Fossil Sciaroidea (Diptera) in Cretaceous Ambers ...

Copyright q American Museum of Natural History 2004 ISSN 0003-0082

P U B L I S H E D B Y T H E A M E R I C A N M U S E U M O F N AT U R A L H I S T O RY

CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024

Number 3433, 76 pp., 81 figures, 7 plates, 7 tables February 27, 2004

Fossil Sciaroidea (Diptera) in Cretaceous Ambers,Exclusive of Cecidomyiidae, Sciaridae, and

Keroplatidae

VLADIMIR BLAGODEROV1 AND DAVID GRIMALDI2

CONTENTS

Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

Materials and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4Systematic Paleontology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

Superfamily Sciaroidea Billberg, 1820 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6Family Diadocidiidae Edwards, 1925 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

Docidiadia, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6Sciaroidea incertae sedis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9Thereotricha, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

Family Lygistorrhinidae Edwards, 1925 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12Archaeognoriste, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12Lebanognoriste, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14Plesiognoriste, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14Protognoriste, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17Leptognoriste, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

Family Mycetophilidae Newman, 1834 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22Subfamily Manotinae Edwards, 1925 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

Alavamanota Blagoderov and Arillo, 2002 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

1 Division of Invertebrate Zoology (Entomology), American Museum of Natural History; Paleontological Instituteof Russian Academy of Sciences. e-mail: [email protected]

2 Division of Invertebrate Zoology (Entomology), American Museum of Natural History. e-mail: [email protected]

2 NO. 3433AMERICAN MUSEUM NOVITATES

Subfamily Sciophilinae Winnertz, 1863 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22Tribe Sciophilini Winnertz, 1863 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22Neuratelia Rondani, 1856 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22Allocotocera Mik, 1886 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24Pseudomanota, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

Tribe Gnoristini Edwards, 1925 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26Apolephthisa Grzegorgzek, 1885 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26Synapha Meigen, 1818 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28Dziedzickia Johannsen, 1909 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28Saigusaia Vockeroth, 1980 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31Syntemna Winnertz, 1863 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32Gregikia, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32Gaalomyia, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34Tribe Leiini Edwards, 1925 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36Nedocosia, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36Ectrepesthoneura Enderlein, 1911 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39Izleiina, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41Zeliinia, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43Temaleia, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45Lecadonileia, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47Disparoleia, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48Hemolia, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48Protragoneura, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52

Analyses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52Lygistorrhinidae and the Heterotricha Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52Mycetophilidae s.s. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54

Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56Phylogeny . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56Taphonomy and Biogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65

Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65

2004 3BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER

ABSTRACT

The Recent world fauna of Sciaroidea, or fungus gnats, comprises approximately 4000described species in eight families: Bolitophilidae, Cecidomyiidae, Diadocidiidae, Ditomyi-idae, Keroplatidae, Lygistorrhinidae, Mycetophilidae, and Sciaridae. Larvae live primarily indecaying vegetation, feeding on fungal mycelia, and they can be among the most abundantinsects of temperate forests. Stem-group families appeared in the Jurassic, with large Tertiarydeposits being composed almost entirely of living genera, so the Cretaceous is essential forunderstanding the origins and diversification of Recent families. Sixty-six specimens werestudied from six major deposits of Cretaceous amber, spanning 40 million years from the Earlyto Late Cretaceous: Lebanon (ca. 125 Ma), northern Spain (120 Ma), northern Myanmar(Burma) (ca. 105 Ma), northern Siberia (two sites, 105 and 87 Ma), New Jersey (90 Ma), andwestern Canada (80 Ma).

New taxa are the following: Docidiadia burmitica (n.gen., n.sp.) (Diadocidiidae); Thereo-tricha sibirica, (?)T. agapa (n.gen., n.spp.) (Sciaroidea incertae sedis); Archaeognoriste prim-itiva, Lebanognoriste prima, Plesiognoriste carpenteri, P. zherikhini, Protognoriste amplicau-da, P. goeleti, P. nascifoa, Leptognoriste davisi, L. microstoma (n.gen., n.spp.) (Lygistorrhin-idae). In Mycetophilidae sensu stricto: Alavamanota burmitina, n.sp. (Manotinae), Neurateliamaimecha, n.sp., Allocotocera burmitica, n.sp., Pseudomanota perplexa, n.gen., n.sp. (Scio-philinae Sciophilini); Apolephthisa bulunensis, n.sp., Synapha longistyla, n.sp., Dziedzickianashi, n.sp., Saigusaia pikei, n.sp., Syntemna fissurata, n.sp., Gregikia pallida, n.gen., n.sp.,Gaalomyia carolinae, n.gen., n.sp. (Sciophilinae Gnoristini); Nedocosia exsanguis, N. sibirica,N. canadensis, N. novacaesarea, n.gen., n.spp.; Ectrepesthoneura succinimontana, E. swol-enskyi, n.spp.; Izleiina mirifica, I. spinitibialis, n.gen., n.spp.; Zeliina orientalis, Z. occidentalis,n.gen., n.spp.; Temaleia birmitica, n.gen., n.sp., Lecadonileia parvistyla, n.gen., n.sp.; Dis-paroleia cristata, n.gen., n.sp.; Hemolia matilei, H. glabra, n.gen., n.spp.; and Protragoneuraplatycera, n.sp. (Sciophilinae Leiini).

Relationships of the fossil genera are phylogenetically assessed with living genera. TheBurmese amber fauna contains an inordinate abundance and diversity of sciaroids, perhapsbecause of a wetter paleoclimate in that region.

INTRODUCTION

The preservation of fossils in amber is re-nowned for consistently finer preservationthan virtually all other modes of fossilization.Although the geological occurrence of amberbegins in the Triassic, the oldest insect faunasin amber are no older than Early Cretaceous,approximately 120–130 million years ago(Ma). Nonetheless, insects in amber from theCretaceous period (145–65 Ma) afford un-paralleled insight into Mesozoic evolution.The many more preserved characters allowmore accurate phylogenetic inference than ispossible with compression or most otherkinds of fossils, particularly for intricate or-ganisms like insects. Detailed preservationfacilitates not only reconstruction of phylog-eny, but of paleoenvironments as well. In thisregard, the study of extinct Diptera is essen-tial, as Diptera usually comprise the largestproportions by individuals and species of alltypes of arthropods preserved in various am-ber deposits around the world. This paper is

devoted to one particularly diverse group offlies well represented in Cretaceous amber,but which have been virtually neglected: thefungus gnats, or Sciaroidea.

The Sciaroidea is comprised of 14 fami-lies, 6 of them exclusively Mesozoic: Ante-fungivoridae (Middle Jurassic to Early Cre-taceous; Kovalev, 1990), Archizelmiridae(Late Jurassic to Late Cretaceous; Grimaldiet al., 2003), Eoditomyiidae (Early Jurassicto Early Cretaceous; Ansorge, 1996), Meso-sciophilidae (Middle Jurassic to Early Cre-taceous; Kalugina and Kovalev, 1985, Bla-goderov, 1993), Pleciofungivoridae (EarlyJurassic to Early Cretaceous; Kovalev,1987b), and Protopleciidae (Early Jurassic toEarly Cretaceous; Blagoderov, 1996, Koval-ev, 1990). No extinct families occur in theCenozoic. The extant families of the super-family are Bolitophilidae, Cecidomyiidae,Diadocidiidae, Ditomyiidae, Keroplatidae,Lygistorrhinidae, Mycetophilidae (s.s.), andSciaridae, all of which but Ditomyiidae also


occur in the Cretaceous. Phylogenies andcomprehensive revisions of some of thesefamilies were provided by Grimaldi and Bla-goderov (2001) (Lygistorrhinidae), Matile(1981a, 1990, 1997) (Keroplatidae, Sciaro-idea), Munroe (1974) (Ditomyiidae), Soli(1997) (Mycetophilidae s.s), and Vaisanen(1984) (Mycomyiini). There are approxi-mately 4100 described species of Sciaroideaexclusive of the poorly studied but diversefamilies Cecidomyiidae and Sciaridae (Be-chev, 2000). At least 10,000 and perhaps asmany as 20,000–30,000 living species ofSciaroidea exist. Such diversity reflects thesignificance of the group in terrestrial eco-systems, particularly forests, the very habitatthat produces amber.

Larvae of Sciaroidea are abundant in thesporophores of basidiomycete fungi—in de-caying wood, leaf litter, and humus, wherethey feed on fungal mycelia. Where humusand decaying wood are particularly thick inmature forests, sciaroids are among the mostabundant and diverse insects, with their lar-vae inhabiting 80–90% of the macrofungi innatural habitats (Yakovlev, 1988). Larvalhabits are not restricted, though, to mycoph-agy (Matile, 1997). The larvae of some gen-era and species of Keroplatidae are actuallypredators of insects they snare in their slimetrails, the most famous example of which isthe New Zealand ‘‘glow-worm’’, (Arachno-campa Edwards). Larvae of Arachnocampaare luminescent and suspend themselves inmucous strands from the ceilings of caves.Small midges attracted to the light becomeentangled in the strands, which the larvaethen devour. There is even a keroplatid,Planarivora Hickman, whose larva is para-sitic on land planarians in Tasmania (Hick-man, 1964; Matile, 1981b).

The fossil record of the Sciaroidea beginsin the Early Jurassic (Ansorge, 1996; Kalu-gina and Kovalev, 1985; Kovalev, 1987a),and by the Early Cretaceous modern familiesreplaced older, stem-group families (Kovalev,1987a). The Cenozoic Sciaroidea are partic-ularly well studied, especially those in Balticamber (approximately 270 species of Myce-tophilidae, as reviewed in the catalogue offossil Diptera [Evenhuis, 1994]). The Balticamber sciaroid fauna, like most of the otherCenozoic faunas of these flies, is essentially

modern. Prior to the work of the senior authoron extensive insect Lagersatte from Asia (Bla-goderov, 1995, 1997, 1998a, 1998b, 2000),there were only seven species of sciaroidsknown from the Cretaceous, which wereplaced in the Mesosciophilidae or Myceto-philidae (Kovalev, 1986, 1990; Hong, 1992;Ren et al., 1995; Jell and Duncan, 1986;Westwood, 1854). Sciaroids have been knownfor years to occur in the major deposits ofCretaceous ambers from Taimyr, Siberia(Zherikhin and Sukacheva, 1973; Zherikhin1978), Burma (Zherikhin and Ross, 2000),New Jersey (Grimaldi et al., 1989), and west-ern Canada (McAlpine and Martin, 1969;Pike, 1994). Only three species, however,were described: Burmacrocera petiolataCockerell, Sciara burmitina Cockerell (inBurmese amber, thought at the time to be Ce-nozoic), and Schlueterimyia cenomanica Ma-tile (in Cenomanian French amber). Sincethen, major new Cretaceous outcrops of am-ber have been discovered from northern Spain(Alonso et al., 2000), Lebanon (Azar, 2000),New Jersey (Grimaldi, 2000), and Myanmar(Grimaldi et al., 2002), which have producedsignificant numbers of new specimens andtaxa of sciaroids. Based on study of the oldercollections and these newer ones, we signifi-cantly extend the known Cretaceous diversity,based on 27 genera (19 of them new) and 39species (all of them new).

MATERIALS AND METHODS

Sixty-six specimens from the Early andLate Cretaceous of six countries, and some10 localities, were examined. We did notstudy inclusions of Diadocidiidae in Burmeseamber from the Natural History Museum,London, since these were on loan. Adults ofmost extant genera of Mycetophilidae (s.s),and all species of Lygistorrhinidae were ex-amined. Material is housed in the followinginstitutions:

AMNH American Museum of Natural History(Division of Invertebrate Zoology),New York, USA

MCNA Museo de Ciencias Naturales de Alava/Arabako Natur Zientzien Museoa, Vi-toria-Gasteiz, Spain

MCZ Museum of Comparative Zoology (De-partment of Entomology), HarvardCambridge, Massachusetts, USA


PIN Paleontological Institute of the RussianAcademy of Sciences (ArthropodaLaboratory), Moscow, Russia

TMPD Tyrrell Museum of Palaeontology, Al-berta, Canada

Specimens were derived from the follow-ing localities and formations:

Alava, Spain: The amber deposit is lo-cated on the northern slope of Sierra de Can-tabria (Alava), about 30 km southeast of thecity of Vitoria-Gasteiz, near the village ofPenacerrada. A comprehensive description ofthe amber deposit was given by Alonso et al.(2000). It was assigned to the Nograro For-mation, of Aptian–middle Albian age (120–110 Ma). All of the material is housed inMCMA.

Burma (Myanmar): Historically andpresently all amber from Burma derives fromthe northern state of Kachin. Zherikhin andRoss (2000) reviewed historical records ofBurmese amber mining and locations. Formany years Burmese amber was consideredCenozoic in age; however recently it hasbeen found to be Cretaceous after the dis-covery of insects in it that are exclusivelyCretaceous, like Serphitidae and Stigma-phronidae (Hymenoptera). A collection ofapproximately 1200 arthropod inclusions re-sides at the NHM, London, rich in types de-scribed by T.D.A. Cockerell and others. Re-cently, a very large, diverse collection ofBurmese amber has been assembled at theAMNH from material recently excavated inKachin, near villages close to the town ofMyitkyina. This material was acquired byLeeward Capital (Calgary, Canada), whoprovided it to the AMNH. Study of some3500 organisms in this collection revealedmore than 20 Mesozoic insect taxa, confirm-ing the Cretaceous age and even indicatingan age of approximately Cenomanian (98–92Ma) (Grimaldi et al., 2002). Burmese amberis by far the most prolific and diverse Cre-taceous amber for Sciaroidea.

Canada: Material from Grassy Lake, Al-berta, derives from an abandoned coal mine8 km south and 1.6 km east of the village ofGrassy Lake (Pike, 1994). This was collectedby T. Pike and housed at the RTMP. Thislocality was previously cited as ‘‘near Med-icine Hat’’ by McAlpine and Martin (1969),who deposited their material at the Canadian

National Collection (CNC) of Insects andSpiders, Ottawa. The Grassy Lake amber isfrom the Judith River Group of the ForemostFormation, dated as Campanian (Pike, 1994;Borkent, 1995), although Grimaldi and Cum-ming (citing David Eberth, personal com-mun.) indicated the amber to be Santonian.Amber from Cedar Lake, Manitoba was col-lected by F.M. Carpenter in 1938 amongbeach debris (in the MCZ). Although rede-posited and somewhat distant from theGrassy Lake deposit, the two deposits arethought to be contemporaneous (Borkent,1995). An age of 85–80 myo is a reasonableestimate for Canadian amber.

Lebanon: Material studied here is fromthe Acra Collection (collected by Fadi andAftim Acra near Jezzine, 30 km east of Sai-da, Lebanon) and the Estephan Collection(collected by Antoun Estephan, near Bcharre,northern Lebanon). Both of these collectionsare housed in the AMNH. Most recently, ex-tensive collections and stratigraphic studiesof Lebanese amber have been made by Azar(2000). Lebanese amber was originally re-ported to be Neocomian in age, pertaining tobasal periods of the Early Cretaceous (Schleeand Dietrich, 1970), 145–129 Ma. A youngerage of Aptian-Albian (ca. 120–110 Ma) hasalso been mentioned (Whalley, 1976; Zheri-khin, 1978). Azar’s study, which is the mostcomprehensive, indicates a great range ofages of Lebanese amber, from uppermost Ju-rassic (152 Ma) to Albian (112 Ma), but mostof the deposits yielding insect inclusions areBarremian-Aptian (approximately 125 Ma).Sciaroids in Lebanese amber are the oldestknown amber fossils of the superfamily.

New Jersey: Rich amber deposits fromthe central county of Middlesex have beenknown for decades, summarized by Grimaldiet al. (1989). Recently, an extraordinary out-crop was discovered in the town of Sayre-ville, with a smaller one being discowered inthe adjacent town of East Brunswick (Gri-maldi, 2000). This material has produced theoldest definitive ants, tardigrades, mush-rooms, parasitiform mites, and flowers(among many others) preserved in amber.Chemical analyses of the amber and stratig-raphy of the outcrops have been made. Theamber occurs in the South Amboy Fire Clayof the Magothy Formation, Turonian in age


(Grimaldi, 2000). All of the New Jersey am-ber is deposited in the AMNH.

Siberia (Taimyr): Reviews of the arthro-pod fauna in the Siberian amber were pro-vided by Zherikhin and Sukacheva (1973)and Zherikhin (1978). Sciaroidea were foundat four localities, ages of which vary. Mate-rial was collected by a PIN expedition in1973 to Nizhnaya Agapa, which is on thenorth shore of the Agapa River, 40 km down-stream from Lake Ladonnakh, Yst’-Eniseydepression. This material is in the DolganFormation, dated palynologically as Albian–Cenomanian (Saks and Ronkina, 1957). Am-ber from Yantardakh (‘‘amber mountain’’)occuring 3–5 km upstream from the mouthof the Maimecha River, Khatanga depres-sion, in the Kheta Formation (dated as Con-iacian–Santonian [Saks et al., 1959], ca. 87myo) was collected in 1970 and 1971. A1976 PIN expedition collected a contempo-raneous material from Bulun (middle courseof Bulun River [right tributary of Kheta Riv-er], 18 km S of post Novaya, Taimyr). Am-ber from Baikura-Neru (Taimyr Lake, Baik-ura-Neru Bay) contains the only incompletespecimen of Sciaroidea. Baikura-Neru, ageof which is unclear, has been assigned to theOgneva Formation (Saks et al., 1959; Ap-tian-Albian, 120–110 Ma), but arthropod in-clusions indicate a Late Cretaceous age(Dlussky, 1987; Zherikhin and Eskov, 1999).All Siberian amber is in PIN.

Specimens originally stored in small boxeswere prepared according to the method de-scribed in Nascimbene and Silverstein(2000). This involved embedding the speci-men in a stable epoxy (Buehler) under vac-uum. The vacuum extracts air in fine cracks,which the epoxy then permeates, thus im-proving visibility and fragility of the piece.Pieces were then trimmed to thin pieces, withsurfaces often to within fractions of a milli-meter of the inclusion, in order to optimizeobservation of details. Prepared in this way,pieces can be mounted on microscope slidesand the inclusions observed under 100–4003 magnification with a compound micro-scope. Specimens were measured with a dig-ital stage micrometer mounted under a ZeissSV8 stereoscope, and photographed using anInfinity K-2 lens attached to a Nikon D-1camera and illuminated with focusable fiber-

optic flash wands (ML-1000 from Microp-tics).

Morphological terminology used followsMcAlpine (1981) and Soli (1997) with mod-ification after Kovalev (Kalugina and Koval-ev, 1985). To compare relative position ofveins in sciaroid wings, Kovalev defined bas-al (from the base of Rs to the base of r-m),middle (from the base of r-m to the base ofR4), and apical (equivalent to R5) sections inradius stem as RS1, RS2, and RS3 sections,respectively. Also, the basal (from the baseof the wing to tb), middle (from tb to r-m),and apical (from r-m to the base of the M1

and M2 fork, equivalent to the stem of M112)sections of media stem were defined as M1,M2, and M3 (see also figs. 80, 81). The term‘‘stem of M’’ refers to M1, M2, and M3 sec-tions together. The term ‘‘base (or stem) ofM1 and M2 fork’’ refers to the point of fur-cation of the veins. The terms ‘‘stem of M1

and M2 fork’’ or ‘‘stem of M112’’ refer to M3section. The terms ‘‘R5’’ and ‘‘stem of M112’’are used to refer to the veins themselves, and‘‘RS3’’ and ‘‘M3’’ are used in reference tothe relative length of sections. All the veinsand section names are summarized in table1, which shows comparison of the vein no-menclature used by various authors. In somecases, for the sake of stability, we preferredto use traditional vein names in descriptionsof species rather than use their names basedon presumed, primary homology. Theseveins are denoted in boldface type in the ta-ble. In the discussion of phylogeny, however,it seemed advisable to use names of veinsbased on homology for establishing transfor-mation series. We used the term ‘‘not visi-ble’’ when a structure could not be observedclearly.

SYSTEMATIC PALEONTOLOGY

SUPERFAMILY SCIAROIDEA BILLBERG, 1820

FAMILY DIADOCIDIIDAE EDWARDS, 1925

Docidiadia, new genus

DIAGNOSIS: Head round. Flagellum 14-seg-mented. First flagellomere length slightlymore than width. Fore tibial comb absent.Wing membrane without macrotrichia. Cends beyond tip of R5; Sc long, ends free;RS base at the middle of R1; crossveins r-m,


TABLE 1Nomenclature and Homology of Sciaroid Wing Veins According to Authors

See also figures 80 and 81.

tb, and m-cu in one line; M3 section and baseof M fork absent; CuA strongly curved backat the apex. A short. Male 9th tergite withoutmarginal bristles, with one large, acute, tri-angular medial appendage and two small lat-eral ones; gonostyli do not bifurcate at apex.

TYPE SPECIES: Docidiadia burmitica, n.sp.ETYMOLOGY: The name is a feminine an-

agram of Diadocidia.COMMENTS: The genus is close to Diado-

cidia Ruthe, but differs in having the firstflagellomere short; wing membrane withoutmacrotrichia, Sc ending free, base of RSrather distal, M3 section and base of M forkreduced; CuA curved at apex rather thanwith two straight sections; and male tergiteIX narrow, triangular, and with two lateralappendages. Diadocidia consists of two sub-genera and includes 10 Holarctic species(Chandler, 1994; Lastovka and Matile, 1972;Polevoi, 1996; Wu, 1995; Zaitzev, 1994) anda Neotropical one (Edwards, 1940; Papavero,1977a), as well as undescribed Australianspecies (Tonnoir, 1929; Colless, 1963). One

species is known from Baltic amber (Even-huis, 1994).

Docidiadia burmitica, new speciesFigures 1–3, Plate 1A

DIAGNOSIS: As for genus.DESCRIPTION: Body length 5 1.88 mm (ho-

lotype)/1.61 (paratype); wing length 5 1.61/1.55 mm. Head. Eyes bare, facets round.Clypeus setose. Flagellum 14-segmented;first flagellomere cylindrical, width slightlyless than length; apical flagellomere twice thewidth. Apical flagellomere of male second-arily segmented in two parts. Scape and ped-icel turbinate. Only 3 segments of palpi seen,palpomeres cylindrical, subequal in length,basal wider than the rest. Thorax. Scutumsetose, dome-shaped, with long protrudingsetae. Metepisternum bare, height equal towidth, shallow incision anteriorly. Wingmembrane without macrotrichia. Costa endsbeyond tip of R5, midway between tips of R5

and M1. Sc ends free, slightly beyond base


Figs. 1–3. Docidiadia burmitica, n.sp. 1. Holotype AMNH Bu-033. 2. Tip of the antenna of theholotype. 3. Male genitalia of the holotype.

of RS. R1 setulose, R5 with sparse setae, al-most straight. R1 short, about 0.63 winglength. M3 absent. M1 and M2 weakened,their bases absent. M314 weakened. Abdo-men. Female cerci wide, subtriangular withacute ventral angle. Male tergite IX narrow,triangular with two lateral appendages. Gon-

ocoxites short, with length about the width.Gonostyli massive, length 2.53 the length ofgonocoxites, hairy, without apical teeth orspines.

MATERIAL: Holotype AMNH Bu-033,male; paratype B-002, female. Myanmar:Katchin, from amber mines near Myitkyina.


ETYMOLOGY: The species epithet is a ref-erence to Burma, the former name of thecountry where the amber originates.

SCIAROIDEA INCERTAE SEDIS

Thereotricha, new genus

DIAGNOSIS: Eyes forming incomplete eyebridge, facets large, round. Ocelli three. Fla-gellomeres barrel-shaped, length no morethan 1.53 the width. Antepronotum andproepisternum subequal, setose. Proepimerontouches episternum at the episternal suture.Anepisternum smaller than katepisternum.Anepisternal cleft distinct, narrow. Midpleur-al pit present. Metepisternum setose. Ante-rior parapsidal suture distinct. Insertion ofabdomen wide. Wing membrane with orwithout macrotrichia. Sc short, ends free. Rsbase, r-m, base of M314, and CuA fork verybasal. Section M2 connects r-m and base ofM314 and CuA fork. M314 reduced.

TYPE SPECIES: Thereotricha sibirica, n.sp.ETYMOLOGY: The genus name is a femi-

nine anagram of Heterotricha. The name isfeminine.

COMMENTS: The new genus is close to theHeterotricha group of genera, which havebeen included in the Sciaridae or Diadoci-diidae. Recently, Chandler (2002) reviewedknown taxa of the group and described sevenmore genera from all zoogeographic regionsexcept Nearctic. These taxa seem to repre-sent a stem group of Recent families of Sci-aroidea, but monophyly of the group is notapparent. The new genus differs from all taxaof the Heterotricha group in having eyeswith large facets, that form an eye bridge,short antennae, the scape and pedicel not dif-fering from flagellomeres in length, palpivery short, anepisternite and katepisternitesubequal, the base of RS in basal position,and r-m and the section of M2 subequal,where eyes and palpi demonstrate an apo-morphic condition. The new genus resemblesSciaropota Chandler in the porrect antennawith short flagellomeres, an absence of aclearly differentiated series of scutellar bris-tles, large katepisternum, mesepimeponbroader below, but it differs by the short Sc,long stem of M112, and reduced M314.

Distinct synapomorphies separate thegroup from other Mesozoic Sciaroidea: M1

section completely reduced; M2 sectionfused with tb in one vein meeting the baseof M314, and that oblique vein is shifted dis-tad. The same structure of the basal veins isobserved in advanced representatives of theMesozoic family Mesosciophilidae, whichare thought to represent a sister group to My-cetophilidae (Kalugina and Kovalev, 1985;Blagoderov, 1993). Similar conditions occurin the peculiar Mesozoic family Archizel-miridae (Grimaldi et al., 2003), but Archi-zelmiridae have crossvein r-m aligned withM2 1 tb and the basal portion of M314, form-ing one horizontal vein and the base of RSis shifted distad. Diadocidiidae s.str. (Dia-docidia and Docidiadia n.gen.) also havethese veins aligned (r-m through the base ofM314), but they form a vertical vein. In My-cetophilidae the combined vein M2 1 tb lostcontact with the base of M314 and meets thebase of CuA or MA (arculus) (see Shcher-bakov et al., 1995). Some Mycetophilidae(Drepanocercus, Ectrepesthoneura, Creta-ceous Paradzickia, Drepanorzeckia, Ekhiri-tus, Zazicia) have the fork of M314 and CuAsessile or short-stalked, but the base of thefork is situated more basally that in MesozoicSciaroidea and the Heterotricha group.Moreover, at least in Ectrepesthoneura thesessile fork of M314 and CuA is secondary(see Analyses below and fig. 78). Obviously,reduction of M1 and fusion of M2 with tbmight have originated several times in thehistory of Sciaroidea. Although monophylyof the group combining Recent Heterotricha-like taxa and Cretaceous Thereotricha is notproven, position of these taxa in sciaroidphylogeny should be at the base of lineagesleading to Mesosoic Mesosciophilidae andArchizelmiridae and Recent Sciaridae on theone hand and higher sciaroids such as My-cetophilidae and Lygistorrhinidae on the oth-er.

Thereotricha sibirica, new speciesFigure 4, Plate 1B

DIAGNOSIS: Wing membrane without mac-rotrichia, RS base very basal, oblique, ;23the length of r-m; M3 section long; base ofM1 and M2 fork at level of tip of R1; veinsR1, R5, M1, M2, and CuA with long setae.

DESCRIPTION: Body length 5 1.83 mm (ho-


→

Figs. 4–6. Thereotricha, n.gen. 4. Thereotricha sibirica, n.sp., drawing of amber piece with fourembedded specimens, including holotype PIN 3130/183. 5.? T. agapa, n.sp., holotype PIN 3426/256.6.? T. agapa, head of the holotype.

lotype)/1.84–1.85 mm (paratypes); winglength 5 1.46 mm (holotype)/1.25 mm (para-type PIN 3130/185). Head. Three ocelli pre-sent, equal in size, distance of lateral ocellusfrom median ocellus and eye margin 1.53 itsdiameter. Occiput with numerous small setae.Eyes large, bare, forming incomplete dorsalbridge (separated by twice the facet diame-ter). Scape and pedicel short, rounded. Fla-gellum 14-segmented, flagellomeres barrel-like, width 1.33 the length, covered by shorttrichia. Palpi 4-segmented. Basal segmentsmall, bacilliform, others short, rounded; an-tepenultimate segment with round sensorypit dorsomedially, slightly longer than oth-ers.Thorax. Antepronotum with one strongseta, proepisternum with six. Proepimerontouches episternum at episternal suture. Me-sonotum uniformly setose. Midpleural pitpresent. Katepistrnum large, expanded cau-dally. Ventral part of mesepimeron not verynarrow, parallel-sided. Laterotergites bare,small. Metepisternum setose, with narrowanteriodorsal cleft. Wing membrane withoutmacrotrichia. Costa ends beyond tip of R5,one-third the distance between tips of R5 andM1. Sc very short, ends free just beyond RSbase. RS base proximad, oblique, 1.53 r-mlength. R1, R5, M1, M2, and CuA with longsetae. R1 about 0.73 wing length. RS baseoblique. Crossvein r-m fused with tb and m-cu in one horizontal vein. M3 section 1.53as long as fork of M1 and M2. M314 reduced,seen as a fold at base only. CuA ends beforemiddle of the wing. Legs. Fore coxae withnumerous setae on anterior and distal surfac-es, mid coxae with distal rows and severalsetae on apical part, hind coxae with caudo-distal row of setae. Fore tibia with anteroap-ical pit having two combs of setae. Abdo-men setose. Eighth segment slightly shorterthan preceding ones. Ninth tergite ovoid,with apical comb of short blunt setae. Gon-ocoxites stout, triangular in lateral view.Gonostyli small, flat, rectangular, with sev-eral apical setae.

MATERIAL: Holotype PIN 3130/183, male;

paratypes, PIN 3130/182, 3130/184, 3130/185, males; all specimens in the same pieceof amber. Russia: Taimyr Peninsula, Yantan-darkh, coll. 1970.

ETYMOLOGY: The species epithet is in ref-erence to Siberia, the region where the orig-inal locality is situated.

?Thereotricha agapa, new speciesFigures 5, 6, Plate 1C

DIAGNOSIS: Three ocelli, touching. Wingmembrane with scattered macrotrichia. RSbase transverse, shorter than r-m. M2 proba-bly absent.

DESCRIPTION: Body length 5 1.56 mm(rest, estimated length 1.87 mm); winglength 1.48 mm. Partly preserved specimen.Head: Occiput and vertex with straight andcurved setae. Three ocelli present, each onits own mound, the latter fused by contactingedges. Eyes bare, with incomplete bridge,deeply emarginate at antennal base. Frontalfurrow and keels well developed. Scape andpedicel as long as flagellomeres. Flagellum14-segmented, flagellomeres barrel-shaped,length about 1.53 the width, with few strongsetae and numerous fine trichia. Clypeus se-tose. Palpi 4-segmented, short. Antepenulti-mate palpomere with dosomedial round sen-sory pit. Thorax: Proepisternum with 3 longand and 11 shorter setae. Proepimeron touch-es episternum at the episternal suture, kate-pisternum with shallow excavation below thepoint. Anepisternum less than katepisternum,the latter with several trichia in lower part.Laterotergite bare, divided with katepister-num by anepimeron. Mediotergite bare. Me-tepisternum setose. Scutum with scattered se-tae. Scutellum with 6 long setae. Wing mem-brane with microtrichia and macrotrichiamostly in apical and hind area. Costa, R1, andR5 with long setae. Sc very short, ends free.R1 and R5 almost straight. RS base short,;0.53 r-m length, transverse; r-m fused withtb and m-cu, slightly oblique. Crossvein m-


cu twice the length of r-m. M2 probably ab-sent. M314 absent. M1 straight, weak.

MATERIAL: Holotype PIN 3426/256, sexunknown. Russia, Taimyr Peninsula, Nizh-nyaya Agapa, coll. 1973.

ETYMOLOGY: The species is an epithet inreference to the locality, the Nizhnyaya Aga-pa River.

COMMENTS: Structure of ocelli and a trans-verse RS base are apparent characters of ge-neric level, distinguishing the species fromT. sibirica, but incomplete preservation ofthe specimen from Nizhnyaya Agapa doesnot allow description of a new genus. Bothspecies share such synapomorphies as an in-complete eye bridge, proepimeron apex atthe episternal suture, setose metepisternum,and M314 reduced. Until more specimens ofthis peculiar group will be found and phy-logenetic analysis is done we prefer to keepthe species in this genus.

FAMILY LYGISTORRHINIDAE EDWARDS, 1925

DIAGNOSIS: Eyes round; pedicel bears bris-tles apically; mouthparts reduced and formproboscis in some genera; Anepimeron fusedto katepisternum; Sc very short; R1 short;stem of M and M112 fork base absent; r-maligned with tb; M2 1 tb subhorizontal; hindcoxae short, hind tibiae and tarsi swollen;metepisternum with deep anterodorsal cleft;gonostyli of simple shape (presumably re-versed in Plesiognoriste and Protognoriste).

Archaeognoriste, new genus

DIAGNOSIS: Palpi long, three-segmented,mouthparts reduced. Base of RS present,crossvein r-m distinct. All the coxae approx-imately equal in length. Hind tibiae slightlyswollen. Gonostyli with long inner process.

TYPE SPECIES: Archaeognoriste primitiva,n.sp.

ETYMOLOGY: The name is a combinationof archaeos (Greek arxa , or ancient) andio§the genus name Gnoriste. The name is fem-inine.

Archaeognoriste primitiva, new speciesFigures 7, 8, Plate 1D


lotype)/1.47–1.61 mm (paratypes); winglength 5 0.98/0.95–1.10 mm. Head: Poste-rior surface flat, with scattered setulae. Fronsmembranous. Eyes large, hairy. Ocelli notseen. Scape and pedicel slightly wider thanflagellomeres, subconical. Flagellum 14-seg-mented, flagellomeres cylindrical, widthabout equal to length. Palpi 3-segmented,segments cylindrical. Penultimate segmentabout 0.753 the length of apical one and 23the length of antepenultimate. Thorax: Scu-tum irregularly covered with short setae andbearing long lateral and acrostichal setae.Scutellum with two pairs of long setae andseveral short ones. Antepronotum and pre-episternum with 6–8 long setae. Anepisternalsuture declines posteriorly. Anepisternumwith small cleft. Mediotergite setose, latero-tergites bare. Dorsal edge of mesepimeronprotruding toward laterotergite. Metepister-num with deep cleft, height more than width.Wing membrane without macrotrichia. C notreaching wing tip, ends one-fourth distancebetween tips of R5 and M1. All veins bearsetae. Sc very short, free. R1 about 0.63 thelength of wing. RS base distinct, at the levelof Sc tip. Crossvein r-m fused with tb,oblique. Stem of M and base of M1 and M2

fork absent. M314 base very weak. M andCuA veins weak. Legs: Fore coxae denselysetose, mid and hind coxae with setae on api-cal half, all coxae about equal in size. Tibialand tarsal trichia not in rows. Abdomen se-tose, with exception of 1st and 2nd sternites.Seventh sternite short, 8th small and retract-able. Ninth tergite small, rectangular, withwell-developed, short, one-segmented cerciapically. Gonocoxites straight. Gonostyli sa-berlike, thinner to apex, with one apical andone very long curved basal processes.

MATERIAL: Holotype AMNH Bu-1539,male; paratypes AMNH Bu-412a and Bu-412b, 2 males in the same piece, Bu-485, Bu-693, males. Myanmar: Katchin, from ambermines near Myitkyina.

ETYMOLOGY: The species epithet is fromLatin primitivus meaning ‘‘the first or earliestof its kind’’ and is a reference to the basalposition of the species within Lygistorrhinidae.

COMMENTS: Venation of the species is al-most identical to Palaeognoriste sp. (Gri-maldi and Blagoderov, 2001: figs. 5a, 6a),with bases of RS and r-m somewhat reduced


Figs. 7–11. Archaeognoriste, n.gen. and Lebanognoriste, n.gen. 7. A. primitiva, n.sp., holotypeAMNH Bu-1539 8. A. primitiva, male genitalia. 9. L. prima, n.sp. holotype AMNH JG268/1. 10. L.prima, male genitalia dorsally. 11. L. prima, male genitalia ventrally.


and shifted proximally. Characters such asreduced palpi, membranous frons, the wingvenation, simplified genitalic structure, and alarge metepisternum (presumably plesio-morphic) with deep anteriodorsal cleft obvi-ously refer the species to the Lygistorrhini-dae, but compared to more derived speciesthe new species has many plesiomorphies:presence of RS base and a distinct r-m, un-modified legs, and gonostyli with long pro-cesses.

Lebanognoriste, new genus

DIAGNOSIS: Wing membrane without mac-rotrichia, R1 short, crossvein r-m fused withbase of M314, fork of M314 and CuA not ses-sile; preapical palpomere attached at the tipof antepreapical, inner mid and hind tibialspurs longer than outer ones.

TYPE SPECIES: Lebanognoriste prima, n.sp.ETYMOLOGY: The name is a combination

of Lebanon, source country of the amber, andthe genus name Gnoriste. The name is fem-inine.

COMMENTS: Lebanognoriste together withArchaeognoriste form the most basal groupof Lygistorrhinidae. Synapomorphies such asrather long fore coxae; short vein R1; short,transverse RS1 section; reduced base of theM1 and M2 fork; and crossvein r-m shiftedproximally and aligned to M2 1 tb fusedvein, which is oblique or longitudinal andshifted proximally, undoubtedly ally thesetwo genera with the lygistorrhinids (see alsoAnalyses). Both genera lack autapomorphies,and comparison to recent representatives ofthe family indicates various plesiomorphies.

Lebanognoriste prima, new speciesFigures 9–11, Plate 1E

DIAGNOSIS: As for genus.DESCRIPTION: Body length 5 1.53 mm;

wing length 5 1.64 mm. Head: Postocciputflat, with several strong setae behind eyemargin. Eyes setose. Ocelli not visible. Scapeand pedicel twice as wide as flagellomeres,scape with one and pedicel with two apicalrows of setae. Flagellum with 14 cylindricalsegments, covered by hairs as long as 1/2flagellomere diameter. Face setulose. Onlythree apical segments of palpi visible; ante-penultimate and penultimate knoblike; apical

long, slender, straight, its length approxi-mately the combined length of preceding twopalpomeres. Thorax: Mesonotum with fewlong lateral and scattered shorter setae. An-terior parapsidal suture distinct. Scutellumwith 6 long setae and sparse short ones. Lat-erotergites and mediotergite bare. Wingmembrane without macrotrichia, microtrichianot arranged in rows. All longitudinal veinswith setae. Costa ends far beyond tip of R5.Humeral cross-vein transverse. R1 short,about 0.43 wing length. Sc short, ends in Rat middle of r-m. R5 slightly curved caudally.M1 slightly curved forward at tip. M forkbase and M stem absent. Crossvein r-m short,oblique, fused with base of M314. M314 andCuA stem short, but distinct. M314 almoststraight, CuA curved gently back. Legs: Foretibiae with simple anteroapical depressedarea bearing one row of short bristles. Midtibia with inner spur 1.53 as long as outerone, inner hind tibia spur 1.33 as long asouter one. Tibial setulae in distinct rows. Tib-ial spurs 2–33 the tibial diameter. Tarsalclaw with one tooth. Abdomen short, setose,with 7 visible segments. Genitalia rounded,rotated at 1808. Gonocoxites massive, fusedventrally. Gonostyli hemicylindrical, slightlycurved, shovel-like at apex. Ninth tergitelength equals the width, with two deep lateralincisions.

MATERIAL: Holotype AMNH JG268/1,male. Lebanon: near Jezzine, coll. Aftim andFadi Acra.

ETYMOLOGY: The species epithet is a Latinword primus meaning ‘‘first’’ and is a refer-ence to the oldest find of the family in thefossil record.

Plesiognoriste, new genus

DIAGNOSIS: Wing membrane without mac-rotrichia. Palpi short, two-segmented. Facewide, about one-third of head width. Eyes ofmale relatively small, setose. Proboscis notdistended. Laterotergites bare. All veins withsetae. R5 runs very close to R1. RS base ab-sent. M2 absent. M314 and CuA fork short.Middle and hind tibiae spurs almost equal inlength. Tibial setulae forming more or lessregular rows apically.

TYPE SPECIES: Plesiognoriste carpenteri,n.sp.


ETYMOLOGY: The genus name is a combi-nation of plesion (Greek plhs on, or neigh-ibor) and the genus name Gnoriste. The nameis feminine.

COMMENTS: See under Protognoriste.

Plesiognoriste carpenteri, new speciesFigures 12, 13, Plate 1F

DIAGNOSIS: Apical palpal segment curved,bulbous; flagellomeres longer than wide,anepisternite with small dorsal cleft; M314

gently curved, M314 and CuA stem weak-ened, fore and hind tibiae and hind femorabroadened.

DESCRIPTION: Body length 5 1.32 mm;wing length 5 1.27 mm. Head: Antennaeinserted near middle of head. Scape cylin-drical; pedicel subconical, apex wide withoblique margin. Flagellum 14-segmented,flagellomeres cylindrical, length of eachtwice the width, covered with setulae as longas flagellomere diameter. Face wide, pentan-gular, setose. Frons and occiput uniformly se-tulose. Three ocelli present, equal in size,distance from middle to lateral ocelli thesame length as from lateral ocellus to eyemargin (distance of lateral ocellus from eyemargin equal to diameter of ocellus). Eyessetose, slightly emarginate at antennal bases.Palpi with two visible segments; apical seg-ment slightly curved, bulbous, setulose; basalone with a few short setae. Thorax: Scutumwith long lateral and dorsocentral setae. Scu-tellum with 6 long setae. Antepronotum se-tose, divided from proepisternum by a dis-tinct suture. Anepisternum with small dorsalcleft. Mediotergite bare. Laterotergite notfully visible. Wing membrane without mac-rotrichia; microtrichia not arranged in regularrows. Costa ends slightly beyond tip of R5.All veins bear setae. Sc very short, ends free,humeral cross-vein almost transverse. R1

about 0.63 wing length. RS base absent. R5

runs very close to R1, virtually straight; baseweak, so that vein invisible proximal to Sctip. M1 weak, originates midway betweentips of R1 and R5. M2 and stem of M absent.M314 curved gently. CuA has distinct kink atbase of M314. M314 and CuA stem weak.Legs rather short and stout. Fore coxae withlong bristles on anterior edge. Fore tibia wid-ened. Hind coxae equal in length to middle

ones, but broader, with scattered setae. Hindfemora swollen, without ventral spines. Hindtibiae with apical comb of setae. Tarsal clawwith one tooth. Abdomen shorter thanwings, setose. Gonocoxites quadrate, flat, se-tose, with dorsal protruding appendage. Gon-ostyli simple, length ;33 the width, apicallywith two black inner spines and outer combof short setae.

MATERIAL: Holotype MCZC 6927, male.Canada: Manitoba, Cedar Lake, coll. F.M.Carpenter.

ETYMOLOGY: The species epithet is a pa-tronym for the late Professor Frank Carpen-ter of the MCZ, who collected this and manyothers Canadian amber specimens.

Plesiognoriste zherikhini, new speciesFigure 14, Plate 2A

DIAGNOSIS: Scape and pedicel not en-larged, flagellomeres short, eyes form incom-plete bridge, apical palpal segment rounded,M314 strongly curved, legs not modified.

DESCRIPTION: Body length 5 1.40 mm;wing length 5 1.22 mm. Head: Postocciputflat. Antennae inserted near middle of face.Scape and pedicel not enlarged, widthsequal to width of average flagellomere. Fla-gellum 14-segmented, flagellomeres cylin-drical, width about equal to length, setulose.Frons setose. Eyes setulose, emarginate atantennal bases, forming lobes toward mid-dle of head 4–5 facets wide, but dorsalbridge incomplete, separated by distancemore than 43 facet diameter. Ocelli three,equal in size to the distance from middle tolateral ocelli, twice the length as from lateralocellus to eye margin (distance of lateralocellus from eye margin equal to diameterof ocellus). Face setose. Palpi short, only 2visible, rounded segments, with numeroushairs. Clypeus bare. Thorax: Scutum withlong lateral, dorsocentral, and acrostichalsetae. Scutellum with 6 long setae. Anteriorparapsidal suture distinct. Suture betweenantepronotum and proepisternum oblique.Proepimeron touches mesepisternum at ane-pisternal suture. Anepisternal cleft narrow,oblique. Mediotergite and laterotergitesbare. Metepisternum touches katepisternum.Wing membrane without macrotrichia, withscattered microtrichia. All veins setose.


Figs. 12–14. Plesiognoriste, n.gen. 12. P. carpenteri, n.sp. holotype MCZC 6927. 13. P. carpenteri,male genitalia. 14. P. zherikhini, n.sp., holotype PIN 3311/664


Costa ends beyond tip of R5, one-third thelength between tips of R5 and M1. Sc veryshort, ends free. R1 short, about 0.5 winglength. R5 about 0.853 wing length, curvedslightly, runs very close to R1. M1 confinedto apical third of wing. RS base, M1 andstem of M absent. Crossvein r-m fused withR5, horizontal, fused with CuA base. CuAbase weak. Base of M314 and CuA fork atlevel of tip of R1. M314 curved and wellrounded. CuA with two straight segmentsforming obtuse angle. Legs: Coxae almostequal in length. Hind coxae bare at base.Fore tibia distally with apical hemisphericalanteroapical depression with comb of finesetae. Tarsal claw without teeth. Abdomenshorter than wings, with 7 visible segments,setose. Sixth and 7th segments twice asshort as 5th, 8th retracted. Cerci one-seg-mented.

MATERIAL: Holotype PIN 3311/664, fe-male. Russia: Taimyr Peninsula, Yantardakh,coll. 1971.

ETYMOLOGY: The species epithet is a pa-tronym in honor of the late Dr. VladimirZherikhin of the Paleontological Institute inMoscow, prominent paleoentomologist, whocollected the specimen.

Protognoriste, new genus

DIAGNOSIS: Wing membrane without mac-rotrichia, occiput without row of strong se-tae. Palpi short. Eyes bare. Face wide. Stemof M and M2 absent. RS base distinct. Cross-vein r-m meets M314 and CuA stem. CuAgently curved. Abdomen insertion broad.Fore tibiae shorter than femora.

TYPE SPECIES: Protognoriste amplicauda,n.sp.

ETYMOLOGY: The name is a combinationof protos (Greek pr to§, or first) and thevgenus name Gnoriste. The name is feminine.

COMMENTS: The following apomorphiccharacters refer the genera Plesiognoristeand Protognoriste to the Lygistorrhinidae:short palpi; small dorsal cleft of anepister-num; anepisternum and metepimeron not di-vided by laterotergite (possibly plesiomorph-ic); short, incomplete Sc; short R1; RS baseand M stem reduced; r-m horizontal andfused with R5; hind coxa smaller and broaderthan mid coxa; abdomen insertion narrow;

cerci simple. Face wide, with three ocelli, sit-uated almost in straight line, suggestive ofManotinae, but structures of palpi, katepi-sternum and metepimeron, and absence ofstrong setae, show no apomorphies with re-spect to that subfamily. Eye size, face width,vein vestiture, tibial spurs of equal size, andcomplicated shape of the gonostyli are moreprimitive conditions than occur in Recent ly-gistorrhinids, although some features of ve-nation and gonostyli are apomorphic. Theshort fork of M314 and CuA and absence ofa long proboscis are most similar to the ge-nus Seguyola Matile. It is possible that thesetwo genera should be treated as a separatesubfamily of Lygistorrhinidae. We prefer toexpand the definition of the family to includenewly described taxa.

Protognoriste amplicauda, new speciesFigure 15, Plate 2B

DIAGNOSIS: R1 length 33 that of r-m. Gon-ocoxite massive. Gonostyli straight, flatteneddorsoventrally.

DESCRIPTION: Body length 5 1.25 mm;wing length 5 1.11 mm. Head: Eyes setu-lose, setae very short. Ocelli three, equal, al-most in straight line, distance of lateral ocel-lus from eye margin equal to distance frommid ocellus to lateral one. Vertex and fronssetose. Scape very small. Pedicel spherical,obscure. Flagellum 14-segmented, flagello-meres cylindrical, lengths about equal towidths, setose. Palpi short, 3-segmented, api-cal segment rounded, penultimate cylindri-cal, length 23 the width. Face wide, quad-rate. Thorax: Scutum with lateral, dorsocen-tral and acrostichal setae. Scutellum withseveral long setae. Mediotergite with shorttrichia. Metepisternum with anterodorsalcleft, height about equal to width. Wingmembrane without macrotrichia. Costa endsbeyond tip of R5, at 1/6 the length betweentips of R5 and M1. Sc short, ends free at thelevel of RS base. Humeral cross-vein trans-verse. R1 short, about 0.43 wing length. R5

about 0.83 wing length, almost straight. RSbase very short, transverse. Crossvein r-mhorizontal, meets M314 and CuA stem, itslength 33 less than R1 length. M1 almoststraight, weakened at base. M2 and M stemabsent. Base of M314 and CuA fork between


Figs. 15–18. Protognoriste, n.gen. 15. P. amplicauda, n.sp., holotype PIN 3426/257. 16. P. goeleti,n.sp., holotype AMNH Bu-406. 17. P. nascifoa, n.sp., holotype AMNH Bu-434. 18. P. nascifoa, malegenitalia.

levels of RS base and R1 tip. CuA and M314

curved caudally. Legs: Coxae almost equalin length, hind coxae bare. Tibial setulae indistinct rows. Tibiae, especially fore, short.Tarsal claw with small obtuse tooth. Abdo-men as long as wing, with 8 segments, se-

tose. Eighth segment short, retracted. Gono-coxite massive. Gonostyli straight, flatteneddorsoventrally.

MATERIAL: Holotype PIN 3426/257, male.Russia: Taimyr Peninsula, Nizhnyaya Agapa,coll. 1973.


ETYMOLOGY: The species epithet is derivedfrom Latin words amplus meaning ‘‘large,distinguished’’ and cauda meaning ‘‘tail’’ inreference to large terminalia of the species.

Protognoriste goeleti, new speciesFigure 16, Plate 2C

DIAGNOSIS: R1 length 1.83 that of r-m.DESCRIPTION: Body length 5 1.21 mm;

wing length 5 0.95 mm. Head: ovate, height;1.53 width in profile. Occiput and postgenadensely setose. Antennae attached below themiddle of the head. Scape and pedicel sub-conical, wider than flagellum. Flagellomerescylindrical, length 1.1–1.53 the width. Eyessetose, ovate, without incision, facets round,close. Clypeus bare. Two palpomeres seen:basal one ovate, length 23 the width, apicalround, 43 shorter than basal. Thorax: Scu-tum irregularly setose. Anterior parapsidal su-ture distinct. Scutellum small. Anepisternumwith several setae. Proepimeron touches me-sepisternum at anepisternal suture. Anepister-num with deep cleft in the middle. Lateroter-gite shifted dorsocaudally, so that metepister-num contacts katepisternum for some dis-tance, not in a point. Laterotergites andmetepisternum with short hairs. Mediotergiteshort. Meron at mid coxae large. Wing mem-brane with microtrichia not arranged in rows.Costa ends beyond tip of R5, at one-fourthlength between tips of R5 and M1. Sc veryshort, ends free. Humeral cross-vein oblique.R1 short, about 0.43 wing length. RS baseshort, transverse. R5 about 0.83 wing length,almost straight. M1 slightly sigmoid, confinedto apical third of wing, ends at tip of wing.Stem of M and M2 absent. Crossvein r-m hor-izontal, very weak, meets M314 and CuAstem, its length 1.23 more than length of R1.M314 and CuA fork 1.73 stem length, its basebetween levels of RS base and tip of R1. M314

and CuA curved gently. Legs: Fore tibia1.253 shorter than fore femora. Tibial andtarsal setulae not in rows. Hind tibia with 15dorsal bristles. Tibial spurs 23 the diameterof tibia. Hind tibiae with apical comb of shortsetae. Fore coxae with dense anterior setae,mid and hind coxae with setae in apical part,hind coxae without posterior setae. Abdomeninsertion broad. Abdomen shorter than wings.

Cerci with large basal and small round apicalsegments.

MATERIAL: Holotype AMNH Bu-406, fe-male. Myanmar: Katchin, from amber minesnear Myitkyina.

ETYMOLOGY: The species epithet in honorof Mr. Robert Goelet, for his generosity infunding purchase of specimens and the au-thors’ work.

Protognoriste nascifoa, new speciesFigures 17, 18, Plate 2D

DIAGNOSIS: RS base and the base of theM314 and CuA fork very basally. R1 length43 r-m.

DESCRIPTION: Body length 5 1.09 mm;wing length 5 1.05 mm. Head: Occiput andfrons setose. Pedicel and scape wider thanflagellomeres. Flagellum 14-segmented, fla-gellomeres cylindrical, length about equal towidth. Face wide, quadrate. Clypeus setose,palpi 2-segmented, basal palpomere swollen,apical one very small. Eyes with incision,forming incomplete eye bridge (possibly ar-tifact, face deformed). Thorax: Proepimeroncontacts katepisternum. Anepisternum width1.53 height, with distinct cleft posteriorly.Laterotergites, mediotergite and metepister-num with very short trichia. Scutum with nu-merous erect setae. Metepisternum with longanteriodorsal process touching katepister-num. Wing: Sc very short, free. Costa pro-duced beyond R5 one-third distance betweenR5 and M1 apices. RS base situated proxi-mally, faint, oblique. R1 length 43 that ofr-m. Crossvein r-m weak, 43 shorter than R1.M1 originates at distal third of wing, curvedat base. Base of the M314 and CuA fork atthe level of RS base. Legs: Hind coxae short-er than mid ones. Fore coxae with long an-terior setae, mid and hind ones with apicalsetae. Tibial setulae not in rows except apicalhalf of hind tibiae. Hind tibiae long, swollensomewat at apex, with dorsal row of bristlesand apical comb of setae. Tibial spur length1.0–1.33 tibial diameter. Abdomen setose,segments short. Genital complex wider thanlong. Gonocoxites fused. Gonostyli with oneouter and two inner lobes.

MATERIAL: Holotype AMNH Bu-434,male. Myanmar: Katchin, from amber minesnear Myitkyina.


→

Figs. 19–24. Leptognoriste, n.gen. 19. L. davisi, n.sp., holotype AMNH Bu-126a. 20. L. davisi, wingof the holotype. 21. L. davisi, head of the holotype. 22. L. davisi, male genitalia of the holotype. 23. L.microstoma, n.sp., holotype AMNH Bu-429. 24. L. microstoma, male genitalia.

ETYMOLOGY: The species epithet derivedfrom U.S. National Science Foundation, agenerous sponsor of this and other fossil in-sect research at the AMNH.

Leptognoriste, new genus

DIAGNOSIS: Palpi short, 4-segmented.Mouthparts form short proboscis or reduced.Laterotergites and mediotergite setose. Wingmembrane with macrotrichia. Sc long, endsat C. Costa produced beyond R5, not reach-ing wing apex. M1 absent. M2 reduced at thebase. RS base transverse. M314 and CuA forkstalked. Hind coxae shorter than fore andmid ones. Male genitalia simple.

TYPE SPECIES: Leptognoriste davisi, n.sp.ETYMOLOGY: The name is a combination

of leptos (Greek lept s, or thin, lean) andothe genus name Gnoriste. The new name isfeminine.

Leptognoriste davisi, new speciesFigures 19–22, Plate 2E

DIAGNOSIS: The base of M314 and CuAfork at level of base of RS. Mouthparts formshort proboscis. M1 reaching wing margin.Ninth tergite small, longer than wider, rect-angular.

DESCRIPTION: Body length 5 1.94 mm (ho-lotype)/1.69 mm (paratype); wing length 51.26 mm (holotype)/1.27 mm (paratype).Head round with protruding ocelli. Ocelli intriangle, lateral separated from medial byocellus diameter and from eye margin by 2ocellus diameters. Eyes large, rounded, withlight emargination, with large facets. Clypeusnarrow, triangular, setose. Palpi 4-segmented,palpomere length ratio 1:2:2:3. Apical pal-pomere attached preapically. Short proboscis,one-half head height. Wing length equal toabdomen length. Sc produced slightly be-yond RS base. R1 length 1.23 r-m. M2 orig-inates in distal half of wing. Base of M314

and CuA fork at level of RS base. Macrotri-chia numerous in basal part of wing. Tho-rax: Scutum with long lateral, dorsocentral

and acrostichal and shorter irregular setae.Anterior parapsidal suture distinct. Antepron-otum and proepisternum setose. Anepister-num wider than its height. Proepimerontouches katepisternum slightly below anepi-sternal suture. Mesepisternum with long an-terodorsal process, touching anepisternum.Laterotergites with row of long setae. Me-diotergite irregularly setose with short setae.Legs: Coxae with relatively short setae. Hindtibiae with bristles in dorsal row, 1.53 longerthan femora. Abdomen setose, except firststernite. Abdomen insertion very narrow.Tergite 8 shorter than sternite. Tergite 9 nar-row, rectangular with numerous setae atapex. Gonocoxites rather slender. Gonostylislightly curved.

MATERIAL: Holotype AMNH Bu-126a,male; paratype AMNH Bu-126b, male, in thesame piece. Myanmar: Katchin, from ambermines near Myitkyina.

ETYMOLOGY: The species epithet is a pa-tronym for Mr. Jim Davis, who supplied theAMNH with fossiliferous amber from Myan-mar.

Leptognoriste microstoma, new speciesFigures 23, 24, Plate 2F

DIAGNOSIS: Mouthparts reduced. R1 shorterthan r-m. M2 weak at apex. Base of M314 andCuA fork beyond the level of RS base. Male8th tergite wider than long, rounded.

DESCRIPTION: Body length 5 1.33 mm;wing length 5 1.24 mm. Head: Occiput withlong setae. Pedicel slightly wider then scapeand flagellum, flagellomeres barrel-shaped,as long as wide. Palpi 4-segmented, palpo-mere length ratio 1:2:2:3.5. Thorax: Scutumwith long lateral, acrostichal, and dorsocen-tral setae and short setae between rows. Pro-epimeron touches katepisternum below theepisternal suture. Anepisternum width aboutequal to height. Laterotergites with long se-tae. Mediotergite with several long setae ven-trally. Mesepisternum with long anterodorsalprocess, touching anepisternum. Wing length


1.53 the abdomen length. R1 slightly shorterthan r-m. Macrotrichia in basal part of wingabsent. M2 weak at apex. Base of M314 andCuA fork beyond the level of RS base. Legs:Fore coxae with dense long anterior setae,mid and hind ones with apical. Hind tibiaewithout dorsal bristles, only slightly longerthan femora. Abdomen. Tergite 9 wider thanlong, rounded on apex. Sternite I bare. Gon-ostyli curved at apex.

MATERIAL: Holotype AMNH Bu-429.Myanmar: Katchin, from amber mines nearMyitkyina.

ETYMOLOGY: The species epithet is a com-bination of micros (Greek mikroz, or small)and stoma (Greek st ma, or mouth), in ref-oerence to the reduced mouthparts.

FAMILY MYCETOPHILIDAE NEWMAN, 1834

SUBFAMILY MANOTINAE EDWARDS, 1925

Alavamanota Blagoderov and Arillo, 2002

Alavamanota Blagoderov and Arillo, 2002: 34.

DIAGNOSIS: Antepronotum and proepister-num completely divided; mediotergite andlaterotergites bare; wing membrane with orwithout macrotrichia; R1 relatively short,length of R1 about the length of r-m; R4 pre-sent; M314 and CuA fork with stem.

TYPE SPECIES: Alavamanota hispanica Bla-goderov and Arillo, 2002.

COMMENTS: Closest to the recent genusManota Williston, 1896, distinguished by hav-ing the fourth palpomere attached preapicallybut very close to the apex of the third; twocrossveins instead of one between R1 and R5

(RS1 and R4); tibial trichia irregularly ar-ranged; sternite 9 separate; sternite 8 without4 strong protuberances bearing long setae; andthe basal segment of each cercus small.

Alavamanota burmitina, new speciesFigure 25, Plate 3A

DIAGNOSIS: Flagellum compressed dorso-ventrally; mesonotum with long lateral setae;wing membrane with macrotrichia; length ofsmall radial cell 63 width; base of M1 and M3section weak; tibial setulae arranged in rows.

DESCRIPTION: Body length 5 2.50 mm (ho-lotype)/1.62–2.5 mm (paratypes); winglength 5 1.69 mm (holotype)/1.08–1.85 mm(paratypes). Head: Postocciput with row of

long protruding setae behind eye margin.Eyes slightly emarginate, densely setose, se-tae length 23 facet diameter. Facets round,densely set. Three ocelli in triangle, close toeach other. Frons and face setulose. Antennaeinserted above middle of head. Scape andpedicel subconical, with apical setae. Flagel-lum 14-segmented, flagellomeres com-pressed, widths 1.2–23 length, apical oneconical. Texture of flagellomeres polygon-like. Only 3 segments of palpi visible, ante-penultimate and penultimate with strong se-tae; penultimate 2.53 as long as precedingone, ovate; apical segment 1.73 length ofpenultimate one, narrow. Thorax: Scutumuniformly densely setose with short setae,bearing long lateral and posterior setae. An-terior parapsidal suture distinct. Suture be-tween antepronotum and proepisternum com-plete, both segments uniform and coveredwith bristles. Scutellum with long setae.Anepisternum wider than higher, with widedorsal cleft. Anepisternal suture declinesposteriorly. Katepisternum setose ventrally.Mediotergite and laterotergites bare. Met-episternum with several light hairs postero-ventrally. Wing membrane with macrotri-chia; microtrichia not arranged in rows. Cos-ta ends slightly beyond the tip of R5, C andR5 run very close to each other in apical part.R1, R5, r-m, M1, and M2 with setae ventrallyand dorsally. Sc very short, ends free. Hu-meral vein oblique. R1 about equal to lengthof r-m. Ratio of r-m and RS2 section is 1:1.4. Base of M1 and M2 fork at the level ofR4. Small radial cell with length 63 width.Crossvein r-m horizontal, fused to tb, meetsMA. M314 and CuA fork long, but not sessile.CuA curved caudally. Legs. Hind coxae bearapical only, not posterior setae. Tibial spurlengths ca. 43 tibial diameter. Tibial trichiain rows. Abdomen setose, with 6 visible seg-ments. Gonocoxites fused, lighter then ab-domen, setose, swollen. Gonostyli not seen.

MATERIAL: Holotype AMNH Bu-1271,male; paratypes AMNH Bu-428a, Bu 279a,males. Myanmar: Katchin, from amber minesnear Myitkyina.

SUBFAMILY SCIOPHILINAE WINNERTZ, 1863

TRIBE SCIOPHILINI WINNERTZ, 1863

Neuratelia Rondani, 1856Neuratelia Rondani, 1856: 195.Anaclinia Winnertz, 1863:770.


Figs. 25, 26. Alavamanota and Neuratelia. 25. A. burmitina, n.sp., holotype AMNH Bu-1271. 26.N. maimecha, n.sp., holotype PIN 3311/661.


Fig. 27. Allocotocera burmitica, n.sp., holo-type AMNH B-056, reconstruction of wing ve-nation.

Proanaclinia Meunier, 1904:145.Odontopoda Aldrich, 1897: 187.

DIAGNOSIS: As given by Vockeroth (1972):Tibiae with distinct bristles, length of tibialspurs twice the tibia diameter; anepisternum,mesepimeron, and metepisternum bare; wingmembrane with macrotrichia; sternite 8 ofmale large; gonocoxites partly or completelyfused ventrally, gonostyli complex, subdivid-ed or with elaborate processes.

TYPE SPECIES: Mycetophila nemoralis Mei-gen, 1818: 256 (orig. des.).

COMMENTS: Two species of the genus weredescribed from Baltic amber (Meunier, 1904)

Neuratelia maimecha, new speciesFigure 26, Plate 3B

DIAGNOSIS: Sc short, ends before RS base;length of RS1 section 2.53 that of crossveinr-m; base of M314 and CuA fork at the middleof r-m; mediotergite with sparse, fine trichia.

DESCRIPTION: Body length 5 2.11 mm(rest); wing length 5 2.05 mm. Head: Fla-gellum 14-segmented, flagellomeres cylindri-cal, with length about equal to width. Scapeand pedicel rounded. Mouthparts form pro-boscis slightly shorter than head height. Palpi4-segmented, basal and antepenultimate seg-ments oval, penultimate and apical segmentbacilliform, apical slightly longer and nar-rower than penultimate, length ratio 1:3:4:5.5. Thorax: Scutum with numerous, short,scattered setae and long lateral ones. Ante-pronotum with three setae, proepisternumwith five. Anepisternum bare, with deep cleftin posterior part. Anepisternal suture declinesposteriorly. Metepisternum quadrate, touch-ing katepisternum. Laterotergites with sev-eral setae, mediotergite with sparse, fine tri-chia caudally. Wing membrane with micro-trichia and few macrotrichia. Costa ends be-yond tip of R5. Sc meets C just before RSbase. Sc2 absent. Longitudinal veins with se-tae. M314 and CuA base slightly distad of M3base. Legs: Tibiae with distinct bristles. Foretibia longer than first tarsomere.

MATERIAL: Holotype PIN 3311/661, end ofabdomen not preserved, sex unknown. Rus-sia: Taimyr Peninsula, Yantardakh, coll.1971.

ETYMOLOGY: Species epithet is derivedfrom the Maimecha river on the Taimyr Pen-

insula, Siberia, where the amber deposit islocated.

Allocotocera Mik, 1886

Eurycera Dziedzicki, 1885:166.Allocotocera Mik, 1886: 102.Euryceras Marshall, 1896: 291.

DIAGNOSIS: Laterotergites and mediotergitesetose. Wing membrane with macrotrichiaand with or without microtrichia. Sc2 preap-ical or medial. Base of M314 and CuA forkbefore the base of the fork of M1 and M2.

TYPE SPECIES: Eurycera flava Dziedzicki,1885: 167 [5 pulchella (Curtis, 1837)] (bymonotypy).

COMMENTS: Another fossil species occursin Lower Cretaceous Spanish amber (Bla-goderov and Arillo, 2002).

Allocotocera burmitica, new speciesFigure 27, Plate 3C

DIAGNOSIS: Wing membrane with macro-and microtrichia. Segment M3 2.53 lengthof r-m, 0.33 that of M fork. Sc ends at levelof RS base; Sc2 in the apical one-fourth ofSc, distal to midpoint of Sc.

DESCRIPTION: Body length 5 2.86 mm;wing length 5 2.46 mm. Head: Flagellum14-segmented, thickened to apex. Flagello-meres almost equal in length, length of thefirst one 23 its width, apical one 53 itswidth. Scape and pedicel subconical, withsmall apical setae, 23 wider than flagello-meres. Eyes pubescent. Two palpomeresseen, cylindrical, length ca. 43 the width.Thorax: Scutum irregularly covered withshort setae and with strong, long lateral setae.Antepronotum with long setae. Scutellumwith two pairs of long setae and several more


short ones. Laterotergites and mediotergitewith setae. Metepisternum with two short se-tae posteroventrally.

Wing membrane clear, with macro- andmicrotrichia. Costa ends at tip of R5. Scmeets C beyond RS base. Sc2 at level of baseof section M3. Section of RS1 transverse. R1

rather short, about 0.73 wing length. R5

curved caudally, not reaching wing apex.Crossvein r-m approximately 2.53 as long assection RS1 and 2.53 length of section M3.Fork of M1 and M2 3.33 as long as sectionM3. Base of M314 and CuA fork slightly be-fore base of section M3. Legs: Hind coxaewith long posterior setae. Fore tibiae longerthan first tarsomere. Cerci 1-segmented, 23as long as 8th tergite, setose.

MATERIAL: Holotype AMNH B-056, fe-male. Myanmar: Katchin, from amber minesnear Myitkyina.

ETYMOLOGY: The species epithet is a ref-erence to the country of origin of the amber.

COMMENTS: The species is very close to A.xavieri, but differs in having a shorter Scvein and a longer M1 and M2 fork.

Pseudomanota, new genus

DIAGNOSIS: Infraorbital setae absent. Threepalpomeres, penultimate one with long thinapical appendage. Antepronotum wide. Pro-episternum rodlike. Laterotergites and me-diotergite bare. Sc short, free. Wing mem-brane with macrotrichia. Base of M1 and M2

fork reduced. M1 reaches wing margin beforewing apex. M314 free at the base. Gonostylisimple, slightly curved.

TYPE SPECIES: Pseudomanota perplexan.sp.

ETYMOLOGY: The genus name is a combi-nation of pseudos (Greek ce dos, or a lie, ayfraud) and the genus name Manota. Thename is feminine.

COMMENTS: Some structures, especially thewide quadrate face, long and flexible apicalpalpomere, short Sc and R1, horizontal r-m,and reduced venation are like Manotinae.Also, unlike typical Sciophilinae, the newgenus has laterotergites and especially themediotergite bare. Nevertheless, the genus isattributed to the subfamily Sciophilinaebased on a long, rodlike proepimeron, therather distal position of humeral vein, and

long and slender gonocoxites (which are notknown for any manotines). Within Sciophil-inae the new genus might be close to theAzana group of genera (Matile, 1998), dem-onstrating reduction of median veins. All ofthe genera in the group have reduced vena-tion, with CuA simple and not forming a forkwith M314. Matile (1998) supposed that M314

in this group was reduced completely, whilethe fork of M1 and M2 lost its base. Jugdingon the position of vein apices, at least Par-atrizygia, Neoaphelomera, and Neotrizygiamay have lost M2, and an incomplete veinbetween M1 and CuA may be homologous toM314. The new species has venation moreprimitive than the genera in the Azana group,preserving all the longitudinal veins, al-though without bases.

Pseudomanota perplexa, new speciesFigures 28–30, Plate 3D

DIAGNOSIS: As for the genus.DESCRIPTION: Body length 5 1.44 mm;

wing length 5 1.07 mm. Head: Eyes large,setulose, with large facets. Occiput and post-gena setose, without erect bristles. Ocelli notseen. Antennae attached above middle ofhead. Frons, face and clypeus setose. Pedicelsubconical. Flagellum 14-segmented, slightlythicker toward apex; flagellomeres cylindri-cal, width 1–1.53 length. Three palpomeresseen, basal one heart-shaped, penultimateone with long thin apical appendage aboutone-third its length, apical one long and slen-der. Thorax: Antepronotum and proepister-num wide, completely divided, setose. Pro-epimeron narrow, rodlike. Scutum with lat-eral, acrostichal, and dorsocentral setae withbare strips in between. Anterior parapsidalsuture distinct. Anepisternal suture declinesposteriorly. Anepisternum with wide cleft.Metepisternum with long, narrow anterodor-sal process touching laterotergite. Lateroter-gites and mediotergite bare. Wing membranewith macrotrichia and microtrichia. Humeralvein slightly distad of MA. Costa extends toR5 apex at two-thirds distance between tipsof R5 and M1, not reaching wing apex.Length of R1 is 0.63 wing length. Small ra-dial cell length 53 width. M1 originates atapical one-third of wing, reaching wing mar-gin before wing apex. M2 originates at apical


Figs. 28–30. Pseudomanota perplexa, n.sp. 28. Holotype AMNH Bu-599a. 29. Male genitalia dor-sally. 30. Male genitalia ventrally.

half of wing. M314 free at the base. M1, M2,M314, and CuA apices weak. Legs: Tibial se-tulae not arranged in rows except on apex ofmid and hind tibiae. Tibial spurs ca. 3.53tibia diameter. Hind tibia with a dorsal rowof bristles. Abdomen setose, with 6 seg-ments visible. Genital complex as long as4th–6th segments combined, rotated 1808.Tergite 9 small, apex round, covering basalpart of gonocoxites only. Aedeagus with twoteeth apically. Gonocoxites separated, mas-sive, setose. Gonostyli simple, sticklike,slightly curved, apically flat, shovel-like.

MATERIAL: Holotype AMNH Bu-599a,male. Myanmar: Katchin, from amber minesnear Myitkyina.

ETYMOLOGY: The species epithet is fromthe Latin word perplexus, meaning ‘‘mud-dled, intricate’’, in reference to the intriguingcombination of sciophiline and manotinecharacters.

TRIBE GNORISTINI EDWARDS, 1925

Apolephthisa Grzegorgzek, 1885

Apolephthisa Grzegorgzek, 1885: 205.

DIAGNOSIS: Mediotergite bare; lateroter-gites setose; Sc meets C; R4 present; base ofM314 distad of r-m, faint; M2 several timeslonger than r-m.

TYPE SPECIES: A. rara Grzegorgzek, 1885:


Figs. 31–34. Apolephthisa and Synapha. 31. A. bulunensis, n.sp., holotype PIN 3963/4, reconstruc-tion of wing venation. 32. Male genitalia. 33. S. longistyla, n.sp., holotype MCZC 6944, reconstructionof wing venation. 34. Male genitalia.

206 (by monotypy) [5 subincana (Curtis,1837) (Sciophila)]

COMMENTS: This genus is widely distrib-uted in the Holarctic but is of low diversity,including only one described Palaearctic andone undescribed Nearctic species (Vocker-oth, 1981). Apolephthisa mesozoica is de-scribed from the Lower Cretaceous of Mon-golia (Blagoderov, 1998a), and one unde-scribed specimen is known from PalaeoceneSakhalin amber (Blagoderov, in prep.).

Apolephthisa bulunensis, new speciesFigures 31, 32, Plate 3E

DIAGNOSIS: Sc short, ends before RS base.Gonostyli with distinct rounded apical lobeand long inner process.

DESCRIPTION: Body length 5 2.24 mm;wing length 5 1.46 mm. Head: Eyes slightlyemarginate near antennal base. Flagellumwith 14 cylindrical flagellomeres, covered bytrichia equal to 0.53 flagellomere diameter.Palpi rather long; antepenultimate segmentoval, broadened; penultimate and apical seg-ments long and slender, together equal to forecoxa in length; apical segment slightlyclubbed at apex, with several short setae. Pal-pomere length ratio 1:2:4. Thorax: Scutumwith lateral, dorsocentral, and acrostichalrows of short setae. Mediotergite bare.Wing: Veins R1, R5, M1, and M2 with setae.Sc meets C before RS base. R5 almoststraight. Costa ends beyond tip of R5 one-fifth the distance between tips of R5 and M1.


M1, M2, and M stem weak. Base of M314 ab-sent. Legs: Hind coxae without posterior se-tae. Abdomen setose, with 7 visible seg-ments, 8th one small and retracted. Gono-coxites long, stout, thinner apically. Gono-styli with rounded apical lobe, bearingdorsoventral row of bristles and long innerprocess, ending in a short spur.

MATERIAL: Holotype PIN 3963/4, male.Russia: Taimyr Peninsula, Bulun, coll. 1976.

ETYMOLOGY: The species epithet is topo-nymic.

COMMENTS: Given the current low diver-sity of the genus, it would seem highly im-probable for three fossil species to occur un-less the genus was more diverse and abun-dant in the past.

Synapha Meigen, 1818

Synapha Meigen, 1818: 227.Sinapha Rondani, 1856: 196, misspelling.Empalia Winnertz, 1863: 762.

DIAGNOSIS: As given by Freeman (1951):Setae on stems and forks of both M and Cupresent, flagellomere lengths not more thanwidth, Sc long, penultimate palpomere short-er than apical and antepenultimate ones, la-terotergites bare or setose, midtibial organpresent.

TYPE SPECIES: S. fasciata Meigen, 1818(by monotypy).

COMMENTS: About 25 extant species ofSynapha are known from all biogeographicregions. Two fossil species of the genus weredescribed from Eocene Baltic amber (Meu-nier, 1904) and the Lower Cretaceous ofMontsec, Spain (Blagoderov and Martınez-Delclos, 2001). The related genus Austrosy-napha Tonnoir, 1929 has 23 species fromSouth America, which were cataloged by Pa-pavero (1977b); 7 occur in New Zealand(Tonnoir and Edwards, 1927) and one isfrom Australia (Tonnoir, 1929).

Synapha longistyla, new speciesFigures 33, 34, Plate 3F

DIAGNOSIS: Distinguished from living andfossil members by the structure of genitalia,having gonostyli very long and with numer-ous inner spurs.

DESCRIPTION: Body length 5 2.15 mm;wing length 5 1.58 mm. Head round. Eyes

large, rounded, with round facets. Flagellum14-segmented, flagellomeres cylindrical,length of each about 1.53 its width. Two pal-pomeres seen. Penultimate 0.73 length ofapical one, which is slender and flexible.Thorax: Scutum irregularly setose. Medi-otergite and laterotergites bare. Wing: R1

and R5 almost straight, with setae, remainingveins bare, except apex of M314, a few setaepresent on M2 and CuA1 dorsally. Length ra-tio of sections RS1, RS2, and RS3 is 1:1.4:13. Crossvein r-m slightly sigmoid, its length23 length of M3 section. Base of M314 ab-sent, the vein originates at level of M3 mid-point. CuA and M314 gently curved caudally.Legs: Midtibial organ absent. Abdomenwith 6 visible segments, 7th and 8th seg-ments small and retractable. Gonocoxitesfused. Gonostyli long, straight, with numer-ous stiff spines arranged in rows on innersurface and several dark, curved ones atbase.

MATERIAL: Holotype MCZC 6944, male,Canada: Manitoba, Cedar Lake.

ETYMOLOGY: The species epithet refers tothe long gonostyli of the species.

COMMENTS: This species lacks distinct syn-apomorphies of Synapha (i.e., midtibial sen-sory organ and short gonostyli) as well asstrong setae on the medial and cubital forks.The fossil has moderately long flagellomeresand subequal palpomeres, considered to besynapomorphies of Austrosynapha (see table2). Long, slender gonostyli, which are char-acteristic for Austrosynapha and many otherMesozoic fungus gnats, is considered to beplesiomorphic. Most likely, the new speciestogether with the Lower Cretaceous S. rubie-sensis Blagoderov and Martınez-Delclos,2001 represent a Synapha-Austrosynaphastem group, although monophyly of thegroup and included genera is still to be es-tablished. Matile (1991) noted that Synaphawas probably polyphyletic. The definition ofboth genera is vague, and often characters ofdifferent subgroups overlap. For example,species of A. (Paraaustrosynapha) Durethave short flagellomeres, while species of A.(Neoaustrosynapha) Duret retain some mac-rotrichia on the medial fork (Duret, 1980).

Dziedzickia Johannsen, 1909Hertwigia Dziedzicki, 1885: 166 (preocc. Schmidt,

1880).


TABLE 2Comparison of Synapha, Austrosynapha, and A. longistyla n.sp.

Dziedzickia Johannsen, 1909: 44.

DIAGNOSIS: After Freeman (1951): Eyesemarginate at level of antennae; 3 ocelli in astraight line; laterotergal hairs present or ab-sent; wing membrane without macrotrichia;Sc ending at R before or beyond base of RS,Sc1 may be present as a short stump; base ofM314 and CuA fork considerably basal to M1

and M2 fork; R4 may be present or absent.TYPE SPECIES: Hertwigia marginata Dzied-

zicki, 1885: 165 (by monotypy).COMMENTS: About 50 extant species of the

genus are described, mostly from the NewWorld, especially South America, with sev-eral species from Holarctic and Afrotropics.Four species are known from Baltic amber(Meunier, 1917a, 1917b, 1922) and one fromthe Oligocene of Rott (Statz, 1944). Someauthors (Chandler, 1999; Hutson, 1979; Ma-tile, 1992; Vockeroth, 1980) noted that thegenus in present definition was plausiblypolyphyletic since some of the species be-long to other gnoristine genera, such as Syn-temna Winnertz, 1863 and HadroneuraLundstrom, 1906.

Dziedzickia nashi, new speciesFigures 35, 36, Plate 4A

DIAGNOSIS: Laterotergites setose. Forkedveins without setae. Tergite 9 short. Gono-coxites simple. Gonostyli with strong innerprocess.

DESCRIPTION: Body length 5 2.96 mm (ho-lotype)/2.92–3.37 mm (paratypes); winglength 5 2.50 mm (holotype)/2.28–3.32 mm(paratypes). Brown gnats, thorax dark brown,

occiput greyish brown, tibiae and tarsi yel-lowish. Head: Eyes large, bare, slightlyemarginate; three ocelli in line. Median ocel-lus slightly sunken, distance from middle tolateral ocellus and from lateral one to eyemargin ;1.53 ocellus diameter. Scape andpedicel shorter than flagellomeres, bare. Fla-gellomeres cylindrical, length ;1.53 width,length of apical one ;23 the width. Threepalpomeres seen, short, gradually thinner to-ward apex. Mouthparts less than one-fourthheight of head. Thorax: Mesonotum veryconvex, with dorsocentral, acrostichal, andlateral setae well developed. Antepronotumand laterotergite setose, other sclerites bare.Proepimeron thick, short, touches katepister-num. Anepisternal suture declines posterior-ly. Wing membrane without macrotrichia. Scends at R1 beyond RS base. Sc1 absent. RS1,RS2, and RS3 ratio 1:1.2–1.3:11–15. M3length 23 that of r-m crossvein. Base ofM112 fork at level of R4. Base of M314 andCuA fork before base of r-m. Abdomen se-tose, 7th segment 23 shorter than 6th. Male:Gonocoxites massive, bristly, separated ven-trally by deep cleft. Ninth tergite short, trans-verse, covers only base of gonocoxites. Gon-ostyli with a strong, thin, dark processpointed inward. Female with cerci 2-seg-mented. Basal segment 2.53 apical one, cy-lindrical. Apical segment rounded.

MATERIAL: Holotype AMNH NJ-117a,male. Paratypes: AMNH NJ-117b, c, e, m,females, NJ-117d, g, h, males, NJ-117f(wing), all in the same piece of amber. USA:New Jersey, Sayerville, coll. P. Nascimbene.See figure 38 for a map of syninclusions.


Figs. 35–37. Dziedzickia and Saigusaia. 35. D. nashi, n.sp., holotype AMNH NJ-117a. 36. Malegenitalia of the holotype. 37. Saigusaia pikei, n.sp., holotype TMPD P79.15.7.21

ETYMOLOGY: The species epithet is patron-ymic for Mr. Paul Nascimbene, who collect-ed the piece and who has diligently preparednumerous amber specimens for the AMNH.

COMMENTS: Differs from all other speciesby the structure of male genitalia. A thor-ough comparision will be possible only afterrevision of the genus.


Fig. 38. Map of AMNH NJ-117 syninclusion.

TABLE 3Comparison of Boletina, Saigusaia, and Saigusaia pikei n.sp.

Saigusaia Vockeroth, 1980

Saigusaia Vockeroth, 1980: 541.

DIAGNOSIS: As given by Vockeroth (1980):Face weakly sclerotized, bare; laterotergitesbare; metepisternum with short fine hairs;prosternum bare; sternites of abdomen withmedian fold line. Male with tergite 7 andsternite 7 haired; subequal in length, two-thirds length of segment 6; tergite 9 not fusedwith gonocoxites. Female with sternite 8deeply emarginate posteriorly.

TYPE SPECIES: Boletina cincta Johannsen,1912: 270 (by original designation).

COMMENTS: Three species of the genus areknown from Eastern North America, Europeand Japan (Khonsu), and Taiwan and Nepal.

Saigusaia pikei, new speciesFigure 37, Plate 4B

DIAGNOSIS: Distinguished from the livingspecies by Sc long, meeting C just before RSbase.

DESCRIPTION (see also table 3): Bodylength 5 2.84 mm, wing length 5 1.80 mm.Head: Ocelli three, each slightly raised, veryclose. Eyes pubescent. Vertex and clypeussetulose. Flagellum 14-segmented, flagello-meres cylindrical, with length slightly greaterthan width. Mouthparts form short proboscis,length is one-half the head height. Only threepalpal segments seen: antepenultimate seg-ment swollen, penultimate and apical onesnarrower, bacilliform. Length ratio 1:1.7:2.7.Thorax: Antepronotum with two long setae.Scutum with strong, curved lateral, dorso-central and acrostichal setae. Mediotergiteand laterotergites bare. Scutellum with 5pairs of rather short setae. Metepisternumwith two long setae dorsally. Wing: Costaends beyond tip of R5, one-third distance be-tween tips of R5 and M1. Stem of R, R1 andR5 with rows of dorsal and ventral setae. R1

and R5 straight. Sc meets C just before RSbase. Sc2 apical, proximal to base of M3.Crossvein r-m 1.53 length of RS1 and 0.33length of M3 section. Fork of M1 and M2

2.73 length of M3. M1 and M2 fork campan-ulate, veins straight, with a few ventral setaeapically. Base of M314 and CuA fork proxi-mal to base of M1 and M2 fork, M314 and


CuA curved caudally. Legs: Tibial bristles23 tibial diameter. Tarsal claw with onetooth. Abdomen: Sternite 8 with two trian-gular, rounded, caudal lobes covered by longsetae. Cerci two-segmented. Basal segmentlarge and broad, apical one 0.53 the lengthand 0.53 the width of basal one; both setose.

MATERIAL: Holotype TMPD P79.15.7.21,female. Canada: Alberta, Grassy Lake, coll.T. Pike.

ETYMOLOGY: The species epithet is patron-ymic for Dr. T. Pike, who collected the spec-imen.

COMMENTS: This species has almost all thefeatures of Saigusaia (table 3, see also Vock-eroth, 1980), so despite the long Sc vein itsplacement in the genus is virtually certain.

Syntemna Winnertz, 1863

Syntemna Winnertz, 1863: 767.

DIAGNOSIS: Mediotergite bare; lateroter-gites setose; wing membrane with macrotri-chia; Sc meets R; radial cell very small, su-bquadrate; base of M314 and CuA fork prox-imal to base of M1 and M2 fork; segment 7of abdomen reduced.

TYPE SPECIES: S. morosa Winnertz, 1863:767, by monotypy.

COMMENTS: About 20 extant species areknown from the Holarctic (Zaitzev, 1994),with an additional 12 species described fromBaltic amber (Meunier, 1904, 1917a, 1922)and 4 from the Lower and Upper Cretaceousof northeast Asia (Blagoderov, 1995, 1998a,2000).

Syntemna fissurata, new speciesFigures 39, 40, Plate 4C

DIAGNOSIS: Sc ends at R at the base of RS;R4 present; crossvein r-m 0.53 length of M3section; base of fork of M314 and CuA prox-imal to the base of M1 and M2 fork.

DESCRIPTION: Body length 5 3.06 mm,wing length 5 2.28 mm. Head not fully vis-ible. Flagellum 14-segmented, flagellomeresbarrel-shaped, length of each 1.53 the width,setose, length of setae about equal to flagel-lomere width. Thorax: Scutum with strong,long setae in lateral, dorsocentral, and acros-tichal rows, with short setulae. Anteprono-tum, proepisternum, laterotergites, and me-

diotergite setose. Wing membrane with mi-crotrichia and short macrotrichia. Costa endsbeyond the tip of R5, one-fourth distance be-tween tips of R5 and M1. Sc meets R justbefore base of RS. Sections of RS1 and RS2equal and 1.53 length of r-m. Crossvein r-m0.53 length of M3 section. Fork of M1 andM2 3.53 length of its stem (M3). Base offork of M314 and CuA proximal to the baseof M2 section. Legs: Inner surface of midtibia with fissurelike sensory pit in apicalthird. Tarsal claws with two short teeth at thebase. Abdomen setose, with 7 visible seg-ments, 7th one very small. Cerci 2-segment-ed, basal segment roundish, with numerouslong trichia; apical segment bacilliform, withlength twice the width, bare. Lateral lobes of8th sternite setulose.

MATERIAL: Holotype TMPD P83.15.3.8,female. Canada: Alberta, Grassy Lake, coll.T. Pike

ETYMOLOGY: The species epithet is theLatin word fissures, meaning ‘‘full ofcracks’’, in reference to the state of preser-vation of the specimen.

COMMENTS: The new species differs fromall other known Mesozoic species in havingsection M3 long, at least twice the length ofcrossvein r-m.

Gregikia, new genus

DIAGNOSIS: Costa virtually ends at the apexof R5. Sc bare. Sc2 apical. Section M2 setose.R5 not reaching wing apex. M3 weak. Ac-rostichal setae absent. Tergite 9 short. Gon-ostyli well developed.

TYPE SPECIES: Gregikia pallida, n.sp.ETYMOLOGY: The name is derived from

some letters commonly used in names ofgenera in this complex. The name is femi-nine.

COMMENTS: The genus is very similar toPalaecomoptera Blagoderov, 1997, five spe-cies of which were described from the LowerCretaceous of Transbaikalia and Mongolia(Blagoderov, 1997, 1998a), as well as tothree monotypic genera: Grzegorzekia Ed-wards, Creagdhuhia Chandler, and Phoeni-kiella Chandler. It is distinguished from Pa-laecomoptera by R5 not reaching wing apex,costa not produced after R5 apex, and baseof fork of M1 and M2 weak. The genus dif-


Figs. 39–43. Syntemna and Gregikia. 39. S. fissurata, n.sp., holotype TMPD P83.15.3.8, left wing.40. Female genitalia of the holotype. 41. G. pallida, n.sp., holotype AMNH NJ 117j. 42. Male genitaliadorsally. 43. Male genitalia ventrally.


fers from Grzegorzekia, Creagdhuhia, andPhoenikiella by a short apical palpomere, ab-sence of acrostichal setae, weak M stem andbase of fork of M1 and M2, absence of setaeon A, from Grzegorzekia also by having Scbare. Gonocoxites of the new genus alsohave two lobes as in the last three genera,but the gonostyli are better developed. Thisstate is undoubtedly more plesiomorphic thanthe highly modified genital complex of Grze-gorzekia, Creagdhuhia and Phoenikiella.Species of Palaecomoptera have gonostylideveloped as well, but some (P. shcherba-kovi, P. lukashevichae) have tergite 9 large,fully covering the gonocoxites.

Gregikia pallida, new speciesFigures 41–43, Plate 4D


lotype)/3.09–3.65 mm (paratypes); winglength 5 3.26 mm (holotype)/2.36–3.76 mm(paratypes). Head: Vertex setose. Scape andpedicel small. Flagellum 14-segmented. Fla-gellomeres cylindrical, length 23 the width,covered by trichia, length of trichia aboutone-half flagellomere width. Only three seg-ments of palpi visible; segments short, nearlycylindrical, combined length slightly lessthen head height. Thorax: Scutum with erectsetae, arranged in rows with wide bare stripsbetween them, acrostichal setae absent. An-tepronotum and proepisternum setose. Pro-epimeron touches mesepisternum at anepis-ternal suture. Scutellum with four pairs ofsetae. Anepimeron very narrow ventrally, sothat laterotergite touches katepisternum. Ka-tepisternum larger than anepisternum. Me-diotergite, laterotergites, and metepisternumbare. Wing: Costa ends slightly beyond tipof R5. Sc meets C at level of RS base. Rstem, R1, R5, and distal parts of M1 and M2

with setae. R5 not reaching wing apex. Sec-tion RS1 oblique. R4 transverse. Ratios oflengths of RS1, RS2, and RS3 is 1:1.5:12–15. Length of crossvein r-m about equal toRS1 and 0.253 M3 section. M1 subparallelto R5; veins M1, M2, M314, and CuA diver-gent. Base of fork of M314 and CuA proximalto M3 base. M314 and CuA curved gently.M1, M2, M314, and CuA with a few setae onapical part. Legs: Hind coxae with long dor-

solateral setae. Mid tibiae without sensorypit. Tarsal claw with one basal tooth. Ab-domen densely setose. Segments 7 and 8short, about one-half length of segment 6.Tergite 9 short, does not cover gonocoxites.Gonocoxites densely setose, with long, slimapical and basal lobes, pointed caudally andbearing numerous inner setae. Gonostyliovate, bare, with strong dark apical process-es.

MATERIAL: Holotype AMNH NJ 117j,male. Paratypes: AMNH NJ 117i, male (seefig. 74 for the scheme of syninclusions), coll.P. Nascimbene; AMNH NJ 871a and NJ871b, sex unknown, coll. K. Luzzi. USA:New Jersey, Sayreville.

ETYMOLOGY: The species epithet is a Latinword pallidus meaning ‘‘pale’’ in referenceto state of preservation of the specimen.

Gaalomyia, new genus

DIAGNOSIS: Palpi 3-segmented, short. Scu-tum with long lateral, dorsocentral, and ac-rostichal setae. Laterotergites bare. Sc endsat R. Base of fork of M314 and CuA at thelevel of r-m base. M3 section approximatelyequal to r-m. Gonostyli simple.

TYPE SPECIES: Gaalomyia carolinae, n.sp.ETYMOLOGY: The name is a feminine an-

agram of the genus name Aglaomyia.COMMENTS: The new genus differs form

Aglaomyia Vockeroth in having acrostichalsetae present, palpi short, Sc ending at R ba-sally of posterior fork, and 7th abdominalsegment well developed. PalaeodocosiaMeunier has palpi 4-segmented, vein Scshorter, and the base of fork M314 and CuAmore proximal. Pseudalysiinia Tonnoir haspalpi 4-segmented and incrassate, with nobristles on the thorax and legs, and tibialspurs short. The genus may be close to Ip-saneusidalys Blagoderov, 1998, especially I.longipennis, but the latter differs in havinglong fork of M314 and CuA and M3 section,and R5 is sinuous. Preservation of I. longi-pennis seems to be insufficient to decide ifboth species are congeneric.

Gaalomyia carolinae, new speciesFigure 44, Plate 4E

DIAGNOSIS: As for genus.DESCRIPTION: Body length 5 2.68 mm;


Fig. 44. Gaalomyia carolinae, n.sp., holotype AMNH Bu-390.

wing length 5 2.34 mm. Head: Occiput andfrons densely setose. Eyes large, with roundfacets, slightly emarginate, setulose. Ocelliabsent. Scape and pedicel rounded, flagellum14-segmented, filiform. Length of 1st andapical flagellomeres 2.53 width, with others23 width. Palpi 3-segmented, palpomeresequal in length and subsequently narrower toapex. Length of penultimate one 1.53 width,apical one 33 width. Clypeus setose. Tho-rax: Scutum with long lateral, dorsocentral,and acrostichal setae. Proepimeron toucheskatepisternum at shallow incision. Katepis-ternum larger than anepisternum. Anepister-nal suture horizontal. Antepronotum andproepisternum with long setae, other thoracicsclerites bare. Wing membrane without mi-crotrichia. R, R1, R5, and M1 with dorsal se-tae. Sc ends at R at the level of base of r-m.

R1 and R5 straight. Crossvein r-m equal toM3 section. Length of fork of M1 and M2 43M3 section. M1 and M2 not reaching wingmargin. Legs: Fore and mid coxae denselysetose, hind coxae with long posterolateralsetae. Femora with ventral row of setae. Tib-ial spurs 2–2.33 tibial diameter. Abdomendensely setose. Sternite I bare. Eighth tergiteshort, length 0.53 7th tergite. Tergite IX re-niform, width 33 length. Gonocoxites slen-der, length 3.53 width, widely separated.Gonostyli saberlike, curved inside.


ETYMOLOGY: The species epithet is a pa-tronym honoring our friend and colleagueCaroline S. Chaboo, specialist on chryso-melid beetles.


TRIBE LEIINI EDWARDS, 1925

Nedocosia, new genus

DIAGNOSIS: Three ocelli, lateral ones donot contact eye margin. Sc merges with R.R1 2–33 length of r-m. Veins of median andcubital forks lightly sclerotized. Base of M314

and CuA fork lies between levels of base ofr-m and M1–M2 fork. Tibial bristles absent.

TYPE SPECIES: Nedocosia exsangius, n.sp.ETYMOLOGY: The genus name is derived

from the prefix ne-, ‘‘negation’’, and the ge-nus Docosia. The name is feminine.

COMMENTS: This genus is most similar toDocosia Winnertz, 1863, which consists of25 extant Palaearctic, 15 Nearctic, and 2Neotropical species (Zaitzev, 1994). Sevenspecies of Docosia are known from Balticamber (Meunier, 1904, 1916, 1922, 1923)and one from the Oligocene shales of Rott,Germany (Statz, 1944). The new genus dif-fers by having lateral ocelli not touching theeye margins (undoubtedly plesiomorphic),and by the lack of strong tibial bristles. Dif-ferences in sclerotization of anterior veins(Sc, R, R5) and posterior veins (forks) in thenew genus are not as great as in Docosiaspecies. Two species of Docosia describedfrom the Lower Cretaceous of Transbaikalia(Blagoderov, 1995) may also belong to thenew genus, in fact showing affinities to N.novacaesarea.

Nedocosia exsanguis, new speciesFigures 45, 46, Plate 4F

DIAGNOSIS: Sc meets R before level ofbase of r-m. Laterotergites bare. Cerci withtwo combs of dark setae. Apices of gonostylisharp, curved inward at right angle.

DESCRIPTION: Body length 5 1.29 mm;wing length 5 0.67 mm (rest). Head oval,with rounded occiput, occiput and frons withscattered setae. Ocelli three, lateral one aboutits own diameter from eye margin and twodiameters from medial ocellus. Eyes slightlyemarginate, with round facets and short in-terfacetal trichia. Scape subconical, pedicelcylindrical. Flagellum 14-segmented, flagel-lomeres of hexagonal surface texture, cylin-drical, lengths about equal to width. Face andclypeus setose. Thorax: Scutum with setaein lateral, dorsocentral, and acrostichal rows.Antepronotum with 4 long and several short-

er setae. Proepimeron very small, touches ka-tepisternum at incision. Laterotergites andmediotergite bare. Metepisternum broaderthan high. Wing: Sc meets R before M3base. Costa ends beyond tip of R5, more thanone-half length between tips of R5 and M1.Crossvein r-m 0.373 length of R1, and 0.63length of M3 section. Base of fork of M314

and CuA between levels of bases of M3 sec-tion and M1 and M2 fork. M314 very weak.M1 and M2 fork campanulate. Longitudinalveins with setae, except Sc. Legs: Fore coxaewith anterior setae, mid coxa with apical se-tae. Fore leg with first tarsomere 23 longerthan second and 0.553 length of tibia. Ab-domen: Tergites setose, sternites bare. Cerciwith two comblike rows of black bristles.Apices of gonostyli sharp, squarish, curvedinward.

MATERIAL: Holotype PIN 3130/193, male.Russia: Taimyr Peninsula, Yantardakh, coll.1970.

ETYMOLOGY: Latin, meaning pale orbloodless, referring to the pale color of thespecimen.

Nedocosia sibirica, new speciesPlate 5A

DIAGNOSIS: Sc meets R at the level of thebase of the stem of M112. Laterotergites se-tose. Anterior edge of fore coxa bare. Hindlegs stout.

DESCRIPTION: Body length 5 2.37 mm,wing length 5 1.82 mm. Head: Occiput andfrons setose. Ocelli three, equal in size; dis-tance of median to lateral ocellus and fromlateral ocellus to eye margin equal to twicetheir diameter. Eye with slight incision at an-tennal bases. Antennae short, only 10 flagel-lomeres preserved; flagellomeres cylindrical,widths 1.1–1.53 lengths. Face triangular, se-tose, clypeus ovate, bare. Palpi 5-segmented,short, antepenultimate segment swollen, withmedial sensory pit on inner surface. Apicalpalpomere shorter than penultimate one, bothslender, with sparse setulae. Length ratio 1:3:3:5:5. Thorax: Mesonotum with short tri-chia and setae of various sizes; setae ar-ranged in 5 rows. Antepronotum with threesetae; proepisternum with two on lower part.Scutellum with two very long setae and twosmaller ones. Laterotergites with a few tri-


Figs. 45–47. Nedocosia n.gen. 45. N. exsanguis, n.sp., holotype PIN 3130/193. 46. Male genitalia.47. N. novacaesarea, n.sp., holotype AMNH NJ-117k.


chia in lower part. Mediotergite bare. Wing:Sc meets R at the level of the base of thestem of M112. R1 twice the length of r-m. R5

curved slightly back. C ends far beyond tipof R5, but does not reach wing apex. Stemof R, R1, and R5 with setae. Legs: Anterioredge of fore coxae without setae. Fore tibiaewith semicircular anteroapical depressedarea, covered with numerous small trichia.Mid tibial spur length 2.53 apical diameterof tibia, hind tibial spur 1.53 diameter. Hindlegs stout, longer than mid legs. First foretarsomere 2.53 longer than second and0.673 length of tibia. Cerci simple.

MATERIAL: Holotype PIN 3311/665, fe-male. Russia: Taimyr Peninsula, Yantardakh,coll. 1971.

ETYMOLOGY: The species epithet is a ref-erence to the region of origin of the amber.

Nedocosia canadensis, new speciesPlate 5B

DIAGNOSIS: Sc meets R before level ofbase of the stem of M112. R1 2.53 length ofr-m. M314 and CuA fork at the level of r-m.M3 section shorter than r-m crossvein. La-terotergites and mediotergite bare.

DESCRIPTION: Body length 5 1.50 mm(preserved part). Head: Occiput with setaeof different length. Ocelli not seen. Eyes withround facets, setulose. Scape small, pedicelsubconical. Flagellum 14-segmented, flagel-lomeres cylindrical, lengths equal to widths,densely covered by trichia no longer thanhalf width of basal flagellomere. Three seg-ments of palpi seen, combined length equalto head height. Thorax: Scutum uniformlysetose with long and short setae. Antepron-otum with two very long setae and severalshort ones. Laterotergites and mediotergitebare. Wing: Sc meets R before level of baseof the stem of M112. R1 2.53 length of r-m.Crossvein r-m 1.33 length of M3 section.Base of fork of M314–CuA proximal to baseof fork of M1–M2.

MATERIAL: Holotype MCZC 6897, incom-plete specimen, sex unknown. Canada: Man-itoba, Cedar Lake, coll. F. M. Carpenter.

ETYMOLOGY: The species epithet is a ref-erence to the country of origin of the amber.

Nedocosia novacaesarea, new speciesFigure 47, Plate 5C

DIAGNOSIS: Flagellum 11-segmented. Sclong, ends at R between RS and r-m bases.Base of M314 and CuA fork at level of r-m.Fore leg with tibia equal to femur or slightlylonger.

DESCRIPTION: Body length 5 2.34 mm (ho-lotype)/2.88 mm (paratype); wing length 51.75 mm (holotype)/2.20 mm (paratype).Head: Three ocelli in line, distances betweenmedian and lateral ones, and between lateraland eye margin, are equal. Face rectangular,bare. Clypeus bare. Scape and pedicel round-ed, shorter than flagellomeres. Flagellum 11-segmented, flagellomeres cylindrical, withlengths 23 width. Two palpomeres seen, api-cal one 23 length of basal. Thoracic scleritesbare. Scutum with long lateral setae and nu-merous short ones, not arranged in rows.Proepimeron small, fits in shallow incision onkatepisternum. Anepisternal suture declinesbackward. Metepisternum touches anepime-ron. Meron large. Wing: Costa ends beyondtip of R5, more than one-third length betweentips of R5 and M1. Sc long, ends at R betweenRS and r-m bases. Length of R1 2.5–33 thatof r-m, M3 section twice that of r-m. Lengthof fork of M1 and M2 2.5–33 M3 section.Base of fork of M314 and CuA at level of r-m base. Legs: Fore coxae with short anteriorsetae, hind coxae with long posterolateral se-tae. Fore leg with tibia equal to femur. Tibialspurs 1.2–1.53 tibial diameter. Abomen se-tose. Seventh segment very narrow. Gonocox-ites massive, with apices blunt.

MATERIAL: Holotype AMNH NJ-117k,male; paratype NJ-117l, male in the samepiece. USA: New Jersey, Sayreville, coll. P.Nascimbene.

ETYMOLOGY: The species epithet is derivedfrom Latin Nova Caesarea meaning ‘‘NewJersey’’ and is a reference to the state of or-igin of the amber.

COMMENTS: The species is similar in ve-nation to Docosia baisae Blagoderov, 1998and D. zaza Blagoderov, 1998, by proximalposition of the forked base of M314 and CuA,but differs from the latter species in havingSc long and M3 section short.


Ectrepesthoneura Enderlein, 1911

Willistoniella Meunier, 1904: 74 (preoccupied byMik, 1895).

Meunieria Johannsen, 1909: 87 (preoccupied byKieffer, 1904).

Ectrepesthoneura Enderlein, 1911: 115.

DIAGNOSIS: As given by Chandler (1980):Lateral ocelli remote from eye margin; later-otergites bare; Sc ending in R before base ofRS; R4 present; R1 short, at most twice aslong as r-m; R5 straight; C prolonged wellbeyond tip of R5; veins of medial and cubitalsectors weak and faint; posterior fork sessile.Males with ‘‘sensory pit’’ near the base ofmid tibia; hind tibial comb absent.

TYPE SPECIES: Tetragoneura hirta Win-nertz, 1846: 19 (orig. designation)

COMMENTS: There are about 10 extant spe-cies from the Holarctic region. The genus isvery close to Tetragoneura Winnertz (about100 widespread living species) and is some-times included in it or treated within the tribeGnoristini (Tuomikoski, 1966; Vaisanen,1986). There are two fossil species describedfrom Baltic amber (Meunier, 1904) and theOligocene of Rott, Germany (Statz, 1944).Chandler (1999) suggested that Ectrepestho-neura was paraphyletic.

Ectrepesthoneura succinimontana,new species

Figures 48, 49, Plate 5D

DIAGNOSIS: Fore tibiae slightly shorter thanfore femur. Section M3 and bases of M1 andM2 very weak. Length of small radial celltwice the width.

DESCRIPTION: Body length 5 2.39 mm;wing length 5 2.02 mm. Head: Occiput andfrons with short curved setae. Ocelli three,distance of lateral one to eye margin morethan twice the ocellus diameter. Scape sub-conical, pedicel rounded, width twice flagel-lomere width. Eleven segments of flagellumpreserved, flagellomeres cylindrical, withlength about equal to width. Face setose. Pal-pi 4-segmented; 1st palpomere very small;antepenultimate one oval, its length twice thewidth, with medial sensory pit; penultimateone with length 33 width, attached to thesecond one preapically; apical segment longand slender. Clypeus triangular, setose. Tho-rax: Scutum with numerous long setae most-

ly curved forward and shorter ones curvedback, arranged in rows. Antepronotum with3 long setae. Scutellum with two pairs oflong setae. Other thoracic sclerites bare.Wing: All longitudinal veins with short se-tae. Sc meets R proximal to base of M3.Length ratio of sections RS1, RS2, and RS3is 1:2:13. RS1 transverse, 0.33 length ofr-m. Crossvein r-m 0.73 length of R1 and0.53 length of M3 section. Costa ends be-yond tip of R5, one-third distance betweentips of R5 and M1. Section M3 and M1 andM2 bases very weak. Legs: Fore coxae withnumerous anteromedial setae, hind coxaewith posterolateral setae. Fore and mid tibiaeslightly shorter than fore femora. Mid andhind tibiae and tarsi with rows of black bris-tles. Tibial spurs 1.5–23 tibial diameter. Tar-sal claw with one long tooth. Abdomen se-tose. Eighth sternite almost triangular. Cerci2-segmented, basal segment stick-shaped,with length twice the width, extends beyond8th sternite; apical one discoidal, attached atthe apex of the basal one, 2.23 shorter. Dor-sal border of 8th sternite (gonocoxite 8 ofMartinsen and Soli, 2000) straight.


ETYMOLOGY: From Latin succinum mean-ing ‘‘amber’’ and montanus meaning ‘‘moun-tainous’’. Yantardakh means ‘‘amber moun-tain’’ in Dolgan language.

COMMENTS: Comparision with recent spe-cies is difficult, because nongenital charactersused for distinguishing species have been con-sidered to be unreliable for this genus (Mar-tinsen and Soli, 2000). For species where thefemale genitalia are known the new species ismost similar to E. hirta Winnertz.

Ectrepesthoneura swolenskyi, new speciesFigures 50, 51, Plate 5E

DIAGNOSIS: Sc long, meets R just beforebase of M3. Small radial cell relatively long.Gonocoxites with triangular ventral append-ages bearing 2 long, dark apical spurs.

DESCRIPTION: Body length 5 2.71 mm;wing length 5 2.46 mm. Head: Ocelli three,from lateral ocellus to eye margin twice ocel-lus diameter. Flagellomeres cylindrical,length 1.53 width. Three palpomeres seen,


Figs. 48–51. Ectrepesthoneura End. 48. E. succinimontana, n.sp., holotype PIN 3311/662. 49. Fe-male genitalia. 50. E. swolenskyi, n.sp., holotype AMNH NJ-824. 51. Male genitalia.


apical one slender, length ratio 1:2:3.5. Tho-rax: Antepronotum with 2 setae, proepister-num with 4. Scutum with setae curved for-ward, lateral setae very long. Wing: All lon-gitudinal veins except Sc with short setae.Costa ends beyond tip of R5, one-half dis-tance between tips of R5 and M1. Sc meetsR just before M3 base. Length of R1 2.23that of r-m. Length of M3 2.73 that of r-m.Ratios of lengths of sections RS1, RS2, andRS3 1:2.3:7. RS1 oblique, 0.583 length ofr-m. M3, M1, and M2 weak, M1 slightlycurved forward. M314 and CuA curved gent-ly caudad. Legs: Fore and mid tibiae longerthan femora. Mid tibia with anterolateral,posterolateral, and posteromedial rows ofshort bristles, hind tibia with two rows ofposterior setae. Basal third of mid tibia withlarge, dark, narrow sensory pit. Tibial spurs1.3–1.73 tibial diameter. Abdomen. Ninthtergite large, conceals gonocoxites and gon-ostyli almost completely; caudal edgestraight, without medial cleft. Ninth tergitestraight caudally, without cleft, with numer-ous curved setae apically. Gonocoxites withtriangular ventral appendages bearing 2 long,dark apical spurs. Gonostyli short, pointed,curved S-like.

MATERIAL: Holotype AMNH NJ 824,male. USA: New Jersey, Sayreville; coll. S.Swolensky, 1997–1998 (mentioned in Gri-maldi, 2000: fig. 48h).

ETYMOLOGY: The species epithet is patron-ymic for the late Steve Swolensky, who col-lected the specimen.

COMMENTS: The new species differs fromother species of the genus in the structure ofmale genitalia, especially by ventral append-ages of gonocoxites bearing dark sclerotizedspurs.

Izleiina, new genus

DIAGNOSIS: Facets round. Lateral ocelli nottouching eye margins. Clypeus setose. Scmeets C just beyond level of base of M3; Sc2

absent. Length of R1 23 that of r-m. SectionM3, and fork of M1 and M2–M314 reduced;CuA weakened. M314 interrupted at base.

TYPE SPECIES: Izleiina mirifica, n.sp.ETYMOLOGY: Name is derived from the

tribe Leiini.COMMENTS: We assume that in the new ge-

nus vein M1 is entirely reduced. As a rule, inmost leiine genera, vein M1 reaches the wingmargin at or even before the wing apex. Theyhave retained a complete vein that is homol-ogous to CuA based on the length and shapeof the vein, and which is strongly curvedback and ends at the level of RS1, as in mostLeiini. The new genus differs from Neoclas-tobasis Ostroverchova by the lateral ocellitouching eye margins, and from ClastobasisSkuse and Neoclastobasis by C ending be-yond the tip of R5, and by the presence ofthe base of M314. Having R1 relatively longand Sc ending at C makes the genus similarto Rondaniella Johannsen and Indoleia Ed-wards, but differs from them by reduction ofthe veins in the medial sector. Such a reduc-tion also occurs in other leiine genera, forexample Novakia Strobl and Sigmoleia Ton-noir and Edwards, but they have Sc veryshort and free. Species of Cycloneura Mar-shall and Paracycloneura Tonnoir and Ed-wards also have M unbranched and M314 freeor reduced at the base, the anepisternumlarge, metepisternum narrow like Izleiina,but they differ strongly in the shape of CuA,by having M2 long, Sc short and free, and bythe presence of strong tibial bristles.

Izleiina mirifica, new speciesFigures 52–54, Plate 5F

DIAGNOSIS: Apical palpomere short. Tibiaewithout bristles. Gonostyli conical, curvedinward.

DESCRIPTION: Body length 5 1.44 mm,wing length 5 1.23 mm. Head: Ocelli three,distance from eye margin equal to two ocel-lus diameters; distance between each otherequal to one diameter. Flagellum 14-seg-mented, flagellomeres cylindrical, lengthsabout equal to widths, covered by setulae nolonger than one-half flagellomere width. Pal-pi 4-segmented, three basal segments bacil-liform, apical one short and rounded. Occi-put, frons, face, and clypeus with short se-tulae. Thorax: Prescutum and anterior par-apsidal suture distinct. Antepronotum with 6setae, proepisternum with two. Scutumarched. Scutellar setae in rows. Scutellumwith two pairs of long setae. Cavity on an-terior edge of katepisternum shallow. Later-otergites and mediotergite bare. Wing: Sc


Figs. 52–57. Izleiina, n.gen. 52. I. mirifica, n.sp., holotype PIN 3130/187. 53. Reconstruction ofwing venation. 54. Male genitalia. 55. I. spinitibialis, n.sp., holotype AMNH NJ-346, reconstruction ofwing venation. 56. Head. 57. Male genitalia.


meets C just beyond level of base of M3. Cends far beyond apex of R5. R1, R5, and r-mwith setae. R1 length 23 that of r-m. R5

straight. Length of crossvein r-m 0.53 lengthof section M3. M3 weak, can be seen onlyby absence of microtrichia on wing mem-brane. M314 and CuA fork base proximal tothe base of r-m. M314 interrupted at base, notreaching wing margin. Legs: Fore coxaewith anterior setae, mid coxae with lateroap-ical ones. Tibiae without bristles. Genitalia:Gonocoxites massive, fused. Gonostyli horn-like and slightly curved, with long setae, col-ored lighter than gonocoxites.


ETYMOLOGY: The species epithet is a Latinword mirificus meaning ‘‘amazing’’ in ref-erence to unusual combination of characters.The name is feminine.

Izleiina spinitibialis, new speciesFigures 55–57, Plate 6A

DIAGNOSIS: Tibiae with bristles. Gonostylicylindrical, length 2.53 the width, with 3dark teeth at the apex

DESCRIPTION: Body length 5 2.07 mm;wing length 5 1.65 mm. Head: Ocelli three,almost in a straight line. Lateral ocelli distantby two diameters from medial ocellus andtheir own diameter from eye margin. Frontalfurrow distinct. Eyes emarginate at antennalbase, bare. Frons bare. Face setose. Scapeand pedicel without strong setae, rounded,wider than flagellomeres. Eight flagellomeresseen, cylindrical in shape, lengths 1.53width. Clypeus rounded, setulose. Palpi 4-segmented, basal segment rounded, antepen-ultimate and penultimate one bacilliformwith lengths 3–43 width; apical palpomerelong. Thorax: Scutellum with 4 long setae.Laterotergites and mediotergite bare. Wing:Sc long, meets C at level of M3 base. R1

straight, 23 length of r-m. R, R1, R5, and r-mwith a dorsal row of setae. Crossvein r-malmost horizontal. R5 curved caudally. Cends beyond tip of R5. M2 and M314 weak.Base of CuA weak. CuA curved caudally,not sigmoid. Legs: Tibiae with bristles. Ab-domen: Genitalia covered by long setae.Ninth tergite rectangular, length 2.33 the

width. Gonocoxites slightly swollen. Gon-ostyli almost straight, length 2.53 width,with 3 dark teeth at the apex.

MATERIAL: Holotype AMNH NJ-346,male. USA: New Jersey, Sayreville, coll. Y.Goldman, 1995.

ETYMOLOGY: The species epithet is derivedfrom Latin spina, meaning ‘‘spine’’, and tib-ia, meaning ‘‘leg’’.

Zeliinia, new genus

DIAGNOSIS: Close to Izleiina n.gen., butdiffers in having palpi long, Sc free, M1 re-duced at base, M2 absent, and base of pos-terior fork proximally with r-m touching thebase of M314. Setulae on flagellomeres ar-ranged in rings.

TYPE SPECIES: Zeliinia orientalis, n.sp.ETYMOLOGY: The genus name is an ana-

gram for Izleiina. The name is feminine.

Zeliinia orientalis, new speciesFigures 58–61, Plate 6B

DIAGNOSIS: Genitalia longer than wide;gonostyli bilobate. Tergite 9 obovate, withnumerous setae at apex.

DESCRIPTION: Male. Body length 5 1.25mm; wing length 5 1.07 mm. Head: Ocellithree, close to each other. Eyes strongly in-cised, forming incomplete bridge. Pedicelslightly wider than scape and flagellum; 14cylindrical flagellomeres with length 23width. Three palpomeres seen, sticklike,length ratio 1:1.3:2. Thorax: Scutum withlong lateral, dorsocentral, and acrostichal se-tae. Katepisternum larger than anepisternum,proepimeron touches katpisternum at smallexcavation. Antepronotum and proepimeronsetose. Laterotergites and mediotergite bare.Wing: Costa slightly produced beyond apexof R5. Sc free, ends at the middle of r-m. R1

and R5 with long dorsal and ventral setae.Crossvein r-m forming one vein with tb andM2 section, equal to 1.53 length of R1.Length of R5 2.83 R1. Base of M3 faint. M1

ends at wing apex, base curved posteriorly.Base of M314 and CuA fork contact r-m.Legs: Tibial setulae irregular. Tibia II withapical comb of setae. Tibial spurs 1–1.53tibial diameter, spur formula 1:1:1. Abdo-men setose, 7th and 8th segments short, nar-row, retractable. Genital complex large,


Figs. 58–63. Zeliinia, n.gen. 58. Z. orientalis, n.sp., holotype AMNH Bu-315. 59. Wing. 60. Malegenitalia laterally. 61. Male genitalia dorsally. 62. Z. occidentalis, n.sp., holotype MCZC 6943. 63. Malegenitalia.


length equal to 4th–8th segments combined.Ninth tergite fiddle-shaped, with two apicalcombs of dark setae. Gonocoxites rectangu-lar in lateral view. Gonostyli bilobate.


ETYMOLOGY: The species epithet is a Latinword orientalis, meaning ‘‘eastern’’, refer-ring to origin of the amber.

Zeliinia occidentalis, new speciesFigures 62, 63, Plate 6C

DIAGNOSIS: Clypeus with long setae; me-diotergite and laterotergites bare, wing mem-brane without macrotrichia, Sc short, endsfree, R1 short, 1.53 longer than r-m, endsproximally at tip of CuA. Genital complexwider than long; gonostyli unilobate.

DESCRIPTION: Body length 5 1.79 mm;wing length 5 1.22 mm. Head: Eyes bare,forming dorsal bridge, possibly incomplete(vertex and occiput not fully seen due to de-formation). Scape and pedicel short, round-ed. Flagellum 14-segmented, flagellomeresequal in length, apical flagellomeres morenarrow than basal ones, with length abouttwice the length of pedicel, covered withshort microtrichia. First flagellomere length1.63 width, preapical is 2.53 width. Clypeussetose. Three visible palpal segments, twobasal ones about equal in length; apical pal-pomere long and slender, about twice thelength of the penultimate. Thorax: Ante-pronotum with two setae, proepisternum withone. Scutum with long lateral, dorsocentral,and acrostichal setae. Scutellum with 4 pairsof long setae. Katepisternum larger than ane-pisternum. Metepisternum small, quadrate,bare. Mediotergite and laterotergites bare.Wing membrane without macrotrichia. R,R1, and R5 setulose. Sc short, ends free. R1

1.5 length of r-m, R5 more than 4.53 lengthof r-m. Crossveins r-m, tb, and m-cu fusedinto one oblique vein, with distinct kink atthe base of the stem of M112 (M3 section).Stem of M112 and base of M314 weak. M ap-parently without fork. M314 and CuA curvedgently caudad. Legs: Femora with ventralrow of long setae. Coxae relatively short,fore coxa as long as mid and hind ones. Midand hind tibiae with short bristles. Tibial

spurs 1.2–1.53 tibial diameter. Tarsal clawswith very small tooth. Abdomen densely se-tose, with long setae. Genitalia rectangular,width 1.63 length. Gonocoxites stout, gon-ostyli simple, pointed, with inner surfaceslightly curved apically.

MATERIAL: Holotype MCZC 6943, male.Canada: Manitoba, Cedar Lake, coll. F.M.Carpenter.

ETYMOLOGY: The species epithet is Latinmeaning ‘‘western’’, referring to origin of theamber

COMMENTS: Z. occidentalis differs fromthe type species in the structure of genitalia,and although it possesses the same arrange-ment of flagellar setulae and venation pat-tern, both species may not be congeneric.

Temaleia, new genus

DIAGNOSIS: Wing membrane with macro-trichia; Sc long, meets C before base of M3section; C ends at tip of R5; M3 section 33length of r-m; M314 not interrupted at thebase; tibial setulae irregular; tibial bristlesabsent; abdominal segment 6 not reduced;7th and 8th segments telescopic.

TYPE SPECIES: Temaleia birmitica, n.sp.ETYMOLOGY: The genus name is a femi-

nine anagram of Metaleia.COMMENTS: The genera Clastobasis Skuse,

1896, Neoclastobasis Ostroverchova, 1970,and Metaleia Baxter, 1994 have M314 inter-rupted at the base and/or CuA sinuous. Thegenus Sticholeia Soli, 1996, besides havingextremely long cerci and other details of themale terminalia, differs by having a short M3section.

Temaleia birmitica, new speciesFigure 64, Plate 6D

DIAGNOSIS: As for the genus.DESCRIPTION: Body length 5 2.17 mm (ho-

lotype)/2.03–2.38 mm (paratype); winglength 5 1.96 mm (holotype)/1.50–2.07 mm(paratype). Head: Pedicel rounded, lengthabout equal to width. Flagellum 14-segment-ed; flagellomere length about twice thewidth. Eyes large, densely setose. Occiputcovered with numerous short setae. Clypeussetose. Palpi 3-segmented, very short, pal-pomeres with length about equal to width.Labella longer than palpi. Thorax: Scutum


Figs. 64–68. Temaleia and Lecadoniella. 64. T. birmitica, n.sp., holotype AMNH Bu-483a. 65. L.parvistyla, n.sp., holotype MCZC 6941. 66. Wing. 67. Male genitalia ventrally. 68. Male genitaliadorsally.


irregularly covered with short setae and withlong lateral, dorsocentral, and acrostichal se-tae. Scutellum with three pairs of long bris-tles. Laterotergites and mediotergite withseveral protruding bristles. Wing membranewith microtrichia and macrotrichia, locatedmostly in distal half of wing. Costa ends attip of R5, not reaching wing apex. Sc long,meets C before base of M3. Sc2 apical. R1

very short, 1.53 length of r-m. M3 section33 length of r-m. M1 meets wing edge atwing apex. M1 and M2 fork 2.53 as long asM3 section. Base of M314 and CuA fork atlevel of M3 base. M314 and CuA curvedgently caudad. Legs: Fore coxae with ante-rior and anterolateral setae. Mid coxa withlateroapical setae, hind one with posteroapi-cal setae. Tibial spurs short, no longer thanmaximum tibial diameter. Tibial and tarsalbristles absent. Tibial setulae irregular. Ab-domen pubescent, with 7 visible segments,8th small and retracted. Ninth tergite large,oval. Female: Cerci one-segmented, bacilli-form, setose. Male: Gonocoxites and gon-ostyli long, slender, bacilliform. Gonostylislightly curved inside at apex, saberlike.

MATERIAL: Holotype AMNH Bu-483a, fe-male; paratypes AMNH Bu-060, male, Bu-054, male. Myanmar: Katchin, from ambermines near Myitkyina.

ETYMOLOGY: The species epithet is a ref-erence to Burma, the former name of thecountry where the amber originates.

Lecadonileia, new genus

DIAGNOSIS: Palpi reduced, laterotergitessetose; wing membrane with macrotrichia;Sc free, Sc2 absent; M314 interrupted at base,M314 and CuA curved gently caudad, 9th ter-gite small or fused with synsclerite; gono-coxites widely separated.

TYPE SPECIES: Lecadonileia parvistyla,n.sp.

ETYMOLOGY: The genus name is a femi-nine anagram of the Caledonileia Matile,1993.

COMMENTS: The genus is close to the ge-nus Caledonileia, from which it differs innumber and position of ocelli, number of pal-pomeres, setation of scutum, and in havingSc long, the radial cell wide, M3 long, andgonostyli unilobate (Matile, 1993). The ge-

nus differs from Megophthalmidia Dzied-zicki in having M314 interrupted at the baseand the fork of M1 and M2 not shifted ante-riorly, so that M1 reaches the wing marginbeyond the wing apex.

Lecadonileia parvistyla, new speciesFigures 65–68, Plate 6E

DIAGNOSIS: As for genus.DESCRIPTION: Body length 5 2.24 mm,

wing length 5 2.05 mm. Head: Vertex se-tose. Ocelli three, lateral one twice its di-ameter from eye margin. Frons bare, facewith tiny trichia. Scape small, roundish. Ped-icel rounded, with dorsal setae, wider thanflagellomeres. Flagellum 14-segmented, cy-lindrical, flagellomere length equal to width.Palpi 4-segmented. Basal palpomere veryshort, apical three subequal, lengths 2.5–33their width. Clypeus setose. Thorax: Scutumirregularly setose. Antepronotum with 4 se-tae, proepisternum with 5. Ventral process ofproepimeron just opposite anapleural suture.Anepisternum with distinct oblique cleft.Anepimeron very narrow ventrally. Latero-tergites setose. Mediotergite bare. Wingmembrane with macrotrichia. Costa ends be-yond tip of R5, at one-fourth length betweentips of R5 and M1. R1 and R5 with two rowsof setae; R stem, r-m, M1, M2, M314, and CuAwith one row. Sc ends free at the level of thebase of M3 section. R1 equal to M3 sectionand 2.53 length of r-m. Fork of M1 and M2

1.83 length of M3. M314 base absent, veinbegins at level of RS base. Legs: Fore coxaewith numerous setae on anterior and lateralsurfaces. Mid coxae with setae on apical half.Hind coxae with posterolateral row of setae.Tibial bristles short. Fore tibiae with 3 apicalbristles; mid one with 2 anterior and 6 apicalbristles; hind tibia with 4–6 bristles in ante-rior, anteroventral, and anterodorsal rows andseveral apical bristles. Anteroapical depres-sion on fore tibiae well developed, with sin-gle row of short setae. Tarsal claws withoutteeth. Abdomen setose. Eighth and 9th seg-ments very short and retracted. Gonocoxitessemicylindrical, widely separated at base.Gonostyli small, with one apical appendageturned medially.

MATERIAL: Holotype MCZC 6941, male.


Canada: Manitoba, Cedar Lake, coll. F.M.Carpenter.

ETYMOLOGY: The species epithet is a com-bination of the Latin word parvus, meaning‘‘small’’, and stylus, meaning ‘‘spike, stem,pen’’, in reference to simple small gonostyliof the species.

Disparoleia, new genus

DIAGNOSIS: Ocelli absent; anepisternumwith very short setae and trichia; medioter-gite and laterotergites bare; wing membranewith micro- and macrotrichia; Sc ends at C;r-m 23 length of R1, M3 base at basal one-sixth of wing, M3 weak, M2 absent.

TYPE SPECIES: Disparoleia cristata, n.sp.ETYMOLOGY: The genus name is a combi-

nation of the Latin word dispar, meaning‘‘unequal, unlike’’, and the genus Leia, inreference to the distinctiveness of the newgenus. The name is feminine.

COMMENTS: The genus differs from all Re-cent Leiini in the unique combination of suchcharacters as absence of ocelli, very long r-m with the proximal position of the base ofM3, and reduction of veins in the medial sec-tor. It differs from Novakia Strobl in havingthe base of RS distinct, M2 absent, and Scending at C. It differs from Zeliinia, n. gen.in having flagellomeres of smooth texture,ocelli absent, vein M1 and M3 section presentthough weak, and fork of M314 and CuA longbut not sessile.

Disparoleia cristata, new speciesFigures 69, Plate 7A

DIAGNOSIS: As for genus.DESCRIPTION: Female. Body length 5 2.73

mm, wing length 5 2.08 mm. Head: Vertexwith group of long erect setae. Scape andpedicel wider than flagellum. Flagellum 14-segmented, flagellate, flagellomeres longerand narrower apicad. Eyes setose, slightlyemarginate at antennal base. Ocelli absent.Clypeus setose. Palpi 3-segmented, 1st pal-pomere short, 2nd longer, cut obliquely, 3rdlonger than length of 1st and 2nd, narrow.Thorax: Scutum with lateral and dorsocen-tral rows of long setae and short irregularsetae. Scutellum with numerous short setae.Antepronotum with long curved setae andseveral short ones. Anepisternum with 4 light

short setae in upper part and very fine hairsat lower apical part. Proepimeron touches ka-tepisternum slightly ventral to anepisternalsuture. Mediotergite and laterotergites bare.Metepisternum large, with dorsal marginwider than ventral one. Wing membranewith microtrichia and short macrotrichia.Costa ends beyond tip of R5, at one-thirdlength between tips of R5 and M1. Humeralvein located beyond the MA. Sc short, endsat C before level of M3 base, apex faint. R1

very short, r-m 33 length of R5. Medianveins weak. M1, base of M1 and M3 veryweak. Base of M3 very proximal, slightlydistal to apex of Sc. Crossvein r-m 33 lengthof M2 section, M3 section 3.53 that of M2.Base of M314 and CuA fork slightly distal tolevel of base of M3 section. R, R1, R5, M314,and CuA with setae dorsally and ventrally.Leg: Tibial spur formula 1:2:2. Spurs long,4–53 tibia diameter. Mid and hind tibiaewith bristles. Empodium absent; tarsal clawswith one tooth. Abdomen: Tergites of ab-domen setose. Seventh tergite 1.53 shorterthan 6th, 8th is very short. Cerci 2-segment-ed, slightly shorter than 6th and 7th segmentstogether. Apical segment round, basal onebacilliform, 2.53 apical.

MATERIAL: Holotype AMNH B-0125, fe-male. Myanmar: Katchin, from amber minesnear Myitkyina.

ETYMOLOGY: The species epithet is Latinword cristatus, meaning ‘‘tufted, plumed’’, inreference to the group of setae on the vertex.

Hemolia, new genus

DIAGNOSIS: Three ocelli in straight line,middle one small, lateral ones not touchingeye margin; wing membrane without macro-trichia; Sc short, ends at R; R1 short, no morethan 23 length of r-m; r-m short, oblique;M3 short; tibial setae in rows, mid and hindtibiae with inner spurs longer than outer one.

TYPE SPECIES: Hemolia matilei, n.sp.ETYMOLOGY: The genus name is a femi-

nine anagram of Mohelia Matile, 1978.COMMENTS: The genus is close to Mohelia

in the structure of the ocelli, and wing ve-nation, and it differs in tibial spur length anddetails of male genitalia structure. It differsfrom Mohelia in having setae on the medi-otergite, tibial setae in rows, mid and hind


Figs. 69–72. Disparoleia and Hemolia. 69. D. cristata, n.sp., holotype AMNH B-0125. 70. H.matilei, n.sp., holotype AMNH B-0132, wing. 71. Male genitalia ventrally. 72. H. glabra, n.sp., holotypeAMNH B-0112, wing.


←

Figs. 73–75. Protragoneura platycera, n.sp. 73. Holotype AMNH Bu-135. 74. Male genitalia later-odorsally. 75. Male genitalia lateroventrally.

tibiae with inner spurs longer rather than theouter ones, and C ending close to the tip ofR5; Sc is longer, R1 longer than r-m; and M3is short.

Hemolia matilei, new speciesFigures 70, 71, Plate 7B

DIAGNOSIS: Scutum densely setose; later-otergites setose; Sc rather long, ends at thelevel of M314 and CuA fork base.

DESCRIPTION: Body length 5 2.19 mm (ho-lotype)/2.39 mm (paratype); wing length1.49 mm (holotype)/2.21 mm (paratype).Head: Eyes setose. Occiput and frons dense-ly setose. Three ocelli in straight line, middleone small; lateral one separated from eyemargin by its own diameter. Flagellum 14-segmented, flagellomeres cylindrical; lengthof flagellomeres about equal to their width.Clypeus setose. Palpi 4-segmented, basalsegment small, penultimate and apical onessituated preapically, antepenultimate withround sensory pit on inner surface. Thorax:Scutum with long setae not arranged in dis-tinct rows. Laterotergites with long setae.Scutellum protruding, with 6 pairs of longsetae. Antepronotun and proepisternum withlong setae. Wing rather wide, membranewithout macrotrichia. Costa ends beyond tipof R5 at one-fifth length between tips of R5

and M1. Sc ends in R at the level of M314

and CuA fork base. R, R1, and R5 with setaedorsally. R1 length 1.83 length of r-m.Crossvein r-m about the length of M3, withconstriction in the middle. Length of M1 andM2 fork 5.33 length of M3. Legs: Tibial se-tae in rows. Tibial spur formula 1:2:2; midand hind tibial spurs differ in length: innerspur 1.73 outer one. Fore leg basitarsomereshorter than tibia. Abdomen setose, with 6visible segments, 7th and 8th ones small andretractable. Gonocoxites divided by completesuture ventrally. Male: Gonostyli large, tri-angular, flat, with inner, bilobed appendage.Aedeagus bilobate. Female: Eighth sternitestriangular; 10th sternite with 4 dark, thick,wavy, blunt setae. Cerci 2-segmented, round-

ed, flat; apical segment 0.243 basal one,semicircular.

MATERIAL: Holotype AMNH B-0132,male; paratype AMNH B-0133a, female.Myanmar: Katchin, from amber mines nearMyitkyina.

ETYMOLOGY: The species epithet is a pa-tronym in honor of the late Professor LoıcMatile, a world authority on Sciaroidea, whowas also very generous and helpful to bothauthors.

Hemolia glabra, new speciesFigure 72, Plate 7C

DIAGNOSIS: Laterotergites bare; scutum al-most bare, dark and shining, Sc meets R wellbefore the level of base of M314 and CuAfork.

DESCRIPTION: Body length 5 3.16 mm;wing length 5 2.05 mm. Head: Occiput withshort setae. Three ocelli in line, of the samesize, distant from each other and from eyemargin by two ocellus diameters. Eyes large,setose. Pedicel with round sensory organ.Flagellum 14-segmented, apical segmentconical, secondarily divided. Thorax: Scu-tum almost bare; dark, shiny. Anterior par-apsidal suture distinct. Scutellum and later-otergites bare. Mediotergite with a few finesetae anteroventrally. Mid and hind tibiaewith inner spurs 1.53 longer than outer ones.Wing membrane without macrotrichia. Hu-meral vein at the level of MA. Costa endsbeyond tip of R5, at one-third the length be-tween tips of R5 and M1. Sc short, ends at R.R1 1.33 the length of r-m. Length of M3about equal to length of r-m. Length of M1

and M2 fork 4.33 the length of M3. Base ofM314 and CuA fork between tip of Sc andM3 base. Legs: Coxae without setae. Hindtibiae with posterior bristles. Abdomen withscattered setae. Cerci 2-segmented, apicalsegment conical, basal one cylindrical, 33length of apical one. Gonocoxites 8 long,narrow, slightly curved, with long setae.

MATERIAL: Holotype AMNH B-0112, fe-


male. Myanmar: Katchin, from amber minesnear Myitkyina.

ETYMOLOGY: The species epithet is Latinfor ‘‘hairless, smooth’’, in reference to theabsence of thoracic setae and setulae.

Protragoneura, new genus

DIAGNOSIS: Small, dark gnats with anten-nae shorter than thorax, flagellomeres shorterthan wide. Mesonotum densely covered withshort trichia, with or without a few setae lat-erally. Sc short, free. Costa is not producedbeyond the tip of R5, which is long, curved,and almost reaching the tip of M1. Fork ofM1 and M2 shifted anteriorly, M1 ends beforewing apex. Base of M314 and CuA fork at thelevel of base of M3, in basal one-sixth ofwing. M3 length about equal to that of M1

and M2 fork.TYPE SPECIES: Protragoneura platycera,

n.sp.ETYMOLOGY: The genus name is a combi-

nation of the prefix pro- (Greek pro-, be-forehand) and Tetragoneura. The name isfeminine.

COMMENTS: Closest to Tetragoneura Win-nertz, 1846, about 100 extant species ofwhich are known from the Holarctic (17spp.), Neotropical (ca. 60 spp.), and Austra-lasian (23 spp.) regions. The new genus dif-fers in having C not extended beyond the tipof R5, RS2 section (small radial cell) long,and base of M314 and CuA fork very basal;M1 and M2 fork shifted anteriorly. Setationof mesonotum is also distinctive and sepa-rates the new genus from Tetragoneura.

Protragoneura platycera, new speciesFigures 73–75, Plate 7D


lotype)/2.39 mm (paratype); wing length 51.63 mm (holotype)/1.93 mm (paratype).Head ovate, with row of long erect setae be-hind eye margin. Scape and pedicel subcon-ical, with numerous apical setae, both black.Flagellum compressed laterally, flagello-meres transverse, widths 1.5–2.53 length,densely covered with short trichia. Clypeussetose. Three palpomeres seen; antepenulti-mate one swollen, obovate, length 23 width.Penultimate one attached to antepenultimate

palpomere preapically; length 43 width,knoblike. Apical palpomere long and narrow.Length ratio 1:1.7:2.5. Thorax: Scutumdensely covered with short erect trichia,bearing some setae laterally. Antepronotumand proepisternum with long strong setae.Anepisternum with deep cleft. Katepisternumwithout distinct excavation on anterior mar-gin. Laterotergites, mediotergite bare. Met-episternum with few short trichia ventrally.Wing membrane with macrotrichia in distaland posterior part. Sc extremely short, free.All longitudinal veins except R base with se-tae. M3 reduced, can be traced by setae only.M1 base reduced. Base of M314 and CuA forkweakened. Small radial cell length 73 width.Halters dark, with few short setae. Legs:Coxae and femora dark, densely setose, tib-iae and tarsi yellowish. Hind tibiae with nu-merous posterior bristles. Tibial spurs 43 tib-ial diameter. Abdomen with 6 visible seg-ments, 7th and 8th very short, retracted. Ter-gite 9 oval, short, covering the base ofgonocoxites. Gonocoxites fused at base, mas-sive, narrowing to apex in distal half. Gon-ostyli with long ventral lobe bearing a combof short, knoblike setae; a somewhat shorter,dorsal, bare lobe and internal wide lobe witha row of curved bristles.

MATERIAL: Holotype AMNH Bu-135,male; paratype AMNH Bu-1076, male.Myanmar: Katchin, from amber mines nearMyitkyina.

ETYMOLOGY: The species epithet is derivedfrom platy (Greek pl ty§, broad) and cerusa(Greek k ra§, horn), in reference to the com-epressed antennae.

ANALYSES

LYGISTORRHINIDAE AND THE

HETEROTRICHA GROUP

Figure 76

Forty-two morphological characters werecoded for all known Mesozoic Lygistorrhin-idae, Palaeognoriste sp. from Baltic amber,and exemplar species of all living genera ofthe family. The genera Bolitophila, Chiletri-cha, Drepanocercus, and Paratinia wereused as outgroups. In previous studies My-cetophilidae was suggested to be a sistergroup to the Lygistorrhinidae, while Bolito-philidae was the sister group to those fami-


Fig. 76. Phylogeny of Lygistorrhinidae based on cladistic analisys of data presented in table 4 (length84, CI 0.65, RI 0.81). n unreversed changes; ▫ homoplastic character transitions. The character numberis indicated above the branch and the state change is indicated below. Bolitophila, Paratinia, Drepan-ocercus, Chiletricha and Thereotricha are the outgroup.

lies plus Sciaridae (Matile, 1990, 1997; Soli,1997). The position of the Heterotrichagroup is disputed, showing affinities withDiadocidiidae, Sciaridae, and Mycetophili-dae, as well as the extinct families Archizel-miridae and Mesosciophilidae (Grimaldi etal., 2003). A data matrix used for cladisticanalysis is presented in table 4. Character de-scriptions are provided in table 5. Analysisof the data matrix using Winnona, (version2.0; Goloboff, 1999) resulted in four trees oflength 83, CI of 0.66, and RI of 0.82. Thestrict consensus of the trees is shown on fig-ure 76.

A previous cladistic analysis of Lygistor-rhinidae (Grimaldi and Blagoderov, 2001)included as the only fossil Palaeognoriste,

considered a sister group to all living lygis-torrhinids. The present analysis yielded thesame topology for Recent genera: basal po-sition of ‘‘L. asiatica’’, and sister-group re-lationships between Seguyola-Loyugesa andLygistorrhina (with Probolaeus as a subge-nus of Lygistorrhina). The Cretaceous generaArchaeognoriste and Lebanognoriste are atthe base of the lygistorrhine clade and rep-resent a basal, paraphyletic stem group.While having superficial similarity with theHeterotricha group of genera in venationpattern, they show strong apomorphies ofLygistorrhinidae in the structure of trans-verse veins, antennae, thorax, and male gen-italia (characters 3, 8, 12, 25, 26, 29, 36, 38,44). The other Cretaceous genera Plesiog-


TABLE 4Data Matrix for Analysis of Lygistorrhinidae

Character descriptions presented in table 5.

noriste, Protognoriste, and Leptognoriste oc-cupy intermediate positions between primi-tive and recent lygistorrhinids. Interestingly,although these genera do not form a mono-phyletic entity, all species have only one veinin the medial fork preserved. The indepen-dent loss of a median vein in various Me-sozoic lygistorrhinids suggests rapid radia-tion of the family in the beginning of its his-tory, followed by replacement of Mesozoicgroups by recent ones.

MYCETOPHILIDAE S.S.Figures 77–79

Sixty-one characters were coded for 38species representing three main lineages ofCretaceous fungus gnats: Sciophilinae (in-cluding 3 recent and 3 fossil species), Lei-inae (3 recent and 13 fossil species), andGnoristinae (9 recent and 7 fossil species).Macrocera and Bolitophila were used as out-groups. A data matrix used for cladistic anal-ysis is presented in table 6. Character de-scriptions are provided in table 7.

Winnona (version 2.0; Goloboff, 1999)was used to search for the most parsimonious

tree (MPT). An heuristic search (tree bisec-tion-reconnection), cutting trees up to fourpoints, yielded eight MPTs of length 311, CIof 0.21 and RI of 0.51. The low indices ofthe trees are due to the very homoplastic da-taset, reflected by numerous cases of inde-pendent occurrence of structures. The strictconsensus of these MPTs (fig. 77a) is an un-resolved bush for Gnoristinae, with Leiinaeparaphyletic with respect to Sciophilinae.Such an unexpected result can be explainedby including in the parsimony analysis twopeculiar fossil genera, Lecadonileia and Dis-paroleia, which share some characters (5, 10,17, 18) with Sciophilinae. Nonetheless, in allthe analyses conducted, Lecadonileia andDisparoleia nested within Leiinae, havingsynapomorphies with other genera of thesubfamily (e.g., characters 12, 29, 47, 50, 51,52). Due to taxon sampling and the limitednumber of characters used, the most parsi-monious result hypothesized synapomor-phies of Sciophilinae as reversals. The alter-native, when characters 5, 10, 17, and 18 ap-pear independently in different clades, is lessparsimonious by two steps (see below). In-


TABLE 5Characters Used in Cladistic Analysis of Lygistorrhinidae

All characters are nonadditive.

terestingly, analyzing the dataset withWinXpiwe (version 1.3; Goloboff, 1997), aprogram intended for analysis of homoplasticdata (see Goloboff, 1993), yields a topologywith Sciophilinae and Leiinae each as mono-phyletic (fig. 77b). The same result occurswhen Lecadonileia and Disparoleia are ex-cluded from the analysis.

Since the monophyly of Sciophilinae andLeiinae is strongly supported (Soli, 1997),the final analysis was conducted in Winnonawith Sciophilinae and Leiinae constrained asmonophyletic. It resulted in 11 trees oflength 313 (only two steps longer than theMPTs found), CI of 0.24 and RI of 0.58. Thestrict consensus tree (fig. 78) of length 333,CI of 0.22 and RI of 0.54 again indicates

Gnoristinae as basal and paraphyletic toSciophilinae and Leiinae. All the fossil spe-cies assigned to extant genera except Syna-pha longistyla form monophyletic cladeswith recent species. Pseudomanota is closelyallied with Paratrizygia and probably repre-sents a stem group to the Azana group ofgenera (Matile, 1998). Drepanocercus andGregikia are at the base of mycetophilids.Tetragoneura and Ectrepesthoneura shouldbe classified in Leiinae, not Gnoristinae, aswas suggested by Vaisanen (1986). On theother hand, his conclusion about Syntemna,traditionally placed in Sciophilinae, is prob-ably right, for on the cladogram it is nestedwithin Gnoristinae.

In general, including fossils into an anal-


Fig. 77. Unconstrained cladistic analyses of Mycetophilidae using Winnona (a) and WinXpiwe (b).a. Strict consensus tree of eight equally parsimonious trees (length 359, CI 0.21, RI 0.51). b. Fittesttree (total fit 5 3056.9, concavity 3).

ysis dramatically affects results (fig. 79). Toa large extent this effect is a result of exel-lent preservation in amber, which has pre-served characters virtually completely. Thetopology of the MPTs is particularly affect-ed, probably due to extensive homoplasy.Homoplasious characters in plesiomorphicfossils not only decrease resolution and in-crease length of the tree but make taxon def-initions diffuse. Nonetheless, including fos-sils in a cladistic analysis has proven to beextremely useful (Gauthier et al., 1988), andthis study is an another example of that.First, fossils are the only way to establishexistence and position of entirely extinctlineages, like Mesozoic lygistorrhinids orleiines from Burmese amber. Secondly, theyallow the identification of stem groups for

extant lineages. Basal taxa have a funda-mental influence on topology. This is espe-cially true for ancient groups that had a rap-id initial diversification, where unusualcombinations of characters occur.

DISCUSSION

PHYLOGENY

A systematic review of Bibionomorphaand Sciaroidea was not the purpose of thiswork and is well beyond the scope of ourproject. We concur with the composition andevolutionary history of the infraorder as es-tablished by Shcherbakov et al. (1995), andalso grant that fossils afford unique insightinto sciaroid phylogeny.

The traditional nomenclature of venation


TABLE 6Data Matrix for Analysis of Gnoristinae, Sciophilinae, and Leiinae

Character descriptions presented in table 7.

in Sciariodea is controversial and somewhatmisleading. Fortunately, early fossils providea strong basis for vein homology (Rasnitsyn,1980; Shcherbakov et al., 1995), since theyshow intermediate conditions. The short veinof the radial sector is usually thought to beR4 (Chandler, 2002). There are four veins inthe radial sector of the dipteran ground plan.

All representatives of Bibionomorpha s.l. (in-cluding Axymyiiformia and Anisopodifor-mia of Shcherbakov et al., 1995) have nomore than three radial veins posterior to R1,which can be interpreted as R2, R3, and R415

or R213, R4, and R5 according to the authors’concepts of vein homology. We prefer thesecond scheme, assuming that a long vein


TABLE 7Characters Used in Cladistic Analysis of Gnoristinae, Sciophilinae, and Leiinae

All characters are nonadditive.

(R4) is less likely to be atrophied and lostthan is a shorter vein (R2).

Basal Bibionomorpha (Procramptonomyi-idae, Protorrhyphidae) have crossvein r-msituated between the bases of R213 and R4.The earliest Bibionomorpha have two oppos-ing tendencies in the transformation of radialveins. In one group (for example, Triassic

Yala), the base of the forks of radial veinsare shifted basally, so bases of the second(R213) and the third (R4) radial veins are veryclose, and the veins may originate even be-fore the base of r-m crossvein. These veinsare parallel and close to each other along thewhole length, which makes fusion of theseveins more likely. The fusion is completed,


Fig. 78. Phylogeny of Mycetophilidae based on cladistic analysis of data presented in table 6 (length342, CI 0.22, RI 0.53). n unreversed changes; ▫ homoplastic character transitions. The character numberis indicated above the branch and the state change is indicated below. Macrocera and Bolitophila arethe outgroups.


Fig. 79. Comparison of topologies of trees for fossil and recent species (right) and recent speciesonly (left). Tree on the right is the same as on figure 77. Tree on the left is a strict consensus of 6 MPT(length 157, CI 0.40, RI 0.43).


for example, in Pachyneuridae (where veinsare fused basally) and Cramptonomyiidae(apically). Thus, a fore fork in the radial sec-tor of Pachyneuridae, Axymyiidae, and Per-issommatidae should be considered as R213

1 R4.The other group of Bibionomorpha has a

distal shifting of the bases of these forks, ac-companied by shortening and reduction of R4

(e.g., Alinka, or an undescribed Protorrhy-phidae from the Upper Triassic of NorthAmerica) and shifting of the base of R213

close to the base of r-m (Complecia, Eodi-tomyia) or beyond it (Alinka, Eomycetophi-la). Protopleciidae, a presumed ancestor ofSciaroidea, has only one long intermediatevein in its radial sector originating distad ofthe base of crossvein r-m. Having evidenceof distal shifting of the base of the radialveins in the earliest history of the Bibiono-morpha, and considering that R4 as a shortervein is more likely be reduced than a longR213, we prefer to homologize an intermedi-ate vein in the radial sector of Sciaroideawith R213. A transformation series wouldthus include: from two long veins (in Pro-cramptonomyiidae, Protorhyphidae, and Par-axymyiidae [Paraxymyiinae]), to one longand one short vein (Paraxymyiidae [Eomy-cetophilinae] and Eodytomyiidae), to onelong vein proximad (Crosaphis) or distad ofr-m (Protopleciidae, Pleciofungivoridae, Bol-itophilidae, Ditomyiidae, and Keroplatidae),to finally a short crossvein joining R1 or ab-sent (all other Sciaroidea) (figs. 80, 81).

The posterior fork of Sciaroidea wings issometimes denoted as CuA1 and CuA2 or M4

1 CuA1 and CuA2. However, the fore veinin the posterior fork is undoubtedly a rem-nant of the posterior vein in medial sector,namely M4 (fig. 80). With reduction of thediscal cell and M3 (Heterorhyphidae, Parax-ymyiidae) and basal displacement of tb andM stem (Mesosciophilidae, Heterotrichagroup, Mycetophilidae), M4 lost contact withthe base of M. Vein homology in Diptera wasthoroughly discussed by Shcherbakov et al.(1995).

The monophyly of Sciaroidea is supportedby several synapomorphies (Matile, 1997),including loss of vein R4, abbreviation of thecosta at the apex of the wing, long coxae,and narrow insertion of the abdomen (re-

versed in Sciaridae). The last character (Ma-tile, 1990: 386; 1997: 300) was proposedpresumably to separate conditions observedin Sciaridae and some representatives of theHeterotricha group, when the mesothoracicphragma is greatly enlarged and stronglyprojected into the abdomen. Actually, thenarrow insertion of the abdomen, coupledwith reduced phagma, is limited to Myceto-philidae, Keroplatidae, and Lygistorrhinidae,and apparently developed independently.Other Sciaroidea have a phragma that issomewhat developed, though not projectingdeeply into the abdomen (Chandler, 2002).Chandler (2002) hypothesized that a largephragma protruding into the abdomen maybe a synapomorphy of Sciaridae and the Het-erotricha group (except Pterogymnus andSciaropota). Unfortunately, the structure ofthe phagma is rarely seen on fossil speci-mens, even ones in amber, so in most caseswe can only speculate on it. That is why thisundoubtedly important phylogenetic charac-ter was not included in the analyses. How-ever, it is clear that at least in Lygistorrhini-dae, acquiition of a narrow insertion of theabdomen and reduced phragma was not abrief event: early representatives have theplesiomorphic condition (Lebanognoriste,Protognoriste, Plesiognoriste) as well as theapomorphic one (Archaeognoriste, Leptog-noriste). It is possible that the two latter gen-era constitute a monophyletic group with re-cent Lygistorrhinidae.

Reduction of the stem of M (section M1of Kovalev [Kalugina and Kovalev, 1985]),proposed as a synapomorphy of Sciaroidea,actually occurred independently many timesin the history of the group. This section ispresent in Protopleciidae, Pleciofungivori-dae, Bolitophilidae, some Mesosciophilidae,and Keroplatidae. If we accept the reductionof the section M1 as a synapomorphy of Sci-aroidea, the superfamily becames polyphy-letic or leads to an assumption of loss andnumerous independent restorations of thevein, which is highly improbable. Other syn-apomorphies mentioned above combine re-cent families of Sciaroidea (Ditomyiidae,Diadocidiidae, Keroplatidae, Bolitophilidae,Mycetophilidae, Lygistorrhinidae, Sciaridae)with several extinct families (Antefungivor-idae, Archizelmiridae, Eoditomyiidae, Me-


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sosciophilidae, Pleciofungivoridae, and Pro-topleciidae) into one monophyletic groupwith Cecidomyiidae, probably, as a sistergroup to all others.

The phylogeny of recent families of Sci-aroidea was proposed by Matile (1990, 1997)based on imaginal and preimaginal charac-ters, to which extinct groups could be added.Ditomyiidae has been proposed as a sistergroup to all other living Sciaroidea exceptCecidomyiidae; however, no Ditomyiidae areknown in the fossil record before the Eocene(Evenhuis, 1994). Eoditomyia primitiva An-sorge from the Lias of Germany possiblyrepresents a stem-group ditomyiid (Ansorge,1996). The presence of R4 in Eoditomyiidaemay be evidence for an independent originof Ditomyiidae from Paraxymyiidae, whichwould make Sciaroidea polyphyletic. In anycase, discovery of Eoditomyia allows us topropose a Mesozoic, possible Jurassic, originof true, crown-group Ditomyiidae, despitethe absence of fossils.

Matile (1997) provided the only synapo-morphy for Diadocidiidae 1 Keroplatidae,which is the absence or closure of larval ab-dominal spiracles. Chandler (2002) arguedthat the abdominal spiracles may be second-arily lost in Diadocidia due to the tube-livinghabit of larvae. The other putative synapo-morphy of Diadocidiidae 1 Keroplatidaemight be the proximal position of the baseof RS with respect to tb. The same conditionis observed in Australosymmerus, but thisgenus is not considered as basal (Munroe,1974). All other groups of Sciaroidea have(at least in the ground plan) crossvein tbproximal to or at the level of the base of RS.The vertical alignment of veins r-m, M2, andtb in Diadocidiidae and Ditomyiidae, thoughsimilar in shape, probably appeared indepen-dently. Rigidity of the wing is provided by along R1 in Ditomyiidae and by the length-ened base of RS in Diadocidiidae. Thus,crossvein r-m was pulled basad with the baseof RS in Diadocidiidae, while in Ditomyiidaecrossveins r-m, tb, and m-cu were pulled dis-tad together with the base of RS.

Bolitophilidae are represented by only onegenus in the Recent fauna. Another genus—Mangas Kovalev—was described from theLower Cretaceous of Mongolia, and closerelatives were reported from the Lower Cre-

taceous of Transbaikalia (Kovalev, 1986).Chandler (2002) doubted that the position ofMangas was within Bolitophilidae, althoughhe admitted another undescribed but figuredspecimen could belong to the family. Hewrote (2002: 136) ‘‘There is nothing to in-dicate that it is not allied to Heterotricha’’.One of us (V.B.) had the opportunity to studythe type of M. exilis in the collection of PIN.Despite obscure venation, Kovalev’s drawingseems to be accurate and some characters areapparent: long curved base of RS is situateddistally, not basally; r-m is vertical, notoblique; section M2 (not tb, as noted onChandler’s fig. 92) is long and longitudinal(not oblique), although not apparent basallydue to preservation; and there is a short sec-tion M3 (M112 stem) (it is much shorter thanM1 and M2 fork and almost the same lengthas visible part of section M2). This combi-nation of characters is not known in the Het-erotricha group (where the base of Rs andcrossvein r-m are not shifted distally, andsection M2 is much shorter than section M3),but it is characteristic of Bolitophilidae. Bol-itophilidae and the Heterotricha group are arelicts of Mesozoic sciaroids like Protople-ciidae and Pleciofungivoridae, though the re-lationships between bolitophilids and generaallied to Heterotricha are not perfectly ap-parent.

Mycetophilidae and Lygistorrhinidae aresister groups when only Recent taxa are an-alyzed. Including fossils in the analysis in-dicates an independent origin of Bolitophil-idae, Mycetophilidae, Lygistorrhinidae, andpossibly Sciaridae from within the Mesoscio-philidae-Pterogymnus-Heterotricha com-plex. All these recent families as well as ex-tinct Archizelmiridae share synapomorphies,which are also found in Mesosciophilidaeand Heterotricha group. Pterogymnus Free-man is probably related to Mesosciophilidae(Chandler, 2002). One of the Heterotricha-like genera, Sciaropota Chandler, 2002,shows an alliance with an archizelmirid ge-nus, Archimelzira from New Jersey amber(Grimaldi et al., 2003), and probably shouldbe included in Archizelmiridae. Relation-ships within the Heterotricha group cannotbe resolved easily because of the extensivelyparaphyletic nature of this complex.


TAPHONOMY AND BIOGEOGRAPHY

Sciaroidea today are most abundant anddiverse in moist to wet temperate forests ofthe Holarctic and south temperate (e.g., Aus-tral) regions—wherever thick humus andfungal mycelia are abundant. Although sci-aroids are diverse in tropical forests, they arediffuse in numbers there. Among taxa knownin the Cretaceous, Leiinae and Lygistorrhin-idae are better represented in the tropics thanGnoristinae, which are common in temperateregions but scarce in tropical forests.

The most striking aspect of the Cretaceousamber faunas is the one in Burmese amber.Of all insect inclusions in other Cretaceousambers, sciaroids comprise 1% or less ineach deposit: For example, Taymyr 12 sci-aroids/3450 inclusions; Canada 5/1281; Leb-anon 2/600, 2/917, 15/1258 (all of the latterare archizelmirids); and New Jersey 17/1800.Of all insects in Burmese amber, by contrast,5% of them are sciaroids. This is a propor-tion very similar to that found in Baltic andDominican ambers. The proportions of Balticamber sciaroids have been grossly overesti-mated based on museum collections (whichare highly biased) at 15–20%. Based oncrude Baltic amber samples analyzed by PINat the mine sites in Kaliningrad in 1992, sci-aroids (without Sciaridae) comprise 4% of allinsect inclusions. The proportion of Myce-tophilidae in Dominican amber based on un-biased samples is 3.3% of all insect inclu-sions (data of D.G.). Among Cretaceous de-posits the Burmese sciaroid fauna is mostsimilar to Early Cretaceous orictocenoses ofBaisa and Bon Tsagan (Lower Cretaceous ofTransbaikalia and Mongolia), which have1.9% and 1.3%, respectively, of their entireinsect faunas comprised of sciaroids. TheTaimyr, New Jersey, and Canadian ambersciaroid faunas seem the most similar to eachother.

The Burmese amber sciaroid fauna prob-ably reflects a particularly wet paleoenviron-ment, although not necessarily warmer thanthe other Cretaceous deposits. Based on alarge, new assemblage of more than 3000 in-sect inclusions in Burmese amber, Grimaldiet al. (2002) concluded that Burmese amberreflected the most tropical environmentamong all Cretaceous amber deposits. Sci-

aroids present conflicting evidence. On theone hand, sciaroids in Burmese amber wereabundant, as in temperate forests. However,no species is particularly abundant, and Lei-inae and Lygistorrhinidae are quite diverse,as would be found in tropical forests today.

ACKNOWLEDGMENTS

This work would not have been possiblewithout the generous support of Robert G.Goelet, Chairman Emeritus of the AMNHBoard of Trustees, to the senoior author. Weare sincerely grateful to his patronage of col-lection-based research and for his fundingBurmese amber acquisition. The work wasgreatly facilitated by assistance and advicereceived from Jeffrey Cumming, Michael S.Engel, Martin Ramirez, Alexandr P. Rasnit-syn, Andrew J. Ross, Valerie Schawaroch,Toby Shuh, Alexander I. Zaitzev, F. ChristianThompson, and especially from our late col-leagues Loıc Matile and Vladimir V. Zheri-khin. We wish to thank also Peter Chandlerand Andre Nel for their helpful comments ofthe manuscript.

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Plate 1. Families Diadocidiidae and Lygistorrhinidae. A. Docidiadia burmitica Blagoderov and Gri-maldi, holotype AMNH Bu-033. B. Thereotricha sibirica Blagoderov and Grimaldi, holotype PIN 3130/183. C.? T. agapa Blagoderov and Grimaldi, holotype PIN 3426/256. D. Archaeognoriste primitivaBlagoderov and Grimaldi, holotype AMNH Bu-1539. E. Lebanognoriste prima Blagoderov and Gri-maldi, holotype AMNH JG268/1. F. Plesiognoriste carpenteri Blagoderov and Grimaldi, holotypeMCZC 6927.


Plate 2. Family Lygistorrhinidae. A. Plesiognoriste zherikhini Blagoderov and Grimaldi, holotypePIN 3311/664. B. Protognoriste amplicauda Blagoderov and Grimaldi, holotype PIN 3426/257. C. P.goeleti Blagoderov and Grimaldi, holotype AMNH Bu-406. D. P. nascifoa Blagoderov and Grimaldi,holotype AMNH Bu-43. E. Leptognoriste davisi Blagoderov and Grimaldi, holotype AMNH Bu-126a.F. L. microstoma Blagoderov and Grimaldi, holotype AMNH Bu-429.


Plate 3. Family Mycetophilidae: Manotinae, Sciophilini, Gnoristini. A. Alavamanota burmitina Bla-goderov and Grimaldi, holotype AMNH Bu-1271, B. Neuratelia maimecha Blagoderov and Grimaldi,holotype PIN 3311/661. C. Allocotocera burmitica Blagoderov and Grimaldi, holotype AMNH B-056.D. Pseudomanota perplexa Blagoderov and Grimaldi, holotype AMNH Bu-599a. E. Apolephthisa bu-lunensis Blagoderov and Grimaldi, holotype PIN 3963/4. F. Synapha longistyla Blagoderov and Gri-maldi, holotype MCZC 6944.


Plate 4. Family Mycetophilidae: Gnoristini and Leiini. A. Dziedzickia nashi Blagoderov and Gri-maldi, holotype AMNH NJ-117a. B. Saigusaia pikei Blagoderov and Grimaldi, holotype TMPDP79.15.7.21. C. Syntemna fissurata Blagoderov and Grimaldi, holotype TMPD P83.15.3.8. D. Gregikiapallida Blagoderov and Grimaldi, holotype AMNH NJ 117j. E. Gaalomyia carolinae Blagoderov andGrimaldi, holotype AMNH Bu-390. F. Nedocosia exsanguis Blagoderov and Grimaldi, holotype PIN3130/193.


Plate 5. Family Mycetophilidae: Leiini. A. Nedocosia sibirica Blagoderov and Grimaldi, holotypePIN 3311/665. B. N. canadensis Blagoderov and Grimaldi, holotype MCZC 6897. C. N. novacaesareaBlagoderov and Grimaldi, holotype AMNH NJ-117k. D. Ectrepesthoneura succinimontana Blagoderovand Grimaldi, holotype PIN 3311/662. E. E. swolenskyi Blagoderov and Grimaldi, holotype AMNH NJ-824. F. Izleiina mirifica Blagoderov and Grimaldi, holotype PIN 3130/187.


Plate 6. Family Mycetophilidae: Leiini. A. I. spinitibialis Blagoderov and Grimaldi, holotype AMNHNJ-346. B. Zeliinia orientalis Blagoderov and Grimaldi, holotype AMNH Bu-315. C. Z. occidentalisBlagoderov and Grimaldi, holotype MCZC 6943. D. Temaleia birmitica Blagoderov and Grimaldi,holotype AMNH Bu-483a. E. Lecadonileia parvistyla Blagoderov and Grimaldi, holotype MCZC 6941.


Plate 7. Family Mycetophilidae: Leiini. A. Disparoleia cristata Blagoderov and Grimaldi, holotypeAMNH B-0125. B. Hemolia matilei Blagoderov and Grimaldi, holotype AMNH B-0132. C. H. glabraBlagoderov and Grimaldi, holotype AMNH B-0112. D. Protragoneura platycera Blagoderov and Gri-maldi, holotype AMNH Bu-135.

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