Taxonomic revision of Plyomydas Wilcox & Papavero, 1971 with the description of two new species and its transfer to Mydinae (Insecta: Diptera: Mydidae)

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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

Taxonomic

revision

of

Plyomydas

Wilcox

&

Papavero,

1971

with

the

description

of

two

new

species

and

its

transfer

to

Mydinae

(Insecta:

Diptera:

Mydidae)

Stephanie

Castillo

a

_,

_Torsten

_Dikow

b,∗

a_San_Diego_State_University,_Department_of_Biology,_San_Diego,_United_States

b_National_Museum_of_Natural_History,_Smithsonian_Institution,_Department_of_Entomology,_Washington,_United_States

a

r

t

i

c

l

e

i

n

f

o

Articlehistory:

Received22December2016 Accepted9March2017 Availableonline24March2017 AssociateEditor:MarciaSoutoCouri Keywords: Mydasﬂies NeotropicalRegion Cybertaxonomy Openaccess

a

b

s

t

r

a

c

t

ThemonotypicNeotropicalMydidaegenusPlyomydasWilcox&Papavero,1971,todateconﬁnedto coastalPeru,isreviewed.Twonewspecies,Plyomydasadelphesp.nov.andPlyomydasphalarossp.nov.,are describedfrommid-elevationalwesternArgentina,whichextendsthedistributionofthegenus consid-erably.Distribution,occurrenceinbiodiversityhotspotssensuConservationInternational,andseasonal incidencearediscussed.Descriptions/re-descriptions,photographs,illustrations,andidentiﬁcationkeys areprovidedandmadeopenlyaccessibleindatadepositoriestosupportfuturestudiesoftheincluded taxa.PlyomydasistransferredfromtheLeptomydinaetotheMydinae:Messiasiinibasedontheabsence ofacanthophoritespinesonabdominaltergite10infemalesandthepresenceofveinM3+M4terminating inthecostalveinC.LeptomydinaeisthereforerestrictedtotheNorthernHemispherewiththeexception ofHessemydasKondratieff,Carr&Irwin,2005knownfromMadagascar.Messiasianotospila(Wiedemann, 1828)iscomparedtoPlyomydasspecies.

).

Introduction

Mydidaeﬂiesarea conspicuouspartoftheDipterafaunaof SouthAmerica.ThelargestknownMydidae,andpossiblyDiptera ingeneral,isGauromydasheros(Perty,1833)fromsouthernBrazil

(see Wilcox and Papavero, 1971; Calhau et al., 2015). Several

other species of this genus and species of Protomydas Wilcox, Papavero&Pimentel,1989arealsoverylarge,butrarelyobserved innature. Thediscoveryof relatively largespecimens of Mydi-daefromArgentinathatdonotﬁtthedescriptionofGauromydas Wilcox,Papavero&Pimentel,1989orProtomydasinseveralnatural historycollectionsinitiatedthepresentstudy.

Therearesome476Mydidaespeciesknownworld-wideand thelargestspeciesandgenericdiversityisfoundinsouthernAfrica

(Dikow,inpress).TheNeotropicalRegionhascurrentlysome85

speciesin 22generain sixsubfamilytaxa (Papavero,2009,see

alsoNeotropicaltaxoncatalogatasiloidﬂiesweb-site).Note that

theNeotropicalcatalogbyPapavero(2009)includesNearctictaxa ofEctyphinae,Leptomydinae,andRhaphiomidinae.

∗ Correspondingauthor.

E-mail:DikowT@si.edu(T.Dikow).

The Neotropical Leptomydinae include Plyomydas Wilcox & Papavero,1971,agenuswithasinglespeciesknownfromPeru, Nemomydas Curran, 1934 with eight central American species reachingasfarsouthasColombia,andpossiblyPseudonomoneura Bequaert,1961insouthernMexico.Ingeneral,Leptomydinaeis aNorthernHemispheretaxonwithonlyHessemydasKondratieff, Carr&Irwin,2005fromMadagascarandPlyomydasfromPeru pen-etratingtheSouthernHemisphere.

ThegenusPlyomydaswasdescribedfroma singlespecies, P. peruviensis Wilcox &Papavero, 1971, occurringin coastal Peru

(Wilcox and Papavero, 1971)and hasnot beenstudied further

since. New material of this genus hasbeen discovered in sev-eralnaturalhistorycollectionsthatextendthedistributionofthis taxontotheeasternAndesinwesternArgentinaaswellas low-elevationParaguay. These ﬂiesare relativelylargewitha wing lengthbetween15and21mm,butareveryrarein entomologi-calcollectionsasonly16specimens(ofatotalof29Plyomydas) werelocated.Inthisreview,wepresentataxonomicrevisionof PlyomydaswiththedescriptionoftwonewspeciesfromArgentina andthetransferofthegenustotheMydinae.

AlmeidaandLamas(2014)recordPlyomydasfromArgentinafor

theﬁrsttimebymakingreferencetoanundescribedspeciesand provideakeytogeneraof MydidaeoccurringinArgentina.The

http://dx.doi.org/10.1016/j.rbe.2017.03.002

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specimenshowninAlmeidaandLamas(2014,p.425)isPlyomydas phalarossp.nov.(MZSP-MZ003846).

Materialsandmethods

MorphologicalfeatureswereexaminedusinganOlympusSZ60 andaZeissSteREODiscovery.V12stereomicroscope.Winglength ismeasuredfromthetegulatothedistaltipofthewing.Thefemale andmaleterminaliawereﬁrstexcisedandmaceratedin10% potas-siumhydroxide(KOH)at55◦Cfollowedbyneutralizationinacetic acid(glacial,CH3COOH)andrinsingindistilledwater(H2O).They

weretemporarilystoredin75% ethanol(C2H5OH)for

examina-tionandillustrationand eventuallysealedinpolyethylenevials containing100%glycerine(C3H8O)andattachedtothespecimen’s

pin.

Terminology

TerminologyfollowsDikow(2009)and Cummingand Wood

(2009) (general morphology and abbreviations for setae),

Stuckenberg(1999)(antennae), and WoottonandEnnos(1989)

(wingvenation).Abdominaltergitesareabbreviatedinthe descrip-tionswith‘T’,andsternitesare abbreviatedwith‘S’. Theterms prothoracic,mesothoracic,andmetathoracicareabbreviated‘pro’, ‘mes’, and ‘met’, respectively. The term pubescence (adjective pubescent)referstotheshort,ﬁnemicrotrichiadenselycovering certainbodyparts.OthergeneralizedtermsfollowtheTorre-Bueno GlossaryofEntomology(Nichols,1989).

Speciesdescriptionsandre-descriptions

Speciesdescriptionsarebasedoncompositesofallspecimens and not exclusively onthe holotype and are compiled from a charactermatrixof144featuresand179characterstates

assem-bledwithLucidBuilder(version3.5)andeventuallyexportedas

natural-language descriptions. These species descriptions have beendepositedintheZenododatadepositoryandcanbeaccessed inXML-formatfollowingtheSDD(StructureofDescriptiveData) standard.The structureof terminaliais onlydescribed oncefor thegenusandadditionalspecies-speciﬁcfeaturesshouldbe inter-preted from the provided illustrations. All taxon names have been registered in ZooBank (Pyle and Michel, 2008). Previous taxondescriptionshavebeenmarked-upinTaxonXXMLlanguage

(Catapano,2010)anduploadedtothePlaziTreatmentBankfrom

wheretheyareaccessibleforhumanandmachinereading.

Specimenoccurrencedata

Thefollowing data onspecies occurrencesaregiven (where available): country, state/province, county, locality, geographic co-ordinates (formatted in both degrees minutes seconds and decimal latitude/longitude), elevation (in meters), date of col-lection (format: yyyy-mm-dd), habitat information, sampling protocol(if otherthan handnetting),collector,catalog number (a unique specimen number and any other identifying num-ber), depository (institution and collection code), number of specimens, sex, life stage, name of person who identified the specimen,andanyotherpreviousidentifications.Eachspecimen is listed with a unique specimen number (either an institu-tional catalog number or an AAM-XXXXXX number used by the junior author) that will allow the re-investigation as well asprovideauniqueLifeScienceIdentifier(LSID).Theoccurrence of all species is illustrated in distribution maps plotted with

http://www.simplemappr.netwithallofthoselocalitiesforwhich

co-ordinatesareavailable.Typelocalitiesareplottedwithasquare

symbolwhileallotherspecimensareplottedwithacircular sym-bol.

The distribution map includes Biodiversity Hotspots sensu

ConservationInternational (Mittermeier, 1998; Myers et al.,

2000; Mittermeier et al., 2005). The specimen occurrence

data are deposited as a DarwinCoreArchive(DwC-A) in

the GlobalBiodiversityInformationFacility(GBIF) using the

IntegratedPublishingToolkit(IPT)attheNMNH.

Photographsandillustrations

Whole habitus photographs of pinned specimens were taken using a VisionaryDigitalPassportII system (base and StackShot only), an OlympusOM-DEM-5 Micro 4/3 camera, a

60mmf2.8macrolens (equivalent to 120mm focal length in

35mm photography). The specimens were illuminated by a

FalconFLDM-i200LED dome-light for even and soft light.

Indi-vidualRAW-formatimageswerestackedusingHeliconFocusPro

(version 6.7) and exported in AdobeDNG-format. For further information on theimaging and post-production workﬂow see

Photographingasiloidﬂiesblogpost.Allphotographshave been

depositedinMorphbank::BiologicalImagingandimagesof addi-tionalspecimensareavailableonthatplatform,too.Theseimages willbeautomaticallyharvestedbytheEncyclopediaofLife(EOL)

andareavailableundertherespectivespeciespage.

Morphologicalfeatureswereillustratedusinga10×10ocular gridontheOlympusSZ60stereo microscopeandlaterdigitally convertedtovectorgraphicsusingAdobeIllustratorsoftware.The illustrationsofmaleandfemaleterminaliahavebeendepositedin digitalformat(svg-format)inﬁgshare.

Key

The dichotomous, interactive key has been built with

LucidPhoenix and the multi-access, matrix-based key with

LucidBuilderand bothcanbeaccessedonLucidcentraland the

juniorauthor’sresearchweb-site.

Institutionsprovidingspecimens

Institutions providing specimens are listed below, together withtheabbreviationsusedinthetextwhencitingdepositories (institutionCode),alinktotherecordintheGlobalRegistryof Bio-diversityRepositories(GRBio),andthepeoplewhokindlyassisted:

AMNH –American Museumof NaturalHistory, New YorkCity, NewYork,USA(D.Grimaldi);BMNH–NaturalHistoryMuseum, London,UK(E.McAlister);CAS–CaliforniaAcademyofSciences, SanFrancisco, California, USA(M. Trautwein); CNC – Canadian NationalCollectionofInsects,ArachnidsandNematodes,Ottawa, Ontario,Canada(J.Skevington);CSCA–CaliforniaStateCollection ofArthropods,Sacramento,California,USA(M.Hauser,E.Fisher);

IMLA–FundacioneInstitutoMiguelLillo,UniversidadNacional deTucumán,Tucumán,Argentina(E.ConstanzaPerez);IADIZA– InstitutoArgentinodeInvestigazionesdelasZonasÁridas, Men-doza,Argentina(C.Dominguez,S.Roig);MNHN–Museumnational d’Histoirenaturelle,Paris,France(C.Daugeron,E.Delfosse);MUSM

–MuseodeHistoriaNatural,UniversidadNacionalMayordeSan Marcos,Lima,Peru(G.Lamas,P.SanchezFlores);MZSP–Museude Zoologia,UniversidadedeSãoPaulo,SãoPaulo,Brazil(C.Lamas);

SMNS – Staatliches Museum für Naturkunde,Stuttgart, Baden-Württemberg, Germany(H.-P. Tschorsnig);ZSMC–Zoologische Staatssammlung,München,Bayern,Germany(M.Kotrba).

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Dataresources

GBIF: specimen occurrence data-set –

7f2de377-9f9f-4276-b0b8-5c792b499223–DOI10.15468/tiymja.

ﬁgshare: illustration collection ID – DOI 10.6084/m9.

ﬁgshare.c.3650510.

LucidBuilder:illustrated,multi-entry,matrix-based

identiﬁca-tionkey–keys.lucidcentral.org/keys/v3/plyomydas/.

LucidPhoenix: illustrated, dichotomous identiﬁcation key –

keys.lucidcentral.org/keys/phoenix/plyomydas/.

Morphbank:imagecollectionID–860614.

PlaziTreatmentBanktaxontreatments:

WilcoxandPapavero,1971–AD29FFB3FF8E2724FFC56619FFF

DFFF6

SimpleMappr:distributionmap–6102(asinFig.27)–Google

EarthKMLﬁle6102.

Zenodo: natural-language species descriptions from Lucid

BuilderinSDDformat–DOI10.5281/zenodo.154750.

ZooBanknomenclatorialacts:

WilcoxandPapavero,1971B57257ED-AF72-4AEB-AF09-9047

AEDF7D3F

CastilloandDikow2017FF635991-5F13-4B7D-A3E4-D314ABB

E9E98.

Taxonomy

PlyomydasWilcoxandPapavero,1971

ZooBank43AF4DE6-F7DE-4F35-801F-BF4248D531BB

Plazi TreatmentBank 43AF4DE6-F7DE-4F35-801F-BF4248D53 1BB

Plyomydas Wilcox and Papavero, 1971: 106. Type-species: PlyomydasperuviensisWilcoxandPapavero,1971,byoriginal des-ignation.

Diagnosis:ThegenusisdistinguishedfromotherMydinaebythe absenceofaventralmetathoracictibialkeel(Figs.2and20),only slightlyenlargedmetathoracicfemora(Figs.2and20),andapartly pubescentscutum(eitherstripesFig.12orspotsFig.19).Messiasia notospila(Wiedemann,1828)(seeFigs.28–33)issomewhat simi-lartoPlyomydasbecauseitexhibitsaweaklydevelopedtibialkeel andagreypubescencepatternonthescutum,butcanbeseparated fromPlyomydasbasedonthemoreexpandedmetathoracicfemur andmaleterminalia.Thedistinctventro-posteriorgonocoxal pro-cessismuchmoredevelopedinPlyomydasthanitisinMessiasia andthetwootherposteriorgonocoxalprocesses(seeFig.7)are uniquetoPlyomydaswithinMydinae(J.Calhau,pers.comm.).See alsoDiscussionfortransferofPlyomydastoMydinae:Messiasiini.

Description:F*abdomenandgenitalia(Figs.10and11):densely arrangedanteriorlydirected setaeabsentonposterior Tand S; T8 with broad anterior rectangular apodeme; T9 simple, rect-angular; T9+10 entirely fused, T10 formed by single sclerite, acanthophoritespinesabsent;3spermathecae,allequallylarge, formedby±expandedweaklysclerotizedducts;individual sper-mathecal duct short; S9 (furca) formed by 1 sclerite, inverted U-shaped(joinedanteriorly,separatedposteriorly),anterior fur-calapodemeabsent,lateralfurcalapodemepresent,medianfurcal bridgeabsent.

M* abdomen and terminalia (Figs. 7–9, 21–26): T1–7 well-developed, entirely sclerotized, T8 postero-medially weakly sclerotized,withanteriortransversesclerotizedbridgeconnecting lateralsclerites;T7–8anteriorlywith2lateralapodemes;S6 regu-lar,withoutanyspecialsetationpostero-medially;S8simpleplate, entire(undivided)ventro-medially,withhorn-likeantero-lateral processes, not fused to T8 dorso-laterally; epandrium formed bytwo sclerites,separatedmediallyand onlyjoininganteriorly, distally in dorsal view pointed postero-laterally; subepandrial sclerite without lateral or median protuberances; hypandrium

concave,cup-shaped,entirelysclerotizedventrally,entirelyfused with gonocoxite, forming a gonocoxite-hypandrial complex, supra-hypandrialscleriteabsent;gonocoxitesimple,short, hook-like,withoutmedianorlateralprotuberance,gonocoxalapodeme present,short(atmostslightlyextendinghypopygiumanteriorly); 1functionalphallicprong,shortandwide,phallicepimerepresent, distallysimple,evenlyrounded;lateralejaculatoryprocessabsent; ejaculatoryapodeme formedby single dorso-ventrally oriented plate;ventro-medianmarginofparameralsheathheavily sclero-tized(appearingentirelyclosed); parameralsheathlong,sperm sacentirelycovered;spermsacappearing±heavilysclerotized.

Plyomydasadelphesp.nov.

ZooBank27B0A7A0-0548-4F53-97AF-BCFD98B057FC

Figs.1–11,27

Diagnosis:Thespeciesisdistinguishedfromcongenersbythe scutalpubescencepatternwithonlythreepairsofgreypubescent spots(Figs.1and5)andtheabsenceofagreypubescentstripeon theposterioranepisternum(Fig.2).

Etymology: From Greek adelphe=sister, referring to overall morphologicalsimilaritytoPlyomydasphalarossp.nov.

Description: Head: black, in general grey pubescent; width distinctly greater than thorax, interocular distance on vertex largerthanatventraleyemargin,vertexbetweencompoundeyes slightly depressed, parafacialarea very narrow,facial gibbosity nearlytouchingmedianeyemargin;facialgibbositydistinct, well-developedanddiscernibleinlateralview;mystaxbrown,covering facialgibbosityexceptdorso-medially;fronsnotelevated,laterally greypubescent,mediallyapubescent;vertexapubescent;postgena lightlygreypubescent; setation: vertexblack, fronsbrown, ocp setaeyellow,poclmacrosetaeblack;ocellartriangleapubescent; proboscisbrown, long,reachingfronto-clypealsuture; labellum large,muchwiderthanprementum,longerthanprementumandas longasoralcavity,unsclerotizedlaterally;maxillarypalpus cylin-drical,lightbrown,about1/3lengthofproboscis.

Antenna:brown,bulbouspartofpostpedicellightbrown,scape andpedicelblacksetosedorsallyandventrally,rarelybrownsetose dorsallyandventrally;postpedicelcylindricalinproximal½, sym-metricallybulbousin distal ½,≥5.0 timesas longascombined lengthofscapeandpedicel,bulbouspartbrownsetosedorsally andventrally;apicalseta-likesensoryelementsituatedapicallyin cavityonpostpedicel.

Thorax:blackorbrown,predominantlyapubescentwith dis-tinctgreypubescentscutalandpleuralspots;scutumuniformly black,surfacepredominantlysmooth,postero-paramediallyrivose, lightlygreypubescentwithdistinctgreypubescentspots:1pair antero-paramedially,2 pairs laterally(1 attransverse suture,1 postero-laterally), scutalsetation comprised ofpubescent spots long black or brown setose; dc setae pre- and postsuturally black,acr setae absent,lateralscutalsetaeblack or brown,npl setae0,spalsetae0,palsetae0;antepronotumdorso-medially smooth (without anyindentation); postpronotallobebrownto black,lightlygreypubescent;proepisternum,lateral postprono-tum,andpostpronotallobelongblacksetose;scutellumlightlygrey pubescent,brownsetose,apicalscutellarsetaeabsent; mesopost-notum,anatergite,andkatatergitelightlygreypubescent,asetose, katatergiteelevatedandsmoothlyconvex;anterioranepisternum lightly grey pubescent, asetose, supero-posterior anepisternum apubescent, asetose; posterior anepimeron long brown setose, katepimeron asetose; metanepisternum lightly grey pubescent, asetose, metepimeron evenly elevated, same color as T1, grey pubescent,asetose;infra-halterscleriteabsent.

Leg: light brown to brown, setation predominantly black; pro, mes, and met coxa lightly grey pubescent, black setose; met trochanter setose medially; femur black or brown, met

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Figs.1–6.PhotographsofmaleandfemalePlyomydasadelphesp.nov.:(1)M*holotype(USNMENT01242234),dorsal(Morphbank#860756);(2)same,lateral(#860758);

(3)same,headanterior(#860760);(4)F*paratype(AAM-004053),headanterior(#860784);(5)same,dorsal(#860780);(6)same,lateral(#860782).Scaleline=5mm.

femur±cylindricalonlyslightlywider thanproandmesfemur, indistal ½ macrosetose,1antero-ventraland 1postero-ventral rowofmacrosetae,postero-ventrallylongbrownsetose;proand mestibialaterally arched,mettibiastraight,mettibia cylindri-cal,ventralkeelabsent,postero-laterallyregularsetose;proand mestarsomere1longerthantarsomere2,butlessthancombined lengthoftarsomeres2–3,mettarsomere1aslongascombined lengthoftarsomeres2–3;pulvilluswell-developed,aslongas well-developedclaw,andaswideasbaseofclaw;setiformempodium absent.

Wing:length=15.2–18.5mm; lightbrownstained,especially alongveins,distal¼andposteriormarginunstained,veinsbrown, microtrichiaabsent;cellsr1,r4,r5,m3,+cupclosedexceptr5open;

Cwell-developed,aroundentirewing;R4terminatesinR1;R5

ter-minatesinR1;stumpvein(R3)atbaseofR4present,longbutnot

reachingR2;R4andR5±parallelmedially;r-mdistinct,R4+5and

M1 apart,connectedbycrossvein; M1 curves slightly anteriorly

atr-m,M1(orM1+M2)terminatesinC;M4andCuAsplit

prox-imallytom-cu(cellm3 narrowproximally); M3+M4 terminate

togetherinC(reachingwingmargin);CuPstraight,cellcupwide, CuPandwingmarginfurtherapartproximallythandistally;alula well-developed,verylargeandpartlyoverlappingwithscutellum medially;halterbrown.

Abdomen: brown to metallic bluish-black; setation com-prised of scattered black setae, surface entirely smooth; T1–7 brownwithnarrowyellowposterior margin;T1 longbrownto blacksetose,T2–7shortblacksetose;T1lightlygreypubescent, T2–7apubescent;S1brownwithlightbrownposteriormargin, S2–6 brown withnarrow yellow posterior margin; S1 asetose, S2–7 sparsely black setose; S apubescent; T2–4 parallel-sided and not constrictedwaist-like; bullae onT2 black,transversely elongate, surface entirelysmooth, T2surface anterior tobullae smooth.

Typelocality:Argentina:Mendoza:Rt.142atMendozaRiver, 32◦3529S068◦1728W(−32.59139−68.29111).

Materialexamined:Argentina:Mendoza:1M*Rt.142at Men-dozaRiver,21kmNECostadeAraujo,32◦3529S068◦1728W, −32.59139−68.29111,585m,2015-12-04,Steiner,W., Swearin-gen, J. (USNMENT01242234, Holotype, IADIZA); 1F* Mendoza, 35◦5200S 068◦4900W, −32.88944 −68.84583, 1907-00-00 (AAM-004053,Paratype,ZSMC);LaRioja:1M*CuestadeMiranda, 29◦1900S067◦3900W,_−29.31667_−67.65,1700m,1977-11-29, Willink,A.,Fidalgo(AAM-002719,Paratype,MZSP).

Distributionandbiodiversityhotspot:WesternArgentinainthe provincesofLaRiojaandMendoza(Fig.27).Notknowntooccurin anybiodiversityhotspot.

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Figs.7–11.MaleandfemaleterminaliaofPlyomydasadelphesp.nov.:(1)M*paratype(MZSP-MZ003844),lateral(posteriorgonocoxalprocesseshighlightedingrey);(2)

same,dorsal;(3)same,ventral;(4)F*paratype(AAM-004053),dorsal;(5)same,ventral.Scaleline=1mm,setationomitted.FigshareDOI10.6084/m9.ﬁgshare.4490462.

Remarks:Sexualdimorphismisminimal.Basedonthethree knownspecimens,femalesarelarger(winglength18.5mminthe solefemale paratype, 15.2–16.6mm in two males) and dimor-phismonlyoccursincoloroflegsandsclerites.Theholotypewas collectedinlowﬂatmesquitedesertanddryriverbed.

PlyomydasperuviensisWilcoxandPapavero,1971:108

ZooBankF954F6B0-BC56-4C16-BB82-0D2734030993

Plazi TreatmentBank 511087CB-FFCB-276D-FC9D-67B4FB8EF D41

Figs.12–14,21–23,27

Diagnosis: The species is distinguished from congeners by thescutalpubescencepatternwiththreedistinctgreypubescent stripes(nospots,Fig.12),thepresenceofagreypubescentstripeon theposterioranepisternum(Fig.13),anditsdistributionincoastal Peru(Fig.27).

Description: Head: brown, facial gibbosity light brown, in generalgreypubescent;widthdistinctlygreaterthanthorax, inte-roculardistanceonvertexlargerthanatventraleyemargin,vertex betweencompoundeyesslightlydepressed,parafacialareavery narrow,facialgibbositynearlytouchingmedianeyemargin;facial gibbositydistinct,well-developedanddiscernibleinlateralview; mystaxyellow,sparse;fronsnotelevated,entirelygreypubescent; vertexentirelygreypubescent;postgenalightlygreypubescent; setation: vertex yellow, frons brownor light brown, ocp setae yellow,poclmacrosetaelightbrown;ocellartriangleapubescent; proboscisbrown,long, reachingfronto-clypealsuture; labellum large,muchwiderthanprementum,longerthanprementumandas longasoralcavity,unsclerotizedlaterally;maxillarypalpus cylin-drical,lightbrown,about¼lengthofproboscis.

Antenna:brown,bulbouspartofpostpedicellightbrown,scape andpedicelbrownsetosedorsallyandventrally;postpedicel cylin-dricalinproximal2/5,symmetricallybulbousindistal3/5,≥5.0 timesaslongascombinedlengthofscapeandpedicel,bulbous

partbrownsetosedorsallyandventrally;apicalseta-likesensory elementsituatedapicallyincavityonpostpedicel.

Thorax:lightbrown,predominantlyapubescentwithdistinct greypubescentscutalstripesandpleuralspots;scutumuniformly brown, surface predominantly smooth, postero-paramedially rivose, predominantly grey pubescent, apubescent paramedian andmedianstripes(notreachingposteriormargin),scutal seta-tioncomprisedofpubescentstripesshortblacksetose;dcsetae pre-andpostsuturallybrown,acrsetaeabsent,lateralscutalsetae brown,nplsetae0,spalsetae0,palsetae0;antepronotum dorso-medially smooth (without any indentation); postpronotal lobe lightbrown,greypubescent;proepisternum,lateralpostpronotum, andpostpronotallobeshortbrownsetose;scutellumlightlygrey pubescent,laterallydenselygreypubescent,brownsetose,apical scutellarsetaeabsent;mesopostnotum,anatergite,andkatatergite lightlygreypubescent,asetose,katatergiteelevatedandsmoothly convex; anterior anepisternum lightly grey pubescent, asetose, supero-posterioranepisternumgreypubescent,asetose;posterior anepimeronshortbrownsetose,katepimeronasetose; metanepis-ternumgrey pubescent, asetose, metepimeron evenly elevated, same color as T1, grey pubescent, asetose;infra-halter sclerite absent.

Leg: light brown to brown, setation predominantly brown; pro, mes,and met coxa lightly grey pubescent, brown setose; met trochanter setose medially; femur light brown, met femur±cylindricalonlyslightlywider thanproandmesfemur, indistal ½ macrosetose,1antero-ventraland 1postero-ventral rowofmacrosetae,postero-ventrallylongbrownsetose;proand mestibialaterally arched,mettibiastraight, mettibia cylindri-cal,ventralkeelabsent,postero-laterallyregularsetose;proand mestarsomere1longerthantarsomere2,butlessthancombined lengthoftarsomeres2–3,mettarsomere1aslongascombined lengthoftarsomeres2–3;pulvilluswell-developed,aslongas well-developedclaw,andaswideasbaseofclaw;setiformempodium absent.

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Figs.12–14.PhotographsofmalePlyomydasperuviensis(AAM-003731):(12)dorsal(Morphbank#860772);(13)same,lateral(#860774);(14)same,headanterior(#860776). Scaleline=5mm.

Wing:length=10.7–13.0mm;slightlybrownstained through-out, veins brown, microtrichia absent; cells r1, r4, r5, m3,+cup

closedexceptr5 open;Cwell-developed,aroundentirewing;R4

terminatesinR1;R5terminatesinR1;stumpvein(R3)atbaseofR4

present,longbutnotreachingR2;R4andR5±parallelmedially;

r-mdistinct,R4+5andM1apart,connectedbycrossvein;M1curves

slightlyanteriorlyatr-m,M1(orM1+M2)terminatesinC;M4and

CuAsplitproximallytom-cu(cellm3narrowproximally);M3+M4

terminatetogetherinC(reachingwingmargin);CuPstraight,cell cupwide,CuPandwingmarginfurtherapartproximallythan dis-tally;alulawell-developed,verylargeandpartlyoverlappingwith scutellummedially;halterlightbrown.

Abdomen:browntometallicbluish-black;setationcomprised ofsparselyscatteredshortbrownsetae,surfaceentirelysmooth; T1–7 brown with narrow yellow posterior margin; T1 long lightbrownsetose,T2–7sparselybrownsetose;T1lightlygrey pubescent,T2–7apubescent;S1brownwithlightbrownposterior margin,S2–6brownwithnarrowyellowposteriormargin;S1 ase-tose,S2–7sparselyblacksetose;Sapubescent;T2–4parallel-sided and notconstricted waist-like; bullae onT2black, transversely elongate, surfaceentirely smooth,T2 surface anterior tobullae smooth.

F*abdomen:Nofemalewasavailablefordetailedstudy. Type locality: Peru: Lima: Lima, 12◦0300S 077◦0200W (−12.05−77.03333).

Materialexamined:Peru:Ica:1F*EglisedePisco,13◦4236S 076◦1212W, −13.71 −76.20333, 1954-12-29, Dorst (AAM-001235, MNHN); La Libertad: 1M* Cartavio, 07◦5330S 079◦1322W,−7.89167−79.22278,0000-00-00,Smyth,E. (AAM-003729,Paratype,AMNH); 1M*Cartavio,0000-00-00,Smyth,E. (AAM-003730, Paratype, AMNH); Lima: 1M* Lima, 12◦0300S 077◦0200W, −12.05 −77.03333, 1939-02-08, Weyrauch, W. (MZSP-MZ001668, Holotype, MZSP); 1M* Lima, 1968-01-15, Picho, H. (MZSP-MZ003836, Paratype, MZSP); 1F* Lima, 1969-02-11,Razuri,V.(MZSP-MZ003837,Paratype,MZSP);1M*Lima, 1939-02-08,Weyrauch,W.(CNCDiptera198085,CNC);1M*Lima,

1948-04-00(AAM-003005,MUSM); 1M*Hacienda Chuquitanta, 12◦0432S076◦5700W,−12.07556−76.95,1974-03-10,Lamas, G.,Medina,N.(AAM-003742,MUSM);1M*Antioquia,12◦0449S 076◦3036W, −12.08028−76.51,1975-01-11,Garcia, R. (AAM-003737, MUSM); 1M* La Molina, 12◦0454S 076◦5545W, −12.08167 −76.92917, 280 m, 1966-03-11, Garcia, R. (AAM-003769,MUSM);1M* LaMolina, 280m,1966-03-11,Garcia, R. (AAM-003731, MUSM);1F*nolocality data (BMNH(E)1202927, BMNH).

Distributionandbiodiversityhotspot:CoastalPeru(Fig.27).Not knowntooccurinanybiodiversityhotspot.

Remarks:Thedescriptionaboveisbasedonmalespecimens only because no females were available for detailed study to provideanupdatedF*descriptionandinclusioninthismanuscript althoughthree specimens(see above)had beenstudiedseveral years ago and can be unambiguously assigned to this species. Sexualdimorphismismostlikelyminimalasintheotherspecies andthefemaleterminaliawillmostlikelybeidenticaltoP.adelphe sp.nov.andP.phalarossp.nov.ThefemalespecimenintheBMNH (BMNH(E)1202927) was apparently studied by Macquart and labeledMydasperuvianus,butneverdescribedasanewspeciesby him.

Plyomydasphalarossp.nov.

ZooBankAA0C21E1-0420-4A8A-93F3-40DC17864BCE

Figs.15–20,24–27

Diagnosis:Thespeciesisdistinguishedfromcongenersbythe scutalpubescencepatternwithfourpairsandapostero-median greypubescentspot(Figs.15–16,19–20),andthepresenceofagrey pubescentstripeontheposterioranepisternum(Figs.16and20).

Etymology:FromGreekphalaros=white-spotted,referring to thedistinctivegreypubescentspotsonthescutum.

Description:Head:brown, in generalgrey pubescent;width distinctly greater than thorax, interocular distance on vertex largerthanatventraleyemargin,vertexbetweencompoundeyes

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Figs.15–20. PhotographsofmaleandfemalePlyomydasphalarossp.nov.:(15)M*holotype(AAM-005629),dorsal(Morphbank#860834);(16)same,lateral(#860836);(17)

same,headanterior(#860838);(18)F*paratype(MZSPMZ003846),headanterior(#860752);(19)same,dorsal(#860524);(20)same,lateral(#860750).Scaleline=5mm.

Figs.21–26. MaleterminaliaofPlyomydasspecies:(21)Plyomydasperuviensis(AAM-003729),lateral;(22)same,dorsal;(23)same,ventral;(24)Plyomydasphalarossp.nov. paratype(CASENT8380022),lateral;(25)same,dorsal;(26)same,ventral.Scaleline=1mm,setationomitted.FigshareDOI10.6084/m9.ﬁgshare.4490474.

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Fig.27.MapofSouthAmericawithelevationalreliefandbiodiversityhotspots(ingrey)showingdistributionofPlyomydasadelphesp.nov.(orange),P.peruviensis (yellow),andP.phalarossp.nov.(blue).Typelocalitiesplottedwithsquaresymbols.MapdataavailableinGoogleEarthKMLﬁle6102andalsothroughGBIF(data-set 7f2de377-9f9f-4276-b0b8-5c792b499223,DOI10.15468/tiymja).

slightly depressed, parafacialarea very narrow,facial gibbosity nearlytouchingmedianeyemargin;facialgibbositydistinct, well-developedanddiscernibleinlateralview;mystaxbrown,covering facialgibbosityexceptdorso-medially;fronsnotelevated,laterally greypubescent,mediallyapubescent;vertexapubescent;postgena lightlygreypubescent;setation:vertexbrown,fronsbrown,ocp setaeyellow,poclmacrosetaebrown;ocellartriangleapubescent; proboscisbrown, long,reachingfronto-clypealsuture; labellum large,muchwiderthanprementum,longerthanprementumandas longasoralcavity,unsclerotizedlaterally;maxillarypalpus cylin-drical,lightbrown,about1/3lengthofproboscis.

Antenna:brown,bulbouspartofpostpedicellightbrown,scape andpedicelblacksetosedorsallyandventrally,rarelybrownsetose dorsallyandventrally;postpedicelcylindricalinproximal½, sym-metricallybulbousin distal ½,≥5.0 timesas longascombined lengthofscapeandpedicel,bulbouspartbrownsetosedorsally andventrally;apicalseta-likesensoryelementsituatedapicallyin cavityonpostpedicel.

Thorax:brown,predominantlyapubescent withdistinctgrey pubescentscutalandpleuralspots;scutumuniformlyblack, sur-facepredominantlysmooth,postero-paramediallyrivose,lightly

greypubescentwithdistinctgreypubescentspots:1pair antero-paramedially,3pairslaterally(1antero-laterally,1attransverse suture,1postero-laterally),1spotpostero-medially,scutal seta-tioncomprised of pubescentspots longblack orbrownsetose; dc setae pre- and postsuturally black, acr setae absent, lat-eral scutal setaebrown, nplsetae 0, spal setae 0, palsetae 0; antepronotumdorso-mediallysmooth(withoutanyindentation); postpronotallobebrown,predominantlygreypubescent; proepis-ternum,lateralpostpronotum, andpostpronotallobelongblack setose; scutellumlightly grey pubescent,laterally densely grey pubescent,brownsetose,apicalscutellarsetaeabsent; mesopost-notum,anatergite,andkatatergitelightlygreypubescent,asetose, katatergiteelevatedandsmoothlyconvex;anterioranepisternum lightly grey pubescent, asetose, supero-posterior anepisternum greypubescent,asetose;posterioranepimeronlongbrownsetose, katepimeron asetose;metanepisternum lightly grey pubescent, asetose, metepimeron evenly elevated, same color as T1, grey pubescent,asetose;infra-halterscleriteabsent.

Leg:brown,setationpredominantlyblack;pro,mes,andmet coxalightlygreypubescent,blacksetose;mettrochantersetose medially;femurbrown,metfemur±cylindricalonlyslightlywider

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thanproandmesfemur,indistal½macrosetose,1antero-ventral and1postero-ventralrowofmacrosetae,postero-ventrallylong brownsetose;proandmestibialaterallyarched,mettibiastraight, mettibiacylindrical,ventralkeelabsent,postero-laterallyregular setose;proandmestarsomere1longerthantarsomere2,butless thancombinedlengthoftarsomeres2–3,mettarsomere1aslong ascombinedlengthoftarsomeres2–3;pulvilluswell-developed,as longaswell-developedclaw,andaswideasbaseofclaw;setiform empodiumabsent.

Wing:length=16.7–20.8mm;light brownstained,especially alongveins,distal¼andposteriormarginunstained,veinsbrown, microtrichiaabsent;cellsr1,r4,r5,m3,+cupclosedexceptr5open;

Cwell-developed,aroundentirewing;R4terminatesinR1;R5

ter-minatesinR1;stumpvein(R3)atbaseofR4present,longbutnot

reachingR2;R4andR5±parallelmedially;r-mdistinct,R4+5and

M1 apart,connectedbycrossvein;M1 curves slightlyanteriorly

atr-m,M1(orM1+M2)terminatesinC;M4 andCuAsplit

prox-imallytom-cu(cellm3 narrowproximally); M3+M4 terminate

togetherinC(reachingwingmargin);CuPstraight,cellcupwide, CuPandwingmarginfurtherapartproximallythandistally;alula well-developed,verylargeandpartlyoverlappingwithscutellum medially;halterbrown.

Abdomen:browntometallicbluish-black;setationcomprised of scattered black setae, surface entirely smooth; T1–7brown withnarrow yellow posterior margin; T1 long brown to black setose,T2–7shortblacksetose;T1lightlygreypubescent,T2–7 apubescent;S1brownwithlight brownposterior margin,S2–6 brownwith narrow yellowposterior margin; S1 asetose, S2–7 sparselyblacksetose;Sapubescent;T2–4parallel-sidedandnot constrictedwaist-like;bullaeonT2black,transverselyelongate, surfaceentirelysmooth,T2surfaceanteriortobullaesmooth.

Type locality: Argentina: Catamarca: Belén, 27◦3859S 067◦0158W(−27.64972−67.03278).

Material examined: Argentina: Catamarca: 1M* Belén, 27◦3859S 067◦0158W, _−27.64972 _−67.03278, 1975-11-25, Bohart,R.(AAM-005629,Holotype, CSCA);1F*1M*Belén,6km S, 27◦4347S 067◦0115W, −27.72972 −67.02083, 1975-11-25, Stange, L. (M* AAM-002714, F* AAM-002718, Paratypes, IMLA); 1F* 4M* Belén, 6km S, 1975-11-25, Stange, L. (MZSP-MZ003838–MZSP-MZ003841, MZSP-MZ003846, Paratypes, MZSP);Salta: 1M*Cafayate,22kmN, 26◦0256S065◦5401W, −26.04889−65.90028, 1578m, 2003-11-13,whitesand dunes, Irwin,M.,Parker,F.(CASENT8380021–CASENT8380022,Paratypes, CAS); 1M* Cafayate, 26◦0406S 065◦5829W, −26.06833 −65.97472,1700m, 1991-01-05,dune,Osten,T.(AAM-003864, Paratype, SMNS); 1M* Cafayate, 1993-11-14, Rozen, J., Rozen, B. (AAM-004054, Paratype, AMNH); Paraguay: San Pedro: 1M* Carumbé, 24◦0846S 056◦3730W, −23.95 −56.6, 1973-01-10–1973-02-10, Golbach, R. (MZSP-MZ003843, Paratype, MZSP).

Distributionandbiodiversityhotspot:WesternArgentina (Cata-marcaandSalta)andParaguay(SanPedro)(Fig.27).Thegeographic co-ordinatesforCarumbé(Paraguay)aretakenfromGBIFrecord

416921317. The easternmost distribution of this species in

ParaguayoverlapswiththeAtlanticForestbiodiversityhotspot. Remarks:Sexualdimorphismisminimalwithonlyslight differ-encesincolorofscleritesorsetae.Thetwofemalesstudiedhave awinglengthof19.3–19.4mmwhilemaleshave awinglength of 16.7–20.8mm. Two specimens (AAM-003864, AAM-002715) lackthepostero-mediangreypubescentspotdirectlyanteriorto thescutellum.WebelievethatthesetwospecimensbelongtoP. phalarossp.nov.though.ThedisjunctdistributionofP.phalaros sp.nov. in mid-elevationwestern Argentina and low-elevation Paraguay is interesting.We haveno reasonto believethat the Paraguayan specimen (AAM-002720) is mislabeled as a search ontheinternetrevealsthatotherinsectshavebeencollectedby

R.GolbachinCarumbéinthe1970s(Googlesearch)providing evi-dencethatthespecimeniscorrectlylabeled.

Keytospecies

1. Scutumpredominantlydenselygreypubescent,onlymedian stripe(notreachingposteriormargin)andparamedianstripes apubescent(Fig.12);coastalPeruvianAndes...peruviensis - Scutumpredominantlyapubescent,withonlywidelyseparated

anteriorandlateralgreypubescentspots(Fig.1);western ArgentinaandParaguay...2

2. Scutumwithonlythreepairsofgreypubescentspots (antero-lateralspotposteriortopostpronotallobeabsent, Figs.1–2)andwithoutpostero-medianpubescentspot(Fig.1); scutellumonlylightlypubescent;anepisternumwithoutposterior pubescentstripe(Fig.2);postpronotallobelightlygreypubescent ...adelphesp.nov.

- Scutumwithfourpairsofgreypubescentspots(antero-lateral spotposteriortopostpronotallobedistinct,Figs.15and16)and withpostero-mediangreypubescentspot(inadditiontoother spotsalthoughabsentintwostudiedspecimens,Figs.15and19); scutellumdenselygreypubescentlaterally;anepisternumwith narrowposteriorpubescentstripe(Figs.16and20);postpronotal lobedistinctlygreypubescent...phalarossp.nov.

Discussion

Maleterminaliaareoftenreliableindelimitingspecies bound-aries,buttheycanbeconservedamongspeciesofagenusoreven acrossgenericboundariessuchasinAfrotropicalSyllegomydinae

(Dikow,inpress).WithinPlyomydas,themaleterminaliaarevery

similarinallthree speciesand cannoteasilybedescribed with wordssothat wedeferthereader tocomparetheillustrations

(Figs. 7–9, 21–26).Wepostulate that inthis taxonthethoracic

pubescencepattern isreliable forspecies delimitationand that comparisonofmaleterminaliaisunnecessaryatthisstage.

TransferfromLeptomydinaetoMydinae:Messiasiini

WilcoxandPapavero(1971),Papavero(2009),andPapaveroand

Artigas(2009)considerPlyomydastobelongtoLeptomydinae.This

ischieﬂybasedontheabsenceofakeelontheventral metatho-racictibia,whichis absentinPlyomydasandLeptomydinae,but presentinvirtuallyallMydinae(anexceptionisMessiasianotospila (Wiedemann,1828)knownfromArgentina,Brazil,Paraguay,and Uruguaywithaweaklydevelopedkeel(Figs.28–33)).Weregard theabsenceofthemetathoracictibialkeeltobeasimplelossofthis featureandnotanimportantfeaturetoplacethegenus Plyomy-dasinLeptomydinae.Instead,asinallMydinae,andseveralother Mydidaegenera,veinM3+M4reachesthecostalveinCinPlyomydas

(Fig. 12) and which is absentin all Leptomydinae genera. Fur-thermore,thefemaleterminaliaofPlyomydaslackacanthophorite spinesandtergite10isasinglesclerite(Fig.10)asfoundinall Mydinae,Cacatuopyginae,andthreegeneraofSyllegomydinaein theAfrotropicalRegion(Dikow,inpress).AllknownLeptomydinae, incontrast,utilizeadividedtergite10withacanthophoritespines toovipositinsandorsoil.Fromtheshapeofthefemaleterminalia wecanpostulatethatPlyomydasspeciesovipositineitherdecaying woodorantnestsashasbeenobservedinotherMydinaetaxa(see

PapaveroandArtigas,2009).

NotethatthekeytoNeotropicalsubfamiliesinPapaveroand

Artigas(2009)isincorrectincouplet11whereitshouldread“Hind

tibiawithventralkeel(carinate).SubfamilyMydinae...12”and “Hindtibiacylindrical.SubfamilyLeptomydinae...23”.Thiskey willkeyoutPlyomydascorrectly,butwithinLeptomydinaeandnot withinMydinaeasproposedhere.

TheMydinaearecurrentlydividedintofourtribes(Papavero

andWilcox, 1974;Wilcoxet al.,1989)and weplacePlyomydas

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Figs.28–33. PhotographsofmaleandfemaleMessiasianotospila(Argentina:EntreRios:Pronunciamiento,32◦₂₀₄₅_S₀₅₈◦₂₆₄₀_W,_1970-03-00,_Bolle,_S.):₍₂₈₎_M*

(CNCDiptera198880),dorsal(Morphbank#860859);(29)same,lateral(#860861);(30)same,headanterior(#860863);(31)F*(CNCDiptera198880),headanterior(#860870);

(32)same,dorsal(#860866);(33)same,lateral(#860868).Scaleline=5mm.

includedgenussofar,inantennal,proboscis,andfemaleterminalia morphology.WithPlyomydasnowinMydinae,Leptomydinaeisa taxonrestrictedtotheNorthernHemispherewiththeexceptionof HessemydasfromMadagascar.

Seasonalincidence

SpeciesofPlyomydashaveonlybeencollectedduringthe South-ernHemispheresummertoearlyautumn.Plyomydasadelphesp. nov.hasbeencollectedinNovember–December,Plyomydas peru-viensisbetweenDecember–April,andPlyomydasphalarossp.nov. inNovemberinArgentinaandJanuary–FebruaryinParaguay.

Biodiversityhotspots

WiththeexceptionofthesinglespecimenofPlyomydasphalaros sp.nov.fromParaguay,whichisdistributedintheAtlanticForest biodiversityhotspot,thegenusisnotconﬁnedtoanddoesnotoccur inanybiodiversityhotspotsensuConservationInternational.

Conﬂictsofinterest

Theauthorsdeclarenoconﬂictsofinterest.

Acknowledgments

Wewouldliketothankthemuseumcuratorswhomade speci-mensavailablethroughloansandfortheirhospitalitywhenvisiting thecollections.WarrenSteiner(NMNH,Washington,DC,USA)is thanked for bringing the newly collectedspecimen, now holo-type, of Plyomydasadelphe sp.nov. toour attentionand Donat Agosti(Plazi,Bern,Switzerland)forhelp withaddingthe origi-naltaxondescriptionstothePlaziTreatmentBank.WethankJulia Calhaufor insight intothemale terminaliaof Mydinaespecies. Wefurthermorethankthetwopeerreviewerswhoprovided con-structivecommentsthatenhancedthemanuscript.Wegratefully acknowledgesupportfromtheU.S.NationalScienceFoundation forfundingtheResearchExperienceforUndergraduatessitegrant calledNaturalHistoryResearchExperience(NHRE)(OCE1560088; PIE.Cottrell,Co-PIE.Hunt)throughwhichtheseniorauthor par-ticipatedin this researchproject. Special thanksto Gene Hunt,

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ElizabethCottrell,andVirginiaPowerfortheirsupportand admin-istrationoftheNHREProgram.Partofthisprojectwasfundedby aU.S.NationalScienceFoundationREVSYSaward(DEB0919333; PIT.Dikow,Co-PIDavidYeates).Anyopinions,ﬁndings,and con-clusionsor recommendations expressedin this manuscript are thoseoftheauthorsand donotnecessarily reﬂecttheviewsof theNationalScienceFoundation.

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