Psyllobora picta (Germain) species complex (Coleoptera: Coccinellidae), with descriptions of two new species from Chile

REVISTA

BRASILEIRA

DE

Entomologia

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

Systematics,

Morphology

and

Biogeography

Psyllobora

picta

(Germain)

species

complex

(Coleoptera:

Coccinellidae),

with

descriptions

of

two

new

species

from

Chile

Guillermo

González

_,

_Lúcia

_M.

_Almeida

a_{LaReina,Santiago,Chile}

b_{UniversidadeFederaldoParaná,DepartamentodeZoologia,LaboratóriodeSistemáticaeBioecologiadeColeoptera,Curitiba,PR,Brazil}

a

r

t

i

c

l

e

i

n

f

o

Received21November2016 Accepted4August2017 Availableonline19August2017 AssociateEditor:AdrianaMarvaldi Keywords: Coccinellini Genitalapparatus Newspecies SouthAmerica Taxonomy

a

b

s

t

r

a

c

t

ThePsylloborapictaspeciescomplex(Coleoptera:Coccinellidae:Coccinellini)wasrevised,withthe descriptionoftwonewspecies,Psylloboraluerisp.nov.andPsylloborapaulinesp.nov.,bothfromChile. PsylloborabicongregatawasrecordedforthefirsttimefromBrazil,andthereisnoconfirmedrecordof thisspeciesforChile.Also,P.pictawasnotrecognizedfromParaguayandUruguay.Maleandfemale genitaliacharactersareillustratedforallthespecies.

©2017SociedadeBrasileiradeEntomologia.PublishedbyElsevierEditoraLtda.Thisisanopen accessarticleundertheCCBY-NC-NDlicense(http://creativecommons.org/licenses/by-nc-nd/4.0/).

Introduction

Membersof thegenusPsylloboraChevrolataredistinguished amongthecoccinellidbeetlesbytheirlightcolors,white,yellow orbrownandwithdarkspots,rangingfromlightbrowntoblack. Thecharacteristicmultidentatemandiblesareassociatedwiththe micophagoushabit,sharedwithasmallnumberofgenera previ-ouslygroupedinthetribeHalyziini.Thegenushasaworldwide distribution,althoughmostofthespeciesareNeotropical.

Most of the South American species were described in the nineteenthandthefirsthalfofthetwentiethcenturies,withan additional 17 species described since then, totaling 55 species (Blackwelder, 1945;Mader,1957;AlmeidaandMarinoni,1983; Almeida,1985,1991a,1991b,1991c,1992).

Exceptforthespeciesdescribedorrevisedinrecentstudies, whichcoveredlessthanhalfoftheknowntaxa,thegenital appara-tusofmaleshasnotbeenstudied.Traditionally,elytralspotshave beenusedforspeciesidentification,althoughthischaracterisnot alwaysstableoraccurate.

Psylloborapicta(Germain)hasbeencitedfromseveralcountries ofSouth America andlinkedto otherspeciesof thegenus dis-tributedinthesameareas.ThehistoryofspeciesrelatedtoP.picta beganwithitsdescriptionbyGermainin1854.Soonthereafter,

∗ Correspondingauthor.

E-mail:lalmeida@ufpr.br(L.M.Almeida).

Boheman(1859)describedPsylloborabicongregataBohemanfrom Argentina.Mulsant(1866),basedonChileanspecimens,described Psylloboraferalis,whichPhilippi(1887)incorporatedintohis cat-alogof beetlesin Chileas Halyciafemoralis(sic). Crotch(1874)

recognizedP.bicongregatafromArgentinaandP.feralisfromChile anddidnotmentionP.picta.Weise(1904)recognizedthepresence ofP.feralisandP.bicongregataforArgentinaanddescribed Psyllob-orapavidaWeisefromthatcountry.Brèthes(1923)synonymized theChileanspeciesunderthenameP.picta(=P.feralis=P.femoralis).

Korschefsky(1932)synonymizedP.bicongregatawithP.pictaas well,adecision followedbyCostaLima(1937)andBlackwelder (1945).

Finally,Mader(1957)revisingspecimensfrombothcountries recognized them as the two different species P. picta (=P. fer-alis,=P.femoralis)andP.bicongregata(=P.pavida),establishingP. pictainChile,whileconsideringP.bicongregataasalsofoundin ArgentinaandUruguay,butwithdoubtfulrecordsfromChile. How-ever,Gordon(1987),whoreviewed theCrotchcollection,again consideredP.bicongregataassynonymouswithP.picta,basedon

Korschefsky(1932), and probablydidnotconsider theworkof

Mader(1957).González(2008,2010,2013,2014)andSerraetal. (2013)consideredthesetwospeciesaspresentinChile,Argentina, ParaguayandUruguay,basedontheliterature.

These species share similar characteristics in the pattern of spots and in thegenital parts, and are referred to herein as a species complex of P. picta. González (2006) conducted analy-sesanddissectionsofspecimensfromseveralcollectionsofthis

http://dx.doi.org/10.1016/j.rbe.2017.08.002

0085-5626/©2017SociedadeBrasileiradeEntomologia.PublishedbyElsevierEditoraLtda.Thisis anopenaccessarticleundertheCCBY-NC-NDlicense(http:// creativecommons.org/licenses/by-nc-nd/4.0/).

group.Thisinformation, togetherwithobservations inthe cur-rent study of the genital parts, geographical distribution and morphological aspects, have made it possible to recognize the describedspecies,aswellastoestablishtheirgeographical dis-tributionandtodescribenewspeciesofthiscomplexwithsome confidence.

Materialandmethods

Theterminology for adultstructures,especially for the gen-italia,corresponds tothat usedby ´Slipi ´nski (2007).The genital partsofallspeciesincludedinthisstudywerepreservedin glyc-erininmicrovialsalongwiththerespectivespecimen.Themethod of immersing the entire abdomen in a solution of 10% KOH, heatedby thermal convectionin a test tubeimmersed in boil-ingwaterwasused.Theabdomenwasthenwashedindistilled waterandthegenitalstructurewasseparatedundera stereomi-croscope with thehelp of pins. All specimens and parts were photographed.

Thelengthofthepenisguideismeasuredfromtheapextothe insertionof theterminal strut,andthemaximumlengthofthe penisismeasuredfromtheapextothefarthestpointofthepenis tube.

The material studied belongs to the following collections: CFDH:Federicod’Hervéprivatecollection,RioNegro,Argentina; CPAA:AlfonsoAguilera,privatecollection,Temuco,Chile;CPAL: Alfredo Lüer, private collection, Santiago, Chile; CPCF: Claudia Funes,privatecollection,Catamarca, Argentina;CPFR:Francisco Ramírez, private collection, Santiago, Chile; CPGG: Guillermo González, private collection, Santiago, Chile; CPMD: Manuel Diéguez,privatecollection,Santiago,Chile; CPRH:Richard Hon-our,privatecollection,Santiago,Chile;CPSL:SilvioLanati,private collection,Mendoza,Argentina;CPUD:UlfDrechsel,private col-lection, Asunción,Paraguay; DZUP: Colec¸ãoEntomológica Pe. J. S. Moure, Departamento de Zoologia, Universidade Federal do Paraná,Curitiba,Brazil(LúciaM.Almeida);JEBC:JuanEnrique Bar-riga,privatecollection,Curicó,Chile;MAPA:MuseuAnchietade CiênciasNaturais, Porto Alegre,RS,Brazil (FernandoR.Meyer); MHNP: MuseoNacionaldeHistoriaNaturalde Paraguay, Asun-ción(JohnKochalka);MHNS:MuseoNacionaldeHistoriaNatural de Santiago, Chile (Mario Elgueta); MNRJ: Museu Nacional do RiodeJaneiro,Brazil(MiguelA.Monné);UMCE:Colección Ento-mológica.DepartamentodeBiología.UniversidadMetropolitana deCienciasdelaEducación(DaniloCepeda);UNAB:Museo Ento-mológicoFacultaddeAgronomía,UniversidadNacionalde Colom-bia, Sede Bogotá, Bogotá, Cundinamarca, Colombia (Francisco Serna).

The labels of the type materials are arranged in sequence from top to bottom, where the data for each label are within double quotes (“”), a slash (/) separates the rows, and infor-mation between brackets ([]) provides the source and deposit collection.

Results

SpeciescomplexassociatedwithPsylloborapicta(Germain) SpeciesassociatedwithP.pictaarecharacterizedbytheirsize (1.6–3.1mm),pronotalplatewithfourspotsinasemicirclearound afifthsmallspotnearthemiddleofthebase,sometimesoneortwo spotsoneachside,allindependentorfusedtogether.Theelytral patternconsistsofapproximatelytenspots(3:1:2:1:2:1),always recognizablebut sometimespartially fused,separate orabsent, or smallsupernumerary spots are present (Figs. 1, 19, 26, 40).

3 4 1 2 5 10 11 12 13 14 15 17 16 18 21 22 23 14 25 20 19 9 8 7 6

Figs.1–25.1–14:Psylloborapaulinesp.nov.:1–5:maleholotypehabitus(dorsal, lateral,frontal,posteriorandventraldetail);6–9:variations(dorsalview);10:male sipho;11–13:maletegmen(dorsal,lateralandbasallobedetail);14:female gen-italia(spermathecaandinfundibulum).15–25:Psylloboraluerin.sp.:15–18:male holotypehabitus(dorsal,lateral,frontalandposterior);19–20:variations(dorsal view);21:malepenis;22–24:maletegmen(dorsal,lateralandbasallobedetail). 25:femalegenitalia(spermathecaandinfundibulum).

The clover-shaped apical spot has three subquadrate points (Figs.4,29).

Themostdistinctivecharacterofthegroupis thevery thick tubeofthemalepenis,itswidth1/6ofthemaximumlength,with thepeniscapsulewithaconvexoutercornerandtheapexabout asthickasthetube,withacharacteristicsub-rhomboidalshape, throughwhichappearsthesinuousejaculatoryduct,andanoblique scleroticthornontheapex.Thestructureofthispenisisnearlythe sameinthedifferentspeciesofthecomplex(Figs.10,21,35,48). Thetegmenindorsalviewhasthesidesofthepenisguidestraight orsinuate,slightlyconvergent,sometimeswithatriangular mamil-liformtip,characteristicineachspecies(Figs.11,22,36,49).In lateralview,therearemembranousventralpenisguideprojections, foldedandcrossinginward,moreorlesswide(Figs12,23,37,50). Twosubapicalsemicircularmembranousandgloboseprojections, shapedasfacetedeyes,arepresentbut aredifficulttoexamine becausetheyareusuallycrushedanddeformed(Fig.38).The sper-mathecaofthefemaleis“C”shaped(Figs14,25,39,52).

Thecolorpatternofthesespeciesisverysimilar,whichexplains why they were synonymized. The characteristics of the penis, whichhasawidthof1/6ofthemaximumlength(notmorethan1/8 inallotherPsylloboraspecies)andthetruncatedapexofthesame thicknessasthetubeareuniquewithinthegenus,anddistinguish thiscomplexfromallotherknownspeciesofPsyllobora.

28 26 29 27 31 32 42 43 40 41 48 49 50 51 52 47 46 45 44 33 34 39 30 35 36 37 38

Figs.26–52.26–39:Psylloborapicta(Germain):26–30:habitus(dorsal,lateral, frontal,posteriorandventraldetail);31–33:variations(dorsalview);34:specimen alive;35:malepenis;36–38:maletegmen(dorsal,lateralandbasallobedetail);39: femalegenitalia(spermathecaandinfundibulum).40–52:Psylloborabicongregata Boheman:40–43:habitus(dorsal,lateral,frontalandposterior);44–47:variations (dorsalview);48:malepenis;49–51:maletegmen(dorsal,lateralandbasallobe detail).52:femalegenitalia(spermathecaandinfundibulum).

KeytospeciescomplexassociatedwithPsylloborapicta

1.Penisguidewide,3.5timeslongerthanwide,lateralmarginsconvergingtotriangularapexwithstronglyconcavesidesformingdistinctmamilliformprojection (Figs.49,51).Dorsalcolorlightbrown,withbrownspotswithdarkedges,annular,rarelywithhomogeneouscolor.Apicalspotsusuallymissingordisaggregatedintosmall irregularspots(Figs.40–47).Argentina,Brazil,ParaguayandUruguay(Fig.53)...PsylloborabicongregataBoheman Penisguideelongated,morethan4.0timeslongerthanwide,sidesstraightandconvergingorsinuate,apextriangularwithslightlyconcavesides,withlittle-differentiated terminalprojection.Dorsalcolorwhitetolightbrown,withblack,dark-brownorlight-brownspotswithorwithoutdarkedges.Apicalspotsseparatedornot.Argentina andChile...2 2.Penisguidenearlystraightwithslightlyconverginglateralmargins(Figs.22,24).Dorsalcolorlightbrown,withbrownspotswithdarkedges,annular,sometimes homogeneous,browntodarkbrown.Elytralspots2–1–4–5–6–7almostalwayslinkedtogether,forming“S”shape.Apicalspotalmostalwayspresent,rarelymissingor disaggregatedintosmallirregularspots(Figs.15–20).Chile(ChileChicoProvince)(Fig.53)...Psylloboraluerisp.nov.

Penisguidesinuate,sideswithevidentnarrowingat2/3oflength.Dorsalcolorwhitetolightbrown,withblack,dark-brownorlight-brownspotswithorwithoutdark edges.Elytralspotsrarelyjointed,especiallyfirstandfourthspotsalwaysseparate,never“S”-shaped.SouthernArgentinaandChile(fromCopiapótoLlanquihueprovinces)3 3.Penisguidewithverywidepigmentedventralprojections,twiceaswideasparamereandsemicircularborder(Fig.37).Dorsalcolorwhitewithblackspots,rarely yellowishorwithdark-brownspots.Pronotalspotssometimesfusedtogether.Spot8ofbothelytrafusedinacommonspot,sometimesspot4alsofusedwithitscounterpart (Figs.26–35).Legsoftenwithbrownorblackareas.Meso-andmetaventritegenerallyblack(Fig.30).SouthernArgentinaandChile(SanAntoniotoLlanquihueprovinces) (Fig.53)...PsylloborapictaBoheman Penisguidewithunpigmentedventralprojectionsofirregularorstraightedge,rarelysemicircular,narrow,atmost1½timeswidthofparamera(Fig.12).Dorsalcolor yellowishbrown,withlight-brownspotswithdarkborder,ringed.Pronotalalwayswithseparatespots.Suturalspots4and8separatefromsuture,rarelytogetherinsuture andneverformingacommonspotwithcounterpart(Figs.1–9).Legsyellowishbrown.Meso-andmetaventritebrown(Fig.5).Chile(fromCopiapótoTalcaprovinces) (Fig.53)...Psylloborapaulinesp.nov.

Psylloborapaulinesp.nov.

(Figs.1–14,53)

Holotype ♂: “CHILE, Prov. [Provincia] Elqui/Tongoy/1-15-octubre-1996/leg.G.González”“♂576”(mv)[MHNS].

Paratypes(9♂and10♀):2♀:samedataofholotype[MHNS]. 2 ♂: “El Melocotón/Cajón del Maipo/Stgo Chile/enero 1977/G.

Fig.53.DistributionmapofPsylloborapictaspeciescomplex.

GonzálezF.”“♂♂567”(mv)[CPGG].2♂and1♀:“FrayJorge/Prov. Limarí/Chile/16-II-1992, leg./G. González F.” “♂♀ 1645” (mv) [CPMD].1♂and 1♀:“Chile,Elquiprov,Que-/bradaElArrayán, 10km/S.LaVilla,Malaisetrap,/400m,16-dic-20033-ene/2004,leg. MEIrvin,30◦/0426S,71◦004W”“♂1647_{“♀1647”(mv)[JEBC].2♀}

Vázquez” “Chile” “Psyllobora/bicongregata/Bohem. 1859./L.M. de Almeidadet. 1990” “♂1655”“♀1655”(mv) [CPAA]. 1♂: “Chile, IV-R/Limarí/LasRojas/26-IX-2006/leg.A.Lüer”(mv)[CPAL].1♂: “Chile,IIIR/Huasco/Vallenar/12-X-2008/leg.A.Lüer”(mv)[CPAL]:1 ♂and2♀:“16-IX-2006Hacienda/LosAndes–Hurtado/RMen algar-robo/Chile,Coll: R.Honour S.”“Macho/146”, “Hembra/142”and “Hembra/190”[CPRH];1♂:“11-II-2008/LaSerena/IVRegión/Chile, Coll: R. Honour S.” “Hembra/142” and “Macho/146” [CPRH]; 1 ♂:“16a19-IX-2006/HaciendaLosAndes/Hurtado-IVRegión/En Prosopischilensis/Chile,Coll:R.HonourS.”“Macho/191”[CPRH].

Additional material (44 specimens): Chile: Copiapó: Bahía Inglesa,1-XI-1994,leg.J.E.Barriga,1specimen[JEBC].Cordillera:El Melocotón,I-1977,G.GonzálezF.3specimens[CPGG].Elqui: Pai-huano,23-VIII-1994,leg.G.González,1specimen[CPGG],Camino Tongoy-Tambillo,6-XII-1985,G.GonzálezF.9specimens[CPGG], Quebrada El Arrayán, 10km. S. La Villa, 400m, 16-XII-2004 a 3-I-2004,leg.MEIrwin,5specimens[JEBC],Rivadavia,24a 26-XII-1946,coll.L.E.Pe ˜na,1specimen[CPRH],Tongoy,1a15-X-1966, leg.G.González,8specimens[CPGG],Vicu ˜na,3-VII-1975,coll.E. Loo,enzapallo,1specimen[JEBC].Limarí:FrayJorge,Quebrada Honda,7a24-XI-2003,legMEIrwinandF.Parker,2specimens [JEBC],FrayJorgePN,ElMineral,224m,12a31-XII-2003,legME IrwinandF.Parker,2specimens[JEBC];LaSerena,11-feb-2008, coll:R.HonourS,1specimen[CPRH];LaSerena,ReservaSanta Gra-cia,feb-2018,coll:R.Honour,1specimen[CPRH],Limarí:Hurtado, HaciendaLosAndes,16a19/IX/2006,coll:R.Honour,enProsopis chilensis, 1 specimen[CPRH]; same data except “en algarrobo” “macho”,3specimens[CPRH];CaminoVicu ˜na–Hurtadokm14,16 a23/IX/2006,coll:R.Honour,1specimen[CPRH];Maipo:Rangue, 6-4-1996,Coll:A.Lüer,1specimen[CPAL];Quillota:Granizo, 17-XI-1990,leg.G.González,1specimen[CPGG].Santiago:II-1964, G.GonzálezF.1specimen[CPGG],QuebradadeMacul.Talca:San Javier,Porter,1specimen[MHNS].

Diagnosis

Thedorsalcoloryellowishbrownwithtenlightbrownspots withdarkborder,ringed,andtheverythicktubeofthemalepenis (itswidth1/6ofthemaximumlength),separateP.paulinefrom themajorityofthegeneraspecies.FromthesimilarlookingP.lueri isdistinguishedbythepenisguidewithsinuatesideswith evi-dentnarrowingat2/3oflengthandtheelytralspotsrarelyjointed, whilefromtheothersimilarlookingspeciesP.bicongregatais dis-tinguishedbythepenisguideelongatemorethan4.0timeslonger thanwide.Otherwise,P.lueriisrestrictedtoChileChicoprovince fromChile,andP.bicongregatatoArgentina,Brasil,Paraguayand Uruguay,regionswhereP.paulineisnotpresent.

Holotypedescription

Male.Length4.0mm;width3.4mm.Bodyoval,withmaximum widthnearmiddleofelytra;yellowishbrown,with5dark-brown spotsonpronotum,elytrawith10dark-brownirregularspots,with darkborders. Legs,antennae and mouthparts yellowish brown. Ventralsurfaceslightlybrownish,metaventritedarkbrown.Dorsal punctationfineandsparse,opaque(Figs.1–5).

Head:Lightbrown,except2dark-brownspotsonfrons. Anten-nae,labrum,maxillarypalpsandlabiumbrown.Eyesrounded,head three timeswidthof eye.Punctation irregular,denserlaterally, wherepointsareseparatedby2ormorediameters(Fig.3).

Thorax: Pronotum light brown, with lateral and anterior borderstranslucentyellow.Smallbrownspotnearbase,with semi-circleoffourbrownspotsaroundcenterspot,twoposteriorspots touchingbaseandtwoanteriorspotsatmidlength,allequidistant fromeachother.Scutellumsmall,triangular,darkbrown(Figs.1,3). Hypomeronandmesoventriteyellowishbrown,metaventritedark

brown.Legsyellowishbrown(Fig.5).Pronotumwithverysparse, almostimperceptiblepunctation.

Elytra:Lightyellowishbrown,withtenbrownspots,withpartly darkenedborders,especiallythosespotsclosesttolateralborder. Spots1and2large,about1/6lengthofelytra,adjacenttobaseand joinedtogetheratmiddle;spot3almostpunctiform,nearhumeral angle;spot4,1/5to2/5length;spots5and6similarinsizetospots 1and2,spot5discalandspot6lateral,attachedtomargin,darker thanremainingspots;spot7immediatelybehind5,connectedat outerborder,formingsinglelarge“C”shapedspot,opentosuture; spot8nearsuture,halfsizeofspot4;spot9smaller,nearlateral border,darker,similartospot6;spot10behind7,largeand clover-leafinshape.Punctationsmall,irregularlydistributed,separatedby aboutfourdiametersonaverage.Epipleuronpaleyellow,strongly descendant(Figs.1–4).

Abdomen:Lightbrownwithfirsttwoventritesbrownin mid-dle;ventrites3–5yellowishinmiddle.Postcoxallinesdiagonal, descending45◦,bentposteriorborder attachedtoit.Ventrite3 slightlytruncate,ventrite6halflengthof5,roundedandweakly notchedincenter.

Malegenitalia:Tegmen4.5timeslongerthanwide, subrect-angularphallobase1.2timeslongerthanwideandwithslightly rounded posterior margin. Terminal strut short, less than half lengthoftegmen.Penisguideparallelatbasal2/5,narrowingin widthfrom7/10to3/5long,apextriangularwithconcavesidesin apical1/9;inlateralviewsubtriangular,sinuous,distal1/3 bend-ingtowardparameres,membranousprojectionsnarrowandwith irregularborders.Parameresindorsalviewnarrow,curvingslightly tophallobase,1/8shorterthanbasallobe,atdistal2/3withlong pubescence,longerthanparamere,inlateralviewwiderindistal third(Figs.11–13).Penismaximumwidthapproximately1/5of maximumlength,basalhalfsemicircular,curvedto2/3oflength, distal1/3straight.Apex1/6oflengthandslightlywiderthanpenis tube,separatedfromitbysclerotizedconnectioninwidelyobtuse angle,withejaculatoryduct“S”shapedandsclerotizedspinenear apex;peniscapsulewithinternalbranchacute,outerbranchtwice aswideastheinternalbranch,basalmarginconvexwithslight notchincenter(Fig.10).

Female:Similartomale.Spermatheca“C”shaped,abouteight timeslongerthanwide,cornushort,slightly expandedatapex, withlateralnodulusandramus1/6oftotallength,veryshort semi-nalduct,anchor-shapedinfundibulumhalfsizeofspermathecal (fig.14).Coxitespaddle-shaped,withverywidepubescentapex anddistinctstylus.

Variation: Length 1.6–2.5mm. The background color varies fromlightyellowtolightbrown.Thespotsarebrownwithdark bordersgivingtheappearanceofrings,orwithoutborders.Rarely, specimenshavehomogeneousdark-brownspots.Thesizeofthe spots variesslightly;sometimestheheadspots canoccupythe entireareabetweeneyes.Thepronotalspotsarealwaysseparate, althoughsometimesthecentralspotmaybemissing;whileelytral spotscansometimesbeincontactwitheach other,butwithout fusing,andtheposteriorspotsaresometimesmissing.Sometimes there aresomesmallsupernumeraryspots indifferentareasof thepronotumandelytra.Somespecimenshaveathindark bor-deratthebaseoftheelytra,alongsutureoralongelytralmargins (Figs.1,6–9).

Discussion

Psylloborapaulinesp.nov.,fromChile,canbeconfusedwithP. bicongregatafromArgentinaandothercountries,duetoitsvery similarcolorpattern,althoughthemalegenitaliaareverydifferent (seespecieskey).OnlyP.pictasharesitsdistributionwithP.pauline

sp.nov.Inadditiontothedifferencesinthegenitaliaandhabitus, indicatedinthekey,bothspecieshavebeenfrequentlyobservedin

natureinareaswheretheirdistributionsoverlap,usuallyingroups of5–30specimensoneachplant,butneverindividualsofthetwo speciesonthesameplant.

Remarks

Psylloborapaulinesp.nov.andP.pictahaveverysimilarmale genitalia,makingsomespecimensdifficulttodistinguishbasedon thischaracter,buttheoverallappearanceisvery different.This situationisnotunusualinspeciesofCoccinellidaeandreinforces thecombineduseofcharacters.AremarkableexampleisAdalia anguliferaMulsantandAdaliadeficiens Mulsant,bothfromChile andArgentina, which have indistinguishablemale genitaliabut areverydifferentintheirhabitus.Incontrasttothepresentcase ofPsylloboraspecies,bothspeciesofAdaliaarefrequentlyfound inabundanceonthesameplant,oftenincopulation,butalways betweenthesamespecies,andarethereforesympatricspecies.

Geographicaldistribution:Chile,CopiapótoTalcaprovinces (Fig.53).

Etymology: This species is named in memory of Pauline América Aguilera Phillips (1970–2015), daughter of Alfonso AguileraPuente,agronomistengineerandspecialistinthestudy ofladybirds ofChile, author of numerous works onthegroup, friendandcollaboratorindevelopingthewebsite“Coccinellidae deChile”.Thenameisproposedasanouninapposition.

Psylloboraluerisp.nov.

(Figs.15–25,53)

Holotype♂:“Chile,XI-R[Región]/Gral.[General]Carrera/Chile Chico/14-2-2005/coll.[leg.]A.Lüer”“♂1657”(mv)[MHNS].

Paratypes(20specimens):1♂:samedataasholotypeexcept “♂148”(mv)[CPAL].1♂and2♀:samedataexceptdate16-2-2005 (mv)[MHNS].12specimens:samedataexceptseveraldatesfrom 12to17–2-2005[CPAL].2♂:“10/02/2016/13kmWChileChico/XI Región/Chileleg.R.HonourS.”“♂♂1733 (mv)[CPRH]and2♀: samedataexcept“♀♀1733”(ab)[CPMD].

Diagnosis

Thedorsalcoloryellowishbrownwithtenlight-brownspots withdarkborder,ringed,andtheverythicktubeofthemalepenis (itswidth1/6ofthemaximumlength),separateP.luerifromthebig majorityofthegeneraspecies.FromthesimilarlookingP.pauline isdistinguishedbythepenisguidenearlystraight withslightly convergingsidesandtheelytralspots2–1–4–5–6–7almostalways linkedtogether,formingan“S”shape,whilefromtheothersimilar lookingspeciesP.bicongregataisdistinguishedbythepenisguide elongatemorethan4.0timeslongerthanwide.OtherwiseP.pauline isrestrictedtoCopiapótoLlanquihueprovincesfromChile,andP. bicongregatatoArgentina,Brasil,ParaguayandUruguay,regions whereP.lueriisnotpresent.

Holotypedescription:Male.Length:2.4mm,width:1.9mm. Ovalwithmaximumwidth nearmiddle of elytra.Back yellow-ish brown, with five dark-brown spots on pronotum and two smallsupernumeraryspots nearborder, elytra withtenbrown spotswithdarker,irregular borders,withspottenwidely scat-tered and a clearer supernumerary spot between spots 5 and 7. Spots 3–2–1–4–5–7–8 joined together, forming an “S” pat-tern.Legs, antennae and mouthparts yellowish brown. Ventral surfacelightbrownwithdark-brownmetaventrite.Dorsal puncta-tioninconspicuous,irregular,surfaceopaque,withoutreticulation (Figs.15–20).

Head:Colorlightbrown,withtwobrownspotsbetweeneyesat insertionofantennasandlargedark-browntransversespotabove eyes,extendinglaterally onvertex.Antennas,labrum, maxillary

palpsandlabiumlightbrown.Eyesroundedfronsthreetimeswidth ofeye.Punctationirregular,denserlaterally,withpointsseparated bytwoormorediameters.

Thorax:Pronotumlightbrown,withanteriorandlateral bor-dersyellowish and translucent. Small central brown spot near base,surroundedwithsemicircle offourbrownspots,two pos-teriorspotstouchingbaseandtwo anteriorspotshalflengthof pronotum,allequidistantfromeach other,andtwosmall,pale, somewhatirregularspotsoneachside.Scutellumsmall,triangular, darkbrown(Figs.15,17).Hypomeresandmesosternumyellowish brown,metaventritedarkbrown.Legsyellowishbrown.Pronotum withalmostimperceptiblepunctation,pointswidelyseparated.

Elytra:Slightlyyellowishbrown,withtenbrownspots,with somewhatdarkerborders(3:1:2:1:2:1)andclearer supernumer-aryspotinspacebetweenspots5,7andsuture.Spot1verylarge, almost¼lengthofelytra,spot2slightlysmallerandspot3even smaller,adjacentandincontactwithbase,connectedtogetherat middle;spot4longitudinal-oval,occupying1/5to2/5oflengthof theelytra,closetosuturebutnottouching,nearspots1and5;spots 5and6nearlyhalflengthoftheelytra,spot5largeandconnected tospots4and7,andspot6separateandhalfdividedlongitudinally, homogeneousdarkbrown;spot7immediatelybehindlargespot 5,semicircular,connectedatouterborder,bothforminglarge“C” shapedspot,inspacebetweentheseandsuture,with supernumer-aryspotmuchclearer,smallandirregular;spot8nearsuture,halfof diameterofspot4,spot9onsametransverselinebutstillsmaller, nearlateralborder,darkbrown;spot10disjointedinthreesmall irregularinconspicuousspots,arrangedintriangle(Figs.15–18). Punctationwithlargedistinctpoints,irregularlydistributed, sepa-ratedbyabouttwodiametersonaverage.Paleyellowepipleuron strongly descendant, with small dark spot in middle of outer border.

Abdomen: Brown, with middle of first two ventrites dark brown.Postcoxallinesdescendingdiagonallyat45◦,benttotouch backborderattachedtoit.VentriteVemarginatearoundposterior margin,ventriteVIasshortashalfof5ventrite,gentlyrounded withshallownotchincenter.

Malegenitalia:Tegmen4.5timeslongerthanwide,phallobase subtrapezoidal,1.1timeslongerthanwideandwithposterior bor-derslightlyrounded.Terminalstrut1/3lengthofphallobase.Penis guideindorsalviewwithslightlyconvergingsides,slightly sinu-ous,narrowingto3/4ofwidthat5/6oflength,apical1/6endingin triangle,withconcavesidesandslightapicalprojection;inlateral viewtriangular,sinuous,benttowardparameresindistal 1/6of apex.Parameresindorsalviewnarrowandslightlycurvedtoward basallobe,1/5shorter thanthis,withpubescenceindistalhalf, exceedingparamereinalmosthalfoflength(Figs.22–24).Penis widthabout1/6ofmaximumlength,inasemicircleinbasalhalf, slightlycurvedto2/3length,distal1/3straight.Apex1/6oflength andslightlywiderthanpenistube,separatedfromitbyasclerosed unionatwidelyobliqueangle,withejaculatoryduct“S”shapedand sclerosespineonapex;peniscapsulewithinternalbranch hook-shaped,outerbranchtwiceaswideasinternal,basalmarginconvex (Fig.21).

Female:Similartomale.Spermatheca“C”shaped,aboutten timeslongerthanwide,narrowingregularlytowardapex, nodu-luslateralandshort,ramus1/10oftotallength,seminalductvery short.Infundibulum“Y”shaped,halfsizeofspermathecal(Fig.25). Coxitespaddle-shaped,withverywide,pubescentapexand evi-dentstylus.

Variation:Length2.1–2.6mm.Colornearlyhomogenous,spots varyingfromlightbrowndark-ringedtohomogeneousdarkbrown. Thespots,especiallyonthepronotumandsupernumeraryofthe elytratendtodisappear,whiletheelytralspotsmaybepartially dis-jointedtorarelylosingthe“S”shape,theposteriorspotsometimes complete(Figs.15,19,20).

Discussion

Psylloboraluerisp.nov.differsin thecolorationpattern and inthemale genitalia(seespecieskey).Also,itisgeographically isolatedfrom otherspecies ofthegenus,probably occurringin ArgentinatoChubutProvince,wherenoPsylloboraspecieshasbeen recordedpreviously.

Geographicaldistribution:AustralChile,ChileChicoProvince (Fig.53).

Etymology:ThisspeciesisnamedforAlfredoLüer,dedicated Chileanentomologistandcollectorofmanyoftheknown speci-mensofthisspecies.

Psylloborapicta(Germain,1854)

(Figs.25–37,50)

CoccinellapictaGermain,1854:335;Philippi,1887:173. Psyllobora picta: Brèthes, 1923:454; Costa Lima, 1937:32;

Blackwelder, 1945:455; Mader, 1957:54; González, 2008:95, 2013:52,2014:530;Serraetal.,2013:238.

PsylloboraferalisMulsant,1866:178,Crotch,1874:142;Weise, 1904:195;Gordon,1987:17assynonymofP.picta).

Psylloborapictaa.feralis:Blackwelder,1945:455.

Halyziafemoralis:Philippi,1887:174(transcriptionerror “fer-alis”).

Malegenitalia:Tegmen4.3timeslongerthanwideonaverage, phallobasesubrectangular,1.1timeslongerthanwideand with somewhatroundedposteriorborder.Terminalstrut2/5of phal-lobaselength.Indorsalview,penisguidewithalmostparallelsides inbasalhalf,thensinuousandreducedto3/5widthindistalthird, withmembranoussemicircularlateralexpansions,apical1/6 trian-gularwithconcavesidesandslightapicalprojection;inlateralview withyellowishventralprojections,semicircularandwithverywide border,twicewidthofparamere,in1/3distalapexcurvingtoward parameres.Parameresnarrowindorsalviewandslightlycurved towardpenisguide,1/5shorterthanpenisguide,distalhalfwith longpubescence,whichexceedsparamerebyalmosthalfitslength (Figs.36–38).Peniswide,about1/6ofmaximumlength,basalhalf semicircular,slightlycurveduntil2/3oflength,distal1/3straight. Apexcomprising1/6oflengthandslightlywiderthanpenistube, separatedinstronglyacuteangle,withejaculatoryduct“S”shaped andsclerosespineatapex;peniscapsule,internalbranch hook-shaped,outerbranchtwiceaswideasinternalbranch,basalmargin sometimesslightlyconvex,withnotchedcenter(Fig.35).

Female:Similartomale.Spermatheca“C”shaped,abouteight times longerthan wide, lateralnodulus quiteshort, ramus1/6 oftotallength,seminalductveryshort,infundibulumirregularly anchor-shaped,halfsizeofspermatheca(Fig.39).Coxites paddle-shaped,withverywideapexandevidenthairstylus.

Variation:Length2.2–3.1mm.Dorsalcoloroflivespecimens brightwhitewithblackspots(Fig.34),butsomespecimenswiththe dorsalcolorivoryyellowanddark-brownspots.Spotsofthehead arealwayspresent,sometimesjoined.Pronotalspotsonthesame sidearesometimesfusedinobliquestripes,bothstripessometimes joinedatthebase;somespecimenshaveadditionalspotsonthe pronotumborder,togetherorseparate.Elytralspotsarevariable, withspot4andsometimesspot8formingacommonspotwithits counterpartontheotherelytra,orslightlyseparatedandtouching thesuture,spots1–3normallyattached,otherssometimesslightly touching,andapicalspot10isalwayswellformed(Figs.26,31–34). Ventralsurfaceof metaventriteis usuallyblack,and sometimes pro-,mesosterno-,andpartofabdomenblack.Femursclearorin somespecimenshaveblackringscoveringthemiddlearea(Fig.30).

Distribution: Southern Argentina (Neuquén and Río Negro provinces) and Chile (San Antonio to Llanquihue provinces) (Fig.53).ThereportofthisspeciespresenceinParaguay(González,

2013)shouldbediscarded,sinceitcomesfromacitationofBertoni (1925)thatprobablyreferredtoP.bicongregata,atthattime con-sideredasynonymofP.picta.Forthesamereason,itislikelynot presentinUruguay,whereitwascitedbyRuffinelliandCarbonell (1954).ThepresenceofthespeciesinArgentinaisbasedonasingle citationofP.feralisWeise(1904)inNeuquénandonefemalefrom ElBolsón,RíoNegroProvince,soitspresenceshouldbeconfirmed byexaminationofmalespecimens.

Typematerialexamined:Holotype:“Chile”“Holotipo” “Coc-cinellapictaGermain”“M.N./H.N./TipoN◦2162”[MHNS].

Dissected Material examined (11 ♂ and 12 ♀): 1 ♂ and 1 ♀: “Chile, Prov. San/Antonio,Algarrobo/16-enero-1989/leg. G. González” [CPGG]. 2 ♂ and 1 ♀: “[Chile] Maullín a/Las Que-mas/Llanquihue/17, -dic.1972/Coll:L.E.Pe ˜na” [UMCE].2♂and 1 ♀: “Chile, Prov. Melipilla/Pollocabe 300 msnm/7–29-marzo-2004/leg.V.M.Diéguez”“gen.1648”[CPMD].1♀:“[Chile]Quillota/7 1894” “Colección/P.Germain”“Psyllobora/picta/(Germain)/Mader det.” “gen. 1649” [MHNS]. 1 ♂: same data except “gen. 1650” [MHNS].1♂and1♀:“Contulmo/Arauco/IX-Reg-Chile/10-I-1988” [CPGG].1♂and1♀:“[Chile]S.IsidroXII”“ColecciónPhilippi”“Gen 1654”[MHNS].1♀:“ChileX-R/Valdivia/SanJoséde la/Mariquina/2-2-1985/Coll: C. Fierro” “Gen 1658” [CPAL]. 1 _♀: “Chile – R.M./Chacabuco/Chacabuco/15-II-2008/Coll:A.Lüer”“Gen1659” [CPAL].1♂:samedataexcept“12-XI-2007”[CPAL].1♀:“Argentina, RioNegro/ElBolsón/41,939242◦S71,544006◦W/Febrero2002/Leg. P.Sackman”(ab)[MHNS];1♀:“marzo2007/P.N.LaCampana/V Región/ChileColl:R.HonourS”“Hembra187”[CPRH];2♀: “23-IV-1993/CuestaBarriga/PadreHurtadoR.M./ChileColl:R.HonourS” “Hembra139”and“Hembra145[CPRH];2♂:samedataexcept “macho140”and“macho144”[CPRH].

Other material examined (101 specimens): Chile: without data,coll.P.Germain,2specimens[MHNS];Bio-Bio:Trapa-Trapa, 16-12-1996,coll.ALüer,1specimen[CPAL];Cachapoal:Pencahue, 24-10-1991,coll.ALüer,1specimen[CPAL];LaderaS.Cerro Llivi-Llivi,1-9-2012,leg.ALüer,1specimen[CPAL];Cautín:Villarica, 12-II-1964,T.Cekalovic,41specimens[DZUP];Temuco,III-1922, D.S.Bullock,1specimen[MNRJ];Caburga,16-2-2008,coll.ALüer, 2specimens[CPAL];Lican-Ray,I-1982,G.GonzálezF.1specimen [CPGG]; Curacautín, Lag. Conguillio, 6-I-1979,leg. D.Jackson, 1 specimen[CPGG];Chacabuco:Chacabuco,ElRoble,6-III-2011,leg. ALüer,1specimen[CPAL];Choapa:Limahuida,17-X-2009,leg.A Lüer,1specimen[CPAL];Cordillera:SanJosédeMaipo,Lagunillas, 2100m,leg.R.HonourS,1specimen[CPRH];Curicó:Cerro Hueca-Huecan,Zapallar,15kmECuricó,12a21-XII-1997,Malaisetrap, leg.J.E.Barriga,1specimen[JEBC];15kmE.PotreroGrande, 29-I-1995,leg.J.E.Barriga,1specimen[JEBC];Llanquihue:Todoslos Santos,Cayutue,6-III-(19)50,coll.Kuschel,2specimens[MHNS]; Fresia,6-I-(19)45,coll.Kuschel,Ribesnigrum,1specimen[MHNS]; Malleco: Conguillio, 3-II-1983, E. Krahner, 1 specimen[MHNS]; El Manzanar, Lonquimay, IV-1988, leg. J.E.Barriga, 1 specimen [JEBC];Lonquimay,12-I-2008,foggings/Nothofagusdombeyi,leg. J.E.Barriga,1specimen[JEBC]:Pemehue,SelvaOscura,22-12-1996, coll.ALüer,1specimen[CPAL];Victoria,Fdo.LasMariposas, 18-10-1995,coll.ALüer,2specimens[CPAL]:Melipilla:Melipilla,C◦ Cantillana,2-8-1998,leg.ALüer,1specimen[CPAL];Chi ˜nihue, 21-II-2007, leg.A Lüer,1 specimen[CPAL]; Quillota:3-XI-1963, M. Cerda,1specimen[MNRJ]; ˜Nuble:SanFabián,21-11-1994,Coll. ALüer,1specimen[CPAL];Chillán,LasComadres,6-I-1996,coll. S. Roitman,2specimens [CPGG];LosLleuques,12-12-2008,leg. ALüer, 1specimen[CPAL];Las Trancas,11a 16-II-2011,leg.R. Honour S, 1 specimen [CPRH];Petorca: LaCanela, 20-dic-2008, leg.ALüer,1specimen[CPAL];Quillota:Quillota,7-1894, Colec-ción P. Germain,1 specimen[MHNS]; P.N. LaCampana, marzo 2007,Coll:R.HonourS.1specimen[CPRH];SanAntonio:Algarrobo. 16-II-1969, G. González F, 6 specimens [CPGG]; Santiago: Val-lecito,19-2-2011,leg.ALüer,1specimen[CPAL];QuebradaMacul,

4-VIII,J-C.Sáez,1specimen[JEBC];Renca,IV-1964,leg.L.E.Pe ˜na, 1specimen[JEBC];Talagante:Chi ˜nihue,20-8-2009,leg.ALüer,1 specimen[CPAL];Maipú,26-XI-2011,coll:R.HonourS,4specimens [CPRH];Talagante:/PadreHurtado,CuestaBarriga,Coll:R.Honour S.2specimens [CPRH].Talca:VilchesAlto,10-XII-1988,Trampa malaise,1specimen[CPGG];AltosdeVilches,I-1997,leg.J.E. Bar-riga,1specimen[JEBC];Valdivia:PuertoPirihueico,16-1-1998,leg. ALüer,1specimen[CPAL];Valparaíso:4-X-1921,417/FCC-567,1 specimen[MNRJ];LasZorras(nearValparaiso),25-II-1932,3 spec-imens[MNRJ];Limache,II-1922,418/FCC-568,1specimen[DZUP]; LoVazquez,X-1988,leg.J.E.Barriga,1specimen[JEBC].

Discussion

Psylloborapictaiseasilyrecognizablebyitswhitecolorwith well-definedblackspots(Fig.34)(seespecies key).Afew spec-imens are ivory- yellow or with dark-brownspots, somewhat similartosome specimens ofP. paulinesp.nov., but retainthe othercharacteristicsof thespecies. Probablythecolorof these specimensistheresultofsomepost-mortemprocesses,because theycomefromoldcollections.ThepopulationsofsouthernChile havelargerandmorepigmentedspecimens,butcorrespondwell withthespecies(seenoteunderP.paulinesp.nov.).

PsylloborabicongregataBoheman,1858

(Figs.40–53)

PsylloborabicongregataBoheman,1858:203,Crotch,1874:141; Weise,1926:11;Blackwelder,1945:455assynonymofP.picta;

Mader,1957:24;Gordon,1987:17assynonymofP.picta(González, 2008:95;2013:52,2014:530;Serraetal.,2013:237.

Psylloborapavida Weise, 1904:195; Blackwelder, 1945: 455;

Mader,1957:24.

Malegenitalia:Tegmen3.6timeslongerthanwideonaverage, phallobasesubrectangular,aslongaswideandwithrounded bor-der.Terminalstrutshort,1/3ofphallobaselength.Indorsalview, penisguidewithgentlysinuoussides,withasinuosityat3/5of lengthandalmostparallelsides,reducedto4/5ofbasalwidthat 5/6oflength,thentaperingsharplyintriangleinapical1/6toend inmamilliformprojectioninapical1/12;inlateralviewwith yel-lowishventralsemicircularprojectionsverywide,twicewidthof paramere,in1/3distalapexcurvingtoparameres.Parameresin dorsalviewnarrowandslightlycurved,1/6shorterthanbasallobe, distalhalfwithlongpubescence,whichexceedsparamerebyover halfitslength(Figs.49–51).Penis wide,about1/6ofmaximum length,basalhalfsemicircular,thengentlycurvingto2/3length withstraightdistal1/3.Apexoccupying1/6oflengthandslightly widerthanpenistube,separatedfromitbyalmost perpendicu-larunion,withejaculatoryduct“S”shapedandsclerosedspineon innerapex;peniscapsule,innerbranchhook-shaped,outerbranch twiceaswideasinternal,basalmarginconvex,sometimes some-whatemarginatetocenter(Fig.48).

Female:Similartomale.Spermatheca“C”shaped,abouteight timeslongerthanwide,lateralnodulusquiteshort,ramus1/6of totallength,seminalductshort,infundibulumirregularlyanchor shaped,halfsizeofspermathecal(Fig.52).Coxitespaddle-shaped, withverywideapexandevidentstylus.

Variation:Length 1.9–2.7mm.Dorsalcolorvaries fromlight browntoyellowish ivory, spots brownwith darkenedborders, outerspotsusuallyhomogeneouslydarkbrown,sometimessome ofdiscalspotsdarkbrown.Headspotsvaryinginsizeandcolor. Pronotalspotsvaryinginsize,sometimesjoinedonsameside,some specimenswithsmallspotsonborderofpronotum.Elytralspots quitevariable,alwaysseparatefromsuture,usuallyspots1–3and spots5–7joined,otherspotssometimeslightlytouching. Super-numeraryspotsusuallybetweenspots5,7andsuture,sometimes

verylightincolor,apicalspotusuallydistinct,verylightincolor, sometimesblurry.Spot6oftenseparatedintotwosmall longitudi-nalspots.Ventralsurfacelightbrown,usuallywithdarkerbrown metaventrite(Figs.40,44–47).Inonecase,weobservedonlyspots 1,2and4andanentirelyclearventralsurface,perhapsanimmature specimen.

Geographical distribution: Argentina (from Neuquén north), Uruguay, Paraguay and southern Brazil (Paraná) (Fig.53).

Dissectedmaterialexamined(12♂and13♀):3♂(mv)and 1 ♀ (ab): “Argentina, prov. Formosa/Estancia La/Marcela,35km E.El/Colorado/21dic2004,/26◦17.35S59◦8.6 Wleg/J.Williams JEBarriga”[JEBC].2 ♂(mv) and3 ♀(mv):“[Argentina]Trampa agua. Cult. Hinojo/Pocito, San Juan/23-IX-2005. Lanati S.J. col” [CPSL].1♂(mv)and1♀(mv):“Argentina,Prov./Cordoba.Capilla del/Monte,7-mar-2006,luz,/Leg.D.Carpintero[JEBC].2♀(mv): “Argentina, La Rioja/Villa Mazán, Finca/Dahba, 11-IX-2009/Leg. Claudia Funes” [CPCF]. 1 ♂ (mv) and 2 ♀ (ab): “Argentina, La Rioja/Villa Mazán, Finca de/La Fuente, 51-II-2009/Leg. Clau-diaFunes” [CPCF]. 1 ♂ (mv) and 2 ♀ (ab, mv), “Argentina, Rio Negro/Cervantes Chacra/Metallier, 5/V/2015, leg./F. d’Hervé, en perales/fajas de cartón” [CFDH]. 1 _♂ (mv): “Bariloche/Rio Negro/II-1976” “Colección/Dr. Carpintero/Argentina” “Psyllob-orabicongregata/det. Dr. Carpintero” [JEBC]. 2 ♂(mv) and 1 ♀ (mv), “Argentina,Rio Negro/FrayLuis Beltrán, Chacra/Castagno, 29/III/2014, leg./F. d’Hervé, en manzanos/fajas de cartón” [CFDH]. 1 ♂ (mv) and 1 ♀(mv): Paraguay, Paraguari/Sapucay, Chircal,/S25◦43 W56◦59/38-31-I-2005/leg. Ulf Drechsel [CPUD].

Othermaterialexamined(222specimens):Argentina:Buenos Aires:BuenosAires,XII-(19)78,leg.J.E.Barriga,1specimen[JEBC]; Burzaco,I-(19)40,leg.H.C.Hepper,1specimen[JEBC];Delta, X-1959,M.J.Viana,11specimens[JEBC],LaPlata,Villena,05/10/2011, M.Lattari,1specimen[CPGG];Miramar,II-2004,leg.D.Carpintero, 1specimen[JEBC];Palermo,XI-1976,leg.J.E.Barriga,1specimen [JEBC];Pilar,13-XI-1976,leg.J.E.Barriga,1specimen[JEBC];Punta Lara,XI-1959,M.J.Viana,5specimens[JEBC];Quilmes,I-2002,leg. D.Carpintero,luz,1specimen[JEBC];Tandil,II-2001,leg.Dellais, luz,1specimen[JEBC];Tigre,IX-1959,M.J.Viana, 10specimens [JEBC];Tigre,X-1959,M.J.Viana,5specimens[JEBC];Tigre,I-1961, M.Viana,5specimens[JEBC];VillaDevoto,X-(19)40,leg.H.C. Hep-per,1specimen[JEBC];Zárate,I(Isla)Talavera,30-X-2004,leg.J.E. Barriga,1specimen[JEBC];Zelaya,I-1967,col.H.C.Hepper,2 spec-imens[JEBC];Catamarca:ValleCentral,CerroGuacho,24-IV-2009, 2specimens[CPCF];samedataexcept26-VI-2009, 2specimens [CPCF];Córdoba: Calamuchita, ElSauce, III-1959, M.J. Viana, 25 specimens [JEBC]; same data except 17-XI-1970, 5 specimens [JEBC];Corrientes:Lavalle,SantaLucía,31-X-1961,M.A.Roisilloand M.M.Portillo,s/Malvacea,2specimens[JEBC];Estación experimen-talBellaVista,1-VIII-(19)62,1specimen[JEBC];EntreRíos:1de Mayo,VI-1960,leg.H.C.Hepper,1specimen[JEBC];Colón,III-2003, leg.D.Carpintero,luz,1specimen[JEBC];Concordia,22-III-(19)77, M.M.Portillo,s/Zinnia, 4specimens[JEBC];Formosa:Estancia La Marcela,35km. E.ElColorado,21-VII-2003 leg.J.Williams and J.E.Barriga-Tu ˜nón,1specimen[JEBC]; samedataexcept 21-XII-2004,5specimens[JEBC];LaRioja:Patqui,XI-1959,M.J.Viana,20 specimens[JEBC];VillaMazán,FincadeLaFuente,5-II-2009,leg. ClaudiaFunes,4specimens[CPCF];samedataexcept11-IX-2009, 14specimens[CPCF];LaPampa:SantaRosa,I-2000,leg.Stortl,luz,3 specimens[JEBC];Mendoza:LaConsulta,3-X-2003,leg.S.J.Lanati, trampaagua,1specimen[CPGG],samedataexcept19-IX-2003, 25-IX-2003and1-X-2003, 3specimens[CPGG];Misiones:Arroyo Itaembe,II-1996,leg.D.Carpintero,1specimen[JEBC];RíoNegro: Bariloche,leg.D.Carpintero,1 specimen[JEBC]; Salta:Cafayate, Corralito,X-1986,F.J.Viana,1specimen[JEBC];RosariodeLerma, I-1983,M.J.Viana,1specimen[JEBC];RosariodeLerma,XI-1982,

M.J.Viana,3specimens[JEBC];SanJuan:Pocito,9-IX-2005,leg.S.J. Lanati,cultivohinojo,trampaagua,2specimens[CPSL];samedata except1-X-2003,3-X-2003and17-XI-2005,3specimens[CPSL], Sapuncay,same data except 6–7/11/2005 and 28–31/1/2005,3 specimens[CPUD].Paraguay:Boquerón:Mistolar,22◦40S62◦08 W,3–12/10/1997,leg.UlfDrechsel,1specimen[CPUD],Transchaco Hwykm420,22◦24S59◦50W,19–28/4/1995,leg.UlfDreschel,1 specimen[CPUD].Caazapa:Tava.Cordillera:Naranjo,30-XII-2005, leg.C.Aguilar,1specimen[MHNP].Paraguarí:Sapuncay,Chircal, 23◦43S66◦69W,28a31-I-2005,leg.Ulf.Dreschel,2specimens [CPUD], Presidente Hayes: LolltaYaragui, 14/1/2001, Ulf Drech-sel,1specimen[CPUD];SanPedro:Rio Ypane,XII-1993,leg.M. Viana,3specimens[JEBC];Cororo,PuertoRosario,16–21/10/2006, Ulf Drechsel, 1 specimen [CPUD]. Uruguay: Canelones: Arroyo Vizcaíno (34◦3703.72 S 56◦0232.56 W), 26/I/2012, leg. W.S. Serra, A. Duarte, M. Zarucki, M. Loureiro/MNHN 897/6 spec-imens [URMU]; Las Toscas (34◦4554.22 S 55◦4417.93 W), 11/I/2012,leg.S.Greco/MNHN,939/1specimen[URMU];Florida: CiudaddeFlorida.(34◦0555.20S56◦1310.78W),18/XII/2011, leg. W.S. Serra/MNHN 865/2 specimens [URMU]; same data except 14/I/2012/MNHN 862/2 specimens [URMU]; same data except25/VII/2011/MNHN705/1specimen[URMU];Montevideo: Casavale,430/FCC-580;430/FCC581,1-3-XII-1934,429/FCC-579,3 specimens;429/FCC-579,1specimen“Homeotype”,Korschefsky [MNRJ];Río Negro:Las Ca ˜nas(33◦0957.08 S58◦2132.43 W), 21/IX/2011,leg.W.S.Serra,C.ClavijoandV.Bianchinotti/MNHN 750/5 specimens [URMU]; same data except 22/IX/2011, leg. W.S.Serra,C.ClavijoandV.Bianchinotti/MNHN751/9specimens [URMU], same data except 7/XII/2011,leg. W.S. Serra,C. Clav-ijoandC. deMello/MNHN849/7specimens [URMU],samedata exceptIII/2012,leg.W.S.Serra,C.ClavijoandS.González/MNHN 1013/4 specimens [URMU]; San José: Paso del arroyoMauricio (34◦4056.87S56◦4120.69W),9/I/2012,leg.W.S.Serra/MNHN 834/13 specimens [URMU]; Treinta y Tres: La Charqueada, Caba ˜nas de la Intendencia (33◦1213.28 S 53◦4758.09 W), 17/VIII/2011,leg.W.S.Serra/MNHN742/1specimen[URMU],same data except28/IX/2011, leg. W.S. Serra/MNHN 814/1 specimen [URMU].

Newrecords from Brazil (31specimens): Paraná: Maringa, Escola Milton Santos, S 26◦2330.83 O 52◦132.65, 484malt. 22jul2012,C.CamachoHuerta,1specimen[UNAB].Santa Cata-rina:Concordia(EntreRios),84/FCC-216,1specimen[MNRJ];Nova Teutônia,X-1980,F.Plaumann,4specimens[DZUP].RioGrandedo Sul:PortoAlegre,III-1928,N.B.Fagundes,1specimen[MNRJ]; 16-IX-1932,1specimen[MAPA];I-1940,A.Bento,1specimen[MNRJ]; 18-X-1944,1 specimen; 29-III-1948, 2 specimens; 6-X-1948, 2 specimens;11-VII-1949,1 specimen;12-VIII-1949, 1specimen; 5-VII-1950, 1 specimen; 31-V-1950, 1 specimen; 14-V-1952, 1 specimen; 27-VIII-1952, 1 specimen; 23-XII-1952, 1 specimen; 29-I-1964,2 specimens[MAPA[; Bagé,30-IX-1972, A.R.Jobin, 1 specimen[DZUP];CerroLargo(SerroAzul),XII-1943,2specimens; Viamão,11-XI-1964,4specimens[MAPA];XII-1926,Deslandes,1 specimen[MNRJ].

Discussion

The taxon P. bicongregata was long maintained in syn-onymywithP.picta.However,theirmorphologicalcharacteristics, especiallythemalegenitalia,allowthespeciestobeclearly distin-guished(seespecieskey).ThepresenceofP.bicongregatainChileis discarded,sincespecimenspreviouslyassignedtothistaxon actu-allyrepresentthenewspeciesdescribedhere,P.paulinesp.nov.

andP.luerisp.nov.

Conclusion

ThisrevisionofthePsylloborapictaspeciescomplexallowedus toestablishthepresenceoffourdistinctandgeographicallyquite separatespecies.

Theexternalsimilarityofthesespeciesandtheapparent transi-tionsbetweenthemhaveledspecialiststoconfuseandsometimes synonymizethemwitheachother,makingthelocalitiescitedin theliteraturedubious.Studyofabundantmaterialallowedusto discardtherecordsof P.picta fromParaguay and Uruguayand ofP.bicongregatafromChile,andtoextendthedistributionofP. bicongregatatoBrazilandalsotodescribetwonewspeciesfrom Chile.Analysisofthemalegenitaliawasessentialtorecognizethe species,although thedifferencesobservedfor somespeciesare minorandsometimesdifficulttoevaluateinisolatedspecimens. Fortunately,thecombinationofthiselementwiththefeaturesof design,coloration,geographicaldistributionandfieldobservations hasenabledustoproperlyidentifythespecies.Thefemalegenital tracthasnotshownanyusefulcharacteristicsinthisgroup,dueto itsintraspecificvariability,exceptperhapsthelargest spermathe-caelength/widthratioobservedinP.lueriwithrespecttotheother species.Wetakethisopportunitytoacknowledgethesignificant contributionofMader(1957),who,aloneamongtheauthorswho treatedthesespeciesinthepast,clearlyintuitedtheiridentityand theircorrectgeographicaldistributions.

Conflictsofinterest

Theauthorsdeclarenoconflictsofinterest.

Acknowledgments

We thanks toAlfonso Aguilera (CPAA), Alfredo Lüer (CPAL), Claudia Funes (CPCF), Francisco Ramírez (CPFR), Richard Hon-our(CPRH),GerardoArriagadaandManuelDiéguez(CPMD),Ulf Drechsel(CPUD),JuanEnriqueBarriga(JEBC),FernandoR.Meyer (MAPA), JohnKochalka (MHNP),MarioElgueta(MHNS), Miguel AngelMonné(MNRJ),andDaniloCepeda(UMCE)fortheloansof material.ToFranciscoSernaandTakumasaKondoforsupportthe visitfromthecollectionoftheUniversidadNacionaldeColombia (UNAB)andtoJanetReidforrevisingtheEnglish.Specialthanksto AlfonsoAguilera,RichardHonourandthetwoanonymous review-ersforcriticalreviewofthemanuscript.WealsothanktheConselho NacionaldeDesenvolvimentoCientíficoeTecnológico(CNPq)for theresearchfellowshiptoLMA(306772/2006-0).Thisis contri-butionnumber1939oftheDepartmentofZoology,Universidade FederaldoParaná.

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