SystematicEntomology(2015),40,242–267 Long-proboscid brachyceran flies in Cretaceous amber (Diptera: Stratiomyomorpha: Zhangsolvidae) A N TO N I O A R I L L O1, E N R I QU E P EÑA LV E R2, R I C A R D O PÉ R E Z - D E L A F U E N T E3, X AV I E R D E L C L Ò S4, J U L I A C R I S C I O N E5, P H I L L I P M . BA R D E N5, M A R K L . R I C C I O6and DAV I D A . G R I M A L D I5 1DepartamentodeZoologíayAntropologíaFísica,FacultaddeBiología,UniversidadComplutense,Madrid,Spain,2Museo Geominero,InstitutoGeológicoyMinerodeEspaña,Madrid,Spain,3MuseumofComparativeZoology,HarvardUniversity, Cambridge,MA,U.S.A.,4Departamentd’Estratigrafia,PaleontologiaiGeociènciesMarines,FacultatdeGeologia,Universitatde Barcelona,Barcelona,Spain,5DivisionofInvertebrateZoology,AmericanMuseumofNaturalHistory,NewYork,NY,U.S.A.and 6InstituteofBiotechnology,CornellUniversity,Ithaca,NY,U.S.A. Abstract. The monophyletic family Zhangsolvidae comprises stout-bodied brachyc- eranflieswithalongproboscisandoccurringonlyintheCretaceous,originallyknown in shale from the Early Cretaceous Laiyang Formation (Fm.) in China (Zhangsolva Nagatomi & Yang), subsequently from limestones of the Early Cretaceous Crato Fm. of Brazil. Cratomyoides Wilkommen is synonymized with Cratomyia Mazzarolo & Amorim,bothfromtheCratoFm.;CratomyiidaeissynonymizedwithZhangsolvidae. TwogeneraandthreespeciesofZhangsolvidaearedescribed:Buccinatormyiamagnifica Arillo,Peñalver&Pérez-delaFuente,gen.etsp.n.andB.soplaensisArillo,Peñalver& Pérez-delaFuente,sp.n.,inAlbianamberfromLasPeñosasFm.inSpain;andLingua- tormyiateletactaGrimaldi,gen.etsp.n.,inUpperAlbian–LowerCenomanianamber fromHukawngValleyinMyanmar.BuccinatormyiasoplaensisandLinguatormyiatele- tacta are unique among all Brachycera, extant or extinct, by their remarkably long, flagellateantennae,about1.6×thebodylengthinthelatterspecies.Aphylogeneticanal- ysisof52morphologicalcharactersfor35taxaispresented,11taxabeingCretaceous species,whichsupportsplacementofthefamilywithinStratiomyomorpha,althoughnot toanyparticularfamilywithintheinfraorder. This published work has been registered in Zoobank, http://zoobank.org/urn:lsid: zoobank.org:pub:F32CF887-7C37-45D5-BD6B-135FE9B729A7. Introduction Wilkommen & Grimaldi, 2007). The genus Cratomyia was placed in the infraorder Stratiomyomorpha by Mazzarolo & The Zhangsolvidae is an extinct family of brachyceran flies Amorim (2000), a basal lineage of Brachycera that includes erectedbyNagatomi&Yang(1998)forZhangsolvacupressa threelivingfamilies:theStratiomyidaeLatreille,acosmopoli- (Zhang, Zhang & Li), preserved in shale from the Early tan group comprising 2651 species in 375 genera (Woodley, Cretaceous of China. The family was subsequently found to 2001); the Xylomyidae Verrall, a small cosmopolitan fam- includetwospeciesofCratomyiafromtheAptian-agedCrato ily comprising approximately 138 species in four genera Formation limestone of Brazil (Mazzarolo & Amorim, 2000; (Woodley, 2011); and the Pantophthalmidae Bigot, a small neotropical family of very large flies with 20 species in two Correspondence:RicardoPérez-delaFuente,MuseumofCompara- genera (Val, 1976). Several other late Mesozoic families have tiveZoology,HarvardUniversity,26OxfordSt.,Cambridge,MA02138, been suggested to be within this infraorder, but their rela- U.S.A.E-mail:[email protected] tionships remain poorly understood. These families include EremochaetidaeUssatchov,TethepomyiidaeGrimaldi&Arillo Theauthorsdeclarenoconflictofinterest. 242 ©2015TheRoyalEntomologicalSociety CretaceousbrachyceranfamilyZhangsolvidae 243 and, perhaps, Oligophrynidae Rohdendorf (Krzemin´ski & SieghardEllenberger,Germany,whoobtainedtheamberfrom Krzemin´ska, 2003; Grimaldi & Arillo, 2008), although the minesnearMyitkyina,KachinProvince,northernMyanmar(see focusofthispaperisonthepositionofZhangsolvidae. Grimaldietal.,2002).Theambersurfaceslyingoverthedor- In this paper we describe four new Cretaceous specimens salandventralportionsoftheflywereslightlyflattenedusing of Zhangsolvidae, three of them originally found in the same a lapidary wheel; there was no other preparation. The amber amber piece from the Spanish locality of El Soplao, plus one piece is occluded with a turbid suspension of particles, so in specimenfromBurmeseamber.TheElSoplaooutcrop,Albian ordertovisualizecriticaldetailsitwasCTscannedattheCornell inage,wasrecentlydiscoveredinSpain(Najarroetal.,2009, BiotechnologyResourceCenterImagingFacility. 2010; Peñalver & Delclòs, 2010). To date, insects belonging TheBurmesespecimenscanningwasperformedusingaZeiss to13ordersandabout40familiesarepreservedinthisamber VERSA XRM-520 instrument. For each dataset, 2400 X-Ray (Pérez-de la Fuente, 2012). In the order Diptera, the families ∘ ∘ projections were obtained at 0.15 intervals over 360 using Atelestidae, Cecidomyiidae, Ceratopogonidae, Chimeromyi- 60kV, 32μA and 5000ms exposure time, with a 2048×2048 idae, Chironomidae, Dolichopodidae, Hybotidae, Phoridae, pixeldetectorusinga1.5μmpixelsizethatwas2×2binnedinto PsychodidaeandRhagionidaehavebeenfound(ibid.),although aneffectivepixelsizeof3.0μm.Properplacementofthespeci- only ceratopogonids have been studied (Pérez-de la Fuente menwasaccomplishedwiththeassistanceofX-Rayfluoroscopy etal.,2011). mode,toensurethattheareaofinterestwasincludedwithinthe Burmese amber contains probably the most diverse palaeo- scanningfieldofview.Theprojectionviewswereusedtorecon- biotaofthesevenmajorworlddepositsofamberfromtheCre- structaCTimageusingaconvolutionback-projectionapproach taceousPeriod,withapproximately228familiesoforganisms implementedin3D,givinga3×3×3mmvolumeofimagedata (primarily arthropods) thus far reported, Diptera being espe- with3μmisotropicvoxelsinarbitrarydensityunits(ADU)and ciallydiverse(Zherikhin&Ross,2000;Grimaldietal.,2002; exported as DICOM or TIFF stacks. Reconstructed data files Ross etal., 2010). Burmese amber contains, for instance, the wereimportedintothesoftwareprogramsOsiriX(64bit,v5.5) oldest definitive Mesozoic record of mosquitoes (Culicidae) (Borkent & Grimaldi, 2004). Estimated age of ‘burmite’ was andAvizo8.0(VSG),wheretheywerevisualizedandanalysed recently established as the Albian–Cenomanian boundary, c. usingavarietyof2Dand3Dtechniques. 99Ma,basedonradiometricdatingofzirconsfromtheamber Fly morphological terminology generally follows McAlpine matrix(Shietal.,2012). (1981).Wedonotusethetermpostpedicel,usedbysomedipter- The new zhangsolvids are the first specimens preserved as iststorefertothatportionoftheantennadistaltothepedicel, amber inclusions and the youngest occurrences in the fossil because by definition this is the flagellum. Szucsich & Krenn recordofthefamily,significantlyexpandingitspalaeobiogeo- (2000, 2002) introduced the term ventral rostral membrane, graphicrange.Theexquisitepreservationinamberallowsobser- whichweusehere,referringtothepleatedmembraneatthebase vationofminutemorphologicaldetails,providingsubstantially oftheproboscis. more data to test the proposed relationships of Zhangsolvidae Although the three Spanish amber specimens are syninclu- withintheStratiomyomorpha.Thesefindingsalsoprovidesig- sions, the gender of only one of them is definitive. Because nificantpalaeoecologicaldata,butthesearepresentedelsewhere anotherspecimenissubstantiallylargerbutlacksterminalia(and (E.Peñalveretal.,inpreparation). hassignificantdifferencesinvenationandantennalstructure), thelengthsofthethoraxandwingin34exemplarextantspecies ofStratiomyidae,XylomyidaeandPantophthalmidaeweremea- Materialandmethods suredforatleastonemaleandonefemale,totestforthepres- ence/extentofsexualdimorphisminbodysize.Weusedpinned The Spanish specimens were discovered in a large piece of specimensfromtheAMNHcollectionforthesemeasurements. ambercontainingnumerousinclusionsfromtheElSoplaoout- Forthefirsttwoofthesefamiliesthemeasurementsweremade crop, near the municipality of Rábago (Autonomous Commu- using a Nikon SMZ1500 stereomicroscope and NIS Elements nityofCantabria).Thispiecewastrimmedintoseveralsmaller software;measurementsofPantophthalmidaeweremadeusing pieces to better allow examination of individual inclusions. amillimetrerulerbecauseoftheirlargesizes. Each of these smaller pieces was embedded in synthetic resin Phylogenetic analysis used morphological characters from (EPO-TEK301)andthenpolished(seeNascimbene&Silver- exemplar extant and fossil stratiomyomorphans, as well as stein, 2000). Fossils were examined with an Olympus BX51 all species of Zhangsolvidae, analysed under equal-weights compoundmicroscope(OlympusOpticalCo.,Ltd.,Japan).Pho- tomicrographs were taken using an attached camera, some of parsimony with the program TNT (Goloboff etal., 2008). themz-stacked.DrawingsweremadeusinganOlympusU-DA Parametersemployedan‘XMULT’search,whichusessectorial drawingtubeattachedtothecompoundmicroscope.TheSpan- searching, tree drifting, ratcheting and fusing. The search ran ishspecimensareprovisionallyhousedattheMuseoGeominero until the optimal tree score was found 100× independently, (IGME)inMadrid(Spain);theirfinaldepositwillbeinthecol- whereupon all lowest-cost topologies were consolidated into lectionoftheElSoplaoCave(GovernmentofCantabria).The a strict consensus. Consistency and retention indices were BurmeseamberspecimenisintheAmericanMuseumofNat- calculated, along with Bremer and bootstrap support values ural History, New York, and was purchased in 2012 from Mr. (with10000pseudoreplicates). ©2015TheRoyalEntomologicalSociety,SystematicEntomology,40,242–267 244 A.Arilloetal. Systematicpalaeontology were: (i) long basal section of Rs, (ii) antennal flagellum 12-segmented,(iii)proboscisconspicuouslylongand(iv)costal OrderDipteraLinnaeus,1758 cellconcavebeforeRs,possiblyrepresentingasc-rcrossvein. InfraorderStratiomyomorphaHennig,1973 The holotype was not studied by Nagatomi & Yang, so they assumed that the original illustrations were accurately drawn. Unfortunately,theholotypeofZ.cupressa(auniquespecimen) FamilyZhangsolvidaeNagatomi&Yang,1998 is currently lost (Dr H.-C. Zhang, personal communication to CratomyiidaeMazzarolo&Amorim,2000:94.Typegenusand A.A.,2011),whichiswhywewereunabletorestudytheholo- species:CratomyiamacrorrhynchaMazzarolo&Amorim(by type.AccordingtotheICZNcode,dramatictaxonomicchanges originaldesignation).NEWSYNONYMY. wouldbenecessaryforthedesignationofanewtypespecimen ofthefamily(fromadifferenttypegenus),whichwehavenot Type genus and species. Zhangsolva Nagatomi & Yang, madeincasethelostspecimeneventuallyappearsinaChinese 1998.HolotypeofthetypespeciesArchisolvacupressaZhang, collection,oranewspecimen(andpotentialneotype)isexca- Zhang&Li,1993,iscurrentlylost(DrH.-C.Zhang,personal vatedfromtheLaiyangFormation. communicationtoA.A.,2011).Archisolvawaspreoccupiedin Later, Mazzarolo & Amorim (2000) described Cratomyia Stratiomyidae,soNagatomi&Yang1998providedthereplace- macrorrhyncha, from the Early Cretaceous Crato Formation, mentgenericnameZhangsolva. Brazil,forwhichtheyproposedthefamilyCratomyiidae.Their studyrevealedthatCratomyiawasasistergroupto(Xylomyi- Taxonomic composition. Zhangsolva cupressa (Zhang, dae+Stratiomyidae), but these authors had not taken into Zhang & Li, 1993); Cratomyia macrorrhyncha Mazzarolo & accountZhangsolvacupressa.Themostrecentcontributionon Amorim, 2000; Cratomyia cretacica Wilkommen, comb.n.; thisgroupofflieswasthedescriptionofCratomyoidescretaci- Buccinatormyia magnifica Arillo, Peñalver & Pérez-de la cusWilkommen(inWilkommen&Grimaldi,2007),afossilfly Fuente, gen. et sp.n.; Buccinatormyia soplaensis Arillo, alsofoundintheCratoFormationandverysimilartoCratomyia. Peñalver&Pérez-delaFuente,sp.n.;Linguatormyiateletacta Inthatpapertheauthorsrecognizedthecloserelationshipamong Grimaldi,gen.etsp.n. Zhangsolva,CratomyiaandCratomyoides.Recently,Grimaldi etal.(2011)suggestedthatallofthesefossilsshouldbeincluded inthefamilyZhangsolvidae(withCratomyiidaeasajuniorsyn- Diagnosis (emended). Proboscis rather long to very long onym).TheyalsosuggestedthatCratomyoidesisajuniorsyn- (from1toabout4×theheadlength),slender,apexwithsmall onymofCratomyia,buttheydidnotformallysynonymizethese labellum; veinCendingatornearapex ofwing,M strongly 1 taxa. In the same work the authors also indicated that several arched, M fused to CuA and CuA fused to A near wing 3 1 2 1 featuresintheoriginaldescriptionofZhangsolvawereprobably margin. erroneous:(i)partsofthewingvenationareobviousartefacts, suchasasc-rcrossvein,whichispresentinonlyonewing(as Remarks. We have refrained from including the following seen in the photograph of the holotype in the original paper), charactersinthefamilydiagnosisabove,astheyarenotknown and(ii)theantennaeprobablyhavenomorethaneightflagel- fromallthezhangsolvidgenera (onlyfromhalforlessofthe lomeres(thegroundplaninBrachycera),amisinterpretationper- species,mainlytheonespreservedinamber),butwepositthat hapsoriginatingfrompreservation(e.g.transversecracksincar- theycouldbeofimportancetocharacterizethefamily:(i)maxil- bonaceousfilmscommonlyoccurincompression-fossilinsects larypalptwo-segmented,longandprojecting(observedinBuc- preserved in rock). Furthermore, the long basal section of Rs cinatormyiamagnificagen.etsp.n.andLinguatormyiateletacta proposedasdiagnosticforZhangsolvidaeforNagatomi&Yang gen.etsp.n.,tentativelyobservedinCratomyiamacrorrhyncha, (1998)isquiteshortinthenewBurmeseamberspecies.Hence, seeMazzarolo&Amorim,2000),(ii)tibialspurs0,2,2(only theconspicuouslylongproboscisistheonlycharacterthatwe observedinB.magnifica),and(iii)pulvilliandempodiumwell haveretainedherefromtheoriginaldiagnosisofthefamily. developed,pulvilliform(idem).Somezhangsolvids,i.e.Bucci- natormyia soplaensis sp.n. and L.teletacta, are readily recog- GenusCratomyiaMazzarolo&Amorim nizablebyshowingremarkableflagellateantennaethatcanbe Cratomyia Mazzarolo & Amorim, 2000: 94. Type species: morethan1.5×longerthanthebody(inthelatterspecies). C.macrorrhyncha Mazzarolo & Amorim. By original The species Archisolva cupressa was described by Zhang designation. etal. (1993) from the Early Cretaceous Laiyang Formation CratomyoidesWilkommen,2007,inWilkommen&Grimaldi, in Laiyang, Shandong Province, China, originally placed in 2007:375.Typespecies:CratomyoidescretacicusWilkommen. the family Solvidae (=Xylomyidae). Later, Nagatomi & Yang Byoriginaldesignation.NEWSYNONYMY. (1998) proposed a replacement name, as the generic name was already occupied by Archisolva Enderlein (Stratiomyi- dae). They provided the new name Zhangsolva and consid- Diagnosis (emended). Cratomyia is readily separated from eredthat,eventhoughthegenuswasclosetoStratiomyoidea, other(Laurasian)generaofZhangsolvidaebytheuniquepres- it represented a new family. Diagnostic characters of the new enceintheformerofastemtoveinsM andM distaltocelld 1 2 family Zhangsolvidae proposed by Nagatomi & Yang (1998) (insteadofdivergingdirectlyfromcelld). ©2015TheRoyalEntomologicalSociety,SystematicEntomology,40,242–267 CretaceousbrachyceranfamilyZhangsolvidae 245 GenusBuccinatormyiaArillo,Peñalver&Pérez-dela levelofr-mcrossvein,cellm narrow(W/L=0.21),M andM 3 2 3 Fuente,gen.n nearlyparallel,CuPwellsclerotized. http://zoobank.org/urn:lsid:zoobank.org:act:D45D7DD7- 89E2-450F-BA5E-46F92F66611A Description(♀;basedlargelyontheholotypeexceptforwing features entirely based on the paratype). Body 9.20mm long Typespecies. Buccinatormyiamagnificasp.n. (estimatedfromreconstruction). Head:Broad,1.82mmwide.Eyeslarge,bulging,occupying Etymology. Derived from buccinator- (L.), trumpeter, and nearlyentirelateralsurfaceofhead(genaeexposedventrally), -myia(Gr.),fly,inreferencetothelongproboscis. widely separated, ventrally 0.73mm wide between inner mar- gins; bare, devoid of interfacetal setulae; facets undifferenti- ated;innermarginsinfrontalviewwithoutemarginationaround Diagnosis. Antenna with six articles (four flagellomeres), antennalbase.Eyessignificantlybroaderdorsally.Dense,long, withminuteapicalstylus;stemofRslong(severaltimesther-m fine,stiffsetaearoundeyemargin.Threesmallocellipresent, crossveinlength),CveinendingbeyondR butwithoutreaching 5 lying in a clearly demarcated, densely setose ocellar triangle. M ,veinsM andM divergingdirectlyfromcelld(notfroma 1 1 2 Vertexofheadwithparamedianemarginations,centrallyraised portionofM distadtocelld). 1+2 intomediandorsaltubercle;vertexwiththreerangesofpilos- ity:(i)vestitureoffine,densesetulaeasontherestofthehead, BuccinatormyiamagnificaArillo,Peñalver&Pérez-dela (ii)sparse,longersetae,and(iii)long,fine,stiffsetaelikethose Fuente,sp.n aroundeyemargin.Areassurroundingantennalbasesraisedinto http://zoobank.org/urn:lsid:zoobank.org:act:21A4E7D7-D8F6- bare, rounded mounds (=facial lobes), nearly meeting medi- 4FBB-9736-73F5F80F6066 ally.Frontofheadwithwell-developed,invertedY-shapedsul- (Figs1–8) cus. Clypeus large, bulging, covered by fine, dense setulae, raised into low mound between facial lobes and palpal bases, Type material. Holotype CES-349.1 and paratype in two fully frontal in position; with short vertical and median sulci parts, CES-015.1 and CES-392.3, from El Soplao amber andatransverse,mediandepression;ventralsurfaceofclypeus (Spain). These specimens and holotype of B. soplaensis sp.n. membranous,oval;membraneconnectedtobaseofproboscis. were found in the same amber piece as syninclusions, which Frontoclypealsulci(demarcatinglateraledgesofclypeus)very was prepared in three sample pieces for study (CES-349: deep.Genaedevelopedintopairofrounded,ventrallobes.Neck amberfragment10×5×5mminaresinprism16×8×5mm; entirelymembranous. CES-015:amberfragment13×11×6mminaresintrapezoid Proboscis: Long (virtually all of it haustellum), 3.85mm 15×19×6mm;andCES-392:amberfragment24×10×7mm long from base to apex (0.42× the body length), 0.05mm in a resin prism 30×11×8mm). Holotype is in a good state wide, 0.17mm high at middle. Proboscis laterally flattened, of preservation, virtually complete, but wings are incomplete 0.05mm wide, 0.17mm high at the middle (width 0.31× the and a dorsal portion of the thorax is slightly altered. Also, cross-sectionaldepth)anddirectedforwardinrelationtohead. proboscis,legsandabdomenarecrossedbyafracturethatpar- Proboscisinsertedbelowfacialmargin.Baseofproboscisven- tiallydestroyedsomesmallportionsoftheabdomen.Paratype trally with annulated membrane (ventral rostral membrane), is strongly compressed but virtually complete, lacking only five-foldsvisible,eachwithnumerousannulations(membrane thedistalportionsoftheproboscisandtheabdomen;itliesin probablytelescoped).Probosciscoveredbydensemicrotrichia two prepared sample pieces, the smaller portion (CES-392.3) andadensevestitureoflarge,stiffdeeply-ribbedsetaewithstel- consistingofacostalfragmentofwingwiththehalter,several latetransversesectionanddirectedforward.Haustellumexter- legportionsandasmallportionoftheabdominalcuticle. nally composed by a tube-like structure with a dorsal groove (=theca), subellipsoidal in cross-section and with straight lat- eral walls (well preserved in holotype; laterally collapsed Syninclusions. Holotype and paratype of the new species in paratype, ventrally forming an artifactual surface). Dorsal andtheholotypeofB.soplaensissp.n.werepreservedinone grooveinternallyclosedbyanarrow(0.05mmwide),semitrans- largepieceofamber,whichwasdividedintoseveralpiecesto parentmembranewiththickenedmarginsandcovereddorsally allowstudyofthe48arthropodsinit,includingtenadditional by a fine vestiture of minute setae, interpreted as a modified dipterans,mainlysmallbrachycerans. labrum (most visible in the basal half of the holotype’s pro- boscis,partiallybrokenandprotrudingduetodownwardbend- Etymology. Namedafterthewonderfulstateofpreservation ingoftheproboscis).Labrumexternallycoveringtheproboscis andtheimpressivehabitusofthetwospecimens. baseandbasallyenteringintotheproboscis.Epi/hypopharynx not visible, but most likely forming floor of proboscis. Food Diagnosis. Proboscis very elongate (about 0.40× the body canalpreserved,0.03mmwideand0.07mmhighaspreserved) length); antenna not flagellate and short (about 0.30× the (visibleintheambersurfacethatcross-sectionstheparatype’s wing length), flagellomere II with dorsal surface concave, proboscis). Labellum very small, about as wide as proboscis, flagellomereIIIabout0.30×theantennallength;Scendingat oval in cross-section. Labellar lobes closely pressed together ©2015TheRoyalEntomologicalSociety,SystematicEntomology,40,242–267 246 A.Arilloetal. Fig.1. Buccinatormyiamagnificagen.etsp.n.,fromElSoplaoamber.EntireholotypeCES-349.1(ventro-obliqueview),shownaspreserved. medially,0.22mmlong,0.13mmhigh;eachlobehasaround13 basally and gradually narrowing distally, 0.15mm high at the pseudotracheaewithtransversepseudotrachealringsonexposed base (greatest height), 0.06mm high at apex (lowest height); surfaces. distal2/3ofitslengthlaterallyflattened,0.11mmwideatthe Maxillary palp: Elongate, 1.45mm long; projected forward; base(greatestwidth).Basalpalpomerewithabundant,verylong, ∘ two-segmented. Bases of palps flank base of proboscis. Basal fine, stiff setae along its length, oriented at 90 -angle to the palpomerestraightandlong,0.60×thetotalpalplength;inflated antennal axis. Apical palpomere long, club-like, pedunculate, ©2015TheRoyalEntomologicalSociety,SystematicEntomology,40,242–267 CretaceousbrachyceranfamilyZhangsolvidae 247 comprisingdense,smallfoveae(probablysensillamuchsmaller thanotherfoveaeonantennaandpalp).FlagellomereIVacumi- natedistally.Antennawithminuteapicalstylus. Thorax: Narrow and deep as preserved, 2.34mm long, 1.20mm wide (estimated), 1.88mm high, slightly narrower than head (thoracic cuticle of holotype partially lost, mainly anterodorsally; thorax of paratype slightly collapsed, pleuron not well seen). Thorax devoid of macrosetae, exocuticle with fine,shortsetaeonly;colorationappearsuniformlydarkbrown- ish (possibly preservational). Anterior part of thorax obscure. Mesothorax large, expansive. Katepisternum ventrally broad andslightlybulging.Anepisternumlarge;anepimeronsmaller, subdivided by partial cleft; mesothoracic spiracle not visible; meronslightlylargerthanmesocoxa.Coxaandventralsurfaces of femur with sparse, short setulae. Procoxa slender, 1.02mm long;mesocoxa0.64mmlong,0.23mmwidebasally;metacoxa 0.60mm long. Anterodistal surfaces of each coxa with small, protruding lobe. Trochanter small, slender, fully articulated with femur and coxa (no fusion). Legs slender. Prothoracic leg about as long as other legs: femur 1.58mm long, tibia 1.63mm long and tarsomere I 2.38mm long. Mesothoracic leg 5.85mm long, as estimated: femur 1.70mm long as esti- mated,tibia1.95mmlong,tarsomere(ta)I1.22mmlong,taII 0.45mmlong,taIIImm0.23long,taIV0.13mmlongandtaV 0.18mmlong.Metathoracicleg6.83mmlong:femur2.03mm long, tibia 2.50mm long, tarsomere (ta) I 1.30mm long, taII 0.48mmlong,taIII0.25mmlong,taIV0.13mmlongandtaV 0.15mm long. Femora with recumbent, dense setulae. Tibiae withdenser,strongersetae.Protibiawithoutspurs.Onepairof Fig.2. Buccinatormyiamagnificagen.etsp.n.,fromElSoplaoamber. setose distal spurs each on meso- and metatibia. Tarsomeres Photomicrographoftheholotype,CES-349.1. covered with abundant, short, strong setae. Tarsi more setose thantibiae,especiallystrongsetaedistallyandeachtarsomere strongly laterally flattened, very broad distally, apex rounded; having several distal, spine-shaped setae. Claws covered by with several fine foveae (probably sensilla) distributed mainly microtrichia. Pulvilli paddle-shaped; flat, paddle-shaped (pul- distallybutwithoutapparentpattern. villiform)empodiaonallpretarsi,ofsimilarsizetopulvilli;all Antenna:Six-articled(flagellumwithfourflagellomeres),pro- threepadscoveredbymicrotrichia. jectingforward;1.85mmlong(0.20×estimatedbodylength). Wing(basedonparatype,includinghalter):Length6.00mm, Antennal bases narrowly separated by distance subequal to 2.09mm wide (greatest width) (W/L=0.35). Costal vein diameter of antennal socket (c. 0.80× distance between palp ending between R and M . Veins Sc and R fused basally to 5 1 bases). Scape short, c. twice as long as wide, 0.18mm long, crossvein h; Sc 2.57mm long (0.43× the wing length). Vein 0.10mm wide, 0.14mm high; both scapes oriented antero- Scmeetswingmarginbeyondmiddleofwinglength,atlevel laterally (divergent) as preserved. Scape and pedicel covered of vein r-m. Vein R 3.15mm long, gradually thickened api- 1 by sparse, relatively strong setae projected forward. Antenna cally; stem of Rs relatively long, 0.89mm long (ca. 4.5× the entirely covered by a vestiture of dense, fine setulae. Pedicel length of r-m crossvein); R 1.62mm long; stem of R 2+3 4+5 short,0.75×thescapelength,aswideasscape.Flagellumrel- 1.51mm long, R and R short, asymmetrical, R 0.57mm 4 5 4 ativelyshort,laterallyflattened,comprisingfourflagellomeres: long, R 1.05mm long; tip of R preapical to wing tip. Cell 5 5 I 0.27mm long, 0.06mm wide, 0.09mm high, approximately d short and thick, 1.03mm long (greatest length), 0.40mm as long as scape+pedicel; II 0.56mm long, 0.05mm wide, wide (W/L=0.39). Stem of M 0.86mm long; M arched 1+2 1 0.09mmhigh(greatestheight);IIIaslongasflagellomereIIbut costad, tip of M ending posterior to wing tip; M 1.18mm 1 1 withdifferentmorphology,0.53mmlong,0.12mmwide(great- long (linear, not along curvature), M 0.74mm long; M and 2 1 estwidth),0.03mmhigh;IVshort,aslongasscape,0.18mm M diverge directly from cell d, not from a portion of M 2 1+2 long,0.03mmwide,0.06mmhigh(greatestheight).Flagellom- distad to cell d; veins M and M nearly parallel. M ends 2 3 2 ereIIwithdorsalsurfaceconcave;IIandIIIinsertedventrally before wing margin. Cell m long and slender, 1.02mm long, 3 totheirproximalflagellomere.FlagellomereIIIproximallywith 0.21mm wide (W/L=0.21); veins CuA and M joined in 1 3 afewfinefoveae(probablysensilla),similartothoseondistal very short vein before meeting wing margin. Cell br slightly palpomere,distributionwithoutapparentpattern;thisflagellom- narrower than bm; cell bm slightly longer (2.72mm long, vs. erewithnarrowbandonventrolateralsurface(exceptonapex) 2.54mm).VeinCuAwithbaseapproximatelyhalfthethickness ©2015TheRoyalEntomologicalSociety,SystematicEntomology,40,242–267 248 A.Arilloetal. Fig.3. Spatial relation of the paratype of Buccinatormyia magnifica gen. et sp.n., the holotype of B.soplaensis sp.n., and some other close syninclusionsintegratingdrawingsfromtwopreparationsofthesameamberpiece(CES-015andCES-392).(1)Buccinatormyiasoplaensispartially reconstructed;(2)Buccinatormyiamagnifica;(3–4)Diptera:Brachycera;(5)Neuroptera:Coniopterygidae;(6)Lepidoptera:Micropterigidae;(7)Acari: Actinedida. of base of R; CuA fusing with A prior to wing margin, thorax; 5.30mm long (without cerci); strongly flattened 2 1 forming closed, narrow cell cup; cup 2.97mm long dorsoventrally, especially on margins. Abdomen compris- (W/L=0.12); CuP well developed and sclerotized, parallel ing eight visible segments; dorsal surface (tergites) flat, to CuA and separated by distance equal to vein thickness. virtually without relief. Coloration appears uniformly dark Vein CuA strongly curved before connection with A . Vein brownish(possiblypreservational).Sternitescoverentiretrans- 2 1 A entirely straight; A preserved, basal sclerotized portion verse width of abdomen; no abdominal pleural membrane 1 2 0.95mm long. Apices of veins CuA +M and CuA +A exposed. Segments V–VIII laterally form very thin planar 1 3 2 1 very thin, faint. Anal lobe, alula and calypter well developed; expansions (=‘lateral flanges’ sensu Stuckenberg (2000) for calypter 0.52mm long, with marginal vein. Wing membrane Vermileonidae), but very small on segment VIII; segment entirelyhyaline(nopatterningorpterostigmaticsclerotization), IV apparently with a small distal section as lateral flanges. evenlycoveredwithminutemicrotrichia.Halter1.00mmlong, Abdominal cuticle showing a vestiture constituted by two with slender apical knob 0.41mm long, 0.21mm wide. Stem different types of specialized setae: (i) ‘stellate hairs’, cor- and knob covered by dense microtrichia and knob with a few responding to microtrichia arranged in bunches of c. five, additional longer setae. Halter with sensory area at least in present on dorsal surface of lateral flanges and also laterally middle of its stem, having campaniform-like sensilla; pilosity at least on segment III, and (ii) large, stiff deeply-ribbed ofbaseobscure. setae (stellate in transverse section), similar to those on pro- Abdomen (virtually complete in holotype): Broad, broader boscis, sparsely covering the entire abdominal cuticle. Only distally, 1.45mm wide (greatest width), slightly broader than larger lateral flanges (segments V–VII) have dorso-marginal, ©2015TheRoyalEntomologicalSociety,SystematicEntomology,40,242–267 CretaceousbrachyceranfamilyZhangsolvidae 249 Diagnosis. Probosciselongate(about0.30×thebodylength, estimated); antenna flagellate and elongate (about 0.60× the winglength),flagellomereIIwithdorsalsurfacestraight,flag- ellomereIIIabout0.65×theantennallength;Scendingbeyond levelofr-mcrossvein,cellm broad(W/L=0.34),M andM 3 2 3 highlydivergent,CuPweaklydeveloped. Description (sex unknown). Body about 14mm long (esti- mate,basedoncomparisonofwinglengthsbetweenthisspecies andB.magnifica). Head: Incomplete (only a portion of eye preserved as an impression, showing that eyes are large in this species, though insufficiently preserved to determine if holoptic or dichoptic). Proboscis:Incomplete,preservedportion1.07mmlong(prox- imal end obliquely sectioned by surface of the preparation, estimatedtohaveoriginallybeensimilarorslightlygreaterin length than that of B.magnifica). Proboscis laterally flattened, 0.22mm high, directed forward in relation to head. Proboscis covered by dense microtrichia and a dense vestiture of large, stoutsetaewithdeeplongitudinalribbing(stellateintransverse section)anddirectedforward.Haustellumexternallycomposed by a tube-like structure with a dorsal groove (=theca), subel- lipsoidalincross-sectionandwithstraightlateralsides.Labrum Fig.4. PhotomicrographoftheparatypeofBuccinatormyiamagnifica narrow,0.06mmwide,withthickenedmargins(visiblebycutic- gen. et sp.n. and the wing of the holotype of B.soplaensis sp.n., ulartransparencyinadorsal,slightlyobliquepositionandalso CES-015. asanobliquesectioncreatedbythesurfaceoftheamberprepa- ration).Foodcanalnotvisible.Labellumverysmall,narrower longitudinal,curvedlinescomprisingminuteellipsoidplaques than proboscis (preserved in fine detail, located very close to of variable size (=abdominal muscle plaques, see Stoffolano preparationsurface,thusobservableunderhighmagnification). etal., 1988). Female terminalia preserved: cercus strongly Labellarlobescushion-like,0.26mmlong,c.0.16mmwide,c. flatteneddorsoventrally,two-segmented,coveredbyfinesetae; 0.08mmhigh;eachlobehas10–12pseudotracheaewith30–40 basal segment 0.21mm long, 0.08mm wide; distal segment transversepseudotrachealrings(onelabellarlobewellexposed, 0.15mmlong,0.04mmwide. slightly oblique; other in perpendicular position, obscured to observation).Labellarlobeswithsparse,strongsetaeirregularly BuccinatormyiasoplaensisArillo,Peñalver&Pérez-dela distributed,asfigured. Fuente,sp.n. Antenna: Six-articled (flagellum with four flagellomeres), http://zoobank.org/urn:lsid:zoobank.org:act:C7BDF2D4-F83E- very elongate and flagellate, projecting forward, with same 4BC4-AAC0-F7562F656A4A configurationasinB.magnificasp.n.butwithtwoflagellom- (Figs3,4,8,9) eres elongated (c. 2× for flagellomere II and c. 7× for flag- ellomere III). Scapes not preserved, only apices of pedicels preserved; scape+pedicel of same length as in B.magnifica Type material. Holotype in two fragments CES-015.2 and (inferred by the close position of the eye remains); antenna CES-392.2,fromElSoplaoamber(Spain).Specimenpreserved 5.62mm long (estimated from reconstruction). Preserved por- asacompletewingwithasmallportionofthethoraciccuticle tionofpedicelcoveredbylongsetaeprojectingforward.Flag- (CES-015.2) and distal parts of the antennae and proboscis ellum covered by vestiture of dense, fine setulae. Flagellum includingtracesoftheeyes(CES-392.2). long, laterally flattened, comprising four articles: flagellom- ereI0.34mmlong,0.03mmwide,0.12mmhigh;II1.18mm Syninclusions. The holotype of the new species and the long,0.03mmwide,0.17mmhigh(greatestheight);III3.03× holotypeandparatypeofB.magnificasp.n.werepreservedin the length of flagellomere II but with similar morphology, onelargepieceofamber,whichwasdividedintoseveralpieces 3.57mm long, 0.02mm wide, 0.10mm high basally, 0.04mm toallowstudyofthe48arthropodsinit,includingtenadditional highdistally,0.64×theantennallength;IVshort,0.19mmlong, dipterans,mainlysmallbrachycerans. 0.02mmwide,0.04mmhigh.FlagellomereIIwithdorsalsur- facestraight.FlagellomereIIIwithabundant,fine,subcircular Etymology. NamedafterElSoplao,theamberlocalitywhere foveae(probablysensilla)c.12μmwide(greatestwidth),iden- theholotypewasfound. tical to those on same flagellomere in B.magnifica, occurring ©2015TheRoyalEntomologicalSociety,SystematicEntomology,40,242–267 250 A.Arilloetal. Fig.5. Anatomicalfeaturesoftheholotype(CES-349.1)ofBuccinatormyiamagnificagen.etsp.n.fromElSoplaoamber.(A)Headindorsalview aspreserved;(B)reconstructedlabellumindorsal(left)andlateral(right)views;(C)reconstructedpalpindorsal(top)andlateral(bottom)views;(D) thoraxinlateralview. on the proximal third but without apparent pattern; this flag- Thorax:Onlyasmallportionofthedorsalcuticleispreserved, ellomere with a narrow ventrolateral band comprising dense, havinglongsetae(suchsetaeabsentinB.magnifica). abundant, minute, elongate foveae (7μm long) (also probably Wing:Incomplete[smallbasalportionandpartofanallobe sensilla,butabouthalfthesizeofotherfoveae).Flagellomeres notpreserved(cellcupincomplete)].Length9.07mm,3.10mm II and III broadly articulated with proximal article, especially wide(estimatedgreatestwidth)(W/L=0.34).Costalveinend- flagellomere II, indicating significant lateral mobility. Flagel- ingbetweenR andM .VeinsScandRfusedbasaltocrossvein 5 1 lomeres III and IV acuminate distally, but III only on ventral h; Sc 4.46mm long (0.49× the wing length). Vein Sc meets margin. Minute apical stylus present, slightly longer than in wing margin beyond middle of wing length, beyond level of B.magnifica. r-m crossvein. Vein R 5.26mm long; stem of Rs relatively 1 ©2015TheRoyalEntomologicalSociety,SystematicEntomology,40,242–267 CretaceousbrachyceranfamilyZhangsolvidae 251 veins CuA +M and CuA +A not preserved. Cell br nar- 1 3 2 1 rowerthanbm;cellbmslightlylonger,4.20mmlong(estimated) vs.4.04mmanddistallybroaderduetoveinMarchedcostad. Vein CuA with base approximately half the thickness of base of R; CuP weakly developed, not sclerotized, parallel to CuA andseparatedbydistanceequaltoveinthickness.VeinsA and 1 A notpreserved.Alulaandcalypternotpreserved.Wingmem- 2 brane entirely hyaline (no patterning or pterostigmatic sclero- tization),evenlycoveredwithminutemicrotrichia.Haltersnot preserved. LinguatormyiaGrimaldi,gen.n. http://zoobank.org/urn:lsid:zoobank.org:act:B279141E-9204- 4B56-9504-16E7F43D7663 Typespecies. Linguatormyiateletactagen.etsp.n. Etymology. Derived from lingua- (L.), tongue, and -myia (Gr.),fly,inreferencetothelongproboscis. Diagnosis. Antenna with five articles (three flagellomeres), withoutminuteapicalstylus;stemofRsshort(shorterthanr-m crossvein),M andM divergingdirectlyfromcelld(notfrom 1 2 aportionofM distadtocelld),veinA absent. 1+2 2 LinguatormyiateletactaGrimaldi,sp.n. http://zoobank.org/urn:lsid:zoobank.org:act:8A386070-4F23- 45F9-A6A6-DD527B6A55BC (Figs10–13) Type material. Holotype AMNH Bu-SE13, in Burmese amber(Myanmar).Thespecimenisina36×30×11mmpiece ofdark,turbidamber containing fineparticulate debris,trans- versefracturesandflowlines(especiallynearthecentre).These obscuresomefeaturesofthespecimen,especiallydorsally.The entirehead,palp,proboscisandevenantennaearepreserved,as Fig.6. ReconstructionofBuccinatormyiamagnificagen.etsp.n.in isthethoraxandmostoftheabdomen(exceptforitstip).Lost dorsalandlateralviewstoshowitslifeaspectbasedontheholotypeand atthesurfaceoftheamberaretipoftherightwingandapices paratype.Thecolourpatterningisconjecturalbutbasedonthecommon ofalltibiaeandalltarsi.Distalthirdofrightwingisfolded3× black-and-yellow patterns found in Xylomyidae and Stratiomyidae (Artist:J.A.Peñas). andrequiredcarefulreconstruction. Syninclusions. Two thrips (Thysanoptera), a small beetle, long,1.55mmlong(c.5.5×thelengthofr-mcrossvein);R 2+3 an empidoid fly, a portion of a neuropteran wing, and insect 2.85mm long; stem of R 2.33mm long, R and R short 4+5 4 5 frass. but relatively longer in comparison to B.magnifica, asymmet- rical, R 1.19mm long, R 1.55mm long; tip of R preapical 4 5 5 to wing tip. Wing tip less acute than in B.magnifica. Cell d Etymology. Combination of tele (G.), far, and tactus (L.), sense of touch, meaning ‘far touching’, in reference to the shortandthick,1.50mmlong(greatestlength),0.70mmwide strikinglyelongateantennae. (W/L=0.47).StemofM 1.17mmlong;M archedcostad, 1+2 1 tipofM endingposteriortowingtip;M 2.02mmlong(linear, 1 1 not along curvature), M 1.42mm long; M and M virtually Diagnosis. As for genus, with the following additions: 2 1 2 diverge directly from cell d, not from a portion of M dis- proboscis very long (about 0.45× the body length); antenna 1+2 tadtocelld;veinsM andM highlydivergent.VeinM ends flagellate and extremely elongate (about 1.80× the wing 2 3 2 beforewingmargin.Cellm longandbroad(smalldistalportion length), flagellomere III about 0.80× the entire antennal 3 lost), 1.66mm long (estimated), 0.57mm wide (W/L=0.34); length. ©2015TheRoyalEntomologicalSociety,SystematicEntomology,40,242–267