PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3433, 76 pp., 81 ®gures, 7 plates, 7 tables February 27, 2004

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

VLADIMIR BLAGODEROV1 AND DAVID GRIMALDI2

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 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

Copyright ᭧ American Museum of Natural History 2004 ISSN 0003-0082 2 AMERICAN MUSEUM NOVITATES NO. 3433

Subfamily Winnertz, 1863 ...... 22 Tribe Sciophilini Winnertz, 1863 ...... 22 Rondani, 1856 ...... 22 Mik, 1886 ...... 24 Pseudomanota, new genus ...... 25 Tribe Gnoristini Edwards, 1925 ...... 26 Grzegorgzek, 1885 ...... 26 Meigen, 1818 ...... 28 Johannsen, 1909 ...... 28 Vockeroth, 1980 ...... 31 Winnertz, 1863 ...... 32 Gregikia, new genus ...... 32 Gaalomyia, new genus ...... 34 Tribe Leiini Edwards, 1925 ...... 36 Nedocosia, new genus ...... 36 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 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 3

ABSTRACT The Recent world fauna of Sciaroidea, or fungus gnats, comprises approximately 4000 described species in eight families: Bolitophilidae, Cecidomyiidae, Diadocidiidae, Ditomyi- idae, Keroplatidae, Lygistorrhinidae, Mycetophilidae, and Sciaridae. Larvae live primarily in decaying vegetation, feeding on fungal mycelia, and they can be among the most abundant of temperate forests. Stem-group families appeared in the , with large Tertiary deposits being composed almost entirely of living genera, so the Cretaceous is essential for understanding the origins and diversi®cation of Recent families. Sixty-six specimens were studied from six major deposits of Cretaceous amber, spanning 40 million years from the Early to : 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); Thereo- tricha sibirica, (?)T. agapa (n.gen., n.spp.) (Sciaroidea incertae sedis); Archaeognoriste prim- itiva, Lebanognoriste prima, Plesiognoriste carpenteri, P. zherikhini, Protognoriste amplicau- da, P. goeleti, P. nascifoa, Leptognoriste davisi, L. microstoma (n.gen., n.spp.) (Lygistorrhin- idae). In Mycetophilidae sensu stricto: Alavamanota burmitina, n.sp. (Manotinae), Neuratelia maimecha, n.sp., Allocotocera burmitica, n.sp., Pseudomanota perplexa, n.gen., n.sp. (Scio- philinae Sciophilini); Apolephthisa bulunensis, n.sp., Synapha longistyla, n.sp., Dziedzickia nashi, n.sp., Saigusaia pikei, n.sp., Syntemna ®ssurata, n.sp., Gregikia pallida, n.gen., n.sp., Gaalomyia carolinae, n.gen., n.sp. (Sciophilinae Gnoristini); Nedocosia exsanguis, N. sibirica, N. canadensis, N. novacaesarea, n.gen., n.spp.; Ectrepesthoneura succinimontana, E. swol- enskyi, n.spp.; Izleiina miri®ca, I. spinitibialis, n.gen., n.spp.; Zeliina orientalis, Z. occidentalis, n.gen., n.spp.; Temaleia birmitica, n.gen., n.sp., Lecadonileia parvistyla, n.gen., n.sp.; Dis- paroleia cristata, n.gen., n.sp.; Hemolia matilei, H. glabra, n.gen., n.spp.; and 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 devoted to one particularly diverse group of ¯ies well represented in Cretaceous amber, The preservation of fossils in amber is re- but which have been virtually neglected: the nowned for consistently ®ner preservation fungus gnats, or Sciaroidea. than virtually all other modes of fossilization. The Sciaroidea is comprised of 14 fami- Although the geological occurrence of amber lies, 6 of them exclusively Mesozoic: Ante- begins in the , the oldest faunas fungivoridae (Middle Jurassic to Early Cre- in amber are no older than Early Cretaceous, approximately 120±130 million years ago taceous; Kovalev, 1990), Archizelmiridae (Ma). Nonetheless, insects in amber from the (Late Jurassic to Late Cretaceous; Grimaldi Cretaceous period (145±65 Ma) afford un- et al., 2003), Eoditomyiidae (Early Jurassic paralleled insight into Mesozoic evolution. to Early Cretaceous; Ansorge, 1996), Meso- The many more preserved characters allow sciophilidae (Middle Jurassic to Early Cre- more accurate phylogenetic inference than is taceous; Kalugina and Kovalev, 1985, Bla- possible with compression or most other goderov, 1993), Pleciofungivoridae (Early kinds of fossils, particularly for intricate or- Jurassic to Early Cretaceous; Kovalev, ganisms like insects. Detailed preservation 1987b), and Protopleciidae (Early Jurassic to facilitates not only reconstruction of phylog- Early Cretaceous; Blagoderov, 1996, Koval- eny, but of paleoenvironments as well. In this ev, 1990). No extinct families occur in the regard, the study of extinct Diptera is essen- Cenozoic. The extant families of the super- tial, as Diptera usually comprise the largest family are Bolitophilidae, Cecidomyiidae, proportions by individuals and species of all Diadocidiidae, , Keroplatidae, types of preserved in various am- Lygistorrhinidae, Mycetophilidae (s.s.), and ber deposits around the world. This paper is Sciaridae, all of which but Ditomyiidae also 4 AMERICAN MUSEUM NOVITATES NO. 3433 occur in the Cretaceous. Phylogenies and modern. Prior to the work of the senior author comprehensive revisions of some of these on extensive insect LagersaÈtte from Asia (Bla- families were provided by Grimaldi and Bla- goderov, 1995, 1997, 1998a, 1998b, 2000), goderov (2001) (Lygistorrhinidae), Matile there were only seven species of sciaroids (1981a, 1990, 1997) (Keroplatidae, Sciaro- known from the Cretaceous, which were idea), Munroe (1974) (Ditomyiidae), SoÈli placed in the Mesosciophilidae or Myceto- (1997) (Mycetophilidae s.s), and VaÈisaÈnen philidae (Kovalev, 1986, 1990; Hong, 1992; (1984) (Mycomyiini). There are approxi- Ren et al., 1995; Jell and Duncan, 1986; mately 4100 described species of Sciaroidea Westwood, 1854). Sciaroids have been known exclusive of the poorly studied but diverse for years to occur in the major deposits of families Cecidomyiidae and Sciaridae (Be- Cretaceous ambers from Taimyr, Siberia chev, 2000). At least 10,000 and perhaps as (Zherikhin and Sukacheva, 1973; Zherikhin many as 20,000±30,000 living species of 1978), Burma (Zherikhin and Ross, 2000), Sciaroidea exist. Such diversity re¯ects the New Jersey (Grimaldi et al., 1989), and west- signi®cance of the group in terrestrial eco- ern Canada (McAlpine and Martin, 1969; systems, particularly forests, the very habitat Pike, 1994). Only three species, however, that produces amber. were described: Burmacrocera petiolata Larvae of Sciaroidea are abundant in the Cockerell, Sciara burmitina Cockerell (in sporophores of basidiomycete fungiÐin de- Burmese amber, thought at the time to be Ce- caying wood, leaf litter, and humus, where nozoic), and Schlueterimyia cenomanica Ma- they feed on fungal mycelia. Where humus tile (in Cenomanian French amber). Since and decaying wood are particularly thick in then, major new Cretaceous outcrops of am- mature forests, sciaroids are among the most ber have been discovered from northern Spain abundant and diverse insects, with their lar- (Alonso et al., 2000), Lebanon (Azar, 2000), vae inhabiting 80±90% of the macrofungi in New Jersey (Grimaldi, 2000), and Myanmar natural habitats (Yakovlev, 1988). Larval (Grimaldi et al., 2002), which have produced habits are not restricted, though, to mycoph- signi®cant numbers of new specimens and agy (Matile, 1997). The larvae of some gen- taxa of sciaroids. Based on study of the older era and species of Keroplatidae are actually collections and these newer ones, we signi®- predators of insects they snare in their slime cantly extend the known Cretaceous diversity, trails, the most famous example of which is based on 27 genera (19 of them new) and 39 the New Zealand ``glow-worm'', (Arachno- species (all of them new). campa Edwards). Larvae of Arachnocampa are luminescent and suspend themselves in MATERIALS AND METHODS mucous strands from the ceilings of caves. Sixty-six specimens from the Early and Small midges attracted to the light become Late Cretaceous of six countries, and some entangled in the strands, which the larvae 10 localities, were examined. We did not then devour. There is even a keroplatid, study inclusions of Diadocidiidae in Burmese Planarivora Hickman, whose larva is para- amber from the Natural History Museum, sitic on land planarians in Tasmania (Hick- London, since these were on loan. Adults of man, 1964; Matile, 1981b). most extant genera of Mycetophilidae (s.s), The fossil record of the Sciaroidea begins and all species of Lygistorrhinidae were ex- in the Early Jurassic (Ansorge, 1996; Kalu- amined. Material is housed in the following gina and Kovalev, 1985; Kovalev, 1987a), institutions: and by the Early Cretaceous modern families replaced older, stem-group families (Kovalev, AMNH American Museum of Natural History 1987a). The Cenozoic Sciaroidea are partic- (Division of Invertebrate Zoology), ularly well studied, especially those in Baltic New York, USA MCNA Museo de Ciencias Naturales de AÂ lava/ amber (approximately 270 species of Myce- Arabako Natur Zientzien Museoa, Vi- tophilidae, as reviewed in the catalogue of toria-Gasteiz, Spain fossil Diptera [Evenhuis, 1994]). The Baltic MCZ Museum of Comparative Zoology (De- amber sciaroid fauna, like most of the other partment of Entomology), Harvard Cenozoic faunas of these ¯ies, is essentially Cambridge, Massachusetts, USA 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 5

PIN Paleontological Institute of the Russian National Collection (CNC) of Insects and Academy of Sciences (Arthropoda Spiders, Ottawa. The Grassy Lake amber is Laboratory), Moscow, Russia from the Judith River Group of the Foremost TMPD Tyrrell Museum of Palaeontology, Al- Formation, dated as Campanian (Pike, 1994; berta, Canada Borkent, 1995), although Grimaldi and Cum- Specimens were derived from the follow- ming (citing David Eberth, personal com- ing localities and formations: mun.) indicated the amber to be Santonian. Alava, Spain: The amber deposit is lo- Amber from Cedar Lake, Manitoba was col- cated on the northern slope of Sierra de Can- Â lected by F.M. Carpenter in 1938 among tabria (Alava), about 30 km southeast of the beach debris (in the MCZ). Although rede- city of Vitoria-Gasteiz, near the village of posited and somewhat distant from the PenÄacerrada. A comprehensive description of Grassy Lake deposit, the two deposits are the amber deposit was given by Alonso et al. thought to be contemporaneous (Borkent, (2000). It was assigned to the Nograro For- 1995). An age of 85±80 myo is a reasonable mation, of Aptian±middle Albian age (120± estimate for Canadian amber. 110 Ma). All of the material is housed in Lebanon: Material studied here is from MCMA. the Acra Collection (collected by Fadi and Burma (Myanmar): Historically and Aftim Acra near Jezzine, 30 km east of Sai- presently all amber from Burma derives from da, Lebanon) and the Estephan Collection the northern state of Kachin. Zherikhin and (collected by Antoun Estephan, near BcharreÂ, Ross (2000) reviewed historical records of northern Lebanon). Both of these collections Burmese amber mining and locations. For are housed in the AMNH. Most recently, ex- many years Burmese amber was considered tensive collections and stratigraphic studies Cenozoic in age; however recently it has of Lebanese amber have been made by Azar been found to be Cretaceous after the dis- (2000). Lebanese amber was originally re- covery of insects in it that are exclusively ported to be Neocomian in age, pertaining to Cretaceous, like Serphitidae and Stigma- basal periods of the Early Cretaceous (Schlee phronidae (Hymenoptera). A collection of and Dietrich, 1970), 145±129 Ma. A younger approximately 1200 inclusions re- age of Aptian-Albian (ca. 120±110 Ma) has sides at the NHM, London, rich in types de- also been mentioned (Whalley, 1976; Zheri- scribed by T.D.A. Cockerell and others. Re- khin, 1978). Azar's study, which is the most cently, a very large, diverse collection of comprehensive, indicates a great range of Burmese amber has been assembled at the ages of Lebanese amber, from uppermost Ju- AMNH from material recently excavated in rassic (152 Ma) to Albian (112 Ma), but most Kachin, near villages close to the town of of the deposits yielding insect inclusions are Myitkyina. This material was acquired by Barremian-Aptian (approximately 125 Ma). Leeward Capital (Calgary, Canada), who Sciaroids in Lebanese amber are the oldest provided it to the AMNH. Study of some known amber fossils of the superfamily. 3500 organisms in this collection revealed New Jersey: Rich amber deposits from more than 20 Mesozoic insect taxa, con®rm- the central county of Middlesex have been ing the Cretaceous age and even indicating known for decades, summarized by Grimaldi an age of approximately Cenomanian (98±92 et al. (1989). Recently, an extraordinary out- Ma) (Grimaldi et al., 2002). Burmese amber crop was discovered in the town of Sayre- is by far the most proli®c and diverse Cre- ville, with a smaller one being discowered in taceous amber for Sciaroidea. the adjacent town of East Brunswick (Gri- Canada: Material from Grassy Lake, Al- maldi, 2000). This material has produced the berta, derives from an abandoned coal mine oldest de®nitive ants, tardigrades, mush- 8 km south and 1.6 km east of the village of rooms, parasitiform mites, and ¯owers Grassy Lake (Pike, 1994). This was collected (among many others) preserved in amber. by T. Pike and housed at the RTMP. This Chemical analyses of the amber and stratig- locality was previously cited as ``near Med- raphy of the outcrops have been made. The icine Hat'' by McAlpine and Martin (1969), amber occurs in the South Amboy Fire Clay who deposited their material at the Canadian of the Magothy Formation, Turonian in age 6 AMERICAN MUSEUM NOVITATES NO. 3433

(Grimaldi, 2000). All of the New Jersey am- optic ¯ash wands (ML-1000 from Microp- ber is deposited in the AMNH. tics). Siberia (Taimyr): Reviews of the arthro- Morphological terminology used follows pod fauna in the Siberian amber were pro- McAlpine (1981) and SoÈli (1997) with mod- vided by Zherikhin and Sukacheva (1973) i®cation after Kovalev (Kalugina and Koval- and Zherikhin (1978). Sciaroidea were found ev, 1985). To compare relative position of at four localities, ages of which vary. Mate- veins in sciaroid wings, Kovalev de®ned bas- rial was collected by a PIN expedition in al (from the base of Rs to the base of r-m), 1973 to Nizhnaya Agapa, which is on the middle (from the base of r-m to the base of north shore of the Agapa River, 40 km down- R4), and apical (equivalent to R5) sections in stream from Lake Ladonnakh, Yst'-Enisey radius stem as RS1, RS2, and RS3 sections, depression. This material is in the Dolgan respectively. Also, the basal (from the base Formation, dated palynologically as Albian± of the wing to tb), middle (from tb to r-m),

Cenomanian (Saks and Ronkina, 1957). Am- and apical (from r-m to the base of the M1 ber from Yantardakh (``amber mountain'') and M2 fork, equivalent to the stem of M1ϩ2) occuring 3±5 km upstream from the mouth sections of media stem were de®ned as M1, of the Maimecha River, Khatanga depres- M2, and M3 (see also ®gs. 80, 81). The term sion, in the Kheta Formation (dated as Con- ``stem of M'' refers to M1, M2, and M3 sec- iacian±Santonian [Saks et al., 1959], ca. 87 tions together. The term ``base (or stem) of myo) was collected in 1970 and 1971. A M1 and M2 fork'' refers to the point of fur- 1976 PIN expedition collected a contempo- cation of the veins. The terms ``stem of M1 raneous material from Bulun (middle course and M2 fork'' or ``stem of M1ϩ2'' refer to M3 of Bulun River [right tributary of Kheta Riv- section. The terms ``R5'' and ``stem of M1ϩ2'' er], 18 km S of post Novaya, Taimyr). Am- are used to refer to the veins themselves, and ber from Baikura-Neru (Taimyr Lake, Baik- ``RS3'' and ``M3'' are used in reference to ura-Neru Bay) contains the only incomplete the relative length of sections. All the veins specimen of Sciaroidea. Baikura-Neru, age and section names are summarized in table of which is unclear, has been assigned to the 1, which shows comparison of the vein no- Ogneva Formation (Saks et al., 1959; Ap- menclature used by various authors. In some tian-Albian, 120±110 Ma), but arthropod in- cases, for the sake of stability, we preferred clusions indicate a Late Cretaceous age to use traditional vein names in descriptions (Dlussky, 1987; Zherikhin and Eskov, 1999). of species rather than use their names based All Siberian amber is in PIN. on presumed, primary homology. These Specimens originally stored in small boxes veins are denoted in boldface type in the ta- were prepared according to the method de- ble. In the discussion of phylogeny, however, scribed in Nascimbene and Silverstein it seemed advisable to use names of veins (2000). This involved embedding the speci- based on homology for establishing transfor- men in a stable epoxy (Buehler) under vac- mation series. We used the term ``not visi- uum. The vacuum extracts air in ®ne cracks, ble'' when a structure could not be observed which the epoxy then permeates, thus im- clearly. proving visibility and fragility of the piece. Pieces were then trimmed to thin pieces, with SYSTEMATIC PALEONTOLOGY surfaces often to within fractions of a milli- SUPERFAMILY SCIAROIDEA BILLBERG, 1820 meter of the inclusion, in order to optimize observation of details. Prepared in this way, FAMILY DIADOCIDIIDAE EDWARDS, 1925 pieces can be mounted on microscope slides Docidiadia, new genus and the inclusions observed under 100± 400ϫ magni®cation with a compound micro- DIAGNOSIS: Head round. Flagellum 14-seg- scope. Specimens were measured with a dig- mented. First ¯agellomere length slightly ital stage micrometer mounted under a Zeiss more than width. Fore tibial comb absent. SV8 stereoscope, and photographed using an Wing membrane without macrotrichia. C

In®nity K-2 lens attached to a Nikon D-1 ends beyond tip of R5; Sc long, ends free; camera and illuminated with focusable ®ber- RS base at the middle of R1; crossveins r-m, 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 7

TABLE 1 Nomenclature and Homology of Sciaroid Wing Veins According to Authors See also ®gures 80 and 81.

tb, and m-cu in one line; M3 section and base species is known from Baltic amber (Even- of M fork absent; CuA strongly curved back huis, 1994). at the apex. A short. Male 9th tergite without marginal bristles, with one large, acute, tri- Docidiadia burmitica, new species angular medial appendage and two small lat- Figures 1±3, Plate 1A eral ones; gonostyli do not bifurcate at apex. TYPE SPECIES: Docidiadia burmitica, n.sp. DIAGNOSIS: As for genus. ETYMOLOGY: The name is a feminine an- DESCRIPTION: Body length ϭ 1.88 mm (ho- agram of Diadocidia. lotype)/1.61 (paratype); wing length ϭ 1.61/ COMMENTS: The genus is close to Diado- 1.55 mm. Head. Eyes bare, facets round. cidia Ruthe, but differs in having the ®rst Clypeus setose. Flagellum 14-segmented; ¯agellomere short; wing membrane without ®rst ¯agellomere cylindrical, width slightly macrotrichia, Sc ending free, base of RS less than length; apical ¯agellomere twice the rather distal, M3 section and base of M fork width. Apical ¯agellomere of male second- reduced; CuA curved at apex rather than arily segmented in two parts. Scape and ped- with two straight sections; and male tergite icel turbinate. Only 3 segments of palpi seen, IX narrow, triangular, and with two lateral palpomeres cylindrical, subequal in length, appendages. Diadocidia consists of two sub- basal wider than the rest. Thorax. Scutum genera and includes 10 Holarctic species setose, dome-shaped, with long protruding (Chandler, 1994; LasÏtovka and Matile, 1972; setae. Metepisternum bare, height equal to Polevoi, 1996; Wu, 1995; Zaitzev, 1994) and width, shallow incision anteriorly. Wing a Neotropical one (Edwards, 1940; Papavero, membrane without macrotrichia. Costa ends

1977a), as well as undescribed Australian beyond tip of R5, midway between tips of R5 species (Tonnoir, 1929; Colless, 1963). One and M1. Sc ends free, slightly beyond base 8 AMERICAN MUSEUM NOVITATES NO. 3433

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. R1 setulose, R5 with sparse setae, al- ocoxites short, with length about the width. most straight. R1 short, about 0.6ϫ wing Gonostyli massive, length 2.5ϫ the length of length. M3 absent. M1 and M2 weakened, gonocoxites, hairy, without apical teeth or their bases absent. M3ϩ4 weakened. Abdo- spines. men. Female cerci wide, subtriangular with MATERIAL: Holotype AMNH Bu-033, acute ventral angle. Male tergite IX narrow, male; paratype B-002, female. Myanmar: triangular with two lateral appendages. Gon- Katchin, from amber mines near Myitkyina. 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 9

ETYMOLOGY: The species epithet is a ref- section completely reduced; M2 section erence to Burma, the former name of the fused with tb in one vein meeting the base country where the amber originates. of M3ϩ4, and that oblique vein is shifted dis- tad. The same structure of the basal veins is SCIAROIDEA INCERTAE SEDIS observed in advanced representatives of the Thereotricha, new genus Mesozoic family Mesosciophilidae, which are thought to represent a sister group to My- DIAGNOSIS: Eyes forming incomplete eye cetophilidae (Kalugina and Kovalev, 1985; bridge, facets large, round. Ocelli three. Fla- Blagoderov, 1993). Similar conditions occur gellomeres barrel-shaped, length no more in the peculiar Mesozoic family Archizel- than 1.5ϫ the width. Antepronotum and miridae (Grimaldi et al., 2003), but Archi- proepisternum subequal, setose. Proepimeron zelmiridae have crossvein r-m aligned with touches episternum at the episternal suture. M2 ϩ tb and the basal portion of M3ϩ4, form- Anepisternum smaller than katepisternum. ing one horizontal vein and the base of RS Anepisternal cleft distinct, narrow. Midpleur- is shifted distad. Diadocidiidae s.str. (Dia- al pit present. Metepisternum setose. Ante- docidia and Docidiadia n.gen.) also have rior parapsidal suture distinct. Insertion of these veins aligned (r-m through the base of abdomen wide. Wing membrane with or M3ϩ4), but they form a vertical vein. In My- without macrotrichia. Sc short, ends free. Rs cetophilidae the combined vein M2 ϩ tb lost base, r-m, base of M3ϩ4, and CuA fork very contact with the base of M3ϩ4 and meets the basal. Section M2 connects r-m and base of base of CuA or MA (arculus) (see Shcher-

M3ϩ4 and CuA fork. M3ϩ4 reduced. bakov et al., 1995). Some Mycetophilidae TYPE SPECIES: Thereotricha sibirica, n.sp. (Drepanocercus, Ectrepesthoneura, Creta- ETYMOLOGY: The genus name is a femi- ceous Paradzickia, Drepanorzeckia, Ekhiri- nine anagram of Heterotricha. The name is tus, Zazicia) have the fork of M3ϩ4 and CuA feminine. sessile or short-stalked, but the base of the COMMENTS: The new genus is close to the fork is situated more basally that in Mesozoic Heterotricha group of genera, which have Sciaroidea and the Heterotricha group. been included in the Sciaridae or Diadoci- Moreover, at least in Ectrepesthoneura the diidae. Recently, Chandler (2002) reviewed sessile fork of M3ϩ4 and CuA is secondary known taxa of the group and described seven (see Analyses below and ®g. 78). Obviously, more genera from all zoogeographic regions reduction of M1 and fusion of M2 with tb except Nearctic. These taxa seem to repre- might have originated several times in the sent a stem group of Recent families of Sci- history of Sciaroidea. Although monophyly aroidea, but monophyly of the group is not of the group combining Recent Heterotricha- apparent. The new genus differs from all taxa like taxa and Cretaceous Thereotricha is not of the Heterotricha group in having eyes proven, position of these taxa in sciaroid with large facets, that form an eye bridge, phylogeny should be at the base of lineages short antennae, the scape and pedicel not dif- leading to Mesosoic Mesosciophilidae and fering from ¯agellomeres in length, palpi Archizelmiridae and Recent Sciaridae on the very short, anepisternite and katepisternite one hand and higher sciaroids such as My- subequal, the base of RS in basal position, cetophilidae and Lygistorrhinidae on the oth- and r-m and the section of M2 subequal, er. where eyes and palpi demonstrate an apo- morphic condition. The new genus resembles Thereotricha sibirica, new species Sciaropota Chandler in the porrect antenna Figure 4, Plate 1B with short ¯agellomeres, an absence of a clearly differentiated series of scutellar bris- DIAGNOSIS: Wing membrane without mac- tles, large katepisternum, mesepimepon rotrichia, RS base very basal, oblique, ϳ2ϫ broader below, but it differs by the short Sc, the length of r-m; M3 section long; base of long stem of M1ϩ2, and reduced M3ϩ4. M1 and M2 fork at level of tip of R1; veins Distinct synapomorphies separate the R1,R5,M1,M2, and CuA with long setae. group from other Mesozoic Sciaroidea: M1 DESCRIPTION: Body length ϭ 1.83 mm (ho- 10 AMERICAN MUSEUM NOVITATES NO. 3433 lotype)/1.84±1.85 mm (paratypes); wing paratypes, PIN 3130/182, 3130/184, 3130/ length ϭ 1.46 mm (holotype)/1.25 mm (para- 185, males; all specimens in the same piece type PIN 3130/185). Head. Three ocelli pre- of amber. Russia: Taimyr Peninsula, Yantan- sent, equal in size, distance of lateral ocellus darkh, coll. 1970. from median ocellus and eye margin 1.5ϫ its ETYMOLOGY: The species epithet is in ref- diameter. Occiput with numerous small setae. erence to Siberia, the region where the orig- Eyes large, bare, forming incomplete dorsal inal locality is situated. bridge (separated by twice the facet diame- ter). Scape and pedicel short, rounded. Fla- ?Thereotricha agapa, new species gellum 14-segmented, ¯agellomeres barrel- Figures 5, 6, Plate 1C like, width 1.3ϫ the length, covered by short trichia. Palpi 4-segmented. Basal segment DIAGNOSIS: Three ocelli, touching. Wing small, bacilliform, others short, rounded; an- membrane with scattered macrotrichia. RS tepenultimate segment with round sensory base transverse, shorter than r-m. M2 proba- pit dorsomedially, slightly longer than oth- bly absent. ers.Thorax. Antepronotum with one strong DESCRIPTION: Body length ϭ 1.56 mm seta, proepisternum with six. Proepimeron (rest, estimated length 1.87 mm); wing touches episternum at episternal suture. Me- length 1.48 mm. Partly preserved specimen. sonotum uniformly setose. Midpleural pit Head: Occiput and vertex with straight and present. Katepistrnum large, expanded cau- curved setae. Three ocelli present, each on dally. Ventral part of mesepimeron not very its own mound, the latter fused by contacting narrow, parallel-sided. Laterotergites bare, small. Metepisternum setose, with narrow edges. Eyes bare, with incomplete bridge, anteriodorsal cleft. Wing membrane without deeply emarginate at antennal base. Frontal furrow and keels well developed. Scape and macrotrichia. Costa ends beyond tip of R5, one-third the distance between tips of R and pedicel as long as ¯agellomeres. Flagellum 5 14-segmented, ¯agellomeres barrel-shaped, M1. Sc very short, ends free just beyond RS base. RS base proximad, oblique, 1.5ϫ r-m length about 1.5ϫ the width, with few strong length. R ,R,M,M, and CuA with long setae and numerous ®ne trichia. Clypeus se- 1 5 1 2 tose. Palpi 4-segmented, short. Antepenulti- setae. R1 about 0.7ϫ wing length. RS base oblique. Crossvein r-m fused with tb and m- mate palpomere with dosomedial round sen- cu in one horizontal vein. M3 section 1.5ϫ sory pit. Thorax: Proepisternum with 3 long and and 11 shorter setae. Proepimeron touch- as long as fork of M1 and M2.M3ϩ4 reduced, seen as a fold at base only. CuA ends before es episternum at the episternal suture, kate- middle of the wing. Legs. Fore coxae with pisternum with shallow excavation below the numerous setae on anterior and distal surfac- point. Anepisternum less than katepisternum, es, mid coxae with distal rows and several the latter with several trichia in lower part. setae on apical part, hind coxae with caudo- Laterotergite bare, divided with katepister- distal row of setae. Fore tibia with anteroap- num by anepimeron. Mediotergite bare. Me- ical pit having two combs of setae. Abdo- tepisternum setose. Scutum with scattered se- men setose. Eighth segment slightly shorter tae. Scutellum with 6 long setae. Wing mem- than preceding ones. Ninth tergite ovoid, brane with microtrichia and macrotrichia with apical comb of short blunt setae. Gon- mostly in apical and hind area. Costa, R1, and ocoxites stout, triangular in lateral view. R5 with long setae. Sc very short, ends free. Gonostyli small, ¯at, rectangular, with sev- R1 and R5 almost straight. RS base short, eral apical setae. ϳ0.5ϫ r-m length, transverse; r-m fused with MATERIAL: Holotype PIN 3130/183, male; tb and m-cu, slightly oblique. Crossvein m-

→ Figs. 4±6. Thereotricha, n.gen. 4. Thereotricha sibirica, n.sp., drawing of amber piece with four embedded specimens, including holotype PIN 3130/183. 5.? T. agapa, n.sp., holotype PIN 3426/256. 6.? T. agapa, head of the holotype. 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 11 12 AMERICAN MUSEUM NOVITATES NO. 3433

cu twice the length of r-m. M2 probably ab- lotype)/1.47±1.61 mm (paratypes); wing sent. M3ϩ4 absent. M1 straight, weak. length ϭ 0.98/0.95±1.10 mm. Head: Poste- MATERIAL: Holotype PIN 3426/256, sex rior surface ¯at, with scattered setulae. Frons unknown. Russia, Taimyr Peninsula, Nizh- membranous. Eyes large, hairy. Ocelli not nyaya Agapa, coll. 1973. seen. Scape and pedicel slightly wider than ETYMOLOGY: The species is an epithet in ¯agellomeres, subconical. Flagellum 14-seg- reference to the locality, the Nizhnyaya Aga- mented, ¯agellomeres cylindrical, width pa River. about equal to length. Palpi 3-segmented, COMMENTS: Structure of ocelli and a trans- segments cylindrical. Penultimate segment verse RS base are apparent characters of ge- about 0.75ϫ the length of apical one and 2ϫ neric level, distinguishing the species from the length of antepenultimate. Thorax: Scu- T. sibirica, but incomplete preservation of tum irregularly covered with short setae and the specimen from Nizhnyaya Agapa does bearing long lateral and acrostichal setae. not allow description of a new genus. Both Scutellum with two pairs of long setae and species share such synapomorphies as an in- several short ones. Antepronotum and pre- complete eye bridge, proepimeron apex at episternum with 6±8 long setae. Anepisternal the episternal suture, setose metepisternum, suture declines posteriorly. Anepisternum and M3ϩ4 reduced. Until more specimens of with small cleft. Mediotergite setose, latero- this peculiar group will be found and phy- tergites bare. Dorsal edge of mesepimeron logenetic analysis is done we prefer to keep protruding toward laterotergite. Metepister- the species in this genus. num with deep cleft, height more than width. Wing membrane without macrotrichia. C not FAMILY LYGISTORRHINIDAE EDWARDS, 1925 reaching wing tip, ends one-fourth distance between tips of R and M . All veins bear DIAGNOSIS: Eyes round; pedicel bears bris- 5 1 setae. Sc very short, free. R about 0.6ϫ the tles apically; mouthparts reduced and form 1 length of wing. RS base distinct, at the level proboscis in some genera; Anepimeron fused of Sc tip. Crossvein r-m fused with tb, to katepisternum; Sc very short; R short; 1 oblique. Stem of M and base of M and M stem of M and M fork base absent; r-m 1 2 1ϩ2 fork absent. M base very weak. M and aligned with tb; M2 tb subhorizontal; hind 3ϩ4 ϩ CuA veins weak. Legs: Fore coxae densely coxae short, hind tibiae and tarsi swollen; setose, mid and hind coxae with setae on api- metepisternum with deep anterodorsal cleft; cal half, all coxae about equal in size. Tibial gonostyli of simple shape (presumably re- and tarsal trichia not in rows. Abdomen se- versed in Plesiognoriste and Protognoriste). tose, with exception of 1st and 2nd sternites. Seventh sternite short, 8th small and retract- Archaeognoriste, new genus able. Ninth tergite small, rectangular, with DIAGNOSIS: Palpi long, three-segmented, well-developed, short, one-segmented cerci mouthparts reduced. Base of RS present, apically. Gonocoxites straight. Gonostyli sa- crossvein r-m distinct. All the coxae approx- berlike, thinner to apex, with one apical and imately equal in length. Hind tibiae slightly one very long curved basal processes. swollen. Gonostyli with long inner process. MATERIAL: Holotype AMNH Bu-1539, TYPE SPECIES: Archaeognoriste primitiva, male; paratypes AMNH Bu-412a and Bu- n.sp. 412b, 2 males in the same piece, Bu-485, Bu- ETYMOLOGY: The name is a combination 693, males. Myanmar: Katchin, from amber of archaeos (Greek ␣␳␹␣␫Â␱␵, or ancient) and mines near Myitkyina. the genus name . The name is fem- ETYMOLOGY: The species epithet is from inine. Latin primitivus meaning ``the ®rst or earliest of its kind'' and is a reference to the basal Archaeognoriste primitiva, new species position of the species within Lygistorrhinidae. OMMENTS Figures 7, 8, Plate 1D C : Venation of the species is al- most identical to Palaeognoriste sp. (Gri- DIAGNOSIS: As for genus. maldi and Blagoderov, 2001: ®gs. 5a, 6a), DESCRIPTION: Body length ϭ 1.58 mm (ho- with bases of RS and r-m somewhat reduced 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 13

Figs. 7±11. Archaeognoriste, n.gen. and Lebanognoriste, n.gen. 7. A. primitiva, n.sp., holotype AMNH Bu-1539 8. A. primitiva, male genitalia. 9. L. prima, n.sp. holotype AMNH JG268/1. 10. L. prima, male genitalia dorsally. 11. L. prima, male genitalia ventrally. 14 AMERICAN MUSEUM NOVITATES NO. 3433 and shifted proximally. Characters such as long, slender, straight, its length approxi- reduced palpi, membranous frons, the wing mately the combined length of preceding two venation, simpli®ed genitalic structure, and a palpomeres. Thorax: Mesonotum with few large metepisternum (presumably plesio- long lateral and scattered shorter setae. An- morphic) with deep anteriodorsal cleft obvi- terior parapsidal suture distinct. Scutellum ously refer the species to the Lygistorrhini- with 6 long setae and sparse short ones. Lat- dae, but compared to more derived species erotergites and mediotergite bare. Wing the new species has many plesiomorphies: membrane without macrotrichia, microtrichia presence of RS base and a distinct r-m, un- not arranged in rows. All longitudinal veins modi®ed legs, and gonostyli with long pro- with setae. Costa ends far beyond tip of R5. cesses. Humeral cross-vein transverse. R1 short, about 0.4ϫ wing length. Sc short, ends in R Lebanognoriste, new genus at middle of r-m. R5 slightly curved caudally. M slightly curved forward at tip. M fork DIAGNOSIS: Wing membrane without mac- 1 base and M stem absent. Crossvein r-m short, rotrichia, R short, crossvein r-m fused with 1 oblique, fused with base of M .M and base of M , fork of M and CuA not ses- 3ϩ4 3ϩ4 3ϩ4 3ϩ4 CuA stem short, but distinct. M almost sile; preapical palpomere attached at the tip 3ϩ4 straight, CuA curved gently back. Legs: Fore of antepreapical, inner mid and hind tibial tibiae with simple anteroapical depressed spurs longer than outer ones. area bearing one row of short bristles. Mid TYPE SPECIES: Lebanognoriste prima, n.sp. tibia with inner spur 1.5ϫ as long as outer ETYMOLOGY: The name is a combination one, inner hind tibia spur 1.3ϫ as long as of Lebanon, source country of the amber, and outer one. Tibial setulae in distinct rows. Tib- the genus name Gnoriste. The name is fem- ial spurs 2±3ϫ the tibial diameter. Tarsal inine. claw with one tooth. Abdomen short, setose, COMMENTS: Lebanognoriste together with with 7 visible segments. Genitalia rounded, Archaeognoriste form the most basal group rotated at 180Њ. Gonocoxites massive, fused of Lygistorrhinidae. Synapomorphies such as ventrally. Gonostyli hemicylindrical, slightly rather long fore coxae; short vein R ; short, 1 curved, shovel-like at apex. Ninth tergite transverse RS1 section; reduced base of the length equals the width, with two deep lateral M and M fork; and crossvein r-m shifted 1 2 incisions. proximally and aligned to M2 ϩ tb fused MATERIAL: Holotype AMNH JG268/1, vein, which is oblique or longitudinal and male. Lebanon: near Jezzine, coll. Aftim and shifted proximally, undoubtedly ally these Fadi Acra. two genera with the lygistorrhinids (see also ETYMOLOGY: The species epithet is a Latin Analyses). Both genera lack autapomorphies, word primus meaning ``®rst'' and is a refer- and comparison to recent representatives of ence to the oldest ®nd of the family in the the family indicates various plesiomorphies. fossil record. Lebanognoriste prima, new species Figures 9±11, Plate 1E Plesiognoriste, new genus

DIAGNOSIS: As for genus. DIAGNOSIS: Wing membrane without mac- DESCRIPTION: Body length ϭ 1.53 mm; rotrichia. Palpi short, two-segmented. Face wing length ϭ 1.64 mm. Head: Postocciput wide, about one-third of head width. Eyes of ¯at, with several strong setae behind eye male relatively small, setose. Proboscis not margin. Eyes setose. Ocelli not visible. Scape distended. Laterotergites bare. All veins with and pedicel twice as wide as ¯agellomeres, setae. R5 runs very close to R1. RS base ab- scape with one and pedicel with two apical sent. M2 absent. M3ϩ4 and CuA fork short. rows of setae. Flagellum with 14 cylindrical Middle and hind tibiae spurs almost equal in segments, covered by hairs as long as 1/2 length. Tibial setulae forming more or less ¯agellomere diameter. Face setulose. Only regular rows apically. three apical segments of palpi visible; ante- TYPE SPECIES: Plesiognoriste carpenteri, penultimate and penultimate knoblike; apical n.sp. 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 15

ETYMOLOGY: The genus name is a combi- ones, but broader, with scattered setae. Hind nation of plesion (Greek ␲␭␩␴␫Â ␱␯, or neigh- femora swollen, without ventral spines. Hind bor) and the genus name Gnoriste. The name tibiae with apical comb of setae. Tarsal claw is feminine. with one tooth. Abdomen shorter than COMMENTS: See under Protognoriste. wings, setose. Gonocoxites quadrate, ¯at, se- tose, with dorsal protruding appendage. Gon- Plesiognoriste carpenteri, new species ostyli simple, length ϳ3ϫ the width, apically Figures 12, 13, Plate 1F with two black inner spines and outer comb of short setae. DIAGNOSIS: Apical palpal segment curved, MATERIAL: Holotype MCZC 6927, male. bulbous; ¯agellomeres longer than wide, Canada: Manitoba, Cedar Lake, coll. F.M. anepisternite with small dorsal cleft; M3ϩ4 Carpenter. gently curved, M3ϩ4 and CuA stem weak- ETYMOLOGY: The species epithet is a pa- ened, fore and hind tibiae and hind femora tronym for the late Professor Frank Carpen- broadened. ter of the MCZ, who collected this and many DESCRIPTION: Body length ϭ 1.32 mm; others Canadian amber specimens. wing length ϭ 1.27 mm. Head: Antennae inserted near middle of head. Scape cylin- Plesiognoriste zherikhini, new species drical; pedicel subconical, apex wide with Figure 14, Plate 2A oblique margin. Flagellum 14-segmented, ¯agellomeres cylindrical, length of each DIAGNOSIS: Scape and pedicel not en- twice the width, covered with setulae as long larged, ¯agellomeres short, eyes form incom- as ¯agellomere diameter. Face wide, pentan- plete bridge, apical palpal segment rounded, gular, setose. Frons and occiput uniformly se- M3ϩ4 strongly curved, legs not modi®ed. tulose. Three ocelli present, equal in size, DESCRIPTION: Body length ϭ 1.40 mm; distance from middle to lateral ocelli the wing length ϭ 1.22 mm. Head: Postocciput same length as from lateral ocellus to eye ¯at. Antennae inserted near middle of face. margin (distance of lateral ocellus from eye Scape and pedicel not enlarged, widths margin equal to diameter of ocellus). Eyes equal to width of average ¯agellomere. Fla- setose, slightly emarginate at antennal bases. gellum 14-segmented, ¯agellomeres cylin- Palpi with two visible segments; apical seg- drical, width about equal to length, setulose. ment slightly curved, bulbous, setulose; basal Frons setose. Eyes setulose, emarginate at one with a few short setae. Thorax: Scutum antennal bases, forming lobes toward mid- with long lateral and dorsocentral setae. Scu- dle of head 4±5 facets wide, but dorsal tellum with 6 long setae. Antepronotum se- bridge incomplete, separated by distance tose, divided from proepisternum by a dis- more than 4ϫ facet diameter. Ocelli three, tinct suture. Anepisternum with small dorsal equal in size to the distance from middle to cleft. Mediotergite bare. Laterotergite not lateral ocelli, twice the length as from lateral fully visible. Wing membrane without mac- ocellus to eye margin (distance of lateral rotrichia; microtrichia not arranged in regular ocellus from eye margin equal to diameter rows. Costa ends slightly beyond tip of R5. of ocellus). Face setose. Palpi short, only 2 All veins bear setae. Sc very short, ends free, visible, rounded segments, with numerous humeral cross-vein almost transverse. R1 hairs. Clypeus bare. Thorax: Scutum with about 0.6ϫ wing length. RS base absent. R5 long lateral, dorsocentral, and acrostichal runs very close to R1, virtually straight; base setae. Scutellum with 6 long setae. Anterior weak, so that vein invisible proximal to Sc parapsidal suture distinct. Suture between tip. M1 weak, originates midway between antepronotum and proepisternum oblique. tips of R1 and R5.M2 and stem of M absent. Proepimeron touches mesepisternum at ane-

M3ϩ4 curved gently. CuA has distinct kink at pisternal suture. Anepisternal cleft narrow, base of M3ϩ4.M3ϩ4 and CuA stem weak. oblique. Mediotergite and laterotergites Legs rather short and stout. Fore coxae with bare. Metepisternum touches katepisternum. long bristles on anterior edge. Fore tibia wid- Wing membrane without macrotrichia, with ened. Hind coxae equal in length to middle scattered microtrichia. All veins setose. 16 AMERICAN MUSEUM NOVITATES NO. 3433

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 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 17

Costa ends beyond tip of R5, one-third the cerci simple. Face wide, with three ocelli, sit- length between tips of R5 and M1.Scvery uated almost in straight line, suggestive of short, ends free. R1 short, about 0.5 wing Manotinae, but structures of palpi, katepi- length. R5 about 0.85ϫ wing length, curved sternum and metepimeron, and absence of slightly, runs very close to R1.M1 con®ned strong setae, show no apomorphies with re- to apical third of wing. RS base, M1 and spect to that subfamily. Eye size, face width, stem of M absent. Crossvein r-m fused with vein vestiture, tibial spurs of equal size, and

R5, horizontal, fused with CuA base. CuA complicated shape of the gonostyli are more base weak. Base of M3ϩ4 and CuA fork at primitive conditions than occur in Recent ly- level of tip of R1.M3ϩ4 curved and well gistorrhinids, although some features of ve- rounded. CuA with two straight segments nation and gonostyli are apomorphic. The forming obtuse angle. Legs: Coxae almost short fork of M3ϩ4 and CuA and absence of equal in length. Hind coxae bare at base. a long proboscis are most similar to the ge- Fore tibia distally with apical hemispherical nus Seguyola Matile. It is possible that these anteroapical depression with comb of ®ne two genera should be treated as a separate setae. Tarsal claw without teeth. Abdomen subfamily of Lygistorrhinidae. We prefer to shorter than wings, with 7 visible segments, expand the de®nition of the family to include setose. Sixth and 7th segments twice as newly described taxa. short as 5th, 8th retracted. Cerci one-seg- mented. Protognoriste amplicauda, new species MATERIAL: Holotype PIN 3311/664, fe- Figure 15, Plate 2B male. Russia: Taimyr Peninsula, Yantardakh, coll. 1971. DIAGNOSIS:R1 length 3ϫ that of r-m. Gon- ETYMOLOGY: The species epithet is a pa- ocoxite massive. Gonostyli straight, ¯attened tronym in honor of the late Dr. Vladimir dorsoventrally. Zherikhin of the Paleontological Institute in DESCRIPTION: Body length ϭ 1.25 mm; Moscow, prominent paleoentomologist, who wing length ϭ 1.11 mm. Head: Eyes setu- collected the specimen. lose, setae very short. Ocelli three, equal, al- most in straight line, distance of lateral ocel- Protognoriste, new genus lus from eye margin equal to distance from mid ocellus to lateral one. Vertex and frons DIAGNOSIS: Wing membrane without mac- setose. Scape very small. Pedicel spherical, rotrichia, occiput without row of strong se- obscure. Flagellum 14-segmented, ¯agello- tae. Palpi short. Eyes bare. Face wide. Stem meres cylindrical, lengths about equal to of M and M2 absent. RS base distinct. Cross- widths, setose. Palpi short, 3-segmented, api- vein r-m meets M3ϩ4 and CuA stem. CuA cal segment rounded, penultimate cylindri- gently curved. Abdomen insertion broad. cal, length 2ϫ the width. Face wide, quad- Fore tibiae shorter than femora. rate. Thorax: Scutum with lateral, dorsocen- TYPE SPECIES: Protognoriste amplicauda, tral and acrostichal setae. Scutellum with n.sp. several long setae. Mediotergite with short ETYMOLOGY: The name is a combination trichia. Metepisternum with anterodorsal of protos (Greek ␲␳␻Â ␶␱␵, or ®rst) and the cleft, height about equal to width. Wing genus name Gnoriste. The name is feminine. membrane without macrotrichia. Costa ends

COMMENTS: The following apomorphic beyond tip of R5, at 1/6 the length between characters refer the genera Plesiognoriste tips of R5 and M1. Sc short, ends free at the and Protognoriste to the Lygistorrhinidae: level of RS base. Humeral cross-vein trans- short palpi; small dorsal cleft of anepister- verse. R1 short, about 0.4ϫ wing length. R5 num; anepisternum and metepimeron not di- about 0.8ϫ wing length, almost straight. RS vided by laterotergite (possibly plesiomorph- base very short, transverse. Crossvein r-m ic); short, incomplete Sc; short R1; RS base horizontal, meets M3ϩ4 and CuA stem, its and M stem reduced; r-m horizontal and length 3ϫ less than R1 length. M1 almost fused with R5; hind coxa smaller and broader straight, weakened at base. M2 and M stem than mid coxa; abdomen insertion narrow; absent. Base of M3ϩ4 and CuA fork between 18 AMERICAN MUSEUM NOVITATES NO. 3433

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 R1 tip. CuA and M3ϩ4 tose. Eighth segment short, retracted. Gono- curved caudally. Legs: Coxae almost equal coxite massive. Gonostyli straight, ¯attened in length, hind coxae bare. Tibial setulae in dorsoventrally. distinct rows. Tibiae, especially fore, short. MATERIAL: Holotype PIN 3426/257, male. Tarsal claw with small obtuse tooth. Abdo- Russia: Taimyr Peninsula, Nizhnyaya Agapa, men as long as wing, with 8 segments, se- coll. 1973. 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 19

ETYMOLOGY: The species epithet is derived Cerci with large basal and small round apical from Latin words amplus meaning ``large, segments. distinguished'' and cauda meaning ``tail'' in MATERIAL: Holotype AMNH Bu-406, fe- reference to large terminalia of the species. male. Myanmar: Katchin, from amber mines near Myitkyina. Protognoriste goeleti, new species ETYMOLOGY: The species epithet in honor of Mr. Robert Goelet, for his generosity in Figure 16, Plate 2C funding purchase of specimens and the au- thors' work. DIAGNOSIS:R1 length 1.8ϫ that of r-m. DESCRIPTION: Body length ϭ 1.21 mm; wing length ϭ 0.95 mm. Head: ovate, height Protognoriste nascifoa, new species ϳ1.5ϫ width in pro®le. Occiput and postgena Figures 17, 18, Plate 2D densely setose. Antennae attached below the middle of the head. Scape and pedicel sub- DIAGNOSIS: RS base and the base of the conical, wider than ¯agellum. Flagellomeres M3ϩ4 and CuA fork very basally. R1 length cylindrical, length 1.1±1.5ϫ the width. Eyes 4ϫ r-m. setose, ovate, without incision, facets round, DESCRIPTION: Body length ϭ 1.09 mm; close. Clypeus bare. Two palpomeres seen: wing length ϭ 1.05 mm. Head: Occiput and basal one ovate, length 2ϫ the width, apical frons setose. Pedicel and scape wider than round, 4ϫ shorter than basal. Thorax: Scu- ¯agellomeres. Flagellum 14-segmented, ¯a- tum irregularly setose. Anterior parapsidal su- gellomeres cylindrical, length about equal to ture distinct. Scutellum small. Anepisternum width. Face wide, quadrate. Clypeus setose, palpi 2-segmented, basal palpomere swollen, with several setae. Proepimeron touches me- apical one very small. Eyes with incision, sepisternum at anepisternal suture. Anepister- forming incomplete eye bridge (possibly ar- num with deep cleft in the middle. Lateroter- tifact, face deformed). Thorax: Proepimeron gite shifted dorsocaudally, so that metepister- contacts katepisternum. Anepisternum width num contacts katepisternum for some dis- 1.5ϫ height, with distinct cleft posteriorly. tance, not in a point. Laterotergites and Laterotergites, mediotergite and metepister- metepisternum with short hairs. Mediotergite num with very short trichia. Scutum with nu- short. Meron at mid coxae large. Wing mem- merous erect setae. Metepisternum with long brane with microtrichia not arranged in rows. anteriodorsal process touching katepister- Costa ends beyond tip of R5, at one-fourth num. Wing: Sc very short, free. Costa pro- length between tips of R5 and M1.Scvery duced beyond R one-third distance between short, ends free. Humeral cross-vein oblique. 5 R5 and M1 apices. RS base situated proxi- R1 short, about 0.4ϫ wing length. RS base mally, faint, oblique. R1 length 4ϫ that of short, transverse. R5 about 0.8ϫ wing length, r-m. Crossvein r-m weak, 4ϫ shorter than R1. almost straight. M1 slightly sigmoid, con®ned M originates at distal third of wing, curved to apical third of wing, ends at tip of wing. 1 at base. Base of the M3ϩ4 and CuA fork at Stem of M and M2 absent. Crossvein r-m hor- the level of RS base. Legs: Hind coxae short- izontal, very weak, meets M3ϩ4 and CuA er than mid ones. Fore coxae with long an- stem, its length 1.2ϫ more than length of R1. terior setae, mid and hind ones with apical M3ϩ4 and CuA fork 1.7ϫ stem length, its base setae. Tibial setulae not in rows except apical between levels of RS base and tip of R1.M3ϩ4 half of hind tibiae. Hind tibiae long, swollen and CuA curved gently. Legs: Fore tibia somewat at apex, with dorsal row of bristles 1.25ϫ shorter than fore femora. Tibial and and apical comb of setae. Tibial spur length tarsal setulae not in rows. Hind tibia with 15 1.0±1.3ϫ tibial diameter. Abdomen setose, dorsal bristles. Tibial spurs 2ϫ the diameter segments short. Genital complex wider than of tibia. Hind tibiae with apical comb of short long. Gonocoxites fused. Gonostyli with one setae. Fore coxae with dense anterior setae, outer and two inner lobes. mid and hind coxae with setae in apical part, MATERIAL: Holotype AMNH Bu-434, hind coxae without posterior setae. Abdomen male. Myanmar: Katchin, from amber mines insertion broad. Abdomen shorter than wings. near Myitkyina. 20 AMERICAN MUSEUM NOVITATES NO. 3433

ETYMOLOGY: The species epithet derived and acrostichal and shorter irregular setae. from U.S. National Science Foundation, a Anterior parapsidal suture distinct. Antepron- generous sponsor of this and other fossil in- otum and proepisternum setose. Anepister- sect research at the AMNH. num wider than its height. Proepimeron touches katepisternum slightly below anepi- Leptognoriste, new genus sternal suture. Mesepisternum with long an- terodorsal process, touching anepisternum. DIAGNOSIS: Palpi short, 4-segmented. Laterotergites with row of long setae. Me- Mouthparts form short proboscis or reduced. diotergite irregularly setose with short setae. Laterotergites and mediotergite setose. Wing Legs: Coxae with relatively short setae. Hind membrane with macrotrichia. Sc long, ends tibiae with bristles in dorsal row, 1.5ϫ longer at C. Costa produced beyond R , not reach- 5 than femora. Abdomen setose, except ®rst ing wing apex. M absent. M reduced at the 1 2 sternite. Abdomen insertion very narrow. base. RS base transverse. M and CuA fork 3ϩ4 Tergite 8 shorter than sternite. Tergite 9 nar- stalked. Hind coxae shorter than fore and row, rectangular with numerous setae at mid ones. Male genitalia simple. apex. Gonocoxites rather slender. Gonostyli TYPE SPECIES: Leptognoriste davisi, n.sp. slightly curved. ETYMOLOGY: The name is a combination MATERIAL: Holotype AMNH Bu-126a, of leptos (Greek ␭⑀␲␶␱Â ␴, or thin, lean) and male; paratype AMNH Bu-126b, male, in the the genus name Gnoriste. The new name is same piece. Myanmar: Katchin, from amber feminine. mines near Myitkyina. ETYMOLOGY: The species epithet is a pa- Leptognoriste davisi, new species tronym for Mr. Jim Davis, who supplied the Figures 19±22, Plate 2E AMNH with fossiliferous amber from Myan-

DIAGNOSIS: The base of M3ϩ4 and CuA mar. fork at level of base of RS. Mouthparts form short proboscis. M1 reaching wing margin. Leptognoriste microstoma, new species Ninth tergite small, longer than wider, rect- Figures 23, 24, Plate 2F angular.

DESCRIPTION: Body length ϭ 1.94 mm (ho- DIAGNOSIS: Mouthparts reduced. R1 shorter lotype)/1.69 mm (paratype); wing length ϭ than r-m. M2 weak at apex. Base of M3ϩ4 and 1.26 mm (holotype)/1.27 mm (paratype). CuA fork beyond the level of RS base. Male Head round with protruding ocelli. Ocelli in 8th tergite wider than long, rounded. triangle, lateral separated from medial by DESCRIPTION: Body length ϭ 1.33 mm; ocellus diameter and from eye margin by 2 wing length ϭ 1.24 mm. Head: Occiput with ocellus diameters. Eyes large, rounded, with long setae. Pedicel slightly wider then scape light emargination, with large facets. Clypeus and ¯agellum, ¯agellomeres barrel-shaped, narrow, triangular, setose. Palpi 4-segmented, as long as wide. Palpi 4-segmented, palpo- palpomere length ratio 1:2:2:3. Apical pal- mere length ratio 1:2:2:3.5. Thorax: Scutum pomere attached preapically. Short proboscis, with long lateral, acrostichal, and dorsocen- one-half head height. Wing length equal to tral setae and short setae between rows. Pro- abdomen length. Sc produced slightly be- epimeron touches katepisternum below the yond RS base. R1 length 1.2ϫ r-m. M2 orig- episternal suture. Anepisternum width about inates in distal half of wing. Base of M3ϩ4 equal to height. Laterotergites with long se- and CuA fork at level of RS base. Macrotri- tae. Mediotergite with several long setae ven- chia numerous in basal part of wing. Tho- trally. Mesepisternum with long anterodorsal rax: Scutum with long lateral, dorsocentral process, touching anepisternum. Wing length

→ 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. 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 21 22 AMERICAN MUSEUM NOVITATES NO. 3433

1.5ϫ the abdomen length. R1 slightly shorter long protruding setae behind eye margin. than r-m. Macrotrichia in basal part of wing Eyes slightly emarginate, densely setose, se- absent. M2 weak at apex. Base of M3ϩ4 and tae length 2ϫ facet diameter. Facets round, CuA fork beyond the level of RS base. Legs: densely set. Three ocelli in triangle, close to Fore coxae with dense long anterior setae, each other. Frons and face setulose. Antennae mid and hind ones with apical. Hind tibiae inserted above middle of head. Scape and without dorsal bristles, only slightly longer pedicel subconical, with apical setae. Flagel- than femora. Abdomen. Tergite 9 wider than lum 14-segmented, ¯agellomeres com- long, rounded on apex. Sternite I bare. Gon- pressed, widths 1.2±2ϫ length, apical one ostyli curved at apex. conical. Texture of ¯agellomeres polygon- MATERIAL: Holotype AMNH Bu-429. like. Only 3 segments of palpi visible, ante- Myanmar: Katchin, from amber mines near penultimate and penultimate with strong se- Myitkyina. tae; penultimate 2.5ϫ as long as preceding ETYMOLOGY: The species epithet is a com- one, ovate; apical segment 1.7ϫ length of bination of micros (Greek ␮␫␬␳␱␨, or small) penultimate one, narrow. Thorax: Scutum and stoma (Greek ␴␶␱Â ␮␣, or mouth), in ref- uniformly densely setose with short setae, erence to the reduced mouthparts. bearing long lateral and posterior setae. An- terior parapsidal suture distinct. Suture be- FAMILY MYCETOPHILIDAE NEWMAN, 1834 tween antepronotum and proepisternum com- plete, both segments uniform and covered SUBFAMILY MANOTINAE EDWARDS, 1925 with bristles. Scutellum with long setae. Alavamanota Blagoderov and Arillo, 2002 Anepisternum wider than higher, with wide Alavamanota Blagoderov and Arillo, 2002: 34. dorsal cleft. Anepisternal suture declines posteriorly. Katepisternum setose ventrally. DIAGNOSIS: Antepronotum and proepister- Mediotergite and laterotergites bare. Met- num completely divided; mediotergite and episternum with several light hairs postero- laterotergites bare; wing membrane with or ventrally. Wing membrane with macrotri- without macrotrichia; R1 relatively short, chia; microtrichia not arranged in rows. Cos- length of R1 about the length of r-m;R4 pre- ta ends slightly beyond the tip of R5, C and sent; M3ϩ4 and CuA fork with stem. R5 run very close to each other in apical part. TYPE SPECIES: Alavamanota hispanica Bla- R1,R5, r-m, M1, and M2 with setae ventrally goderov and Arillo, 2002. and dorsally. Sc very short, ends free. Hu- COMMENTS: Closest to the recent genus meral vein oblique. R1 about equal to length Manota Williston, 1896, distinguished by hav- of r-m. Ratio of r-m and RS2 section is 1: ing the fourth palpomere attached preapically 1.4. Base of M1 and M2 fork at the level of but very close to the apex of the third; two R4. Small radial cell with length 6ϫ width. crossveins instead of one between R1 and R5 Crossvein r-m horizontal, fused to tb, meets (RS1 and R4); tibial trichia irregularly ar- MA. M3ϩ4 and CuA fork long, but not sessile. ranged; sternite 9 separate; sternite 8 without CuA curved caudally. Legs. Hind coxae bear 4 strong protuberances bearing long setae; and apical only, not posterior setae. Tibial spur the basal segment of each cercus small. lengths ca. 4ϫ tibial diameter. Tibial trichia in rows. Abdomen setose, with 6 visible seg- Alavamanota burmitina, new species ments. Gonocoxites fused, lighter then ab- Figure 25, Plate 3A domen, setose, swollen. Gonostyli not seen. MATERIAL: Holotype AMNH Bu-1271, DIAGNOSIS: Flagellum compressed dorso- male; paratypes AMNH Bu-428a, Bu 279a, ventrally; mesonotum with long lateral setae; males. Myanmar: Katchin, from amber mines wing membrane with macrotrichia; length of near Myitkyina. small radial cell 6ϫ width; base of M1 and M3 section weak; tibial setulae arranged in rows. SUBFAMILY SCIOPHILINAE WINNERTZ, 1863 DESCRIPTION: Body length ϭ 2.50 mm (ho- TRIBE SCIOPHILINI WINNERTZ, 1863 lotype)/1.62±2.5 mm (paratypes); wing Neuratelia Rondani, 1856 length ϭ 1.69 mm (holotype)/1.08±1.85 mm Neuratelia Rondani, 1856: 195. (paratypes). Head: Postocciput with row of Anaclinia Winnertz, 1863:770. 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 23

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 AMERICAN MUSEUM NOVITATES NO. 3433

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

DIAGNOSIS: 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, subdivid- Fig. 27. Allocotocera burmitica, n.sp., holo- ed or with elaborate processes. type AMNH B-056, reconstruction of wing ve- TYPE SPECIES: Mycetophila nemoralis Mei- nation. gen, 1818: 256 (orig. des.). COMMENTS: Two species of the genus were insula, Siberia, where the amber deposit is described from Baltic amber (Meunier, 1904) located.

Neuratelia maimecha, new species Allocotocera Mik, 1886 Figure 26, Plate 3B Eurycera Dziedzicki, 1885:166. DIAGNOSIS: Sc short, ends before RS base; Allocotocera Mik, 1886: 102. length of RS1 section 2.5ϫ that of crossvein Euryceras Marshall, 1896: 291. r-m; base of M and CuA fork at the middle 3ϩ4 DIAGNOSIS: Laterotergites and mediotergite of r-m; mediotergite with sparse, ®ne trichia. setose. Wing membrane with macrotrichia DESCRIPTION: Body length ϭ 2.11 mm and with or without microtrichia. Sc preap- (rest); wing length ϭ 2.05 mm. Head: Fla- 2 ical or medial. Base of M and CuA fork gellum 14-segmented, ¯agellomeres cylindri- 3ϩ4 before the base of the fork of M and M . cal, with length about equal to width. Scape 1 2 TYPE SPECIES: Eurycera ¯ava Dziedzicki, and pedicel rounded. Mouthparts form pro- 1885: 167 [ϭ pulchella (Curtis, 1837)] (by boscis slightly shorter than head height. Palpi monotypy). 4-segmented, basal and antepenultimate seg- COMMENTS: Another fossil species occurs ments oval, penultimate and apical segment in Lower Cretaceous Spanish amber (Bla- bacilliform, apical slightly longer and nar- goderov and Arillo, 2002). rower than penultimate, length ratio 1:3:4: 5.5. Thorax: Scutum with numerous, short, Allocotocera burmitica, new species scattered setae and long lateral ones. Ante- Figure 27, Plate 3C pronotum with three setae, proepisternum with ®ve. Anepisternum bare, with deep cleft DIAGNOSIS: Wing membrane with macro- in posterior part. Anepisternal suture declines and microtrichia. Segment M3 2.5ϫ length posteriorly. Metepisternum quadrate, touch- of r-m, 0.3ϫ that of M fork. Sc ends at level ing katepisternum. Laterotergites with sev- of RS base; Sc2 in the apical one-fourth of eral setae, mediotergite with sparse, ®ne tri- Sc, distal to midpoint of Sc. chia caudally. Wing membrane with micro- DESCRIPTION: Body length ϭ 2.86 mm; trichia and few macrotrichia. Costa ends be- wing length ϭ 2.46 mm. Head: Flagellum yond tip of R5. Sc meets C just before RS 14-segmented, thickened to apex. Flagello- base. Sc2 absent. Longitudinal veins with se- meres almost equal in length, length of the tae. M3ϩ4 and CuA base slightly distad of M3 ®rst one 2ϫ its width, apical one 5ϫ its base. Legs: Tibiae with distinct bristles. Fore width. Scape and pedicel subconical, with tibia longer than ®rst tarsomere. small apical setae, 2ϫ wider than ¯agello- MATERIAL: Holotype PIN 3311/661, end of meres. Eyes pubescent. Two palpomeres abdomen not preserved, sex unknown. Rus- seen, cylindrical, length ca. 4ϫ the width. sia: Taimyr Peninsula, Yantardakh, coll. Thorax: Scutum irregularly covered with 1971. short setae and with strong, long lateral setae. ETYMOLOGY: Species epithet is derived Antepronotum with long setae. Scutellum from the Maimecha river on the Taimyr Pen- with two pairs of long setae and several more 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 25 short ones. Laterotergites and mediotergite long and slender gonocoxites (which are not with setae. Metepisternum with two short se- known for any manotines). Within Sciophil- tae posteroventrally. inae the new genus might be close to the Wing membrane clear, with macro- and Azana group of genera (Matile, 1998), dem- microtrichia. Costa ends at tip of R5.Sc onstrating reduction of median veins. All of meets C beyond RS base. Sc2 at level of base the genera in the group have reduced vena- of section M3. Section of RS1 transverse. R1 tion, with CuA simple and not forming a fork rather short, about 0.7ϫ wing length. R5 with M3ϩ4. Matile (1998) supposed that M3ϩ4 curved caudally, not reaching wing apex. in this group was reduced completely, while

Crossvein r-m approximately 2.5ϫ as long as the fork of M1 and M2 lost its base. Jugding section RS1 and 2.5ϫ length of section M3. on the position of vein apices, at least Par- Fork of M1 and M2 3.3ϫ as long as section atrizygia, Neoaphelomera, and Neotrizygia

M3. Base of M3ϩ4 and CuA fork slightly be- may have lost M2, and an incomplete vein fore base of section M3. Legs: Hind coxae between M1 and CuA may be homologous to with long posterior setae. Fore tibiae longer M3ϩ4. The new species has venation more than ®rst tarsomere. Cerci 1-segmented, 2ϫ primitive than the genera in the Azana group, as long as 8th tergite, setose. preserving all the longitudinal veins, al- MATERIAL: Holotype AMNH B-056, fe- though without bases. male. Myanmar: Katchin, from amber mines near Myitkyina. Pseudomanota perplexa, new species ETYMOLOGY: The species epithet is a ref- Figures 28±30, Plate 3D erence to the country of origin of the amber. COMMENTS: The species is very close to A. DIAGNOSIS: As for the genus. xavieri, but differs in having a shorter Sc DESCRIPTION: Body length ϭ 1.44 mm; vein and a longer M1 and M2 fork. wing length ϭ 1.07 mm. Head: Eyes large, setulose, with large facets. Occiput and post- Pseudomanota, new genus gena setose, without erect bristles. Ocelli not seen. Antennae attached above middle of DIAGNOSIS: Infraorbital setae absent. Three head. Frons, face and clypeus setose. Pedicel palpomeres, penultimate one with long thin subconical. Flagellum 14-segmented, slightly apical appendage. Antepronotum wide. Pro- thicker toward apex; ¯agellomeres cylindri- episternum rodlike. Laterotergites and me- cal, width 1±1.5ϫ length. Three palpomeres diotergite bare. Sc short, free. Wing mem- seen, basal one heart-shaped, penultimate brane with macrotrichia. Base of M1 and M2 one with long thin apical appendage about fork reduced. M1 reaches wing margin before one-third its length, apical one long and slen- wing apex. M3ϩ4 free at the base. Gonostyli der. Thorax: Antepronotum and proepister- simple, slightly curved. num wide, completely divided, setose. Pro- TYPE SPECIES: Pseudomanota perplexa epimeron narrow, rodlike. Scutum with lat- n.sp. eral, acrostichal, and dorsocentral setae with ETYMOLOGY: The genus name is a combi- bare strips in between. Anterior parapsidal nation of pseudos (Greek ␺⑀␷Æ ␦␱␴, or a lie, a suture distinct. Anepisternal suture declines fraud) and the genus name Manota. The posteriorly. Anepisternum with wide cleft. name is feminine. Metepisternum with long, narrow anterodor- COMMENTS: Some structures, especially the sal process touching laterotergite. Lateroter- wide quadrate face, long and ¯exible apical gites and mediotergite bare. Wing membrane palpomere, short Sc and R1, horizontal r-m, with macrotrichia and microtrichia. Humeral and reduced venation are like Manotinae. vein slightly distad of MA. Costa extends to

Also, unlike typical Sciophilinae, the new R5 apex at two-thirds distance between tips genus has laterotergites and especially the of R5 and M1, not reaching wing apex. mediotergite bare. Nevertheless, the genus is Length of R1 is 0.6ϫ wing length. Small ra- attributed to the subfamily Sciophilinae dial cell length 5ϫ width. M1 originates at based on a long, rodlike proepimeron, the apical one-third of wing, reaching wing mar- rather distal position of humeral vein, and gin before wing apex. M2 originates at apical 26 AMERICAN MUSEUM NOVITATES NO. 3433

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

half of wing. M3ϩ4 free at the base. M1,M2, ETYMOLOGY: The species epithet is from M3ϩ4, and CuA apices weak. Legs: Tibial se- the Latin word perplexus, meaning ``mud- tulae not arranged in rows except on apex of dled, intricate'', in reference to the intriguing mid and hind tibiae. Tibial spurs ca. 3.5ϫ combination of sciophiline and manotine tibia diameter. Hind tibia with a dorsal row characters. of bristles. Abdomen setose, with 6 seg- ments visible. Genital complex as long as TRIBE GNORISTINI EDWARDS, 1925 4th±6th segments combined, rotated 180Њ. Tergite 9 small, apex round, covering basal Apolephthisa Grzegorgzek, 1885 part of gonocoxites only. Aedeagus with two Apolephthisa Grzegorgzek, 1885: 205. teeth apically. Gonocoxites separated, mas- sive, setose. Gonostyli simple, sticklike, DIAGNOSIS: Mediotergite bare; lateroter- slightly curved, apically ¯at, shovel-like. gites setose; Sc meets C; R4 present; base of

MATERIAL: Holotype AMNH Bu-599a, M3ϩ4 distad of r-m, faint; M2 several times male. Myanmar: Katchin, from amber mines longer than r-m. near Myitkyina. TYPE SPECIES: A. rara Grzegorgzek, 1885: 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 27

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

206 (by monotypy) [ϭ subincana (Curtis, DESCRIPTION: Body length ϭ 2.24 mm; 1837) ()] wing length ϭ 1.46 mm. Head: Eyes slightly COMMENTS: This genus is widely distrib- emarginate near antennal base. Flagellum uted in the Holarctic but is of low diversity, with 14 cylindrical ¯agellomeres, covered by including only one described Palaearctic and trichia equal to 0.5ϫ ¯agellomere diameter. one undescribed Nearctic species (Vocker- Palpi rather long; antepenultimate segment oth, 1981). Apolephthisa mesozoica is de- oval, broadened; penultimate and apical seg- scribed from the Lower Cretaceous of Mon- ments long and slender, together equal to fore golia (Blagoderov, 1998a), and one unde- coxa in length; apical segment slightly scribed specimen is known from Palaeocene clubbed at apex, with several short setae. Pal- Sakhalin amber (Blagoderov, in prep.). pomere length ratio 1:2:4. Thorax: Scutum with lateral, dorsocentral, and acrostichal Apolephthisa bulunensis, new species rows of short setae. Mediotergite bare. Figures 31, 32, Plate 3E Wing: Veins R1,R5,M1, and M2 with setae. DIAGNOSIS: Sc short, ends before RS base. Sc meets C before RS base. R5 almost Gonostyli with distinct rounded apical lobe straight. Costa ends beyond tip of R5 one- and long inner process. ®fth the distance between tips of R5 and M1. 28 AMERICAN MUSEUM NOVITATES NO. 3433

M1,M2, and M stem weak. Base of M3ϩ4 ab- large, rounded, with round facets. Flagellum sent. Legs: Hind coxae without posterior se- 14-segmented, ¯agellomeres cylindrical, tae. Abdomen setose, with 7 visible seg- length of each about 1.5ϫ its width. Two pal- ments, 8th one small and retracted. Gono- pomeres seen. Penultimate 0.7ϫ length of coxites long, stout, thinner apically. Gono- apical one, which is slender and ¯exible. styli with rounded apical lobe, bearing Thorax: Scutum irregularly setose. Medi- dorsoventral row of bristles and long inner otergite and laterotergites bare. Wing: R1 process, ending in a short spur. and R5 almost straight, with setae, remaining MATERIAL: Holotype PIN 3963/4, male. veins bare, except apex of M3ϩ4, a few setae Russia: Taimyr Peninsula, Bulun, coll. 1976. present on M2 and CuA1 dorsally. Length ra- ETYMOLOGY: The species epithet is topo- tio of sections RS1, RS2, and RS3 is 1:1.4: nymic. 13. Crossvein r-m slightly sigmoid, its length COMMENTS: Given the current low diver- 2ϫ length of M3 section. Base of M3ϩ4 ab- sity of the genus, it would seem highly im- sent, the vein originates at level of M3 mid- probable for three fossil species to occur un- point. CuA and M3ϩ4 gently curved caudally. less the genus was more diverse and abun- Legs: Midtibial organ absent. Abdomen dant in the past. with 6 visible segments, 7th and 8th seg- ments small and retractable. Gonocoxites Synapha Meigen, 1818 fused. Gonostyli long, straight, with numer- ous stiff spines arranged in rows on inner Synapha Meigen, 1818: 227. surface and several dark, curved ones at Sinapha Rondani, 1856: 196, misspelling. base. Empalia Winnertz, 1863: 762. MATERIAL: Holotype MCZC 6944, male, DIAGNOSIS: As given by Freeman (1951): Canada: Manitoba, Cedar Lake. Setae on stems and forks of both M and Cu ETYMOLOGY: The species epithet refers to present, ¯agellomere lengths not more than the long gonostyli of the species. width, Sc long, penultimate palpomere short- COMMENTS: This species lacks distinct syn- er than apical and antepenultimate ones, la- apomorphies of Synapha (i.e., midtibial sen- terotergites bare or setose, midtibial organ sory organ and short gonostyli) as well as present. strong setae on the medial and cubital forks. TYPE SPECIES: S. fasciata Meigen, 1818 The fossil has moderately long ¯agellomeres (by monotypy). and subequal palpomeres, considered to be COMMENTS: About 25 extant species of synapomorphies of Austrosynapha (see table Synapha are known from all biogeographic 2). Long, slender gonostyli, which are char- regions. Two fossil species of the genus were acteristic for Austrosynapha and many other described from Eocene Baltic amber (Meu- Mesozoic fungus gnats, is considered to be nier, 1904) and the Lower Cretaceous of plesiomorphic. Most likely, the new species Montsec, Spain (Blagoderov and MartõÂnez- together with the Lower Cretaceous S. rubie- DelcloÂs, 2001). The related genus Austrosy- sensis Blagoderov and MartõÂnez-DelcloÂs, napha Tonnoir, 1929 has 23 species from 2001 represent a Synapha-Austrosynapha South America, which were cataloged by Pa- stem group, although monophyly of the pavero (1977b); 7 occur in New Zealand group and included genera is still to be es- (Tonnoir and Edwards, 1927) and one is tablished. Matile (1991) noted that Synapha from Australia (Tonnoir, 1929). was probably polyphyletic. The de®nition of both genera is vague, and often characters of Synapha longistyla, new species different subgroups overlap. For example, Figures 33, 34, Plate 3F species of A.(Paraaustrosynapha) Duret have short ¯agellomeres, while species of A. DIAGNOSIS: Distinguished from living and (Neoaustrosynapha) Duret retain some mac- fossil members by the structure of genitalia, rotrichia on the medial fork (Duret, 1980). having gonostyli very long and with numer- ous inner spurs. Dziedzickia Johannsen, 1909 DESCRIPTION: Body length ϭ 2.15 mm; Hertwigia Dziedzicki, 1885: 166 (preocc. Schmidt, wing length ϭ 1.58 mm. Head round. Eyes 1880). 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 29

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

Dziedzickia Johannsen, 1909: 44. occiput greyish brown, tibiae and tarsi yel- lowish. Head: Eyes large, bare, slightly DIAGNOSIS: After Freeman (1951): Eyes emarginate at level of antennae; 3 ocelli in a emarginate; three ocelli in line. Median ocel- straight line; laterotergal hairs present or ab- lus slightly sunken, distance from middle to sent; wing membrane without macrotrichia; lateral ocellus and from lateral one to eye Sc ending at R before or beyond base of RS, margin ϳ1.5ϫ ocellus diameter. Scape and Sc may be present as a short stump; base of pedicel shorter than ¯agellomeres, bare. Fla- 1 gellomeres cylindrical, length ϳ1.5ϫ width, M3ϩ4 and CuA fork considerably basal to M1 and M fork; R may be present or absent. length of apical one ϳ2ϫ the width. Three 2 4 palpomeres seen, short, gradually thinner to- TYPE SPECIES: Hertwigia marginata Dzied- zicki, 1885: 165 (by monotypy). ward apex. Mouthparts less than one-fourth height of head. Thorax: Mesonotum very COMMENTS: About 50 extant species of the genus are described, mostly from the New convex, with dorsocentral, acrostichal, and World, especially South America, with sev- lateral setae well developed. Antepronotum eral species from Holarctic and Afrotropics. and laterotergite setose, other sclerites bare. Four species are known from Baltic amber Proepimeron thick, short, touches katepister- (Meunier, 1917a, 1917b, 1922) and one from num. Anepisternal suture declines posterior- the Oligocene of Rott (Statz, 1944). Some ly. Wing membrane without macrotrichia. Sc authors (Chandler, 1999; Hutson, 1979; Ma- ends at R1 beyond RS base. Sc1 absent. RS1, tile, 1992; Vockeroth, 1980) noted that the RS2, and RS3 ratio 1:1.2±1.3:11±15. M3 genus in present de®nition was plausibly length 2ϫ that of r-m crossvein. Base of polyphyletic since some of the species be- M1ϩ2 fork at level of R4. Base of M3ϩ4 and long to other gnoristine genera, such as Syn- CuA fork before base of r-m. Abdomen se- temna Winnertz, 1863 and Hadroneura tose, 7th segment 2ϫ shorter than 6th. Male: LundstroÈm, 1906. Gonocoxites massive, bristly, separated ven- trally by deep cleft. Ninth tergite short, trans- Dziedzickia nashi, new species verse, covers only base of gonocoxites. Gon- ostyli with a strong, thin, dark process Figures 35, 36, Plate 4A pointed inward. Female with cerci 2-seg- DIAGNOSIS: Laterotergites setose. Forked mented. Basal segment 2.5ϫ apical one, cy- veins without setae. Tergite 9 short. Gono- lindrical. Apical segment rounded. coxites simple. Gonostyli with strong inner MATERIAL: Holotype AMNH NJ-117a, process. male. Paratypes: AMNH NJ-117b, c, e, m, DESCRIPTION: Body length ϭ 2.96 mm (ho- females, NJ-117d, g, h, males, NJ-117f lotype)/2.92±3.37 mm (paratypes); wing (wing), all in the same piece of amber. USA: length ϭ 2.50 mm (holotype)/2.28±3.32 mm New Jersey, Sayerville, coll. P. Nascimbene. (paratypes). Brown gnats, thorax dark brown, See ®gure 38 for a map of syninclusions. 30 AMERICAN MUSEUM NOVITATES NO. 3433

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

ETYMOLOGY: The species epithet is patron- COMMENTS: Differs from all other species ymic for Mr. Paul Nascimbene, who collect- by the structure of male genitalia. A thor- ed the piece and who has diligently prepared ough comparision will be possible only after numerous amber specimens for the AMNH. revision of the genus. 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 31

Saigusaia pikei, new species Figure 37, Plate 4B

DIAGNOSIS: Distinguished from the living species by Sc long, meeting C just before RS base. DESCRIPTION (see also table 3): Body length ϭ 2.84 mm, wing length ϭ 1.80 mm. Head: Ocelli three, each slightly raised, very close. Eyes pubescent. Vertex and clypeus setulose. Flagellum 14-segmented, ¯agello- meres cylindrical, with length slightly greater than width. Mouthparts form short proboscis, length is one-half the head height. Only three Fig. 38. Map of AMNH NJ-117 syninclusion. palpal segments seen: antepenultimate seg- ment swollen, penultimate and apical ones narrower, bacilliform. Length ratio 1:1.7:2.7. Saigusaia Vockeroth, 1980 Thorax: Antepronotum with two long setae. Scutum with strong, curved lateral, dorso- Saigusaia Vockeroth, 1980: 541. central and acrostichal setae. Mediotergite DIAGNOSIS: As given by Vockeroth (1980): and laterotergites bare. Scutellum with 5 Face weakly sclerotized, bare; laterotergites pairs of rather short setae. Metepisternum bare; metepisternum with short ®ne hairs; with two long setae dorsally. Wing: Costa prosternum bare; sternites of abdomen with ends beyond tip of R5, one-third distance be- median fold line. Male with tergite 7 and tween tips of R5 and M1. Stem of R, R1 and sternite 7 haired; subequal in length, two- R5 with rows of dorsal and ventral setae. R1 thirds length of segment 6; tergite 9 not fused and R5 straight. Sc meets C just before RS with gonocoxites. Female with sternite 8 base. Sc2 apical, proximal to base of M3. deeply emarginate posteriorly. Crossvein r-m 1.5ϫ length of RS1 and 0.3ϫ TYPE SPECIES: cincta Johannsen, length of M3 section. Fork of M1 and M2 1912: 270 (by original designation). 2.7ϫ length of M3. M1 and M2 fork campan- COMMENTS: Three species of the genus are ulate, veins straight, with a few ventral setae known from Eastern North America, Europe apically. Base of M3ϩ4 and CuA fork proxi- and Japan (Khonsu), and Taiwan and Nepal. mal to base of M1 and M2 fork, M3ϩ4 and

TABLE 3 Comparison of Boletina, Saigusaia, and Saigusaia pikei n.sp. 32 AMERICAN MUSEUM NOVITATES NO. 3433

CuA curved caudally. Legs: Tibial bristles diotergite setose. Wing membrane with mi- 2ϫ tibial diameter. Tarsal claw with one crotrichia and short macrotrichia. Costa ends tooth. Abdomen: Sternite 8 with two trian- beyond the tip of R5, one-fourth distance be- gular, rounded, caudal lobes covered by long tween tips of R5 and M1. Sc meets R just setae. Cerci two-segmented. Basal segment before base of RS. Sections of RS1 and RS2 large and broad, apical one 0.5ϫ the length equal and 1.5ϫ length of r-m. Crossvein r-m and 0.5ϫ the width of basal one; both setose. 0.5ϫ length of M3 section. Fork of M1 and MATERIAL: Holotype TMPD P79.15.7.21, M2 3.5ϫ length of its stem (M3). Base of female. Canada: Alberta, Grassy Lake, coll. fork of M3ϩ4 and CuA proximal to the base T. Pike. of M2 section. Legs: Inner surface of mid ETYMOLOGY: The species epithet is patron- tibia with ®ssurelike sensory pit in apical ymic for Dr. T. Pike, who collected the spec- third. Tarsal claws with two short teeth at the imen. base. Abdomen setose, with 7 visible seg- COMMENTS: This species has almost all the ments, 7th one very small. Cerci 2-segment- features of Saigusaia (table 3, see also Vock- ed, basal segment roundish, with numerous eroth, 1980), so despite the long Sc vein its long trichia; apical segment bacilliform, with placement in the genus is virtually certain. length twice the width, bare. Lateral lobes of 8th sternite setulose. Syntemna Winnertz, 1863 MATERIAL: Holotype TMPD P83.15.3.8, female. Canada: Alberta, Grassy Lake, coll. Syntemna Winnertz, 1863: 767. T. Pike DIAGNOSIS: Mediotergite bare; lateroter- ETYMOLOGY: The species epithet is the gites setose; wing membrane with macrotri- Latin word ®ssures, meaning ``full of chia; Sc meets R; radial cell very small, su- cracks'', in reference to the state of preser- vation of the specimen. bquadrate; base of M3ϩ4 and CuA fork prox- COMMENTS: The new species differs from imal to base of M1 and M2 fork; segment 7 of abdomen reduced. all other known Mesozoic species in having TYPE SPECIES: S. morosa Winnertz, 1863: section M3 long, at least twice the length of 767, by monotypy. crossvein r-m. COMMENTS: About 20 extant species are known from the Holarctic (Zaitzev, 1994), Gregikia, new genus with an additional 12 species described from Baltic amber (Meunier, 1904, 1917a, 1922) DIAGNOSIS: Costa virtually ends at the apex and 4 from the Lower and Upper Cretaceous of R5. Sc bare. Sc2 apical. Section M2 setose. of northeast Asia (Blagoderov, 1995, 1998a, R5 not reaching wing apex. M3 weak. Ac- 2000). rostichal setae absent. Tergite 9 short. Gon- ostyli well developed. Syntemna ®ssurata, new species TYPE SPECIES: Gregikia pallida, n.sp. Figures 39, 40, Plate 4C ETYMOLOGY: The name is derived from some letters commonly used in names of DIAGNOSIS: Sc ends at R at the base of RS; genera in this complex. The name is femi-

R4 present; crossvein r-m 0.5ϫ length of M3 nine. section; base of fork of M3ϩ4 and CuA prox- COMMENTS: The genus is very similar to imal to the base of M1 and M2 fork. Palaecomoptera Blagoderov, 1997, ®ve spe- DESCRIPTION: Body length ϭ 3.06 mm, cies of which were described from the Lower wing length ϭ 2.28 mm. Head not fully vis- Cretaceous of Transbaikalia and Mongolia ible. Flagellum 14-segmented, ¯agellomeres (Blagoderov, 1997, 1998a), as well as to barrel-shaped, length of each 1.5ϫ the width, three monotypic genera: Grzegorzekia Ed- setose, length of setae about equal to ¯agel- wards, Creagdhuhia Chandler, and Phoeni- lomere width. Thorax: Scutum with strong, kiella Chandler. It is distinguished from Pa- long setae in lateral, dorsocentral, and acros- laecomoptera by R5 not reaching wing apex, tichal rows, with short setulae. Anteprono- costa not produced after R5 apex, and base tum, proepisternum, laterotergites, and me- of fork of M1 and M2 weak. The genus dif- 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 33

Figs. 39±43. Syntemna and Gregikia. 39. S. ®ssurata, 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 AMERICAN MUSEUM NOVITATES NO. 3433 fers from Grzegorzekia, Creagdhuhia, and solateral setae. Mid tibiae without sensory Phoenikiella by a short apical palpomere, ab- pit. Tarsal claw with one basal tooth. Ab- sence of acrostichal setae, weak M stem and domen densely setose. Segments 7 and 8 base of fork of M1 and M2, absence of setae short, about one-half length of segment 6. on A, from Grzegorzekia also by having Sc Tergite 9 short, does not cover gonocoxites. bare. Gonocoxites of the new genus also Gonocoxites densely setose, with long, slim have two lobes as in the last three genera, apical and basal lobes, pointed caudally and but the gonostyli are better developed. This bearing numerous inner setae. Gonostyli state is undoubtedly more plesiomorphic than ovate, bare, with strong dark apical process- the highly modi®ed genital complex of Grze- es. gorzekia, Creagdhuhia and Phoenikiella. MATERIAL: Holotype AMNH NJ 117j, Species of Palaecomoptera have gonostyli male. Paratypes: AMNH NJ 117i, male (see developed as well, but some (P. shcherba- ®g. 74 for the scheme of syninclusions), coll. kovi, P. lukashevichae) have tergite 9 large, P. Nascimbene; AMNH NJ 871a and NJ fully covering the gonocoxites. 871b, sex unknown, coll. K. Luzzi. USA: New Jersey, Sayreville. Gregikia pallida, new species ETYMOLOGY: The species epithet is a Latin Figures 41±43, Plate 4D word pallidus meaning ``pale'' in reference to state of preservation of the specimen. DIAGNOSIS: As for genus. DESCRIPTION: Body length ϭ 3.77 mm (ho- Gaalomyia, new genus lotype)/3.09±3.65 mm (paratypes); wing DIAGNOSIS: Palpi 3-segmented, short. Scu- length ϭ 3.26 mm (holotype)/2.36±3.76 mm tum with long lateral, dorsocentral, and ac- (paratypes). Head: Vertex setose. Scape and rostichal setae. Laterotergites bare. Sc ends pedicel small. Flagellum 14-segmented. Fla- at R. Base of fork of M and CuA at the gellomeres cylindrical, length 2ϫ the width, 3ϩ4 level of r-m base. M3 section approximately covered by trichia, length of trichia about equal to r-m. Gonostyli simple. one-half ¯agellomere width. Only three seg- TYPE SPECIES: Gaalomyia carolinae, n.sp. ments of palpi visible; segments short, nearly ETYMOLOGY: The name is a feminine an- cylindrical, combined length slightly less agram of the genus name Aglaomyia. then head height. Thorax: Scutum with erect COMMENTS: The new genus differs form setae, arranged in rows with wide bare strips Aglaomyia Vockeroth in having acrostichal between them, acrostichal setae absent. An- setae present, palpi short, Sc ending at R ba- tepronotum and proepisternum setose. Pro- sally of posterior fork, and 7th abdominal epimeron touches mesepisternum at anepis- segment well developed. ternal suture. Scutellum with four pairs of Meunier has palpi 4-segmented, vein Sc setae. Anepimeron very narrow ventrally, so shorter, and the base of fork M and CuA that laterotergite touches katepisternum. Ka- 3ϩ4 more proximal. Pseudalysiinia Tonnoir has tepisternum larger than anepisternum. Me- palpi 4-segmented and incrassate, with no diotergite, laterotergites, and metepisternum bristles on the thorax and legs, and tibial bare. Wing: Costa ends slightly beyond tip spurs short. The genus may be close to Ip- of R . Sc meets C at level of RS base. R 5 saneusidalys Blagoderov, 1998, especially I. stem, R ,R, and distal parts of M and M 1 5 1 2 longipennis, but the latter differs in having with setae. R not reaching wing apex. Sec- 5 long fork of M and CuA and M3 section, tion RS1 oblique. R transverse. Ratios of 3ϩ4 4 and R is sinuous. Preservation of I. longi- lengths of RS1, RS2, and RS3 is 1:1.5:12± 5 pennis seems to be insuf®cient to decide if 15. Length of crossvein r-m about equal to both species are congeneric. RS1 and 0.25ϫ M3 section. M1 subparallel to R ; veins M ,M,M , and CuA diver- 5 1 2 3ϩ4 Gaalomyia carolinae, new species gent. Base of fork of M and CuA proximal 3ϩ4 Figure 44, Plate 4E to M3 base. M3ϩ4 and CuA curved gently. M1,M2,M3ϩ4, and CuA with a few setae on DIAGNOSIS: As for genus. apical part. Legs: Hind coxae with long dor- DESCRIPTION: Body length ϭ 2.68 mm; 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 35

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

wing length ϭ 2.34 mm. Head: Occiput and R1 and R5 straight. Crossvein r-m equal to frons densely setose. Eyes large, with round M3 section. Length of fork of M1 and M2 4ϫ facets, slightly emarginate, setulose. Ocelli M3 section. M1 and M2 not reaching wing absent. Scape and pedicel rounded, ¯agellum margin. Legs: Fore and mid coxae densely 14-segmented, ®liform. Length of 1st and setose, hind coxae with long posterolateral apical ¯agellomeres 2.5ϫ width, with others setae. Femora with ventral row of setae. Tib- 2ϫ width. Palpi 3-segmented, palpomeres ial spurs 2±2.3ϫ tibial diameter. Abdomen equal in length and subsequently narrower to densely setose. Sternite I bare. Eighth tergite apex. Length of penultimate one 1.5ϫ width, short, length 0.5ϫ 7th tergite. Tergite IX re- apical one 3ϫ width. Clypeus setose. Tho- niform, width 3ϫ length. Gonocoxites slen- rax: Scutum with long lateral, dorsocentral, der, length 3.5ϫ width, widely separated. and acrostichal setae. Proepimeron touches Gonostyli saberlike, curved inside. katepisternum at shallow incision. Katepis- MATERIAL: Holotype AMNH Bu-390, ternum larger than anepisternum. Anepister- male. Myanmar: Katchin, from amber mines nal suture horizontal. Antepronotum and near Myitkyina. proepisternum with long setae, other thoracic ETYMOLOGY: The species epithet is a pa- sclerites bare. Wing membrane without mi- tronym honoring our friend and colleague crotrichia. R, R1,R5, and M1 with dorsal se- Caroline S. Chaboo, specialist on chryso- tae. Sc ends at R at the level of base of r-m. melid beetles. 36 AMERICAN MUSEUM NOVITATES NO. 3433

TRIBE LEIINI EDWARDS, 1925 er setae. Proepimeron very small, touches ka- Nedocosia, new genus tepisternum at incision. Laterotergites and mediotergite bare. Metepisternum broader DIAGNOSIS: Three ocelli, lateral ones do than high. Wing: Sc meets R before M3 not contact eye margin. Sc merges with R. base. Costa ends beyond tip of R5, more than R1 2±3ϫ length of r-m. Veins of median and one-half length between tips of R5 and M1. cubital forks lightly sclerotized. Base of M3ϩ4 Crossvein r-m 0.37ϫ length of R , and 0.6ϫ and CuA fork lies between levels of base of 1 length of M3 section. Base of fork of M3ϩ4 r-m and M1±M2 fork. Tibial bristles absent. and CuA between levels of bases of M3 sec- TYPE SPECIES: Nedocosia exsangius, n.sp. tion and M1 and M2 fork. M3ϩ4 very weak. ETYMOLOGY: The genus name is derived M1 and M2 fork campanulate. Longitudinal from the pre®x ne-, ``negation'', and the ge- veins with setae, except Sc. Legs: Fore coxae nus . The name is feminine. with anterior setae, mid coxa with apical se- COMMENTS: This genus is most similar to tae. Fore leg with ®rst tarsomere 2ϫ longer Docosia Winnertz, 1863, which consists of than second and 0.55ϫ length of tibia. Ab- 25 extant Palaearctic, 15 Nearctic, and 2 domen: Tergites setose, sternites bare. Cerci Neotropical species (Zaitzev, 1994). Seven with two comblike rows of black bristles. species of Docosia are known from Baltic Apices of gonostyli sharp, squarish, curved amber (Meunier, 1904, 1916, 1922, 1923) inward. and one from the Oligocene shales of Rott, MATERIAL: Holotype PIN 3130/193, male. Germany (Statz, 1944). The new genus dif- Russia: Taimyr Peninsula, Yantardakh, coll. fers by having lateral ocelli not touching the 1970. eye margins (undoubtedly plesiomorphic), ETYMOLOGY: Latin, meaning pale or and by the lack of strong tibial bristles. Dif- bloodless, referring to the pale color of the ferences in sclerotization of anterior veins specimen. (Sc, R, R5) and posterior veins (forks) in the new genus are not as great as in Docosia species. Two species of Docosia described Nedocosia sibirica, new species from the Lower Cretaceous of Transbaikalia Plate 5A (Blagoderov, 1995) may also belong to the DIAGNOSIS: Sc meets R at the level of the new genus, in fact showing af®nities to N. base of the stem of M1ϩ2. Laterotergites se- novacaesarea. tose. Anterior edge of fore coxa bare. Hind legs stout. Nedocosia exsanguis, new species DESCRIPTION: Body length ϭ 2.37 mm, Figures 45, 46, Plate 4F wing length ϭ 1.82 mm. Head: Occiput and DIAGNOSIS: Sc meets R before level of frons setose. Ocelli three, equal in size; dis- base of r-m. Laterotergites bare. Cerci with tance of median to lateral ocellus and from two combs of dark setae. Apices of gonostyli lateral ocellus to eye margin equal to twice sharp, curved inward at right angle. their diameter. Eye with slight incision at an- DESCRIPTION: Body length ϭ 1.29 mm; tennal bases. Antennae short, only 10 ¯agel- wing length ϭ 0.67 mm (rest). Head oval, lomeres preserved; ¯agellomeres cylindrical, with rounded occiput, occiput and frons with widths 1.1±1.5ϫ lengths. Face triangular, se- scattered setae. Ocelli three, lateral one about tose, clypeus ovate, bare. Palpi 5-segmented, its own diameter from eye margin and two short, antepenultimate segment swollen, with diameters from medial ocellus. Eyes slightly medial sensory pit on inner surface. Apical emarginate, with round facets and short in- palpomere shorter than penultimate one, both terfacetal trichia. Scape subconical, pedicel slender, with sparse setulae. Length ratio 1: cylindrical. Flagellum 14-segmented, ¯agel- 3:3:5:5. Thorax: Mesonotum with short tri- lomeres of hexagonal surface texture, cylin- chia and setae of various sizes; setae ar- drical, lengths about equal to width. Face and ranged in 5 rows. Antepronotum with three clypeus setose. Thorax: Scutum with setae setae; proepisternum with two on lower part. in lateral, dorsocentral, and acrostichal rows. Scutellum with two very long setae and two Antepronotum with 4 long and several short- smaller ones. Laterotergites with a few tri- 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 37

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 AMERICAN MUSEUM NOVITATES NO. 3433 chia in lower part. Mediotergite bare. Wing: Nedocosia novacaesarea, new species Sc meets R at the level of the base of the Figure 47, Plate 5C stem of M1ϩ2.R1 twice the length of r-m. R5 curved slightly back. C ends far beyond tip DIAGNOSIS: Flagellum 11-segmented. Sc of R5, but does not reach wing apex. Stem long, ends at R between RS and r-m bases. of R, R1, and R5 with setae. Legs: Anterior Base of M3ϩ4 and CuA fork at level of r-m. edge of fore coxae without setae. Fore tibiae Fore leg with tibia equal to femur or slightly with semicircular anteroapical depressed longer. area, covered with numerous small trichia. DESCRIPTION: Body length ϭ 2.34 mm (ho- Mid tibial spur length 2.5ϫ apical diameter lotype)/2.88 mm (paratype); wing length ϭ of tibia, hind tibial spur 1.5ϫ diameter. Hind 1.75 mm (holotype)/2.20 mm (paratype). legs stout, longer than mid legs. First fore Head: Three ocelli in line, distances between tarsomere 2.5ϫ longer than second and median and lateral ones, and between lateral 0.67ϫ length of tibia. Cerci simple. and eye margin, are equal. Face rectangular, MATERIAL: Holotype PIN 3311/665, fe- bare. Clypeus bare. Scape and pedicel round- male. Russia: Taimyr Peninsula, Yantardakh, ed, shorter than ¯agellomeres. Flagellum 11- coll. 1971. segmented, ¯agellomeres cylindrical, with ETYMOLOGY: The species epithet is a ref- lengths 2ϫ width. Two palpomeres seen, api- erence to the region of origin of the amber. cal one 2ϫ length of basal. Thoracic sclerites bare. Scutum with long lateral setae and nu- Nedocosia canadensis, new species merous short ones, not arranged in rows. Plate 5B Proepimeron small, ®ts in shallow incision on katepisternum. Anepisternal suture declines DIAGNOSIS: Sc meets R before level of backward. Metepisternum touches anepime- ron. Meron large. Wing: Costa ends beyond base of the stem of M1ϩ2.R1 2.5ϫ length of tip of R , more than one-third length between r-m. M3ϩ4 and CuA fork at the level of r-m. 5 M3 section shorter than r-m crossvein. La- tips of R5 and M1. Sc long, ends at R between terotergites and mediotergite bare. RS and r-m bases. Length of R1 2.5±3ϫ that DESCRIPTION: Body length ϭ 1.50 mm of r-m, M3 section twice that of r-m. Length (preserved part). Head: Occiput with setae of fork of M1 and M2 2.5±3ϫ M3 section. of different length. Ocelli not seen. Eyes with Base of fork of M3ϩ4 and CuA at level of r- round facets, setulose. Scape small, pedicel m base. Legs: Fore coxae with short anterior subconical. Flagellum 14-segmented, ¯agel- setae, hind coxae with long posterolateral se- lomeres cylindrical, lengths equal to widths, tae. Fore leg with tibia equal to femur. Tibial densely covered by trichia no longer than spurs 1.2±1.5ϫ tibial diameter. Abomen se- half width of basal ¯agellomere. Three seg- tose. Seventh segment very narrow. Gonocox- ments of palpi seen, combined length equal ites massive, with apices blunt. to head height. Thorax: Scutum uniformly MATERIAL: Holotype AMNH NJ-117k, setose with long and short setae. Antepron- male; paratype NJ-117l, male in the same otum with two very long setae and several piece. USA: New Jersey, Sayreville, coll. P. short ones. Laterotergites and mediotergite Nascimbene. bare. Wing: Sc meets R before level of base ETYMOLOGY: The species epithet is derived of the stem of M1ϩ2.R1 2.5ϫ length of r-m. from Latin Nova Caesarea meaning ``New Crossvein r-m 1.3ϫ length of M3 section. Jersey'' and is a reference to the state of or-

Base of fork of M3ϩ4±CuA proximal to base igin of the amber. of fork of M1±M2. COMMENTS: The species is similar in ve- MATERIAL: Holotype MCZC 6897, incom- nation to Docosia baisae Blagoderov, 1998 plete specimen, sex unknown. Canada: Man- and D. zaza Blagoderov, 1998, by proximal itoba, Cedar Lake, coll. F. M. Carpenter. position of the forked base of M3ϩ4 and CuA, ETYMOLOGY: The species epithet is a ref- but differs from the latter species in having erence to the country of origin of the amber. Sc long and M3 section short. 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 39

Ectrepesthoneura Enderlein, 1911 ly curved forward and shorter ones curved Willistoniella Meunier, 1904: 74 (preoccupied by back, arranged in rows. Antepronotum with Mik, 1895). 3 long setae. Scutellum with two pairs of Meunieria Johannsen, 1909: 87 (preoccupied by long setae. Other thoracic sclerites bare. Kieffer, 1904). Wing: All longitudinal veins with short se- Ectrepesthoneura Enderlein, 1911: 115. tae. Sc meets R proximal to base of M3. Length ratio of sections RS1, RS2, and RS3 DIAGNOSIS: As given by Chandler (1980): is 1:2:13. RS1 transverse, 0.3 length of Lateral ocelli remote from eye margin; later- ϫ r-m. Crossvein r-m 0.7 length of R and otergites bare; Sc ending in R before base of ϫ 1 0.5 length of M3 section. Costa ends be- RS; R present; R short, at most twice as ϫ 4 1 yond tip of R , one-third distance between long as r-m;R straight; C prolonged well 5 5 tips of R and M . Section M3 and M and beyond tip of R ; veins of medial and cubital 5 1 1 5 M bases very weak. Legs: Fore coxae with sectors weak and faint; posterior fork sessile. 2 numerous anteromedial setae, hind coxae Males with ``sensory pit'' near the base of with posterolateral setae. Fore and mid tibiae mid tibia; hind tibial comb absent. slightly shorter than fore femora. Mid and TYPE SPECIES: hirta Win- hind tibiae and tarsi with rows of black bris- nertz, 1846: 19 (orig. designation) tles. Tibial spurs 1.5±2ϫ tibial diameter. Tar- COMMENTS: There are about 10 extant spe- sal claw with one long tooth. Abdomen se- cies from the Holarctic region. The genus is tose. Eighth sternite almost triangular. Cerci very close to Tetragoneura Winnertz (about 2-segmented, basal segment stick-shaped, 100 widespread living species) and is some- with length twice the width, extends beyond times included in it or treated within the tribe 8th sternite; apical one discoidal, attached at Gnoristini (Tuomikoski, 1966; Vaisanen, the apex of the basal one, 2.2 shorter. Dor- 1986). There are two fossil species described ϫ sal border of 8th sternite (gonocoxite 8 of from Baltic amber (Meunier, 1904) and the Martinsen and SoÈli, 2000) straight. Oligocene of Rott, Germany (Statz, 1944). MATERIAL: Holotype PIN 3311/662, male. Chandler (1999) suggested that Ectrepestho- Russia: Taimyr Peninsula, Yantardakh, coll. neura was paraphyletic. 1971. ETYMOLOGY: From Latin succinum mean- Ectrepesthoneura succinimontana, ing ``amber'' and montanus meaning ``moun- new species tainous''. Yantardakh means ``amber moun- Figures 48, 49, Plate 5D tain'' in Dolgan language. DIAGNOSIS: Fore tibiae slightly shorter than COMMENTS: Comparision with recent spe- fore femur. Section M3 and bases of M1 and cies is dif®cult, because nongenital characters M2 very weak. Length of small radial cell used for distinguishing species have been con- twice the width. sidered to be unreliable for this genus (Mar- DESCRIPTION: Body length ϭ 2.39 mm; tinsen and SoÈli, 2000). For species where the wing length ϭ 2.02 mm. Head: Occiput and female genitalia are known the new species is frons with short curved setae. Ocelli three, most similar to E. hirta Winnertz. distance of lateral one to eye margin more than twice the ocellus diameter. Scape sub- Ectrepesthoneura swolenskyi, new species conical, pedicel rounded, width twice ¯agel- Figures 50, 51, Plate 5E lomere width. Eleven segments of ¯agellum preserved, ¯agellomeres cylindrical, with DIAGNOSIS: Sc long, meets R just before length about equal to width. Face setose. Pal- base of M3. Small radial cell relatively long. pi 4-segmented; 1st palpomere very small; Gonocoxites with triangular ventral append- antepenultimate one oval, its length twice the ages bearing 2 long, dark apical spurs. width, with medial sensory pit; penultimate DESCRIPTION: Body length ϭ 2.71 mm; one with length 3ϫ width, attached to the wing length ϭ 2.46 mm. Head: Ocelli three, second one preapically; apical segment long from lateral ocellus to eye margin twice ocel- and slender. Clypeus triangular, setose. Tho- lus diameter. Flagellomeres cylindrical, rax: Scutum with numerous long setae most- length 1.5ϫ width. Three palpomeres seen, 40 AMERICAN MUSEUM NOVITATES NO. 3433

Figs. 48±51. Ectrepesthoneura End. 48. E. succinimontana, n.sp., holotype PIN 3311/662. 49. Fe- male genitalia. 50. E. swolenskyi, n.sp., holotype AMNH NJ-824. 51. Male genitalia. 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 41

apical one slender, length ratio 1:2:3.5. Tho- nus vein M1 is entirely reduced. As a rule, in rax: Antepronotum with 2 setae, proepister- most leiine genera, vein M1 reaches the wing num with 4. Scutum with setae curved for- margin at or even before the wing apex. They ward, lateral setae very long. Wing: All lon- have retained a complete vein that is homol- gitudinal veins except Sc with short setae. ogous to CuA based on the length and shape

Costa ends beyond tip of R5, one-half dis- of the vein, and which is strongly curved tance between tips of R5 and M1. Sc meets back and ends at the level of RS1, as in most R just before M3 base. Length of R1 2.2ϫ Leiini. The new genus differs from Neoclas- that of r-m. Length of M3 2.7ϫ that of r-m. tobasis Ostroverchova by the lateral ocelli Ratios of lengths of sections RS1, RS2, and touching eye margins, and from Clastobasis RS3 1:2.3:7. RS1 oblique, 0.58ϫ length of Skuse and Neoclastobasis by C ending be- r-m. M3, M1, and M2 weak, M1 slightly yond the tip of R5, and by the presence of curved forward. M3ϩ4 and CuA curved gent- the base of M3ϩ4. Having R1 relatively long ly caudad. Legs: Fore and mid tibiae longer and Sc ending at C makes the genus similar than femora. Mid tibia with anterolateral, to Johannsen and Indoleia Ed- posterolateral, and posteromedial rows of wards, but differs from them by reduction of short bristles, hind tibia with two rows of the veins in the medial sector. Such a reduc- posterior setae. Basal third of mid tibia with tion also occurs in other leiine genera, for large, dark, narrow sensory pit. Tibial spurs example Strobl and Sigmoleia Ton- 1.3±1.7ϫ tibial diameter. Abdomen. Ninth noir and Edwards, but they have Sc very tergite large, conceals gonocoxites and gon- short and free. Species of Cycloneura Mar- ostyli almost completely; caudal edge shall and Paracycloneura Tonnoir and Ed- straight, without medial cleft. Ninth tergite wards also have M unbranched and M3ϩ4 free straight caudally, without cleft, with numer- or reduced at the base, the anepisternum ous curved setae apically. Gonocoxites with large, metepisternum narrow like Izleiina, triangular ventral appendages bearing 2 long, but they differ strongly in the shape of CuA, dark apical spurs. Gonostyli short, pointed, by having M2 long, Sc short and free, and by curved S-like. the presence of strong tibial bristles. MATERIAL: Holotype AMNH NJ 824, male. USA: New Jersey, Sayreville; coll. S. Izleiina miri®ca, new species Swolensky, 1997±1998 (mentioned in Gri- Figures 52±54, Plate 5F maldi, 2000: ®g. 48h). ETYMOLOGY: The species epithet is patron- DIAGNOSIS: Apical palpomere short. Tibiae ymic for the late Steve Swolensky, who col- without bristles. Gonostyli conical, curved lected the specimen. inward. COMMENTS: The new species differs from DESCRIPTION: Body length ϭ 1.44 mm, other species of the genus in the structure of wing length ϭ 1.23 mm. Head: Ocelli three, male genitalia, especially by ventral append- distance from eye margin equal to two ocel- ages of gonocoxites bearing dark sclerotized lus diameters; distance between each other spurs. equal to one diameter. Flagellum 14-seg- mented, ¯agellomeres cylindrical, lengths Izleiina, new genus about equal to widths, covered by setulae no longer than one-half ¯agellomere width. Pal- DIAGNOSIS: Facets round. Lateral ocelli not pi 4-segmented, three basal segments bacil- touching eye margins. Clypeus setose. Sc liform, apical one short and rounded. Occi- meets C just beyond level of base of M3; Sc2 put, frons, face, and clypeus with short se- absent. Length of R1 2ϫ that of r-m. Section tulae. Thorax: Prescutum and anterior par-

M3, and fork of M1 and M2±M3ϩ4 reduced; apsidal suture distinct. Antepronotum with 6 CuA weakened. M3ϩ4 interrupted at base. setae, proepisternum with two. Scutum TYPE SPECIES: Izleiina miri®ca, n.sp. arched. Scutellar setae in rows. Scutellum ETYMOLOGY: Name is derived from the with two pairs of long setae. Cavity on an- tribe Leiini. terior edge of katepisternum shallow. Later- COMMENTS: We assume that in the new ge- otergites and mediotergite bare. Wing: Sc 42 AMERICAN MUSEUM NOVITATES NO. 3433

Figs. 52±57. Izleiina, n.gen. 52. I. miri®ca, n.sp., holotype PIN 3130/187. 53. Reconstruction of wing venation. 54. Male genitalia. 55. I. spinitibialis, n.sp., holotype AMNH NJ-346, reconstruction of wing venation. 56. Head. 57. Male genitalia. 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 43 meets C just beyond level of base of M3. C width. Gonocoxites slightly swollen. Gon- ends far beyond apex of R5.R1,R5, and r-m ostyli almost straight, length 2.5ϫ width, with setae. R1 length 2ϫ that of r-m. R5 with 3 dark teeth at the apex. straight. Length of crossvein r-m 0.5ϫ length MATERIAL: Holotype AMNH NJ-346, of section M3. M3 weak, can be seen only male. USA: New Jersey, Sayreville, coll. Y. by absence of microtrichia on wing mem- Goldman, 1995. brane. M3ϩ4 and CuA fork base proximal to ETYMOLOGY: The species epithet is derived the base of r-m. M3ϩ4 interrupted at base, not from Latin spina, meaning ``spine'', and tib- reaching wing margin. Legs: Fore coxae ia, meaning ``leg''. with anterior setae, mid coxae with lateroap- ical ones. Tibiae without bristles. Genitalia: Zeliinia, new genus Gonocoxites massive, fused. Gonostyli horn- DIAGNOSIS: Close to Izleiina n.gen., but like and slightly curved, with long setae, col- differs in having palpi long, Sc free, M re- ored lighter than gonocoxites. 1 duced at base, M absent, and base of pos- MATERIAL: Holotype PIN 3130/187, male. 2 terior fork proximally with r-m touching the Russia: Taimyr Peninsula, Yantardakh, coll. base of M . Setulae on ¯agellomeres ar- 1970. 3ϩ4 ranged in rings. ETYMOLOGY: The species epithet is a Latin TYPE SPECIES: Zeliinia orientalis, n.sp. word miri®cus meaning ``amazing'' in ref- ETYMOLOGY: The genus name is an ana- erence to unusual combination of characters. gram for Izleiina. The name is feminine. The name is feminine. Zeliinia orientalis, new species Izleiina spinitibialis, new species Figures 58±61, Plate 6B Figures 55±57, Plate 6A DIAGNOSIS: Genitalia longer than wide; DIAGNOSIS: Tibiae with bristles. Gonostyli gonostyli bilobate. Tergite 9 obovate, with cylindrical, length 2.5ϫ the width, with 3 numerous setae at apex. dark teeth at the apex DESCRIPTION: Male. Body length ϭ 1.25 DESCRIPTION: Body length ϭ 2.07 mm; mm; wing length ϭ 1.07 mm. Head: Ocelli wing length ϭ 1.65 mm. Head: Ocelli three, three, close to each other. Eyes strongly in- almost in a straight line. Lateral ocelli distant cised, forming incomplete bridge. Pedicel by two diameters from medial ocellus and slightly wider than scape and ¯agellum; 14 their own diameter from eye margin. Frontal cylindrical ¯agellomeres with length 2ϫ furrow distinct. Eyes emarginate at antennal width. Three palpomeres seen, sticklike, base, bare. Frons bare. Face setose. Scape length ratio 1:1.3:2. Thorax: Scutum with and pedicel without strong setae, rounded, long lateral, dorsocentral, and acrostichal se- wider than ¯agellomeres. Eight ¯agellomeres tae. Katepisternum larger than anepisternum, seen, cylindrical in shape, lengths 1.5ϫ proepimeron touches katpisternum at small width. Clypeus rounded, setulose. Palpi 4- excavation. Antepronotum and proepimeron segmented, basal segment rounded, antepen- setose. Laterotergites and mediotergite bare. ultimate and penultimate one bacilliform Wing: Costa slightly produced beyond apex with lengths 3±4ϫ width; apical palpomere of R5. Sc free, ends at the middle of r-m. R1 long. Thorax: Scutellum with 4 long setae. and R5 with long dorsal and ventral setae. Laterotergites and mediotergite bare. Wing: Crossvein r-m forming one vein with tb and

Sc long, meets C at level of M3 base. R1 M2 section, equal to 1.5ϫ length of R1. straight, 2ϫ length of r-m. R, R1,R5, and r-m Length of R5 2.8ϫ R1. Base of M3 faint. M1 with a dorsal row of setae. Crossvein r-m ends at wing apex, base curved posteriorly. almost horizontal. R5 curved caudally. C Base of M3ϩ4 and CuA fork contact r-m. ends beyond tip of R5.M2 and M3ϩ4 weak. Legs: Tibial setulae irregular. Tibia II with Base of CuA weak. CuA curved caudally, apical comb of setae. Tibial spurs 1±1.5ϫ not sigmoid. Legs: Tibiae with bristles. Ab- tibial diameter, spur formula 1:1:1. Abdo- domen: Genitalia covered by long setae. men setose, 7th and 8th segments short, nar- Ninth tergite rectangular, length 2.3ϫ the row, retractable. Genital complex large, 44 AMERICAN MUSEUM NOVITATES NO. 3433

Figs. 58±63. Zeliinia, n.gen. 58. Z. orientalis, n.sp., holotype AMNH Bu-315. 59. Wing. 60. Male genitalia laterally. 61. Male genitalia dorsally. 62. Z. occidentalis, n.sp., holotype MCZC 6943. 63. Male genitalia. 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 45 length equal to 4th±8th segments combined. spurs 1.2±1.5ϫ tibial diameter. Tarsal claws Ninth tergite ®ddle-shaped, with two apical with very small tooth. Abdomen densely se- combs of dark setae. Gonocoxites rectangu- tose, with long setae. Genitalia rectangular, lar in lateral view. Gonostyli bilobate. width 1.6ϫ length. Gonocoxites stout, gon- MATERIAL: Holotype AMNH Bu-315, ostyli simple, pointed, with inner surface male. Myanmar: Katchin, from amber mines slightly curved apically. near Myitkyina. MATERIAL: Holotype MCZC 6943, male. ETYMOLOGY: The species epithet is a Latin Canada: Manitoba, Cedar Lake, coll. F.M. word orientalis, meaning ``eastern'', refer- Carpenter. ring to origin of the amber. ETYMOLOGY: The species epithet is Latin meaning ``western'', referring to origin of the Zeliinia occidentalis, new species amber Figures 62, 63, Plate 6C COMMENTS: Z. occidentalis differs from the type species in the structure of genitalia, DIAGNOSIS: Clypeus with long setae; me- and although it possesses the same arrange- diotergite and laterotergites bare, wing mem- ment of ¯agellar setulae and venation pat- brane without macrotrichia, Sc short, ends tern, both species may not be congeneric. free, R1 short, 1.5ϫ longer than r-m, ends proximally at tip of CuA. Genital complex Temaleia, new genus wider than long; gonostyli unilobate. DIAGNOSIS: Wing membrane with macro- DESCRIPTION: Body length ϭ 1.79 mm; trichia; Sc long, meets C before base of M3 wing length ϭ 1.22 mm. Head: Eyes bare, section; C ends at tip of R ; M3 section 3ϫ forming dorsal bridge, possibly incomplete 5 length of r-m;M not interrupted at the (vertex and occiput not fully seen due to de- 3ϩ4 base; tibial setulae irregular; tibial bristles formation). Scape and pedicel short, round- absent; abdominal segment 6 not reduced; ed. Flagellum 14-segmented, ¯agellomeres 7th and 8th segments telescopic. equal in length, apical ¯agellomeres more TYPE SPECIES: Temaleia birmitica, n.sp. narrow than basal ones, with length about ETYMOLOGY: The genus name is a femi- twice the length of pedicel, covered with nine anagram of Metaleia. short microtrichia. First ¯agellomere length COMMENTS: The genera Clastobasis Skuse, 1.6ϫ width, preapical is 2.5ϫ width. Clypeus 1896, Neoclastobasis Ostroverchova, 1970, setose. Three visible palpal segments, two and Metaleia Baxter, 1994 have M inter- basal ones about equal in length; apical pal- 3ϩ4 rupted at the base and/or CuA sinuous. The pomere long and slender, about twice the genus Sticholeia SoÈli, 1996, besides having length of the penultimate. Thorax: Ante- extremely long cerci and other details of the pronotum with two setae, proepisternum with male terminalia, differs by having a short M3 one. Scutum with long lateral, dorsocentral, section. and acrostichal setae. Scutellum with 4 pairs of long setae. Katepisternum larger than ane- Temaleia birmitica, new species pisternum. Metepisternum small, quadrate, Figure 64, Plate 6D bare. Mediotergite and laterotergites bare. Wing membrane without macrotrichia. R, DIAGNOSIS: As for the genus.

R1, and R5 setulose. Sc short, ends free. R1 DESCRIPTION: Body length ϭ 2.17 mm (ho- 1.5 length of r-m, R5 more than 4.5ϫ length lotype)/2.03±2.38 mm (paratype); wing of r-m. Crossveins r-m, tb, and m-cu fused length ϭ 1.96 mm (holotype)/1.50±2.07 mm into one oblique vein, with distinct kink at (paratype). Head: Pedicel rounded, length the base of the stem of M1ϩ2 (M3 section). about equal to width. Flagellum 14-segment- Stem of M1ϩ2 and base of M3ϩ4 weak. M ap- ed; ¯agellomere length about twice the parently without fork. M3ϩ4 and CuA curved width. Eyes large, densely setose. Occiput gently caudad. Legs: Femora with ventral covered with numerous short setae. Clypeus row of long setae. Coxae relatively short, setose. Palpi 3-segmented, very short, pal- fore coxa as long as mid and hind ones. Mid pomeres with length about equal to width. and hind tibiae with short bristles. Tibial Labella longer than palpi. Thorax: Scutum 46 AMERICAN MUSEUM NOVITATES NO. 3433

Figs. 64±68. Temaleia and Lecadoniella. 64. T. birmitica, n.sp., holotype AMNH Bu-483a. 65. L. parvistyla, n.sp., holotype MCZC 6941. 66. Wing. 67. Male genitalia ventrally. 68. Male genitalia dorsally. 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 47 irregularly covered with short setae and with nus differs from Megophthalmidia Dzied- long lateral, dorsocentral, and acrostichal se- zicki in having M3ϩ4 interrupted at the base tae. Scutellum with three pairs of long bris- and the fork of M1 and M2 not shifted ante- tles. Laterotergites and mediotergite with riorly, so that M1 reaches the wing margin several protruding bristles. Wing membrane beyond the wing apex. with microtrichia and macrotrichia, located mostly in distal half of wing. Costa ends at Lecadonileia parvistyla, new species tip of R , not reaching wing apex. Sc long, 5 Figures 65±68, Plate 6E meets C before base of M3. Sc2 apical. R1 very short, 1.5ϫ length of r-m. M3 section DIAGNOSIS: As for genus. 3ϫ length of r-m. M1 meets wing edge at DESCRIPTION: Body length ϭ 2.24 mm, wing apex. M1 and M2 fork 2.5ϫ as long as wing length ϭ 2.05 mm. Head: Vertex se- M3 section. Base of M3ϩ4 and CuA fork at tose. Ocelli three, lateral one twice its di- level of M3 base. M3ϩ4 and CuA curved ameter from eye margin. Frons bare, face gently caudad. Legs: Fore coxae with ante- with tiny trichia. Scape small, roundish. Ped- rior and anterolateral setae. Mid coxa with icel rounded, with dorsal setae, wider than lateroapical setae, hind one with posteroapi- ¯agellomeres. Flagellum 14-segmented, cy- cal setae. Tibial spurs short, no longer than lindrical, ¯agellomere length equal to width. maximum tibial diameter. Tibial and tarsal Palpi 4-segmented. Basal palpomere very bristles absent. Tibial setulae irregular. Ab- short, apical three subequal, lengths 2.5±3ϫ domen pubescent, with 7 visible segments, their width. Clypeus setose. Thorax: Scutum 8th small and retracted. Ninth tergite large, irregularly setose. Antepronotum with 4 se- oval. Female: Cerci one-segmented, bacilli- tae, proepisternum with 5. Ventral process of form, setose. Male: Gonocoxites and gon- proepimeron just opposite anapleural suture. ostyli long, slender, bacilliform. Gonostyli Anepisternum with distinct oblique cleft. slightly curved inside at apex, saberlike. Anepimeron very narrow ventrally. Latero- MATERIAL: Holotype AMNH Bu-483a, fe- tergites setose. Mediotergite bare. Wing male; paratypes AMNH Bu-060, male, Bu- 054, male. Myanmar: Katchin, from amber membrane with macrotrichia. Costa ends be- mines near Myitkyina. yond tip of R5, at one-fourth length between tips of R and M .R and R with two rows ETYMOLOGY: The species epithet is a ref- 5 1 1 5 erence to Burma, the former name of the of setae; R stem, r-m, M1,M2,M3ϩ4, and CuA country where the amber originates. with one row. Sc ends free at the level of the base of M3 section. R1 equal to M3 section and 2.5ϫ length of r-m. Fork of M and M Lecadonileia, new genus 1 2 1.8ϫ length of M3. M3ϩ4 base absent, vein DIAGNOSIS: Palpi reduced, laterotergites begins at level of RS base. Legs: Fore coxae setose; wing membrane with macrotrichia; with numerous setae on anterior and lateral surfaces. Mid coxae with setae on apical half. Sc free, Sc2 absent; M3 4 interrupted at base, ϩ Hind coxae with posterolateral row of setae. M3ϩ4 and CuA curved gently caudad, 9th ter- gite small or fused with synsclerite; gono- Tibial bristles short. Fore tibiae with 3 apical coxites widely separated. bristles; mid one with 2 anterior and 6 apical TYPE SPECIES: Lecadonileia parvistyla, bristles; hind tibia with 4±6 bristles in ante- n.sp. rior, anteroventral, and anterodorsal rows and ETYMOLOGY: The genus name is a femi- several apical bristles. Anteroapical depres- nine anagram of the Caledonileia Matile, sion on fore tibiae well developed, with sin- 1993. gle row of short setae. Tarsal claws without COMMENTS: The genus is close to the ge- teeth. Abdomen setose. Eighth and 9th seg- nus Caledonileia, from which it differs in ments very short and retracted. Gonocoxites number and position of ocelli, number of pal- semicylindrical, widely separated at base. pomeres, setation of scutum, and in having Gonostyli small, with one apical appendage Sc long, the radial cell wide, M3 long, and turned medially. gonostyli unilobate (Matile, 1993). The ge- MATERIAL: Holotype MCZC 6941, male. 48 AMERICAN MUSEUM NOVITATES NO. 3433

Canada: Manitoba, Cedar Lake, coll. F.M. short setae in upper part and very ®ne hairs Carpenter. at lower apical part. Proepimeron touches ka- ETYMOLOGY: The species epithet is a com- tepisternum slightly ventral to anepisternal bination of the Latin word parvus, meaning suture. Mediotergite and laterotergites bare. ``small'', and stylus, meaning ``spike, stem, Metepisternum large, with dorsal margin pen'', in reference to simple small gonostyli wider than ventral one. Wing membrane of the species. with microtrichia and short macrotrichia.

Costa ends beyond tip of R5, at one-third Disparoleia, new genus length between tips of R5 and M1. Humeral vein located beyond the MA. Sc short, ends DIAGNOSIS: Ocelli absent; anepisternum at C before level of M3 base, apex faint. R with very short setae and trichia; medioter- 1 very short, r-m 3ϫ length of R . Median gite and laterotergites bare; wing membrane 5 veins weak. M , base of M and M3 very with micro- and macrotrichia; Sc ends at C; 1 1 weak. Base of M3 very proximal, slightly r-m 2ϫ length of R , M3 base at basal one- 1 distal to apex of Sc. Crossvein r-m 3ϫ length sixth of wing, M3 weak, M absent. 2 of M2 section, M3 section 3.5ϫ that of M2. TYPE SPECIES: Disparoleia cristata, n.sp. Base of M and CuA fork slightly distal to ETYMOLOGY: The genus name is a combi- 3ϩ4 level of base of M3 section. R, R ,R,M , nation of the Latin word dispar, meaning 1 5 3ϩ4 and CuA with setae dorsally and ventrally. ``unequal, unlike'', and the genus ,in Leg: Tibial spur formula 1:2:2. Spurs long, reference to the distinctiveness of the new 4±5ϫ tibia diameter. Mid and hind tibiae genus. The name is feminine. with bristles. Empodium absent; tarsal claws COMMENTS: The genus differs from all Re- with one tooth. Abdomen: Tergites of ab- cent Leiini in the unique combination of such domen setose. Seventh tergite 1.5ϫ shorter characters as absence of ocelli, very long r- than 6th, 8th is very short. Cerci 2-segment- m with the proximal position of the base of ed, slightly shorter than 6th and 7th segments M3, and reduction of veins in the medial sec- together. Apical segment round, basal one tor. It differs from Novakia StroÈbl in having bacilliform, 2.5ϫ apical. the base of RS distinct, M absent, and Sc 2 MATERIAL: Holotype AMNH B-0125, fe- ending at C. It differs from Zeliinia, n. gen. male. Myanmar: Katchin, from amber mines in having ¯agellomeres of smooth texture, near Myitkyina. ocelli absent, vein M and M3 section present 1 ETYMOLOGY: The species epithet is Latin though weak, and fork of M and CuA long 3ϩ4 word cristatus, meaning ``tufted, plumed'', in but not sessile. reference to the group of setae on the vertex. Disparoleia cristata, new species Figures 69, Plate 7A Hemolia, new genus

DIAGNOSIS: As for genus. DIAGNOSIS: Three ocelli in straight line, DESCRIPTION: Female. Body length ϭ 2.73 middle one small, lateral ones not touching mm, wing length ϭ 2.08 mm. Head: Vertex eye margin; wing membrane without macro- with group of long erect setae. Scape and trichia; Sc short, ends at R; R1 short, no more pedicel wider than ¯agellum. Flagellum 14- than 2ϫ length of r-m; r-m short, oblique; segmented, ¯agellate, ¯agellomeres longer M3 short; tibial setae in rows, mid and hind and narrower apicad. Eyes setose, slightly tibiae with inner spurs longer than outer one. emarginate at antennal base. Ocelli absent. TYPE SPECIES: Hemolia matilei, n.sp. Clypeus setose. Palpi 3-segmented, 1st pal- ETYMOLOGY: The genus name is a femi- pomere short, 2nd longer, cut obliquely, 3rd nine anagram of Mohelia Matile, 1978. longer than length of 1st and 2nd, narrow. COMMENTS: The genus is close to Mohelia Thorax: Scutum with lateral and dorsocen- in the structure of the ocelli, and wing ve- tral rows of long setae and short irregular nation, and it differs in tibial spur length and setae. Scutellum with numerous short setae. details of male genitalia structure. It differs Antepronotum with long curved setae and from Mohelia in having setae on the medi- several short ones. Anepisternum with 4 light otergite, tibial setae in rows, mid and hind 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 49

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 AMERICAN MUSEUM NOVITATES NO. 3433 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 51 tibiae with inner spurs longer rather than the ed, ¯at; apical segment 0.24ϫ basal one, outer ones, and C ending close to the tip of semicircular.

R5; Sc is longer, R1 longer than r-m; and M3 MATERIAL: Holotype AMNH B-0132, is short. male; paratype AMNH B-0133a, female. Myanmar: Katchin, from amber mines near Hemolia matilei, new species Myitkyina. Figures 70, 71, Plate 7B ETYMOLOGY: The species epithet is a pa- tronym in honor of the late Professor LoõÈc DIAGNOSIS: Scutum densely setose; later- otergites setose; Sc rather long, ends at the Matile, a world authority on Sciaroidea, who level of M and CuA fork base. was also very generous and helpful to both 3ϩ4 authors. DESCRIPTION: Body length ϭ 2.19 mm (ho- lotype)/2.39 mm (paratype); wing length 1.49 mm (holotype)/2.21 mm (paratype). Hemolia glabra, new species Head: Eyes setose. Occiput and frons dense- Figure 72, Plate 7C ly setose. Three ocelli in straight line, middle one small; lateral one separated from eye DIAGNOSIS: Laterotergites bare; scutum al- margin by its own diameter. Flagellum 14- most bare, dark and shining, Sc meets R well segmented, ¯agellomeres cylindrical; length before the level of base of M3ϩ4 and CuA of ¯agellomeres about equal to their width. fork. Clypeus setose. Palpi 4-segmented, basal DESCRIPTION: Body length ϭ 3.16 mm; segment small, penultimate and apical ones wing length ϭ 2.05 mm. Head: Occiput with situated preapically, antepenultimate with short setae. Three ocelli in line, of the same round sensory pit on inner surface. Thorax: size, distant from each other and from eye Scutum with long setae not arranged in dis- margin by two ocellus diameters. Eyes large, tinct rows. Laterotergites with long setae. setose. Pedicel with round sensory organ. Scutellum protruding, with 6 pairs of long Flagellum 14-segmented, apical segment setae. Antepronotun and proepisternum with conical, secondarily divided. Thorax: Scu- long setae. Wing rather wide, membrane tum almost bare; dark, shiny. Anterior par- without macrotrichia. Costa ends beyond tip apsidal suture distinct. Scutellum and later- otergites bare. Mediotergite with a few ®ne of R5 at one-®fth length between tips of R5 setae anteroventrally. Mid and hind tibiae and M1. Sc ends in R at the level of M3ϩ4 with inner spurs 1.5ϫ longer than outer ones. and CuA fork base. R, R1, and R5 with setae Wing membrane without macrotrichia. Hu- dorsally. R1 length 1.8ϫ length of r-m. Crossvein r-m about the length of M3, with meral vein at the level of MA. Costa ends beyond tip of R , at one-third the length be- constriction in the middle. Length of M1 and 5 tween tips of R and M . Sc short, ends at R. M2 fork 5.3ϫ length of M3. Legs: Tibial se- 5 1 tae in rows. Tibial spur formula 1:2:2; mid R1 1.3ϫ the length of r-m. Length of M3 and hind tibial spurs differ in length: inner about equal to length of r-m. Length of M1 spur 1.7ϫ outer one. Fore leg basitarsomere and M2 fork 4.3ϫ the length of M3. Base of shorter than tibia. Abdomen setose, with 6 M3ϩ4 and CuA fork between tip of Sc and visible segments, 7th and 8th ones small and M3 base. Legs: Coxae without setae. Hind retractable. Gonocoxites divided by complete tibiae with posterior bristles. Abdomen with suture ventrally. Male: Gonostyli large, tri- scattered setae. Cerci 2-segmented, apical angular, ¯at, with inner, bilobed appendage. segment conical, basal one cylindrical, 3ϫ Aedeagus bilobate. Female: Eighth sternites length of apical one. Gonocoxites 8 long, triangular; 10th sternite with 4 dark, thick, narrow, slightly curved, with long setae. wavy, blunt setae. Cerci 2-segmented, round- MATERIAL: Holotype AMNH B-0112, fe-

← Figs. 73±75. Protragoneura platycera, n.sp. 73. Holotype AMNH Bu-135. 74. Male genitalia later- odorsally. 75. Male genitalia lateroventrally. 52 AMERICAN MUSEUM NOVITATES NO. 3433 male. Myanmar: Katchin, from amber mines palpomere preapically; length 4ϫ width, near Myitkyina. knoblike. Apical palpomere long and narrow. ETYMOLOGY: The species epithet is Latin Length ratio 1:1.7:2.5. Thorax: Scutum for ``hairless, smooth'', in reference to the densely covered with short erect trichia, absence of thoracic setae and setulae. bearing some setae laterally. Antepronotum and proepisternum with long strong setae. Protragoneura, new genus Anepisternum with deep cleft. Katepisternum without distinct excavation on anterior mar- DIAGNOSIS: Small, dark gnats with anten- gin. Laterotergites, mediotergite bare. Met- nae shorter than thorax, ¯agellomeres shorter episternum with few short trichia ventrally. than wide. Mesonotum densely covered with Wing membrane with macrotrichia in distal short trichia, with or without a few setae lat- and posterior part. Sc extremely short, free. erally. Sc short, free. Costa is not produced All longitudinal veins except R base with se- beyond the tip of R , which is long, curved, 5 tae. M3 reduced, can be traced by setae only. and almost reaching the tip of M . Fork of 1 M base reduced. Base of M and CuA fork M and M shifted anteriorly, M ends before 1 3ϩ4 1 2 1 weakened. Small radial cell length 7ϫ width. wing apex. Base of M and CuA fork at the 3ϩ4 Halters dark, with few short setae. Legs: level of base of M3, in basal one-sixth of Coxae and femora dark, densely setose, tib- wing. M3 length about equal to that of M 1 iae and tarsi yellowish. Hind tibiae with nu- and M fork. 2 merous posterior bristles. Tibial spurs 4ϫ tib- TYPE SPECIES: Protragoneura platycera, ial diameter. Abdomen with 6 visible seg- n.sp. ments, 7th and 8th very short, retracted. Ter- ETYMOLOGY: The genus name is a combi- gite 9 oval, short, covering the base of nation of the pre®x pro- (Greek ␲␳␱-, be- gonocoxites. Gonocoxites fused at base, mas- forehand) and Tetragoneura. The name is sive, narrowing to apex in distal half. Gon- feminine. ostyli with long ventral lobe bearing a comb COMMENTS: Closest to Tetragoneura Win- of short, knoblike setae; a somewhat shorter, nertz, 1846, about 100 extant species of dorsal, bare lobe and internal wide lobe with which are known from the Holarctic (17 a row of curved bristles. spp.), Neotropical (ca. 60 spp.), and Austra- MATERIAL: Holotype AMNH Bu-135, lasian (23 spp.) regions. The new genus dif- male; paratype AMNH Bu-1076, male. fers in having C not extended beyond the tip Myanmar: Katchin, from amber mines near of R , RS2 section (small radial cell) long, 5 Myitkyina. and base of M and CuA fork very basal; 3ϩ4 ETYMOLOGY: The species epithet is derived M and M fork shifted anteriorly. Setation 1 2 from platy (Greek ␲␭␣Â ␶␷␵, broad) and cerus of mesonotum is also distinctive and sepa- (Greek ␬␳␣␵⑀Â , horn), in reference to the com- rates the new genus from Tetragoneura. pressed antennae. Protragoneura platycera, new species ANALYSES Figures 73±75, Plate 7D LYGISTORRHINIDAE AND THE DIAGNOSIS: As for genus. HETEROTRICHA GROUP DESCRIPTION: Body length ϭ 2.49 mm (ho- lotype)/2.39 mm (paratype); wing length ϭ Figure 76 1.63 mm (holotype)/1.93 mm (paratype). Forty-two morphological characters were Head ovate, with row of long erect setae be- coded for all known Mesozoic Lygistorrhin- hind eye margin. Scape and pedicel subcon- idae, Palaeognoriste sp. from Baltic amber, ical, with numerous apical setae, both black. and exemplar species of all living genera of Flagellum compressed laterally, ¯agello- the family. The genera Bolitophila, Chiletri- meres transverse, widths 1.5±2.5ϫ length, cha, Drepanocercus, and Paratinia were densely covered with short trichia. Clypeus used as outgroups. In previous studies My- setose. Three palpomeres seen; antepenulti- cetophilidae was suggested to be a sister mate one swollen, obovate, length 2ϫ width. group to the Lygistorrhinidae, while Bolito- Penultimate one attached to antepenultimate philidae was the sister group to those fami- 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 53

Fig. 76. Phylogeny of Lygistorrhinidae based on cladistic analisys of data presented in table 4 (length 84, CI 0.65, RI 0.81). Ⅲ unreversed changes; ▫ homoplastic character transitions. The character number is indicated above the branch and the state change is indicated below. Bolitophila, Paratinia, Drepan- ocercus, Chiletricha and Thereotricha are the outgroup. lies plus Sciaridae (Matile, 1990, 1997; SoÈli, considered a sister group to all living lygis- 1997). The position of the Heterotricha torrhinids. The present analysis yielded the group is disputed, showing af®nities with same topology for Recent genera: basal po- Diadocidiidae, Sciaridae, and Mycetophili- sition of ``L. asiatica'', and sister-group re- dae, as well as the extinct families Archizel- lationships between Seguyola-Loyugesa and miridae and Mesosciophilidae (Grimaldi et (with Probolaeus as a subge- al., 2003). A data matrix used for cladistic nus of Lygistorrhina). The Cretaceous genera analysis is presented in table 4. Character de- Archaeognoriste and Lebanognoriste are at scriptions are provided in table 5. Analysis the base of the lygistorrhine clade and rep- of the data matrix using Winnona, (version resent a basal, paraphyletic stem group. 2.0; Goloboff, 1999) resulted in four trees of While having super®cial similarity with the length 83, CI of 0.66, and RI of 0.82. The Heterotricha group of genera in venation strict consensus of the trees is shown on ®g- pattern, they show strong apomorphies of ure 76. Lygistorrhinidae in the structure of trans- A previous cladistic analysis of Lygistor- verse veins, antennae, thorax, and male gen- rhinidae (Grimaldi and Blagoderov, 2001) italia (characters 3, 8, 12, 25, 26, 29, 36, 38, included as the only fossil Palaeognoriste, 44). The other Cretaceous genera Plesiog- 54 AMERICAN MUSEUM NOVITATES NO. 3433

TABLE 4 Data Matrix for Analysis of Lygistorrhinidae Character descriptions presented in table 5.

noriste, Protognoriste, and Leptognoriste oc- tree (MPT). An heuristic search (tree bisec- cupy intermediate positions between primi- tion-reconnection), cutting trees up to four tive and recent lygistorrhinids. Interestingly, points, yielded eight MPTs of length 311, CI although these genera do not form a mono- of 0.21 and RI of 0.51. The low indices of phyletic entity, all species have only one vein the trees are due to the very homoplastic da- in the medial fork preserved. The indepen- taset, re¯ected by numerous cases of inde- dent loss of a median vein in various Me- pendent occurrence of structures. The strict sozoic lygistorrhinids suggests rapid radia- consensus of these MPTs (®g. 77a) is an un- tion of the family in the beginning of its his- resolved bush for , with tory, followed by replacement of Mesozoic paraphyletic with respect to Sciophilinae. groups by recent ones. Such an unexpected result can be explained by including in the parsimony analysis two MYCETOPHILIDAE S.S. peculiar fossil genera, Lecadonileia and Dis- Figures 77±79 paroleia, which share some characters (5, 10, Sixty-one characters were coded for 38 17, 18) with Sciophilinae. Nonetheless, in all species representing three main lineages of the analyses conducted, Lecadonileia and Cretaceous fungus gnats: Sciophilinae (in- Disparoleia nested within Leiinae, having cluding 3 recent and 3 fossil species), Lei- synapomorphies with other genera of the inae (3 recent and 13 fossil species), and subfamily (e.g., characters 12, 29, 47, 50, 51, Gnoristinae (9 recent and 7 fossil species). 52). Due to taxon sampling and the limited Macrocera and Bolitophila were used as out- number of characters used, the most parsi- groups. A data matrix used for cladistic anal- monious result hypothesized synapomor- ysis is presented in table 6. Character de- phies of Sciophilinae as reversals. The alter- scriptions are provided in table 7. native, when characters 5, 10, 17, and 18 ap- Winnona (version 2.0; Goloboff, 1999) pear independently in different clades, is less was used to search for the most parsimonious parsimonious by two steps (see below). In- 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 55

TABLE 5 Characters Used in Cladistic Analysis of Lygistorrhinidae All characters are nonadditive.

terestingly, analyzing the dataset with Gnoristinae as basal and paraphyletic to WinXpiwe (version 1.3; Goloboff, 1997), a Sciophilinae and Leiinae. All the fossil spe- program intended for analysis of homoplastic cies assigned to extant genera except Syna- data (see Goloboff, 1993), yields a topology pha longistyla form monophyletic clades with Sciophilinae and Leiinae each as mono- with recent species. Pseudomanota is closely phyletic (®g. 77b). The same result occurs allied with Paratrizygia and probably repre- when Lecadonileia and Disparoleia are ex- sents a stem group to the Azana group of cluded from the analysis. genera (Matile, 1998). Drepanocercus and Since the monophyly of Sciophilinae and Gregikia are at the base of mycetophilids. Leiinae is strongly supported (SoÈli, 1997), Tetragoneura and Ectrepesthoneura should the ®nal analysis was conducted in Winnona be classi®ed in Leiinae, not Gnoristinae, as with Sciophilinae and Leiinae constrained as was suggested by VaÈisaÈnen (1986). On the monophyletic. It resulted in 11 trees of other hand, his conclusion about Syntemna, length 313 (only two steps longer than the traditionally placed in Sciophilinae, is prob- MPTs found), CI of 0.24 and RI of 0.58. The ably right, for on the cladogram it is nested strict consensus tree (®g. 78) of length 333, within Gnoristinae. CI of 0.22 and RI of 0.54 again indicates In general, including fossils into an anal- 56 AMERICAN MUSEUM NOVITATES NO. 3433

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 ®t ϭ 3056.9, concavity 3). ysis dramatically affects results (®g. 79). To extant lineages. Basal taxa have a funda- a large extent this effect is a result of exel- mental in¯uence on topology. This is espe- lent preservation in amber, which has pre- cially true for ancient groups that had a rap- served characters virtually completely. The id initial diversi®cation, where unusual topology of the MPTs is particularly affect- combinations of characters occur. ed, probably due to extensive homoplasy. Homoplasious characters in plesiomorphic DISCUSSION fossils not only decrease resolution and in- PHYLOGENY crease length of the tree but make taxon def- initions diffuse. Nonetheless, including fos- A systematic review of sils in a cladistic analysis has proven to be and Sciaroidea was not the purpose of this extremely useful (Gauthier et al., 1988), and work and is well beyond the scope of our this study is an another example of that. project. We concur with the composition and First, fossils are the only way to establish evolutionary history of the infraorder as es- existence and position of entirely extinct tablished by Shcherbakov et al. (1995), and lineages, like Mesozoic lygistorrhinids or also grant that fossils afford unique insight leiines from Burmese amber. Secondly, they into sciaroid phylogeny. allow the identi®cation of stem groups for The traditional nomenclature of venation 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 57

TABLE 6 Data Matrix for Analysis of Gnoristinae, Sciophilinae, and Leiinae Character descriptions presented in table 7.

in Sciariodea is controversial and somewhat All representatives of Bibionomorpha s.l. (in- misleading. Fortunately, early fossils provide cluding Axymyiiformia and Anisopodifor- a strong basis for vein homology (Rasnitsyn, mia of Shcherbakov et al., 1995) have no

1980; Shcherbakov et al., 1995), since they more than three radial veins posterior to R1, show intermediate conditions. The short vein which can be interpreted as R2,R3, and R4ϩ5 of the radial sector is usually thought to be or R2ϩ3,R4, and R5 according to the authors' R4 (Chandler, 2002). There are four veins in concepts of vein homology. We prefer the the radial sector of the dipteran ground plan. second scheme, assuming that a long vein 58 AMERICAN MUSEUM NOVITATES NO. 3433

TABLE 7 Characters Used in Cladistic Analysis of Gnoristinae, Sciophilinae, and Leiinae All characters are nonadditive.

(R4) is less likely to be atrophied and lost Yala), the base of the forks of radial veins than is a shorter vein (R2). are shifted basally, so bases of the second

Basal Bibionomorpha (Procramptonomyi- (R2ϩ3) and the third (R4) radial veins are very idae, Protorrhyphidae) have crossvein r-m close, and the veins may originate even be- situated between the bases of R2ϩ3 and R4. fore the base of r-m crossvein. These veins The earliest Bibionomorpha have two oppos- are parallel and close to each other along the ing tendencies in the transformation of radial whole length, which makes fusion of these veins. In one group (for example, Triassic veins more likely. The fusion is completed, 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 59

Fig. 78. Phylogeny of Mycetophilidae based on cladistic analysis of data presented in table 6 (length 342, CI 0.22, RI 0.53). Ⅲ unreversed changes; ▫ homoplastic character transitions. The character number is indicated above the branch and the state change is indicated below. Macrocera and Bolitophila are the outgroups. 60 AMERICAN MUSEUM NOVITATES NO. 3433

Fig. 79. Comparison of topologies of trees for fossil and recent species (right) and recent species only (left). Tree on the right is the same as on ®gure 77. Tree on the left is a strict consensus of 6 MPT (length 157, CI 0.40, RI 0.43). 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 61 for example, in Pachyneuridae (where veins versed in Sciaridae). The last character (Ma- are fused basally) and Cramptonomyiidae tile, 1990: 386; 1997: 300) was proposed (apically). Thus, a fore fork in the radial sec- presumably to separate conditions observed tor of Pachyneuridae, Axymyiidae, and Per- in Sciaridae and some representatives of the issommatidae should be considered as R2ϩ3 Heterotricha group, when the mesothoracic ϩ R4. phragma is greatly enlarged and strongly The other group of Bibionomorpha has a projected into the abdomen. Actually, the distal shifting of the bases of these forks, ac- narrow insertion of the abdomen, coupled companied by shortening and reduction of R4 with reduced phagma, is limited to Myceto- (e.g., Alinka, or an undescribed Protorrhy- philidae, Keroplatidae, and Lygistorrhinidae, phidae from the Upper Triassic of North and apparently developed independently.

America) and shifting of the base of R2ϩ3 Other Sciaroidea have a phragma that is close to the base of r-m (Complecia, Eodi- somewhat developed, though not projecting tomyia) or beyond it (Alinka, Eomycetophi- deeply into the abdomen (Chandler, 2002). la). Protopleciidae, a presumed ancestor of Chandler (2002) hypothesized that a large Sciaroidea, has only one long intermediate phragma protruding into the abdomen may vein in its radial sector originating distad of be a synapomorphy of Sciaridae and the Het- the base of crossvein r-m. Having evidence erotricha group (except Pterogymnus and of distal shifting of the base of the radial Sciaropota). Unfortunately, the structure of veins in the earliest history of the Bibiono- the phagma is rarely seen on fossil speci- morpha, and considering that R4 as a shorter mens, even ones in amber, so in most cases vein is more likely be reduced than a long we can only speculate on it. That is why this

R2ϩ3, we prefer to homologize an intermedi- undoubtedly important phylogenetic charac- ate vein in the radial sector of Sciaroidea ter was not included in the analyses. How- with R2ϩ3. A transformation series would ever, it is clear that at least in Lygistorrhini- thus include: from two long veins (in Pro- dae, acquiition of a narrow insertion of the cramptonomyiidae, Protorhyphidae, and Par- abdomen and reduced phragma was not a axymyiidae [Paraxymyiinae]), to one long brief event: early representatives have the and one short vein (Paraxymyiidae [Eomy- plesiomorphic condition (Lebanognoriste, cetophilinae] and Eodytomyiidae), to one Protognoriste, Plesiognoriste) as well as the long vein proximad (Crosaphis) or distad of apomorphic one (Archaeognoriste, Leptog- r-m (Protopleciidae, Pleciofungivoridae, Bol- noriste). It is possible that the two latter gen- itophilidae, Ditomyiidae, and Keroplatidae), era constitute a monophyletic group with re- to ®nally a short crossvein joining R1 or ab- cent Lygistorrhinidae. sent (all other Sciaroidea) (®gs. 80, 81). Reduction of the stem of M (section M1 The posterior fork of Sciaroidea wings is of Kovalev [Kalugina and Kovalev, 1985]), sometimes denoted as CuA1 and CuA2 or M4 proposed as a synapomorphy of Sciaroidea, ϩ CuA1 and CuA2. However, the fore vein actually occurred independently many times in the posterior fork is undoubtedly a rem- in the history of the group. This section is nant of the posterior vein in medial sector, present in Protopleciidae, Pleciofungivori- namely M4 (®g. 80). With reduction of the dae, Bolitophilidae, some Mesosciophilidae, discal cell and M3 (Heterorhyphidae, Parax- and Keroplatidae. If we accept the reduction ymyiidae) and basal displacement of tb and of the section M1 as a synapomorphy of Sci- M stem (Mesosciophilidae, Heterotricha aroidea, the superfamily becames polyphy- group, Mycetophilidae), M4 lost contact with letic or leads to an assumption of loss and the base of M. Vein homology in Diptera was numerous independent restorations of the thoroughly discussed by Shcherbakov et al. vein, which is highly improbable. Other syn- (1995). apomorphies mentioned above combine re- The monophyly of Sciaroidea is supported cent families of Sciaroidea (Ditomyiidae, by several synapomorphies (Matile, 1997), Diadocidiidae, Keroplatidae, Bolitophilidae, including loss of vein R4, abbreviation of the Mycetophilidae, Lygistorrhinidae, Sciaridae) costa at the apex of the wing, long coxae, with several extinct families (Antefungivor- and narrow insertion of the abdomen (re- idae, Archizelmiridae, Eoditomyiidae, Me- 62 AMERICAN MUSEUM NOVITATES NO. 3433 Fig. 80. Scheme of wing venation of different taxa of Bibionomorpha and possible ways of vien transformation. 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 63 Fig. 81. Scheme of wing venation (continued). 64 AMERICAN MUSEUM NOVITATES NO. 3433 sosciophilidae, Pleciofungivoridae, and Pro- taceous of Transbaikalia (Kovalev, 1986). topleciidae) into one monophyletic group Chandler (2002) doubted that the position of with Cecidomyiidae, probably, as a sister Mangas was within Bolitophilidae, although group to all others. he admitted another undescribed but ®gured The phylogeny of recent families of Sci- specimen could belong to the family. He aroidea was proposed by Matile (1990, 1997) wrote (2002: 136) ``There is nothing to in- based on imaginal and preimaginal charac- dicate that it is not allied to Heterotricha''. ters, to which extinct groups could be added. One of us (V.B.) had the opportunity to study Ditomyiidae has been proposed as a sister the type of M. exilis in the collection of PIN. group to all other living Sciaroidea except Despite obscure venation, Kovalev's drawing Cecidomyiidae; however, no Ditomyiidae are seems to be accurate and some characters are known in the fossil record before the Eocene apparent: long curved base of RS is situated (Evenhuis, 1994). Eoditomyia primitiva An- distally, not basally; r-m is vertical, not sorge from the Lias of Germany possibly oblique; section M2 (not tb, as noted on represents a stem-group ditomyiid (Ansorge, Chandler's ®g. 92) is long and longitudinal 1996). The presence of R4 in Eoditomyiidae (not oblique), although not apparent basally may be evidence for an independent origin due to preservation; and there is a short sec- of Ditomyiidae from Paraxymyiidae, which tion M3 (M stem) (it is much shorter than would make Sciaroidea polyphyletic. In any 1ϩ2 M1 and M2 fork and almost the same length case, discovery of Eoditomyia allows us to as visible part of section M2). This combi- propose a Mesozoic, possible Jurassic, origin nation of characters is not known in the Het- of true, crown-group Ditomyiidae, despite erotricha group (where the base of Rs and the absence of fossils. crossvein r-m are not shifted distally, and Matile (1997) provided the only synapo- section M2 is much shorter than section M3), morphy for Diadocidiidae ϩ Keroplatidae, but it is characteristic of Bolitophilidae. Bol- which is the absence or closure of larval ab- itophilidae and the Heterotricha group are a dominal spiracles. Chandler (2002) argued relicts of Mesozoic sciaroids like Protople- that the abdominal spiracles may be second- ciidae and Pleciofungivoridae, though the re- arily lost in Diadocidia due to the tube-living lationships between bolitophilids and genera habit of larvae. The other putative synapo- allied to Heterotricha are not perfectly ap- morphy of Diadocidiidae ϩ Keroplatidae might be the proximal position of the base parent. of RS with respect to tb. The same condition Mycetophilidae and Lygistorrhinidae are is observed in Australosymmerus, but this sister groups when only Recent taxa are an- genus is not considered as basal (Munroe, alyzed. Including fossils in the analysis in- 1974). All other groups of Sciaroidea have dicates an independent origin of Bolitophil- (at least in the ground plan) crossvein tb idae, Mycetophilidae, Lygistorrhinidae, and proximal to or at the level of the base of RS. possibly Sciaridae from within the Mesoscio- The vertical alignment of veins r-m, M2, and philidae-Pterogymnus-Heterotricha com- tb in Diadocidiidae and Ditomyiidae, though plex. All these recent families as well as ex- similar in shape, probably appeared indepen- tinct Archizelmiridae share synapomorphies, dently. Rigidity of the wing is provided by a which are also found in Mesosciophilidae and Heterotricha group. Pterogymnus Free- long R1 in Ditomyiidae and by the length- ened base of RS in Diadocidiidae. Thus, man is probably related to Mesosciophilidae crossvein r-m was pulled basad with the base (Chandler, 2002). One of the Heterotricha- of RS in Diadocidiidae, while in Ditomyiidae like genera, Sciaropota Chandler, 2002, crossveins r-m, tb, and m-cu were pulled dis- shows an alliance with an archizelmirid ge- tad together with the base of RS. nus, Archimelzira from New Jersey amber Bolitophilidae are represented by only one (Grimaldi et al., 2003), and probably should genus in the Recent fauna. Another genusÐ be included in Archizelmiridae. Relation- Mangas KovalevÐwas described from the ships within the Heterotricha group cannot Lower Cretaceous of Mongolia, and close be resolved easily because of the extensively relatives were reported from the Lower Cre- paraphyletic nature of this complex. 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 65

TAPHONOMY AND BIOGEOGRAPHY aroids present con¯icting evidence. On the one hand, sciaroids in Burmese amber were Sciaroidea today are most abundant and abundant, as in temperate forests. However, diverse in moist to wet temperate forests of no species is particularly abundant, and Lei- the Holarctic and south temperate (e.g., Aus- inae and Lygistorrhinidae are quite diverse, tral) regionsÐwherever thick humus and as would be found in tropical forests today. fungal mycelia are abundant. Although sci- aroids are diverse in tropical forests, they are ACKNOWLEDGMENTS diffuse in numbers there. Among taxa known in the Cretaceous, Leiinae and Lygistorrhin- This work would not have been possible idae are better represented in the tropics than without the generous support of Robert G. Gnoristinae, which are common in temperate Goelet, Chairman Emeritus of the AMNH regions but scarce in tropical forests. Board of Trustees, to the senoior author. We The most striking aspect of the Cretaceous are sincerely grateful to his patronage of col- amber faunas is the one in Burmese amber. lection-based research and for his funding Of all insect inclusions in other Cretaceous Burmese amber acquisition. The work was ambers, sciaroids comprise 1% or less in greatly facilitated by assistance and advice each deposit: For example, Taymyr 12 sci- received from Jeffrey Cumming, Michael S. aroids/3450 inclusions; Canada 5/1281; Leb- Engel, Martin Ramirez, Alexandr P. Rasnit- anon 2/600, 2/917, 15/1258 (all of the latter syn, Andrew J. Ross, Valerie Schawaroch, are archizelmirids); and New Jersey 17/1800. Toby Shuh, Alexander I. Zaitzev, F. Christian Of all insects in Burmese amber, by contrast, Thompson, and especially from our late col- 5% of them are sciaroids. This is a propor- leagues LoõÈc Matile and Vladimir V. Zheri- tion very similar to that found in Baltic and khin. We wish to thank also Peter Chandler Dominican ambers. The proportions of Baltic and Andre Nel for their helpful comments of amber sciaroids have been grossly overesti- the manuscript. mated based on museum collections (which are highly biased) at 15±20%. Based on REFERENCES crude Baltic amber samples analyzed by PIN at the mine sites in Kaliningrad in 1992, sci- Alonso, J., A. Arillo, E. Barron, J. C. Corral, J.L. Grimalt, J.F. Lopez, R. Lopez, X. MartõÂnez-Del- aroids (without Sciaridae) comprise 4% of all cloÂs, V. Ortuno, E. Penalver, and P.R. Trincao. insect inclusions. The proportion of Myce- 2000. A new fossil resin with biological inclu- tophilidae in Dominican amber based on un- sions in Lower Cretaceous deposits from Alava biased samples is 3.3% of all insect inclu- (northern Spain, Basque-Cantabrian Basin). sions (data of D.G.). Among Cretaceous de- Journal of Paleontology 74(1): 158±178. posits the Burmese sciaroid fauna is most Ansorge, J. 1996. Insekten aus dem oberen Lias similar to Early Cretaceous orictocenoses of von Grimmen (Vorpommern, Norddeutsch- Baisa and Bon Tsagan (Lower Cretaceous of land). Neue Palaeontologische Abhandlungen Transbaikalia and Mongolia), which have 2: 1±132. Azar, D. 2000. Les ambres MeÂsozoõÈques du Li- 1.9% and 1.3%, respectively, of their entire ban. Ph.D. dissertation, University of Paris, Or- insect faunas comprised of sciaroids. The say, 164 pp. Taimyr, New Jersey, and Canadian amber Bechev, D. 2000. World distribution of the genera sciaroid faunas seem the most similar to each of fungus gnats (Diptera: Sciaroidea, excluding other. Sciaridae). Studia Dipterologica 7(2): 543±552. The Burmese amber sciaroid fauna prob- Blagoderov, V.A. 1993. Dipterans (Mesosciophil- ably re¯ects a particularly wet paleoenviron- idae) from the Lower Cretaceous of Transbai- ment, although not necessarily warmer than kal. Paleontological Journal (Paleontologiches- the other Cretaceous deposits. Based on a kii Zhurnal) 27(1A): 123±130. Blagoderov, V.A. 1995. Fungus gnats of the tribe large, new assemblage of more than 3000 in- Sciophilini (Diptera, Mycetophilidae) from the sect inclusions in Burmese amber, Grimaldi Early Cretaceous of Transbaikalia. Paleontolog- et al. (2002) concluded that Burmese amber ical Journal (Paleontologicheskii Zhurnal) re¯ected the most tropical environment 29(1): 72±83. among all Cretaceous amber deposits. Sci- Blagoderov, V.A. 1996. Revision of the nemato- 66 AMERICAN MUSEUM NOVITATES NO. 3433

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tomology. The Quarterly Journal of the Geo- (trakheinye i khelitzeratovye) [Development logical Society of London 10: 378±96. and changes of the Cretaceous and Caenozoic Whalley, P.E.S. 1976. Lower Cretaceous Lepidop- faunistic assemblages (Tracheata and Chelicer- tera. Nature 266(5602): 526. ata)]. Trudy Paleontologicheskogo Instituta Wu, H. 1995. Diptera: Diadocididae [Diadocidi- 165: 1±198. [in Russian] idae] and Keroplatidae. In H. Wu (editor), In- Zherikhin, V.V., and K.Yu. Eskov. 1999. Meso- sects of Baishanzu Mountain, eastern China: zoic and lower Tertiary resins in former USSR. 423±434.Beijing: China Forestry Publishing In J. Alonso, J.C. Corral, and R.V. Lopez (con- House. veners). Proceedings of the World Congress on Yakovlev, E.B. 1988. Plodonoshenie gribov i se- amber inclusions. Estudios del Museo de Cien- zonnaya aktivnost' dvukrylykh nasekomykh v cias Naturales de Alava 14, Numero especial 2: sosnovykh i ocinovykh molodnyakakh. [Mush- 119±131. Zherikhin, V.V., and A.J. Ross. 2000. A review of room fructi®cation and seasonal activity of dip- the history, geology and age of Burmese amber terans in pine and aspen underwoods]. Petro- (burmite). Bulletin of the Natural History Mu- zavodsk, 127 pp. [in Russian] seum Geology Series 56(1): 3±10. Zaitzev, A.I. 1994. Gribnye komary fauny Rossii Zherikhin, V.V., and I.D. Sukacheva. 1973. On the i sopredelnykh regionov. Chast 1. [Fungus Cretaceous insect-bearing ``ambers'' (retinites) gnats of the fauna of Russia and adjacent re- from North Siberia. In Voprosy paleontologii gions. Part 1]. Moscow: Nauka, 288 pp. [in nasekomykh [Problems of Insect Paleontolo- Russian] gy]. Doklady na XXIV Ezhegodnom Chtenii Zherikhin, V.V. 1978. Razvitie i smena melovykh Pamyati N.A. Kholodkovskogo 1±2 Apr. 1971: i kainozoiskikh faunisticheskikh kompleksov 3±48. Leningrad: Nauka. [in Russian] 70 AMERICAN MUSEUM NOVITATES NO. 3433

Plate 1. Families Diadocidiidae and Lygistorrhinidae. A. Docidiadia burmitica Blagoderov and Gri- maldi, holotype AMNH Bu-033. B. Thereotricha sibirica Blagoderov and Grimaldi, holotype PIN 3130/ 183. C.? T. agapa Blagoderov and Grimaldi, holotype PIN 3426/256. D. Archaeognoriste primitiva Blagoderov and Grimaldi, holotype AMNH Bu-1539. E. Lebanognoriste prima Blagoderov and Gri- maldi, holotype AMNH JG268/1. F. Plesiognoriste carpenteri Blagoderov and Grimaldi, holotype MCZC 6927. 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 71

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 AMERICAN MUSEUM NOVITATES NO. 3433

Plate 3. Family Mycetophilidae: Manotinae, Sciophilini, Gnoristini. A. Alavamanota burmitina Bla- goderov and Grimaldi, holotype AMNH Bu-1271, B. Neuratelia maimecha Blagoderov and Grimaldi, holotype PIN 3311/661. C. Allocotocera burmitica Blagoderov and Grimaldi, holotype AMNH B-056. D. Pseudomanota perplexa Blagoderov and Grimaldi, holotype AMNH Bu-599a. E. Apolephthisa bu- lunensis Blagoderov and Grimaldi, holotype PIN 3963/4. F. Synapha longistyla Blagoderov and Gri- maldi, holotype MCZC 6944. 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 73

Plate 4. Family Mycetophilidae: Gnoristini and Leiini. A. Dziedzickia nashi Blagoderov and Gri- maldi, holotype AMNH NJ-117a. B. Saigusaia pikei Blagoderov and Grimaldi, holotype TMPD P79.15.7.21. C. Syntemna ®ssurata 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. 74 AMERICAN MUSEUM NOVITATES NO. 3433

Plate 5. Family Mycetophilidae: Leiini. A. Nedocosia sibirica Blagoderov and Grimaldi, holotype PIN 3311/665. B. N. canadensis Blagoderov and Grimaldi, holotype MCZC 6897. C. N. novacaesarea Blagoderov and Grimaldi, holotype AMNH NJ-117k. D. Ectrepesthoneura succinimontana Blagoderov and Grimaldi, holotype PIN 3311/662. E. E. swolenskyi Blagoderov and Grimaldi, holotype AMNH NJ- 824. F. Izleiina miri®ca Blagoderov and Grimaldi, holotype PIN 3130/187. 2004 BLAGODEROV AND GRIMALDI: SCIAROIDEA IN AMBER 75

Plate 6. Family Mycetophilidae: Leiini. A. I. spinitibialis Blagoderov and Grimaldi, holotype AMNH NJ-346. B. Zeliinia orientalis Blagoderov and Grimaldi, holotype AMNH Bu-315. C. Z. occidentalis Blagoderov and Grimaldi, holotype MCZC 6943. D. Temaleia birmitica Blagoderov and Grimaldi, holotype AMNH Bu-483a. E. Lecadonileia parvistyla Blagoderov and Grimaldi, holotype MCZC 6941. 76 AMERICAN MUSEUM NOVITATES NO. 3433

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