Redescription and first record in South America of Neogriphoneura timida Curran (Diptera, Lauxaniidae)

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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

Redescription

and

ﬁrst

record

in

South

America

of

Neogriphoneura

timida

Curran

(Diptera,

Lauxaniidae)

Ramon

Luciano

Mello

a,∗

_,

_Thiago

_Mahlmann

b

_,

_Vera

_Cristina

_Silva

c

a_Universidade_Federal_de_Mato_Grosso_do_Sul,_Laboratório_de_Sistemática_de_Diptera,_Campo_Grande,_MS,_Brazil b_Instituto_Nacional_de_Pesquisas_da_Amazônia,_Manaus,_AM,_Brazil

c_Universidade_Estadual_Paulista_(UNESP),_Faculdade_de_Ciências_Agrárias_e_{Veterinárias,}_Departamento_de_Morfologia_e_Fisiologia_Animal,_Jaboticabal,_SP,_Brazil

a

r

t

i

c

l

e

i

n

f

o

Articlehistory:

Received4March2016

Accepted17October2016

Availableonline2November2016

AssociateEditor:MarciaSoutoCouri

Keywords: Distribution Schizophora Spermcompetition Spermathecae Spermatozoa

a

b

s

t

r

a

c

t

ThegenusNeogriphoneuraMalloch,1924iscurrentlycomposedof11specieswithNewWorld distribu-tion.NeogriphoneuratimidaCurran,1942isrecordedfortheﬁrsttimeinSouthAmerica,withoccurrences inBrazil,ColombiaandTrinidadandTobago;andnewCentralAmericanrecordsarepresented:Belize, HondurasandMexico.Hereweredescribethespeciesandpresent,fortheﬁrsttime,illustrationsofthe maleterminaliaandfemalespermathecae,anddiscussthemaindiagnosticcharacteristicsofexternal morphology.Abriefdiscussionaboutthepostcopulatorysexualselectioninthisspeciesisproposed basedonmorphologyofthespermathecae.

Introduction

LauxaniidaeisalargefamilyofSchizophoraﬂiescomprisedof over2000speciesinabout200generaworldwide.Inthe Neotrop-icalRegioniscomposedbynearly400speciesinabout70genera

(GaimariandSilva,2010).

ThegenusNeogriphoneurawaserectedbyMalloch(inMalloch

andMcAtee,1924)toincludethespeciesSapromyzasordida

Wiede-mann, 1830, originally recorded from West Indies. The genus Rhabdolauxania was described by Hendel (1925) for his two new species R. laevifrons from Peru, and R. schnusei (the type species)fromBoliviaandPeru.Neogriphoneurastriatifronswaslater describedbyHendel(1932)basedonmaterialfromBolivia.Ina subsequentpublication,Hendel(1933)describedRhabdolauxania immaculatafromParaguay.Neogrphiphoneuratimidawasdescribed

byCurran(1942)basedonfourmalesandsixfemalesfromPanama;

inadditiondescribingtwoothernewspecies:N.strigaandN.tertia bothbasedonmaterialfromPanamaandBrazil.

After66 years withoutany additional species or taxonomic modiﬁcation, Mello and Silva (2008a) presented a taxonomic reviewofNeogriphoneura.Inthisreview,theydescribedthespecies N.bispoifromBrazil,N.corrugatafromtheBritishVirginIslands,and

∗ Correspondingauthor.

E-mail:ramon.mello@ufms.br(R.L.Mello).

N.pacatafromBolivia;thegenusRhabdolauxaniawassynonymized underNeogriphoneura,anditsthreespecieswerecombinedinto Neogriphoneura;lectotypesforR.immaculataandR.laevifronswere designated;anidentiﬁcationkeytothespecieswaspresented;and newoccurrencerecordswereregisteredforthefollowingspecies: N.sordida,N.striatifrons,N.striga,N.tertia,andN.timida.According withthisinformation,Neogriphoneuraiscomprisedofthe follow-ing11species:N.bispoi,N.corrugata,N.immaculata,N.laevifrons, N.pacata,N.schnusei,N.sordida,N.striatifrons,N.striga,N.tertia, andN.timida.

Inthisstudy,N.timidaisredescribed,withphotographsof exter-nalmorphology,andillustrationsofmaleterminalia,andfemale spermathecae;newrecordsareaddedtoitsdistribution, includ-ingtheﬁrstrecordforthespeciesinSouthAmerica,inadditionto newrecordswithinCentralAmericaandMexico.Thedifferentiated morphologyofthespermathecaeofthisspecies,sharedwithtwo othercongeners,N.bispoiandN.corrugata,isbrieﬂydiscussedin thelightofapossiblecorrelatedevolutionwithsomemaletraits.

Materialandmethods

Thematerialanalyzedin thisstudybelongs tothefollowing collections:AmericanMuseumofNaturalHistory(AMNH),New York,USA; Colec¸ãoZoológica daUniversidadeFederal de Mato GrossodoSul(ZUFMS),CampoGrande,Brazil;Colec¸ãoZoológica doMaranhão,UniversidadeEstadualdoMaranhão(CZMA),Caxias,

http://dx.doi.org/10.1016/j.rbe.2016.10.001

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Figs.1–4.NeogriphoneuratimidaCurran.(1)Habitus,lateralview(female);(2)habitus,dorsalview(female);(3)head,dorsalview(male);(4)wing(female).Scale:2mm.

Brazil;InstitutodeInvestigacióndeRecursosBiológicos Alexan-dervonHumboldt(IAvH),Bogotá,Colombia;InstitutoNacionalde PesquisasdaAmazônia(INPA),Manaus,Brazil;NationalMuseumof NaturalHistory(USNM),Washington,DC,USA;TheNaturalHistory Museum(BMNH),London,England.

ThemorphologicalterminologyfollowedCummingandWood

(2009).Thedissectionofthemaleterminaliafollowsthe

proce-durespresentedinMelloandSilva(2008b).

Taxonomy

NeogriphoneuratimidaCurran(Figs.1–9)

NeogriphoneuratimidaCurran,1942:75.Typelocality:Panama, PatillaPoint.Holotypemale,AMNH.

Diagnosis:Thisspeciescanbeseparatedfromitscongenersbythe followingcombinationofcharacters:fronsunmarked; mesono-tumwitha central darkstripeanteriorly,and anovalspoton posteriormarginextendingontoscutellum;wingunspotted. Redescription

Body(Figs.1and2):Yellowtoyellowishbrownororange;length:

4–4.5mm;wing:3.2mm.

Head(Figs.1–3):Vertexrounded;ocellartuberclebrown,closeto

vertex,welldeveloped;fronsyellow,withoutdarkmarkings, rect-angular,widerthanlong,concaveonmiddlepartofanteriorhalf; eyesuboval,higherthanlong,posteroventralmarginconcave;face yellowishtolightbrown,unmarked,lowerhalfslightlyconvexin profile;parafacialpale,whitishyellow;genayellow;occiputand medianoccipitalscleriteyellowish,unmarked;antennayellowto yellowishbrown,scapeshorterthanpedicel,firstflagellomeretwo timeslongerthan wide,aristaabout2.5timeslongerthanthe lengthof theantennalarticles,blackand plumose;palpus and labellumyellowtoyellowish brown.Chaetotaxy:outervertical

setalateroclinate,1/3lengthofinnerverticalseta;innervertical setareclinate,postocellarsetaconvergent;ocellarseta diminu-tive;anteriororbitalsetainclinate,slightlylongerthanreclinate posteriorseta.

Thorax (Fig.2): Mesonotum yellowish brown to orange, with a blackstripeanteromedially and arounded spotonposterior part,extendingontoscutellum;scutellumﬂat,otherwiseyellow; pleurayellowish.Chaetotaxy:1postpronotalseta;0+3 dorsocen-tralsetae;1prescutellaracrostichalseta;1presuturalsupra-alar seta;1postsuturalsupra-alarseta;2notopleuralsetae;postalar setaabsent;1intra-alarseta;1proepisternalseta;1anepisternal seta;1katepisternalseta;2scutellarsetaedivergent.Legs:whitish yellow,tarsiyellow.Chaetotaxy:forecoxawithatransversalrow ofanterodorsalsetae;forefemurwithalongitudinalrowof pos-terodorsalsetae,alongitudinalrowof3apicalsetaeonventral surface,and2apicalsetaeonposteriorsurface;foretibiawith1 preapicaldorsalseta;midcoxawithatransversalrowofsetaeon dorsalsurface;midfemurwithalongitudinalrowof anterodor-salsetaeand1apicalsetaonposteriorsurface;midtibiawith1 preapicaldorsalsetaand1strongtibialspur;hindfemurwith2 longitudinalsetaeonapicalmarginofdorsalsurface;hindtibia with1apicalsetaondorsalsurface.Wing(Fig.4):hyaline;veins yellow;costasapromyziform;Rbare;r-mslightlybasalto mid-pointofdiscalmedialcell;crossveindm-cuatthemidpointofcell r4+5.Halteryellowishbrowntoorange.

Abdomen(Fig.2):Yellowishbrown, somespecimenswithone ovaldarkspotmediallyoneach oftergites3–5.Male termina-lia:Epandrialcomplex(Figs.5and6):epandriumsaddle-shapein posteriorview(Fig.5),loweranduppermarginsovalandstraight respectively,inlateralview(Fig.6),coveredbylargeandmedial setae;surstylusarticulatedtoepandrium,conicalinposteriorview (Fig.5), trapezoidalonlateralview(Fig.6), lowermargin cov-eredbythinhairsinlateralview.Phalliccomplex(Figs.7and8): phallapodemetubular,welldeveloped(Fig.7);phalluscylindrical, foldedinthemiddle,withsclerotizedspinesinternallyonapical region(Figs.7and8);hypandrialarmsdeveloped,almostthree

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Figs.5–9.NeogriphoneuratimidaCurran.(5)Epandrialcomplex,posteriorview;(6)epandrialcomplex,lateralview;(7)aedeagalcomplex,dorsalview;(8)aedeagalcomplex, lateralview;(9)spermathecae.Scale:1mm.

timesthelengthofpostgonite(Fig.7);hypandriumforminga com-pletering(Fig.7);postgonitepresent(Fig.7).Femaleterminalia: Spermathecae(Fig.9):three(1+2),wrinkledandtubular. Distribution:Mexico,Guatemala,Belize,Honduras,ElSalvador, CostaRica,Panama,Trinidad andTobago, Colombia,andBrazil (Fig.10).

Type-materialexamined:Holotypemale:PANAMA,PatillaPoint, CanalZone;1/ii/1929;Coll.C.Curran(AMNH).Paratypefemale: idem(AMNH).

Additional material examined: MEXICO, Chiapas, Huixtla; 26/vii/1967; H. Sanches leg.; 1 male (USNM). GUATEMALA, Quirigua;1926;M.Aldrichleg.;1male(USNM).BELIZE,Cayo Dis-trict,Pook’sHillLodge(17◦0915N;88◦5108W);2008;Malaise

trap;leg.V.&R.Snaddon;2malesand1female(BMNH);idem, 22.vi.2008;Malaisetrap;leg.V.&R.Snaddon;2malesand1female (BMNH).HONDURAS, LaCeiba,F.Deyerleg.;1female(USNM). Tegucigalpa;F.Deyerleg.;1male(USNM).PANAMA,PanamaCity, CanalZone,MonsoonForest;15–30/vii/1979;E.Broadheadleg.;1 male(USNM).TRINIDADANDTOBAGO,TobagoIsland,SaintJohn, Charlotteville;14–21/iii/1979;D.Hardy&W.Rowelegs.;1female (USNM).COLOMBIA,Chocó,Riosucio,ParqueNacionalNaturallos Katíos,CentroAdministrativoSautatá,AfueradelBosque(7◦51N; 77◦8W); 13–29/vi/2003; P. Lopes leg.; 2 males (IAvH). Meta, ParqueNacionalNaturaldeLaMacarena,Caba ˜naCerrillo(3◦21N; 73◦56W);21/xii/2002–04/i/2003;A.Herrera&W.Villalbaleg.;1 female(IAvH).BRAZIL,Roraima,RioUraricoera,IlhadeMaracá;

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0.000 –20 000 –40 000 –100 000 –80 000 –60 000 –40 000 –40 000 –20 000 0.000

Fig.10. DistributionalmapofNeogriphoneuratimidaCurran.Yellowcirclescorrespondtopreviouslyknownrecords;blackcircleswithyellowrimcorrespondtothenew

records.

02–13/v/1987;J.Rafael,J.Brasil&L. Aquinoleg.; 5males and 12females (INPA). Idem;1male and1 female(ZUFMS). Ama-zonas,Manaus, INPA (3◦545S; 59◦5921W); 21/viii/2014; T. Mahlmannleg.;1female(ZUFMS).Maranhão,BomJardim,REBIO [Reserva Biológica] do Gurupi (3◦5649.23S; 46◦4923.02W); Armadilha McPhail, 17–27/i/2010, F. Limeira-de-Oliveira, J. T. Câmara&M.B.AguiarNeto,cols.;2malesand8females(CZMA). Ceará,Ubajara,ParqueNacionaldeUbajara,CachoeiradoCafundó (3◦5013S;40◦5435W);ArmadilhaMalaise;18–30/xi/2012;F. Limeira-de-Oliveira,J.S.PintoJuniorcols.;1female(CZMA).

Discussion

NeogriphoneuratimidaissimilartoitscongenerN.pacataMello

andSilva,2008a.Neogriphoneurapacatahasthesamepatternof

marksonthemesonotum,butdiffersfromN.timidabythe pres-enceofalarge,longitudinaldarkstripethroughthefrons.Thethree wrinkledandtubularspermathecaefoundinN.timidacorresponds innumberand shapewithitscongenersN.bispoi andN. corru-gata.AccordingtoMelloandSilva(2008a)thischaracterstateis likeasynapomorphysupportingacladewithinthegenus,asother Neogriphoneuraspeciespresentdifferencesinnumberandshape ofspermathecae(MelloandSilva,unpublisheddata).Thespecies

whichspermathecaewerenotobservedare:N.immaculata,N. lae-vifrons,N.pacata,N.schnusei,N.sordidaandN.striatifrons.

The differences in the morphology of the spermathecae observedinN.bispoi,N.corrugataandN.timidaseemtosuggest aconnectionwithalongerlengthoftheirsperm,indicatingthat spermcompetition(spermatozoaofsuccessivematingsmay com-peteforfertilization,accordingtoParker,1970)orcrypticfemale choice(the spermbedifferentially usedby thefemale,

accord-ingtoEberhard,1996)occuratthespermathecae;inthiscase,as

indicatedbyPitnicketal.(2009),wouldbepreferredtorefertoa mechanismofpostcopulatorysexualselection.

AccordingwithWilkinsonandJohns(2005),maleandfemale Dipterareproductivetraitsshouldexhibitcorrelatedpatternsof evolutionwithgreatdiversiﬁcationamongspeciesofﬂies consid-eringspermsize,spermnumber,andfemalestorageorgans.Future studiesinvolvingdetailedultra-structuralobservationsonthese threespeciescomparativetootherlauxaniidscouldhelptosolve thisquestion.

Neogriphoneurahas a New World distribution (Stuckenberg,

1971;Melloand Silva,2008a).Theoccurrenceof N.timidawas

ﬁrstrecognizedasbeingthroughoutcontinentalCentralAmerica inGuatemala,ElSalvador,CostaRicaandPanama(MelloandSilva,

2008a;GaimariandSilva,2010).Herein,thespeciesisadditionally

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perBrown,2009),Belize,andHonduras.Thisistheﬁrstrecordof thespeciesforSouthAmerica,occurringinBrazil,Colombiaand TrinidadandTobago,basedontheexamofmaterial;additionally, thereisa photoofthis speciesinasite (Cresswell,2010–2014) fromColombia,DepartmentofMagdalena.Thenewrecordsofthe species,intheNorthwestofSouthAmerica,formacontinuous dis-tributionalpatternintotheCentralandSouthAmericas.

Conﬂictsofinterest

Theauthorsdeclarenoconﬂictsofinterest.

Acknowledgments

Weareverygratefultothecuratorsfortheloanofthe mate-rial;toStephenGaimari(CDFA)forsendinglabeldatafromUSNM specimens;toFernandoJoséZara(UNESP/Jaboticabal)for provid-ingbibliographicmaterial;totheProgramadeApoio a Núcleos deExcelência(Pronex)oftheFundac¸ãodeAmparoàPesquisado EstadodoAmazonas(FAPEAM,Proc.1437/2007)coordinatedby JoséAlbertinoRafael,INPA,Manaus,forthephotomicroscopeused inthiswork;toF.Limeira-de-Oliveira(UEMA)forprovidingaccess toZCMAcollection;toRaduanA.Solemanforhelpingusproducing thedrawings;andtoFundac¸ãodeAmparoàPesquisadoEstadode SãoPaulo(FAPESP)grantstoRLM(Proc.03/11909-8).

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