New synonymy to Simulium ( Inaequalium ) diversibranchium Lutz, with comments on the Inaequale species-group (Diptera, Simuliidae)

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RevistaBrasileiradeEntomologia60 (2016)293–296

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

New

synonymy

to

Simulium

(Inaequalium)

diversibranchium

Lutz,

with

comments

on

the

Inaequale

species-group

(Diptera,

Simuliidae)

Leonardo

H. Gil-Azevedo

∗

,

Óscar

Sánchez

Molina

UniversidadeFederaldoRiodeJaneiro,MuseuNacional,DepartamentodeEntomologia,RiodeJaneiro,RJ,Brazil

a

r

t

i

c

l

e

i

n

f

o

Articlehistory: Received14May2016 Accepted22August2016 Availableonline4September2016 AssociateEditor:MarciaSoutoCouri Keywords: Argentina Blackﬂy Brazil NeotropicalRegion Taxonomy

a

b

s

t

r

a

c

t

Simulium(Inaequalium)CoscarónandWygodzinsky,1984isahomogeneoussubgenuswidelyspread ontheNeotropicalRegion,whichtheadultscanonlybereliablyidentiﬁedfromlinked-reared spec-imens.DespitemanyrevisionaryworksonlyasinglecharacterofthepupalgillseparatesSimulium diversibranchiumLutz,1910fromS.mariavulcanoaeCoscarónandWygodzinsky,1984.Specimensofboth speciesshowthatS.mariavulcanoaeisajuniorsynonymofS.diversibranchium.Therelationshipbetween S.diversibranchiumandS.subnigrumLutz,1910needsfurtherinvestigation.Somefeaturesofthepupal gilloftenusedasdiagnosisinSimulium(Inaequalium)species,suchasthedirectionofgillﬁlamentsorthe heightofthebifurcationofthesecondaryortertiarybranchesofthepupalgillarehighlypolymorphic, makingitsuseasdiagnosticcharacternotreliable.

Introduction

Simulium(Inaequalium)CoscarónandWygodzinsky,1984isa subgenuswidelydistributedacrosstheNeotropicalRegion.Shelley

etal.(2010)proposedthesynonymyofSimulium(Psaroniocompsa)

Enderlein,1934andS.(Inaequalium)basedonasubjectivecriterion, whichwasthelackof“subgenericvalue”inthecharactersused inthediagnosisofS.(Inaequalium).IndeedS.(Inaequalium)andS. (Psaroniocompsa)aresistergroups,theysharesimilarities,asthe adultfemalecibariumshape;larvalantennashape;andpresence of1+1ventraltuberclesonthelarvalabdomen(Gil-Azevedoetal., 2012).Butboth subgeneraareprobablymonophyleticandhave easilyrecognizablediagnosticcharacters,asadultwingwithsetae onbasalsectorofradialvein;pupaltrichomeswith3–6branches; andlarvalpostgenalcleftsubtriangularinS.(Inaequalium).Onthe otherhand,S.(Psaroniocompsa)haswingwithoutsetaeonbasal sectorofradialvein;pupaltrichomesonlywithtwobranches;and larvalpostgenalcleftoval(Gil-Azevedoetal.,2012).We donot acceptthissynonymyofthesubgenerabasedoncurrentlyavailable data.

TheS.(Inaequalium)speciesaremorphologicallyhomogeneous andaredividedintwospecies-groups,Botulibranchiumand Inae-quale (Coscarón and Wygodzinsky, 1984), which are broadly accepted(Coscarónetal.,2008;Shelleyetal.,2010;andAdlerand

∗ Correspondingauthor.

E-mail:[email protected](L.H.Gil-Azevedo).

Crosskey,2016).However,thisclassicdivisionwaschallengedin

the2000swiththedescriptionoffourspecieswhich do notﬁt easilyintoanyofbothspecies-groups:S.maranguapense(Pessoa etal.,2005);S.margaritatumPepinellietal.,2006; S.lundi(

Py-Daniel and Barbosa, 2007); and S. marins Pepinelli et al., 2009

(see the original descriptions for more details). Py-Daniel and

Barbosa(2007)proposedanotherspecies-group,Maranguapense,

toaccommodatetheﬁrstthreespecies(fromnowonreferredto asthe‘2000sspecies’),whichwascriticizedbyAdlerandCrosskey

(2011).Anyway,webelievethatthisissueshouldbebetterand

deeperinvestigatedinthefuture.

Adults of the Inaequale species-group (sensu Adler and

Crosskey,2016)arealmostidentical.However,thepupalgillis

aquitevariablestructure,andallthespeciesofthegrouphave diagnosesbasedprimarilyonit.Theadultscanonlybereliably identiﬁedfromlinked-rearedspecimens(Coscarónand

Coscarón-Arias,2007;Shelleyetal.,2010),withprominentexceptionssuch

asthe‘2000sspecies’(Pessoaetal.,2005;Pepinellietal.,2006;

Py-DanielandBarbosa,2007).Thisproblematiccombinationmakes

mostspeciesoftheInaequalespecies-groupidentiﬁedfromonly afewcharactersofgillﬁlaments.Thishasresultedin21named species,ofwhichonlytenarecurrentlyvalid(AdlerandCrosskey,

2016;Coscarónetal.,2008).

Ofthose tenvalidspecies oftheInaequaliumspecies-group, the‘2000s species’ arethe exception,as theyareeasily distin-guishable,andhaverestricteddistribution,toveryisolatedhigh grassland areasofNortheasternandNorthernRegionsofBrazil. Theothersevenspecieshaveawidergeographicaldistributionand

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294 L.H.Gil-Azevedo,Ó.S.Molina/RevistaBrasileiradeEntomologia60(2016)293–296

arefrequentlyfoundinAtlanticForeststreams,andformamuch more morphologicallyhomogenous group (Adler and Crosskey, 2016).Fromthosespecies,onlySimuliumtravassosiVulcanoand d’Andretta, 1947 lackssix gill filaments in their pupa, instead theybearthreethickfilaments.Inthisgroup,theimmaturesof mostspeciespreferforestedloworderstreams,whereasS. inae-quale(Paterson andShannon,1927)preferstreamsinopenand deforestedareas,andS.rappaePy-Daniel&Coscarón,1982seems tobegeneralist.Theselasttwospeciesalsohaveamore distin-guishablepupa:S.inaequale–allgillbranchesbifurcatenearthe baseandcocoonbearsmallanterodorsalprojection;S.rappae– despiteitsknowngillvariation,theyalwayshavethickgill fila-ments(Hernández-Trianaetal.,2007b).

Therefore,fourspeciesofInaequalespecies-groupwith indis-tinctadultsandsimilarpupae(withsixpairedgillﬁlaments)occur instreamsrunninginsideforestedareas:S.clavibranchiumLutz, 1910,S.subnigrumLutz,1910,S.diversibranchiumLutz,1910,and S.mariavulcanoaeCoscarónandWygodzinsky,1984.The imma-turesofthetwoﬁrstspeciesarecommonandabundant,andcan befoundonmostofthecreeksoftheAtlanticForest,whereasthe othertwospeciesarequiterare.

Coscarón and Wygodzinsky (1984) considered S.

diversibranchiumand S.mariavulcanoae closespecies thatcould bedifferentiatedfromeachother,andfromtheremainingspecies ofthesubgenusbytheconﬁgurationofthegillbranches.Those authorsalsopointedoutmilddifferencesbetweenthemaleand femaleterminaliaofbothspecies,andalsoamongtheremaining Inaequaliumspecies-groupspecies.But,mostoftheworks pos-teriorCoscarónandWygoddzinsky(1984)consideredtheadults ofInaequalium species-groupmorphologicallyindistinguishable (e.g.CoscarónandCoscarón-Arias,2007;Hernández-Trianaetal.,

2007b).

InthisworkweproposethatS.mariavulcanoaeisajunior syn-onymofS.diversibranchium.Wealsohighlightthattherelationship betweenotherspeciesofInaequaliumspecies-groupneedsfurther investigationbecausesomefeaturesofthepupalgill,oftenusedas diagnosisofitsspecies,arenotreliable.

Materialandmethods

Inordertoobtainimages,weusedfocusstacking,using Com-bineZP(Hadley,2013),withphotographstakenusingamicroscope. WefollowedthemorphologicalnomenclatureofGil-Azevedoetal.

(2005).Thecollectionsmentionedinthispaperandtheirrespective

acronymsare:MuseodeLaPlata,LaPlata,Argentina(MLP);Museu deZoologiadaUniversidadedeSãoPaulo,SãoPaulo,Brazil(MZSP); InstitutoOswaldoCruz, Fundac¸ãoOswaldoCruz, Riode Janeiro (CSIOC);MuseuNacional,UniversidadeFederaldoRiodeJaneiro, RiodeJaneiro(MNRJ);InstitutoNacionaldePesquisasdaAmazônia, Manaus(INPA).Geographicaldistributioninformationwasbased onthelabelsoftheanalyzedmaterialandrecordsobtainedfrom theliterature(Coscarón,1991;Coscarónetal.,2008;Gil-Azevedo

andMaia-Herzog,2010;Strieder,2004).

Examinedmaterial

SimuliumdiversibranchiumLutz,1910:LECTOTYPE(designated

byHernández-Trianaetal.,2007b):BRAZIL,SP, SãoJosédo

Bar-reiro,SerradaBocaina,FazendodoBonito:1pupa(slide)(CSIOC #12,097Lutzcollection).PARALECTOTYPES:samedataof Lecto-type:4pupae(CSIOC#12,095and12,098Lutzcollection).

ARGENTINA, Misiones, Campi ˜na: 3 pupae (slide) (MLP 21/VIII/1972, S. Coscarón col.). BRAZIL, SP, Boracéia: 1 pupa (spirit)(MLP 31/X/1979, S.Coscarón col.).Campos do Jordão: 4 pupae (spirit) (CSIOC #3,774; and 3,780). Casagrande: 1 pupa

(spirit)(MLP25/VII/1979,S.Coscaróncol.).SP?:2pupae(spirit) (CSIOC#3,737).

Simulium mariavulcanoae Coscarón and Wygodzinsky, 1984: HOLOTYPE:BRAZIL,SP,Salesópolis,Boracéia:1female,withpupal exuvia(slide)(MZSP#115-212).PARATYPES:BRAZIL,SP, Salesópo-lis,Boracéia:1 male,withpupalexuvia(slide) (MZSP#115-47, “allotype”);3pupae(slides)(MLP#115-263and#115-232).

BRAZIL,MG,BocainadeMinas:1pupa(spirit)(MNRJ#60,020). SC,SãoBento:1male,withpupalexuvia(pinned)(CSIOC#4,581). SP, Campos do Jordão: 2 females, withpupal exuviae (pinned) (CSIOC#4,446;and4,448).SerradaBocaina:3adults,withpupal exuviae(pinned)(CSIOC#3,986;4,404;and4,469).SP?:1female, withpupalexuvia(pinned)(CSIOC#4,132).

Simulium subnigrum Lutz, 1910: ARGENTINA, Misiones, Pto. Leoni:1pupa(slide)(MLP07/VII/2002,S.Coscaróncol.).BRAZIL, MG, BocainadeMinas: 3females, withpupalexuviae+1 male, with pupal exuvia+3 pupae (spirit) (MNRJ #60,002; #60,006; #60.007; #60,010; #60,012; and #60,024). Pedralva: 5 pupae (CSIOC#438;440;and442).RJ,Teresópolis:1female,withpupal exuvia+8pupae(spirit)(MNRJ#60,027;60,029;and60,032).RS, ValedosSinos,ArroioCascatinha:1pupa(spirit)(MLP22/X/2000, S.Coscaróncol.).Brazil?:1malewithpupalexuvia(pinned)(CSIOC #5,702);1femalewithpupalexuvia(pinned)(CSIOC#5,727).

Taxonomy

Simulium(Inaequalium)diversibranchiumLutz,1910(Figs.1–5; 10–12)

SimuliummissionumCoscarón,1976

SimuliummariavulcanoaeCoscarón&Wygodzinsky,1984syn.n.

Taxonomic remarks. Despite many revisionary works about Simulium(Inaequalium)(Coscarón,1991;Coscarónand

Coscarón-Arias,2007;CoscarónandWygodzinsky,1984;Hernández-Triana

etal.,2007b; Shelleyetal.,2010; Striederand Py-Daniel,2000,

2002), still only one character of pupal gill shape separates S. diversibranchiumfromS.mariavulcanoae:“secondarybranchesof ventralprimarybranchdownwardsandposteriorlydirected”;in contrastwiththeotherspeciesinwhichthosebranchesare ante-riorlydirected.Butthoseauthorsagreethatbothspeciesareclose, andcouldbegroupedbysharingasinglecharacterofthepupal gill shape: “dorsal primary branch thicker than other primary branches”.Ontheotherhand,S.mariavulcanoaelacksunique exclu-sivecharacters,makingthespeciesmeaningless.

ThissinglecharactercurrentlyseparatingS.diversibranchium andS.mariavulcanoaeisnotconsistentandcouldbecausedby envi-ronmentalinfluencesinthepupaldevelopment.Thereis,however, notmanyspecimensforexamination,thespeciesbeingrare.But wefoundaninterestingpupafromPedralva (MG),which could beidentifiedeitherasS.diversibranchium,orasS.mariavulcanoae dependingiftheleft(Fig.3),ortheright(Fig.4)sideisexamined. Thisisevidencethatthedirectionofgillfilamentsisnotareliable character.

Lutz(1910)describedS.diversibranchiumbasedonpupae

syn-types.In a reviewof Simulium (Inaequalium),Hernández-Triana

etal.(2007b) choseoneofthepupaesyntypes andraiseditto

lectotype status of thespecies. We examined this, and thegill doesnothavethesecondarybranchesofventralprimarybranch downwardandposteriorlydirected,butallbranchesare anteri-orlydirected(Fig.1).ThelectotypeofS.diversibranchiumdoesnot matchthedescriptionofthis speciesintheliterature,but with thedescriptionandthetypesofS.mariavulcanoae(Coscarónand

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Fig.1.PupalgillbranchesofSimulium(Inaequalium)diversibranchium(=S.mariavulcanoae),inslides(1–5;10–12):(1)CSIOC#12,097Lutzcollection(Lectotype);(2)MLP #115,232(paratypeofS.mariavulcanoae);(3)CSIOC#3,109leftgill;(4)CSIOC#3,109rightgill;(5)MNRJ#60,020;(10)CSIOC#3,780;(11)CSIOC#3,986;(12)CSIOC#4,448. PupalgillbranchesofSimulium(Inaequalium)subnigrum,inslides(6–9):(6)MNRJ#60,002;(7)MNRJ#60,007;(8)MNRJ#60,010;(9)MNRJ#60,012.

entity,andweproposethatS.mariavulcanoaeisajuniorsynonym ofS.diversibranchium.

Diagnosis(basedonlyonpupa)(modiﬁedfromCoscarónand

Coscarón-Arias,2007:286).Frontoclypeusandthoraxwith

abun-dant tubercles; trichomes of frontoclypeus with 2–6branches, thoracictrichomeswith5–8branches.Gillwith6ﬁlamentsinthree primarybranches:dorsalprimarybranchdistinctlythickerthan medialandventralprimarybranch;themostventralsecondary branch(from ventral primarybranch) could bedownward and posteriorlydirected,ordirectedastheothersecondarybranches; gilllength2.8–3.5mm.Cocoonwiththinweave,withfewevident threads,reinforcededgeand slowlyprojected ventrally;cocoon lengthonbase3.2–4.0mm,ondorsum2.6–3.5mm.

Distribution.Argentina, Misiones: Campi ña. Brazil,RS:Caxias doSul;BomJesus.SC:SãoBentodoSul;Joinville.SP: Selesópo-lis,EstaçãoBiológicadeBoraceia;Itu;Capivari;BragançaPaulista; CamposdoJordão;SãoJosédoBarreiro,SerradaBocaina,Fazenda do Bonito;Ocauçu (asCasagrande). RJ: Guapimirim;Petrópolis; Itatiaia.MG:BocainadeMinas.

Discussion

Someauthorssuggestedthatthelevelofthebifurcationofthe secondaryortertiarybranchesofthepupaecouldbeagood diag-nosticcharacterforneotropicalSimuliumspecies(e.g.Coscarónand

Coscarón-Arias,2007;Hernández-Trianaetal.,2007a).However,

thischaracterisquestionable,becauseitalwaysshowsmany inter-mediarystates.EspeciallyinS.(Inaequalium)species,inwhichthat problemiswelldocumented,e.g.:S.clavibranchium“Variationin thebranchingofthedorsalprimarybranchandtheenlargement ofthepupalgillﬁlaments(...)havebeenseen”(Hernández-Triana

etal.,2007a:p.18);S.rappae:“Gillconﬁgurationveryvariablewith

ﬁlamentsbranchingatdifferentheightsevenwithinsame speci-men.”(Hernández-Trianaetal.,2007b:p.43;seealsoPepinellietal., 2009:p.735);S.subnigrum:“gillbranchingatdifferentdistances frombase(...)withrelative variabilityofdistancesin different specimens(CoscarónandCoscarón-Arias,2007:p.278;seealso

Hernández-Trianaetal.,2007b:p.47).Andmanyspecieshavebeen

synonymizedbasedontheweaknessofthatfeature,e.g.:S. subclavi-branchiumLutz,1910;S.nogueiraid’AndrettaandDoloresGonzález, 1964;S.beaupertuyiRamírez-Pérez,Rassi,Ramírez,1977;andS. leopoldense(Striederand Py-Daniel,2000), allconsideredjunior synonymofS.subnigrum(Hernández-Trianaetal.,2007a:p.18–20;

seealsoShelleyetal.,2010).Observationoftheavailable

mate-rialshowsagreatvariationintheheightofthebifurcationofthe secondaryortertiarybranchesofthepupalgill(Figs.1–12), bring-ingnewevidencethatgillﬁlamentsinS.(Inaequalium)speciesisa polymorphiccharacteranditsuseasadiagnosticcharacterforthis subgenusisdebatable.

OnlyasinglecharacterseparatesS.diversibranchiumsensuthis paperandS.subnigrum.Theformerspecieshasa“pupalgillwith dorsalprimarybranchthicker thanotherprimarybranches”;in contrastwithS.subnigruminwhichallprimarybranchesare

simi-lar(Coscarón,1991;CoscarónandCoscarón-Arias,2007;Coscarón

andWygodzinsky,1984;StriederandPy-Daniel,2000,2002).We

examinedmaterialofbothspecies,andfoundthatsomespecimens aredifﬁculttodetermineifthedorsalprimarybranchdiameteris thickerorsimilartotheothers.Onceagain,thereare intermedi-arystates,aswecanseeintheexaminedmaterial,especiallyfrom BocainadeMinas(Figs.5–9).

WhereasS.subnigrumhasawidedistributionfromnortheastern ArgentinatoTrinidad(AdlerandCrosskey,2016),andisabundant

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inforestedareasinallitsgeographicalrange,S.diversibranchium haspunctualdistributionand isquiterare.Infact,itdrawsour attentionthatthereareveryfewspecimensofS.diversibranchium incollections,eveninthethreemostrepresentativeNeotropical simuliidcollections(MLP,CSIOC,andINPA).

TheverylowfrequencyofS.diversibranchium,alwaysfoundin areaswhereS.subnigrumisabundant,alliedthatthetwospecies canonlybedifferentiatedbyonepupalgillcharacter,suggeststhat S.diversibranchiumisavariationofS.subnigrum.Maybethoseare aberrantvariationsonthegillthatappearsinsomepopulationsof S.subnigrum,causedbyenvironmentalvariablesand/ormutations. OrS.diversibranchiumisjustaveryrarespecies.Morphometricand molecularstudies,basedonlargenumberofspecimensshouldbe necessarytoanswerit.Additionally,larvaecouldberearedunder differentvelocityconditionstoseewhataffectthathadonthegill.

Conﬂictsofinterest

Theauthorsdeclarenoconﬂictsofinterest.

Acknowledgements

MarilzaMaia-Herzog(CSIOC),SixtoCoscarón(MLP),CarlosE. Lamas(MZSP),NeusaHamadaandVictorPy-Daniel(INPA)kindly permittedaccessand/orprovideinformation totheirrespective collections.AmandaMendes(Universidadedo EstadodoRio de Janeiro),MateusPepinelli(RoyalOntarioMuseum),andDouglas Craig(UniversityofAlberta)contributedmanythoughtful sugges-tionsthat improvedthis work.Thisresearchwassponsored by Fundac¸ãoCarlosChagasFilhodeAmparoàPesquisadoEstadodo Riode Janeiro(FAPERJ; E-26/110.707/2011;E-26/111.767/2012) and by a scholarship granted to OSM by the Brazilian National Council for Scientiﬁc and Technological Development (CNPq).

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