casa-grandedet.1985’,depositedinMNRJ(examined).

Taygetisindecisa: May,1933:120;Euptychiagriseola: May,1933:120;

D’Abrera,1988:789;Pseudodebisgriseola:Forster,1964:77;Lamas,1983:16;

Robbinsetal.,1996:230;Freitas,2003:103;Euptychiapunctata:D’Abrera,

1988:789;Taygetisgriseola: D’Abrera,1988:756;Harjesiagriseola:Lamas,

2004:219; Brévignon,2008: 68; Brévignon andBenmesbah,2012: 39;

Matos-Maravietal.,2013:54;Yphthimoidespunctata:Lamas,2004:223.

Redescription

Male(Figs.1,2A-C-D-E,4A-B-D).Eyesreddishbrown,covered

withsparseblackhairs.Palpus1.5timesaslongashead,brown

withlightbrownhairs(Fig.2C).Antennaofmales9.0–10.0mmin

lengthwith36antennomeres,extendingtomid-costa;shaft

rust-browndorsally,orangebrownventrally,sparselyscaleddorsally;

clubnotconspicuouslydeveloped,includingelevensegments,with

apicalportion(lastﬁvesegments)darkbrown.Forewinglength

23–25mm(n=6); hindwinglength19–20mm(n=6).HWouter

marginslightlyundulate.Male wingvenation showninFig.2A.

Wingswithdorsalgroundcolordarkbrownwithfewmarkings,

restrictedtoa suffused darkbrownoutermarginonDFW, and

todarkdouble marginal line, and a submarginal linefollowing

contours ofmarginal lineonDHW. Ventral wingslight brown;

VFW crossed by two thin zigzag dark brown lines, extending

fromcostato2A,ﬁrstlineone-thirddistancefromwingbaseto Table

2 Comparisons of Sepona punctata with exemplar species from related genera in “Taygetis clade”. For aedeagus and saccus width, the value refers to length/width ratio in the medial portion of each structure; higher numbers therefore represent a narrower structure. Species Eyes Forewing apex Hind wing margin Shape of aedeagus Aedeagus width Valvae Uncus shape Gnathos shape Tegumen Saccus width Sepona punctata 1, 2 Few sparse hairs Rounded Slightly wavy Strongly curved 23 Bearing a long thin terminal projection Curved, broad Elongated, pointed Pronounced 11 Harjesia blanda 4 Hairy Rounded Wavy Straight 8 Projection absent Curved, slender Elongated, pointed Not pronounced 4 Posttaygetis penelea 4, 9 Hairy Rounded Wavy Curved in the distal end 15 Projection absent Straight, broad Short, rounded Not pronounced 4 Pseudodebis valentina 4 Hairy Rounded Slightly wavy Straight 12 Projection absent Curved slender Short, broad Slightly pronounced 5 Pseudodebis celia 4 Hairy Truncate Wavy Straight 10 Projection absent Straight, slender Short, rounded Not pronounced 4 Taygetina kerea 5 Hairy Acute Wavy Straight 13 Projection absent Straight. Broad Elongated, pointed Pronounced 7 Taygetis yphthima 7 Short sparse hairs in lateral portion Acute Wavy Straight 12 Projection absent Straight, slender Elongated, pointed Pronounced 4 Taygetis laches 3, 6 Hairy Truncate Wavy Straight 25 Projection absent Straight. Slender Elongated, pointed Slightly pronounced 8 Taygetis mermeria 4 Hairy Acute Wavy Straight 10 Projection absent Straight, slender Elongated, pointed Not pronounced 6 Taygetis sylvia 4 Hairy Acute Wavy Straight 6 Projection absent Straight, broad Elongated, pointed Not pronounced 4 Taygetis echo 8 Few sparse hairs Rounded Wavy Straight 14 Projection absent Straight, slender Elongated, pointed Not pronounced 5 Source of material (all localities in Brazil) 1, Pindamonhangaba, SP; 2, Jaru, RO; 3, Linhares, ES; 4, Marechal Thaumaturgo, AC; 5, Jatai, SP; 6, Campinas, SP; 7, Campos do Jordao, SP; 8, Alta Floresta, MT; 9, Morro do Diabo, SP.

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Fig.1.AdultmaleofSeponapunctata–Jaru,Rondônia,Brazil.Dorsalabove,ventral below.

apex;secondlineextendingfromcostato2Aattwo-thirds dis-tancefromwingbasetoapex;aconspicuouslighterouterband is adjacent tosecond line, followed by a darker ocellarregion (seenext);athindarkbrownzigzagsubmarginallinewithsingle blackdotsinverticesandabrownregularmarginalline extend-ingfromcostato2A;fourdarkocelliin spacesR5–M1(ocellus 1),M1–M2(2),M2–M3(3) and M3–CuA1(4). VHW crossedby two thin darkbrown lines from costa to anal margin, in sim-ilarposition tothose onforewing; a conspicuouslighter outer band is adjacent to second line, followed by a darker ocellar region (see next); a dark brown zigzag submarginal line with singleblackdotsin verticesand a brownregular marginalline extending from costa to 2A; a series of ﬁve dark ocelli can be found in cells Rs–M1 (ocellus 1), M1–M2 (2), M2–M3 (3), M3–CuA1(4) and CuA1–CuA2 (5).Details aboutocellisize and shapediscussedfurtherbelow.Noconspicuousandroconialscales observed.

Malegenitalia(Fig.3A–E).Saccuselongate;tegumenrounded

andshort;gnathos longandpointed,projectingupwardsabove

uncus;uncuselongated,withlateralexpansionsindorsalview,

giv-inganarrowheadappearance;valvaeelongated,endinginabump

withalongthinpointedprocess;aedeaguscurved;cornutiabsent;

juxtasclerotized,linkingbothvalvaetogether.

Female(Fig.2B,F,4C-E-F).Forewinglength24–26mm(n=2);

hindwinglength20–22mm (n=2). Antenna 11.0mm inlength,

with35antennomeres,extendingtomid-costa.Generalcolorand

patternverysimilartothoseofmales.FemalegenitaliaasinFig.3E,

F.Ductusbursaepartiallysclerotized,corpusbursaerounded;apair

ofconspicuoussignapresent.

Taxonomyandvariation

Weymer (1911) described Euptychia punctata based on an

unstated number of specimens from Minas Gerais, Brazil.

Sev-eralpageslater,hedescribedEuptychiagriseolabasedalsoonan

unstatednumberofspecimensfromMapiri,Bolivia.Later,Ribeiro

(1931)describedathirdtaxonasTaygetisindecisa,basedonone

female from Brazil, Rio Jamari; this taxon was promptly

syn-onymizedwithEuptychiagriseolabyMay(1933).Descriptionsof

bothE.punctataandE.griseolawerealsobasedonfemale

spec-imens,and althoughthis cannotbedeterminedunambiguously

fromtheiroriginaldescriptions,thefactthatthetypesarefemales

inbothcasessuggeststhatmaleswereunknowntotheauthors.

Notype specimen(s)ofpunctatahasbeenfound,but thereis a

singlefemaleinZSMidentiﬁedbyForster(1964)asthe“Typus”

of griseola, and we accept that it indeed representsa syntype

(which we designate hereinas lectotype), since this particular

specimen(examined)matchespreciselytheillustrationprovided

subsequentlybyWeymer(1911:pl.47g,ﬁg.[7]).Thefemale

holo-type(examined)ofTaygetisindecisaRibeiroisdepositedinMNRJ.

Althoughappearingrather differentin wingpattern,thenames

punctataandgriseolaapparentlyrepresentextremesof

geograph-icalvariationwithinasinglespecies.Variationonthedorsalwing

surfacesispracticallyabsentandobviousseasonalvariationshave

notbeendetected.Theventralsurface ofboth wings,however,

showsmuch variation,especiallyinthenumberandsizeofthe

ocelli.MostindividualsfromcentralandsoutheasternBraziland

easternBolivia(“punctata”phenotype)havetheocellireducedto

smallblackdots,sometimeswithatinywhitepupilontheVHW;

theyalsopresentamorehomogeneousventralpattern(Fig.4E,F).

Conversely,individualsfromwesternAmazoniaandGuianas(the

“griseola”phenotype)havemoredevelopedocellicircledby

yel-lowishcreamscalesandwithawhitepupilonbothwings,anda

conspicuousbandedpatternontheventralwings(Fig.4A,B).

Inter-mediatephenotypesbetween“punctata”and“griseola”areknown

fromAcreandRondôniainBrazil,andfromBolivia,andare

usu-allymoresimilartothe“griseola”phenotype(Fig.4C,D).Toour

knowledge,notwoofthesethreephenotypes(“punctata”,

“grise-ola”andintermediate)havebeenrecordedinsympatry.Thetwo

nameswerepublishedseveralmonthsapart,andwethustreatE.

griseolaasajuniorsubjectivesynonymofS.punctatan.syn.

Specimensexamined(34♂,27♀):Bolivia.–Beni:Prov.Mamoré,

SanRamón,EstanciaS/Lorenzo[13◦269.26S,64◦362.79W],13

Oct 2003, T. V. Bosque de Galeria, T17, Yuvinka Gareca leg.

(YUGA); La Paz: ‘Madidi’ [14◦355S,68◦5049W] (Aliaga, W.),

4 Oct 2005, 1 ♀ (MUSM), 5 Oct 2005, 1 ♂ (MUSM); Mapiri

[15◦1927S,68◦650W], 1 ♂[LTgriseola; “Original! / nunaber

heissengriseola//Mapiri//Collection/v.Rosen//grisea/Weym.

//Typus Nr./ EuptychiagriseolaStgr.i. l./Weymer /

Zoologis-che/Staatssammlung/München”](ZSM);SantaCruz:Buenavista

[17◦27S,64◦40W], 750m(Steinbach,J.), Aug 1907–Apr1908,1

♂[BMNH-E-1204766](NHMUK).Brazil.–Acre:50kmNWBujari

[9◦3253S,68◦189W],200m(Mielke,O.,M.Casagrande),25Sep

2003, 1 ♀ (DZUP); AssisBrasil, Estac¸ão Ecológica do AltoAcre

[11◦3S,70◦16W], 300m(Brown, K. S.), 19 Jul 2005, 1 ♂[DNA

voucherYPH-0346](ZUEC-AVLF),21Aug2005,1♀(ZUEC-AVLF),

Aug2005,1♂[DNAvoucherYPH-0502](ZUEC-AVLF);Marechal

Thaumaturgo,ReservaExtrativistadoAltoJuruá,BocadoCaipora

[9◦916S,72◦4021W](Brown,K.S.&A.V.L.Freitas),1Sep1997,

1♂[ZUECLEP-9231],1♀[ZUECLEP-9232](ZUEC);Marechal

Thau-maturgo,upperJuruáRiver,Fozdo[Rio]Tejo[8◦591S,72◦43W]

(Freitas, A.V.L.), 20–27 Aug 1997, 1 ♀ (ZUEC-AVLF); PAD

Humaitá,PortoAcre[9◦3529S,67◦3220W], 200m(Mielke,O.,

M.Casagrande),8Oct2004,1_♂(DZUP);EspíritoSanto:Colatina

[19◦3155S,40◦4042W], Aug–Sep 1937, 3♂ [24/579, 52/439,

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A

C

D

E

F

B

Sc R1 R2 R3 R4 R5 M1 M2 M3 CuA1 CuA2 2A Rs M1 M2 M3 CuA1 CuA2 _6.0mm 1.5mm 0.4mm 1.75mm 0.5mm 2A dc dc h 3A Sc R1 R2 R3 R4 R5 Rs M1 dc h M2 M3 CuA1 CuA2 2A 3A M1 M2 M3 CuA1 CuA2 2A dc Sc+R1 Sc+R1

Fig.2.MorphologicalcharactersofSeponapunctata.A,malewingvenation–forewingaboveandhindwingbelow;B,femalewingvenation–forewingaboveandhindwing below;C,malepalpus;D,maleforeleg;E,malemidleg;Ffemaleforeleg.

Sabiá,Est.[rada]N.[ova]Venecia[19◦820S,40◦378W],Collatina

[sic],Oct1936,1♀,E.Mayleg.[52/336](MNRJ);MatoGrosso do

Sul:Bonito[21◦734S,56◦2925W] (Araújo,P.F.),15 Oct2014,

1 _♀ [ZUEC LEP-9234; DNA voucher YPH-0507] (ZUEC), 5 Nov

2014,1♂[ZUECLEP-9235;DNAvoucherYPH-0506](ZUEC);Mato

Grosso:11–18kmNRibeirãoCascalheira[12◦5047S,51◦4659W]

(Mielke,O.),21–23Aug1997,1♀(DZUP);C[oron]el.Rio Branco

[15◦1424S,58◦649W],400m(Buzzi,Mielke,Elias&Casagrande),

19 Sep 1984, 1 ♂ [DZ-5465; Proj.[eto] Polonoroeste] (DZUP)

(Mielke,O.,K.S.Brown),3Jul1972,1♂(DZUP);Diamantino,Rio

Arinos,Faz.[enda]S.[ão]João[14◦2118S,56◦92W],300–400m

(Ebert,H.&H.D.),5May1978,1♂(DZUP);ChapadadosGuimarães,

Buriti[15◦2331S,55◦5311W],1Jan1969,1♂[ZUECLEP-9246;

Slideno1762, ♂genitália, LeeD. Miller](ZUEC);Minas Gerais:

Conceic¸ãodo MatoDentro[19◦2’S,43◦2531W] (Ramos,G.), 11

Sep2013,1♀[ZUECLEP-9233;DNAvoucherYPH-0494](ZUEC);

nospeciﬁclocality (Boullet),1913,2_♂(MNHN);RiodeJaneiro:

Itabapoana[21◦8’S,41◦41’W],1♀ [BMNH-E-1204794](NHMUK);

Rondônia:67kmSAriquemes,5kmSofCacaulândiaonlinha

C-10atRioPardooffB-65[10◦2315S,62◦5453W](Austin,G.T.),14

Oct1993,1♂(FLMNH),8Oct1993,1♀(FLMNH)(Gomes,O.),15

May1994,1♀(FLMNH);67kmSAriquemes,5kmSof

Cacaulân-diaonlinhaC-10[10◦2313S,62◦5412W],170m(Gomes,O.),1

Aug1993,1_♂(FLMNH);Jaru[10◦27S,62◦30W](Brown,K.S.),30

Sep1975,2♀ (FLMNH),5Oct 1975,1♀ (FLMNH)(Callaghan,C.

J.),10Aug1976,1♂[genitaliavialAVF7841],1♀[genitaliavial

AVF7840](FLMNH);PôrtoVelho[8◦45S,63◦53W],31Jan2010,

1♂[ZUECLEP-9236;DNAvoucherYPH-0511](ZUEC);RioJamari

[10◦12S,63◦15W],1♂[HTindecisa;“’Comm.Rondon/No.43.19.

6.14/Coll.Stolle//HOLOTIPO//N053/645//HolótipoTaygetis/

indecisa/Ribeiro,1931/Mielke&Casa-grandedet.1985](MNRJ);

UpperRioMadeira,Abunã[9◦42S,65◦21W](Nunes,R.V.),1May

2013,1♂[ZUECLEP-9230;DNAvoucherYPH-0240](ZUEC);São

Paulo:Pindamonhangaba, ReservaNatural Municipaldo Trabijú

[22◦5018S,45◦3123W](Rosa,A.H.B.),22Jul2014,1♂[ZUEC

LEP-9237;DNAvoucherYPH-0503](ZUEC).Ecuador.–Napo:Río

Jatunyacu,Pimpilala[1◦431S,77◦5613W],800m(Jasinski,A.),28

May1997,1♀(MZUJ);Orellana:[c.13kmWCoca],RíoPayamino

[0◦2618S,77◦646W](Wolhuter,A.&L.),28Oct2009,1♀(AWLW)

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A

B

C

D

E

F

te gn un va sa bu st 0.5 mm 0.2 mm 0.2 mm 0.2 mm 0.2 mm 0.2 mm

Fig.3. MaleandfemalegenitaliaofSeponapunctata.A,malegenitaliainlateralview;B,malegenitaliaindorsalview;C,malegenitaliainventralview;D,maleaedeagus (lateralview);E,femalegenitaliainventralview;F.femalegenitalia:detailofthesignaincorpusbursae.(sa)saccus,(te)tegumen,(un)uncus,(va)valva,(bu)corpusbursae, (st)sterigma.

RíoBigal,8deDiciembre[0◦3215S,77◦2518W],975m(T.&M.

García),Oct2009(photographlivespecimen);Pastaza:Puyo,Chifa

Restaurant[1◦2913S,78◦0W],1000m(K.Willmott&J.Hall),25

Aug 1993,1 ♂(KWJH);13km N Puyo [1◦2326S,77◦5835W],

1100m(Emmel,T.C.),8Sep1969,1♀(FLMNH).FrenchGuiana.

– Saint-Laurent-du-Maroni: Maroni river, 1 ♀ (FLMNH). Peru. –

Cuzco:Echarate,AltoManugali[12◦27S,73◦02W],812m

(Valen-cia, G.), 5 Apr 2009, 1 ♀ (MUSM); Río Urubamba, Saringabeni

[12◦1105S,72◦4948W],442m(Cerde ˜na,J.,J.Farfán,R.

Gutiér-rez), 31 Aug 2007, 1 ♀ [ﬁsh bait trap] (MUSM); San Martín-3

Camp[11◦47S,72◦42W],475m(Valencia,G.), 11Apr1997,1♀

(MUSM);Loreto:‘CavalloCocha’[=Caballococha][3◦55S,70◦31W],

90m(Mathan,M.de),May-Jul1884,1♂[BMNH-E-786155],1♂

[BMNH-E-786156](NHMUK);Madrede Dios: BocaRío LaTorre

[12◦50S,69◦17W],300m(Covell,C.V.),22Oct1983,1♀(MUSM)

(Lamas, G.), 15 Feb 1982, 1 ♀ (MUSM); Parque Manu, Pakitza

[11◦5640S,71◦1658W], 340m (Mielke, O.), 1 Oct 1991, 1 ♂

(DZUP),24Sep1991,1♂(DZUP),27Sep1991,1♂(DZUP),7Oct

1991,1♂(DZUP);ParqueManu,Pakitza[11◦53S,70◦58W],400m

(Lamas,G.),16Oct1990,1♂(MUSM),20Oct1990,1♂(MUSM),

7Oct1990,1♂(MUSM)(Rowe,W.), 3Nov1990,1♀ (MUSM);

ReservaTambopata[12◦50S,69◦17W],300m(Lamas,G.),31Oct

1990,1♀(MUSM);QuebradaAguaNegra[12◦53S,69◦17W],200m

(Williams,H.),16Sep1995,1♀(MUSM);RíoMadredeDios,

Alber-gueAmazonia[12◦52S,71◦23W],500m(Lamas,G.),24Oct2013,

1♀(MUSM),28Sep2014,1♂(MUSM),29Sep2014,1♀(MUSM);

Salvación[12◦50S,71◦21W],500m(Matthews,M.J.),6Aug1982,

1♂(MUSM);RíoAzul[13◦3S,69◦55W],850m(Pe ˜na,C.),1Oct

2010,1♂[voucherCP24-01](MUSM),23Sep2010,1♂(MUSM);

Río Azul,ReservaComunal Amarakaeri[12◦49S,71◦6W], 507m

(Vílchez,M.),11Oct2010,1♀(MUSM).

Other records: French Guiana. – Cayenne, Roura, Cacao

[4◦35N,52◦28W](Damico,R.),Feb1996,1♀ (LBCB)(Brévignon,

2008: 68, pl. 7, ﬁg. 81, 82); Saint-Laurent-du-Maroni, Saül

[3◦37N,53◦12W], 9 Oct 2011, 1 ♂ (MOBE) (Brévignon &

Benmesbah,2012:46,pl.3,ﬁg.11[adultwings],11a[male gen-italia]).

Biologyanddistribution

SeponapunctataisknownfromeasternEcuadortosoutheastern

Brazil(EspíritoSanto,RiodeJaneiro,MinasGeraisandSãoPaulo).

TherearealsorecordsfromtwositesinFrenchGuiana(Brévignon,

2008;BrévignonandBenmesbah,2012)(Fig.6).IneasternEcuador

thespeciesisextremelyrare,andthefewknownlocalitiesarein

theAndeanfoothillsonthetypesofsandstonesoilsthatfrequently

supportstandsofbamboo.Adultsareusuallyscarceandrarein

collections,althoughtheyweresometimescommoninareaswith

largebamboopatchesintheupperJuruáRiver,inAcre,Brazil(AVLF

andK.S.BrownJr.,pers.obs.).Thespeciesisusuallyassociatedwith

forestedhabitats,butsomepopulationsinSEBrazil(the“punctata”

phenotype)areknownfromriparianforestsinthecerrado.The

immaturestagesandhostplantsareunknown.

Discussion

ThepositionofSeponapunctataasa well-supportedsisterto

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Fig.4. VariationinwingpatternofSeponapunctata(allfromBrazil).A,Abunã,RioMadeira,Rondônia;B,PortoVelho,Rondônia;C,Estac¸ãoEcológicadoAltoAcre,Acre;D, PortoAcre,PADHumaitá,Acre;E,Conceic¸ãodoMatodentro,MinasGerais;F,ParqueMunicipaldoTrabijú,Pindamonhangaba,SãoPaulo(A,B,D–males;C,E,F–females).

natureofHarjesiaasillustratedbyMatos-Maravietal.(2013)and

in thepresent paper (Fig. 5), clearly shows that this species is

notpartofHarjesia(whichhasasitstypespeciesTaygetisblanda

Möschler,1877). Thereasons forerecting a newgenusfor this

taxonarethereforeclear:unlessallspeciesinthe“Taygetisclade”

arelumpedintoasinglegenus,anundesirableoption giventhe

morphologicalvariationandtaxonomicdiversitywithintheclade,

thereisnowaytocircumscribemonophyleticgeneraintheclade

withoutmakingthistaxonamonobasicgenus.Inadditiontoits

phylogenetic position,themale genitaliaofS. punctatais quite

distinct from all known species of Harjesia (Forster, 1964 and

unpublished results of the authors), presenting several unique

characters,includingtheextremelythinandcurvedaedeagusand

theuniqueshapeofthevalvae(seeFig.3AandTable2).

TheknownwingpatternsofS.punctataarehighlyvariable,but

althoughspecimensfromwesternAmazoniaandGuianasarequite

Fig.5.RelationshipsamongSeponapunctataandselectedspeciesinthe“Taygetisclade”andseveraloutgroupsinferredwithmaximumlikelihood.Numbersnearbranch nodesarebootstrapbranchsupport.NamesinparenthesesforSeponapunctatarefertothephenotypeofthevoucherspecimens(seetext).

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Fig.6.RecordedlocalitiesofSeponapunctata(basedonallexaminedmaterial;seetextfordetails).Blackdots=“griseola”phenotype;whitedots=“punctata”phenotype; graydots=intermediatephenotypes.

divergentfromthose fromsoutheasternBrazil,individualswith

intermediatewingpatternareknownfromeastern Bolivia,and

Acreand Rondônia,Brazil.Inaddition,thesedifferencesarenot

relatedtoseasonalformsand,basedonthefewknownindividuals,

thereis lowvariation withinpopulations,includinginthesites

whereintermediatepopulationsareknown.Thesereasonswere

consideredsufﬁcienttonotrecognizesubspeciﬁctaxawithinthis

species.

Theabove-describedvariationinwingpatternsthroughoutthe

distribution of S. punctata easily explains why this taxon was

described as three different species, twiceby the same author

(Weymer,1911),inthreedifferentgenera(seethesynonymiclist

above).Thissituationisaperfectexampleofhowcomplexisthe

taxonomyofEuptychiina,wheremostofthelargegeneraare

non-monophyletic,withspeciesspreadintwoormoredifferentclades

(asisthecaseofSplendeuptychiaForster,1964,CissiaDoubleday,

1848,andParyphthimoidesForster,1964,seePe ˜naetal.,2010).

Hopefully,forthcomingstudiescombiningmorphologicaland

moleculardatawillhelptodisentanglethecomplexand

species-richcladewhichconstitutesthesubtribeEuptychiina,providinga

wellresolvedphylogenythatwillserveasaframeworkforfuture

studiesfocusingondiversiﬁcationpatternsofNeotropical

butter-ﬂies.

Conﬂictsofinterest

Theauthorsdeclarenoconﬂictsofinterest.

Acknowledgements

WewouldliketothankKeithS.BrownJr.,BlancaHuertas,

Mar-cioUehara-Prado,LeeD.Miller,JacquelineY.Miller,DebraMurray,

JasonHall,AxelHausmann,CarlaPenz,GláuciaMarconato,Yuvinka

GarecaandAndreasSegererfortheirhelpindiversephasesofthe

manuscript.WealsothankPolianaF.AraujoandAugustoH.B.Rosa

forprovidingfreshspecimens,LuizaM.Magaldiforhelpingwith

theDNAextractionandTamaraM.C.Aguiarforhelpingspreadthe

specimens.LeeD.MillerandJacquelineY.Millergavegreat

inspi-rationtotheﬁrstauthortoworkonSatyrinaesystematics.EPB

acknowledgesFAPESP(2012/03750-8)foragraduatefellowship.

AVLFacknowledgessupportfromFAPESP(Biota-Fapesp–grants

2011/50225-3,2012/50260-6,2013/50297-0),fromtheBrazilian

ResearchCouncil–CNPq(fellowship302585/2011-7).This

publica-tionispartoftheRedeLep‘RedeNacionaldePesquisaeConservac¸ão

dedeLepidópteros’SISBIOTA-Brasil/CNPq(563332/2010-7),ofthe

project “Identiﬁcac¸ão Molecular de Biodiversidade de

Inverte-bradosTerrestres”(grant564954/2010-1)includedinthe“Rede

Nacionalde Identiﬁcac¸ãoMolecularda Biodiversidade—BR-BoL”

(MCT/CNPq/FNDCT50/2010),thecollaborativegrant“Dimensions

US-Biota-SãoPaulo:AssemblyandevolutionoftheAmazonbiota

anditsenvironment:anintegratedapproach”,USNSF,NASA,and

FAPESP(2012/50260-6),andfromtheNationalScienceFoundation

(DEB-1256742). References

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