REVISTA
BRASILEIRA
DE
Entomologia
AJournalonInsectDiversityandEvolutionw w w . r b e n t o m o l o g i a . c o m
Systematics,
Morphology
and
Biogeography
A
new
species
of
Actinote
Hübner
(Nymphalidae:
Heliconiinae:
Acraeini)
from
southeast
Brazil
André
Victor
Lucci
Freitas
a,∗,
Ronaldo
Bastos
Francini
b,
Márlon
Paluch
c,
Eduardo
Proenc¸
a
Barbosa
aaUniversidadeEstadualdeCampinas,InstitutodeBiologia,DepartamentodeBiologiaAnimal,Campinas,SP,Brazil bUniversidadeCatólicadeSantos,Santos,SP,Brazil
cUniversidadeFederaldoRecôncavodaBahia,CentrodeCiênciasAgrárias,AmbientaiseBiológicas,CruzdasAlmas,BA,Brazil
a
r
t
i
c
l
e
i
n
f
o
Articlehistory:
Received24October2017 Accepted26January2018 Availableonline9February2018 AssociateEditor:LiviaPinheiro Keywords: AtlanticForest AraucariaForest Actinotealalia Speciesredescription Immaturestages
a
b
s
t
r
a
c
t
ThepresentpaperdescribesanewspeciesofActinote(Nymphalidae,Heliconiinae,Acraeini)from south-easternBrazil,anddescribesthemorphologyoftheadultsandimmaturestagesofthisspecies.Actinote mantiqueirasp.nov.occursintheSerradaMantiqueiraandSerradoMarintheAtlanticForest.Adults fromthispopulationareverysimilartootherspeciesofthe“orangishredmimicrycomplex”,including Actinotealalia(C.Felder&R.Felder,1860),itssisterspecies,restrictedtothemountainsofsouthernBrazil. Actinotemantiqueirasp.nov.andA.alaliaaredistinguishablebywingpattern,malegenitaliaandlarval morphology,andhavestronglyallopatricdistributions.AredescriptionofActinotealaliaisalsoprovided. ©2018SociedadeBrasileiradeEntomologia.PublishedbyElsevierEditoraLtda.Thisisanopen accessarticleundertheCCBY-NC-NDlicense(http://creativecommons.org/licenses/by-nc-nd/4.0/).
Introduction
ThegenusActinoteHübner,[1819](sensuLamas,2004,seealso
Silva-Brandãoetal.,2008)iscomposedof35 describedspecies,
occurringintheNeotropicalregionfromsouthernMexico (Guer-rero,Veracruz,OaxacaandChiapas)tonorthernArgentinainthe borderprovincesofLaRioja,CórdodaandBuenosAires(Lamas,
2004;Paluch,2006;Paluchetal.,2006;Neild,2008;Willmottetal.,
2009,2017).The genusreachesits greatest species richness in
themontaneareasof southeasternBrazil(Brown,1992;Lamas,
2004;Paluch,2006;Silva-Brandãoetal.,2008;Neild,2008;
Will-mottetal.,2009).Larvaeofallknownspeciesfeedonhostplants withinthefamilyAsteraceaeandarehighlygregariousinallinstars
(Francini,1989,1992;Freitasetal.,2010andreferencestherein).
Withsomenotableexceptions(e.g.ActinotezikaniR.F.d’Almeida, 1951,seeFrancinietal.,2005,2011),adultsofActinoteare asso-ciatedwithforestedges,clearingsandotheropenenvironments wheretheirhostplantsgrow(Francini,1989;FranciniandFreitas,
2010;Freitasetal.,2009a,b,2010).
SpeciesofActinotearecyanogenicinalllifestagesandrarely preyeduponbynaturalenemies,whichincludevisuallyoriented vertebrates (Brown and Francini, 1990), and adults are usually
∗ Correspondingauthor.
E-mail:baku@unicamp.br(A.V.L.Freitas).
involvedinMüllerianandBatesianmimicrycomplexeswith sev-eralspeciesofbutterfliesandmoths(Brown,1992).Inpartbecause ofthismimicry,Actinotebutterfliesarenotoriouslydifficulttotell apart(e.g.,D’Almeida,1925,1935;Francini,1989,1992;Penzand
Francini,1996),andspeciesidentificationcanbechallenging,often
resultinginmisidentification(especiallyforfemales)andmixed series frequently identified as a single species (Francini, 1989;
Paluch,2006).Conversely,larvaeofmostspeciesareusuallyquite
distinct,evenamongverysimilarspecies,andcanbevery help-fulforspeciesidentification,especiallywithinthe“orange-yellow mimicrycomplex”(Francini,1989;Francinietal.,2004).Molecular data,however,hasproventobeineffectiveindiscriminatingamong speciesinsomeclades,suchasintheActinotepelleneaclade(formed byA.pelleneaHübner,[1821],ActinotecarycinaJordan,1913and ActinotepyrrhaFabricius,1775)(Silva-Brandãoetal.,2008; Silva-Brandãoetal.,inprep.).
Thespeciesinthe“orangishredmimicrycomplex”(Francini, 1989)areclassicexampleoftheseproblemsinidentification.All specieswithinthegroupareverysimilar(mostofthem,infact, shareacommonancestor,seeSilva-Brandãoetal.,2008), charac-terizedbyadarkorangeandbrowndorsalstripedpattern,with a ventralpatternthat issomewhat variableamongspecies and thereforeusefulforidentification.FoursoutheastBrazilianspecies havebeendescribedwithinthegroup,from1860to1913,including Actinotealalia(C.Felder&R.Felder,1860).
IntheoriginaldescriptionofAcraeaalalia,Felder&Felder(1860)
providedjustabriefdescriptionofmalewingpattern;thenumbers
https://doi.org/10.1016/j.rbe.2018.01.003
0085-5626/©2018SociedadeBrasileiradeEntomologia.PublishedbyElsevierEditoraLtda.ThisisanopenaccessarticleundertheCCBY-NC-NDlicense(http:// creativecommons.org/licenses/by-nc-nd/4.0/).
ofindividualsusedforthedescriptionandtheexacttypelocality werenotprovided, withjustthecountryoforigin,Brazil,being mentioned.Lamas(1997)designateda male(withoutthehead) asthe Lectotypeof A. alalia, and proposedthat thetype local-itywassoutheasternBrazil.Inthesameyear,PenzandFrancini
(1996)describedthreenewspeciesbasedonmaterialpreviously
misidentifiedasA.alalia,solvingmostoftheproblemsrelatedto thiscomplexofcrypticspecies.Finally,inarecentunpublished the-sisaboutthegenusActinote,Paluch(2006)redescribedA.alaliain detail,includinghostplant,voltinismanditsgeographical distri-bution.
Detailedmorphologicalstudyof allsoutheasternBrazil pop-ulationsofA.alaliahaverevealedthatthisspecies,aspresently conceived,comprises twodistincttaxa, distinguishablebywing pattern,malegenitaliaand larvalmorphology,and withclearly allopatric distributions (amolecularstudy of the“orangish red mimicry complex” is currentlybeing prepared). In the present paper,anewspeciesofActinote,previouslyidentifiedasA.alalia,is describedbasedonmorphologyofadultsandimmaturestages.Due toseveralincongruencesintheoriginaldescription(not acknowl-edgedbyPaluch,2006),wealsopresentaredescriptionofActinote alalia.
Materialandmethods
Drawingsand measurements of wings, legs and palpi were madeusing a Leica® MZ7.5 stereomicroscope equipped with a micrometricscaleand adrawingtube.Photographsofthemale andfemalegenitaliaandsomemeasurementsweretakenusing aZeiss®DiscoveryV20Stereomicroscope.Dissectionsweremade usingstandardtechniques,wherelegs,palpi,andabdomenswere soakedinhot10%KOHsolutionfor10minbeforedissection,and dissectedpartswerestoredinglycerol.Tostudythevenation,wings werediaphonizedbysoakingtheminalcoholandNaClOsolution (bleach)andmountedinslides.Taxonomicnomenclaturefollows
Lamas(2004), modified after Wahlberget al. (2009). Male and
femalegenitaliaterminologyfollowsKlots(1970).Nomenclature ofvenation usedhereinfollowsWootton(1979)modified from
ComstockandNeedham (1898–99).Specimens ofActinotewere
examinedin14publicandprivatecollections(seebelow)(no spec-imensofA.alaliaorofthenewspecieswerefoundintheMNHNand USNM).TheLamascollectionofNeotropicalbutterflytype spec-imenphotographsattheMUSM(alsoavailableonlineinWarren etal.,2017),representingmostcurrentlyrelevantnamesand rec-ognizedspeciesofActinote(Lamas,2004),wasexamined.
The acronymsfor visited collections are: NHMUK, The Nat-ural History Museum, London, England; CGCM, Colec¸ão Carlos GuilhermeCosta Mielke,Curitiba,Paraná, Brazil; CLDZ,Colec¸ão deLepidoptera,DepartamentodeZoologia,UniversidadeFederal do Rio Grande do Sul, PortoAlegre, Rio Grande do Sul, Brazil; DZUP,DepartamentodeZoologia,UniversidadeFederaldoParaná, Curitiba,Paraná,Brazil;IOC,InstitutoOswaldoCruz,RiodeJaneiro, RJ,Brazil;MC,Colec¸ãoMoser,SãoLeopoldo, RioGrandedoSul, Brazil; MECB, Museu Entomológico Ceslau Biezanko, Departa-mentodeFitossanidade,UniversidadeFederaldePelotas,Pelotas, Rio Grande do Sul, Brazil; MNHN, Muséum National d’Histoire Naturelle,Paris,France;MNRJ,MuseuNacional,Universidade Fed-eraldoRiodeJaneiro,RiodeJaneiro,RiodeJaneiro,Brazil;MZSP, MuseudeZoologia,UniversidadedeSãoPaulo,SãoPaulo,Brazil; OM,Colec¸ãoOlafMielke,Curitiba,Paraná,Brazil;USNM,National Museum of Natural History, Smithsonian Institution, Washing-ton,DC,USA;ZUEC,MuseudeZoologiadaUniversidadeEstadual deCampinas,Unicamp,Campinas,SãoPaulo,Brazil;ZUEC-AVLF, AndréV.L.FreitasCollection,UniversidadeEstadualdeCampinas, Campinas,SãoPaulo,Brazil.
Results
ActinotemantiqueiraFreitas,Francini,Paluch&Barbosa,sp.nov. Actinote alalia;Ebert,1969:35; Lewis,1973:43,fig. 1;Lewis,
1975:43,fig.1;Penz&Francini,1996:313,fig.4a,b,c,317;Francini
&Penz,2006:figs.8,9,Plate12;Silva-Brandãoetal.,2008:519,528,
figs.3,4,5;Freitasetal.,2009b:88;Freitasetal.,2010:100,103.
Actinotealaliaalalia;Eltringham&Jordan,1913:10. Actinoteundescribedspecies;Freitasetal.,2010:103. Diagnosis
Thewing colorand pattern are typicalof the “orangishred mimicrycomplex” (Francini,1989),withdarkbrownstripeson anorangishredbackgroundonthedorsalwingsurfaces(Fig.9), butthenewspeciescanbedistinguishedfromallotherpreviously describedspeciesinthiscomplexbyseveralmorphological
charac-ters(seeFranciniandPenz,2006).ThemalesofActinotemantiqueira
sp.nov.canbedistinguishedfromthoseofActinotealaliabythe usu-allybroadertransversedarkbrownbandsonthedorsalforewing (Fig.2AandE).Onthedorsalforewing,A.mantiqueirasp.nov.can bedistinguishedfromA.alaliabythepresenceofausually con-tinuousdarkbandintheinternalmargin(thisismostlyorangein A.alalia).Ontheventralhindwing,A.mantiqueirasp.nov.canbe distinguishedfromA.alaliabythefaintlymarked,orangev-shaped transverseband(thisbandisconspicuouslymarkedinA.alalia),by theabsenceofacompletedarkbarcrossingthemiddleofthediscal cell(thisdarkbarispresentinA.alalia,delimitingabasalorange area),andbytheabsenceofadistinctpatchoforangescalesinthe humeralangle,basallytothehumeralvein(thisregioniscoveredby creamscalesinA.mantiqueirasp.nov.)(Fig.2BandF).Femalesare verysimilarinbothspecies(Fig.2CandG),andhardlydiscernible; themostconsistentcharacteristhepresenceofadistinctpatchof orangescalesinthehumeralangle,basallytothehumeralvein,in A.alalia(creaminA.mantiqueirasp.nov.)(Fig.2DandH).Themale genitaliaofbothspeciesareverysimilar,andthemaindifferenceis theprofileofthesaccus,whichismoretriangularandpointedinA. mantiqueirasp.nov.,andbumpedandabruptlyenlargingnearthe baseinA.alalia(Fig.3DandL).
Fig.1.Holotypemale(A)andallotypefemale(B)ofActinotemantiqueirasp.nov. (left=dorsal;right=ventral).Scalebar=10mm.
Actinote mantiqueira Actinote alalia
A
C
E
G
B
D
F
H
Fig.2.ComparativeplateshowingthemaindifferencesinwingpatternbetweenActinotemantiqueirasp.nov.andActinotealalia.(A)MaleparatypeofA.mantiqueirasp.nov. (dorsalleft,ventralright);(B)detailofhumeralregioninmaleVHW;(C)femaleparatypeofA.mantiqueirasp.nov.(dorsalleft,ventralright);(D)detailofhumeralregionin femaleVHW;(E)maleofA.alalia(dorsalleft,ventralright);(F)detailofhumeralregioninmaleVHW;(G)femaleofA.alalia(dorsalleft,ventralright);(H)detailofhumeral regioninfemaleVHW.
Description
Male(Figs.1A,2A,B,4A–D,9A):Antenna black,11–12mmin
length(n=11), extendingtomid-costa, with 42 antennomeres, 9 of which form a well-defined club. Palpus length about 2.0 timesheadheight.MalepalpusshowninFig.4B.Forewingnarrow andelongated,length25–28mm(mean=26.8mm,SD=1.08mm, n=11);hindwingrounded,abouttwo-thirdslengthofforewing, length 18–21mm (mean=19.7mm, SD=0.90mm, n=11). Male wingvenationasshowninFig.4A.Maleforeleg(Fig.4C)witha sin-gletarsomere.MalemidlegasshowninFig.4D.Bodydarkbrown, abdomenventrallycoveredbycreamscales.Forewingdorsal back-grounddarkorangewithdarkbrownveinsandstripesasfollows: abroaddarkbrownmargin,broadinapexandnarrowingtoward tornus;entirespacefromanalmarginto2Adarkbrown;three addi-tionalbroaddarkbrowntransversestripes,firstextendingfrom costatoCuA2,twothirdsfrombase,secondv-shapedcrossing dis-calcellinmidportion,andthirdcrossingspaceCuA2-2Ahalfway frombase.Manyindividualspresentadarkcubitalspotadjacent todiscalcellnearwingbase.Hindwingdorsal backgrounddark
orangewithbroaddarkbrownmargin;allveinsdarkbrown,and narrowdarkbrownstripesextendingininter-venalspaces;some individuals canpresenttransversepatches ofdarkscales cross-ingproximallyspacesM1-M2,M2-M3,M3-CuA1 andCuA1-CuA2, insomecasesalmostformingav-shapedtransverseband. Ven-tralforewingwitha pattern verysimilar todorsalpattern, but lackingbroaddarkbrownmargin;ventralhindwingmore homo-geneouslycream,withdarkbrownveinsandstripesininter-venal spaces.
Malegenitalia(Fig.3A–G):Valvaeelongatedandcurved,length aboutfourtimeswidthofmedianportion;broaderatbase;apex roundedwithaconspicuouscurltowardmiddlelineofbody.In dor-salview,basalportionofuncuswide,abruptlynarrowing,ending inapoint.Tegumenlongandbroad,trapezoidalwithaslight con-strictionatmiddle.Gnathosabsent.Inventralview,saccusshaped asanisoscelestriangle,abouthalflengthofgenitalcapsule. Aedea-gusabouthalflengthofgenitalcapsule,endinginasharppoint, inlateralviewstraight.Juxtabroad,aboutsamewidthofbaseof onevalva(inventralview),tear-shaped(withananteriorpointed process);lengthequaltowidth.
Actinote mantiqueira Actinote alalia sa va va ae ae un un un un va va sa 1mm 1mm sa sa ju ju
A
B
D
E
F
G
H
I
J
P
N
L
K
M
O
Fig.3.MaleandfemalegenitalcharactersofActinotemantiqueirasp.nov.(A–H)andActinotealalia(I–P).A.mantiqueirasp.nov.:(A)lateralviewofmalegenitalia;(B)uncus indorsalview;(C)aedeagusdorsal(above)andlateral(below);(D)saccus,ventralview;(E)valva,ventralview;(F)juxta;G.8thsternite;(H)femalesterigma.A.alalia:(I) lateralviewofmalegenitalia;(J)uncusindorsalview;(K)aedeagusdorsal(above)andlateral(below);(L)saccus,ventralview;(M)valva,ventralview;(N)juxta;(O)8th sternite;(P)femalesterigma.ae,aedeagus;ju,juxta;sa,saccus;un,uncus;va,valva.
3A dc h h
A
B
C
D
F
G
H
E
dc Sc Sc R1 R2 R3 R4 R5 M1 M2 M3 CuA1 CuA2 2A Sc+R1 Rs M1 M2 CuA2 1A 2A 3A dc dc CuA1 R1 R2 R3R4 R5 M1 M2 M3 CuA1 CuA2 2A 2A 1A CuA2 CuA1 M3 7.0mm M2 M1 Rs Sc+R1 1.5mm 2.0mm 1.5mm 2.0mmFig.4.Morphologicalcharactersofmales(A–D)andfemales(E–H)ofActinotemantiqueirasp.nov.(A)Wingvenation;(B)Palpus;(C)Foreleg;(D)Midleg;(E)Wingvenation; (F)Palpus;(G)Foreleg;(H)Midleg.
Female(Figs. 1B,2C,D, 4E–H,8B):Antenna black,12mm in length(n=3),extendingtomid-costa,with41antennomeres,10 in club. Female palpus as shown in Fig. 4F. Forewing narrow andelongated,length29–31mm(mean=29.3mm,SD=1.53mm, n=3). Hindwing rounded, not translucent, length 20–22mm
(mean=20.7mm, SD=1.15mm, n=3). Female wing venation is showninFig.4E.Colorpatternoffemalewingssimilartothatof malesbutpaler.Forelegwithsixsegmentsontarsus,withthird segmentbearingoneshortsetae(Fig.4G).Femalemidlegasshown inFig.4H.
Femalegenitalia(Fig.3H):Corpusbursaerounded,signaabsent. Ductus bursae not sclerotized, same length as corpus bursae; sterigmatrapezoidal.
Taxonomyandvariation
AshasbeenreportedformostspeciesofActinote,A.mantiqueira sp.nov.alsopresentsintraspecificvariation,especiallyinfemale colorpattern(malesaremoreuniform).Minorvariationhasalso beenreportedinwingvenationandgenitalia.Themostvariable charactersare theextentof thehumeralvein, whichcan reach ornot thewingedge, andtheshape ofthesaccusapex, which canvaryfrompointedtoalmostrounded.Thepopulationsfrom SerradaBocaina(SãoPauloState)areslightlydifferentonthen ventralhindwing,withtheorangev-shapedtransversebandmore conspicuouslymarked.
Holotype(Fig.1A).Male from Pindamonhangaba, São Paulo, Brazil (22◦463.62S 45◦3127.08W). Deposited in the Museu de Zoologia da Universidade Estadual de Campinas (ZUEC), Unicamp, Campinas, São Paulo, Brazil. Labels on the holo-type (five labels separated by transverse bars): /HOLOTY-PUS/Mirante do Picodo Itapeva, Pindamonhangaba, São Paulo: Brazil28.XII.2016,2000m,22◦463.62S45◦3127.08W,L.M. Mag-aldi,leg./Holotypus—ActinotemantiqueiraFreitas,Francini,Paluch &Barbosadet.2017/DNAvoucher–BLU936/ZUECLEP10004/.
Allotype (Fig. 1B). Same data as Holotype. Deposited in the Museude Zoologiada Universidade Estadual de Campinas (ZUEC), Unicamp, Campinas, São Paulo, Brazil. Labels on the allotype (five labels separated by transverse bars): /ALLOTY-PUS/Mirante do Picodo Itapeva, Pindamonhangaba, São Paulo: Brazil28.XII.2016,2000m,22◦463.62S45◦3127.08W,L.M. Mag-aldi,leg./Allotypus—ActinotemantiqueiraFreitas,Francini,Paluch& Barbosadet.2017/DNAvoucher–BLU939/ZUECLEP10005/.
Paratypes(AllfromBrazil). Minas Gerais:Itamonte,Caminho paraAgulhasNegras,ParqueNacionaldoItatiaia,1800m,1male, 4.XII.1991,A. V.L. Freitas leg.(ZUEC-AVLF). SãoPaulo:Piquete, EstradaparaSãoFranciscodosCamposdoJordão,1800m,1male, 5.XII.2004,R.B.Francinileg.(R.B. Francinicol.#22);Campos do Jordão,AltodaBoaVista,1800m,1male,6.XII.2007(DNAvoucher AC110),A.V.L.Freitasleg.;Pindamonhangaba,PicodoItapeva, 1800m,1female,31.XII.2005,A.V.L.Freitasleg.;6males(DNA vouchersAC111,AC112,AC113,AC114,AC115,AC116),6.XII.2007, A.V.L.Freitasleg.(ZUEC-AVLF);MirantedoPicodoItapeva,2000m, 3 males, 28.XII.2016,22◦463.62S45◦3127.08W,L. M. Maga-ldi,leg.(DNAvouchers BLU937,BLU938, BLU940) (ZUECLEP 10006,ZUECLEP10007,ZUECLEP10008)(ZUEC);SantoAntônio doPinhal,EstradaparaoPicoAgudo,1220m,20.XI.2013,22◦511S 45◦3942W,T.S.Souzaleg(ZUECLEP10003)(ZUEC).
Additional studied material (all from Brazil). DZUP – Minas Gerais:Camanducaia,Monteverde,1650m,3malesand6females, 22.XII.1968, 1 male, 23.XII.1968, 4 males,27.XII.1968, H. Ebert leg.DZ3769,DZ3772,DZ3771,DZ5848,DZ5865,DZ6041,DZ 5961,DZ5937,DZ5896,DZ3773,DZ3774,DZ3474,DZ7869, DZ3770;SantosDumont,RioNovo,850m,1male,13.XI.1953,H. Ebertleg. DZ5849;Barbacena,SerradaMantiqueira, 1100m, 1 female,4.XII.1952,H.Ebertleg.DZ7091;Poc¸osdeCaldas,1250m, 1 male, 10.XII.1966, 2 males, 11.XII.1966, 1 male, 15.XII.1966, 3 males,17.XII.1966,H. Ebert leg.DZ3776, DZ3786,DZ3785, DZ3787, DZ,3767,DZ 3788,DZ 2977;Delfim Moreira, 15 Km SE, 1500–1700m, 2females, 22-23.I.2004,Mielke &Casagrande leg. DZ 9290,DZ 9250. Rio de Janeiro: P. N. Itatiaia, 1600m, 2 females,12.I.1973,Mielkeleg.DZ5969,DZ5857;Imbariê,25m, 1female,29.VII.1964,H.Ebertleg.DZ5928;Itatiaia,Oeste,1400m, 2males,22.XII.1957,H.Ebertleg.DZ3782,DZ3110;Teresópolis, 1000m,1female,5.I.1973,Mielkeleg.DZ7997.SãoPaulo: Cam-posdoJordão,1700m,1female,30.I.1966,H.Ebertleg.DZ5945,
CamposdoJordão,Toriba,2males,12.XI.1922,1male,16.XII.1952, DAlmeida&L.Travassos Filholeg.DZ3781,DZ3762,DZ2974; Bananal,Bocaina,2malesand2females,2.I.1937,Travassosleg.DZ 3780,DZ3779,DZ5913,DZ5904.IOC–MinasGerais:Passa Qua-tro,FazendadosCampos,1600m,1female,2.XII.1915,1female, 5.XII.1915,J.F.Zikánleg.I.O.C.no25195,No.25196.RiodeJaneiro: Itatiaia,CampoBelo,2females,10.XII.1919,1female,12.II.1920, km12,1female,20.I.1925,J.F.Zikánleg.I.O.C.No.25193,No.25177, No.25198,No.25217,1female,30.XI.1920,1female,29.XII.1920, M.Zikánleg.I.O.C.No.25176,No.25178.SãoPaulo:Bananal,3males and3females,8.I.1937,L.TravassosFilholeg.Coll.Travassos.MNRJ –RiodeJaneiro:NovaFriburgo,1female,III.1934,Col.J.Oiticica Filho.SãoPaulo:CamposdoJordão,3females,29.I.1933,Travassos &J.OiticicaFilho,Col.J.OiticicaFilho;Bananal,Bocaina,3females, I.1937,A.Costaleg.Col.Dr.A.CostaNo.4376,No.4377,No.4365. ZUEC-AVLF– MinasGerais:Andradas,PicodoGavião,1600m,1 male,12.XII.2017,22◦11S46◦3737W,A.V.L.Freitas,L.M. Maga-ldi,J.Y.O.Carreira&A.Taciolileg.;Poc¸osdeCaldas,SerradoCristo, 1560m,2males,15.XII.2017,L.M.Magaldi&A.Taciolileg.São Paulo:Silveiras,pontesobreoRiodoHigino,SerradaBocaina,2 males(DNAvouchersAC176,AC180)and2females(DNAvouchers AC174,AC179),17.I.2009,K.L.Silva-Brandãoleg;SerradaBocaina, 1450m,2males,19.XII.2017,22◦4754S44◦4248W,L.M. Maga-ldi,A.Tacioli&A.H.B.Rosaleg.
Immaturestages
Egg(Figs.5A,B,8C).Lightyellowwhenfirstlaid,changing
gradu-allytopinkishredafter24h;barrel-shapedwith13–16verticalribs andseveral(∼14–17)weaklymarkedhorizontalribs;meanheight 0.683mm(range0.67–0.71mm,n=3),meandiameter0.412mm (range0.37–0.44mm,n=9).Aeropyleslocalizedonlynearbaseof egg,intwoirregularrows(noteveryverticalribbearsanaeropyle). Firstinstar(Figs.6and7A).Headbrown,smooth,withoutscoli, meanwidth0.33mm(range0.29–0.36mm,n=3);bodypalecream, withoutscoliandwithlongpalesetaearisingfrompinacula;legs palebrown,prolegspale,analplatepalebrown.Prothoracicplate paleanddifficulttoobserveunderstereomicroscope.T1presents only2subventralsetae,similartoActinotealalia.Headchaetotaxy andbodychaetotaxyarepresentedinFigs.6and7A,respectively. Maximumreportedlength3.0mm.
Lastinstar(Figs.7Band8D).Headdarkbrown, smoothwith thinpalesetaeandwithoutscoli,spinesorchalazae,meanwidth 3.14mm(range3.01–3.24mm,SD=0.073mm,n=4); body dark browndorsally,palecreamlaterallyandventrally,coveredwith mediumsizeddarkbrownscolibearingbrownsetaefromT1to A1 and from A7 to A10, and white setae from A2 to A6; legs black,prolegspalecream;analplatedarkbrown.Maximumlength: 40mm(n=4).ScolidistributionasinFig.7B.Prepupachangescolor, becomingpalecreamwithamorehomogeneouscoloration.
Pupa(Fig.8E).Generalprofileelongated,groundcolorpale yel-lowcreamwithdarkbrownmarkingsinwingcasesandabdomen; abdominalsegmentsmobile,withaseriesoffivepairsof subdor-salblackspinesfromsegmentsA2toA6.Totallength20–21mm (n=2).
Adultandimmaturebehavior,hostplantsandnaturalhistory. LarvaeofA.mantiqueirasp.nov.havebeenrecordedusing Eupa-toriumintermediumandEupatoriumsp.(aspeciesverysimilarto E.intermedium) (Asteraceae) (Fig. 8A)as hostplants (Penz and
Francini,1996;RBFandAVLFpers.obs.).Femaleswereobserved
ovipositingfrom12:00to02:00PMontheundersideofmature leavesoftheirhostplants(Fig.8B).Ovipositionsarelarge(Fig.8C), varyingfrom409to453eggs(n=4).Newlyhatchedlarvaefirst con-sumedthechorion,andafter3–5hbegantofeedonleaftissue.First instarsfedontheundersideleaftissuebyscrapingtheleafsurface, whilelaterinstarswereobservedconsumingtheentireleaf.Infirst
A
B
C
D
100µm 50µm
100µm 50µm
Fig.5.EggsofActinotemantiqueirasp.nov.(A–lateralview,B–dorsalview)andActinotealalia(C–lateralview,D–dorsalview)inscanningelectronmicroscopy.
Fa F1 C2 C1 A1 A2 S1 S3 AF1 A3 P1 L1 P2 MD2 MD1 AF2 AFa Aa La Pb MDa Pa Sb
Fig.6. ChaetotaxyoffirstinstarheadcapsuleofActinotemantiqueirasp.nov.(setaenomenclatureontherightsideandporenomenclatureontheleftside).
instars,frasspelletsweregluedontotheleafbysilk,notfallingto theground.Larvaewereconsistentlygregariousinallinstars,and allactivities,suchasfeeding,resting,ormovingbetweenleaves, occurredsimultaneously.
Adultswereonlyobservedonsunnydays,quickly disappear-ingwhenweatherconditionsbecamecloudy.Malesbegantofly around10:00AM,usuallyflying2–5mhigh.Femaleswereseldom
observed,andterritorialbehaviorcourtship behavioror copula-tionwerenotperceived.AllknownpopulationsofA.mantiqueira sp.nov.wererecordedinwell-preservedsubtropicalwetmontane forestabove1000maltitude,wheretheclimateapproaches tem-perateconditions,includingcoldwinterswithfrequentfrostsand temperaturefrequentlyfallingbelow0◦C.Maleswereobserved feeding on flowers of Croton urucurana Baill. (Euphorbiaceae),
D1 D1 D1 D1 D1 D1 D1 D2 XD2 XD1 SD1 SD1 SD1 SD1 SD1 SD1 SD1 PP1 PP2 SV2 SV3 SV4 Anal proleg Proleg Leg SD2 SD2 SD2 L2 L1 L1 L1 L1 L1 L1 L2 L2 L2 SV1 SV2 SV1 SV2 SV1 SV1 SV2 SV1 SV1 SV1 D2 D2 D2 D2 D2 D2 Spiracle
A
B
SpiracleProleg Anal proleg
Leg
T1 T2-3 A1-2 A3-6 A7-8 A9 A10
Fig.7. LarvalbodydiagramsofActinotemantiqueirasp.nov.(A)Firstinstarbodychaetotaxy;(B)lastinstarscolidistribution.
Chromolaena punctulata(D.C.) R.King & H. Robins(Asteraceae) and on several other species of forest edge plants. Adults are univoltineinallknownpopulations,withonlyoneflightperiod duringthewarmermonths,fromlateNovembertoearly Febru-ary.AdultsofA.mantiqueirasp.nov.aresexuallydimorphic,with maleshavingadeeporange coloration(Figs.1A, 2A,B,9A), and femalesbeingmuchmoretranslucentandsometimeshavingthe subapical spoton theforewing light cream (Figs. 1B, 2C,D). In moststudied sitesindividuals were never abundant,with usu-ally 5–30 individuals observed in a typical day of field work (4–5hof observation),most ofwhich weremales. Thisspecies ispartofthe“orangishredmimicrycomplex”ofActinote(sensu
Francini,1989)(Fig.9),andissympatricwithfiveotherActinote
co-mimics,namelyActinotebonitaPenz,1996(Fig.9B),Actinote conspicua Jordan,1913(Fig.9C), Actinotequadra(Schaus, 1902) (Fig.9D),ActinotedalmeidaiFrancini,1996(Fig.9E),andActinote surima(Schaus,1902)(Fig.9F)andonepieridco-mimic,Dismorphia melia(Godart,[1824])(Pieridae:Dismorphiinae)(seeDiasetal.,
2016).
Habitat.Inallknownlocalities,thespeciesisassociatedwith well-preservedmontaneombrophilicforest,usually ataltitudes above1000m.Malesandfemalesareeasilyobservedinforestedges andinareasofcontactbetweenforestandhighaltitudenatural grasslands(Fig.10AandB).
Geographic distribution. The species is known from the Serra da Mantiqueira region, from the Serra da Bocaina and Serra dos Órgãos, at altitudes from 1000 to 2000m (Fig.11).
Etymology. Mantiqueira is a word in native Tupi language meaning“raindrops”(fromamana=rainandtykyra=drop)andis thenameofalargemountainrangeinSEBrazil(Serrada Man-tiqueira),wheremostoftheknownpopulationsofA.mantiqueira sp.nov.occur.
RedescriptionofActinotealalia(C.Felder&R.Felder,1860) AcraeaalaliaC.Felder&R.Felder,1860:105.Brazil(NHMUK);
Kirby, 1871:136; Müller, 1877:219; Lamas, [1997]:31 (includes
LECTOTYPEdesignation:male[southeastern]Brazil).
Acraea eulalia [sic]; Müller, 1878:296; R. Llano & M. Llano,
1973:70.
Actinote alalia; Mabilde, 1896:64; D’Almeida, 1935:71, 93;
Lewis,1973:227; Lewis,1975:227; D’Abrera,1987:445; Ackery,
1988:136; Mielke, 1994:769; Lamas, 1997:31; Penz &Francini, 1996:309;Paluchetal.,2003:573;Lamas,2004:263;Paluchetal., 2005:416;D’Abrera,2006:247;Iserhardetal.,2010:313,318,319;
Dolibainaetal.,2011:351.
Actinote alalia alalia; Eltringham & Jordan, 1913:10; Jordan, 1913:374;Jordan&Eltringham,1916:19.
Acraea(Actinote)alalia;Pierre,1987:22.
Actinotesp1.;Silva-Brandãoetal.,2008:519,528,figs.3,4,5. Description
Male (Figs. 2E, F, 9G): Antenna black, 11–12mm in length
A
B
C
D
F
G
E
15.0mm 6.0mm 2.5mm 8.0mm 8.0mm 15.0mmFig.8.NaturalhistoryofA.mantiqueirasp.nov.(A–E)andActinotealalia(F–G).(A)ShrubofEupatoriumintermedium,hostplantofA.mantiqueira;(B)FemaleofA.mantiqueira sp.nov.ovipositing;(C)closeviewofanovipositionofA.mantiqueirasp.nov.;(D)LastinstarofA.mantiqueirasp.nov.(lateralview);(E)PupaofA.mantiqueirasp.nov. (latero-ventralview);(F)LastinstarofA.alalia(lateralview);(G)FemaleofA.alaliaovipositing(photobyLucasA.Kaminski).
form a well-defined club. Forewing narrow and elongated, length25–28mm(mean=26.8mm,SD=1.09mm,n=5);hindwing rounded,length18–21mm(mean=19.8mm,SD=1.09mm,n=5). Bodydark brown, abdomenventrally covered by creamscales. Forewingbackground dark orange with dark brownveins and stripesasfollows:abroaddarkbrownmargin,broadinapexand narrowingtowardtornus;spacefromanalmarginto2Adarkbrown infinalthirdonly;fouradditionalnarrowdarkbrowntransverse stripes,firstextendingfromcostatoCuA2,twothirdsfrombase, secondcrossingend ofdiscalcellfromRsectortoM3,third v-shapedcrossingdiscalcellin midportion,and fourthv-shaped crossingspaceCuA2-2Ahalffrombase.Hindwingbackgrounddark orangewithbroaddarkbrownmargin;allveinsdarkbrown,and narrowdarkbrownstripesextendingininter-venalspaces. Trans-versepatches ofdarkscalescrossingproximally spacesM1-M2, M2-M3,M3-CuA1andCuA1-CuA2.Ventralforewingwithapattern verysimilartodorsalpattern,butlackingbroaddarkbrown mar-gin;ventralhindwingbackgroundcream,withawell-markeddark
brownv-shapedbandfromSc+R1 to3A,withanadjacentwell definedbroadv-shapedorangeband.Discalcellusuallycrossedby anobliquedarkbrownstripe,defininganorangebasalhalfanda creamdistalhalfregion.
Malegenitalia(Fig.3I–O):Valvaeelongateandcurved,length aboutfourtimeswidthofmedianportion;broaderatbase;apex roundedwithaconspicuouscurltowardmiddlelineofbody.In dor-salview,basalportionofuncuswide,abruptlynarrowing,ending inapoint.Tegumenlongandbroad,trapeze-shapedwithaslight constrictionatmiddle.Gnathosabsent.Inventralview,saccuswith anabruptslightnarrowingnearbase,thenprojectingasabroad lobewithroundedend;abouthalflengthofgenitalcapsule. Aedea-gusabouthalflengthofgenitalcapsule,endinginasharppoint,in lateralviewslightlycurveddownwards.Juxtabroad,aboutsame widthofbaseofonevalva(inventralview),diamond-shapedwith ananteriorshortpointedprocess;lengthequaltowidth.
Female(Figs.2G,H,8G).Antennablack,13mminlength(n=2), extendingtomid-costa, with41segments,11 of which forma
Fig.9.ComparativeplateshowingallknownBrazilianspeciesofActinotebelongingtothe“orangishredmimicrycomplex”,dorsalleftandventralright(malesonly).(A) Actinotemantiqueirasp.nov.(paratype),(B)Actinotebonita,(C)Actinoteconspicua,(D)Actinotequadra,(E)Actinotedalmeidai,(F)Actinotesurima,(G)Actinotealalia,(H)Actinote catarina.
well-defined club. Forewing narrow and elongated, length 31–33mm (n=2). Hindwing rounded, not translucent, length 23mm(n=2).Colorpatternoffemalewingssimilartothatofmales butpaler.
Femalegenitalia(Fig.3P):Corpusbursaerounded,signaabsent. Ductus bursae not sclerotized, same length as corpus bursae; sterigmarounded.
Taxonomyandvariation
ActinotealaliawasdescribedbyC.Felder&R.Felder(1860)based onanunstatednumberofspecimenswithoutaspecifiedlocalityin Brazil.AmalesyntypedepositedintheNHMUK(figuredinWarren etal.,2017),wasexaminedbyGerardoLamasanddesignatedas thelectotypebyhim(Lamas[1997]).Basedontheprovenanceof mostoftheFelder’smaterial,Lamas([1997])suggestedthatthis specimenshouldhavecomefromsoutheasternBrazil.Indeed,the wingpatternoftheabovelectotypecorrespondstoActinotealalia populationsfromsouthern Brazilinterms ofthefollowing visi-blecharacters:(1)thenarrowtransversedarkbrownbandsonthe dorsalforewing;(2)amostlyorangebandintheinternalmargin ofthedorsalforewing;(3)aconspicuousorangev-shapedband transversallycrossingtheventralhindwing,and (4)a complete
darkbarcrossingthemiddle ofthediscalcelland delimitinga basalorangearea(seeFig.2).Basedontheseobservationsandthe datapresentedinthispaper,thetypelocalityofA.alaliashouldbe changedto“southernBrazil”.Ashasbeenreportedformostspecies ofActinote,A.alaliaalsopresentsintraspecificvariation,especially inthefemalecolorpattern(malesaremuchmoreuniform).Asfor A.mantiqueirasp.nov.,minorvariationhasalsobeenreportedin wingvenationandgenitalia.Themostvariablecharactersarethe extentofthehumeralvein,whichcanreachornotthewingedge, andtheshapeofsaccusapex,thatcanvaryfrompointedtoalmost rounded.
Examinedmaterial(allfromBrazil).DZUP–Paraná:Curitiba, 900m,1maleand2females,28.II.1969,Mielkeleg.DZ3153,DZ 5864,DZ5905,1female,13.XII.1968,Mielkeleg.DZ3119;Palmas, 1100m,7malesand1female,6.II.1976,Mielke&Buzzileg.DZ 3136,DZ3161DZ3133,DZ3132,DZ3121,DZ3129,DZ3160,DZ 5929,2males,I.1930,Stawiarskileg.DZ3765,DZ3784; Pruden-tópolis,1200m,10malesand1female,2.II.1976,Mielke&Buzzi leg.DZ3106,DZ3128,DZ3157,DZ3137,DZ3116,DZ3120,DZ 3108,DZ3148,DZ3134,DZ3138,DZ5872;Castro,1000m,1male and2females,25.I.1971,Mielkeleg.DZ3147,DZ6017,DZ6001; Guarapuava,1200m,2malesand1female,18.II.1978,Mielke& Miersleg.DZ3152,DZ3122,DZ5888;PortoUnião,1maleand
Fig.10.(AandB)HabitatofActinotemantiqueirasp.nov.;(CandD)HabitatofActinotealalia.(A)GeneralviewofthemontaneforestsinCamposdoJordão,SãoPauloState (1400–1600ma.s.l.);(B)regionofcontactbetweengrasslandandhighaltitudeforest,PicodoItapeva,Pindamonhangaba,SãoPauloState(1800ma.s.l.);(C)generalviewof themontaneforestsinSãoFranciscodePaula,RioGrandedoSulState(900–1100ma.s.l.);(D)closeviewofthemontaneforestinSãoFranciscodePaula,RioGrandedoSul State(900ma.s.l.);thecharacteristicAraucariatreesarevisiblenearthetrail(blackarrow).
BRAZIL 0 10 20 30 40 50 60 Tropic of capricorn Atlantic ocean 0 - 800 m 800 - 1000 m > 1000 25ºS A. mantiqueira sp.nov. A. alalia 0 200 Kilometers 45ºW 40ºW
Fig.11. RecordeddistributionofActinotemantiqueirasp.nov.(opencircles)andActinotealalia(solidcircles)inBrazil.LocalitydataincluderecordscompiledfromPaluch (2006),museumdata(seetext)andrecentsightings(AVLF,unpublisheddata).
1female,Stawiarskileg.DZ3775,DZ6071;SãoJosédosPinhais, ColôniaMuricy,1male, 14.XII.2001,Paluch leg.DZ4758; Lapa, 1male,nodata,H.Ebert leg.DZ3768;PontaGrossa, 1female, IV.1956,Coll.F.Justus,DZ8014.SantaCatarina:RiodasAntas,4 malesand2females,I.1953,Camargoleg.DZ3764,DZ3763,DZ 3778,DZ3777,DZ5912,DZ6009;SantaCecília,1000m,1male, 22.II.1973,Mielke leg.DZ3105;PonteAltodo Norte, 1000m, 1 male,12.II.1973,Mielke&Sakakibaraleg.DZ3783;SãoJoaquim, 1250m, 1female,24.II.1973,Mielkeleg.DZ 5897,São Joaquim,
Mantiqueira,8females,26.II.1973,Mielkeleg.DZ5936,DZ5881,DZ 5873,DZ5993,DZ5921,DZ5977,DZ6025,DZ5985,SãoJoaquim, PlanaltodeLages,1300m,1female,2-4.II.1973,H.&H.D.Ebert leg.DZ 5953;Lages, Painel,1000m, 1female,24.II.1983,Mielke &Casagrandeleg.DZ5856,1female,23.II.1973,Mielkeleg.DZ 6033,Lages,ParquePedrasBrancas,920m,2females,13.II.1973, Mielke&Sakakibaraleg.DZ5920,DZ6066.RioGrandedoSul:Serra Geral,Canela,800m,1male,22.I.1973,H&H.D.Ebert,H.DZ3766. ZUEC–SantaCatarina:BomJardimdaSerra,1maleand1female,
1360m,11.I.2011,L.A.Kaminskileg.ZUEC-AVLF–RioGrandedoSul: SãoFranciscodePaula,FlorestaNacionaldoPinho,4males(DNA vouchersAC117,AC118,AC119,AC120),31.XII.2007,L.A.Kaminski leg.;1male,1.I.2009,L.A.Kaminski;1female(DNAvoucherAC210), 7.XII.2009,L.A.Kaminskileg.MZSP–Paraná:Lapa,2males,1female, XI.1940,B.Pohlleg.SantaCatarina:RiodasAntas,1female,I.1953, Camargoleg.
Immaturestages
Egg(Fig.5CandD).Lightyellowwhenfirstlaid,changing grad-uallytopinkishredduringthefirst24h;barrel-shapedwith13–15 vertical ribsand several(∼12) weakly marked horizontal ribs; meanheight0.72mm(range0.71–0.73mm,n=3),mean diame-ter0.42mm(range0.24–0.52mm,n=3).Aeropyleslocalizedonly nearbaseofegg,intwoirregularrows(noteveryverticalribbears anaeropyle).
Firstinstar.Headlightbrown,smooth,withoutscoli;bodypale cream,withoutscoliandwithlongpalesetaearisingfrompinacula; legspalebrown,prolegspale,analplatepalebrown.Prothoracic plate pale and difficult to observeunder stereomicroscope. T1 presentsonly2subventralsetae,thesameconditionasinActinote mantiqueirasp.nov.
Lastinstar(Fig.8F).Headdarkbrown,smoothwiththinpale setaeandwithoutscoli,spinesorchalazae;bodybluishdorsally, greenventrally,withaconspicuouslateralband,thisisorangein thoracicsegmentsandcreaminabdominalsegments.Coveredwith mediumsizedscolibearingbluishsetae;scoliaredarkinT1-T2and A9-A10andbluishinT3throughA8;legsblack,prolegsgreen;anal platedarkbrown.
Pupa.Generalprofileelongated,groundcolorpaleyellowwith darkbrownmarkingsinwingcasesandabdomen;abdominal seg-mentsmobile,withaseriesoffivepairsofsubdorsalblackspines fromsegmentsA2toA6.
Adultandimmaturebehavior,hostplantsandnaturalhistory. OvipositionofA.alaliawasrecordedonGrazieliaserrata(Spreng.)R. M.King&H.Rob(Asteraceae).Femaleswereobservedovipositing attheendoftheafternoon,approximately05:00PM.(L.A. Kamin-ski,pers.comm.)ontheundersideofmatureleavesoftheirhost plants(Fig.8G).Ovipositionsarelarge,varyingfrom411to442eggs (n=2).Newlyhatchedlarvaefirstconsumedthechorion,andafter 3–5hbegantofeedonleaftissue.Firstinstarsfedonthe under-sideleaftissuebyscrapingtheleafsurface,whilelaterinstarswere observedconsumingtheentireleaf.Inthefirstinstar,frass pel-letsweregluedontotheleafbysilk,notfallingtotheground.First instarsweregregariousandallactivities,suchasfeeding,resting, ormovingbetweenleaves,occurredsimultaneously.
AsreportedforA.mantiqueirasp.nov.,adultsofA.alaliawere onlyobservedonsunnydays,quicklydisappearingwhenweather conditionsbecamecloudy.Malesbegantoflyaround10:00AM, usually flying3–5mhigh. Females wereseldom observed, and territorial behavior courtship behavior or copulation were not perceived.AllknownpopulationsofA.alaliawererecordedin pre-servedsubtropicalwetmontaneforestabove800maltitude,where theclimateapproachestemperateconditions,includingcold win-terswithfrequentfrostsandtemperaturefrequentlyfallingbelow 0◦C. Maleswereobservedfeedingonflowersofseveralspecies offorestedgeplants.Adultsareunivoltineinallknown popula-tions,withonlyoneflightperiodduringthewarmermonths,from DecembertoFebruary.AdultsofA.alaliaaresexuallydimorphic, withmaleshavingadeeporangecoloration(Figs.2E,F,9G),and femalesbeingmuchmoretranslucent(Fig.2GandH).Inmoststudy sitesindividualswereneverabundant,withusually2–6 individu-alsobservedinatypicaldayoffieldwork(4–5hofobservation), mostof whichweremales.Thisspeciesispartofthe“orangish redmimicrycomplex”ofActinote(sensuFrancini,1989)(Fig.9),
and is sympatric withthree other Actinote co-mimics, namely, Actinotedalmeidai(Fig.9E),Actinotesurima(Fig.9F)andActinote catarinaPenz,1996(Fig.9H)andonepieridco-mimic,Dismorphia melia(Godart,[1824])(Pieridae:Dismorphiinae)(seeDiasetal.,
2016).
Habitat.Inallknownlocalities,thespeciesisassociatedwith preservedmontanemixedforest(alsoknownasAraucariaForest duetothepresenceofParanápineAraucariaangustifolia),usually inaltitudesabove800m.Malesandfemalesareobservedin for-estedgesandinareasofcontactbetweenforestandhighaltitude naturalgrasslands(Fig.10CandD).
Geographicdistribution.Thespeciesisknownfromthe moun-tainsofsouthernBrazilintheStatesofParaná,SantaCatarinaand RioGrandedoSul,ataltitudesfrom800to1400m(Fig.11).
Discussion
Duetothestrongsimilaritiesinwingpattern,high intraspe-cificvariationandlowdifferentiationinfemalecolorpattern,some species of Actinote are difficult to tell apart (D’Almeida, 1935;
Francini, 1989;Paluch, 2006), andseveralcomplexes of cryptic
specieshavebeenrecognizedwithinthegenus,asisthecaseofthe “orangishredmimicrycomplex”(PenzandFrancini,1996). Conse-quently,severalnewspecieshavebeenrecognizedanddescribedin recentyearsthroughouttheNeotropics(PenzandFrancini,1996;
Francini etal., 2004;Paluch etal., 2006;Neild, 2008;Willmott
etal.,2009,2017).Remarkably,whilesomeofthesenewspecies
arerareandlocalized,andknownfromnomorethan10 individ-uals(e.g.ActinoteebertiFrancini,Freitas&Penz,2004),somehave proventobecommonandwidespread,suchasActinotepratensis Francini,Freitas&Penz,2004.Thislastspecieshasbeenobserved inseveraladditionallocalitiessinceitsdescription,andwasfound breedinginabackyardatCampinasStateUniversity(Freitas,pers. obs.).Conversely,whileadultscanbehardtodistinguish, caterpil-larsaresufficientlydistincttopermitpositiveidentificationstothe species-level,asearlynoticedbyD’Almeida(1935)andafterwards
byFrancini(1989).
In thepresent case, both adults and immature stages of A. mantiqueirasp.nov.aredistinctenoughtosupportthe descrip-tionofthisnewspecies,alsoallowingeasydifferentiationfromA. alalia.Inaddition,A.mantiqueira sp.nov.iscommonand abun-dant in montane forests above 1200m in southeastern Brazil, facilitatingtheacquisitionofsufficientinformationtoconfidently determine itstaxonomic status as a newspecies distinct from A.alalia.
Basedonapreviousmolecularstudy,A. mantiqueirasp.nov. andA.alaliaaresisterspecieswithalowlevelofmolecular
distinc-tion(Silva-Brandãoetal.,2008),andpreliminaryanalysissuggest
thatthesetwospeciesdivergedinLatePleistocene(Silva-Brandão and Freitas unpublished; L. M. MagaldiPhD, inprep.). Climatic changes inthe Pleistoceneareconsideredimportant toexplain thegeographicdistributionanddiversificationoftheNeotropical faunaandflora(Vuilleumier,1971;Simpson,1979;Whitmoreand
Prance,1987),andsimilarexamplesofcloselyrelatedspeciesof
LatePleistoceneoriginthatareendemictodistinctBrazilian moun-tainrangesareknownforbirds(Mataetal.,2009;Freitasetal., 2012),frogs(Leiteetal.,2008),mammals(Gonc¸alvesetal.,2007), andotherbutterflies(AVLF,inprep.).However,additionalstudies areneededtoinvestigatefurtherthisissue.
Insummary,thepresentstudyisagoodexampleofhow increas-ingknowledgeofimmaturestagesand naturalhistorycanhelp betterunderstandingbiodiversity, and in thepresent case, this informationhashelpedintherecognitionofanew,undescribed speciesoflocallycommonbutterfly.
Conflictsofinterest
Theauthorsdeclarenoconflictsofinterest. Acknowledgments
WethankCristianoA.Iserhard,LucasA.Kaminski,JuniaY.O. Carreira,LuizaM.Magaldi,KarinaL.Silva-Brandão,ThadeuS.Souza, AndréTacioliand AugustoH. B.Rosafor providing fresh speci-mensand/orhelpinginfieldwork.TamaraM.C.Aguiarforhelping spreadingthespecimens(includingpartofthetypeseries).Carla Penzreadandcriticizedanearlyversionofthemanuscript and LucasKaminskiandKeithWillmottcarefullyreadthelastversion ofthemanuscript. FábioRaposodoAmaral helpedwith discus-sionsaboutbiogeography andpaleoclimate.EPBthanksFAPESP fora PhD fellowship(2012/03750-8)and apost-doc fellowship (2016/15873-8).MP thanksDr. Gerdt Guenther Hatschbach (in memoriam)andOsmardosSantosRibas(MuseuBotânico Munic-ipal, Curitiba – PR) by the identification of the host plant of Actinote alalia, and to Dr. Jorge Manuel Saraiva Bizarro by the greatincentiveand helpin fieldworkin Paraná.RBFthanksto UniversidadeCatólicadeSantosforlogisticsupport.AVLF acknowl-edgessupportfromFAPESP(Biota-FapespGrants2011/50225-3, 2013/50297-0)andfromtheBrazilianResearch Council–CNPq (Grants302585/2011-7and 303834/2015-3).Thispublicationis partofthe“RedeLep–RedeNacionaldePesquisaeConservac¸ão deLepidópteros”–SISBIOTA-Brasil/CNPq(563332/2010-7). References
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