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

Copyright q American Museum of Natural History 2004 ISSN 0003-0082 PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY 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 BLAGODEROV 1 AND DAVID GRIMALDI 2 CONTENTS Abstract ....................................................................... 3 Introduction .................................................................... 3 Materials and Methods ........................................................ 4 Systematic Paleontology ......................................................... 6 Superfamily Sciaroidea Billberg, 1820 .......................................... 6 Family Diadocidiidae Edwards, 1925 .......................................... 6 Docidiadia, new genus ..................................................... 6 Sciaroidea incertae sedis ...................................................... 9 Thereotricha, new genus .................................................... 9 Family Lygistorrhinidae Edwards, 1925 ....................................... 12 Archaeognoriste, new genus ............................................... 12 Lebanognoriste, new genus ................................................ 14 Plesiognoriste, new genus ................................................. 14 Protognoriste, new genus ................................................. 17 Leptognoriste, new genus ................................................. 20 Family Mycetophilidae Newman, 1834 ........................................ 22 Subfamily Manotinae Edwards, 1925 ........................................ 22 Alavamanota Blagoderov and Arillo, 2002 .................................. 22 1 Division of Invertebrate Zoology (Entomology), American Museum of Natural History; Paleontological Institute of Russian Academy of Sciences. e-mail: [email protected] 2 Division of Invertebrate Zoology (Entomology), American Museum of Natural History. e-mail: grimaldi@ amnh.org 2 NO. 3433AMERICAN MUSEUM NOVITATES Subfamily Sciophilinae Winnertz, 1863 ...................................... 22 Tribe Sciophilini Winnertz, 1863 ............................................ 22 Neuratelia Rondani, 1856 ................................................. 22 Allocotocera Mik, 1886 ................................................... 24 Pseudomanota, new genus ................................................. 25 Tribe Gnoristini Edwards, 1925 ............................................. 26 Apolephthisa Grzegorgzek, 1885 ........................................... 26 Synapha Meigen, 1818 .................................................... 28 Dziedzickia Johannsen, 1909 ............................................... 28 Saigusaia Vockeroth, 1980 ................................................ 31 Syntemna Winnertz, 1863 ................................................. 32 Gregikia, new genus ...................................................... 32 Gaalomyia, new genus .................................................... 34 Tribe Leiini Edwards, 1925 ................................................ 36 Nedocosia, new genus .................................................... 36 Ectrepesthoneura Enderlein, 1911 .......................................... 39 Izleiina, new genus ....................................................... 41 Zeliinia, new genus ....................................................... 43 Temaleia, new genus ..................................................... 45 Lecadonileia, new genus .................................................. 47 Disparoleia, new genus ................................................... 48 Hemolia, new genus ...................................................... 48 Protragoneura, new genus ................................................. 52 Analyses ...................................................................... 52 Lygistorrhinidae and the Heterotricha Group .................................... 52 Mycetophilidae s.s. .......................................................... 54 Discussion .................................................................... 56 Phylogeny .................................................................. 56 Taphonomy and Biogeography ................................................ 65 Acknowledgments ............................................................. 65 References .................................................................... 65 2004 3BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER ABSTRACT The Recent world fauna of Sciaroidea, or fungus gnats, comprises approximately 4000 described species in eight families: Bolitophilidae, Cecidomyiidae, Diadocidiidae, Ditomyiidae, Keroplatidae, Lygistorrhinidae, Mycetophilidae, and Sciaridae. Larvae live primarily in decaying vegetation, feeding on fungal mycelia, and they can be among the most abundant insects of temperate forests. Stem-group families appeared in the Jurassic, with large Tertiary deposits being composed almost entirely of living genera, so the Cretaceous is essential for understanding the origins and diversification of Recent families. Sixty-six specimens were studied from six major deposits of Cretaceous amber, spanning 40 million years from the Early to 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), and western Canada (80 Ma). New taxa are the following: Docidiadia burmitica (n.gen., n.sp.) (Diadocidiidae); Thereotricha sibirica, (?)T. agapa (n.gen., n.spp.) (Sciaroidea incertae sedis); Archaeognoriste primitiva,Lebanognoriste prima,Plesiognoriste carpenteri,P. zherikhini,Protognoriste amplicauda,P. goeleti,P. nascifoa,Leptognoriste davisi,L. microstoma (n.gen., n.spp.) (Lygistorrhinidae). In Mycetophilidae sensu stricto: Alavamanota burmitina, n.sp. (Manotinae), Neuratelia maimecha, n.sp., Allocotocera burmitica, n.sp., Pseudomanota perplexa, n.gen., n.sp. (Sciophilinae Sciophilini); Apolephthisa bulunensis, n.sp., Synapha longistyla, n.sp., Dziedzickia nashi, 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. swolenskyi, 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.; Disparoleia cristata, n.gen., n.sp.; Hemolia matilei,H. glabra, n.gen., n.spp.; and Protragoneura platycera, n.sp. (Sciophilinae Leiini). Relationships of the fossil genera are phylogenetically assessed with living genera. The Burmese amber fauna contains an inordinate abundance and diversity of sciaroids, perhaps because of a wetter paleoclimate in that region. INTRODUCTION The preservation of fossils in amber is renowned for consistently finer preservation than virtually all other modes of fossilization. Although the geological occurrence of amber begins in the Triassic, the oldest insect faunas in amber are no older than Early Cretaceous, approximately 120–130 million years ago (Ma). Nonetheless, insects in amber from the Cretaceous period (145–65 Ma) afford unparalleled insight into Mesozoic evolution. The many more preserved characters allow more accurate phylogenetic inference than is possible with compression or most other kinds of fossils, particularly for intricate organisms like insects. Detailed preservation facilitates not only reconstruction of phylogeny, but of paleoenvironments as well. In this regard, the study of extinct Diptera is essential, as Diptera usually comprise the largest proportions by individuals and species of all types of arthropods preserved in various amber deposits around the world. This paper is devoted to one particularly diverse group of flies well represented in Cretaceous amber, but which have been virtually neglected: the fungus gnats, or Sciaroidea. The Sciaroidea is comprised of 14 families, 6 of them exclusively Mesozoic: Antefungivoridae (Middle Jurassic to Early Cretaceous; Kovalev, 1990), Archizelmiridae (Late Jurassic to Late Cretaceous; Grimaldi et al., 2003), Eoditomyiidae (Early Jurassic to Early Cretaceous; Ansorge, 1996), Mesosciophilidae (Middle Jurassic to Early Cretaceous; Kalugina and Kovalev, 1985, Blagoderov, 1993), Pleciofungivoridae (Early Jurassic to Early Cretaceous; Kovalev, 1987b), and Protopleciidae (Early Jurassic to Early Cretaceous; Blagoderov, 1996, Kovalev, 1990). No extinct families occur in the Cenozoic. The extant families of the superfamily are Bolitophilidae, Cecidomyiidae, Diadocidiidae, Ditomyiidae, Keroplatidae, Lygistorrhinidae, Mycetophilidae (s.s.), and Sciaridae, all of which but Ditomyiidae also 4 NO. 3433AMERICAN MUSEUM NOVITATES occur in the Cretaceous. Phylogenies and comprehensive revisions of some of these families were provided by Grimaldi and Blagoderov (2001) (Lygistorrhinidae), Matile (1981a, 1990, 1997) (Keroplatidae, Sciaroidea), Munroe (1974) (Ditomyiidae), So¨li (1997) (Mycetophilidae s.s), and Va¨isa¨nen (1984) (Mycomyiini). There are approximately 4100 described species of Sciaroidea exclusive of the poorly studied but diverse families Cecidomyiidae and Sciaridae (Bechev, 2000). At least 10,000 and perhaps as many as 20,000–30,000 living species of Sciaroidea exist. Such diversity reflects the significance of the group in terrestrial ecosystems, particularly forests, the very habitat that produces amber. Larvae of Sciaroidea are abundant in the sporophores of basidiomycete fungi—in decaying wood, leaf litter, and humus, where they feed on fungal mycelia. Where humus and decaying wood are particularly thick in mature forests, sciaroids are among the most abundant and diverse insects, with their larvae inhabiting 80–90% of the macrofungi in natural habitats (Yakovlev, 1988). Larval habits are not restricted, though, to mycophagy (Matile, 1997). The larvae of some genera and species of Keroplatidae are actually predators of insects they snare in their slime trails, the most famous example of which is the New Zealand ‘‘glow-worm’’, (Arachnocampa Edwards). Larvae of Arachnocampa are luminescent and suspend themselves in mucous strands from the ceilings of caves. Small midges attracted to the light become entangled in the strands, which the larvae then devour. There is even a keroplatid, Planarivora Hickman, whose larva is parasitic on land planarians in Tasmania (Hickman, 1964; Matile, 1981b). The fossil record of the Sciaroidea begins in the Early Jurassic (Ansorge, 1996; Kalugina and Kovalev, 1985; Kovalev, 1987a), and by the Early Cretaceous modern families replaced older, stem-group families (Kovalev, 1987a). The Cenozoic Sciaroidea are particularly well studied, especially those in Baltic amber (approximately 270 species of Mycetophilidae, as reviewed in the catalogue of fossil Diptera [Evenhuis, 1994]). The Baltic amber sciaroid fauna, like most of the other Cenozoic faunas of these flies, is essentially modern. Prior to the work of the senior author on extensive insect Lagersa¨tte from Asia (Blagoderov, 1995, 1997, 1998a, 1998b, 2000), there were only seven species of sciaroids known from the Cretaceous, which were placed in the Mesosciophilidae or Mycetophilidae (Kovalev, 1986, 1990; Hong, 1992; Ren et al., 1995; Jell and Duncan, 1986; Westwood, 1854). Sciaroids have been known for years to occur in the major deposits of Cretaceous ambers from Taimyr, Siberia (Zherikhin and Sukacheva, 1973; Zherikhin 1978), Burma (Zherikhin and Ross, 2000), New Jersey (Grimaldi et al., 1989), and western Canada (McAlpine and Martin, 1969; Pike, 1994). Only three species, however, were described: Burmacrocera petiolata Cockerell, Sciara burmitina Cockerell (in Burmese amber, thought at the time to be Cenozoic), and Schlueterimyia cenomanica Matile (in Cenomanian French amber). Since then, major new Cretaceous outcrops of amber 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 produced significant numbers of new specimens and taxa of sciaroids. Based on study of the older collections and these newer ones, we significantly extend the known Cretaceous diversity, based on 27 genera (19 of them new) and 39 species (all of them new). M ATERIALS AND M ETHODS Sixty-six specimens from the Early and Late Cretaceous of six countries, and some 10 localities, were examined. We did not study inclusions of Diadocidiidae in Burmese amber from the Natural History Museum, London, since these were on loan. Adults of most extant genera of Mycetophilidae (s.s), and all species of Lygistorrhinidae were examined. Material is housed in the following institutions: AMNH American Museum of Natural History (Division of Invertebrate Zoology), New York, USA MCNA Museo de Ciencias Naturales de A ´lava/ Arabako Natur Zientzien Museoa, Vitoria-Gasteiz, Spain MCZ Museum of Comparative Zoology (Department of Entomology), Harvard Cambridge, Massachusetts, USA 2004 5BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER PIN Paleontological Institute of the Russian Academy of Sciences (Arthropoda Laboratory), Moscow, Russia TMPD Tyrrell Museum of Palaeontology, Alberta, Canada Specimens were derived from the following localities and formations: Alava, Spain: The amber deposit is located on the northern slope of Sierra de Cantabria (A ´lava), about 30 km southeast of the city of Vitoria-Gasteiz, near the village of Pen˜acerrada. A comprehensive description of the amber deposit was given by Alonso et al. (2000). It was assigned to the Nograro Formation, of Aptian–middle Albian age (120– 110 Ma). All of the material is housed in MCMA. Burma (Myanmar): Historically and presently all amber from Burma derives from the northern state of Kachin. Zherikhin and Ross (2000) reviewed historical records of Burmese amber mining and locations. For many years Burmese amber was considered Cenozoic in age; however recently it has been found to be Cretaceous after the discovery of insects in it that are exclusively Cretaceous, like Serphitidae and Stigmaphronidae (Hymenoptera). A collection of approximately 1200 arthropod inclusions resides at the NHM, London, rich in types described by T.D.A. Cockerell and others. Recently, a very large, diverse collection of Burmese amber has been assembled at the AMNH from material recently excavated in Kachin, near villages close to the town of Myitkyina. This material was acquired by Leeward Capital (Calgary, Canada), who provided it to the AMNH. Study of some 3500 organisms in this collection revealed more than 20 Mesozoic insect taxa, confirming the Cretaceous age and even indicating an age of approximately Cenomanian (98–92 Ma) (Grimaldi et al., 2002). Burmese amber is by far the most prolific and diverse Cretaceous amber for Sciaroidea. Canada: Material from Grassy Lake, Alberta, derives from an abandoned coal mine 8 km south and 1.6 km east of the village of Grassy Lake (Pike, 1994). This was collected by T. Pike and housed at the RTMP. This locality was previously cited as ‘‘near Medicine Hat’’ by McAlpine and Martin (1969), who deposited their material at the Canadian National Collection (CNC) of Insects and Spiders, Ottawa. The Grassy Lake amber is from the Judith River Group of the Foremost Formation, dated as Campanian (Pike, 1994; Borkent, 1995), although Grimaldi and Cumming (citing David Eberth, personal commun.) indicated the amber to be Santonian. Amber from Cedar Lake, Manitoba was collected by F.M. Carpenter in 1938 among beach debris (in the MCZ). Although redeposited and somewhat distant from the Grassy Lake deposit, the two deposits are thought to be contemporaneous (Borkent, 1995). An age of 85–80 myo is a reasonable estimate for Canadian amber. Lebanon: Material studied here is from the Acra Collection (collected by Fadi and Aftim Acra near Jezzine, 30 km east of Saida, Lebanon) and the Estephan Collection (collected by Antoun Estephan, near Bcharre´, northern Lebanon). Both of these collections are housed in the AMNH. Most recently, extensive collections and stratigraphic studies of Lebanese amber have been made by Azar (2000). Lebanese amber was originally reported to be Neocomian in age, pertaining to basal periods of the Early Cretaceous (Schlee and Dietrich, 1970), 145–129 Ma. A younger age of Aptian-Albian (ca. 120–110 Ma) has also been mentioned (Whalley, 1976; Zherikhin, 1978). Azar’s study, which is the most comprehensive, indicates a great range of ages of Lebanese amber, from uppermost Jurassic (152 Ma) to Albian (112 Ma), but most of the deposits yielding insect inclusions are Barremian-Aptian (approximately 125 Ma). Sciaroids in Lebanese amber are the oldest known amber fossils of the superfamily. New Jersey: Rich amber deposits from the central county of Middlesex have been known for decades, summarized by Grimaldi et al. (1989). Recently, an extraordinary outcrop was discovered in the town of Sayreville, with a smaller one being discowered in the adjacent town of East Brunswick (Grimaldi, 2000). This material has produced the oldest definitive ants, tardigrades, mushrooms, parasitiform mites, and flowers (among many others) preserved in amber. Chemical analyses of the amber and stratigraphy of the outcrops have been made. The amber occurs in the South Amboy Fire Clay of the Magothy Formation, Turonian in age 6 NO. 3433AMERICAN MUSEUM NOVITATES (Grimaldi, 2000). All of the New Jersey amber is deposited in the AMNH. Siberia (Taimyr): Reviews of the arthropod fauna in the Siberian amber were provided by Zherikhin and Sukacheva (1973) and Zherikhin (1978). Sciaroidea were found at four localities, ages of which vary. Material was collected by a PIN expedition in 1973 to Nizhnaya Agapa, which is on the north shore of the Agapa River, 40 km downstream from Lake Ladonnakh, Yst’-Enisey depression. This material is in the Dolgan Formation, dated palynologically as Albian– Cenomanian (Saks and Ronkina, 1957). Amber from Yantardakh (‘‘amber mountain’’) occuring 3–5 km upstream from the mouth of the Maimecha River, Khatanga depression, in the Kheta Formation (dated as Coniacian–Santonian [Saks et al., 1959], ca. 87 myo) was collected in 1970 and 1971. A 1976 PIN expedition collected a contemporaneous material from Bulun (middle course of Bulun River [right tributary of Kheta River], 18 km S of post Novaya, Taimyr). Amber from Baikura-Neru (Taimyr Lake, Baikura-Neru Bay) contains the only incomplete specimen of Sciaroidea. Baikura-Neru, age of which is unclear, has been assigned to the Ogneva Formation (Saks et al., 1959; Aptian-Albian, 120–110 Ma), but arthropod inclusions indicate a Late Cretaceous age (Dlussky, 1987; Zherikhin and Eskov, 1999). All Siberian amber is in PIN. Specimens originally stored in small boxes were prepared according to the method described in Nascimbene and Silverstein (2000). This involved embedding the specimen in a stable epoxy (Buehler) under vacuum. The vacuum extracts air in fine cracks, which the epoxy then permeates, thus improving visibility and fragility of the piece. Pieces were then trimmed to thin pieces, with surfaces often to within fractions of a millimeter of the inclusion, in order to optimize observation of details. Prepared in this way, pieces can be mounted on microscope slides and the inclusions observed under 100– 400 3 magnification with a compound microscope. Specimens were measured with a digital stage micrometer mounted under a Zeiss SV8 stereoscope, and photographed using an Infinity K-2 lens attached to a Nikon D-1 camera and illuminated with focusable fiberoptic flash wands (ML-1000 from Microptics). Morphological terminology used follows McAlpine (1981) and So¨li (1997) with modification after Kovalev (Kalugina and Kovalev, 1985). To compare relative position of veins in sciaroid wings, Kovalev defined basal (from the base of Rs to the base of r-m), middle (from the base of r-m to the base of R 4 ), and apical (equivalent to R 5 ) sections in radius stem as RS1, RS2, and RS3 sections, respectively. Also, the basal (from the base of the wing to tb), middle (from tb to r-m), and apical (from r-m to the base of the M 1 and M 2 fork, equivalent to the stem of M 1 1 2 ) 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 sections together. The term ‘‘base (or stem) of M 1 and M 2 fork’’ refers to the point of furcation of the veins. The terms ‘‘stem of M 1 and M 2 fork’’ or ‘‘stem of M 1 1 2 ’’ refer to M3 section. The terms ‘‘R 5 ’’ and ‘‘stem of M 1 1 2 ’’ are used to refer to the veins themselves, and ‘‘RS3’’ and ‘‘M3’’ are used in reference to the relative length of sections. All the veins and section names are summarized in table 1, which shows comparison of the vein nomenclature used by various authors. In some cases, for the sake of stability, we preferred to use traditional vein names in descriptions of species rather than use their names based on presumed, primary homology. These veins are denoted in boldface type in the table. In the discussion of phylogeny, however, it seemed advisable to use names of veins based on homology for establishing transformation series. We used the term ‘‘not visible’’ when a structure could not be observed clearly. SYSTEMATIC PALEONTOLOGY SUPERFAMILY SCIAROIDEA BILLBERG, 1820 FAMILY DIADOCIDIIDAE EDWARDS, 1925 Docidiadia,new genus D IAGNOSIS : Head round. Flagellum 14-segmented. First flagellomere length slightly more than width. Fore tibial comb absent. Wing membrane without macrotrichia. C ends beyond tip of R 5 ; Sc long, ends free; RS base at the middle of R 1 ; crossveins r-m, 2004 7BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER TABLE 1 Nomenclature 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 base of M fork absent; CuA strongly curved back at the apex. A short. Male 9th tergite without marginal bristles, with one large, acute, triangular medial appendage and two small lateral ones; gonostyli do not bifurcate at apex. T YPE S PECIES :Docidiadia burmitica, n.sp. E TYMOLOGY : The name is a feminine anagram of Diadocidia. C OMMENTS : The genus is close to Diadocidia Ruthe, but differs in having the first flagellomere short; wing membrane without macrotrichia, Sc ending free, base of RS rather distal, M3 section and base of M fork reduced; CuA curved at apex rather than with two straight sections; and male tergite IX narrow, triangular, and with two lateral appendages. Diadocidia consists of two subgenera and includes 10 Holarctic species (Chandler, 1994; Lasˇtovka and Matile, 1972; Polevoi, 1996; Wu, 1995; Zaitzev, 1994) and a Neotropical one (Edwards, 1940; Papavero, 1977a), as well as undescribed Australian species (Tonnoir, 1929; Colless, 1963). One species is known from Baltic amber (Evenhuis, 1994). Docidiadia burmitica,new species Figures 1–3 , Plate 1A D IAGNOSIS : As for genus. D ESCRIPTION : Body length 5 1.88 mm (holotype)/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 slightly less than length; apical flagellomere twice the width. Apical flagellomere of male secondarily segmented in two parts. Scape and pedicel turbinate. Only 3 segments of palpi seen, palpomeres cylindrical, subequal in length, basal wider than the rest. Thorax. Scutum setose, dome-shaped, with long protruding setae. Metepisternum bare, height equal to width, shallow incision anteriorly. Wing membrane without macrotrichia. Costa ends beyond tip of R 5 , midway between tips of R 5 and M 1 . Sc ends free, slightly beyond base 8 NO. 3433AMERICAN MUSEUM NOVITATES Figs. 1–3. Docidiadia burmitica, n.sp. 1. Holotype AMNH Bu-033. 2. Tip of the antenna of the holotype. 3. Male genitalia of the holotype. of RS. R 1 setulose, R 5 with sparse setae, almost straight. R 1 short, about 0.6 3 wing length. M3 absent. M 1 and M 2 weakened, their bases absent. M 3 1 4 weakened. Abdomen. Female cerci wide, subtriangular with acute ventral angle. Male tergite IX narrow, triangular with two lateral appendages. Gonocoxites short, with length about the width. Gonostyli massive, length 2.5 3 the length of gonocoxites, hairy, without apical teeth or spines. M ATERIAL : Holotype AMNH Bu-033, male; paratype B-002, female. Myanmar: Katchin, from amber mines near Myitkyina. 2004 9BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER E TYMOLOGY : The species epithet is a reference to Burma, the former name of the country where the amber originates. SCIAROIDEA INCERTAE SEDIS Thereotricha,new genus D IAGNOSIS : Eyes forming incomplete eye bridge, facets large, round. Ocelli three. Flagellomeres barrel-shaped, length no more than 1.5 3 the width. Antepronotum and proepisternum subequal, setose. Proepimeron touches episternum at the episternal suture. Anepisternum smaller than katepisternum. Anepisternal cleft distinct, narrow. Midpleural pit present. Metepisternum setose. Anterior parapsidal suture distinct. Insertion of abdomen wide. Wing membrane with or without macrotrichia. Sc short, ends free. Rs base, r-m, base of M 3 1 4 , and CuA fork very basal. Section M2 connects r-m and base of M 3 1 4 and CuA fork. M 3 1 4 reduced. T YPE S PECIES :Thereotricha sibirica, n.sp. E TYMOLOGY : The genus name is a feminine anagram of Heterotricha. The name is feminine. C OMMENTS : The new genus is close to the Heterotricha group of genera, which have been included in the Sciaridae or Diadocidiidae. Recently, Chandler (2002) reviewed known taxa of the group and described seven more genera from all zoogeographic regions except Nearctic. These taxa seem to represent a stem group of Recent families of Sciaroidea, but monophyly of the group is not apparent. The new genus differs from all taxa of the Heterotricha group in having eyes with large facets, that form an eye bridge, short antennae, the scape and pedicel not differing from flagellomeres in length, palpi very short, anepisternite and katepisternite subequal, the base of RS in basal position, and r-m and the section of M2 subequal, where eyes and palpi demonstrate an apomorphic condition. The new genus resembles Sciaropota Chandler in the porrect antenna with short flagellomeres, an absence of a clearly differentiated series of scutellar bristles, large katepisternum, mesepimepon broader below, but it differs by the short Sc, long stem of M 1 1 2 , and reduced M 3 1 4 . Distinct synapomorphies separate the group from other Mesozoic Sciaroidea: M1 section completely reduced; M2 section fused with tb in one vein meeting the base of M 3 1 4 , and that oblique vein is shifted distad. The same structure of the basal veins is observed in advanced representatives of the Mesozoic family Mesosciophilidae, which are thought to represent a sister group to Mycetophilidae (Kalugina and Kovalev, 1985; Blagoderov, 1993). Similar conditions occur in the peculiar Mesozoic family Archizelmiridae (Grimaldi et al., 2003), but Archizelmiridae have crossvein r-m aligned with M2 1 tb and the basal portion of M 3 1 4 , forming one horizontal vein and the base of RS is shifted distad. Diadocidiidae s.str. (Diadocidia and Docidiadia n.gen.) also have these veins aligned (r-m through the base of M 3 1 4 ), but they form a vertical vein. In Mycetophilidae the combined vein M2 1 tb lost contact with the base of M 3 1 4 and meets the base of CuA or MA (arculus) (see Shcherbakov et al., 1995). Some Mycetophilidae (Drepanocercus,Ectrepesthoneura, Cretaceous Paradzickia,Drepanorzeckia,Ekhiritus,Zazicia) have the fork of M 3 1 4 and CuA sessile or short-stalked, but the base of the fork is situated more basally that in Mesozoic Sciaroidea and the Heterotricha group. Moreover, at least in Ectrepesthoneura the sessile fork of M 3 1 4 and CuA is secondary (see Analyses below and fig. 78). Obviously, reduction of M1 and fusion of M2 with tb might have originated several times in the history of Sciaroidea. Although monophyly of the group combining Recent Heterotrichalike taxa and Cretaceous Thereotricha is not proven, position of these taxa in sciaroid phylogeny should be at the base of lineages leading to Mesosoic Mesosciophilidae and Archizelmiridae and Recent Sciaridae on the one hand and higher sciaroids such as Mycetophilidae and Lygistorrhinidae on the other. Thereotricha sibirica,new species Figure 4 , Plate 1B D IAGNOSIS : Wing membrane without macrotrichia, RS base very basal, oblique, ; 2 3 the length of r-m; M3 section long; base of M 1 and M 2 fork at level of tip of R 1 ; veins R 1 ,R 5 ,M 1 ,M 2 , and CuA with long setae. D ESCRIPTION : Body length 5 1.83 mm (ho- 16 NO. 3433AMERICAN MUSEUM NOVITATES 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 2004 17BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER Costa ends beyond tip of R 5 , one-third the length between tips of R 5 and M 1 .Scvery short, ends free. R 1 short, about 0.5 wing length. R 5 about 0.85 3 wing length, curved slightly, runs very close to R 1 .M 1 confined to apical third of wing. RS base, M 1 and stem of M absent. Crossvein r-m fused with R 5 , horizontal, fused with CuA base. CuA base weak. Base of M 3 1 4 and CuA fork at level of tip of R 1 .M 3 1 4 curved and well rounded. CuA with two straight segments forming obtuse angle. Legs: Coxae almost equal in length. Hind coxae bare at base. Fore tibia distally with apical hemispherical anteroapical depression with comb of fine setae. Tarsal claw without teeth. Abdomen shorter than wings, with 7 visible segments, setose. Sixth and 7th segments twice as short as 5th, 8th retracted. Cerci one-segmented. M ATERIAL : Holotype PIN 3311/664, female. Russia: Taimyr Peninsula, Yantardakh, coll. 1971. E TYMOLOGY : The species epithet is a patronym in honor of the late Dr. Vladimir Zherikhin of the Paleontological Institute in Moscow, prominent paleoentomologist, who collected the specimen. Protognoriste,new genus D IAGNOSIS : Wing membrane without macrotrichia, occiput without row of strong setae. Palpi short. Eyes bare. Face wide. Stem of M and M 2 absent. RS base distinct. Crossvein r-m meets M 3 1 4 and CuA stem. CuA gently curved. Abdomen insertion broad. Fore tibiae shorter than femora. T YPE S PECIES :Protognoriste amplicauda, n.sp. E TYMOLOGY : The name is a combination of protos (Greek pr to§ , or first) and the v ´ genus name Gnoriste. The name is feminine. C OMMENTS : The following apomorphic characters refer the genera Plesiognoriste and Protognoriste to the Lygistorrhinidae: short palpi; small dorsal cleft of anepisternum; anepisternum and metepimeron not divided by laterotergite (possibly plesiomorphic); short, incomplete Sc; short R 1 ; RS base and M stem reduced; r-m horizontal and fused with R 5 ; hind coxa smaller and broader than mid coxa; abdomen insertion narrow; cerci simple. Face wide, with three ocelli, situated almost in straight line, suggestive of Manotinae, but structures of palpi, katepisternum and metepimeron, and absence of strong setae, show no apomorphies with respect to that subfamily. Eye size, face width, vein vestiture, tibial spurs of equal size, and complicated shape of the gonostyli are more primitive conditions than occur in Recent lygistorrhinids, although some features of venation and gonostyli are apomorphic. The short fork of M 3 1 4 and CuA and absence of a long proboscis are most similar to the genus Seguyola Matile. It is possible that these two genera should be treated as a separate subfamily of Lygistorrhinidae. We prefer to expand the definition of the family to include newly described taxa. Protognoriste amplicauda,new species Figure 15 , Plate 2B D IAGNOSIS :R 1 length 3 3 that of r-m. Gonocoxite massive. Gonostyli straight, flattened dorsoventrally. D ESCRIPTION : Body length 5 1.25 mm; wing length 5 1.11 mm. Head: Eyes setulose, setae very short. Ocelli three, equal, almost in straight line, distance of lateral ocellus from eye margin equal to distance from mid ocellus to lateral one. Vertex and frons setose. Scape very small. Pedicel spherical, obscure. Flagellum 14-segmented, flagellomeres cylindrical, lengths about equal to widths, setose. Palpi short, 3-segmented, apical segment rounded, penultimate cylindrical, length 2 3 the width. Face wide, quadrate. Thorax: Scutum with lateral, dorsocentral and acrostichal setae. Scutellum with several long setae. Mediotergite with short trichia. Metepisternum with anterodorsal cleft, height about equal to width. Wing membrane without macrotrichia. Costa ends beyond tip of R 5 , at 1/6 the length between tips of R 5 and M 1 . Sc short, ends free at the level of RS base. Humeral cross-vein transverse. R 1 short, about 0.4 3 wing length. R 5 about 0.8 3 wing length, almost straight. RS base very short, transverse. Crossvein r-m horizontal, meets M 3 1 4 and CuA stem, its length 3 3 less than R 1 length. M 1 almost straight, weakened at base. M 2 and M stem absent. Base of M 3 1 4 and CuA fork between 18 NO. 3433AMERICAN MUSEUM NOVITATES 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, male genitalia. levels of RS base and R 1 tip. CuA and M 3 1 4 curved caudally. Legs: Coxae almost equal in length, hind coxae bare. Tibial setulae in distinct rows. Tibiae, especially fore, short. Tarsal claw with small obtuse tooth. Abdomen as long as wing, with 8 segments, setose. Eighth segment short, retracted. Gonocoxite massive. Gonostyli straight, flattened dorsoventrally. M ATERIAL : Holotype PIN 3426/257, male. Russia: Taimyr Peninsula, Nizhnyaya Agapa, coll. 1973. 2004 19BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER E TYMOLOGY : The species epithet is derived from Latin words amplus meaning ‘‘large, distinguished’’ and cauda meaning ‘‘tail’’ in reference to large terminalia of the species. Protognoriste goeleti,new species Figure 16 , Plate 2C D IAGNOSIS :R 1 length 1.8 3 that of r-m. D ESCRIPTION : Body length 5 1.21 mm; wing length 5 0.95 mm. Head: ovate, height ; 1.5 3 width in profile. Occiput and postgena densely setose. Antennae attached below the middle of the head. Scape and pedicel subconical, wider than flagellum. Flagellomeres cylindrical, length 1.1–1.5 3 the width. Eyes setose, ovate, without incision, facets round, close. Clypeus bare. Two palpomeres seen: basal one ovate, length 2 3 the width, apical round, 4 3 shorter than basal. Thorax: Scutum irregularly setose. Anterior parapsidal suture distinct. Scutellum small. Anepisternum with several setae. Proepimeron touches mesepisternum at anepisternal suture. Anepisternum with deep cleft in the middle. Laterotergite shifted dorsocaudally, so that metepisternum contacts katepisternum for some distance, not in a point. Laterotergites and metepisternum with short hairs. Mediotergite short. Meron at mid coxae large. Wing membrane with microtrichia not arranged in rows. Costa ends beyond tip of R 5 , at one-fourth length between tips of R 5 and M 1 .Scvery short, ends free. Humeral cross-vein oblique. R 1 short, about 0.4 3 wing length. RS base short, transverse. R 5 about 0.8 3 wing length, almost straight. M 1 slightly sigmoid, confined to apical third of wing, ends at tip of wing. Stem of M and M 2 absent. Crossvein r-m horizontal, very weak, meets M 3 1 4 and CuA stem, its length 1.2 3 more than length of R 1 . M 3 1 4 and CuA fork 1.7 3 stem length, its base between levels of RS base and tip of R 1 .M 3 1 4 and CuA curved gently. Legs: Fore tibia 1.25 3 shorter than fore femora. Tibial and tarsal setulae not in rows. Hind tibia with 15 dorsal bristles. Tibial spurs 2 3 the diameter of tibia. Hind tibiae with apical comb of short setae. Fore coxae with dense anterior setae, mid and hind coxae with setae in apical part, hind coxae without posterior setae. Abdomen insertion broad. Abdomen shorter than wings. Cerci with large basal and small round apical segments. M ATERIAL : Holotype AMNH Bu-406, female. Myanmar: Katchin, from amber mines near Myitkyina. E TYMOLOGY : The species epithet in honor of Mr. Robert Goelet, for his generosity in funding purchase of specimens and the authors’ work. Protognoriste nascifoa,new species Figures 17, 18 , Plate 2D D IAGNOSIS : RS base and the base of the M 3 1 4 and CuA fork very basally. R 1 length 4 3 r-m. D ESCRIPTION : Body length 5 1.09 mm; wing length 5 1.05 mm. Head: Occiput and frons setose. Pedicel and scape wider than flagellomeres. Flagellum 14-segmented, flagellomeres cylindrical, length about equal to width. Face wide, quadrate. Clypeus setose, palpi 2-segmented, basal palpomere swollen, apical one very small. Eyes with incision, forming incomplete eye bridge (possibly artifact, face deformed). Thorax: Proepimeron contacts katepisternum. Anepisternum width 1.5 3 height, with distinct cleft posteriorly. Laterotergites, mediotergite and metepisternum with very short trichia. Scutum with numerous erect setae. Metepisternum with long anteriodorsal process touching katepisternum. Wing: Sc very short, free. Costa produced beyond R 5 one-third distance between R 5 and M 1 apices. RS base situated proximally, faint, oblique. R 1 length 4 3 that of r-m. Crossvein r-m weak, 4 3 shorter than R 1 . M 1 originates at distal third of wing, curved at base. Base of the M 3 1 4 and CuA fork at the level of RS base. Legs: Hind coxae shorter than mid ones. Fore coxae with long anterior setae, mid and hind ones with apical setae. Tibial setulae not in rows except apical half of hind tibiae. Hind tibiae long, swollen somewat at apex, with dorsal row of bristles and apical comb of setae. Tibial spur length 1.0–1.3 3 tibial diameter. Abdomen setose, segments short. Genital complex wider than long. Gonocoxites fused. Gonostyli with one outer and two inner lobes. M ATERIAL : Holotype AMNH Bu-434, male. Myanmar: Katchin, from amber mines near Myitkyina. 20 NO. 3433AMERICAN MUSEUM NOVITATES → Figs. 19–24. Leptognoriste, n.gen. 19. L. davisi, n.sp., holotype AMNH Bu-126a. 20. L. davisi, wing of 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. E TYMOLOGY : The species epithet derived from U.S. National Science Foundation, a generous sponsor of this and other fossil insect research at the AMNH. Leptognoriste,new genus D IAGNOSIS : Palpi short, 4-segmented. Mouthparts form short proboscis or reduced. Laterotergites and mediotergite setose. Wing membrane with macrotrichia. Sc long, ends at C. Costa produced beyond R 5 , not reaching wing apex. M 1 absent. M 2 reduced at the base. RS base transverse. M 3 1 4 and CuA fork stalked. Hind coxae shorter than fore and mid ones. Male genitalia simple. T YPE S PECIES :Leptognoriste davisi, n.sp. E TYMOLOGY : The name is a combination of leptos (Greek lept s , or thin, lean) and o ´ the genus name Gnoriste. The new name is feminine. Leptognoriste davisi,new species Figures 19–22 , Plate 2E D IAGNOSIS : The base of M 3 1 4 and CuA fork at level of base of RS. Mouthparts form short proboscis. M 1 reaching wing margin. Ninth tergite small, longer than wider, rectangular. D ESCRIPTION : Body length 5 1.94 mm (holotype)/1.69 mm (paratype); wing length 5 1.26 mm (holotype)/1.27 mm (paratype). Head round with protruding ocelli. Ocelli in triangle, lateral separated from medial by ocellus diameter and from eye margin by 2 ocellus diameters. Eyes large, rounded, with light emargination, with large facets. Clypeus narrow, triangular, setose. Palpi 4-segmented, palpomere length ratio 1:2:2:3. Apical palpomere attached preapically. Short proboscis, one-half head height. Wing length equal to abdomen length. Sc produced slightly beyond RS base. R 1 length 1.2 3 r-m. M 2 originates in distal half of wing. Base of M 3 1 4 and CuA fork at level of RS base. Macrotrichia numerous in basal part of wing. Thorax: Scutum with long lateral, dorsocentral and acrostichal and shorter irregular setae. Anterior parapsidal suture distinct. Antepronotum and proepisternum setose. Anepisternum wider than its height. Proepimeron touches katepisternum slightly below anepisternal suture. Mesepisternum with long anterodorsal process, touching anepisternum. Laterotergites with row of long setae. Mediotergite irregularly setose with short setae. Legs: Coxae with relatively short setae. Hind tibiae with bristles in dorsal row, 1.5 3 longer than femora. Abdomen setose, except first sternite. Abdomen insertion very narrow. Tergite 8 shorter than sternite. Tergite 9 narrow, rectangular with numerous setae at apex. Gonocoxites rather slender. Gonostyli slightly curved. M ATERIAL : Holotype AMNH Bu-126a, male; paratype AMNH Bu-126b, male, in the same piece. Myanmar: Katchin, from amber mines near Myitkyina. E TYMOLOGY : The species epithet is a patronym for Mr. Jim Davis, who supplied the AMNH with fossiliferous amber from Myanmar. Leptognoriste microstoma,new species Figures 23, 24 , Plate 2F D IAGNOSIS : Mouthparts reduced. R 1 shorter than r-m. M 2 weak at apex. Base of M 3 1 4 and CuA fork beyond the level of RS base. Male 8th tergite wider than long, rounded. D ESCRIPTION : Body length 5 1.33 mm; wing length 5 1.24 mm. Head: Occiput with long setae. Pedicel slightly wider then scape and flagellum, flagellomeres barrel-shaped, as long as wide. Palpi 4-segmented, palpomere length ratio 1:2:2:3.5. Thorax: Scutum with long lateral, acrostichal, and dorsocentral setae and short setae between rows. Proepimeron touches katepisternum below the episternal suture. Anepisternum width about equal to height. Laterotergites with long setae. Mediotergite with several long setae ventrally. Mesepisternum with long anterodorsal process, touching anepisternum. Wing length 2004 21BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 22 NO. 3433AMERICAN MUSEUM NOVITATES 1.5 3 the abdomen length. R 1 slightly shorter than r-m. Macrotrichia in basal part of wing absent. M 2 weak at apex. Base of M 3 1 4 and CuA fork beyond the level of RS base. Legs: Fore coxae with dense long anterior setae, mid and hind ones with apical. Hind tibiae without dorsal bristles, only slightly longer than femora. Abdomen. Tergite 9 wider than long, rounded on apex. Sternite I bare. Gonostyli curved at apex. M ATERIAL : Holotype AMNH Bu-429. Myanmar: Katchin, from amber mines near Myitkyina. E TYMOLOGY : The species epithet is a combination of micros (Greek mikroz , or small) and stoma (Greek st ma , or mouth), in refo ´ erence to the reduced mouthparts. FAMILY MYCETOPHILIDAE NEWMAN, 1834 SUBFAMILY MANOTINAE EDWARDS, 1925 Alavamanota Blagoderov and Arillo, 2002 Alavamanota Blagoderov and Arillo, 2002: 34. D IAGNOSIS : Antepronotum and proepisternum completely divided; mediotergite and laterotergites bare; wing membrane with or without macrotrichia; R 1 relatively short, length of R 1 about the length of r-m;R 4 present; M 3 1 4 and CuA fork with stem. T YPE S PECIES :Alavamanota hispanica Blagoderov and Arillo, 2002. C OMMENTS : Closest to the recent genus Manota Williston, 1896, distinguished by having the fourth palpomere attached preapically but very close to the apex of the third; two crossveins instead of one between R 1 and R 5 (RS1 and R 4 ); tibial trichia irregularly arranged; sternite 9 separate; sternite 8 without 4 strong protuberances bearing long setae; and the basal segment of each cercus small. Alavamanota burmitina,new species Figure 25 , Plate 3A D IAGNOSIS : Flagellum compressed dorsoventrally; mesonotum with long lateral setae; wing membrane with macrotrichia; length of small radial cell 6 3 width; base of M 1 and M3 section weak; tibial setulae arranged in rows. D ESCRIPTION : Body length 5 2.50 mm (holotype)/1.62–2.5 mm (paratypes); wing length 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, setae length 2 3 facet diameter. Facets round, densely set. Three ocelli in triangle, close to each other. Frons and face setulose. Antennae inserted above middle of head. Scape and pedicel subconical, with apical setae. Flagellum 14-segmented, flagellomeres compressed, widths 1.2–2 3 length, apical one conical. Texture of flagellomeres polygonlike. Only 3 segments of palpi visible, antepenultimate and penultimate with strong setae; penultimate 2.5 3 as long as preceding one, ovate; apical segment 1.7 3 length of penultimate one, narrow. Thorax: Scutum uniformly densely setose with short setae, bearing long lateral and posterior setae. Anterior parapsidal suture distinct. Suture between antepronotum and proepisternum complete, both segments uniform and covered with bristles. Scutellum with long setae. Anepisternum wider than higher, with wide dorsal cleft. Anepisternal suture declines posteriorly. Katepisternum setose ventrally. Mediotergite and laterotergites bare. Metepisternum with several light hairs posteroventrally. Wing membrane with macrotrichia; microtrichia not arranged in rows. Costa ends slightly beyond the tip of R 5 , C and R 5 run very close to each other in apical part. R 1 ,R 5 ,r-m, M 1 , and M 2 with setae ventrally and dorsally. Sc very short, ends free. Humeral vein oblique. R 1 about equal to length of r-m. Ratio of r-m and RS2 section is 1: 1.4. Base of M 1 and M 2 fork at the level of R 4 . Small radial cell with length 6 3 width. Crossvein r-m horizontal, fused to tb, meets MA. M 3 1 4 and CuA fork long, but not sessile. CuA curved caudally. Legs. Hind coxae bear apical only, not posterior setae. Tibial spur lengths ca. 4 3 tibial diameter. Tibial trichia in rows. Abdomen setose, with 6 visible segments. Gonocoxites fused, lighter then abdomen, setose, swollen. Gonostyli not seen. M ATERIAL : Holotype AMNH Bu-1271, male; paratypes AMNH Bu-428a, Bu 279a, males. Myanmar: Katchin, from amber mines near Myitkyina. SUBFAMILY SCIOPHILINAE WINNERTZ, 1863 TRIBE SCIOPHILINI WINNERTZ, 1863 Neuratelia Rondani, 1856 Neuratelia Rondani, 1856: 195. Anaclinia Winnertz, 1863:770. 2004 23BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER Figs. 25, 26. Alavamanota and Neuratelia.25. A. burmitina, n.sp., holotype AMNH Bu-1271. 26. N. maimecha, n.sp., holotype PIN 3311/661. 24 NO. 3433AMERICAN MUSEUM NOVITATES Fig. 27. Allocotocera burmitica, n.sp., holotype AMNH B-056, reconstruction of wing venation. Proanaclinia Meunier, 1904:145. Odontopoda Aldrich, 1897: 187. D IAGNOSIS : As given by Vockeroth (1972): Tibiae with distinct bristles, length of tibial spurs twice the tibia diameter; anepisternum, mesepimeron, and metepisternum bare; wing membrane with macrotrichia; sternite 8 of male large; gonocoxites partly or completely fused ventrally, gonostyli complex, subdivided or with elaborate processes. T YPE S PECIES :Mycetophila nemoralis Meigen, 1818: 256 (orig. des.). C OMMENTS : Two species of the genus were described from Baltic amber (Meunier, 1904) Neuratelia maimecha,new species Figure 26 , Plate 3B D IAGNOSIS : Sc short, ends before RS base; length of RS1 section 2.5 3 that of crossvein r-m; base of M 3 1 4 and CuA fork at the middle of r-m; mediotergite with sparse, fine trichia. D ESCRIPTION : Body length 5 2.11 mm (rest); wing length 5 2.05 mm. Head: Flagellum 14-segmented, flagellomeres cylindrical, with length about equal to width. Scape and pedicel rounded. Mouthparts form proboscis slightly shorter than head height. Palpi 4-segmented, basal and antepenultimate segments oval, penultimate and apical segment bacilliform, apical slightly longer and narrower than penultimate, length ratio 1:3:4: 5.5. Thorax: Scutum with numerous, short, scattered setae and long lateral ones. Antepronotum with three setae, proepisternum with five. Anepisternum bare, with deep cleft in posterior part. Anepisternal suture declines posteriorly. Metepisternum quadrate, touching katepisternum. Laterotergites with several setae, mediotergite with sparse, fine trichia caudally. Wing membrane with microtrichia and few macrotrichia. Costa ends beyond tip of R 5 . Sc meets C just before RS base. Sc 2 absent. Longitudinal veins with setae. M 3 1 4 and CuA base slightly distad of M3 base. Legs: Tibiae with distinct bristles. Fore tibia longer than first tarsomere. M ATERIAL : Holotype PIN 3311/661, end of abdomen not preserved, sex unknown. Russia: Taimyr Peninsula, Yantardakh, coll. 1971. E TYMOLOGY : Species epithet is derived from the Maimecha river on the Taimyr Peninsula, Siberia, where the amber deposit is located. Allocotocera Mik, 1886 Eurycera Dziedzicki, 1885:166. Allocotocera Mik, 1886: 102. Euryceras Marshall, 1896: 291. D IAGNOSIS : Laterotergites and mediotergite setose. Wing membrane with macrotrichia and with or without microtrichia. Sc 2 preapical or medial. Base of M 3 1 4 and CuA fork before the base of the fork of M 1 and M 2 . T YPE S PECIES :Eurycera flava Dziedzicki, 1885: 167 [ 5 pulchella (Curtis, 1837)] (by monotypy). C OMMENTS : Another fossil species occurs in Lower Cretaceous Spanish amber (Blagoderov and Arillo, 2002). Allocotocera burmitica,new species Figure 27 , Plate 3C D IAGNOSIS : Wing membrane with macroand microtrichia. Segment M3 2.5 3 length of r-m, 0.3 3 that of M fork. Sc ends at level of RS base; Sc2 in the apical one-fourth of Sc, distal to midpoint of Sc. D ESCRIPTION : Body length 5 2.86 mm; wing length 5 2.46 mm. Head: Flagellum 14-segmented, thickened to apex. Flagellomeres almost equal in length, length of the first one 2 3 its width, apical one 5 3 its width. Scape and pedicel subconical, with small apical setae, 2 3 wider than flagellomeres. Eyes pubescent. Two palpomeres seen, cylindrical, length ca. 4 3 the width. Thorax: Scutum irregularly covered with short setae and with strong, long lateral setae. Antepronotum with long setae. Scutellum with two pairs of long setae and several more 2004 25BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER short ones. Laterotergites and mediotergite with setae. Metepisternum with two short setae posteroventrally. Wing membrane clear, with macroand microtrichia. Costa ends at tip of R 5 .Sc meets C beyond RS base. Sc 2 at level of base of section M3. Section of RS1 transverse. R 1 rather short, about 0.7 3 wing length. R 5 curved caudally, not reaching wing apex. Crossvein r-m approximately 2.5 3 as long as section RS1 and 2.5 3 length of section M3. Fork of M 1 and M 2 3.3 3 as long as section M3. Base of M 3 1 4 and CuA fork slightly before base of section M3. Legs: Hind coxae with long posterior setae. Fore tibiae longer than first tarsomere. Cerci 1-segmented, 2 3 as long as 8th tergite, setose. M ATERIAL : Holotype AMNH B-056, female. Myanmar: Katchin, from amber mines near Myitkyina. E TYMOLOGY : The species epithet is a reference to the country of origin of the amber. C OMMENTS : The species is very close to A. xavieri, but differs in having a shorter Sc vein and a longer M 1 and M 2 fork. Pseudomanota,new genus D IAGNOSIS : Infraorbital setae absent. Three palpomeres, penultimate one with long thin apical appendage. Antepronotum wide. Proepisternum rodlike. Laterotergites and mediotergite bare. Sc short, free. Wing membrane with macrotrichia. Base of M 1 and M 2 fork reduced. M 1 reaches wing margin before wing apex. M 3 1 4 free at the base. Gonostyli simple, slightly curved. T YPE S PECIES :Pseudomanota perplexa n.sp. E TYMOLOGY : The genus name is a combination of pseudos (Greek ce dos , or a lie, a y ˘ fraud) and the genus name Manota. The name is feminine. C OMMENTS : Some structures, especially the wide quadrate face, long and flexible apical palpomere, short Sc and R 1 , horizontal r-m, and reduced venation are like Manotinae. Also, unlike typical Sciophilinae, the new genus has laterotergites and especially the mediotergite bare. Nevertheless, the genus is attributed to the subfamily Sciophilinae based on a long, rodlike proepimeron, the rather distal position of humeral vein, and long and slender gonocoxites (which are not known for any manotines). Within Sciophilinae the new genus might be close to the Azana group of genera (Matile, 1998), demonstrating reduction of median veins. All of the genera in the group have reduced venation, with CuA simple and not forming a fork with M 3 1 4 . Matile (1998) supposed that M 3 1 4 in this group was reduced completely, while the fork of M 1 and M 2 lost its base. Jugding on the position of vein apices, at least Paratrizygia,Neoaphelomera, and Neotrizygia may have lost M 2 , and an incomplete vein between M 1 and CuA may be homologous to M 3 1 4 . The new species has venation more primitive than the genera in the Azana group, preserving all the longitudinal veins, although without bases. Pseudomanota perplexa,new species Figures 28–30 , Plate 3D D IAGNOSIS : As for the genus. D ESCRIPTION : Body length 5 1.44 mm; wing length 5 1.07 mm. Head: Eyes large, setulose, with large facets. Occiput and postgena setose, without erect bristles. Ocelli not seen. Antennae attached above middle of head. Frons, face and clypeus setose. Pedicel subconical. Flagellum 14-segmented, slightly thicker toward apex; flagellomeres cylindrical, width 1–1.5 3 length. Three palpomeres seen, basal one heart-shaped, penultimate one with long thin apical appendage about one-third its length, apical one long and slender. Thorax: Antepronotum and proepisternum wide, completely divided, setose. Proepimeron narrow, rodlike. Scutum with lateral, acrostichal, and dorsocentral setae with bare strips in between. Anterior parapsidal suture distinct. Anepisternal suture declines posteriorly. Anepisternum with wide cleft. Metepisternum with long, narrow anterodorsal process touching laterotergite. Laterotergites and mediotergite bare. Wing membrane with macrotrichia and microtrichia. Humeral vein slightly distad of MA. Costa extends to R 5 apex at two-thirds distance between tips of R 5 and M 1 , not reaching wing apex. Length of R 1 is 0.6 3 wing length. Small radial cell length 5 3 width. M 1 originates at apical one-third of wing, reaching wing margin before wing apex. M 2 originates at apical 32 NO. 3433AMERICAN MUSEUM NOVITATES CuA curved caudally. Legs: Tibial bristles 2 3 tibial diameter. Tarsal claw with one tooth. Abdomen: Sternite 8 with two triangular, rounded, caudal lobes covered by long setae. Cerci two-segmented. Basal segment large and broad, apical one 0.5 3 the length and 0.5 3 the width of basal one; both setose. M ATERIAL : Holotype TMPD P79.15.7.21, female. Canada: Alberta, Grassy Lake, coll. T. Pike. E TYMOLOGY : The species epithet is patronymic for Dr. T. Pike, who collected the specimen. C OMMENTS : This species has almost all the features of Saigusaia (table 3, see also Vockeroth, 1980), so despite the long Sc vein its placement in the genus is virtually certain. Syntemna Winnertz, 1863 Syntemna Winnertz, 1863: 767. D IAGNOSIS : Mediotergite bare; laterotergites setose; wing membrane with macrotrichia; Sc meets R; radial cell very small, subquadrate; base of M 3 1 4 and CuA fork proximal to base of M 1 and M 2 fork; segment 7 of abdomen reduced. T YPE S PECIES :S. morosa Winnertz, 1863: 767, by monotypy. C OMMENTS : About 20 extant species are known from the Holarctic (Zaitzev, 1994), with an additional 12 species described from Baltic amber (Meunier, 1904, 1917a, 1922) and 4 from the Lower and Upper Cretaceous of northeast Asia (Blagoderov, 1995, 1998a, 2000). Syntemna fissurata,new species Figures 39, 40 , Plate 4C D IAGNOSIS : Sc ends at R at the base of RS; R 4 present; crossvein r-m 0.5 3 length of M3 section; base of fork of M 3 1 4 and CuA proximal to the base of M 1 and M 2 fork. D ESCRIPTION : Body length 5 3.06 mm, wing length 5 2.28 mm. Head not fully visible. Flagellum 14-segmented, flagellomeres barrel-shaped, length of each 1.5 3 the width, setose, length of setae about equal to flagellomere width. Thorax: Scutum with strong, long setae in lateral, dorsocentral, and acrostichal rows, with short setulae. Antepronotum, proepisternum, laterotergites, and mediotergite setose. Wing membrane with microtrichia and short macrotrichia. Costa ends beyond the tip of R 5 , one-fourth distance between tips of R 5 and M 1 . Sc meets R just before base of RS. Sections of RS1 and RS2 equal and 1.5 3 length of r-m. Crossvein r-m 0.5 3 length of M3 section. Fork of M 1 and M 2 3.5 3 length of its stem (M3). Base of fork of M 3 1 4 and CuA proximal to the base of M2 section. Legs: Inner surface of mid tibia with fissurelike sensory pit in apical third. Tarsal claws with two short teeth at the base. Abdomen setose, with 7 visible segments, 7th one very small. Cerci 2-segmented, basal segment roundish, with numerous long trichia; apical segment bacilliform, with length twice the width, bare. Lateral lobes of 8th sternite setulose. M ATERIAL : Holotype TMPD P83.15.3.8, female. Canada: Alberta, Grassy Lake, coll. T. Pike E TYMOLOGY : The species epithet is the Latin word fissures, meaning ‘‘full of cracks’’, in reference to the state of preservation of the specimen. C OMMENTS : The new species differs from all other known Mesozoic species in having section M3 long, at least twice the length of crossvein r-m. Gregikia,new genus D IAGNOSIS : Costa virtually ends at the apex of R 5 . Sc bare. Sc 2 apical. Section M2 setose. R 5 not reaching wing apex. M3 weak. Acrostichal setae absent. Tergite 9 short. Gonostyli well developed. T YPE S PECIES :Gregikia pallida, n.sp. E TYMOLOGY : The name is derived from some letters commonly used in names of genera in this complex. The name is feminine. C OMMENTS : The genus is very similar to Palaecomoptera Blagoderov, 1997, five species of which were described from the Lower Cretaceous of Transbaikalia and Mongolia (Blagoderov, 1997, 1998a), as well as to three monotypic genera: Grzegorzekia Edwards, Creagdhuhia Chandler, and Phoenikiella Chandler. It is distinguished from Palaecomoptera by R 5 not reaching wing apex, costa not produced after R 5 apex, and base of fork of M 1 and M 2 weak. The genus dif- 2004 33BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 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 genitalia dorsally. 43. Male genitalia ventrally. 34 NO. 3433AMERICAN MUSEUM NOVITATES fers from Grzegorzekia,Creagdhuhia, and Phoenikiella by a short apical palpomere, absence of acrostichal setae, weak M stem and base of fork of M 1 and M 2 , absence of setae on A, from Grzegorzekia also by having Sc bare. Gonocoxites of the new genus also have two lobes as in the last three genera, but the gonostyli are better developed. This state is undoubtedly more plesiomorphic than the highly modified genital complex of Grzegorzekia,Creagdhuhia and Phoenikiella. Species of Palaecomoptera have gonostyli developed as well, but some (P. shcherbakovi,P. lukashevichae) have tergite 9 large, fully covering the gonocoxites. Gregikia pallida,new species Figures 41–43 , Plate 4D D IAGNOSIS : As for genus. D ESCRIPTION : Body length 5 3.77 mm (holotype)/3.09–3.65 mm (paratypes); wing length 5 3.26 mm (holotype)/2.36–3.76 mm (paratypes). Head: Vertex setose. Scape and pedicel small. Flagellum 14-segmented. Flagellomeres cylindrical, length 2 3 the width, covered by trichia, length of trichia about one-half flagellomere width. Only three segments of palpi visible; segments short, nearly cylindrical, combined length slightly less then head height. Thorax: Scutum with erect setae, arranged in rows with wide bare strips between them, acrostichal setae absent. Antepronotum and proepisternum setose. Proepimeron touches mesepisternum at anepisternal suture. Scutellum with four pairs of setae. Anepimeron very narrow ventrally, so that laterotergite touches katepisternum. Katepisternum larger than anepisternum. Mediotergite, laterotergites, and metepisternum bare. Wing: Costa ends slightly beyond tip of R 5 . Sc meets C at level of RS base. R stem, R 1 ,R 5 , and distal parts of M 1 and M 2 with setae. R 5 not reaching wing apex. Section RS1 oblique. R 4 transverse. Ratios of lengths of RS1, RS2, and RS3 is 1:1.5:12– 15. Length of crossvein r-m about equal to RS1 and 0.25 3 M3 section. M 1 subparallel to R 5 ; veins M 1 ,M 2 ,M 3 1 4 , and CuA divergent. Base of fork of M 3 1 4 and CuA proximal to M3 base. M 3 1 4 and CuA curved gently. M 1 ,M 2 ,M 3 1 4 , and CuA with a few setae on apical part. Legs: Hind coxae with long dorsolateral setae. Mid tibiae without sensory pit. Tarsal claw with one basal tooth. Abdomen densely setose. Segments 7 and 8 short, about one-half length of segment 6. Tergite 9 short, does not cover gonocoxites. Gonocoxites densely setose, with long, slim apical and basal lobes, pointed caudally and bearing numerous inner setae. Gonostyli ovate, bare, with strong dark apical processes.M ATERIAL : Holotype AMNH NJ 117j, male. Paratypes: AMNH NJ 117i, male (see fig. 74 for the scheme of syninclusions), coll. P. Nascimbene; AMNH NJ 871a and NJ 871b, sex unknown, coll. K. Luzzi. USA: New Jersey, Sayreville. E TYMOLOGY : The species epithet is a Latin word pallidus meaning ‘‘pale’’ in reference to state of preservation of the specimen. Gaalomyia,new genus D IAGNOSIS : Palpi 3-segmented, short. Scutum with long lateral, dorsocentral, and acrostichal setae. Laterotergites bare. Sc ends at R. Base of fork of M 3 1 4 and CuA at the level of r-m base. M3 section approximately equal to r-m. Gonostyli simple. T YPE S PECIES :Gaalomyia carolinae, n.sp. E TYMOLOGY : The name is a feminine anagram of the genus name Aglaomyia. C OMMENTS : The new genus differs form Aglaomyia Vockeroth in having acrostichal setae present, palpi short, Sc ending at R basally of posterior fork, and 7th abdominal segment well developed. Palaeodocosia Meunier has palpi 4-segmented, vein Sc shorter, and the base of fork M 3 1 4 and CuA more proximal. Pseudalysiinia Tonnoir has palpi 4-segmented and incrassate, with no bristles on the thorax and legs, and tibial spurs short. The genus may be close to Ipsaneusidalys Blagoderov, 1998, especially I. longipennis, but the latter differs in having long fork of M 3 1 4 and CuA and M3 section, and R 5 is sinuous. Preservation of I. longipennis seems to be insufficient to decide if both species are congeneric. Gaalomyia carolinae,new species Figure 44 , Plate 4E D IAGNOSIS : As for genus. D ESCRIPTION : Body length 5 2.68 mm; 2004 35BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER Fig. 44. Gaalomyia carolinae, n.sp., holotype AMNH Bu-390. wing length 5 2.34 mm. Head: Occiput and frons densely setose. Eyes large, with round facets, slightly emarginate, setulose. Ocelli absent. Scape and pedicel rounded, flagellum 14-segmented, filiform. Length of 1st and apical flagellomeres 2.5 3 width, with others 2 3 width. Palpi 3-segmented, palpomeres equal in length and subsequently narrower to apex. Length of penultimate one 1.5 3 width, apical one 3 3 width. Clypeus setose. Thorax: Scutum with long lateral, dorsocentral, and acrostichal setae. Proepimeron touches katepisternum at shallow incision. Katepisternum larger than anepisternum. Anepisternal suture horizontal. Antepronotum and proepisternum with long setae, other thoracic sclerites bare. Wing membrane without microtrichia. R, R 1 ,R 5 , and M 1 with dorsal setae. Sc ends at R at the level of base of r-m. R 1 and R 5 straight. Crossvein r-m equal to M3 section. Length of fork of M 1 and M 2 4 3 M3 section. M 1 and M 2 not reaching wing margin. Legs: Fore and mid coxae densely setose, hind coxae with long posterolateral setae. Femora with ventral row of setae. Tibial spurs 2–2.3 3 tibial diameter. Abdomen densely setose. Sternite I bare. Eighth tergite short, length 0.5 3 7th tergite. Tergite IX reniform, width 3 3 length. Gonocoxites slender, length 3.5 3 width, widely separated. Gonostyli saberlike, curved inside. M ATERIAL : Holotype AMNH Bu-390, male. Myanmar: Katchin, from amber mines near Myitkyina. E TYMOLOGY : The species epithet is a patronym honoring our friend and colleague Caroline S. Chaboo, specialist on chrysomelid beetles. 36 NO. 3433AMERICAN MUSEUM NOVITATES TRIBE LEIINI EDWARDS, 1925 Nedocosia,new genus D IAGNOSIS : Three ocelli, lateral ones do not contact eye margin. Sc merges with R. R 1 2–3 3 length of r-m. Veins of median and cubital forks lightly sclerotized. Base of M 3 1 4 and CuA fork lies between levels of base of r-m and M 1 –M 2 fork. Tibial bristles absent. T YPE S PECIES :Nedocosia exsangius, n.sp. E TYMOLOGY : The genus name is derived from the prefix ne-, ‘‘negation’’, and the genus Docosia. The name is feminine. C OMMENTS : This genus is most similar to Docosia Winnertz, 1863, which consists of 25 extant Palaearctic, 15 Nearctic, and 2 Neotropical species (Zaitzev, 1994). Seven species of Docosia are known from Baltic amber (Meunier, 1904, 1916, 1922, 1923) and one from the Oligocene shales of Rott, Germany (Statz, 1944). The new genus differs by having lateral ocelli not touching the eye margins (undoubtedly plesiomorphic), and by the lack of strong tibial bristles. Differences in sclerotization of anterior veins (Sc, R, R 5 ) and posterior veins (forks) in the new genus are not as great as in Docosia species. Two species of Docosia described from the Lower Cretaceous of Transbaikalia (Blagoderov, 1995) may also belong to the new genus, in fact showing affinities to N. novacaesarea. Nedocosia exsanguis,new species Figures 45, 46 , Plate 4F D IAGNOSIS : Sc meets R before level of base of r-m. Laterotergites bare. Cerci with two combs of dark setae. Apices of gonostyli sharp, curved inward at right angle. D ESCRIPTION : Body length 5 1.29 mm; wing length 5 0.67 mm (rest). Head oval, with rounded occiput, occiput and frons with scattered setae. Ocelli three, lateral one about its own diameter from eye margin and two diameters from medial ocellus. Eyes slightly emarginate, with round facets and short interfacetal trichia. Scape subconical, pedicel cylindrical. Flagellum 14-segmented, flagellomeres of hexagonal surface texture, cylindrical, lengths about equal to width. Face and clypeus setose. Thorax: Scutum with setae in lateral, dorsocentral, and acrostichal rows. Antepronotum with 4 long and several shorter setae. Proepimeron very small, touches katepisternum at incision. Laterotergites and mediotergite bare. Metepisternum broader than high. Wing: Sc meets R before M3 base. Costa ends beyond tip of R 5 , more than one-half length between tips of R 5 and M 1 . Crossvein r-m 0.37 3 length of R 1 , and 0.6 3 length of M3 section. Base of fork of M 3 1 4 and CuA between levels of bases of M3 section and M 1 and M 2 fork. M 3 1 4 very weak. M 1 and M 2 fork campanulate. Longitudinal veins with setae, except Sc. Legs: Fore coxae with anterior setae, mid coxa with apical setae. Fore leg with first tarsomere 2 3 longer than second and 0.55 3 length of tibia. Abdomen: Tergites setose, sternites bare. Cerci with two comblike rows of black bristles. Apices of gonostyli sharp, squarish, curved inward. M ATERIAL : Holotype PIN 3130/193, male. Russia: Taimyr Peninsula, Yantardakh, coll. 1970. E TYMOLOGY : Latin, meaning pale or bloodless, referring to the pale color of the specimen. Nedocosia sibirica,new species Plate 5A D IAGNOSIS : Sc meets R at the level of the base of the stem of M 1 1 2 . Laterotergites setose. Anterior edge of fore coxa bare. Hind legs stout. D ESCRIPTION : Body length 5 2.37 mm, wing length 5 1.82 mm. Head: Occiput and frons setose. Ocelli three, equal in size; distance of median to lateral ocellus and from lateral ocellus to eye margin equal to twice their diameter. Eye with slight incision at antennal bases. Antennae short, only 10 flagellomeres preserved; flagellomeres cylindrical, widths 1.1–1.5 3 lengths. Face triangular, setose, clypeus ovate, bare. Palpi 5-segmented, short, antepenultimate segment swollen, with medial sensory pit on inner surface. Apical palpomere shorter than penultimate one, both slender, with sparse setulae. Length ratio 1: 3:3:5:5. Thorax: Mesonotum with short trichia and setae of various sizes; setae arranged in 5 rows. Antepronotum with three setae; proepisternum with two on lower part. Scutellum with two very long setae and two smaller ones. Laterotergites with a few tri- 2004 37BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 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. 38 NO. 3433AMERICAN MUSEUM NOVITATES chia in lower part. Mediotergite bare. Wing: Sc meets R at the level of the base of the stem of M 1 1 2 .R 1 twice the length of r-m. R 5 curved slightly back. C ends far beyond tip of R 5 , but does not reach wing apex. Stem of R, R 1 , and R 5 with setae. Legs: Anterior edge of fore coxae without setae. Fore tibiae with semicircular anteroapical depressed area, covered with numerous small trichia. Mid tibial spur length 2.5 3 apical diameter of tibia, hind tibial spur 1.5 3 diameter. Hind legs stout, longer than mid legs. First fore tarsomere 2.5 3 longer than second and 0.67 3 length of tibia. Cerci simple. M ATERIAL : Holotype PIN 3311/665, female. Russia: Taimyr Peninsula, Yantardakh, coll. 1971. E TYMOLOGY : The species epithet is a reference to the region of origin of the amber. Nedocosia canadensis,new species Plate 5B D IAGNOSIS : Sc meets R before level of base of the stem of M 1 1 2 .R 1 2.5 3 length of r-m. M 3 1 4 and CuA fork at the level of r-m. M3 section shorter than r-m crossvein. Laterotergites and mediotergite bare. D ESCRIPTION : Body length 5 1.50 mm (preserved part). Head: Occiput with setae of different length. Ocelli not seen. Eyes with round facets, setulose. Scape small, pedicel subconical. Flagellum 14-segmented, flagellomeres cylindrical, lengths equal to widths, densely covered by trichia no longer than half width of basal flagellomere. Three segments of palpi seen, combined length equal to head height. Thorax: Scutum uniformly setose with long and short setae. Antepronotum with two very long setae and several short ones. Laterotergites and mediotergite bare. Wing: Sc meets R before level of base of the stem of M 1 1 2 .R 1 2.5 3 length of r-m. Crossvein r-m 1.3 3 length of M3 section. Base of fork of M 3 1 4 –CuA proximal to base of fork of M 1 –M 2 . M ATERIAL : Holotype MCZC 6897, incomplete specimen, sex unknown. Canada: Manitoba, Cedar Lake, coll. F. M. Carpenter. E TYMOLOGY : The species epithet is a reference to the country of origin of the amber. Nedocosia novacaesarea,new species Figure 47 , Plate 5C D IAGNOSIS : Flagellum 11-segmented. Sc long, ends at R between RS and r-m bases. Base of M 3 1 4 and CuA fork at level of r-m. Fore leg with tibia equal to femur or slightly longer. D ESCRIPTION : Body length 5 2.34 mm (holotype)/2.88 mm (paratype); wing length 5 1.75 mm (holotype)/2.20 mm (paratype). Head: Three ocelli in line, distances between median and lateral ones, and between lateral and eye margin, are equal. Face rectangular, bare. Clypeus bare. Scape and pedicel rounded, shorter than flagellomeres. Flagellum 11segmented, flagellomeres cylindrical, with lengths 2 3 width. Two palpomeres seen, apical one 2 3 length of basal. Thoracic sclerites bare. Scutum with long lateral setae and numerous short ones, not arranged in rows. Proepimeron small, fits in shallow incision on katepisternum. Anepisternal suture declines backward. Metepisternum touches anepimeron. Meron large. Wing: Costa ends beyond tip of R 5 , more than one-third length between tips of R 5 and M 1 . Sc long, ends at R between RS and r-m bases. Length of R 1 2.5–3 3 that of r-m, M3 section twice that of r-m. Length of fork of M 1 and M 2 2.5–3 3 M3 section. Base of fork of M 3 1 4 and CuA at level of rmbase. Legs: Fore coxae with short anterior setae, hind coxae with long posterolateral setae. Fore leg with tibia equal to femur. Tibial spurs 1.2–1.5 3 tibial diameter. Abomen setose. Seventh segment very narrow. Gonocoxites massive, with apices blunt. M ATERIAL : Holotype AMNH NJ-117k, male; paratype NJ-117l, male in the same piece. USA: New Jersey, Sayreville, coll. P. Nascimbene. E TYMOLOGY : The species epithet is derived from Latin Nova Caesarea meaning ‘‘New Jersey’’ and is a reference to the state of origin of the amber. C OMMENTS : The species is similar in venation to Docosia baisae Blagoderov, 1998 and D. zaza Blagoderov, 1998, by proximal position of the forked base of M 3 1 4 and CuA, but differs from the latter species in having Sc long and M3 section short. 2004 39BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER Ectrepesthoneura Enderlein, 1911 Willistoniella Meunier, 1904: 74 (preoccupied by Mik, 1895). Meunieria Johannsen, 1909: 87 (preoccupied by Kieffer, 1904). Ectrepesthoneura Enderlein, 1911: 115. D IAGNOSIS : As given by Chandler (1980): Lateral ocelli remote from eye margin; laterotergites bare; Sc ending in R before base of RS; R 4 present; R 1 short, at most twice as long as r-m;R 5 straight; C prolonged well beyond tip of R 5 ; veins of medial and cubital sectors weak and faint; posterior fork sessile. Males with ‘‘sensory pit’’ near the base of mid tibia; hind tibial comb absent. T YPE S PECIES :Tetragoneura hirta Winnertz, 1846: 19 (orig. designation) C OMMENTS : There are about 10 extant species from the Holarctic region. The genus is very close to Tetragoneura Winnertz (about 100 widespread living species) and is sometimes included in it or treated within the tribe Gnoristini (Tuomikoski, 1966; Vaisanen, 1986). There are two fossil species described from Baltic amber (Meunier, 1904) and the Oligocene of Rott, Germany (Statz, 1944). Chandler (1999) suggested that Ectrepesthoneura was paraphyletic. Ectrepesthoneura succinimontana, new species Figures 48, 49 , Plate 5D D IAGNOSIS : Fore tibiae slightly shorter than fore femur. Section M3 and bases of M 1 and M 2 very weak. Length of small radial cell twice the width. D ESCRIPTION : Body length 5 2.39 mm; wing length 5 2.02 mm. Head: Occiput and frons with short curved setae. Ocelli three, distance of lateral one to eye margin more than twice the ocellus diameter. Scape subconical, pedicel rounded, width twice flagellomere width. Eleven segments of flagellum preserved, flagellomeres cylindrical, with length about equal to width. Face setose. Palpi 4-segmented; 1st palpomere very small; antepenultimate one oval, its length twice the width, with medial sensory pit; penultimate one with length 3 3 width, attached to the second one preapically; apical segment long and slender. Clypeus triangular, setose. Thorax: Scutum with numerous long setae mostly curved forward and shorter ones curved back, arranged in rows. Antepronotum with 3 long setae. Scutellum with two pairs of long setae. Other thoracic sclerites bare. Wing: All longitudinal veins with short setae. Sc meets R proximal to base of M3. Length ratio of sections RS1, RS2, and RS3 is 1:2:13. RS1 transverse, 0.3 3 length of r-m. Crossvein r-m 0.7 3 length of R 1 and 0.5 3 length of M3 section. Costa ends beyond tip of R 5 , one-third distance between tips of R 5 and M 1 . Section M3 and M 1 and M 2 bases very weak. Legs: Fore coxae with numerous anteromedial setae, hind coxae with posterolateral setae. Fore and mid tibiae slightly shorter than fore femora. Mid and hind tibiae and tarsi with rows of black bristles. Tibial spurs 1.5–2 3 tibial diameter. Tarsal claw with one long tooth. Abdomen setose. Eighth sternite almost triangular. Cerci 2-segmented, basal segment stick-shaped, with length twice the width, extends beyond 8th sternite; apical one discoidal, attached at the apex of the basal one, 2.2 3 shorter. Dorsal border of 8th sternite (gonocoxite 8 of Martinsen and So¨li, 2000) straight. M ATERIAL : Holotype PIN 3311/662, male. Russia: Taimyr Peninsula, Yantardakh, coll. 1971. E TYMOLOGY : From Latin succinum meaning ‘‘amber’’ and montanus meaning ‘‘mountainous’’. Yantardakh means ‘‘amber mountain’’ in Dolgan language. C OMMENTS : Comparision with recent species is difficult, because nongenital characters used for distinguishing species have been considered to be unreliable for this genus (Martinsen and So¨li, 2000). For species where the female genitalia are known the new species is most similar to E. hirta Winnertz. Ectrepesthoneura swolenskyi,new species Figures 50, 51 , Plate 5E D IAGNOSIS : Sc long, meets R just before base of M3. Small radial cell relatively long. Gonocoxites with triangular ventral appendages bearing 2 long, dark apical spurs. D ESCRIPTION : Body length 5 2.71 mm; wing length 5 2.46 mm. Head: Ocelli three, from lateral ocellus to eye margin twice ocellus diameter. Flagellomeres cylindrical, length 1.5 3 width. Three palpomeres seen, 40 NO. 3433AMERICAN MUSEUM NOVITATES Figs. 48–51. Ectrepesthoneura End. 48. E. succinimontana, n.sp., holotype PIN 3311/662. 49. Female genitalia. 50. E. swolenskyi, n.sp., holotype AMNH NJ-824. 51. Male genitalia. 2004 41BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER apical one slender, length ratio 1:2:3.5. Thorax: Antepronotum with 2 setae, proepisternum with 4. Scutum with setae curved forward, lateral setae very long. Wing: All longitudinal veins except Sc with short setae. Costa ends beyond tip of R 5 , one-half distance between tips of R 5 and M 1 . Sc meets R just before M3 base. Length of R 1 2.2 3 that of r-m. Length of M3 2.7 3 that of r-m. Ratios of lengths of sections RS1, RS2, and RS3 1:2.3:7. RS1 oblique, 0.58 3 length of r-m. M3, M 1 , and M 2 weak, M 1 slightly curved forward. M 3 1 4 and CuA curved gently caudad. Legs: Fore and mid tibiae longer than femora. Mid tibia with anterolateral, posterolateral, and posteromedial rows of short bristles, hind tibia with two rows of posterior setae. Basal third of mid tibia with large, dark, narrow sensory pit. Tibial spurs 1.3–1.7 3 tibial diameter. Abdomen. Ninth tergite large, conceals gonocoxites and gonostyli almost completely; caudal edge straight, without medial cleft. Ninth tergite straight caudally, without cleft, with numerous curved setae apically. Gonocoxites with triangular ventral appendages bearing 2 long, dark apical spurs. Gonostyli short, pointed, curved S-like. M ATERIAL : Holotype AMNH NJ 824, male. USA: New Jersey, Sayreville; coll. S. Swolensky, 1997–1998 (mentioned in Grimaldi, 2000: fig. 48h). E TYMOLOGY : The species epithet is patronymic for the late Steve Swolensky, who collected the specimen. C OMMENTS : The new species differs from other species of the genus in the structure of male genitalia, especially by ventral appendages of gonocoxites bearing dark sclerotized spurs. Izleiina,new genus D IAGNOSIS : Facets round. Lateral ocelli not touching eye margins. Clypeus setose. Sc meets C just beyond level of base of M3; Sc 2 absent. Length of R 1 2 3 that of r-m. Section M3, and fork of M 1 and M 2 –M 3 1 4 reduced; CuA weakened. M 3 1 4 interrupted at base. T YPE S PECIES :Izleiina mirifica, n.sp. E TYMOLOGY : Name is derived from the tribe Leiini. C OMMENTS : We assume that in the new genus vein M 1 is entirely reduced. As a rule, in most leiine genera, vein M 1 reaches the wing margin at or even before the wing apex. They have retained a complete vein that is homologous to CuA based on the length and shape of the vein, and which is strongly curved back and ends at the level of RS1, as in most Leiini. The new genus differs from Neoclastobasis Ostroverchova by the lateral ocelli touching eye margins, and from Clastobasis Skuse and Neoclastobasis by C ending beyond the tip of R 5 , and by the presence of the base of M 3 1 4 . Having R 1 relatively long and Sc ending at C makes the genus similar to Rondaniella Johannsen and Indoleia Edwards, but differs from them by reduction of the veins in the medial sector. Such a reduction also occurs in other leiine genera, for example Novakia Strobl and Sigmoleia Tonnoir and Edwards, but they have Sc very short and free. Species of Cycloneura Marshall and Paracycloneura Tonnoir and Edwards also have M unbranched and M 3 1 4 free or reduced at the base, the anepisternum large, metepisternum narrow like Izleiina, but they differ strongly in the shape of CuA, by having M 2 long, Sc short and free, and by the presence of strong tibial bristles. Izleiina mirifica,new species Figures 52–54 , Plate 5F D IAGNOSIS : Apical palpomere short. Tibiae without bristles. Gonostyli conical, curved inward. D ESCRIPTION : Body length 5 1.44 mm, wing length 5 1.23 mm. Head: Ocelli three, distance from eye margin equal to two ocellus diameters; distance between each other equal to one diameter. Flagellum 14-segmented, flagellomeres cylindrical, lengths about equal to widths, covered by setulae no longer than one-half flagellomere width. Palpi 4-segmented, three basal segments bacilliform, apical one short and rounded. Occiput, frons, face, and clypeus with short setulae. Thorax: Prescutum and anterior parapsidal suture distinct. Antepronotum with 6 setae, proepisternum with two. Scutum arched. Scutellar setae in rows. Scutellum with two pairs of long setae. Cavity on anterior edge of katepisternum shallow. Laterotergites and mediotergite bare. Wing: Sc 48 NO. 3433AMERICAN MUSEUM NOVITATES Canada: Manitoba, Cedar Lake, coll. F.M. Carpenter. E TYMOLOGY : The species epithet is a combination of the Latin word parvus, meaning ‘‘small’’, and stylus, meaning ‘‘spike, stem, pen’’, in reference to simple small gonostyli of the species. Disparoleia,new genus D IAGNOSIS : Ocelli absent; anepisternum with very short setae and trichia; mediotergite and laterotergites bare; wing membrane with microand macrotrichia; Sc ends at C; r-m 2 3 length of R 1 , M3 base at basal onesixth of wing, M3 weak, M 2 absent. T YPE S PECIES :Disparoleia cristata, n.sp. E TYMOLOGY : The genus name is a combination of the Latin word dispar, meaning ‘‘unequal, unlike’’, and the genus Leia,in reference to the distinctiveness of the new genus. The name is feminine. C OMMENTS : The genus differs from all Recent Leiini in the unique combination of such characters as absence of ocelli, very long rmwith the proximal position of the base of M3, and reduction of veins in the medial sector. It differs from Novakia Stro¨bl in having the base of RS distinct, M 2 absent, and Sc ending at C. It differs from Zeliinia, n. gen. in having flagellomeres of smooth texture, ocelli absent, vein M 1 and M3 section present though weak, and fork of M 3 1 4 and CuA long but not sessile. Disparoleia cristata,new species Figures 69 , Plate 7A D IAGNOSIS : As for genus. D ESCRIPTION : Female. Body length 5 2.73 mm, wing length 5 2.08 mm. Head: Vertex with group of long erect setae. Scape and pedicel wider than flagellum. Flagellum 14segmented, flagellate, flagellomeres longer and narrower apicad. Eyes setose, slightly emarginate at antennal base. Ocelli absent. Clypeus setose. Palpi 3-segmented, 1st palpomere short, 2nd longer, cut obliquely, 3rd longer than length of 1st and 2nd, narrow. Thorax: Scutum with lateral and dorsocentral rows of long setae and short irregular setae. Scutellum with numerous short setae. Antepronotum with long curved setae and several short ones. Anepisternum with 4 light short setae in upper part and very fine hairs at lower apical part. Proepimeron touches katepisternum slightly ventral to anepisternal suture. Mediotergite and laterotergites bare. Metepisternum large, with dorsal margin wider than ventral one. Wing membrane with microtrichia and short macrotrichia. Costa ends beyond tip of R 5 , at one-third length between tips of R 5 and M 1 . Humeral vein located beyond the MA. Sc short, ends at C before level of M3 base, apex faint. R 1 very short, r-m 3 3 length of R 5 . Median veins weak. M 1 , base of M 1 and M3 very weak. Base of M3 very proximal, slightly distal to apex of Sc. Crossvein r-m 3 3 length of M2 section, M3 section 3.5 3 that of M2. Base of M 3 1 4 and CuA fork slightly distal to level of base of M3 section. R, R 1 ,R 5 ,M 3 1 4 , and CuA with setae dorsally and ventrally. Leg: Tibial spur formula 1:2:2. Spurs long, 4–5 3 tibia diameter. Mid and hind tibiae with bristles. Empodium absent; tarsal claws with one tooth. Abdomen: Tergites of abdomen setose. Seventh tergite 1.5 3 shorter than 6th, 8th is very short. Cerci 2-segmented, slightly shorter than 6th and 7th segments together. Apical segment round, basal one bacilliform, 2.5 3 apical. M ATERIAL : Holotype AMNH B-0125, female. Myanmar: Katchin, from amber mines near Myitkyina. E TYMOLOGY : The species epithet is Latin word cristatus, meaning ‘‘tufted, plumed’’, in reference to the group of setae on the vertex. Hemolia,new genus D IAGNOSIS : Three ocelli in straight line, middle one small, lateral ones not touching eye margin; wing membrane without macrotrichia; Sc short, ends at R; R 1 short, no more than 2 3 length of r-m;r-m short, oblique; M3 short; tibial setae in rows, mid and hind tibiae with inner spurs longer than outer one. T YPE S PECIES :Hemolia matilei, n.sp. E TYMOLOGY : The genus name is a feminine anagram of Mohelia Matile, 1978. C OMMENTS : The genus is close to Mohelia in the structure of the ocelli, and wing venation, and it differs in tibial spur length and details of male genitalia structure. It differs from Mohelia in having setae on the mediotergite, tibial setae in rows, mid and hind 2004 49BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 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., holotype AMNH B-0112, wing. 50 NO. 3433AMERICAN MUSEUM NOVITATES 2004 51BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER ← Figs. 73–75. Protragoneura platycera, n.sp. 73. Holotype AMNH Bu-135. 74. Male genitalia laterodorsally. 75. Male genitalia lateroventrally. tibiae with inner spurs longer rather than the outer ones, and C ending close to the tip of R 5 ; Sc is longer, R 1 longer than r-m; and M3 is short. Hemolia matilei,new species Figures 70, 71 , Plate 7B D IAGNOSIS : Scutum densely setose; laterotergites setose; Sc rather long, ends at the level of M 3 1 4 and CuA fork base. D ESCRIPTION : Body length 5 2.19 mm (holotype)/2.39 mm (paratype); wing length 1.49 mm (holotype)/2.21 mm (paratype). Head: Eyes setose. Occiput and frons densely setose. Three ocelli in straight line, middle one small; lateral one separated from eye margin by its own diameter. Flagellum 14segmented, flagellomeres cylindrical; length of flagellomeres about equal to their width. Clypeus setose. Palpi 4-segmented, basal segment small, penultimate and apical ones situated preapically, antepenultimate with round sensory pit on inner surface. Thorax: Scutum with long setae not arranged in distinct rows. Laterotergites with long setae. Scutellum protruding, with 6 pairs of long setae. Antepronotun and proepisternum with long setae. Wing rather wide, membrane without macrotrichia. Costa ends beyond tip of R 5 at one-fifth length between tips of R 5 and M 1 . Sc ends in R at the level of M 3 1 4 and CuA fork base. R, R 1 , and R 5 with setae dorsally. R 1 length 1.8 3 length of r-m. Crossvein r-m about the length of M3, with constriction in the middle. Length of M 1 and M 2 fork 5.3 3 length of M3. Legs: Tibial setae in rows. Tibial spur formula 1:2:2; mid and hind tibial spurs differ in length: inner spur 1.7 3 outer one. Fore leg basitarsomere shorter than tibia. Abdomen setose, with 6 visible segments, 7th and 8th ones small and retractable. Gonocoxites divided by complete suture ventrally. Male: Gonostyli large, triangular, flat, with inner, bilobed appendage. Aedeagus bilobate. Female: Eighth sternites triangular; 10th sternite with 4 dark, thick, wavy, blunt setae. Cerci 2-segmented, rounded, flat; apical segment 0.24 3 basal one, semicircular. M ATERIAL : Holotype AMNH B-0132, male; paratype AMNH B-0133a, female. Myanmar: Katchin, from amber mines near Myitkyina. E TYMOLOGY : The species epithet is a patronym in honor of the late Professor Loı¨c Matile, a world authority on Sciaroidea, who was also very generous and helpful to both authors. Hemolia glabra,new species Figure 72 , Plate 7C D IAGNOSIS : Laterotergites bare; scutum almost bare, dark and shining, Sc meets R well before the level of base of M 3 1 4 and CuA fork. D ESCRIPTION : Body length 5 3.16 mm; wing length 5 2.05 mm. Head: Occiput with short setae. Three ocelli in line, of the same size, distant from each other and from eye margin by two ocellus diameters. Eyes large, setose. Pedicel with round sensory organ. Flagellum 14-segmented, apical segment conical, secondarily divided. Thorax: Scutum almost bare; dark, shiny. Anterior parapsidal suture distinct. Scutellum and laterotergites bare. Mediotergite with a few fine setae anteroventrally. Mid and hind tibiae with inner spurs 1.5 3 longer than outer ones. Wing membrane without macrotrichia. Humeral vein at the level of MA. Costa ends beyond tip of R 5 , at one-third the length between tips of R 5 and M 1 . Sc short, ends at R. R 1 1.3 3 the length of r-m. Length of M3 about equal to length of r-m. Length of M 1 and M 2 fork 4.3 3 the length of M3. Base of M 3 1 4 and CuA fork between tip of Sc and M3 base. Legs: Coxae without setae. Hind tibiae with posterior bristles. Abdomen with scattered setae. Cerci 2-segmented, apical segment conical, basal one cylindrical, 3 3 length of apical one. Gonocoxites 8 long, narrow, slightly curved, with long setae. M ATERIAL : Holotype AMNH B-0112, fe- 52 NO. 3433AMERICAN MUSEUM NOVITATES male. Myanmar: Katchin, from amber mines near Myitkyina. E TYMOLOGY : The species epithet is Latin for ‘‘hairless, smooth’’, in reference to the absence of thoracic setae and setulae. Protragoneura,new genus D IAGNOSIS : Small, dark gnats with antennae shorter than thorax, flagellomeres shorter than wide. Mesonotum densely covered with short trichia, with or without a few setae laterally. Sc short, free. Costa is not produced beyond the tip of R 5 , which is long, curved, and almost reaching the tip of M 1 . Fork of M 1 and M 2 shifted anteriorly, M 1 ends before wing apex. Base of M 3 1 4 and CuA fork at the level of base of M3, in basal one-sixth of wing. M3 length about equal to that of M 1 and M 2 fork. T YPE S PECIES :Protragoneura platycera, n.sp. E TYMOLOGY : The genus name is a combination of the prefix pro- (Greek pro -, beforehand) and Tetragoneura. The name is feminine. C OMMENTS : Closest to Tetragoneura Winnertz, 1846, about 100 extant species of which are known from the Holarctic (17 spp.), Neotropical (ca. 60 spp.), and Australasian (23 spp.) regions. The new genus differs in having C not extended beyond the tip of R 5 , RS2 section (small radial cell) long, and base of M 3 1 4 and CuA fork very basal; M 1 and M 2 fork shifted anteriorly. Setation of mesonotum is also distinctive and separates the new genus from Tetragoneura. Protragoneura platycera,new species Figures 73–75 , Plate 7D D IAGNOSIS : As for genus. D ESCRIPTION : Body length 5 2.49 mm (holotype)/2.39 mm (paratype); wing length 5 1.63 mm (holotype)/1.93 mm (paratype). Head ovate, with row of long erect setae behind eye margin. Scape and pedicel subconical, with numerous apical setae, both black. Flagellum compressed laterally, flagellomeres transverse, widths 1.5–2.5 3 length, densely covered with short trichia. Clypeus setose. Three palpomeres seen; antepenultimate one swollen, obovate, length 2 3 width. Penultimate one attached to antepenultimate palpomere preapically; length 4 3 width, knoblike. Apical palpomere long and narrow. Length ratio 1:1.7:2.5. Thorax: Scutum densely covered with short erect trichia, bearing some setae laterally. Antepronotum and proepisternum with long strong setae. Anepisternum with deep cleft. Katepisternum without distinct excavation on anterior margin. Laterotergites, mediotergite bare. Metepisternum with few short trichia ventrally. Wing membrane with macrotrichia in distal and posterior part. Sc extremely short, free. All longitudinal veins except R base with setae. M3 reduced, can be traced by setae only. M 1 base reduced. Base of M 3 1 4 and CuA fork weakened. Small radial cell length 7 3 width. Halters dark, with few short setae. Legs: Coxae and femora dark, densely setose, tibiae and tarsi yellowish. Hind tibiae with numerous posterior bristles. Tibial spurs 4 3 tibial diameter. Abdomen with 6 visible segments, 7th and 8th very short, retracted. Tergite 9 oval, short, covering the base of gonocoxites. Gonocoxites fused at base, massive, narrowing to apex in distal half. Gonostyli with long ventral lobe bearing a comb of short, knoblike setae; a somewhat shorter, dorsal, bare lobe and internal wide lobe with a row of curved bristles. M ATERIAL : Holotype AMNH Bu-135, male; paratype AMNH Bu-1076, male. Myanmar: Katchin, from amber mines near Myitkyina. E TYMOLOGY : The species epithet is derived from platy (Greek pl ty§ , broad) and cerus a ´ (Greek kra§ , horn), in reference to the come ´ pressed antennae. ANALYSES L YGISTORRHINIDAE AND THE H ETEROTRICHA GROUP Figure 76 Forty-two morphological characters were coded for all known Mesozoic Lygistorrhinidae, Palaeognoriste sp. from Baltic amber, and exemplar species of all living genera of the family. The genera Bolitophila,Chiletricha,Drepanocercus, and Paratinia were used as outgroups. In previous studies Mycetophilidae was suggested to be a sister group to the Lygistorrhinidae, while Bolitophilidae was the sister group to those fami- 2004 53BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER Fig. 76. Phylogeny of Lygistorrhinidae based on cladistic analisys of data presented in table 4 (length 84, CI 0.65, RI 0.81). n unreversed changes; ▫ homoplastic character transitions. The character number is indicated above the branch and the state change is indicated below. Bolitophila,Paratinia,Drepanocercus,Chiletricha and Thereotricha are the outgroup. lies plus Sciaridae (Matile, 1990, 1997; So¨li, 1997). The position of the Heterotricha group is disputed, showing affinities with Diadocidiidae, Sciaridae, and Mycetophilidae, as well as the extinct families Archizelmiridae and Mesosciophilidae (Grimaldi et al., 2003). A data matrix used for cladistic analysis is presented in table 4. Character descriptions are provided in table 5. Analysis of the data matrix using Winnona, (version 2.0; Goloboff, 1999) resulted in four trees of length 83, CI of 0.66, and RI of 0.82. The strict consensus of the trees is shown on figure 76. A previous cladistic analysis of Lygistorrhinidae (Grimaldi and Blagoderov, 2001) included as the only fossil Palaeognoriste, considered a sister group to all living lygistorrhinids. The present analysis yielded the same topology for Recent genera: basal position of ‘‘L. asiatica’’, and sister-group relationships between Seguyola-Loyugesa and Lygistorrhina (with Probolaeus as a subgenus of Lygistorrhina). The Cretaceous genera Archaeognoriste and Lebanognoriste are at the base of the lygistorrhine clade and represent a basal, paraphyletic stem group. While having superficial similarity with the Heterotricha group of genera in venation pattern, they show strong apomorphies of Lygistorrhinidae in the structure of transverse veins, antennae, thorax, and male genitalia (characters 3, 8, 12, 25, 26, 29, 36, 38, 44). The other Cretaceous genera Plesiog- 54 NO. 3433AMERICAN MUSEUM NOVITATES TABLE 4 Data Matrix for Analysis of Lygistorrhinidae Character descriptions presented in table 5. noriste,Protognoriste, and Leptognoriste occupy intermediate positions between primitive and recent lygistorrhinids. Interestingly, although these genera do not form a monophyletic entity, all species have only one vein in the medial fork preserved. The independent loss of a median vein in various Mesozoic lygistorrhinids suggests rapid radiation of the family in the beginning of its history, followed by replacement of Mesozoic groups by recent ones. M YCETOPHILIDAE S . S . Figures 77–79 Sixty-one characters were coded for 38 species representing three main lineages of Cretaceous fungus gnats: Sciophilinae (including 3 recent and 3 fossil species), Leiinae (3 recent and 13 fossil species), and Gnoristinae (9 recent and 7 fossil species). Macrocera and Bolitophila were used as outgroups. A data matrix used for cladistic analysis is presented in table 6. Character descriptions 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 bisection-reconnection), cutting trees up to four points, yielded eight MPTs of length 311, CI of 0.21 and RI of 0.51. The low indices of the trees are due to the very homoplastic dataset, reflected by numerous cases of independent occurrence of structures. The strict consensus of these MPTs (fig. 77a) is an unresolved bush for Gnoristinae, with Leiinae paraphyletic with respect to Sciophilinae. Such an unexpected result can be explained by including in the parsimony analysis two peculiar fossil genera, Lecadonileia and Disparoleia, which share some characters (5, 10, 17, 18) with Sciophilinae. Nonetheless, in all the analyses conducted, Lecadonileia and Disparoleia nested within Leiinae, having synapomorphies with other genera of the subfamily (e.g., characters 12, 29, 47, 50, 51, 52). Due to taxon sampling and the limited number of characters used, the most parsimonious result hypothesized synapomorphies of Sciophilinae as reversals. The alternative, when characters 5, 10, 17, and 18 appear independently in different clades, is less parsimonious by two steps (see below). In- 2004 55BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER TABLE 5 Characters Used in Cladistic Analysis of Lygistorrhinidae All characters are nonadditive. terestingly, analyzing the dataset with WinXpiwe (version 1.3; Goloboff, 1997), a program intended for analysis of homoplastic data (see Goloboff, 1993), yields a topology with Sciophilinae and Leiinae each as monophyletic (fig. 77b). The same result occurs when Lecadonileia and Disparoleia are excluded from the analysis. Since the monophyly of Sciophilinae and Leiinae is strongly supported (So¨li, 1997), the final analysis was conducted in Winnona with Sciophilinae and Leiinae constrained as monophyletic. It resulted in 11 trees of length 313 (only two steps longer than the MPTs found), CI of 0.24 and RI of 0.58. The strict consensus tree (fig. 78) of length 333, CI of 0.22 and RI of 0.54 again indicates Gnoristinae as basal and paraphyletic to Sciophilinae and Leiinae. All the fossil species assigned to extant genera except Synapha longistyla form monophyletic clades with recent species. Pseudomanota is closely allied with Paratrizygia and probably represents a stem group to the Azana group of genera (Matile, 1998). Drepanocercus and Gregikia are at the base of mycetophilids. Tetragoneura and Ectrepesthoneura should be classified in Leiinae, not Gnoristinae, as was suggested by Va¨isa¨nen (1986). On the other hand, his conclusion about Syntemna, traditionally placed in Sciophilinae, is probably right, for on the cladogram it is nested within Gnoristinae. In general, including fossils into an anal- 56 NO. 3433AMERICAN MUSEUM NOVITATES 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. Fittest tree (total fit 5 3056.9, concavity 3). ysis dramatically affects results (fig. 79). To a large extent this effect is a result of exellent preservation in amber, which has preserved characters virtually completely. The topology of the MPTs is particularly affected, probably due to extensive homoplasy. Homoplasious characters in plesiomorphic fossils not only decrease resolution and increase length of the tree but make taxon definitions diffuse. Nonetheless, including fossils in a cladistic analysis has proven to be extremely useful (Gauthier et al., 1988), and this study is an another example of that. First, fossils are the only way to establish existence and position of entirely extinct lineages, like Mesozoic lygistorrhinids or leiines from Burmese amber. Secondly, they allow the identification of stem groups for extant lineages. Basal taxa have a fundamental influence on topology. This is especially true for ancient groups that had a rapid initial diversification, where unusual combinations of characters occur. DISCUSSION P HYLOGENY A systematic review of Bibionomorpha and Sciaroidea was not the purpose of this work and is well beyond the scope of our project. We concur with the composition and evolutionary history of the infraorder as established by Shcherbakov et al. (1995), and also grant that fossils afford unique insight into sciaroid phylogeny. The traditional nomenclature of venation 2004 57BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER TABLE 6 Data Matrix for Analysis of Gnoristinae, Sciophilinae, and Leiinae Character descriptions presented in table 7. in Sciariodea is controversial and somewhat misleading. Fortunately, early fossils provide a strong basis for vein homology (Rasnitsyn, 1980; Shcherbakov et al., 1995), since they show intermediate conditions. The short vein of the radial sector is usually thought to be R 4 (Chandler, 2002). There are four veins in the radial sector of the dipteran ground plan. All representatives of Bibionomorpha s.l. (including Axymyiiformia and Anisopodiformia of Shcherbakov et al., 1995) have no more than three radial veins posterior to R 1 , which can be interpreted as R 2 ,R 3 , and R 4 1 5 or R 2 1 3 ,R 4 , and R 5 according to the authors’ concepts of vein homology. We prefer the second scheme, assuming that a long vein 64 NO. 3433AMERICAN MUSEUM NOVITATES sosciophilidae, Pleciofungivoridae, and Protopleciidae) into one monophyletic group with Cecidomyiidae, probably, as a sister group to all others. The phylogeny of recent families of Sciaroidea was proposed by Matile (1990, 1997) based on imaginal and preimaginal characters, to which extinct groups could be added. Ditomyiidae has been proposed as a sister group to all other living Sciaroidea except Cecidomyiidae; however, no Ditomyiidae are known in the fossil record before the Eocene (Evenhuis, 1994). Eoditomyia primitiva Ansorge from the Lias of Germany possibly represents a stem-group ditomyiid (Ansorge, 1996). The presence of R 4 in Eoditomyiidae may be evidence for an independent origin of Ditomyiidae from Paraxymyiidae, which would make Sciaroidea polyphyletic. In any case, discovery of Eoditomyia allows us to propose a Mesozoic, possible Jurassic, origin of true, crown-group Ditomyiidae, despite the absence of fossils. Matile (1997) provided the only synapomorphy for Diadocidiidae 1 Keroplatidae, which is the absence or closure of larval abdominal spiracles. Chandler (2002) argued that the abdominal spiracles may be secondarily lost in Diadocidia due to the tube-living habit of larvae. The other putative synapomorphy of Diadocidiidae 1 Keroplatidae might be the proximal position of the base of RS with respect to tb. The same condition is observed in Australosymmerus, but this genus is not considered as basal (Munroe, 1974). All other groups of Sciaroidea have (at least in the ground plan) crossvein tb proximal to or at the level of the base of RS. The vertical alignment of veins r-m, M2, and tb in Diadocidiidae and Ditomyiidae, though similar in shape, probably appeared independently. Rigidity of the wing is provided by a long R 1 in Ditomyiidae and by the lengthened base of RS in Diadocidiidae. Thus, crossvein r-m was pulled basad with the base of RS in Diadocidiidae, while in Ditomyiidae crossveins r-m, tb, and m-cu were pulled distad together with the base of RS. Bolitophilidae are represented by only one genus in the Recent fauna. Another genus— Mangas Kovalev—was described from the Lower Cretaceous of Mongolia, and close relatives were reported from the Lower Cretaceous of Transbaikalia (Kovalev, 1986). Chandler (2002) doubted that the position of Mangas was within Bolitophilidae, although he admitted another undescribed but figured specimen could belong to the family. He wrote (2002: 136) ‘‘There is nothing to indicate that it is not allied to Heterotricha’’. One of us (V.B.) had the opportunity to study the type of M. exilis in the collection of PIN. Despite obscure venation, Kovalev’s drawing seems to be accurate and some characters are apparent: long curved base of RS is situated distally, not basally; r-m is vertical, not oblique; section M2 (not tb, as noted on Chandler’s fig. 92) is long and longitudinal (not oblique), although not apparent basally due to preservation; and there is a short section M3 (M 1 1 2 stem) (it is much shorter than M 1 and M 2 fork and almost the same length as visible part of section M2). This combination of characters is not known in the Heterotricha group (where the base of Rs and crossvein r-m are not shifted distally, and section M2 is much shorter than section M3), but it is characteristic of Bolitophilidae. Bolitophilidae and the Heterotricha group are a relicts of Mesozoic sciaroids like Protopleciidae and Pleciofungivoridae, though the relationships between bolitophilids and genera allied to Heterotricha are not perfectly apparent. Mycetophilidae and Lygistorrhinidae are sister groups when only Recent taxa are analyzed. Including fossils in the analysis indicates an independent origin of Bolitophilidae, Mycetophilidae, Lygistorrhinidae, and possibly Sciaridae from within the Mesosciophilidae-Pterogymnus-Heterotricha complex. All these recent families as well as extinct Archizelmiridae share synapomorphies, which are also found in Mesosciophilidae and Heterotricha group. Pterogymnus Freeman is probably related to Mesosciophilidae (Chandler, 2002). One of the Heterotrichalike genera, Sciaropota Chandler, 2002, shows an alliance with an archizelmirid genus, Archimelzira from New Jersey amber (Grimaldi et al., 2003), and probably should be included in Archizelmiridae. Relationships within the Heterotricha group cannot be resolved easily because of the extensively paraphyletic nature of this complex. 2004 65BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER T APHONOMY AND B IOGEOGRAPHY Sciaroidea today are most abundant and diverse in moist to wet temperate forests of the Holarctic and south temperate (e.g., Austral) regions—wherever thick humus and fungal mycelia are abundant. Although sciaroids are diverse in tropical forests, they are diffuse in numbers there. Among taxa known in the Cretaceous, Leiinae and Lygistorrhinidae are better represented in the tropics than Gnoristinae, which are common in temperate regions but scarce in tropical forests. The most striking aspect of the Cretaceous amber faunas is the one in Burmese amber. Of all insect inclusions in other Cretaceous ambers, sciaroids comprise 1% or less in each deposit: For example, Taymyr 12 sciaroids/3450 inclusions; Canada 5/1281; Lebanon 2/600, 2/917, 15/1258 (all of the latter are archizelmirids); and New Jersey 17/1800. Of all insects in Burmese amber, by contrast, 5% of them are sciaroids. This is a proportion very similar to that found in Baltic and Dominican ambers. The proportions of Baltic amber sciaroids have been grossly overestimated based on museum collections (which are highly biased) at 15–20%. Based on crude Baltic amber samples analyzed by PIN at the mine sites in Kaliningrad in 1992, sciaroids (without Sciaridae) comprise 4% of all insect inclusions. The proportion of Mycetophilidae in Dominican amber based on unbiased samples is 3.3% of all insect inclusions (data of D.G.). Among Cretaceous deposits the Burmese sciaroid fauna is most similar to Early Cretaceous orictocenoses of Baisa and Bon Tsagan (Lower Cretaceous of Transbaikalia and Mongolia), which have 1.9% and 1.3%, respectively, of their entire insect faunas comprised of sciaroids. The Taimyr, New Jersey, and Canadian amber sciaroid faunas seem the most similar to each other. The Burmese amber sciaroid fauna probably reflects a particularly wet paleoenvironment, although not necessarily warmer than the other Cretaceous deposits. Based on a large, new assemblage of more than 3000 insect inclusions in Burmese amber, Grimaldi et al. (2002) concluded that Burmese amber reflected the most tropical environment among all Cretaceous amber deposits. Sciaroids present conflicting evidence. On the one hand, sciaroids in Burmese amber were abundant, as in temperate forests. However, no species is particularly abundant, and Leiinae and Lygistorrhinidae are quite diverse, as would be found in tropical forests today. ACKNOWLEDGMENTS This work would not have been possible without the generous support of Robert G. Goelet, Chairman Emeritus of the AMNH Board of Trustees, to the senoior author. We are sincerely grateful to his patronage of collection-based research and for his funding Burmese amber acquisition. The work was greatly facilitated by assistance and advice received from Jeffrey Cumming, Michael S. Engel, Martin Ramirez, Alexandr P. Rasnitsyn, Andrew J. Ross, Valerie Schawaroch, Toby Shuh, Alexander I. Zaitzev, F. Christian Thompson, and especially from our late colleagues Loı¨c Matile and Vladimir V. Zherikhin. We wish to thank also Peter Chandler and Andre Nel for their helpful comments of the manuscript. REFERENCES Alonso, J., A. Arillo, E. Barron, J. C. Corral, J.L. Grimalt, J.F. Lopez, R. Lopez, X. Martı´nez-Delclo´s, V. Ortuno, E. 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Alonso, J.C. Corral, and R.V. Lopez (conveners). Proceedings of the World Congress on amber inclusions. Estudios del Museo de Ciencias Naturales de Alava 14, Numero especial 2: 119–131. Zherikhin, V.V., and A.J. Ross. 2000. A review of the history, geology and age of Burmese amber (burmite). Bulletin of the Natural History Museum Geology Series 56(1): 3–10. Zherikhin, V.V., and I.D. Sukacheva. 1973. On the Cretaceous insect-bearing ‘‘ambers’’ (retinites) from North Siberia. In Voprosy paleontologii nasekomykh [Problems of Insect Paleontology]. Doklady na XXIV Ezhegodnom Chtenii Pamyati N.A. Kholodkovskogo 1–2 Apr. 1971: 3–48. Leningrad: Nauka. [in Russian] 70 NO. 3433AMERICAN MUSEUM NOVITATES Plate 1. Families Diadocidiidae and Lygistorrhinidae. A. Docidiadia burmitica Blagoderov and Grimaldi, 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 primitiva Blagoderov and Grimaldi, holotype AMNH Bu-1539. E. Lebanognoriste prima Blagoderov and Grimaldi, holotype AMNH JG268/1. F. Plesiognoriste carpenteri Blagoderov and Grimaldi, holotype MCZC 6927. 2004 71BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER Plate 2. Family Lygistorrhinidae. A. Plesiognoriste zherikhini Blagoderov and Grimaldi, holotype PIN 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. 72 NO. 3433AMERICAN MUSEUM NOVITATES Plate 3. Family Mycetophilidae: Manotinae, Sciophilini, Gnoristini. A. Alavamanota burmitina Blagoderov 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 bulunensis Blagoderov and Grimaldi, holotype PIN 3963/4. F. Synapha longistyla Blagoderov and Grimaldi, holotype MCZC 6944. 2004 73BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER Plate 4. Family Mycetophilidae: Gnoristini and Leiini. A. Dziedzickia nashi Blagoderov and Grimaldi, holotype AMNH NJ-117a. B. Saigusaia pikei Blagoderov and Grimaldi, holotype TMPD P79.15.7.21. C. Syntemna fissurata Blagoderov and Grimaldi, holotype TMPD P83.15.3.8. D. Gregikia pallida Blagoderov and Grimaldi, holotype AMNH NJ 117j. E. Gaalomyia carolinae Blagoderov and Grimaldi, holotype AMNH Bu-390. F. Nedocosia exsanguis Blagoderov and Grimaldi, holotype PIN 3130/193.