REVISTA
BRASILEIRA
DE
Entomologia
AJournalonInsectDiversityandEvolution w w w . r b e n t o m o l o g i a . c o mSystematics,
Morphology
and
Biogeography
Redefinition
and
taxonomic
revision
of
the
“buqueti”
species-group,
Dichotomius
Hope,
1838
(Coleoptera:
Scarabaeidae:
Scarabaeinae)
Jorge
Armando
Arias-Buriticá
a,∗,
Fernando
Z.
Vaz-de-Mello
baUniversidadPedagógicayTecnológicadeColombia,GrupodeinvestigaciónSistemáticaBiológica,EscueladeCienciasBiológicas,Tunja,Colombia bUniversidadeFederaldeMatoGrosso,InstitutodeBiociências,DepartamentodeBiologiaeZoologia,Cuiabá,MT,Brazil
a
r
t
i
c
l
e
i
n
f
o
Articlehistory:Received17August2018 Accepted12November2018 Availableonline23November2018 AssociateEditor:MarcelaMonné Keywords:
Taxonomy Dungbeetles
Dichotomiussensustricto Neotropical
Malegenitalorgan
a
b
s
t
r
a
c
t
Onthebasisofexternalmorphologyandgenitaliaofmales,aswellasacomparisonbetweendifferent speciesbelongingtoothergroupsofDichotomius,weproposearedefinitionofthe“buqueti”species groupseparatedbyLuederwaldt(1929).Sixspeciesareexcludedfromthisgroupandtransferredto othergroups:D.ribeiroi(Pereira,1954)inthe“cotopaxi”group;D.camposeabraiMartínez,1974inthe “bitiensis”group;D.reclinatus(Felsche,1901),D.horridus(Felsche,1911),D.quadrinodosus(Felsche,1901) andD.nimuendaju(Luederwaldt,1925)(revalidatedspecies)formingthenewlydefined“reclinatus” group.Theotherspecieskeptinthe“buqueti”groupincludeD.buqueti(Lucas,1857)fromBrazil(lectotype heredesignated),D.haroldi(Waterhouse,1891)fromArgentinaandD.nutans(Harold,1867)fromBrazil, ArgentinaandUruguay.Thetaxonomicrevisionofthe“buqueti”grouparepresented,includingahistory oftheincludedspecies,adeterminationkey,illustrationsofthestructuresofexternalmorphology,male genitaliaandscleritesoftheinternalsac,andadistributionmap.
©2018SociedadeBrasileiradeEntomologia.PublishedbyElsevierEditoraLtda.Thisisanopen accessarticleundertheCCBY-NC-NDlicense(http://creativecommons.org/licenses/by-nc-nd/4.0/).
Introduction
DichotomiusHope,1838isoneofthemostdiverseandabundant generaofdungbeetles(Scarabaeidae:Scarabaeinae)inneotropical ecosystems,withabout154species,inwhichataxonomic revi-sionisurgentlyneeded(Génier,2000;Vaz-de-Melloetal.,2001; Kohlmann,2003).Although,afewauthorshaveworkedingroups ofspecies(Arias-BuriticáandVaz-de-Mello,2012,2013;Maldaner etal.,2015;Nunesetal.,2016;Valoisetal.,2017),thegenushasnot undergonearecentgeneraltaxonomicrevision.Thelastresearcher tostudy the groupas a whole wasLuederwaldt (1929). Based onexternal morphologicalcharacters, this authorproposedthe distinctionofthespeciesintosections(herepresentedas species-groups)tostudythegenus.Withinthenominotypicalsubgenus, thisauthorproposedthe“buqueti”species-groupforspecieswith thefollowingfeatures:“triangularhead,atleastinthemale,which issometimesextremelylongasinthebuquetispecies.Clypeal mar-ginslightlyemarginate,withtransversewrinkles.Headarmor,in males,betweentheeyesorinfrontofthem,withamediumhorn, fullycompressed,generally lamelliform and witha shortspine
∗ Correspondingauthor.
E-mail:joaariasbu@gmail.com(J.A.Arias-Buriticá).
ortoothoneachsideofthebase;inthefemale,intheposterior clypealmargin,transverselywiththreeorfourprojections.Male thoraxstrongorverystronglyexcavated,inthefemale,thethorax excavationissmoothandwithoutthepresenceofprominences. Usuallyblackincolor,rarelybrown. Striaeofelytranotevident withfinepoints.Spurofmetatibiaeusuallyemarginate.Medium tolargebody size”.SensuLuederwaldt (1929)eight speciesare includedinthegroup:Dichotomiusbuqueti(Lucas,1857)(Minas Gerais,Brazil);Dichotomiusharoldi(Waterhouse,1891)(Chacode Santiago,Argentina);Dichotomiushorridus(Felsche,1911)(Bucay, Ecuador); Dichotomius nimuendaju (Luederwaldt, 1925) (Ama-zonas,Brazil);Dichotomiusquadrinodosus(Felsche,1901)(southern BrazilinthestatesofMinasGerais,SantaCatarina,EspiritoSanto and Paraná); Dichotomius verticalis(Felsche, 1901)(Argentina); Dichotomiusreclinatus(Felsche,1901)(Colombia)andDichotomius gibbosusLuederwaldt(1928)(Argentina).
Martínez(1951) proposesthesynonymyofD. gibbosus with Dichotomiustriangulariceps(Blanchard,1846)andPereira(1953) that of D. haroldi with D. verticalis. Pereira (1954) describes DichotomiusribeiroifromthestateofAmazonasinBrazil;raising theknownspeciesinthegrouptonine(includingD.haroldiand D.verticalisasdistinctspecies);however,heproposedthatithas beenlittlestudiedandthatindividualsarerareincollections:the femalesofatleastfivespeciesareunknown.Finally,heelaborates
https://doi.org/10.1016/j.rbe.2018.11.002
0085-5626/©2018SociedadeBrasileiradeEntomologia.PublishedbyElsevierEditoraLtda.ThisisanopenaccessarticleundertheCCBY-NC-NDlicense(http:// creativecommons.org/licenses/by-nc-nd/4.0/).
akeyforthespeciesofthesectionbasedonbodysculpture char-acteristicsandforthefirsttimethemalegenitalorgansofsomeof thespeciesareillustrated.
Martínez(1974)describesDichotomiuscamposeabraifroman individualcollectedinthestateofEspiritoSanto(Brazil)and pro-posestheexclusionofD.triangularicepsfromthe“buqueti”group and the incorporation of Dichotomius nutans (Harold, 1867), a speciesthathadbeenlocatedbyLuederwaldt(1929)inaseparate group(the“nutans”group).AssumingthesynonymyofD.haroldi andD.verticalis,althoughwithadifferentconfiguration,thegroup remainswithninespecies.
Vulcanoet al.(1976)include withinthesectionDichotomius calcaratus, species described by Arrow (1913) from individuals foundinthestateofMatoGrosso(Brazil),whichwereincludedby Luederwaldt(1929)withinthe“assifer”group,Vulcanoetal.(1976) commentthatthissituationoccursbecauseLuederwaldtassumed asmalesthetypespecimensinthedescriptionofArrow. Subse-quently,throughcollectionsmadeinthetypelocalityoftheArrow individuals,malesandfemalesofthisspecieswerefound,themales beingthosedescribedasD.nimuendajubyLuederwaldt(1925)and theyproposeD.nimuendajuasasynonymofD.calcaratus.
Consequently, the “buqueti” species group includes nine species:D.buqueti,D.nutans,D.haroldi,D.quadrinodosus,D. reclina-tus,D.horridus,D.calcaratus,D.ribeiroiandD.camposeabrai(Pereira, 1954;Martínez,1974;Vulcanoetal.,1976).
Nevertheless,groupinconsistenciesbetweentheexternal mor-phologyofsomeofthespeciesandthecharacteristicsproposedby Luederwaldt(1929)werefound.Therefore,basedonotherexternal morphologicalcharacters, themalegenital organand the com-parisonofthespecieswithotherspeciesofthegenus,thispaper proposestheredefinitionandtaxonomicrevisionofthe“buqueti” species-group.
Materialsandmethods
Thisstudywasbasedontheexaminationof383specimensof ninespecies inthe“buqueti” species-groupand apairfrom 37 speciesinotherspecies-groupsandDichotomiussubgenera.These specimensweredepositedinthefollowingentomological collec-tions(nameofeachcollectioncuratorinparentheses):
BMNH:TheNaturalHistoryMuseum,London,England(Maxwell BarclayandMalcolmKerley)
CEMT: Sec¸ão de Entomologia da Colec¸ão Zoológica da Uni-versidade Federal de Mato Grosso, Cuiabá, Brazil (Fernando Vaz-de-Mello)
CMNC:CanadianMuseum ofNature, Ottawa,Canada(Franc¸ois Génier)
IBSP:Colec¸ãoEntomológica“AdolphHempel”,InstitutoBiológico, SãoPaulo,Brazil(SergioIde)
MAPA:MuseuAnchieta,PortoAlegre,Brazil(FernandoR.Meyer)
MNHN: Muséum National d’Histoire Naturelle, Paris, France (OlivierMontreuilandAntoineMantilleri)
MZSP:MuseudeZoologiadaUniversidadedeSãoPaulo,SãoPaulo, Brazil(SoniaCasariandCarlosCampaner)
SMTD:StaatlichesMuseumfürTierkunde,Dresden,Germany(Olaf Jäger)
Speciesidentificationwasperformedusingtheoriginal descrip-tions(Arrow,1913;Felsche,1901,1911;Harold,1867;Lucas,1857; Luederwaldt,1925;Waterhouse,1891;Pereira,1954)andspecific keys(Luederwaldt, 1929;Pereira, 1954).For thepreparation of specimens,theMedinaetal.(2003)methodologywasfollowed. Malegenitalia(aedeagus)dissectionandpreparationwascarried outundertheZuninomethodology(1978).
Illustrations of the male habitus and differences within the pronotumandheadoffemales,aswellasillustrationsofaedeagus inlateral,dorsalandventralviewsandinternalsacscleriteswere produced.Terminologyoftheexternalmorphologywasassigned accordingtoVaz-de-Melloetal.(2011).Formalegenitalia,Zunino (1978)andLópez-Guerrero(2005)nomenclaturewasfollowed.
For eachspecies, thetaxonomictreatmentdeveloped shows thefollowinginformation:Locationofthetypespecimens, nomen-claturalhistory,materialexamined,redescriptionofthemaleand female,commentsandgeographicaldistribution.Forthelatter, dis-tributionmapsweremadewithgeographicaldataobtainedfrom locallabelsofthespecimens,usingthesoftwareDIVAGISVer7.5 (Hijmansetal.,2012).
Resultsanddiscussion
Vulcano etal. (1976)includedD. calcaratusin the“buqueti” species-group.Afterrevisionoftheholotypeandduetothe mor-phologyoftheheadandthethoraxofthisspecies,itisconcluded thatithasnorelationshipwiththe“buqueti”species-group.Inthe samepaper,theseauthorssynonymizedDichotomiusnimuendaju (Luederwaldt,1925)withD.calcaratus,whichwasamistakeand, followingtherevisionoftheholotypeofD.nimuendaju,thisspecies isrevalidated.Therefore,the“buqueti”species-groupwould con-sistofthefollowingspecies:D.buqueti,D.camposeabrai,D.nutans, D.haroldi,D.horridus,D.quadrinodosus,D.nimuendaju,D.reclinatus andD.ribeiroi.
Bystudyingtheexternalmorphologycharacteristicsproposed by Luederwaldt (1929) for the “buqueti” species-group, it was noticed that the triangularshape of the head and the laminar cephalichorninmaleswerenotrelated,featuresnotexhibitedby D.nimuendaju,D.camposeabrai,D.quadrinodosus,D.horridusand D.reclinatus.Thesespecieshave anovalheadwithtwodefined clypeusteethandasub-cylindricalcephalichorn(Fig.1).
Forthisreason,thestudywasexpandedtoincludemale gen-italiamorphologicalcharacteristics,whichhaveshownagreater consistencythanintraspecificexternalmorphologycharacteristics (Zunino,1985;ZuninoandMonteresino,1990).However, inconsis-tenciesinthe“buqueti”species-group(sensuLuederwaldt)were higheranditisthereforeconcludedthatthisgroupconsistsofa mixtureofspecies.Thespeciesinthe“buqueti”species-group pre-sentedtwomorphologiesinmalegenitalia,whereD.buqueti,D. haroldi,D.nutans,D.ribeiroiandD.camposeabraipresented aedea-guswithtwolaminarprocessesbetweenparameresinventralview andD.nimuendaju,D.quadrinodosus,D.horridusandD.reclinatus donothavethisstructurebetweenparameres(Fig.2).
Theinternalsacscleritesalsoshowdifferencesbetweenspecies, mainlyinthecopulatrixlamella:thestructurethataccordingto López-Guerrero(2005)iswherethebiggestdifferencesbetween Dichotomiusspeciesarefound.Inthe“buqueti”species-groupwe foundthatD.buqueti,D.haroldi,D.nutansandD.ribeiroihavea sub-quadrangularcopulatrixlamellawithtwoprojectionsonthe rightside andsetaeacrossthewholesurface(Fig.3A–D),while D.nimuendaju,D.quadrinodosus,D.horridusandD.reclinatushave awellsclerotizedandirregularcopulatrixlamella(Fig.3F–I)and D.camposeabraihaveadifferentlamellatothosedescribedabove (Fig.3E).
Asa result of the aforementionedand according toa com-parison of the external morphology and male genitalia with other Dichotomius groups, we propose the redefinition of the “buqueti”species-group,inwhichshouldbeD.buqueti,D.haroldi and D. nutans,in the following terms: aedeagus with laminar processes between parameres, parameres with setae at apex, sub-quadrangularcopulatrixlamellawithtwolateralprojections andsetaeoverthewholesurface,triangularhead,malecephalic
A.
E.
F.
G.
H.
I.
B.
C.
D.
Fig.1.Headsof“buqueti”speciesgroup.(A)D.buqueti;(B)D.haroldi;(C)D.nutans;(D)D.ribeiroi;(E)D.camposeabrai;(F)D.horridus;(G)D.nimuendaju;(H)D.quadrinodosus; (I)D.reclinatus.Linescale1mm.
A.
E.
F.
G.
H.
I.
B.
C.
D.
Fig.2. Ventralviewoftheparameresof“buqueti”speciesgroup.(A)D.buqueti;(B)D.haroldi;(C)D.nutans;(D)D.ribeiroi;(E)D.camposeabrai;(F)D.horridus;(G)D. nimuendaju;(H)D.quadrinodosus;(I)D.reclinatus.Linescale1mm.
processat leasttwo-thirds inlaminar form endingin a conical processdirectedforwardandpronotumwithexcavationat1/3of theanteriorregion.
Weproposeanewspeciesgroup,whichwillbecalled “recli-natus”,inwhichD.horridus,D.nimuendaju,D.quadrinodosusand D.reclinatusareincluded,withthefollowingfeatures:malesand femaleswithrounded,widerthanlongheads,withaflange exten-sionacrossclypeus,withtwosoftand emarginatedteethinthe center, male withcentral cephalic processdirected toward the back,long,sub-cylindricalandbluntendingandwithtwominor processesatthebase(exceptD.reclinatusandD.horridus,which areintheclypeus).Pronotumwithsharpantero-lateralangle,the maleshaveamiddleexcavationwhichcontinuesuptotwo-thirds ofitslength.Aedeagusinventralview,withoutlaminarprocesses betweenparameres.Paramereswithstraightinneredge,withbasal areaextendedtothephallobaseandsingleinvaginationoneither
side ofthis projection.Irregularand wellsclerotized copulatrix lamella.
D.ribeiroihasatriangularhead,thepronotumanterior exca-vation andthecopulatrix lamellais asspecies ofthe“buqueti” species-group. However, this species is here transferred to “cotopaxi” species-group based onthe following characteristic: head,pronotumandelytrawithcallosity,dimpledelytralstriae, base ofstriae 1–5with depressions,invagination in the poste-rior area of thelateral portion of the pronotum,the shape of theparameresandinternalsacsclerites.Pereira(1954)proposed thesimilarityofthisspecieswithDichotomiuscotopaxiowingto characteristicsofthebodycallusesbutplaceditinthe“buqueti” species-groupbasedonlyonthetriangularshapeofthehead.
Finally, D. camposeabrai, it seems to be a species near Dichotomiusnemoricola(Pereira,1942)becauseofcharacteristics such as the shape of the parameres, typical meaty copulatrix
A.
E.
F.
G.
H.
I.
B.
C.
D.
Fig.3. Copulatrixlamellaeof“buqueti”speciesgroup.(A)D.buqueti;(B)D.haroldi;(C)D.nutans;(D)D.ribeiroi;(E)D.camposeabrai;(F)D.horridus;(G)D.nimuendaju;(H) D.quadrinodosus;(I)D.reclinatus.Linescale1mm.
lamellaprocesswithlongbristlesontheleft,aswellastheshapeof thepronotum,pronotalpunctuation,elytralstriaeandelytral inter-striae.Therefore,thisspeciesshouldbeincludedinthe“bitiensis” species-grouptowhichD.nemoricolabelongs(Arias-Buriticáand Vaz-de-Mello,2012).
Redefinitionof“buqueti”species-group
Thespeciesofthe“buqueti”species-grouparerecognizedby thefollowingcharacteristics:(1)malesandfemaleswithtriangular head.(2)Clypeusinmostwidelyemarginatedandflat,withstrong transversewrinkles.(3)Maleswithlamelliformcephalicprocess, sub-quadrangularatthebaseandendingattheapicalconical pro-cessdirectedforward;withasmallerprocessoneachsideatthe baseoftheleadingedgeattheeye.(4)Femaleswithafour-pointed processbetween theeyes (except in D. haroldi).(5) Males and femaleswithexcavatedpronotuminanteriorthird,withtwo cen-traltubercles,beingstrongerinmalesthaninfemales.(6)Pygidium withincompleteposteriorridgeinthemiddle.(7)Aedeaguswith basalareaofparameresendinginastrongconstrictiongivingthe appearanceofspinaandlamelliformprocessbetweenparameres. (8)Paramereswithsetaeatapex.(9)Sub-quadrangular copula-trixlamellawithtwo extensionsontherightandsetaepresent throughoutthesurface.
Keyforthespeciesofthe“buqueti”species-groupand speciesthatmaybeconfused*
1.Malesandfemaleswithcallusesonhead,elytraandpronotum givingacorrugatedappearance ... D.ribeiroi*
-Malesandfemaleswithoutcallusesonhead,elytraand prono-tum ...2
2.Males and femaleswithelytral striaeof onethrough five widenedinthebasalarea(Fig.7A)... D.nutans(Fig.7)
-Malesandfemaleswithoutwidenedelytralstriae(Figs.5A–6A) ...3
3.Largesize(27–37mm),elytrawithblueovertones.Maleswith long,acutecephaliclateralprocesses(2mmormore),femaleswith fourpointprocess...D.buqueti(Fig.5)
Fig.4. Dichotomiusbuqueti.LabelsofLectotypeheredesignated.
- Medium size (18–28mm), black elytra. Males with small lateralcephalicprocesses(lessthan1mm),femaleswithcephalic processsimilartotheprocessesofthemale... D.haroldi
(Fig.6)
Dichotomiusbuqueti(Lucas,1857) (Figures4,5A–J,8A)
CoprisbuquetiLucas,1857:Page105(originaldescription). Pinotusbuqueti(Lucas,1857):Page1009;Gemminger,M.,Harold, E.,1869.(TransfertoPinotus).
TypeMaterial:Lectotypemale(heredesigned):[1,quadrangular andredlabel,handwritten]Buquetimihi,Tridens,Buquet, [2,quad-rangularandyellowlabel,handwritten]5089.1♂.[BMNH](Fig.4).
Additionalspecimens(20♂♂.24♀♀.):BRAZIL:1♂.[MZSP].Minas Gerais:PassaQuatro,FazendaDosCampos,6/IV/1914,J.F.Zikan 1♀ [MZSP];IX/1917 1♀. [MZSP]; XI/1918 1♀. [MZSP]; 1919 1♂. [MZSP]. Vila Monte Verde, 15/IV/1960, J. Halik. 1♂. [MZSP]; 17/III/19661♂. [MZSP];18/III/1966,3♂. [MZSP];18/IV/1966,1♀. [MZSP];18/IV/19661♂.[MZSP];21/XI/19661♀.[MZSP];23/II/1969 1♀. [MZSP];26/VII/1968 1♀. [MZSP]. Monte verde, 16/VII/1959, ReichardCol.1♀.[MZSP].PassaQuatro,915m,2/IV/1922,J.F.Zikan 2♂.[1♂.MZSP;1♂.IBSP].RiodeJaneiro:Itatiaia,16/III/1989, God-inho,C. 1♂. [CEMT]; IV/1992, 1♀. [CEMT]; II/1993,1♂. [CEMT]; 1/III/1993, Celso, J. 1♂. [CEMT]. Nova Friburgo, Campo Coelho,
A.
E.
F.
G.
H.
I.
J.
B.
C.
D.
Fig.5.Dichotomiusbuqueti.(A)Dorsalhabitusofthemale.(B)Lateralviewofaedeagus.(C)Dorsalviewofparameres.(D)Ventralviewoftheparameres.(E)Copulatrix lamellae.(F)Centralaccessorylamella.(G)Leftlateralaccessorylamella.(H)Rightlateralaccessorylamella.(I)Headoffemale.(J)Thoraxoffemale.Linescale1mm. I/1996,Penna,F.1♂.1♀.[CEMT].Petrópolis,22/I/1899.1♀.[MZSP]
São Paulo: Campos [do] Jordão, 5/I/1960, J. Halik. 1♀. [MZSP]; CamposdoJordão,I/2004,Almeidaneta,G.P1♀.[CEMT];19/I/2004, Pereira,E.1♀.[CEMT];5/II/2004,Pereira,E.1♂.1♀.[CEMT];1800m, I/2005,Pereira,E.2♀.[CEMT].SerradaBocaina,SanJosedoBarro, I-II/1960,M.A.Vulcano.3♂.8♀.[MZSP].
Males:Length 27–37mm; width 17–21mm. Blackcolor.Elytra withblueovertones(Fig.5A).
Head:Longerthanwide,withprolongedanteriormargingiving theappearanceofablunttriangle.Clypeuswithabrightflange, withoutclypealteeth.Surfaceofclypeuswithstrong,parallel wrin-kles.Genaewithcurvedandpronouncedanterioredge,lateraledge straightandthencurved,surfacewithstrongwrinkles. Clypeus-genalsuture evident,reachingthecephalic process. Fronswith threehorns;thecentralandmostclosetobackedgelookinglike aprolonged“C”,dorsoventrallyflattenedinthebasalregionat lat-eralview,inthesecond-thirdgettingcompressedandendingwith acylindrical,brilliantandblunttip;infrontalviewitshowsa cen-tralkeel.Thelateralhornsareconical,with2mmormore,wide baseandendingwithcylindrical,brilliantandblunttips,directed upwardsanddiagonaltothebaseofthecentralhorn,reachingits middle.
Thorax:Pronotum twiceas wide aslong. Surface withcircular punctures,elongatedtowardthelateralmarginsandareseparated bytwiceorthreetimestheirlength.Anteriorregionwithawideand smoothedge,whichgetscompressedtowardtheanteriorangles. Theanteriorangleiscurvedforminganangleofapproximately 90◦.In sideviewofpronotum,theanterior regionis verticalto thedorsalregion,straightinthecentralregionandcurvedtoward thelateralregion.Pronotumwithtwoshortandcurvedtubercles onthecentralregion;fromthesetuberclescometwo protuber-anceslikehumpstowardthelateralzonesgivingtheappearance ofanexcavationattheanteriorareaofthepronotum.Behindand
betweenbothtubercles,thereisanelongatedanddeepfoveathat reachestheposteriorregionofthepronotum.Bothlateralfoveae are big,deepand dorsiventrally enlarged. Hypomere with sha-greenedsurfaceanddensesetigerouspunctures;lateraledgeswith erect,longandbrownsetaewhichcanbeseenatdorsalviewand lessdenseatthecentralregionwheretheyareseparatedbytwice theirdiameter.Atthecentralregionunderthefemur,thesurface issmooth andbrilliant.Shagreenedprosternumwithsetigerous puncturesprolongedfromitsmediumregionuntilcovering the antero-medialzoneofthemesosternum.Fromtheposterioredge oftheprosternumemergemanyyellowsetaedirectedtowardthe mesosternum.Elytrawithsoftandbicarinatestriae,withocellated puncturesspacedbyaboutfivetimestheirdiameter;thebaseofthe firstfivestriaehasanirregularfovea.Interstriaewithshagreened surfacewithverysoftpuncturesseparatedbetweenthembyseven timestheirdiameterobservedat20×.Insomespecimensthereare transversewrinklesneartothestriae.Mesosternumstrongly nar-rowedmedially,onwhichthereisabrilliantandsmoothprocess thatprecludesseeingthemeso-metasternalsuture,thereisa sha-greenedsurfacetowardthesideswithsetigerouspuncturesand yellowsetae.Mesepisternumwiththesamesurfaceasthelateral edgesofthemesosternum.Themeso-metasternalsutureiserased inthecentralareabyamesosternumprocess,lateralmarginsare evidentand brilliant.Metasternumwithshagreened,shiny cen-tralareawithoutsetae;strongcarinafrom1/3ofmetasternumto thebackendinginadeepfovea.Sideedgesoftheanteriorarea shagreenedwithdensesetigerouspunctures,posteriorarea sha-greenedwithlargesetigerouspunctures.Metepisternumequalto mesepisternum.
Abdomen: Sternites shagreened with somesoft and separated punctures, evident in the 5th and 6th sternites. Lateral edges ofallsterniteswithdenseocelatedpunctures.6thsternitevery narrowed medially. Pygidium shagreened with elongated and
separatedpuncturesatleasttwiceitsdiameter,incompletemargin inthemiddleinferiorregion.
Malegenitalorgan:Sideviewofaedeaguswithsub-quadrangular phallobase,basalzonewithamediumbulgeatbothsides,apexwith a constriction of approximately115◦.Sub-triangular parameres withathinprolongationattheinferiorbase,whichcontinueswith aninvaginationtowardthemiddlezone;apexthin,blunt,witha flatsurfacewherelongandyellowsetaeemerge(Fig.5B).Dorsally symmetricwithenlargedbaseandbecomingthinnertowardapical zoneatwhichendingwithablunttip;inthisviewsetaecanbeseen too.Internalregionwithaconvexinvaginationtowardthemiddle untiltheapicalregionofeachparamere(Fig.5C).Intheventral view,parameresarethinatthebasalzonewithanacute prolonga-tiondescribedinthelateralview:apexbluntandflatwheresetae canbeseen.Betweenparameresandinsertedfromtheapicalto thebasalzonebyamembrane,therearetwolamellatedand over-lappedprocesses,thatoftheleftparamereoverthatoftheright paramereinacharacteristicwaywhichgoesfromtheapicalzoneto beyondtwo-thirdsoftheparameres(Fig.5D).Internalsactubular withraspulesatbasal1/3;atthecenteronebig,sub-quadrangular, browncolorcopulatrixlamellaewithtwolateraltubularanddarker prolongations,wholesurfacecovered bybristleswhich become longertoward lateralmargins(Fig.5E).Apical zonewith three accessorylamellae,onelongandcentralwithundefinedform,more sclerotizedinthecentralareawithasurroundingsemi-quitinized membrane(Fig.5F).Leftlaterallamellahas“C”form,withaflat base,wellsclerotizedwithirregularborders(Fig.5G).Right lat-erallamellahas“N”form,enlarged,sclerotizedinthecenterand surroundedbyasemi-quitinizedmembrane(Fig.5H).
Female:length26.5–37mm;width16.8–21.1mm.Itdiffersfrom malesbypresentingamoredefinedclypeusedge.Fronswithafour pointedcephalicprocess;bothcentraltipshigherandattached, lat-eralonesconicalandsmaller(Fig.5I).Centraltuberclesofpronotum aresofter(Fig.5J).6thabdominalscleritenotshortenedmedially.
Commentaries:Thespeciesisdistinguishedfromtheothersofthe group“buqueti”becausethemalespresentaprocessofthreehorns inthefrontwherethetwolateralhornshavetwoormoremmand thefemaleswithfourpointedcephalicprocess,pronotumwithtwo shortandcurvedtuberclesonthecentralregionandelytrawith blueovertones.ThespeciesisassociatedwithareasofAtlantic for-estwithaltitudesofbetween900mand1200minthemountains ofMantiqueiraandÓrgãosinBrazil.
Distribution:ThisspeciesisdistributedinBrazilintheSãoPaulo, RiodeJaneiroandMinasGeraisstates(Fig.8).
Dichotomiusharoldi(Waterhouse,1891) (Figs.6A–J,8B)
PinotusharoldiWaterhouse,1891:page359(originaldescription). PinotusverticalisFelsche,1901:page136(originaldescription). Dichotomiusverticalis(Felsche,1901):page389.Pereira,F.S.1953 (synonym).
TypeMaterial:PinotusharoldiWaterhouse.Holotype.(♂.) [1,cir-clelabelwithred border]Type[2,squarelabelwithgreen line, handwritten]Cordova18./3[3,squarelabel,handwritten]Pinotus haroldi(Type)Waterh[4,quadrangularandredlabelwithblack borders]HOLOTYPE1♂.[BMNH].PinotusverticalisFelsche. Holo-type.(♂.)[1,quadrangularlabel,secondwordhandwritten]Bahia Blanca[2,quadrangularorangelabel]Typus.[3,quadrangulargreen label]Coll.C.FelscheKauf20,1918[4,squarelabelwithdoble pur-pleline,handwritten]verticalisFelsche[5,quadrangularredlabel withblackborders]HOLOTYPE1♂.[SMTD].
Additional specimens.(27♂♂. 25♀♀.):ARGENTINA: CerroCorta Pie,II/1942, 2♀. [MZSP].Buenos Aires:II/1942, 4♂. 4♀. [MZSP]; XI/1942,1♂.2♀.[MZSP];III/1950,3♂.[CEMT].PartidoPuán,Felipe Solá,I/1948,Martínez,A.2♀.[CEMT]Dto.Puán,FelipeSolá,XI/1959, Martínez, A. 1♂. [MZSP]. Felipe Solá, II/1942, 8♂. 2♀. [MZSP];
A.
E.
F.
G.
H.
I.
J.
B.
C.
D.
Fig.6.Dichotomiusharoldi.(A)Dorsalhabitusofthemale.(B)Lateralviewofaedeagus.(C)Dorsalviewofparameres.(D)Ventralviewoftheparameres.(E)Copulatrix lamellae.(F)Centralaccessorylamella.(G)Leftlateralaccessorylamella.(H)Rightlateralaccessorylamella.(I)Headoffemale.(J)Thoraxoffemale.Linescale1mm.
Martínez,A., II/1942, 2♂. 5♀. [MZSP]; Martínez,A. X/1942, 1♂. 1♀.[MZSP];X/1942,Pohl,B.1♀.[MZSP].XII/1961,Martínez,A.1♀. [MZSP].[BuenosAires]:Saavedra,GoyenaI,1960,Martínez,A.1♂. 2♀.[MZSP].Córdoba:1♀.[MZSP].SanFrancisco,X/1961.1♀.[CEMT].
SantiagodelEstero:Pinto,I/2006,Abadie,E.1♂.[CEMT].RíoSalado WagnerCol.1♂.[MZSP]decembreCollectionWagner1♂.[MZSP]. OjodeAgua,Coll.Martínez,Feb[rero].[1]9741♂.1♀.[CMNC].Ojo deAgua,17.II.1982,H.&A.Howden.1♂.[CMNC].
Males:Length 18–28mm; width 11–14.5mm. Blackcolor with brownsetaeatbothsidesofheadandpronotum(Fig.6A).
Head:Aslongasitiswidewithprolongedanteriormargingiving theappearanceofablunttriangle.Clypeuswithabrilliantflange, withoutclypealteeth.Surfaceofclypeuswithstrongandparallel wrinkles.Genaewithcurvedandpronouncedanterioredge, lat-eraledgestraightandthencurved,surfacewithstrongwrinkles. Clypeus-genalsutureevident,reachingthecephalicprocess.Frons withthreehorns,thecentralislong,sub-quadrangularanddirected backwards,dorsoventrallyflattenedatthebasalarea,surfacewith softerwrinklesthantheclypeusandcompressedontheapicalzone atwhichitendswithacylindrical,bluntandbrillianttipthat con-tinuestotheanteriorregion.Thetwolateralhornsaretriangular, small(lessthan1mm),directedupwardsanddiagonaltothebase ofthecentralhorn.Insmallmalesthethreecephalicprocessesare lessdeveloped.
Thorax:Pronotumtwiceaswideasitislong.Surfacewith elon-gatedpuncturesseparatedbyoncetheirlengthordiameteracross thewholesurface.Anteriorregionwithawideandsmoothedge; thereisanacuteanglebehindthecephalichorn,thenthisedge gets compressedtoward theanteriorangles. Theanteriorangle iscurved.Inthesideview,theanteriorregionofthepronotum is verticaltothe dorsal region,excavatedin thecentralregion andcurvedtowardthelateralzones,givingtheappearancethat the central horn of the head fits into this excavation. In the anterior–dorsalviewthethorax hasa deepexcavation,formed
aroundtothetwolonghumpsthatreachtheanteriorangleofthe pronotum,inthecentralzonethereisasmallcontinuous invagi-nationwithanelongatedfoveatothebackregionofthepronotum. Bothlateralfoveaearebig,deepandelongated.Hypomerewith shagreened surface and ocelated punctures rounded on ante-rioredgesandelongatedontheposterioredgesonwhichbeing separatedbyone timetheirdiameter.Allpunctureswitherect, long brown setae,which can be seen on dorsal view and less denseatthecentralregion.Thecentralregionunderthefemur surfaceis brilliantwiththesamepuncturepattern. Shagreened prosternum,prolongedfromitsmiddleregionuntilcoveringthe antero-medialzoneofthemesosternum.Fromtheposterioredge oftheprosternummanyyellowsetaeemergedirectedtowardthe mesosternum.
Elytrawithsoftandbicarinatestriae,withmediumand ocel-lated punctures spaced by about three times their diameter. Interstriaewithshagreenedsurfacewithsmallandbright punc-turesseparatedbyfivetimestheirdiameter.Mesosternumstrongly narrowedmedially,onwhichthereisabrilliantandsmooth pro-cessthatavoidsthemeso-metasternalsuture,thereisashagreened surfacetoward thesideswithsoftand ocellatedpuncturesand denseyellowsetae.Mesepisternumwithbigandocellated punc-tures,separatedbyaboutonetimeitsdiameter,withasetaoutof eachone.Metasternumwithshagreened,shinycentralarea with-outsetae;strongcarinafrom1/3ofmetasternumtothebackending inadeepfovea.Sideedgesoftheanteriorareashagreenedwith densesetigerouspunctures,posteriorareashagreenedwithlarge setigerouspunctures.Metepisternumequaltomesepisternum.
Abdomen:Sternitesshagreenedwithsmallpuncturesseparated byseventimestheirdiameter,denserinthe5thand6thsternites. Lateralmarginsofsternites1–5withdenseocelatedpunctures.6th sterniteverynarrowedmedially.Pygidiumshagreenedwith elon-gatedpuncturesseparatedbyonetimetheirdiameter,incomplete margininthemiddleinferiorregion.
A.
E.
F.
G.
H.
I.
J.
B.
C.
D.
Fig.7.Dichotomiusnutans.(A)Dorsalhabitusofthemale.(B)Lateralviewofaedeagus.(C)Dorsalviewofparameres.(D)Ventralviewoftheparameres.(E)Copulatrix lamellae.(F)Centralaccessorylamella.(G)Leftlateralaccessorylamella.(H)Rightlateralaccessorylamella.(I)Headoffemale.(J)Thoraxoffemale.Linescale1mm.
Malegenitalorgan:Sideviewofaedeaguswithsub-quadrangular phallobase, basal region with a medium bulge on both sides, apex witha constriction of approximately115◦. Sub-triangular paramereswithathinattheinferiorbase,whichcontinueswith aninvaginationtowardthemiddlezone;thinandbluntapex,with aflatsurfacefromwhichemergeshortsetae(Fig.6B).Dorsally sym-metricwithenlargedbaseandbecomingthinnertowardtheapical zoneinwhichendingwithablunttip;setaecanbeseenin lat-eralview.Internalregionwithaconvexandsoftinvaginationin thefirstthirdoftheapicalareaofparameres(Fig.6C).Inventral view,parameresarethinatbasalzonewithanacute prolonga-tiondescribedinthelateralview:bluntandflatapexwheresetae canbeseen.Betweenparameres,andinsertedfromtheapicalto basalzone,there aretwo lamellatedand overlappedprocesses: theleftparamereprocessovertherightone.Theseprocessesgo fromtheapicalzonefortwo-thirdsofparameres(Fig.6D). Inter-nalsacistubularwithraspulesonthebasaltwo-thirds;thereis abig,sub-quadrangularandbrowncopulatrixlamellae,withtwo darkerlateralprolongations,theentiresurfaceofthelamellae cov-eredbybristleswhichbecomelongertowardthelateralmargins (Fig.6E).Apicalzonewiththreeaccessorylamellae;thecentral oneislongwithundefinedform,moresclerotizedinthecentral areasurroundedbyasemi-quitinizedmembrane(Fig.6F).Left lat-erallamellaehas“C”form,withenlargedandsclerotizedsuperior marginwithdefinededges(Fig.6G).Rightlaterallamellahas“N” form,enlarged,sclerotizedinthemiddlewithsurrounding semi-quitinized membrane that makesit look wider than theapical regionofthesclerite(Fig.6H).
Female: length 19–20mm; width 12–13.1mm. It differs from malesduetohavingasofteranterioredgeofthegenaandfrons withtwolateralhorns(Fig.6I).Theexcavationofthepronotumis softer,sothatthebackcentralfoveaisstrongerandlonger(Fig.6J). 6thabdominalscleritenotshortenedmedially.
Commentaries:Thespeciesisdistinguishedfromtheothersofthe group“buqueti”becausethemalespresentaprocessofthreehorns inthefrontwherethetwolateralhornshavelessthan1mmandthe femalesonlyhavethelateralhornsinthefront.Thoraxwithadeep excavationin thecentralarea,which continuesasanelongated foveaintheposteriorareaofthepronotum.Blackelytrawithsoft andbicarenatestriae.Thisspeciesispreferentiallyassociatedto areasadjacenttotheChacodryforestsandgrasslandsofthePampa.
Distribution: This species is distributed in Argentina in the Buenos Aires, Córdoba and Santiago de Estero provinces (Fig.8).
Dichotomiusnutans(Harold,1867) (Figs.7A–J,8C)
PinotusnutansHarold,1867:page97(originaldescription).
Typematerial:PinotusnutansHaroldHolotype(♂.):[1,squarelabel withredborder,handwritten]UruguayP.nutansHarold.[2,square labelwithblackline] Ex-MusaeoE. Harold[3,quadrangularand greenlabel] MuseumParisex Coll.R.Oberthur[4,quadrangular label,handwritten]RioParana[5,quadrangularlabel,handwritten] UruguayanusoReich.1♂.[MNHN].
Additionalspecimens.(21♂♂.15♀♀.):ARGENTINA:BuenosAires:
Tandil,20/I/1917,Bruch, C.1♀. [MZSP].Tandil,André-legit.Coll. Martínez, Ene[ro], [1]954, 1♂. [CMNC]. Tandil, Bolle-legit, Coll. Martínez,23.I.[19]54.1♀.[CMNC].Cordoba:Colón,NofCordoba WofAscochingaSierradeCordobaN.7.1.20101♀.[CEMT].BRAZIL:
RioGrandedoSul:U.S.Anna,IX/1927,P.Buck,1♂.1♀.[MZSP]. PortoAlegre,12/VIII/1931,1♂.;13/X/1927,1♂.1♀.;15/IX/1937,1♀.; 21/VI/1928,1♂.;27/VI/1935,1♂.;3/IX/1941,1♂.;3/V/1938,1♀.; 3/VIII/1931,1♂.;7/IX/1927,4♂.5♀.;7/VI/1928,4♂.4♀.;23/X/1935, 1♂.[MAPA].VII/1928,1♂.[MZSP].VII/1928,Buck,P.1♂.1♀.[MZSP].
URUGUAY:Rocha,CerrodeLechihuana,Castillos25agosto1995. Leg:J.R.Verdú(00023405,000234409)2♂(00023407)1♀.[CEMT].
Males:Length18.5–26.1mm;width11–16.3mm.Blackcolorwith brownsetaeonbothsidesofheadandpronotum(Fig.7A).
Head:Aslongasitiswidewithprolongedanteriormargingiving theappearanceofablunttriangle.Clypeuswithabrilliantflange, withoutclypealteeth.Surfaceofclypeuswithstrongandparallel wrinkles.Genaewithcurvedandpronouncedanterioredge, lat-eraledgestraightandthencurved,surfacewithstrongwrinkles. Clypeus-genalsutureevident,reachingthecephalicprocess.Frons withalong,sub-quadrangular centralcephalic processdirected backwards,dorsoventrallyflattenedatbasalarea,andcompressed ontheapicalzonewhereendingwithacylindricalbluntand bril-lianttip;hornsurfacewithsofterwrinklesthanthoseofclypeus frombasaltomid-region.Insmallermalesthehornisshortand withoutevidentfinalprocess.Posteriorregionoftheheadwith elongatedholesseparatedbyonetimetheirlength.
Thorax:Pronotumtwiceaswideasitislong.Surfacewith circu-larpunctureselongatedtowardthelateraledgesandareseparated byonetimetheirlength.Anteriorregionwithawideandsmooth edge;thereisaninvaginationtowardtheanteriorregionbehindthe cephalichorn,thisedgethengetscompressedtowardtheanterior angles.Theanteriorangleiscurved.Inthesideview,theanterior regionofthepronotumisverticaltothedorsalregion,straight inthecentralregionandcurvedtowardthelateralzones.There aretwoshortandcurvedtuberclesonthecentraldorsalregion ofpronotum;fromthesetuberclestwoprotuberanceslikehumps continuetowardthelateralzones,givinganappearanceofan exca-vationintheanteriorareaofthepronotum.Behindandbetween bothtubercles,thereisanelongatedanddeepfoveathatdoesnot reachtheposteriorregionofthepronotum.Bothlateralfoveae arebig, deepand rounded. Hypomere withshagreenedsurface anddensesetigerouspuncturesonthelateraledgeswitherect, longbrownsetaewhichcanbeseenfromthedorsalviewandless denseinthecentralregionwereseparatedbytwicetheir diam-eter.Inthecentralregionthesurfaceunderthefemurissmooth andbrilliantwiththesamepuncturepattern.Shagreened proster-nummid-regionwithcorrugatedsurfaceandprolongedfromits mediumregionuntilcoveringtheantero-medialzoneof mesoster-num. From theposterior edge of theprosternum many yellow setaeemergedirectedtowardthemesosternum.Elytrawithstrong striae,thefirstfourofwhichformedbyaslopewithashagreened surface,narrowintheanteriorzoneandbecomingbroadertoward themiddlezonebeforebeingcompressedagaininthefinalareaof theelytra;thefifthtoseventhstriaebicarinatewithuniformwidth through theircomplete length. Interstriaeconvexand elevated withshagreenedsurfaceandsmallpuncturesseparatedbythree timestheirdiameter;someofthem havingtransversewrinkles givingacorrugatedappearance.Mesosternumstronglynarrowed medially,onwhichthere isa brilliantandsmooth processthat preventsthemeso-metasternalsuturefrombeingseen.Thereis ashagreenedsurfacetowardthesideswithdensesetigerous punc-turesandyellowsetae.Mesepisternumhasthesamesurfaceasthe lateraledgesofmesosternum.Meso-metasternalsutureiserasedin centralareabyamesosternumprocess,lateralmarginsevidentand brilliant.Metasternumwithshagreened,shinycentralareawithout setae;strongcarinafrom1/3ofmetasternumtothebackending inadeepfovea.Sideedgesoftheanteriorareashagreenedwith densesetigerouspunctures,posteriorareashagreenedwithlarge setigerouspunctures.Metepisternumequaltomesepisternum.
Abdomen:Sternitesshagreenedwithsomepuncturesinthe mid-dleregioninthe5thand 6thsternites.Allsterniteswithdense ocellatedpuncturesonlateraledges.6thsterniteverynarrowed medially.Pygidiumshagreenedwithelongatedpuncturesand sep-aratedbyonetimetheirlength,incompletemargininthemiddle inferiorregion.
BOLIVIA BRAZIL PARAGUAY URUGUAY ATLANTIC OCEAN Dichotomius buqueti Dichotomius haroldi Dichotomius nutans 0 400 Kilometers ARGENTINA
Fig.8.Distributionmapsofthe“buqueti”speciesgroup.
Malegenitalorgan:Sideviewofaedeaguswithsub-quadrangular phallobase, basal zone with a medium bulge on both sides, apexwitha constrictionofapproximately115◦.Parameres sub-triangularwithathinattheinferiorbase,whichcontinueswithan invaginationtowardthemiddlezone;thinandbluntapexwithflat surfacesetae(Fig.7B);dorsallysymmetricwithenlargedbaseand becomingthinnertowardapicalzonewhereendingwithablunt tip;fromthisview,setaecanbeseentoo.Internalregionwitha convexandsoftinvaginationtowardthemiddleofeachparamere (Fig.7C).Intheventralviewparameresarethinatthebasalzone withanacuteprolongationdescribed in thelateralview: blunt andflatapexwheresetaecanbeseen.Betweenparameresand insertedfromtheapicaltobasalzonebymembrane,therearetwo lamellatedandoverlappedprocesses,thatoftheleftparamereover thatoftherightparamereina characteristicwayrunningfrom theapicalzonetobeyondtwo-thirdsoftheparameres(Fig.7D). Internalsactubularwithraspulesonthebasaltwo-thirds;inthe centeronebig,sub-quadrangular,browncolorcopulatrixlamellae, with two tubular and darker lateral prolongations, entire sur-facecoveredbybristleswhichbecomelongertowardthelateral margins(Fig.7E).Apicalzonewiththreeaccessorylamellae,one longandcentralwithundefinedform,in thecentralareamore sclerotizedsurroundedbysemi-quitinizedmembrane(Fig.7F).Left laterallamellahas“C”form,withanenlargedandlesssclerotized superiormargin,irregularborders,inferiorzonethinnerand scle-rotizedwithdefinedborders(Fig.7G).Rightlaterallamellahas“N” form,enlarged,sclerotized,rightmarginwiderthantheleft,with asemi-quitinizedmembrane(Fig.7H).
Female:Length21.5–28mm; width12–16.5mm.It differsfrom malesduetoexhibitingafour-pointedcephalicprocessonfron; bothcentraltipshigher,lateraltipsconicalandsmaller(Fig.7I). Centraltuberclesofthepronotumaresofter(Fig.7J).6thabdominal scleritenotshortenedmedially.
Commentaries:Thespeciesisdistinguishedfromtheothersofthe group“buqueti”becausethemaleshaveanonlycentralhornon thefronandfemaleswithfour-pointedcephalicprocess.Pronotum withtwoshortandcurvedtuberclesonthecentralregionfrom thesetuberclestwoprotuberanceslikehumpscontinue toward thelateraledges,giving anappearanceof anexcavation inthe anteriorareaofthepronotum.Behindandbetweenbothtubercles, there is an elongated and deep fovea that doesnot reachthe posteriorregionofthepronotum.Blackelytrawithstrongstriae wherethefinalpartofthefirsttothefourtharewidened.This
species isassociated tolow mountains ofthe Pampabiome,in extremelyintervenedareas,wherenaturalvegetationisscarce.
Distribution:ThisspeciesisdistributedinArgentinaintheBuenos Aires province, in Brazilin theRio Grande do Sul stateand in UruguayintheRochadepartment(Fig.8).
Dichotomiusribeiroi(Pereira,1954)*
Diagnosis: Triangularhead. Maleswithcephalic process at the front,flattenedanddirectedbackwards.Femaleswithcephalic pro-cesswithbroadbase,endingintwoprocesses.Malesandfemales withadeepandsmoothexcavationinthecentralzoneofthe prono-tum.Head,thoraxandelytrawithcallusesthatgiveacorrugated appearance(foradetaileddescriptionseeArias-Buriticáand Vaz-de-Mello,2013).
Commentaries:Becauseofthemorphologyoftheheadand prono-tum, this species was included in the“buqueti” species-group, however, due tocharacteristicsof the morphologyof themale genitaliaandcallusesonthehead,pronotumandelytrathiswas transferred to the “cotopaxi” species-group (Arias-Buriticá and Vaz-de-Mello,2013).
Conflictsofinterest
Theauthorsdeclarenoconflictsofinterest.
Acknowledgments
Thankstothecuratorsoftheentomologicalcollectionsvisited fortheirsupportandcollaboration.ToOrlandoRangel-Ch.and the“BiodiversidadyConservación”researchgroup(Universidad Nacional deColombia).JAABexpressesmany thankstoMateus Souza,RicardoVicenteandMiquéiasFerrão(UFMT)fortheir col-laboration and hospitality and to Jhon Neitafor his advice on preparingillustrationsandTimothyBakerfortheEnglishrevision. FZVM issupportedbyCNPq(Conselho Nacionalde Desenvolvi-mento Cientifico eTecnologico 304925/2010-1, 302997/2013-0, 405697/2013-9,484035/2013-4,202327/2013-2,248299/2012-3, 306745/2016-0).PartofthisworkwasgrantedbytheSynthesys Project (http://synthesys3.myspecies.info/), which is financed by the European Community Research Infrastructure Action
under the FP7 (GB-TAF-3855) and L’Institut de Systématique, Évolution, Biodiversité (ISYEB, Muséum National d’Histoire Naturelle,CenterNationaldelarecherchescientifique,France.UMR 7205CNRS/MNHN).
References
Arias-Buriticá,J.,Vaz-de-Mello,F.,2012.RedescripciondeDichotomiuscamposeabrai yD.nemoricola(Coleoptera:Scarabaeidae:Scarabaeinae),conapuntessobresu posicionsistemática.Rev.Mex.Biodivers.83,387–395.
Arias-Buriticá, J., Vaz-de-Mello, F., 2013. Dichotomius ribeiroi (Pereira, 1954) (Coleoptera:Scarabaeidae:Scarabaeinae):redescripción yanotaciones tax-onómicasdelaespecie.Caldasia35(1),209–217.
Arrow,G.,1913.SomenewspeciesofLamellicornBeetlesfromBrazil.Ann.Mag. Nat.Hist.8(11),456–466.
Felsche,C.,1901.BeschreibungencoprophagerScarabaeiden.Dtsche.Entomol.Z., 135–155.
Felsche,C.,1911.CoprophageScarabaeiden(Col.).Dtsche.Entomol.Z.,133–141. Génier,F.,2000.Dichotomiuscomarapensissp.nov.,unenouvelleespècebolivienne
descarabéebrachyptère(Coleoptera:Scarabaeidae:Scarabaeinae).Fabreries25 (2–3),25–31.
Harold,E.,1867.NeuerCoprophagen.ColeopterologischeHefte,94–100. Harold,E.,1869.In:Gemminger,M.,Harold,E.von.(Eds.),Cataloguscoleopterorum.
Hucusquedescriptorum,synonymicusetsystematicus.Tom.IV.Scarabaeidae. SumptuE.H.Gummi,Munich,Germany,pp.979–1346.
Hijmans,R.,Guarino,L.,Bussink,C.,Mathur,P.,2012.DIVAGISVersion7.5:Sistema deInformaciónGeográficaparaelAnálisisdeDatosdeDistribucióndeEspecies. http://www.diva-gis.org.
Kohlmann,B.,2003.TribuCoprini.In:Morón,M.A.(Ed.),Atlasdelos escaraba-josdeMéxico,ColeopteraLamellicornia,v.II,FamiliasScarabaeidae,Trogidae, PassalidaeyLucanidae.Arganiaeditorial,Barcelona,Espa ˜na,pp.45–57. López-Guerrero,I.,2005.LosDichotomius(Coleoptera:Scarabaeidae:Dichotomiini)
delafaunadeMéxico.Bol.SEA36,195–209.
Lucas,M.,1857.Animauxnouveauxouraresreceuillispendantl’expeditiondans lespartiescentralesdel’AmeriqueduSuddeRiodeJaneiroaLima,etdeLima àPara;exécutéeparordreduGovernmentfrancaispendantlesannees1843à 1847,sousladirectionduComiteFranc¸oisdeCastelnau.(Entomologie). Luederwaldt,H.,1925.NeuePinotusarten.Bol.Mus.Nal.2(1),67–69.
Luederwaldt,H.,1929.AsespéciesbrasileirasdogêneroPinotus(Coleoptera: Lamel-licornidae:Coprini)comalgumasconsiderac¸õestambémsôbreoutrasespécies. Rev.Mus.Paul.16,603–775.
Maldaner, M., Nunes, R., Vaz-de-Mello, F., 2015. Taxonomic revision of the Dichotomiusspeciosus(Waterhouse,1891)speciesgroup(Coleoptera: Scarabaei-dae:Scarabaeinae).Zootaxa3986(5),549–560.
Martínez, A., 1951. Notas Coleopterológicas V: La invalidez del nombre genéricoPinotusErichsonydosnuevassinonimias.Ann.Soc.Cient.Argent., 138–142.
Martínez,A.,1974.UnanuevaespeciedeDichotomiuss.str.(Coleoptera, Scarabaei-dae,CopriniAteuchina).Physis33(86),41–45.
Medina,C.,Scholtz,C.,Gill,B.,2003.Morphologicalvariationandsystematicsof Can-thonandrelatedgeneraofnewworldCanthoninidungbeetles.Dtsche.Entomol. Z.50(1),23–68.
Nunes,R.,Carvalho,M.,Vaz-de-Mello,F.,2016.TaxonomicreviewoftheDichotomius (Luederwaldtinia)assifer(Eschscholtz)species-group(Coleoptera: Scarabaei-dae).Zootaxa.4078(1),230–244.
Pereira,F.,1953.Notassinonímicas(Coleoptera:Scarabaeidae).Dusenia4(5–6), 387–402.
Pereira,F.,1954.EspeciesnovasdeScarabaeidaeBrasileiros.Arq.Mus.Nal.42, 462–466.
Valois,M.,Vaz-de-Mello,F.,Silva,F.,2017.TaxonomicrevisionoftheDichotomius sericeus(Harold,1867)speciesgroup(Coleoptera:Scarabaeidae:Scarabaeinae). Zootaxa4277(4),503–530.
Vaz-de-Mello,F.,Louzada,J.,Gavino,M.,2001.NovaespéciedeDichotomiusHope, 1838(ColeopteraScarabaeidae)doEspíritoSanto,Brasil.Rev.Bras.Entomol.45 (2),99–102.
Vaz-de-Mello,F.,Edmonds,W.,Ocampo,F.,Schoolmeesters,P.,2011.Amultilingual keytothegeneraandsubgeneraofthesubfamilyScarabaeinaeoftheNewWorld (Coleoptera:Scarabaeidae).Zootaxa2854,1–73.
Vulcano,M.,Pereira,F.,Martínez,A.,1976.NotassobreScarabaeoideaAmericana (ColLamellicornia).Stud.Entomol.19(1–4),515–530.
Waterhouse,Ch.,1891.XXXVII.NewScarabaeidaeintheBritishMuseum:aFourth Contribution.Ann.Mag.Nat.Hist.6(8),348–363.
Zunino,M.,1978.L’armaturagenitalnegliOnthophaginiTechichedepreparazione ecriteridistudio(ColeopteraScarabaeoidea)Supplemento.Boll.Soc.Entomol. Ital.90,21–26.
Zunino, M., 1985. Las relaciones taxonómicas de los Phanaeina (Coleoptera Scarabaeinae)y sus implicacionesbiogeográficas. Folia Entomol.Mex. 64, 101–115.
Zunino,M.,Monteresino,E.,1990.Ideaspreliminaressobrelaevolucióndelos aparatoscopuladoresenRutelinae(ColeopteraScarabaeoidea)ylafilogeniadel grupo.Rev.Soc.Entomol.Argent.48(89),3–13.
