Phylogeny of the Augochlora clade with the description of four new species (Hymenoptera, Apoidea)

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REVISTA

BRASILEIRA

DE

Entomologia

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

Systematics,

Morphology

and

Biogeography

Phylogeny

of

the

Augochlora

clade

with

the

description

of

four

new

species

(Hymenoptera,

Apoidea)

Rodrigo

Barbosa

Gonc¸

alves

UniversidadeFederaldoParaná,DepartamentodeZoologia,Curitiba,PR,Brazil

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Articlehistory:

Received19September2018 Accepted20December2018 Availableonline3January2019 AssociateEditor:RodrigoFeitosa Keywords: Augochlorini Bees Neotropical Systematics Taxonomy

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TheAugochloraclade includesfour genera:AugochloraSmith, AugochlorellaSandhouse, Ceratalictus Moure,andPereirapisMoure.ThisisoneoftherichestandmostwidespreadgroupsofAugochlorini bees.Thereareabout150species,whichoccurfromArgentinatoCanada.ThespeciesofAugochloraclade areconsideredsolitarytofacultativelysocial,exceptCeratalictusforwhichnothingisknown.Wood nes-tingbehavioraroseonceintheclade,inAugochlorasensustrictu.Theobjectiveofthisstudyistodescribe fournewspeciesandtopresentarevisedphylogeneticanalysisoftheAugochloracladefortheplacement ofthesespecies.Themorphologicalmatrixcomprised77charactersand42terminals,andresultedin twomostparsimonioustrees.ThemonophylyoftheAugochloracladeiscorroborated.Ceratalictusand PereirapisareconsideredassistergroupsandCeratalictusinﬂexussp.nov.cameassistertootherspecies ofCeratalictus.AugochloraandAugochlorellaaremonophyleticandsistergroups.Bothextantsubgenera ofAugochlorawerecorroboratedasmonophyletic.Augochlorellacomisisconsideredassistergrouptothe restofAugochlorellaspecies.AllAugochlorellanewspeciesdescribedbelongtotheAugochlorellaephyra group.Augochlorellakelliaesp.nov.isphylogeneticallyrelatedtoAugochlorellauna.Augochlorellaprocliva sp.nov.andAugochlorellamavricerasp.nov.constituteacladewithAugochlorellaacarinata.Including thenewspecies,Augochlorellahas19speciesandCeratalictus11species.Arevisedkeyforspeciesof AugochlorellaandCeratalictusisalsopresentedintheSupplementaryInformation.

Introduction

Augochlorini is a monophyletic bee tribe and the main lin-eagesdivergencedating around55–20Ma (Hedtke et al.,2013; Gonçalves,2016).Thetribecomprises663describedspecies pri-marilydistributedintheNeotropicalregion(Moure,2007).Species ofAugochlorinihavearemarkablerangeofbehaviorsandhabits, including solitary, semisocial, primitively eusocial, crepuscular, nestinginthesoilandwood,andcleptoparasitism(seeMichener, 2007forareview).Thesediverseandecologicallyinterestinghabits maketheaugochlorinebeesagoodgroupfortheevolutionstudies ofsuchbehaviors.Inarecentmolecularandmorphological phy-logeny,Gonçalves(2016)recognizedsevensupragenericcladesfor thetribeandtheAugochloracladewasrecoveredsistertothe Mega-loptagroup(Gonçalves,2016;MeiraandGonçalves,2018).

TheAugochloracladeincludesAugochloraSmith,Augochlorella Sandhouse, CeratalictusMoure, and Pereirapis Moure (Eickwort, 1969).The taxonomic treatmentof these genera varies among authors,Michener (2007)considered Ceratalictus and Pereirapis

E-mail:goncalvesrb@gmail.com

assubgenera ofAugochlorella,butothers treated themas sepa-rategenera(Eickwort,1969;DanforthandEickwort,1997;Engel, 2000;Coelho,2004; Moure,2007;Coelho andGonçalves,2010; EickwortandEickwort,1973).Thelateclassificationissupported by multiples phylogenies (Danforth and Eickwort, 1997; Engel, 2000;Coelho,2004;Gonçalves,2015,2016)inwhichthegenus Augochlorelladoesnotformamonophyleticgroupwith Ceratalic-tusandPereirapis.Michener’sclassificationcreatesaparaphyletic groupinrelationtoAugochlora.

Augochlora is the most speciose genus of the group, with 115species,rangingfromArgentinaandChiletoCanada(Moure, 2007).Augochlorahasthreesubgenera:Augochloras.s.(87species), ElectraugochloraEngel(onefossilspecies),andOxystoglossella Eick-wort (27 species). No revisionary or phylogenetic studies have been conducted for the genus. Pereirapis is distributed from southernBraziltoMexico,and ithaseight speciesaccordingto Moure(2007),althoughEngel(2000)consideredthegenustobe monotypicwithallspeciessynonymwithP.semiaurata(Spinola, 1853).Coelho(2004)revisedAugochlorella,recognizing16species distributedfromArgentinatoCanada.TheAugochlorellaphylogeny supportstheplacementofAugochlorellaasa sistergrouptothe restoftheAugochloraclade(Coelho,2004).Ceratalictustaxonomy https://doi.org/10.1016/j.rbe.2018.12.003

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92 R.B.Gonc¸alves/RevistaBrasileiradeEntomologia63(2019)91–100

wasrevisedbyCoelhoandGonçalves(2010)whorecognized10 species. Gonçalves (2015) provided a phylogeny of Ceratalictus speciesandalsosupportedtheplacementofAugochlorellaassister to the remaining genera. Recently, however, Gonçalves (2016) recoveredthefollowingdifferenttopologyusingmorphologyand DNA((Augochlora,Augochlorella),(Ceratalictus,Pereirapis)).

Thenestingbiologyandbehaviorforsomespeciesofallgenera areknown,exceptCeratalictus.StudiedspeciesofAugochlora (Oxys-toglossella),Augochlorellaand Pereirapisare soilnestersand can befacultativelyeusocial(Eickwort,1969;EickwortandEickwort, 1972; Danforth and Eickwort, 1997 see Michener, 2007 for a review).Augochloras.s.speciesnestinrotten–woodandarealso solitaryorfacultativelyeusocial(Stockhammer,1966;Dalmazzo and Roig-Alsina, 2012, 2015; Wcislo et al., 2003) and are not exclusivelysolitaryaspreviouslypostulated(Stockhammer,1966; DanforthandEickwort,1997).Interestingly,intraspeciﬁcvariation isalsoknownforsomespeciesoftheAugochloraclade,suchas AugochloraiphigeniaHolmberg,1899,Augochlorellacomis(Vachal, 1911)and Augochlorellaaurata (Smith,1853) (seeDanforth and Eickwort,1997forareview).

Since the taxonomic revisionsof Coelho (2004) and Coelho and Gonc¸alves (2010) new species accumulate in Augochlorella and Ceratalictus. A newCeratalictus was already knownby the authorbutwithuncertaintyongenericplacement,mainlyduea peculiargenitalcapsule.Themaingoalofthisstudyistopresent arevisedphylogenetic analysisfortheAugochloraclade forthe placement of four new species. Other objectives are to com-pileandrevisethemorphologicalcharactersofpreviousstudies into a more complete dataset, and to describe one new Cer-atalictusand threenew Augochlorellaspecies.A revisedkey for speciesofAugochlorellaandCeratalictusisalsopresentedinthe SupplementaryInformation.

Materialandmethods Phylogeneticanalysis

AsingroupIselectedseven speciesof Augochlora represent-ingtheextantsubgenera,onespeciesofPereirapis,andallknown species of Augochlorella and Ceratalictus. The new species of Augochlorella and Ceratalictus were also included as terminals. TheincludedAugochloraspeciesaredistributed fromArgentina, Braziland Paraguay.Thereareseveralgenuslevelnames avail-ableforAugochlorabuttheyarenottestedhere.Allspecieswere directly examined except some of Augochlorella aurata group. For those species, the character coding was the same present byCoelho (2004)withfew modifications based ontheoriginal drawings and descriptions. As outgroupI selected four species of Augochlorini including two species of the Megalopta clade, thesistergroupofAugochloraclade(Gonçalves,2016;Meiraand Gonçalves,2018).Pseudaugochloragraminea(Fabricius,1804)was usedtorootthecladogrambasedonthetopologiesofGonçalves (2016)andMeiraandGonçalves(2018).Forthecharacter state-ments,IfirstlyrevisedtheoriginaldatasetsofCoelho(2004),and Gonçalves(2015,2016).Thelaterincorporatedprevious charac-tersfromEickwort(1969)andEngel(2000).Thecharactercoding fromtheoriginalstudieswerecompletelyrescoredbasedondirect observationand newinterpretations. The second step wasthe inclusionofnewcharactersstatementsbasedonexternal morphol-ogyandmalegenitalia. Thewritingofallcharactersstatements followed the recommendations of Sereno (2007). Inapplicable characterstates for contingent characters are indicated by “–”, missing characterstates by “?”. The final matrix comprised 77 charactersand 42 terminals,both matrix and characterlistare

providedintheSupplementaryInformation.Parsimonyanalyses werecarriedoutinTreeAnalysisusingNewTechnologySearch (TNT, Goloboff et al., 2008), using equal weights, and the fol-lowingsettings(ratchetweightingprobability5%,200iterations), tree–drifting(50cycles),andtree–fusing(ﬁverounds,minimum lengthsettobehit1000times).Allcharacters weretreatedas non-additive.Bootstrap (1000replications)and Bremer support werealsocalculatedwithTNT.Treeswereviewedandeditedwith WinCladaver.1.00.08(©KCNixon)andGIMP2.8.16(©TheGIMP Team).

Taxonomy

Thedescriptions were organizedfollowing thebody organi-zation,tagmata, segments,and sclerites, withspecial attention given to the following:structure (general organization), sculp-turing(impressionsandelevationsonscleritesurface),colorand pubescence.Structureterminology followsEickwort (1969)and Michener(2007)exceptthebasalareaofpropodeumreferredhere asthemetapostnotum.IusetheabbreviationsT1,T2,etc.,todenote themetasomalterga;S1,S2,etc.,todenotemetasomalsterna;and F1,F2,etc.,todenotetheflagellomeres.Forexaminedmaterialthe quotationmarkssurroundexacttranscriptionsofindividuallabels, whilethebackslash(\)indicatesdifferentlinesonthesamelabel. Thematerialcitedhereisdepositedinthefollowinginstitutions: BMNH–BritishMuseumofNaturalHistory,London,UK;DZUP– ColeçãoEntomológicaPe.JesusSantiagoMoure,Curitiba,Brazil; MZSP–MuseudeZoologiadaUniversidadedeSãoPaulo,SãoPaulo, Brazil;SEMC–SnowEntomologicalMuseumCollection,Lawrence, USA;UFMG–UniversidadeFederaldeMinasGerais,BeloHorizonte, Brazil;UFV–UniversidadeFederaldeViçosa,Viçosa,Brazil;USNM –SmithsonianNationalMuseumofNaturalHistory,Washington, USA.

Imeasuredthefollowing:eyenotchwidth(asa measureof headwidth);headlength(distancefrombaseofclypeusto ver-tex apex); glossa length; clypeus maximum width and length; supraclypeal length (distance from clypeus base to lower tan-gent of alveoli); distance between upper tangent alveolus and tangentoflowermedianocellus;intertegulardistance;first ter-gum maximum width; second tergum maximum width; total length (approximate by measuring head and mesosoma plus metasoma). All measurements are in millimeters. Relative size comparisonsusedthe midocellusdiameter(=OD)or eye max-imum width in lateral view (=EW). The punctures interspaces (i) are described in relation to puncture diameter (pd). Sculp-tureterminology follows Harris(1979) includingconflected for crowded punctures. For some sculpturing I use two terms to describemixed or transitionalpatterns, asareolate–rugose. For punctation,sparselypunctateisusedforpuncturesseparatedby more than onepuncture diameter, coarselypunctate for punc-turescomparativelycoarserthanordinarycoveringpunctures.For microsculpture, visibleonly in magnificationsabove 50×, I use theprefixmicro-,thetermpolishedisappliedfortheabsenceof microsculpture.

MostspecimenexaminationwasmadewithOlympusSZ51and SZ61usingwhiteledringillumination.Measurementsweremade withaLeicaStemiDV4andaneyepiecemicrometer.Photographs weretakenwithaNikonD700and107mmSigmaMacrolensusing HeliconRemote forcontrollingimage capture. Illumination fol-lowedthesystemofKawadaandBufﬁngton(2016).Imagestacking wasmadewithHeliconFocus (version6.0.18)–rendermethod basedonMethodC(Pyramid).Finalimageadjustments(unsharp maskﬁltering,levelcontrol,backgroundcolorsetting)werewith GIMP2.8.16(©TheGIMPTeam).

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Fig.1. PhylogenyofAugochloraclade(L:170,Ci:53,Ri:84,fastoptimization).Blackandwhitecirclesindicateuniqueandhomoplasticchanges,respectively.Rednumbers indicatecharacterstatesnotoptimizedinunambiguousoptimization.Numbersabovebranches:BootstrapandBremersupportvalues.

Results

Phylogeneticanalysis

Thephylogeneticanalysisresultedintwomostparsimonious treeswith158steps,theconsensustreeispresentedinFig.1.The Augochloracladeismonophyleticandsupportedbyﬁve synapo-morphies(Fig.1).CeratalictusandPereirapisaresistergroupswith foursynapomorphies infastoptimization(onein unambiguous optimization). Ceratalictus is supported by seven synapomor-phies(sixinunambiguousoptimization)andCeratalictusinﬂexus sp. nov. is sister group to the remaining Ceratalictus species. AugochloraandAugochlorellaaresistergroupssharingfour synapo-morphies,fromtheseonlyonesupportsthegroupunambiguously. Augochlora is monophyleticand the clade shows highsupport values compared toothergenera,itsextantsubgenera arealso corroboratedasmonophyletic.Augochlorellacomisisconsideredas sistergrouptotheremainingAugochlorellaspecies.Thethreenew speciesofAugochlorellabelongtothe“ephyragroup”fromCoelho (2004).Augochlorellakelliaesp.nov.isrelatedtoAugochlorellauna. Augochlorellaproclivasp.nov.andAugochlorellamavricerasp.nov.

constitute a clade with Augochlorella acarinata. Based on these results,thegenericclassiﬁcationfromMoure(2007)was corrob-oratedandthenewspeciesaredescribedunderAugochlorellaand Ceratalictusatthegenericlevel.

Taxonomy

Augochlorellakelliaesp.nov. (Figs.2and3)

Diagnosis.FemalesofthisspeciesaresimilartoAugochlorellauna andcanbeseparatedfromtheremainingAugochlorellabytheolive greenheadandmesosoma(notoliveinotherspecies),elongate head(widerthanlongerinotherspecies),andmicroreticulate pos-teriorandlateralsurfacesofthepropodeum(notmicroreticulatein otherspecies)(Fig.2).FromA.una,thenewspeciescanbeseparated bythemetapostnotumwithstrongposteriormarginalangles,being almosttrapezoidal(Fig.2C),preoccipitalridgecarinate, mesoscu-tumpuncturessparseronthemediansurface(interspacesequalto onepuncturediameteronA.una,andabouttwopuncture diame-teronA.kelliaesp.nov.)andthescutellarpuncturesarenotstrong andcontiguous.Themalecanbeseparatedfromotherspeciesby

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Fig.2.HolotypefemaleofAugochlorellakelliaesp.nov.(A)Habitus,(B)frontalview ofhead,(C)dorsalviewofmesosoma.Scalebar:0.5mm.

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Fig.3.ParatypemaleofAugochlorellakelliaesp.nov.(A)Dorsalview;(B)frontalview ofhead;(C)ventralviewofgenitalia.Scalebar:AandBat0.5mm,Cat0.25mm.

thecombinationofclypeuswithatransverseyellowband,hind legwithshortsetae,preoccipitalridgecarinate,andmetapostnotal carinaealmostreachingtheposteriormargin(Fig.3).

Description. Holotype female. Measurements (mm). Eye notch width:1.3;headlength: 2.0;glossa length:1.2;clypeus width: 0.7;clypeuslength: 0.5;supraclypeallength:0.4;clypeus ocel-lar distance: 0.8; intertegular distance: 1.4; T1 width: 1.8; T2 width:2.0;bodylength:7.5.Head.Labrumbasalelevation orbic-ular,black.Mandibleblackwithsubapicalamberspot;subapical toothnotstronglyproduced;longsetae(2–3OD) onouterand ventral margins. Epistomal angle orthogonal. Clypeus sparsely punctate,microreticulateonbasalportion;erectsetaewith1–1.5 OD; apical surface darkened, most surface olive green. Supra-clypeal area sparsely punctulate, microreticulate near clypeus; fineerectsetae; olivegreen. Paraocularareapunctate,strongly microreticulate;olivegreen,someweakcopperreflectionsneareye notch;branchedsetaewith1–1.5OD.Antennae:scapedarkbrown withveryweakmetallichighlights;flagellomeresamberventrally,

especiallyonF9–10;F1–2andbaseofF3setose,withinconspicuous sensilla,remainingflagellomereswithconspicuoussensilla.Frons notdepressed;conflectedpunctate,stronglymicroreticulate;weak blue iridescence;short and finesetae. Preoccipital ridgecarina notreachingthegena.Vertexcoarselypunctatebecomingrugose nearridge;longplumose(shortbranches)setae.Genapuncticulate, microlineate onpostgena;green withveryweakcopper reflec-tions;whiteplumosesetae(1.5–2OD).Hypostomalridgecarinate. Mesosoma.Pronotumdorsalridgeweaklycarinate;lateralanglenot projected,rounded;lateralridgerounded;dorsalsurfacelineate andpuncticulate;green;withfewtomentumondorsalsurface. Pro-pleuronbrownwithgreenhighlights.Prosternumgreen.Foreleg: coxadarkgreen,trochanterandfemurbrown,tibialightbrownon outersurface,baseoftibiaandpretarsusyellow;longwhitesetae oncoxatofemur,yellowerectsetaeontibiaandtarsus. Mesos-cutum anterior border weakly acuminate; median sulcus well impressed;parapsidallinesweaklyimpressed;punctatelaterally, moresparseonmediansurface,microlineationsmore conspicu-ousonanteriorsurface;olivegreen;short(=OD)andsparsewhite setae.Scutellummediansulcusveryweaklyimpressed; puncticu-late,somescatteredpunctures,polished;brightolivegreen;longer setae(=2.5–3OD)onposteriormargin.Mesepisternumruguloseon preepisternum,irregularlypunctateonhypoepimeralarea, sulci weakly foveate, densely punctate near scrobe, becoming more evenlypunctateventrally;olivegreen,weakcopperreflections; coveredwithplumosesetae(=1.5OD).Midlegbrown;yellowsetae onfemurtotarsus.Tegulaconcentriclineate;darkamber,weak greenreflectionsanteriorly.Forewing:marginalcellapexrounded; membranehyaline;blackmicrotrichia.Metanotummediansulcus veryweaklyimpressed;puncticulatetorugulose;greenwithbluish iridescence;longplumosesetae(2–3OD).Metepisternumlineate abovesuperiormetapleuralpit,rugulose–punctate below;green withcopperandbluishreflections.Hindleg:basitibialplateapex rounded;tibialspurserrate;brown;longplumosesetaeon ven-tralcoxatofemur;ventraltibialsetaeshortandbranched.Hind winghyaline.Metapostnotumweaklyinclined;weaklydepressed; apex rounded; less twice as long as metanotum; rugulose on mostsurface,exceptastripeonposteriormargin,microreticulate; olivegreen.Propodeumposteriorsurfacepuncticulate,few scat-teredpunctures,microreticulate;olivegreen;shortplumoseand someerect(1–2OD)setae;propodeumlateralsurface puncticu-late,fewscatteredpunctures,microreticulate;olivegreen;with thickerandlonger(3–4OD)setae.Metasoma.T1anterioranddorsal surfacesnotangledbetween;sparselypuncticulate,microlineate; olivegreenonanteriorandlateralsurfaces,darkbrownonmedian surface.T2puncticulate;olivegreenonanteriorandlateral sur-faces,darkbrownonmediansurface;short.T3–T4puncticulate; olivegreenwithsomedarkbrownonmediansurface,brownon marginalarea;short.T5punctate;darkbrown;coveredwithblack longerectsetae.Pygidialplateshort,truncate.S1without tuber-cle;lineate;amberwithgreenhighlightsonmediansurface;very longsetae(=3OD)onmediansurface.S2–S4lineate;browntolight brown;subapicalsetaeverylong(about5OD)andwithrecurved apices.S5lineate;brown;longsetae(about4OD),plumose.S6 lineate;brown;yellowishsetaeonapicalmargin.

Paratypemale.Measurements(mm).Eyenotchwidth:1.2;head length:1.8;clypeuswidth:0.6;clypeuslength:0.5;supraclypeal length:0.3; clypeus ocellardistance:0.7;intertegular distance: 1.3;T1width:1.2;T2width:1.3;bodylength:9.Head.Labrum andmandibleyellow.Clypeus,epistomalanglestraight;sparsely punctate,weaklymicroreticulate;whitesetaeonmiddle,longer setaeonapex;yellowstripeonapicalmargin,green.Supraclypeal areasparselypunctate,weakly microreticulate; green. Paraocu-larareasparselypunctate,weaklymicroreticulate;green,coppery alongeyemargins;long(>2OD)branchedsetae,yellow. Anten-naeF20.7xF1length,F3=F1,remainingﬂagellomeresrelatively

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longer;scapeandpedicellightbrown,flagellomeresyellowed ven-trally;F11notflattened;withoutsensillaplacodeapatches.Frons notdepressedaboveantennalsockets;coarselypunctate;bright green,fewcopperyreflectionsnearparaocular;fewerectsetae. Pre-occipitalridgecarinate,someportionslamellate.Vertexpunctate becomingrugosenearridge;verylongplumose(shortbranches) setae. Gena puncticulatebecoming lineate onpostgena; green, somecopperyreflections;whiteplumosesetae(1.5–2OD). Hypos-tomalridgecarinate.Mesosoma.Pronotumdorsalridgecarinate; lateralanglealmostorthogonal;lateralridgerounded;dorsal sur-face rugose; bright olivegreen, weakly metallic laterally; with someplumose setaeondorsal surface.Fore leg:coxa tofemur brightmetallicgreen,tibiaandtarsuslightbrown;shortyellow erectsetae.Mesoscutum anteriorborder slightlyprojected over pronotum,weaklylobed; mediansulcuswellimpressed; parap-sidallines wellimpressed; punctate,sparselyon anteriorlobe; brightolivegreen;posteriormarginweaklycoveredwithplumose setae.Scutellummediansulcusweaklyimpressed;punctate, pol-ished;brightolivegreen;anteriorsurfacecoveredwithplumose setae.Mesepisternumpunctate,polished;brightolivegreen;short plumose setae. Mid leg: coxa brown with metallic reflections, remainingleglightbrown;shortyellowerectsetae.Tegula concen-triclineate;brown,greenreflectionsanteriorly.Forewingweakly infuscateonapex.Metanotummediansulcusweaklyimpressed; puncticulate,polished;brightolivegreen;anteriorsurfacedensely coveredwitherectsetae.Metepisternumlineateabovesuperior metapleuralpit,rugulose–punctatebelow;brightolivegreen.Hind leg: coxa brown with metallic reflections, remaining leg light brown;shortyellowerectsetae,shorterthandiameterofsegments. Metapostnotum weakly inclined and depressed; twice as long asmetanotum;apexrounded;carinate,posteriorlyonlyweakly microreticulate;green.Propodeum:posteriorsurfaceposteriorpit shallow,punctate,microreticulate,olivegreen,withfinesetae; lat-eralsurfacepunctate,olivegreen,withthickersetae.Metasoma.T1 verysparselypuncticulate;lightbrownwithmetallicreflections, yellowmarginalareas.T2–5verysparsepuncticulate;lightbrown withmetallicreflections,yellowmarginalareas;vestiturewhite, withweakrowoflongwhitesetaeonbasis.T6green;longsetae. T7black;shortblacksetae.S1marginstraight;sparselypunctate, microlineate;lightbrown;withveryshortsetae.S2–S5straight; brown;coveredwithshortwhitesetaeonmediansurface,posterior surfacewithlongsetae(>4OD).S4marginconcave;withmedian, triangularpatchofsetae;lateralsetaelonger.S6weaklyconcave; brown;blacksetaeonapicalmargin.S7posteriormarginsinuose. S8posteriormarginnotproduced;0.3timesS8width.Genitalia (Fig.3C).Gonapophysisbroad,longerthangonostylusandwithout dorsalcrest;withlargeventralprongandlateralteeth.Gonobase short(0.6timesgonocoxitelength);dorsallobesweaklyproduced; ventralbridgenarrow.Gonostylusbasal process reduced; para-peniallobereduced;ventralprocesswithouterandinnerlobes, innerlobedividedandasymmetric,setose;dorsallobetransversal. Volsellalongerthanwider;oblique,cornersacuminate;without hook.

Comments.Coelho(2004)listedaA.unaparatypefemaleas“1?m#, MinasGerais,SerradaMoeda,ZonaMetalúrgica,15.x.1997(E.A.B. Almeida)(BHMH)”,theauthorlistedallspecimensofA.unawith malesymbolsbutthespeciesisonlyknownfromfemales.Asthe questionmarkcouldrepresentadubiousidentiﬁcationIspeculate thatthisparatypecanbeaspecimenofA.kelliaesp.nov.Inthis caseA.unaisknownonlyfromCentralBrazilandA.kelliaesp.nov. isknownonlyforMinasGerais.

Type material.Holotypefemale(DZUP): “Brasil, MinasGerais,\ SãoTomédasletras,\2143S,4458W\12.I.2010,K.Ramos\and V.Kanamura”.Paratypes:threefemales(DZUP)withsamedataof holotype;twofemales(DZUP)“Brasil,MinasGerais,SerradoCipó, X.2018,LeticiaGrafleg.”;onemale(UFMG)“Abelhas deCanga\

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Fig.4.HolotypefemaleofAugochlorellamavricerasp.nov.(A)Habitus;(B)frontal viewofhead;(C)dorsalviewofmesosoma.Scalebar:0.5mm.

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Fig.5. ParatypemaleofAugochlorellamavricerasp.nov.(A)Dorsalview;(B)frontal viewofhead;(C)ventralviewofgenitalia.Scalebar:AandBat0.5mm,Cat0.25mm.

ParqueEstadualda\Serrado Rola–Moc¸a\25745–75506” “Nova LimaMG\BRASIL14/03/2013\J.E.SantosJr”.

Etymology.InhonorofDr.KelliS.Ramos,friend,melittologist, andcollectorofthetype.

Augochlorellamavricerasp.nov. (Figs.4–6C,D)

Diagnosis.ThefemaleofthisspeciesisverysimilartoA. acari-nata.ThebodyofA.mavricerasp.nov.ismostlydarkgreenwith somevioletreﬂectionsonclypeusandsupraclypealareaandblack dorsalsurfacesofterga(Fig.4)whileA.acarinataisbrightgreen withsomecopperreﬂections.Also,A.mavricerasp.nov. puncta-tionissparserthanonA.acarinata(i>2pdonlowerparaocularand i>pdonmesoscutum).ThemalecanbeeasilyseparatedofA. acar-inatabythedarkbody,includinglegs(Figs.5and6D),compared withyellowishcolorofA.acarinata(Fig.6A,B).TheS4isslightly bilobed(Fig.6C)inthenewspecies,whileA.acarinatamaleshave S4stronglybilobed(Fig.6A).

Description. Holotype female. Measurements (mm). Eye notch width: 1.3;headlength:1.6;glossalength:0.6;clypeus width: 0.7;clypeuslength:0.4;supraclypeallength:0.3;clypeusocellar distance:0.9;intertegulardistance:1.1;T1width:1.7;T2width: 1.8; body length: 7.0. Head. Labrum basal elevation orbicular, black.Mandibleblackwithsubapicalamberspot;subapicaltooth

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B

Fig.6. Ventralviewofmalemetasomaandhindleg.(A)MetasomaofA.acarinata; (B)hindlegofA.acarinata;(C)metasomaofA.mavricerasp.nov.;(D)hindlegofA. mavricerasp.nov.S4apicalmarginindicatedinred.Scalebar:0.5mm.

notstronglyproduced;longsetae(2–3OD)onouterandventral margins.Epistomalangleorthogonal.Clypeussparselypunctate, weaklymicroreticulateonbasalportion;erectsetaewith1–1.5OD; apicalsurfacedarkened,subapicalareaviolet,remainingsurface brightgreen.Supraclypealareasparselypunctulate,polished;fine erectsetae;greenwithvioletreflections.Paraocularareapunctate, strongly microreticulate; olive green, someweak violet reflec-tionsneareyenotch;branchedsetaewith1–1.5OD.Antennae; scapedarkbrownwithveryweakmetallichighlights, flagellom-eres weakly amber ventrally; F1–3 setose, with inconspicuous sensilla,remainingflagellomereswithconspicuoussensilla.Frons notdepressed;conflectedpunctate,stronglymicroreticulate;olive green;shortandfinesetae.Preoccipitalridgenotcarinate.Vertex coarselypunctatebecomingrugosenearridge;longplumose(short branches)setae.Genapuncticulate,microlineateonpostgena;dark greentoblack;whiteplumosesetae(1.5–2OD).Hypostomalridge carinate.Mesosoma.Pronotumdorsalridgeweaklycarinate; lat-eralanglenotprojected,rounded;lateralridgerounded;dorsal surface lineate and puncticulate; darkgreen; withfew tomen-tumondorsalsurface.Propleuronbrownwithgreen highlights. Prosternumgreen.Foreleg:coxadarkgreen,trochanterandfemur brown,tibiabrownonoutersurface,baseoftibiaandpretarsuslight brown;longyellowishsetaeoncoxatofemur,yellowerectsetae ontibiaandtarsus.Mesoscutumanteriorborderweaklyacuminate; mediansulcuswellimpressed;parapsidallinesweaklyimpressed; punctatelaterally,moresparseonmediansurface(i=pd), micro-lineationsmoreconspicuousonanteriorsurface;greenwithviolet

reflections;short(=OD)andsparsewhitesetae.Scutellummedian sulcusveryweaklyimpressed;punctatetoconflected punctate, microreticulate;brightolivegreen;longersetae(=2.5–3OD) on posteriormargin.Mesepisternumconflectedpunctate, microretic-ulate;olivegreen,weakvioletreflections;coveredwithplumose setae(=1.5OD). Mid legbrown; yellowsetaeonfemur to tar-sus.Tegulaconcentriclineate;darkamber,weakgreenreflections anteriorly.Forewing:marginalcellapexrounded;membrane hya-line;blackmicrotrichia.Metanotum mediansulcusveryweakly impressed; rugulose, microreticulate; green with bluish irides-cence;longplumosesetae(2–3OD).Metepisternumlineateabove superiormetapleuralpit,conflectedpunctatebelow;greenwith violetreflections. Hindleg: basitibialplateapex rounded;tibial spurserrate;brown;longplumosesetaeonventralcoxatofemur, ventraltibial setaenot toolongand withshortbranches.Hind winghyaline.Metapostnotumweaklyinclined;weaklydepressed; apexrounded totrapezoidal;less twiceaslongasmetanotum; rugulose onmedian surface, carinate laterally, microreticulate; darkolivegreenwithsomevioletreflections;Propodeum poste-riorsurfacepuncticulate,fewscatteredpunctures,microreticulate togranular;darkolivegreen;shortplumoseandsomeerect(1–2 OD)setae.Propodeumlateralsurfacepuncticulate,microreticulate; olivegreen;withthickerandlonger(3–4OD)setae.Metasoma.T1 anterioranddorsalsurfacesnotangledbetween;sparsely punctic-ulate,microlineate;olivegreenreflectionsonlateralsurfaces,black onmedian surface,few copperreflections elsewhere; short.T2 puncticulate;olivegreenreflectionsonlateralsurfaces,blackon mediansurface, few copperreflections elsewhere; short.T3–T4 puncticulate;olivegreenreflectionsonlateralandmarginal sur-faces,blackonmediansurface,fewcopperreflectionselsewhere; short.T5punctate;darkbrown;coveredwithblacklongerectsetae. Pygidialplateshort,truncate.S1withouttubercle;lineate;amber withgreenhighlightsonmediansurface;verylongsetae(=3OD)on mediansurface.S2–4lineate;browntolightbrown;subapicalsetae verylong(about5OD)andwithrecurvedapices.S5lineate;brown; longsetae(about4 OD),plumose.S6lineate;brown;yellowish setaeonapicalmargin.

Paratypemale.Measurements(mm).Eyenotchwidth:1.2;head length:1.6;clypeuswidth:0.6;clypeuslength:0.4;supraclypeal length:0.3; clypeus ocellardistance:0.9;intertegular distance: 1.2;T1width:1.4;T2width:1.6;bodylength:7.0.Head.Labrum and mandible amber; Clypeus: epistomalangle straight; punc-tate,veryweaklymicroreticulate;whitesetaeonmiddle,longer setaeon apex;yellow markingon latero–apical margin, green. Supraclypealareasparselypunctate,veryweaklymicroreticulate; green.Paraocularareasparselypunctate,weaklymicroreticulate; olivegreen; long(>2 OD) branchedsetae,yellow. Antennae F2 0.8xF1length,F3=F1,remainingflagellomeresrelativelylonger; scape and pedicel light brown, flagellomeres mostly yellowed; F11flatteneddorso–ventrally;withoutsensillaplacodeapatches. Fronsnot depressedaboveantennalsockets;coarselypunctate; olivegreen;fewerectsetae.Preoccipitalridgenotcarinate. Ver-texpunctatebecomingrugosenearridge;verylongplumose(short branches)setae.Genapuncticulate,almostpolishedonpostgena; olivegreen,somecopperyreflectionsonpostgena;whiteplumose setae (1.5–2 OD). Hypostomal area, ridge carinate. Mesosoma. Pronotumdorsalridgecarinate;lateralanglealmostorthogonal; lateralridgerounded;dorsal surfacerugose;brightolivegreen, weaklymetalliclaterally;withsomeplumosesetaeondorsal sur-face.Foreleg:coxatofemurwithsomemetallicreflections,tibia and tarsus dark brown; short yellow erect setae. Mesoscutum anteriorborderslightlyprojectedoverpronotum,weaklylobed; mediansulcuswellimpressed;parapsidallineswellimpressed; punctate,sparselyonanteriorlobe;brightolivegreen;posterior margin weakly covered withplumose setae. Scutellummedian sulcusweaklyimpressed;punctate,polished;brightolivegreen;

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anteriorsurfacecoveredwithplumosesetae.Mesepisternum punc-tate,polished;brightolivegreen;shortplumosesetae.Midleg: coxabrownwithmetallicreflections,remainingleglightbrown; shortyellowerectsetae.Tegulaconcentriclineate;brown,green reflectionsanteriorly.Forewingweaklyinfuscateonapex. Metan-otummedian sulcus weakly impressed; puncticulate, polished; brightolive green; anteriorsurface densely covered witherect setae.Metepisternumlineateabovesuperiormetapleuralpit, punc-tatebelow;brightolivegreen;Hindleg:coxabrownwithmetallic reflections,remainingleglightbrown;apexofventralsurfaceof femurwithapicalverylongthicksetae(7OD), apexofventral surfaceoftibiawithlongsetae(4OD),ventralsurfaceof basitar-suswithtuftoferectsetae(3OD)onaconvexity.Metapostnotum weaklyinclined;weaklydepressed;twiceaslongasmetanotum; apexrounded;rugose, somecarinalaterally;green. Propodeum posteriorsurfaceposteriorpitshallow,punctate,microreticulate, olivegreen,withfinesetae;lateralsurfacepunctate,olivegreen, withthickersetae.Metasoma.T1verysparselypuncticulate;brown withlateralmetallicreflections;short.T2verysparse puncticu-late;lightbrownwithmetallicreflections,yellowmarginalareas; vestiturewhite,withweakrowoflongwhitesetaeonbasis.T3–T5 as T2,some bluish iridescence. T6 green;long setae. T7 black; shortblacksetae.S1marginstraight;sparselypunctate, microlin-eate;lightbrown;withveryshortsetae.S2–S3andS5straight; brown;covered withfewshortwhite setaeonmediansurface, posteriorsurfacewithsomelongsetae(>4OD).S4weaklybilobed (Fig.6C);withapicalrowofthicksetae;lateralsetaeveryshort. S6weaklyconcave;brown;blacksetaeonapicalmargin.S7 poste-riormarginsinuose.S8posteriormarginnotproduced:0.3times S8 width. Genitalia (Fig. 5C). Gonapophysis broad; longer than gonostylus;withoutdorsalcrest;withlargeventralprong;with lateralteeth.Gonobaseshort(0.6gonocoxitelength);dorsallobes weaklyproduced;ventralbridgenarrow.Gonostylus basal pro-cessreduced;parapeniallobereduced;ventralprocesswithouter andinnerlobes,innerlobedividedandasymmetric,setose; dor-sallobetransversal.Volsellalongerthanwider;oblique,corners acuminate;withouthook.

Type material.Holotypefemale(DZUP): “Parque Saint–Hilaire\ Brazil:RS,Viamão, 02/03/2017\Coletado em\Euphorbiaselloi (11–14h)\SaibreiraFogo Recente”.Paratype: one male (DZUP) “ParqueSaint–Hilaire\Brazil:RS,Viamão,21/12/2015\Coletado em\Richardiagrandiﬂora(08–11h)\NOVAFogoIntermediário”. Etymology.FromGreektermsthatmeanblack,mavros,andhorn, ceros,inreferencetotheblackF11,feminine.

Augochlorellaproclivasp.nov. (Fig.7)

Diagnosis.FemalesofA.proclivasp.nov.canbeseparatedfromthe similarA.ephyraandA.iopoecilabythecombinationoftheweakly carinatepreoccipitalridge,greenheadandmesosoma(Fig.7), rugu-loseanddeclivitousmetapostnotum,and thegranularposterior andlateralsurfacesofpropodeum.Thecarinaeofmetapostnotum reachtheedgeposteriorlydifferentlyfromA.urania,othersimilar specieswhichpresentscarinatepreoccipitalridge.Malesarenot known.

Description. Holotype female. Measurements (mm). Eye notch width:1.4;headlength:1.9;glossalength:0.8;clypeuswidth:0.7; clypeuslength:0.7;supraclypeallength:0.3;clypeusocellar dis-tance:0.8;intertegulardistance:1.4;T1width:2.0;T2width:2.2; bodylength:7.6.Head.Labrumbasalelevationweaklyorbicular, amber.Mandibleblackwithsubapicalandapicalamberspot; sub-apicaltoothnotstronglyproduced;longsetae(2–3OD)onouter andventralmargins.Epistomalangleorthogonal.Clypeussparsely punctate, polished; erect setae with 1–1.5 OD; apical surface darkened,mostsurface brightgreen.Supraclypealareasparsely punctulate,polished;ﬁneerectsetae;brightgreen.Paraoculararea

A

B

C

Fig.7.HolotypefemaleofAugochlorellaproclivasp.nov.(A)Habitus;(B)frontal viewofhead;(C)dorsalviewofmesosoma.Scalebar:0.5mm.

punctate,sometimesconflected,polished;green;branchedsetae with 1–1.5 OD.Antenna: scape dark brown with weak metal-lichighlights,flagellomeresamberventrally,especiallyonF9–10; F1–3setose,withinconspicuoussensilla,remainingflagellomeres withconspicuoussensilla.Fronsnotdepressed;conflected punc-tate;brightgreen.shortandfinesetae.Preoccipitalridgeweakly carinate,carinanot reachingthegena.Vertexcoarselypunctate becomingrugosenearridge;longplumose(shortbranches)setae. Genapuncticulate,microlineateonpostgena;greenwithveryweak bluishreflections;whiteplumosesetae(1.5–2 OD).Hypostomal ridge carinate. Mesosoma. Pronotum dorsal ridge weakly cari-nate;lateralanglenotprojected,rounded;lateralridgerounded; dorsalsurfacelineateandpuncticulate;green;withfew tomen-tum ondorsal surface. Propleura brownwith green highlights. Prosternumgreen.Forelegcoxadarkgreen,trochanterandfemur brown,tibialightbrownonoutersurface,baseoftibiaand pre-tarsusyellow; longwhite setaeoncoxato femur,yellowerect setae on tibia and tarsus. Mesoscutum anterior border weakly acuminate;mediansulcuswellimpressed;parapsidallinesweakly impressed; punctate laterally, more sparse on median surface, microlineate;green,fewcopperreflections;short(=OD)andsparse white setae. Scutellum median sulcus very weakly impressed; punctate,becomingruguloseposteriorly;brightolivegreen;longer setae(=2.5–3OD)onposteriormargin.Mesepisternumrugulose, sulci weaklyfoveate;olivegreen, weakcopperreflections; cov-eredwithplumose setae(=1.5OD). Midlegbrown,tarsuslight brown; yellowsetaeonfemur totarsus.Tegula concentric lin-eate;darkamber, weakgreen reflectionsanteriorly.Fore wing, marginalcellapexrounded;membranehyaline;blackmicrotrichia. Metanotummedian sulcusveryweaklyimpressed;puncticulate torugulose; green withbluish iridescence;long plumose setae (2–3OD).Metepisternumlineateabovesuperiormetapleuralpit, rugulose–punctate below;green withcopperand bluish reflec-tions.Hindleg,basitibialplateapexrounded;tibialspurserrate; brown; long plumose setae on ventral coxa to femur, ventral tibialsetaenottoolongandwithshortbranches.Hindwing hya-line.Metapostnotuminclined;weaklydepressed;apexroundedto trapezoidal;lesstwiceaslongasmetanotum;entirelyrugulose, somelateralcarinaextendingtopropodeum;green.Propodeum

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A

B

C

Fig.8.HolotypefemaleofCeratalictusinﬂexussp.nov.(A)Habitus;(B)frontalview ofhead;(C)dorsalviewofmesosoma.Scalebar:0.5mm.

posteriorsurfacepunctaterugulose,microreticulate;green;short plumoseandsomeerect(1–2OD)setae.Propodeumlateral sur-facepunctaterugulose,microreticulate;green;withthicker and longer(3–4OD)setae.Metasoma.T1anterioranddorsalsurfaces notangledbetween;sparselypuncticulate,microlineate;copper greenonanteriorandlateralsurfaces,darkbrownonmedian sur-face;short.T2puncticulate;coppergreenonanteriorandlateral surfaces,darkbrownonmediansurface;short.T3–T4puncticulate; coppergreenwithsomedarkbrownonmediansurface,brownon marginalarea;short.T5punctate;darkbrown;coveredwithamber longerectsetae.Pygidialplateshort,truncate.S1without tuber-cle;lineate;amberwithgreenhighlightsonmediansurface;very longsetae(=3OD)onmediansurface.S2–S4lineate;browntolight brown;subapicalsetaeverylong(about5OD)andwithrecurved apices.S5lineate;brown;longsetae(about4OD),plumose.S6 lineate;brown;yellowishsetaeonapicalmargin.

Typematerial.Holotypefemale(DZUP):“MANAUS–AM\BrasilI 61\C.ELIASleg.”.Paratype:onefemale(DZUP),withsamedataas holotype.

Etymology.FromLatintermproclivus,meaninginclined. Ceratalictusinﬂexussp.nov.

(Figs.8and9)

Diagnosis.Ceratalictusinﬂexussp.nov.isdistinguishedfromother Ceratalictusbythecombinationoflateralangleofpronotumobtuse; surfaceofmesoscutaldiskweaklymicroreticulate;carinaeof meta-postnotumnotreachingedgeposteriorly;metapostnotumsimilar withmetanotuminlengthandshape,andbythemetasomawith bluereﬂections(Figs.8and9).Thefemalehasthebasalelevation occupyingallthelongitudinalportionoflabrum,themalehasthe antennaewithsensillaplacodeapatches.

Description. Holotype female. Measurements (mm). Eye notch width:1.5;headlength: 1.7;glossa length:0.4;clypeus width: 0.8;clypeuslength: 0.4;supraclypeallength:0.3;clypeus ocel-lar distance: 1.1; intertegular distance: 1.5; T1 width: 2.0; T2 width:2.2;bodylength:8.0.Head.Labrumbasalelevation orbic-ular,black.Mandibleblackwithsubapicalamberspot;subapical

A

B

C

Fig.9. ParatypemaleofCeratalictusinﬂexussp.nov.(A)Dorsalview;(B)frontalview ofhead;(C)ventralviewofgenitalia.Scalebar:AandBat0.5mm,Cat0.25mm.

tooth not strongly produced; long setae (2–3 OD) on outer and ventral margins.Epistomal angle obtuse. Clypeus coarsely punctate, microreticulate; erect setae with 1–1.5 OD; apical surface darkened,blue green. Supraclypeal areasparsely punc-tate, microreticulate; fine erect setae; blue green. Paraocular areacoarselypunctatebecomingconflected,strongly microretic-ulate; blue green; branched setae with 1–1.5 OD. Antennae, scapedarkbrownwithveryweakmetallichighlights, flagellom-eresamberventrally,especially onF9–10;F1–2 andbaseof F3 setose,withinconspicuoussensilla,remainingflagellomereswith conspicuoussensilla. Fronsnot depressed; conflected punctate, strongly microreticulate; darkbluegreen; shortand fine setae. Preoccipital ridge carinate, carina not reaching the gena. Ver-texcoarselypunctatebecomingrugosenearridge;longplumose (shortbranches)setae.Genapuncticulate,microlineateon post-gena;bluegreen,darkgreen onpostgena;whiteplumosesetae (1.5–2OD).Hypostomalridgecarinate.Mesosoma.Pronotumdorsal ridgeweaklycarinate;lateralanglenotprojected,rounded;lateral ridgerounded;dorsalsurfacelineateandpuncticulate;bluegreen; withfewtomentumondorsalsurface.Propleurabrownwithgreen highlights.Prosternumgreen.Forelegcoxadarkgreen,remaining legdarkbrown;longwhitesetaeoncoxatofemur,yellowerect setae ontibia and tarsus. Mesoscutum anterior border weakly acuminate;mediansulcuswellimpressed;parapsidallinesweakly impressed; punctate laterally, more sparse on median surface, punctureswith different diameters intermixed,microlineations only on anterior surface and near median sulcus; blue green; short(=OD)andsparsewhitesetae.Scutellummediansulcusvery weaklyimpressed;irregularlypunctate,ruguloseposteriorly;blue green;longersetae(2–2.5OD)onposteriormargin.Mesepisternum ruguloseonpreepisternum, sulci foveate,densely toconflected punctate,microreticulate;bluegreen,blueventrally;coveredwith plumosesetae(1.5OD).Midleglightbrown;yellowsetaeonfemur totarsus.Tegulaconcentriclineate;darkamber,weakgreen reflec-tionsanteriorly.Forewing,marginalcellapexrounded;membrane hyaline;blackmicrotrichia.Metanotummediansulcusveryweakly impressed;puncticulate torugulose; blue green;long plumose setae(2–3OD).Metepisternumlineateabovesuperior metapleu-ralpit,rugulose–punctate below;green withcopperand bluish reflections. Hind leg, basitibial plate apex rounded; tibial spur serrate;brown;longplumosesetaeonventralcoxatofemur, ven-traltibialsetaenottoolongandwithshortbranches.Hindwing hyaline.Metapostnotumweaklyinclined;weaklydepressed;apex rounded;lesstwiceaslongasmetanotum;carinatetoirregularly carinate,carinaenotreachingposteriormargin,microreticulate; bluegreen.Propodeumposteriorsurfacepuncticulate,few scat-teredsetosepunctures,microreticulate;bluegreen;shortplumose

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andsomeerect(1–2OD)setae.Propodeumlateralsurface punctic-ulate,fewscatteredpunctures,microreticulate;bluegreen;with thickerandlonger(3–4OD)setae.Metasoma.T1anteriorand dor-salsurfacesnotangledbetween;sparselypunctate,microlineate; bluegreenwithsomeambertobrownportionsonmediansurface; short.T2puncticulate,weaklymicroreticulate;bluegreen,black onmarginalarea;short.T3–T4puncticulate,weakly microreticu-late;bluegreen,blackonmarginalarea;short.T5punctate;dark brown;coveredwithblacklongerectsetae.Pygidialplateshort, truncate.S1withouttubercle;lineate;amberwithgreenhighlights onmediansurface;verylongbranchedsetae(=3OD)onmedian surface.S2–S4lineate;browntolightbrown;subapicalbranched setaeverylong(about4OD).S5lineate;brown;longsetae(about4 OD),plumose.S6lineate;brown;yellowishsetaeonapicalmargin. Paratypemale.Measurements(mm).Eyenotchwidth:1.3;head length:1.8;clypeuswidth:0.6;clypeuslength:0.5;supraclypeal length:0.3; clypeus ocellardistance:1.1;intertegular distance: 1.6;T1width:1.9;T2width:2.0;bodylength:8.0.Head.Labrum andmandiblebrown.Clypeus,epistomalangleobtuse;punctate, weaklymicroreticulate; whitesetaeonmiddle, longersetaeon apex; yellowstripe on apical margin, green. Supraclypeal area sparselypunctate,microreticulate;green.Paraocularareasparsely punctate,microreticulate;green;long(>2OD)branchedsetae, yel-low.AntennaeF2=F1,remainingflagellomeresrelativelylonger; scapeandpedicellightbrown,flagellomeresyellowedventrally; F11 not flattened;F3–11 with sensilla placodea patches. Frons notdepressedaboveantennalsockets;coarselypunctate;green withbluishiridescence;fewerectsetae.Preoccipitalridge cari-nate,someportionslamellate.Vertexpunctatebecomingrugose near ridge; very long plumose (short branches) setae. Gena puncticulateweakly lineate on postgena; green, copper reflec-tionsonpostgena;whiteplumosesetae(1.5–2OD).Hypostomal area, ridgecarinate. Mesosoma.Pronotum dorsal ridgecarinate; lateralanglealmostorthogonal;lateralridgerounded;dorsal sur-facerugose;green;withsomeplumose setaeondorsalsurface. Fore leg coxa tofemur withgreen reflections, tibiaand tarsus brown; short yellow erect setae. Mesoscutum anterior border slightly projected overpronotum, weaklylobed; mediansulcus wellimpressed;parapsidallineswellimpressed;punctate,sparsely mediansurface,microreticulationsonlyonlateralsurface;green; posteriormarginweaklycoveredwithplumosesetae.Scutellum median sulcusweakly impressed;punctate,rugulose on poste-riormargin, polished;green, blue iridescenceposteriorly,weak copper reflections on middle surface; anterior surface covered withplumosesetae.Mesepisternumpunctate,densely microretic-ulate;greenwithsomecopperreflections;shortplumose setae. Mid leg coxa brown with metallic reflections, remaining leg brown;shortyellowerectsetae.Tegulaconcentriclineate;brown, greenreflectionsanteriorly.Forewingweaklyinfuscateonapex. Metanotummediansulcusweaklyimpressed;puncticulate, pol-ished;brightblue green;anteriorsurface denselycovered with erectsetae.Metepisternumlineateabovesuperiormetapleuralpit, rugulose–punctatebelow;bluegreen.Hindlegcoxawith metal-lic reflections, remaining leg brown; short yellow erect setae, shorterthandiameterofsclerites.Metapostnotumweaklyinclined; weaklydepressed;1.5×aslongasmetanotum;apexrounded; car-inatetoirregularlycarinate,carinaenotreachingposteriormargin, microreticulate;bluegreen.Propodeumposteriorsurface poste-rior pit shallow, densely punctate, microreticulate, blue green, withfinesetae;lateralsurfacepunctate,bluegreen,withthicker setae.Metasoma.T1anteriorsurfacenotpunctate,dorsalsurface punctate,polished; bluegreen, darkmarginal areashort. T2–5 punctate,microreticulate; darkbluegreen,marginalareablack; vestiturewhite,withweakrowoflongwhitesetaeonbasis.T6 green;longsetae.T7black;shortblacksetae.S1marginstraight; sparselypunctate,microlineate;lightbrown;withveryshortsetae.

S2–3straight;brown;coveredwithshortwhitesetaeonmedian surface,posteriorsurfacewithlongsetae(>4OD).S4apical mar-ginstraight;withoutmedianpatchofsetae;lateralsetaeshort. S5asS2.S6weaklyconcave;brown;blacksetaeonapical mar-gin.S7withmedianprocess.S8apicalmarginbilobed.Genitalia. Gonapophysisbroad;longerthangonostylus;withoutdorsalcrest; withlargeventralprong;withlateralteeth.Gonobaseshort(0.6 timesgonocoxitelength);dorsallobesweaklyproduced;ventral bridgenarrow.Gonostylusbasalprocessabsent;parapeniallobe present,short;ventralprocesswithouterandinnerlobes,inner lobetransversalsetoseonapex;dorsallobetransversal.Volsella longerthanwider;oblique,cornersmostlyrounded;withouthook. Variation.Mostexaminedmaterialisbluegreenbutsome speci-mensfromParanáandEspíritoSantoaregreen.Thebluereﬂections vary among specimens, including some violet reﬂections. The mesoscutumpunctuationissparserinsomespecimens.

Type material. Holotype female (DZUP), “PEVV\ 0439” “Brasil, Paraná, Parque\ Estadual de Vila Velha,\ 25◦14S 49◦59‘W,\ 12.XII.2003,G.A.R.Melo&\R.B.Gonçalves”.Paratypes(DZUP),one female, “PEVV\1056” “Brasil, Paraná, Parque\ Estadualde Vila Velha,\ 25◦ 14S49◦59‘W,\7.iii.2003,G. Melo&\R.Gonçalves”, onefemale,“PEVV_\0072”“Brasil,Paraná,Parque_\EstadualdeVila Velha,\25◦ 14S49◦59‘W,\05.x.2003,G.Melo&\R.Gonçalves& A.Aguiar”,twofemales“Brasil,Paraná,Parque\EstadualdeVila Velha,\25◦14S49◦59‘W,\16.iii.2002,G.Melo&R.B.\Gonçalves”, onemale“PEVV\1161”“Brasil,Paraná,Parque\EstadualdeVila Velha,\25◦14S49◦59‘W,\7.iii.2003,G.Melo&\R.Gonçalves”. Additionalexaminedmaterial.BRAZIL,EspíritoSanto:onefemale (DZUP),“SANTA TERESA– ES\BRASIL 22/X/966\C.and T.Elias leg.”.MinasGerais:fourfemales(UFV),“LocalMEUFV.MG\Data 19/02/85\ Col. L. Campos”; one female (UFV) with same data except“18/12/85\A.A.Soares”;onefemale(UFV),“MEUFVMG\ 11/10/1987\A.A.Soares”.Paraná:onefemale(DZUP),“ANTONINA PR\Brasil20–I–1966\Marinoni[and]Azevedo”.Rio deJaneiro: one female (USNM), “RdJaneiro_\ Brazil_\ X 1938””YelFevServ_\ MESBrazil\ RCShannon“. Santa Catarina: one female (SEMC), “BRAZIL\SantaCatarina\NovaTeutonia\15Sept.1955\(F. Plau-mann)“; twofemales (SEMC),“NovaTeutonia,\SantaCatarina,\ BrazilIX–1951\L.E.Plaumann).SãoPaulo:twofemales(MZSP), “Brasil, São Paulo, Iporanga, 10.ix.2000, B. H. Dietz col.”, one female(DZUP),“Brasil,SãoPaulo\Cajuru,Faz.Rio\Grande,21◦12S\ 47◦09‘W02.xii.1999,\GabrielA.R.Melo”.

Etymology.FromtheLatinterminﬂexusmeansbend,curve,in ref-erencetotheformoftheinnerlobeofventralprocessofgonostylus.

Discussion

ThisstudycorroboratesthemonophylyoftheAugochloraclade recovered by previous studies (Eickwort, 1969; Danforth and Eickwort,1997;Engel,2000;Coelho,2004;Gonçalves,2015,2016). ThetopologyofGonçalves(2016)iscorroboratedincontrastto pre-vioushypothesis(Eickwort,1969;Danforth andEickwort,1997; Engel, 2000;Coelho,2004;Gonçalves,2015)(Fig.1).Inspiteof thepeculiargenitalcapsule,Ceratalictusinflexussp.nov.issister lineagetotherestofthespeciesofthegenus.Ceratalictusis sup-portedbysevensynapomorphies.Thephylogeneticrelationships amongtheremainingCeratalictusspecieswerethesameasfound byGonçalves(2015).Ceratalictus’distributionshows biogeograph-icalduplicationsinsoutheasternSouthAmerica(Gonçalves,2015) andthebasalpositionofthenewspeciesreinforcesthehypothesis oftheoriginofthegenusinthisregion.

ThetopologyfoundforAugochlorellaisdifferentfromthatof Coelho(2004)duetheplacementofA.comis.Thisspeciesis consid-eredheresistertotherestofthegenus,incontrasttotheprevious studywhereitwasconsideredasrelatedtothe“auratagroup”.The

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maleF2shorterthanF1andtheconcaveS4aresynapomorphiesfor AugochlorellaexcludingA.comis.Theconvexfaceandthealmost obtuseepistomalangleofA.comisarealsoplesiomorphicfeatures fromotherAugochlorellaspeciesandsupportthisnewphylogenetic hypothesis.Coelho(2004)postulateavicariancebetweenA.ephyra andA.aurata(includingA.comis)groups.Inthepresentstudy,the hypothesispoint toa differentinterpretationwiththeoriginof AugochlorellainCentralAmericaandonlylateroccupyingNorth andSouthAmerica.Moreeffortisnecessarytounderstandthe bio-geographicalevolutionofAugochlorella.Thisstudydescribedfour newspeciesoftheAugochloracladeandprovidedarevised phy-logenyfortheplacementofthesespecies.Thiswastheﬁrstattempt toincludeallspeciesofAugochlorellaandCeratalictusina phyloge-neticanalysis.Thereisstillalotofsystematicworktobedonewith theclade:Pereirapishasneverbeenstudiedindetail;Augochlora taxonomyand systematicsis still very poorly investigated due tolargenumberofundescribedspecies;andtherelationshipsof Augochlorellaspeciesremainswithunresolvednodes.

Funding

This study was partially supported by Brazilian Conselho Nacional de Desenvolvimento Cientíﬁco e Tecnológico, number 443069/2014-0.

Conﬂictsofinterest

Theauthorsdeclarenoconﬂictsofinterest. Acknowledgements

ThankstoCarlosRobertoBrandãoandGabrielMelofor assis-tance during the execution of this study.I am grateful to ﬁve anonymousreviewersforsuggestionsonthemanuscript.

AppendixA. Supplementarydata

Supplementarydataassociatedwiththisarticlecanbefound,in theonlineversion,atdoi:10.1016/j.rbe.2018.12.003.

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