©EntomologicaFennica. 7March2008 Archiconiocompsa prisca Enderlein (Neuroptera: Coniopterygidae): The first neuropteran fossil in Rovno amber (Ukraine) JanuszKupryjanowicz&VladimirN.Makarkin Kupryjanowicz,J.&Makarkin,V.N.2008:ArchiconiocompsapriscaEnderlein (Neuroptera: Coniopterygidae): The first neuropteran fossil in Rovno amber (Ukraine). —Entomol. Fennica 19: 25—31. ArchiconiocompsapriscaEnderlein,previouslyknownfromBalticamber,isre- corded in Rovno amber. A discussion ofthe systematic position ofthis mono- typic genus is provided; it is tentatively assigned to Coniocompsini (Aleuro- pteryginae). J.Kupryjanowicz,InstituteofBiology, UniversityofBialystok, Swierkowa20B, 15—950Bialystok, andMuseumoftheEarthPolishAcademyofSciences,Al.Na Sharpie26/27, 00—488 Warszawa, Poland;E—mail.‘ [email protected] V. N. Makarkin (correspondingauthor), Institute ofBiologyandSoilSciences, FarEasternBranch oftheRussianAcademyofSciences, Vladivostok, 690022, Russia;E—mail.‘ [email protected] Received22March2007, accepted19May2007 1. Introduction Eocene in the age (Engel & Perkovsky 2006a, Perkovskyetal. 2007). Amber andits insect inclusions fromthe Rovno Inthis paper, we reportthe firstneuropteran RegionofUkraineiscurrentlyattractingmuchat- from Rovno amber, the coniopterygid species tention, andithasrecentlybeenintensivelystud- Archiconiocompsaprisca Enderlein, previously ied (e.g., Dlussky 2002, Dlussky & Perkovsky known from Baltic amber, and discuss the sys- 2002, Simutnik 2002, Kononova 2003, Perkov- tematicpositionofthegenus. sky et al. 2003a,b, 2007, Putshkov & Popov The family Coniopterygidae today is com- 2003, Fedotova & Perkovsky 2004, 2005, Per- prisedofminuteinsectswithreducedwingvena— kovsky & Fedotova2004, Polilov & Perkovsky tion; it has relatively few species, distributed 2004, Gumovsky & Perkovsky 2005, Tolkanitz worldwide. Despitethis, theyarecomparatively etal.2005a,b,Engel&Perkovsky2006a—c,Per- well studied, largely stimulatedbytheauthorita- kovsky 2006a,b, Simutnik & Perkovsky 2006, tivemonographofMeinander(1972). Sixspecies Zhantiev2006). Sixteeninsectordersandabouta ofthefamilyhavebeendescribedfromBalticam- hundred families have now been reported from ber(seelistinEngel2004),belongingtofivegen- Rovno amber (Perkovsky etal. 2003b, Engel & era, three of which are extinct: Archiconio— Perkovsky 2006a, Perkovsky 2006a). This in- compsa Enderlein, 1910, Archiconis Enderlein, cludes, however, no species ofNeuroptera. The 1930,HeminiphetiaEnderlein, 1930,allthreeare amber is chemically identical with Baltic amber monotypic. (Kosmowska-Ceranowicz 1999), and both are The presence of the Baltic amber Archi— consideredtoberoughlycontemporaneous,Late coniocompsaprisca in Rovno amber is not sur- 26 Kupryjanowicz&Makarkin ° ENTOMOL. FENNICAVol. 19 prising. SpeciesofDolichopodidae(Diptera)and senschaftliches Zentrum der Universitat Got- Rhinoterrnitidae (Isoptera) have been identified tingen [=Geoscience Centre oftheUniversity of in common between Rovno and Baltic ambers Gottingen], Germamy; MZ, Muzeum Ziemi (Grichanov 2000, Perkovsky et al. 2007). Fur- Polskiej AkademiiNauk [=MuseumoftheEarth ther, 74% of ant species and all 6 species of ofthePolishAcademyofSciences],Warsaw,Po- Psocoptera recorded from Rovno amber were land. previouslydescribedfromBalticamber(Dlussky & Perkovsky 2002, Engel & Perkovsky 2006b,c). Such shared species in other insect 3. Systematic paleontology familiesarenotyetknown,butmostoftheseare represented in Rovno amber by few species or 3.1.ArchiconiocompsapriscaEnderlein, 1910 havenotyetbeencloselyexamined.Therelation- (Figs 1—4) shipbetweentheBaltic andRovno amberinsect faunas is therefore onlybeginning to be evident Archiconiocompsaprisca Enderlein, 1910: 676, astheRovnoassemblagebecomesbetterknown. figs 1—4 (original description); Enderlein 1930: Nevertheless, it is currently apparent that the 101 (keyed);Meinander1972:34(systematicpo- Rovno amber insect assemblage is distinct from sition); Meinander 1975: 53 (taxonomic re- that ofBaltic amber by differing relative abun- marks); Meinander 1979: 20, 23 (systematicpo- dancesoforders,familiesandgroupswithinfam- sition, biogeography); Keilbach 1982: 284 ilies(Perkovskyetal. 2003a,b, Engel&Perkov- (listed); Nel 1990: 343 (listed); Spahr 1992: 71 sky2006a, Perkovskyetal. 2007). (listed); Weitschat&Wichard 1998: 146,pl. 53, figs c, e (photographs oftwo additional speci- mens);Dobosz&Krzeminski2000:220(listed); 2. Material and methods Engel2004: 134 (listed, systematicposition). Hololype(female):Numberunknown,earlier The material examined comes from the Klesov deposited “in the collection ofProfessor Dr. R. locality, about 90 km NNE of Rovno in the Klebs” in Konigsberg [=Kaliningrad], now lost Rovno Region, Ukraine. The specimen was do- orpossiblydestroyed,notfoundinthepartofhis nated to the Museum of the Earth by Andrzej collectionpreservedinthe GZG (Ch. Neumann, Wiszniewski (A.Wiszniewski collection), and M.Reich,pers.comm.).Balticamber(preciselo- preparedforthisstudybyJanuszKupryjanowicz. calitycollectedunknown). LateEocene. It is embedded in a translucent lump ofamber Additionalmaterial(apparentlyfemales): (1) withmeasurements 14x 12x5mm. Thereareno No. 5907, depositedinthe SZG, labelled“Phys. syninclusions. Photographs and measurements Oek. Ges. [=Physikalisch—Okonomischen Ge- weredoneusing aZeiss SV 11 microscope. sellschaft, Konigsberg]/Nr. 9267./11.1. Nr. 4.”; Terminology mainly follows Meinander “5907/ Coniopterix [sic]/ 5277-8”. Baltic amber (1972).Abbreviationsusedinthetextandfigures (preciselocalitycollectedunknown);(2)No.MZ are: A1, A2, 1st and 2nd anal veins (A); Cul, 24667, deposited in the MZ. Ukraine: Rovno Cu2, anterior and posterior branches of the [=Rivne] Region: Klesov [=Klesiv] locality cubitalvein(Cu); M1+2, M3+4, branches ofthe [51.315N 26.907E] (Rovno amber). Late Eo- medialvein(M);R1,anteriorbranchoftheradial cene. vein(R);R2+3,R3+4,branchesoftheradialsec- Description. (Rovno specimen): Body, ap- tor(Rs);Sc1,Sc2,anteriorandposteriorbranches pendages fuscous. Head with very large eyes ofthe subcostalvein (Sc). Crossveins are desig- (Fig. 2a); vertexnotespeciallyprominent, cove- natedafterthelongitudinalveinswhichtheycon- redwithratherscarce,finehairs.Antennaeshort, nect, and numbered in sequence from the wing 16-segmented; scape,pedicelsizesimilar, length base (ifmorethan one crossveinbetween avein ca. 1.5 timeswidth, stouterthanother segments; pair is present), e.g., 2m—cu1, distal crossvein 1stsegmentofflagellum(3rdantennal segment) connectingMandCu1. longer than other flagellar segments, length ca. Institutional abbreviations: GZG, Geowis- 2.2 times width; flagellar segments increasingly ENTOMOL. FENNICAVol. 19 - Archiconiocompsaprisca inRovnoamber 27 . . i I ,; v% vm 0.3mm pllc Fig.2.Archiconiocompsapn'sca, Rovnoamberspeci- Fig. 1 Archiconiocompsapn'sca, Rovnoamberspeci- men MZ24667.—a., Head.—b.Abdomen(lateral . view). mxpl,maxillarypalpus; plic, plicaturae;vtoix, men MZ24667.—a. Leftlateralview.—b. Rightlateral 5thto9thsternites. view. shorter,widertowardapex;terminalflagellomere sitebasalsc-r.Crossvein-likepartofSc2meeting tapering. Antennae covered with moderately R1 distadrl-rs.Basalsc-rveryshort. Sc2slightly long,densehairs.Terminalsegmentofmaxillary curvedtoward Scl, butapically separate. Origin palps much longer, broaderthanothers. Structu- ofRsatproximal1/3thofwinglength.Rsbasally res ofthoraxpoorlyvisibleby preservation, but sinuouswithdistinctknee(bend,asinMeinander as evident appear normal for family (Fig. 3b). 1972) at lrs-m;divisionintoR2+3, R4+5 shifted Legsofusualmorphologyforfamily,slender;fe- distally, rl-rsjoining Rs strongly proximad this mur offore leg stouter than other leg segments. fork. R2+3, R4+5 rather short, broadly spaced, Femur,tibiaofforelegmuchshorterthanthoseof anglebetweenthemalmost90 degrees. lrs-mat mid, hind legs. All femora covered with short, kneeofRs notdetected, butcrossvein-likeweak sparsesetae;tibiaewithlonger,densersetae.Tar- structurevisible slightlydistad forkofM; 2rs-m si five-segmented, all basitarsi longest oftarso- long, nearlyoppositerl-rs. McoalescentwithR meres, as long as, or slightly longer than four for short distance or connecting with it by very othertarsomeres together. Abdomencovered(at short crossvein proximally (this portion not most) with microtrichia. Abdominal plicaturae clearly visible; possibly former variant in left onthirdtosixthsternites,mostprominentonfifth wing, latter in right), deeply bifurcate distally, sternite (Fig. 2b). Seventh segment relatively with branches very long, relatively closely long,eighthsegmentappearingveryshort(indis- spaced;longsetaeontwolargethickeningsofM. tinct preservation). Genital structures suggests lm-cul very short, connecting M, Cul slightly specimenfemale,butnotclearlyvisiblebyclou- distal to fork ofCu; 2m—cu1 very strong, short, dyamber. slightly proximad fork of M. Cu dividing into Forewing(Fig.3).Length3.3mm,maximum Cul, Cu2 close to wing base; cul-cu2 short, width 1.2 mm. Scparallelto costalmargin. Two stronglyproximad2m—cu1. Twolongcrossveins basal crossveins (=subcostal veinlets) in costal between Cul, A1; lcul-al strongly curved. No space: weaker basad, rather strong distad oppo- crossveinbetweenA1, A2. ProximalhalfofA2, 28 Kupryjanowicz&Makarkin - ENTOMOL. FENNICAVol. 19 either intraspecific variation, possible errors in the original description, or different interpreta- tionsofthefeatures.Theseincludethefollowing fourdifferences. [1] Themostdistinctdifference in venation is the position ofthe crossvein-like partofSc2, which is situated opposite to thera- dial crossvein in the holotype, and distad that crossvein in the Rovno specimen (Fig. 3a). We interpret this as intraspecific variation. Similar variationsoccurwithinaspeciesintheclosestex- tant genus Coniocompsa, for example in C. sil- vestriana Enderlein [=C. smithersi Meinander] (seeMeinander 1972: figs 51Aand51G)widely distributed in Africa. Moreover, the position of the crossvein-like part of Sc2 in the specimen Fig.3.Archiconiocompsapn'sca, Rovnoamberspeci- 5907isidenticaltothatfoundintheRovnospeci- men MZ24667.—a., Rightforewing.—b.Thorax(dor- men. [2] The Rovno specimen is larger than the salview)andanalareaofrightforewing.a2-hm, Baltic amber holotype: the holotype forewing is crossveinbetweenA2andhind margin. 2.4 mm long, the Rovno specimen 3.3 mm. Al- thoughsuchsizedifference seemsgreat, itisnot hindmarginconnectedbyrather shortcrossvein exceptional within the Coniopterygidae. More- nearlyperpendiculartoA2(Fig.3b).Alllongitu- over, specimen 5907 is even larger than the dinal veins with one row ofshort dense setae, Rovnospecimen:theforewingoftheformerisca. prominent particularly on Scl, Sc2. Marginal 3.7mmlong. [3]AccordingtoEnderlein(1910), fringes short. Membrane hyaline, slightly plicaturae are present on the second to fifth fuscous, withoutdistinctcolourpatterning. sternites.Butweguessthattheyactuallyoccuron Hindwing. Length ca. 2.5 mm. Sc2 slightly thethirdtothesixthsternites,basedontheobser- curvedtowardSc1,butnotjoined.Crossvein-like vationthatbetween mostdistal segmentbearing partofSc2joinsR1 muchdistadr1-rs. Configu- plicaturae (fifth ofEnderlein, sixth in our inter- rationofRs forkas in forewing. 2rs-mnearly at pretation) and genital segments (ninth-tenth) in wingmid-point. StemofM, M3+4 formstraight theRovnospecimentherearetwosegments: one line; M1+2 originates atveryobtuseangletoM; relatively large (seventh) and one small (eighth) M1+2, M3+4 parallel. Other portions ofhind- (Fig.2b).Moreover,theplicaturaeonthesecond wings obscuredby forewings, venationnotvisi- abdominalsegmentingeneraofAleuropterygini ble. and Coniocompsini (to which this genus most Remarks. Although the Baltic amber speci- closelyrelated) areusuallyabsent,orthereareat men GZG 5907 was formerly deposited in most only traces of them present (Meinander Konigsberg(judgingbyits label), itdoesnotap- 1972). [3] Enderlein (1910) stated that the third pear to be the holotype of Archiconiocompsa antennalsegmentisalmostthreetimesaslongas prisca. Based on the original description ofthe wide, and is the narrowestofany segment. This holotypeofA.prisca, thesizeofspecimenGZG segmentintheRovnospecimenappearstobenot 5907ismuchgreaterthanthatoftheholotypeand particularlymoreslenderthanthenextdistaland itdiffersinvenationaldetails. TheholotypeofA. onlyca. 2.5 times longerthanwide (Fig. 2a). [4] prisca was described with enough detail by Enderlein (1910: fig. 1) showed in the forewing Enderlein(1910)toconfidentlyidentifytheRov- A2 connected with the hind margin by a long no specimen as conspecific, based on almost obliquecrossvein,sothatitappearsbifurcated.In complete morphological agreement between Rovno specimen, this crossvein is not so long, them, particularly in finedetails ofvenation, the and it is nearly perpendicular to A2. Unfortu- antennae and the abdomen. However, there are nately, it is impossible to checkthe accuracy of someminordifferences,whichwepresumetobe Enderlein’sdrawingbecauseoflossofthetype;it 30 Kupryjanowicz&Makarkz'n ° ENTOMOL. FENNICAVol. 19 (the latter ispossiblyparaphyletic: Engel2004). apomorphic features, which we consider to be Thevastmajority offossilAleuropteryginaebe- significant. If this is correct, then the Conio- longtoFontenelleini,including12species(Engel compsini should includeArchiconiocompsa and 2004, Nel et al. 2005). Exceptions areArchico— Coniocompsa, with the former possessesing a niocompsa andthepoorlyknownJuraconiopte— number of plesiomorphic character states not ryxMeinander,whichwastentativelyassignedto sharedwiththelatter(seeabove). this subfamily (Engel 2004). Archiconiocompsa surely does not belong to Fontenelleini, as its Acknowledgements.WethankAndrzejWiszniewski(Bia- hindwingcrossveinr1-rsjoiningRsstronglypro- lystok) for donation ofamberwith inclusions, including thatexaminedinthis study, to theMuseumoftheEarth; ximad the fork ofRs is distinctive. This genus ChristianNeumann(MuseumfiirNaturkunde,Berlin)for was originallyassignedto thetribeAleuroptery- providing information on the holotype ofArchiconio- gini(Enderlein 1910).Later,itwasconsideredas compsa prised; Mike Reich (Geowissenschaftliches having “an intermediatepositionbetweenAleu- ZentrumderUniversitatGottingen)forprovidinguswith ropterygini and Coniocompsini” (Meinander thephotographsoftheBalticamberNeuropteradeposited intheZentrum;EvgenyE. Perkovsky(SchmalhausenIn- 1972: 34), andsubsequentlywas referredto Co- stituteofZoology,Kiev),S.BruceArchibald(Museumof niocompsini (Meinander 1979). Engel (2004) Comparative Zoology, Massachusetts); Sonja Wedmann statedtheopinionthatthisgenushasanuncertain (InstitutfurPalaontology, Bonn), andananonymous re— tribe levelpositionwithintheAleuropteryginae. viewer for helpful discussions and comments that im— Ifourinterpretationoftheplicaturaepositionon provedthispaper;S.BruceArchibaldandJohnD.Oswald (TexasA&MUniversity)forcorrectionofEnglish;Galina abdominal segments (third to sixth) is correct, Sinelnikova (Institute of Biology and Soil Science, thentheprobabilityofassignmentofthisgenusto Vladivostok,Russia)forhelpwithdrawing. Aleuropterygini is increased, because plicaturae areunknowntooccuronthesixthsegmentinCo- niocompsini. The structures ofthe female abdo- References minal apex in the Rovno specimen appear most similar to those ofspecies ofConiocompsa (see Dlussky,G.M.2002:AntsofthegenusDolichoderus(Hy— for example Meinander 1972: figs 45C, 46F, menoptera: Formicidae) from the Baltic and Rovno 51C), and therefore would associate it with the ambers.7Paleontol.J. 36: 50763. Coniocompsini. Venational characters are con- Dlussky,G.M.&Perkovsky,E.E.2002:Ants(Hymenop— tradictory, allowing assignment to either taxon. tera, Formicidae) from the Rovno amber. 7 Vestn. Z001.36(5):3720.(InRussian,withEnglishabstract). As Meinander (1972) correctly mentioned, the Dobosz,R. &Krzeminski,W.2000:Anewspeciesofthe hindwingvenationismoresimilartothatofAleu- Coniopterygidae(Neuroptera)fromBalticamber.7 ropterygini than ofConiocompsini. Archiconio— P01. PismoEntomol. 69: 2197224. compsashareswithAleuropteryginiallfourfore- Enderlein,G. 1910:UberdeBeziehungenderfossilenCo— wing venational states by which Archiconio— niopterygidenzudenrecentenundfiberArchiconio- compsa and Coniocompsa are distinguished compsapriscanov.gen.nov.spec.72001.Anz. 35: 6737677. (above). These states, however, are plesiomorp- Enderlein, G. 1930: DieKlassifikationderConiopterygi- hic. On the other hand, the clearly apomorphic denaufGrundderrecentenundfossilenGattungen.7 venationalfeatures sharedbyArchiconiocompsa Arch.Klassifikat.Phylogenet.Entomol. 1(2):987114. andConiocompsa(above),the 16-segmentedan- Engel,M.S.2004:ThedustywingsinCretaceousBurmese tennaeandpossiblegenitalresemblanceofArchi— amber (Insecta: Neuroptera: Coniopterygidae). 7J. Syst. Palaeontol. 2: 1337136. coniocompsato Coniocompsa, are suggestiveof Engel,M. S.&Perkovsky,E.E.2006a:AnEocenebeein theConiocompsini. Rovno amber, Ukraine (Hymenoptera, Megachili- In summary, although an argumentmade by dae).7Am. Mus.Novit. 3506: 1711. previous authors thatthetribeposition ofArchi— Engel,M.S.&Perkovsky,E.E.2006b:Psocoptera(Insec— coniocompsa to be rather uncertain has some ta)inEoceneRovnoamber(Ukraine).7Vestn.Zool. 40: 1757179. strengths,insomerespectsindeeditisintermedi- Engel, M. S. &Perkovsky, E. E. 2006c: Sphaeropsocus ate between the Aleuropterygini andthe Conio- kuenowii Hagen in Rovno amber from the Ukraine compsini, still, we tentatively assigned it to the (Psocoptera: Sphaeropsocidae). 7 Entomol. News Coniocompsini, based on the presence ofthese 117: 2437245. ENTOMOL. FENNICAVol. 19 ° Archiconiocompsaprisca inRovnoamber 31 Fedotova,Z.A.&Perkovsky,E.E.2004:Newgallmidges amber: subfamily Lestremiinae, tribes Micromyiini (Diptera,Cecidomyiidae)fromtheRovnoamber:sub— andPeromyiini.7Paleontol. J. 38: 3967106. familyLestremiinae,tribesStroblielliniandCampylo- Perkovsky,E. E.,Zosimovich,V. 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J. 37: 2757 Simutnik,S.A.&Perkovsky,E.E.2006:Adescriptionof 279. the encyrtid male (Hymenoptera, Chalcidoidea, En- Kosmowska—Ceranowicz, B. 1999: Succinite and some cyrtidae) with archaic structure of metasoma from other fossil resins in Poland and Europe (deposits, f1nds,featuresanddifferencesinIRS).7Estud.Mus. Rovnoamber.7Vestn. Zool. 40:2837286. Cienc.Nat.Alava 14(num. espec. 2): 737117. Spahr, U. 1992: Erganzungen und Berichtigungen zu R.Keilbachs Bibliographie und Liste der Bemstein- Meinander,M. 1972:ArevisionofthefamilyConioptery- gidae(Planipennia).7Acta2001.Fenn. 136: 17357. fossilien7KlasseInsecta [Ausgenomen: “Apterygo— ta”, Hemipteroidea, Coleoptera, Hymenoptera, Me- Meinander, M. 1975: Fossil Coniopterygidae (Neuropte— ra).7Not. Entomol. 55: 53757. copteroidea]. 7 Stuttg. Beitr. Natkd. Ser. B (Geol. Palaeontol.) 182: 17102. Meinander, M. 1979: The phylogeny and geographical distributionoftheAleuropteryginae(Neuroptera,Co— SZiraki,G. 1998:ZoogeographicrelationsofSouthAsian niopterygidae).7Ann.Entomol. Fenn. 45: 16723. coniopterygids (Neuroptera, Coniopterygidae). 7 Nel, A. 1990 [1991]. Nouveaux insectes neuroptero'1'des Acta2001.Fenn. 209:2497254. fossilesdel’OligocenedeFrance(NeuropteraetMe— Tolkanitz,V.I.,Narolsky,N.B.&Perkovsky,E.E.2005a: galoptera).7Bull.Mus.Natl.Hist.Nat.(Paris).Sect. A new species ofparasitic wasp ofthe genusPher- C(Sci. Terre)(4) 12: 3277349. hombus (Hymenoptera, Ichneumonidae, Pherhombi— Nel, A., Perrichot, V. &Azar, D. 2005: Newandpoorly nae)fromtheRovnoamber.7Paleontol.J.39:5117 knownfossilConiopterygidaeinCretaceousandCe— 513. nozoic ambers (Insecta: Neuroptera). 7Ann. 2001. Tolkanitz,V.I.,Narolsky,N.B.&Perkovsky,E.E.2005b: (Wars.)55: 177. 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