The Frit (, Diptera) of Fennoscandia and Denmark Fauna Entomologica Scandinavica

Editor-in-chief: N.P. Kristensen

Desk editor: V. Michelsen

VOLUME 43

The titles published in this series are listed at brill.com/fes The Frit Flies (Chloropidae, Diptera) of Fennoscandia and Denmark

By Emilia P. Nartshuk & Hugo Andersson

LEIDEN · BOSTON 2013 Cover illustration: frit (Linnaeus, 1758), female.

Library of Congress Cataloging-in-Publication Data

Nartshuk, E.P. The frit flies (Chloropidae, Diptera) of Fennoscandia and Denmark / by Emilia P. Nartshuk and Hugo Andersson. pages cm. – (Fauna entomologica scandinavica ; volume 43) Includes index. ISBN 978-90-04-16710-0 (hardback : acid-free paper) – ISBN 978-90-04-19066-5 (e-book) 1. Frit –Scandinavia–Classification. 2. Frit fly–Scandinavia–Identification. 3. Chloropidae–Scandinavia– Classification. 4. Chloropidae–Scandinavia–Identification. I. Andersson, Hugo, 1927–2008. II. Title.

QL537.C46N37 2013 595.77–dc23 2013006443

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This book is printed on acid-free paper. Contents

Abstract...... 1 Listofnewtaxa,synonymsandnomenclaturalchanges...... 1 Introduction...... 2 Materialandacknowledgements...... 3 ReviewofpreviousstudiesofChloropidaeinFennoscandiaandDenmark...... 4 Methodsofcollecting,preparingandpreserving...... 7 Morphology,diagnosticcharactersandanatomy...... 9 Adults...... 9 Preimaginalstages...... 22 Eggs...... 23 Larvae...... 24 Anatomyofadultsandlarvae...... 27 Bionomicsandecology...... 28 Habitatsandadulthabits...... 28 Modesoflarvalfeeding...... 28 Seasonaloccurrence...... 29 Roleinlandecosystemsandspeciesofeconomicimportance...... 32 Parasitoids,predatorsandparasites...... 37 Distributionandzoogeography...... 47 ...... 49 Monophylyandrelationships...... 49 Classificationandnomenclature...... 50 Key to subfamilies ...... 52 KeytoFennoscandiangenera...... 53 Preliminarykeytogenerafor3rdinstarlarvae...... 57 Subfamily Rhodesiellinae ...... 59 Genus Aspistyla Duda...... 59 Subfamily ...... 61 Genus Duda...... 61 Genus Enderlein...... 68 Genus Colliniella Nartshuk&Andersson,gen.nov...... 77 Genus Duda...... 78 Genus Loew...... 85 Genus Macquart...... 90 Genus Becker...... 96 Genus Schiner...... 100 Genus Loew...... 101 Genus Duda...... 108 Genus Sabrosky...... 109 Genus Anonymous...... 114 Genus Meigen...... 118 Genus Microcercis Beschovski...... 122 Genus Oscinella Becker...... 124 Genus Oscinimorpha Lioy...... 133 Genus Lioy...... 135 Genus Polyodaspis Duda...... 138

v Genus Malloch...... 139 Genus Duda...... 141 Genus Macquart...... 145 Genus Rondani...... 146 Genus Speccafrons Sabrosky...... 148 Genus Enderlein...... 148 Genus Lioy...... 149 SubfamilyChloropinae...... 154 Genus Camarota Meigen...... 154 Genus Centorisoma Becker...... 155 Genus Cetema Hendel...... 155 Genus Chlorops Meigen...... 159 Genus Chloropsina Becker...... 185 Genus Cryptonevra Lioy...... 186 Genus Diplotoxa Loew...... 190 Genus Diplotoxoides Andersson...... 190 Genus Epichlorops Becker...... 192 Genus Eurina Meigen...... 193 Genus Eutropha Loew...... 194 Genus Lasiosina Becker...... 196 Genus Melanum Becker...... 200 Genus Meromyza Meigen...... 200 Genus Neohaplegis Beschovski...... 218 Genus Parectecephala Becker...... 218 Genus Platycephala Fallén...... 219 Genus Strobl...... 223 Genus Thaumatomyia Zenker...... 226 Genus Trichieurina Duda...... 230 Abbreviations...... 235 Catalogue...... 237 Literature...... 247 Taxonomicindex...... 279

vi Abstract

This volume deals with North European species From Fennoscandia and Denmark are known 48 of the family Chloropidae, known as frit flies or genera and 209 species of frit flies: Rhodesiellinae chloropid flies, a large family of acalyptrate flies 1 species, Oscinellinae 111 species, 97 including some notarial, economically pests of ce- species. Among them 119 species are known from reals and fodder grasses. The taxonomy, biology Denmark, 97 from Norway, 189 from Sweden, 144 and faunistics of all NW European species are re- from Finland, and 115 from included provinces of vised. Keys are given to subfamilies, genera and Russia (Karelian Isthmus, Karelia, and the Kola species for the adults, for the larvae to generic level Peninsula). One new genus and 11 new species are as far as possible, and to specific level for a few ge- described, and some new synonyms and nomen- nera. Brief descriptions of the adult flies are given clatural changes are proposed. Some other new for all genera and species. Species distributions species, new synonyms, and overlooked specific in Fennoscandia and Denmark and elsewhere are names resulting from this revision have been pub- briefly outlined and further tabulated in a cata- lished earlier (Nartshuk, 1992, 1998, 1999, 2002a; logue. The known biology and ecology is summa- Nartshuk & Andersson, 2002; Nartshuk & Przhi- rized in general chapters and for each species. Il- boro, 2009; Nartshuk & Tschirnhaus, 2012). lustrations are given of the male genitalia and also of other characters of diagnostic importance.

List of new taxa, synonyms and nomenclatural changes

New taxa: Rhopalopterum tomentosum sp. nov. from North Colliniella gen. nov.; type species: Conioscinella mei- Sweden. jerei Duda, 1933. Restored from synonymy: Aphanotrigonum norrbotticum sp. nov. from North Oscinis ephippium Zetterstedt, 1848. Sweden. New synonyms: Calamoncosis halterata sp.nov.fromSweden. Aphanotrigonum brachypterum (Zetterstedt, 1848) Chlorops bjerkanderi sp.nov.fromSweden. = Aphanotrigonum trilineatum var. micropte- Conioscinella abiskoi sp.nov.fromNorthSweden. rum Duda, 1932. Conioscinella messaurea sp.nov.fromNorthSwe- Aphanotrigonum cinctellum (Zetterstedt, 1848) den. = Oscinis fasciella Zetterstedt, 1855. Conioscinella tornensis sp.nov.fromNorthSweden. Dicraeus nitidus Wahlgren, 1913 = Dicraeus na- Dicraeus tjederi sp. nov. from southern Sweden. paeus Collin, 1946. Eribolus danicus sp. nov. from Denmark and Swe- Eribolus slesvicensis Becker, 1910 = Eribolus cras- den. sipes Nartshuk, 1972. Gaurax norvegicus sp. nov. from Norway and Swe- Gaurax ephippium (Zetterstedt, 1848) = Gaurax den. strobilum Karps, 1981. Gaurax suecicus sp.nov.fromSweden. Trachysiphonella rufijiceps (Macquart, 1835) = Chlo- Lasiosina danielssoni sp.nov.fromGotland,Swe- rops pygmaeus Meigen, 1838 = Oscinis flavella den. Zetterstedt, 1848 = Siphonella diplotoxoides Meromyza bifurcata sp.nov.fromSweden. Strobl, 1893.

1 New combination and status: Aphanotrigonum trilineatum var. micropterum Microcercis kroeberi (Duda, 1933). Duda, 1932. Lectotype designations: Gaurax borealis Duda, 1933.

Introduction

The purpose of the present work is to present a re- first level consumers. Their role as pests of cere- vision of the species of Chloropidae or “frit flies” in als and cultivated grasses should also not be ne- a broad sense, also known as chloropid flies or grass glected. Increased abundance of certain species of flies, occurring in NW Europe with special empha- Chloropidae is used as a bioindicator that cereals sis on Denmark, Sweden, Norway, Finland and the and grasses are under environmental stress, e.g. adjacent Russian provinces of the Fennoscandian from draught or industrial pollution. Peninsula. It provides keys to subfamilies, genera The studies made in preparation of the present and species, descriptions of adult and larval mor- book had some spin-off in terms of descriptions phology, biology, and local and Palearctic distribu- of new species from Fennoscandia and Denmark, tion. Chloropidae is a large family of small aca- new synonymies, finds of overlooked names, and lyptrate flies usually 2.0–4.0 mm in length, excep- a proposal for conservation of a specific name tionally 5.0–8.0 mm. They are morphologically and (Nartshuk, 1992, 1998, 1999, 2002a, 2004; Nartshuk ecologically very diverse. The flies may be found in & Andersson, 2002). Still, not all taxonomical prob- practically all kinds of terrestrial habitats. Greatest lems concerning Scandinavian Chloropidae have diversity and abundance of species is usually found been settled with the present book. Especially the in open landscapes ranging from sandy coasts, over taxonomic problems within Oscinella may not be dry to humid grasslands to bogs, fens, marshes and fully elucidated on basis of morphology alone. other types of wetlands. Fewer chloropids are spe- The morphological terminology follows that pro- cialized forest inhabitants. The common name of posed in the Manual of Nearctic Diptera (McAlpine, these flies contains in many languages the word 1981). The treatment of each species begins with “frit”: frit flies, Fritfliegen, fritflugor, fritfluer. This the valid name followed by the original combina- word was used by C. Linnaeus, when he described tion followed by synonyms established after the the first species of the family as Musca frit (now Catalogue of Palearctic Diptera, vol. 10 (Soós & Papp known as ), a serious pest of cereals in many European countries. (eds), 1984) and listed chronologically with their The very first studies on chloropid flies came bibliographic references. Then follows a diagnosis from Swedish entomologists: C. Linnaeus (1707– and description to ensure correct differentiation 1778), C. Bjerkander (1735–1795), C.F. Fallén (1764– from related species. These are supplemented with 1830), J.W. Zetterstedt (1785–1874) and dealt with illustrations, mainly line drawings, of male geni- the local fauna. Their studies proved particularly talia and sometimes other body parts for nearly all important because they both contained descrip- the species. A few species cannot be safely identi- tions of new species and contributed information fied without examination of the male and female on the biology and damage to agricultural crops genitalia. Each description is followed by an out- caused by these flies. Many European and Palearc- line of the known distribution in Denmark and the tic species of Chloropidae were first described on Fennoscandian countries, and worldwide. Then basis of specimens originating from Sweden. Very follows a brief outline of bionomics in terms of phe- many chloropids in Europe have their northern- nology, habitat preferences and larval host plants. most distributional limit running through Sweden That piece of information, especially about host and the neighboring countries Norway and Fin- plants, is often based on extralimital observations land. in the absence of local data. Most frit flies, especially those belonging to the A Catalogue giving the known distribution by subfamily Chloropinae, have phytophagous larvae. faunistic province for each species in Denmark and In spite of their small size, their abundance gives Fennoscandia is provided in the format practiced them an important role in grass ecosystems as in previous volumes of the series.

2 Altogether 209 species of Chloropidae in 48 ge- ily of flies. Similar studies of the canopy fauna nera have been found in Fennoscandia and Den- of Diptera in Scandinavia has only just started mark, but a few more species can be expected. A lot (Thunes et al., 2004). remains however to be found out about the bio- The authors hope that this volume of Fauna Ento- nomics of Scandinavian frit flies, including such ba- mologica Scandinavica will make the identification sics as the larval host plants of many phytophagous of NW European species of Chloropidae easier and species. An examination of the commu- thus promote the general knowledge about this im- nity of a rain forest canopy (Floren, 2003) indicated portant family of flies. that the Chloropidae were the most abundant fam-

Material and acknowledgements

The present work is based on examination of more Chloropidae from Norway were borrowed from than 100 000 specimens of Chloropidae from dif- the Zoological Museum of Oslo University and the ferent collections in Scandinavian countries and Zoological Museum of Bergen University, includ- material collected by the authors: H. Andersson ing the collection of H. Siebke. The Chloropidae mainly in Sweden from 1954; E.P. Nartshuk on collections in the Oslo Museum were also stud- the Karelian Isthmus (1956–2000) and in Finland ied by E.P. Nartshuk on a visit in 2008. Material (1995). Most material from Sweden is deposited from the Karelian Isthmus, including the collection in the Zoological museum of Lund University and of A.A. Stackelberg, Karelia and the Kola Penin- consists of two main parts: the collections of C.F. sula deposited in the Zoological Institute of Rus- Fallén, J.W. Zetterstedt, E. Wahlgren and O. Ring- sian Academy in St. Petersburg were studied by E.P. dahl with numerous type-specimens for their new Nartshuk as well. Additional material from Finland species, and the material collected by H. Andersson was received from Dr M. Koponen, from Norway from different parts of Sweden. Material from other from Dr L. Greve Jensen, and K. Thunes, from Kare- Swedish collections were also examined, first of all lia from Dr A. Polevoi and Dr A. Przhiboro, from the collection of the Naturhistoriska Riksmuseet Sweden from Dr B. Vicklund. All records listed are in Stockholm, including old material of C.H. Bo- based on specimens determined by the authors. heman and P.F. Wahlberg. Further, a collection at Distribution records from literature are not used, the Zoological Museum of Göteborg University and except the type localities of species described from two collections in Uppsala, deposited respectively the territory investigated. in the Zoological Museum of the Uppsala Univer- For the loan of material and/or for informa- sity and in the Agriculture University. tion the authors thank Lita Greve Jensen, Karl E.P. Nartshuk visited the Zoological Museum of Thunes, Pekka Vilkamaa, Bernard Lindeberg, Gu- the Helsinki University four times (1984, 1985, 1990, nilla Ståhls, Leif Lyneborg, Verner Michelsen, Mi- 2004) to study the large collection of chloropids chael von Tschirnhaus, Alexei Polevoi, Andrei from Finland, Karelia and the Kola Peninsula, in- Przhiboro, John W. Ismay, Walter Hackman, Martti cluding material of E.J. Bonsdoff, J. Sahlberg, R. Koponen, Bert Vicklund. Hans Silverberg (Helsinki) Tuomikosky and types of O. Duda, R. Frey and located many collecting sites and interpreted old F. Reuter. H. Andersson also borrowed types and spellings of locality names in Finnish material. interesting species from the Helsinki Museum. Our sincere thanks to Thomas Pape and Roy The collections of the Zoological Museum (now Danielsson for constant support and help, espe- Natural History Museum of Denmark), University cially during visits of E.P. Nartshuk to Stockholm, of Copenhagen with material from Denmark, in- Lund and Copenhagen, and to Vladimir I. Gusarov cluding material of W.H.C.F. Wüstnei, R.C. Stæger, who organized the visit of E.P. Nartshuk to Oslo in R.W. Schlick, W. Lundbeck, and L. Lyneborg, and 2008 to study Chloropidae collections from Nor- also from other Scandinavian countries were exam- way. ined by both authors (H. Andersson in 1996, E.P. We are indebted to S. Riebe (Botanical Museum Nartshuk in 2004). of the Lund University) for advice on biotic regions

3 and vegetation zones of northern Fennoscandia, MNHN – Muséum d’Histoire Naturelle, Paris, and Anders Peltz (Klassiska och semitiska institu- France. tionen, Lund University) for advise on the Latin NHMH – Natural History Museum, Helsinki, Fin- name of Musca frit. land. Visits by E.P. Nartshuk to Helsinki (1984, 1990), NHMW–NaturhistorischesMuseum,Vienna,Aus- Stockholm and Lund (1997, 1999, 2002) were sup- tria. ported by the Russian Academy of Sciences and NMBA – Naturhistorisches Museum der Benedikti- Svenska institutet (2004). Financial support from ner-Abbei, Admont, Austria. Russian foundation of Basic Researcn (No. 11-08- NMID – National Museum of Ireland, Dublin, Ire- 00186, 13-04-00639). My sincere thanks go to land. V. Michelsen for his extensive editorial work on NMPC – National Museum, Natural History, Prague, my MS. Czech Republic. Types of many species were examined by E.P. NRMS – Naturhistoriska Riksmuseet, Stockholm, Nartshuk in Deutsche Entomologische Institut, Sweden. Eberswalde [now Müncheberg] (1993), Museum OXUM – Hope Entomological Collections, Univer- für Naturkunde an der Humboldt Universität, Ber- sity Museum, Oxford, England. lin (1993), Staatliches Museum für Naturkunde, SMNS – Staatliches Museum für Naturkunde, Stutt- Stuttgart (1993), Hope Entomological Collection, gart, Germany. University Museum, Oxford (1998), Zoölogisch Mu- ZISP – Zoological Institute, Russian Academy of seum, Universiteit van Amsterdam (2001), Natur- Sciences, St. Petersburg, Russia. historisches Museum, Vienna (1996), Hungarian ZIWP – Zoological Institute, Polish Academy of Sci- Natural History Museum, Budapest (1968, 1972, ences, Warzaw, Poland. 1976, 1986) and Muséum National d’Histoire Na- ZMAN – Zoologisches Museum, Universiteit van turelle, Paris by H. Andersson in 1960 and E.P. Amsterdam, The Netherlands. Nartshuk in 2001. ZMBA – Zoological Museum, Bulgarian Academy The following abbreviations are used for the lo- of Sciences, Sofia, Bulgaria. cation of type specimens and other material men- ZMHU – Museum für Naturkunde an der Hum- tioned in the text. boldt Universität, Berlin, Germany. ZMLU – Zoological Museum, Lund University, BMNH – The Natural History Museum, London, Lund, Sweden. England. ZMMU – Zoological Museum, Moscow University, DECU – Department of Entomology, Charles Uni- Moscow, Russia. versity, Prague, Czech Republic. ZMOC – Zoological Museum, Opava, Czech Repub- DEIM – Deutsche Entomologische Institut, Mün- lic. cheberg, Germany. ZMUC – Natural History Museum, University of HNHM – Hungarian Natural History Museum, Bu- Copenhagen, Denmark. dapest, Hungary. ZMUU – Zoological Museum, Uppsala University, MHNB – Museé d’Histoire Naturelle, Bruxelles, Bel- Sweden. gium. MHNL – Musée d’Histoire Naturelle, Lille, France.

Review of previous studies of Chloropidae in Fennoscandia and Denmark

Sweden Linnaeus described a fly which damaged grains of oats in Sweden. He formally described this fly as C. Linnaeus published in 1750 the first data on Musca frit in the 10th edition of his Systema nat- Scandinavian Chloropidae in a paper on in- urae from 1758. The word “frit” is maintained in jurious to cereals. Under the name “korn-flugan” the vernacular names of the family Chloropidae

4 in many languages: frit flies, Fritfliegen, fritfluer, way, Denmark and Finland. Andersson (1966) re- fritflugor. vised all Zetterstedt’s species with lectotype desig- Two opinions have been expressed on the origin nations. 15 species described by Zetterstedt are still of this word. One opinion, repeated in many agri- valid. cultural books, suggests with reference to C. Lin- Only a few new taxa of Chloropidae were de- naeus that the word frit originates from Swedish scribed from Sweden after Zetterstedt. Holmgren language and means “light grain” (Schaufuss, 1893; (1887) described Chlorops taeniopus gotlandica and Schander & Meyer, 1925). Schaufuss (1893) wrote its bionomics and parasites from Gotland, but this “Frit bedeutet im Schwedischen ‘Leichte Waare’, subspecific name has been omitted in most subse- der Name weist also auf die von der Sommer- quent literature on Chloropidae. Wahlgren (1913) Generation angefressenen Körner hin” (p. 82). In- described 3 new species from Öland. Andersson deed grains damaged by larvae of frit flies are un- (1966) described two new species in his revision of ripe and therefore light. The other, better substan- Zetterstedt’s collections. tiated opinion expressed by Jablonowski (1911) sug- Records of chloropids new to Sweden and of gestsanoldLatinoriginofthiswordandrefersto new data on distribution were published in fau- M. Terentius Varro. This is supported by the Ox- nistic papers by Boheman (1852, 1864, 1868), Wal- ford Latin Dictionary (Glare (ed.), 1968–1982) that lengren (1867), Roth (1896, 1897), Wahlgren (1915) gives the following explanation of the word “frit”: and Poppius et al. (1916). Wahlgren (1919) summed “A tiny particle, perh. undeveloped grain, at the top up existing data and gave a review and a key to of an ear of corn; (as a type of minuteness) a ‘grain’ ” 19 genera and 65 species of Swedish Chlorop- (p. 736) and cites two works using the word: Mostel- idae in Svensk insektfauna. In succeeding years laria by T. Maccius Plautus (d. 184 B.C.) and Res it was first of all O. Ringdahl who in numerous Rusticae by M. Terentius Varro (116–27 B.C.). publications (1921, 1931, 1939, 1941, 1947, 1950, Another species of Chloropidae Musca saltatrix 1951, 1954, 1959a, 1959b, 1960) added species and was described by Linnaeus (1761) in Fauna svecica. distribution data to Wahlgren’s list of Swedish Bjerkander (1777, 1778a, 1778b, 1779, 1781, 1789) de- Chloropidae. Other contributions to the knowl- scribed 3 species with biological observations on edge about Swedish Chloropidae are given in pa- another harmful species, now known as Chlorops pers by Jansson (1922), Lindroth (1943), Hedström pumilionis (Bjerkander). The identity of the lat- (1964), Lundberg (1964), Andersson & Daniels- ter species was discussed by Lampa (1888) and son (1980) and Nartshuk (2002a). New synonyms Wahlberg (1918) and Bjerkander’s name of this well based on study of types specimens of Fallén, Zetter- known pest of and barley has been placed on stedt and others were published by Nartshuk & the Offfijicial list of specifijic names in zoology (Name Andersson (2002). Hedström (1994) stated that 100 No. 868). (actually 101) species of Chloropidae were known The first monograph of Swedish Chloropidae from Sweden but estimated that the total number was published in 1820 by C.F. Fallén. He listed 20 of species might rather be 150. species in his family Oscinides. Only 16 species The foregoing papers about Chloropidae con- belong to Chloropidae in the modern sense, but tains mainly taxonomic and faunistic data with these include 1 genus and 12 species proposed as brief notes on habitats and phenology for some new to science. Andersson (1963) revised Fallén’s species. Ardö (1957) on the contrary carried out species with lectotype designations, and 1 genus a study with emphasis on the ecology of Diptera and 10 species are still valid. One more species of occurring in marine shore dunes in Sweden, Nor- Chloropidae was described by Fallén (1823) in his way and Denmark. He recognized 49 species of Phytomyzides. J.W. Zetterstedt dealt with Chlorop- Chloropidae occurring in the dune ecosystems idae from northern Sweden in his Insecta Lappon- investigated. Ardö specifically studied the influ- ica from 1838, and from all parts of Scandinavia in ence of some ecological factors on three species several volumes of his Diptera Scandinaviae (1848, of Chloropidae being stenotopic inhabitants of 1852, 1855, 1860). He carried out an enormous marine shores: Colliniella meijerei, Conioscinella amount of faunistic and taxonomic research. All zetterstedti (as C. brachyptera)andEutropha ful- 6 species of Chloropidae described by Zetterstedt vifrons. Insects, including two chloropid species, (1838) are still valid. 28 species of Chloropidae were associated with the polypore Fomitopsis pinicola described as new in different volumes of Diptera across Fennoscandia were studied by Komonen et Scandinaviae based on material from Sweden, Nor- al. (2004).

5 Chloropid species injurious to cereals and agri- Diptera from marine dune ecosystems in Scandi- cultural grasses in Sweden have been most thor- navia records several species of Chloropidae from oughly studied. Applied aspects connected with Denmark. Lyneborg (1965, 1968) gave lists of Chlo- Oscinella frit, O. pusilla and Chlorops pumilionis ropidae from the Hansted Reservation, NW Jut- are treated in numerous papers. Linnaeus (1750) land and, based on identifications by H. Andersson, and Bjerkander (1777, 1778a, 1778b, 1779, 1781, 1789) from the Faroes. The first list includes 32 species were the first to describe the damage to cereals in 18 genera and the second 3 species in 3 genera. caused by Oscinella frit and Chlorops pumilionis, Petersen (2001) published a check list of Danish respectively. Both pest species were considered Chloropidae that even mentions species expected in Dahlbom’s (1837) review of harmful Scandina- to be present based on known distributions. vian insects. Numerous papers dealing with these Chloropidae as agricultural pests and applied species were published later on, some of which are problems with frit fly in Denmark were considered listed in the chapter ‘Role in land ecosystems and in numerous publications, some of them listed in species of economic importance’. The whole com- the chapter ‘Role in land ecosystems and species of plex of harmful species of Chloropidae in Sweden economic importance’. was analyzed by Johansson (1960). Finland Norway Some species of Chloropidae from Finland were re- Relatively little information is available on Norwe- corded by Zetterstedt (1848, 1860) based mainly on gian Chloropidae. The first records were published material of J.E. Bonsdorff. Bonsdorff (1861) listed by Zetterstedt (1838, 1848). A first list of Norwe- under “Oscinides” only species of the present ge- gian Chloropidae was published by Siebke (1872). nera Platycephala, Meromyza, Oscinis and Madiza. Later on Siebke (1877) treated the Chloropidae in Chydenius & Furuhjelm (1858–1859) recorded his Enumeratio insectorum norvegicum which in- some species. Reuter (1902a) described one species cluded 35 species belonging to 15 genera as present- of Meromyza. Frey studied Finnish Chloropidae in ly defined. New faunistic data on Norwegian Chlo- some detail, resulting in the description of 3 new ropidae were published later on by Bidenkap (1901) species (1908a, 1909, 1947), and many new faunis- from northern provinces, Strand (1903), Kolstad tic records (1908a, 1908b, 1944, 1948, 1950). Becker (1965), Greve (1983, 1999), Skrzypcznska (1987) and (1910) recorded Diplotoxoides dalmatina from Fin- Økland & Hågvar (1994). Some species from Nor- land. The rich collection of Finnish Chloropidae way were also recorded by Ardö (1957) in his re- wasidentifiedbyO.Dudaandbasedonhisre- search on marine dune ecosystems. Rygg (1966a) sults Frey (1933) published a list of 82 species in studied species of Chloropidae livings as larval 30 genera. All earlier records of Finnish Chlorop- pests on cereals and agricultural grasses in Norway idae were critically reviewed in this work. Duda and added some species new to the country. The further used the material from the Finnish collec- arthropod canopy community of Scots pine (Pinus tion in his monograph on Chloropidae in E. Lind- silvestris) in Norway was studied by Thunes et al. ner’s Die Fliegen der Paläarktischen Region (1932– (2004) resulting in 2 chloropid species new to the 1933). The monograph contains descriptions of 3 Norvegian fauna. Nartshuk (2008) published a pa- new species and several new records of Chloropi- per on Gaurax flavomaculatus (Duda) in Norway. dae from Finland. A list of Finnish Chloropidae was published again by Frey (1941) in Enumeratio Insec- torum Fenniae. New faunistic data were also pub- lished by Lindberg & Saris (1952) and Hedström Denmark (1964). Some ecological observations together with new faunistic records were done by R. Krogerus Chloropidae from Denmark were also recorded for (1932, 1960), Kontkanen (1935, 1939), H. Krogerus the first time by Zetterstedt (1848) based on mate- (1948), Luther (1946), Tiensuu (1948), and Hack- rial collected by R.C. Stæger. Some meager faunis- man (1963, 1968). Gall producing Lipara and ac- tic records are represented in papers by Kemner companying species were studied by Rable (1951) (1937), Klefbeck (1951), both investigating the en- and Valkeila (1956, 1959a). Haartman (1950) re- tomofauna of Anholt Is. Ardö’s (1957) study on corded mass occurrences Thaumatomyia notata

6 indoors. A check list of the insects of Finland, (1858). Chydenius & Furuhjelm (1858–1859) listed including Chloropidae, was regularly updated by 11 species from Karelia. Frey (1933a) published a W.E. Hellén from 1926 to 1971. Kallio (1950) stud- short list (5 species) found near Sosnovo (Met- ied species of Chloropidae associated with cereals. säpirtti) on the Karelian Isthmus and in subse- Vappula (1965), in a monograph on pests quent papers (1933b, 1941) recorded some species of cultivated plants in Finland, treated 8 species from all these territories. Duda (1933) described a of Chloropidae that to various extent damage ce- new species of Chlorops from Karelia. Chloropidae reals and cultivated grasses in Finland. Hackman from the Karelian Isthmus were recorded as a part (1980) published a check list of Finnish Diptera of studies of the Diptera fauna of the St. Petersburg with Chloropidae represented by 91 species in 36 Province by Stackelberg (1921, 1958, 1965). Addi- genera. Nartshuk (1988, 1992) revised the Finnish tional species and biological data were published species of Meromyza andaddedsomespeciesnew by Nartshuk (1962a, 1978a, 1978b, 1992). Nartshuk to the list. Later on she revised the remaining ge- (1984) studied Chloropidae of the Bol’shoi Bere- nera of the subfamily Chloropinae from Finland zovyi Is. near Primorsk. Nartshuk & Przhiboro (1998, 1999). A new check list of Finnish Chlorop- (2009) published some new data on Karelian Chlo- idae by Kaharpää is available from the website ropidae. Znamenskaya (1941, 1962) recorded Os- http://www.iki.fi/kahanpaa/diptera/list/. cinella frit from the Kola Peninsula. Zvereva (1993a, Publications on Oscinella frit, Chlorops pumilio- 1993b) and Kozlov & Zvereva (1997) studied the im- nis and other harmful species of frit flies in Finland pact of the industrial pollution in the Kola Penin- are listed in the chapter ‘Role in land ecosystems sula on the insect fauna, including Chloropidae. and species of economic importance’. A list of Chloropidae is given in the latter paper, but some records seem doubtful though. Nartshuk & Przhiboro (2009) described a new species of In- North-West European Russia certella Sabrosky from the coast of the White Sea in Karelia. The Chloropidae of the Russian Karelia, Kola Penin- As a result of the Scandinavian tradition for sula and Karelian Isthmus have never been studied taxonomic study of insects, no less than 49 valid specifically. The first records of Chloropidae from species of Chloropidae have their type localities in the Karelian Isthmus were given by Osten-Sacken Fennoscandia.

Methods of collecting, preparing and preserving

Most Chloropidae are very small to small flies. The method to collect small flies living in very low veg- adults of most species are found in the herb stra- etation. Arnold et al. (1973) described a portable tum, especially among grasses, sedges and reeds. vacuum sampler powered by a small two-stroke Some species visit flowers, especially Apiaceae. petrol engine. Malaise traps are probably advanta- Species of Gaurax, Hapleginella and Aspistyla may geous over other automated methods of collecting be found crawling on foliage of trees or bushes. flying insects including Chloropidae by day. Win- In general no special techniques are required to dow traps may also be useful, but are less effective. collect Chloropidae. Sweeping is a very produc- Chloropidae and most other Diptera are not very tive method of Chloropidae collecting. Herbage in meadows, beaches, swamps, fens, bogs and active at night. Accordingly, light traps normally marshes, hedges and clearings in forests, agricul- give poor results. Different kinds of pitfall traps tural fields, and also low twigs of trees and shrubs are very effective for wingless or brachypterous should be examined. The flies are most easily chloropids, but even collect fully winged species picked up from the net by use of an aspirator. living near the ground. Pitfall traps are especially Sweeping of wet vegetation should be avoided. Us- recommended to use in habitats with low and ing a pouter or vacuum sampler is an effective dense vegetation. Leponte (1956) collected many

7 Chloropidae from the canopy of trees by using a & Beroza, 1967). Octyl butanoate is attractive to modified umbrella. Siphonella sp. (Sugawara & Muto, 1974). Fifteen Use of brightly coloured pan traps or Moericke chloropid species, among them kleptoparasitic traps is a productive way to collect adult Chlorop- flies of the genus (predominantly females), idae (Lauva & Shutele, 1976; Barták, 1997; Kibík were found attracted to alkenas: (E)-2-Octenal, et al., 1999). Some authors found that white and (E)-Hexenal and (E)-2-Decenal (Aldrich & Bar- yellow traps are the most effective in attracting ros, 1995). Adult frit flies are attracted to oils from Oscinella and other chloropids (Koch, 1960; Tešic, sprouted seeds of cereals (Kryshtal et al., 1973). 1971; Ibbotson, 1989), others blue (Ibbotson, 1958; Zhang & Aldrich (2004) studied the attraction of Musolff, 1959; Mayer, 1961; De Bruyn, 1986) or vio- female scavenging Chloropidae to hexyl butyrate let (Oschmann, 1974, 1979). It is thus possible that and (E)-2-hexeny butyrate, both scent gland com- different species prefer different colours. Moreau pounds of plant bugs (Miridae). Their list of species (1963) found that O. frit prefers blue colour and includes species of the genera Conioscinella, La- O. pusilla yellow. Tschirnhaus (1981), who used siochaeta, Oscinella, Siphonella,andChlorops,all coloured pan traps to collect Chloropidae, consid- occurring in Europe. ered white pans to be the most efficient. Insecticidal knock-down fogging is now exten- Sticky traps are also used to collect frit flies in sively used to collect flies and other insects in the agricultural fields. Ibbotson (1958) used panes of forest canopy. The method proved a fairly effective glass coated with an adhesive. Kharchenko & Burov method for Chloropidae in tropical rainforest (Flo- (1990) received a good result with coloured sticky ren, 2003). Also canopy fogging of pine trees has traps to study the flight dynamics of flies in corn been practiced with interesting results in Norway fields. Nielsen (1985) used blue adhesive traps to (Thunes et al., 2004). We predict that this method study flight behavior of Oscinella spp. Roháček et is most efficient for species of Gaurax, Hapleginella al. (1998) found that sweeping was the most pro- and Lasiambia. Using cap traps in the canopy of ductive method to collect Chloropidae. They com- trees in Warsaw parks (Poland) gave a new species pared sweeping, yellow pan traps and Malaise traps of the genus Gaurax (Nartshuk, 1980). as alternative methods of collecting flies in a peat- Wilbur & Fritz (1939), Southwood & Jepson bog. Comparison of different methods of collect- (1962), Emden (1963) and Clements (1980, 1982) ing has been made in more detail for Agromyzidae described several methods for collecting inverte- (Scheirs et al., 1997) than for Chloropidae. brates from grasslands, where most Chloropidae It is known that various natural as well as arti- live, including methods for collecting frit fly lar- ficial odourous substances attract chloropid flies, vae and rearing adults from grasses. Chloropidae but chemical baits are nevertheless rarely used for occurring in leaf litter, debris from birds’ nests, collecting purposes. Both sexes of species of Os- mammals’ borrows etc. are preferably extracted by cinella, Conioscinella, Olcella, , Siphonella placing the substrates in a box or dark plastic bag and females of Tricimba are attracted to 2,4- allowing exit to a light transparent vessel or bag. Al- hexadienyl butyrate, 2,4-hexadienyl propionate, ternative methods of collecting Chloropidae from heptyl and hexenyl butyrate, coproic and valeric grasslands, the preferred habitat of most Chloropi- acids (Fluno et al., 1972; Rogoff et al., 1973; Aldrich dae, such as emergence traps and vacuum suckers & Barros, 1996; Hibbard et al., 1997; Zwang & are described by Clements (1980, 1982). Aldrich, 2004) and Hippelates also to trimetil amine Larvae of many phytophagous chloropids can ammonia acetic acid (Hwang et al., 1976), oleic be found by dissecting stems of grasses, sedges, and linoleic acids (Hwang & Mulla, 1971). Males reeds and other plants. Mature larvae should be of Thaumatomyia glabra are attracted to methyl allowed to stay within their host plant for pupa- anthranilate (Landolt et al., 2000; James, 2005). tion. Most flies emerge from puparia in about two Species belonging to Lasiochaeta, Oscinella, Chlo- weeks, but some species (Lipara, Dicraeus) hiber- rops and Eutropha came to baits of pyrrolizine alka- nate and appear only the following spring. Adults loid and ingested these secondary plant chemicals of Dicraeus preferably emerge after the puparia (Boppre & Pitkin, 1988). Only females of Tricimba have been in contact with water. Some authors cincta seem attracted to trans-5 dodecenol (Nye, 1959; Clements, 1982) applied irritants, oth- (Myrsina et al., 1977; Sybchev & Myrsina, 1977; ers (Goodhue & Emden, 1960) X-rays to accelerate Verba et al., 1991). Different caproic acids and the emergence of flies, but these methods appear ethyl caproate attract some Chloropidae (Jantz to be of limited value.

8 If Chloropidae are collected dry it is preferable microvial with a stopper that can be mounted on to pin them when still soft after having been killed. the same pin as the fly. This method was described Double mounting is usually used for small Chlorop- by H. Andersson (1972) and Nartshuk (1975). idae. Taxonomic characters are easier to observe Some light-coloured species of Chloropinae have in dry-mounted flies than in fluid-preserved spec- weakly sclerotized male genitalia, especially the imens. Flies that have dried out may be glued onto postgonites and aedeagus. Some methods of stain- cardboard triangles. Alternatively, dry flies must be ing are recommended to better examine transpar- softened in a high humidity relaxing chamber be- ent parts. Fairly good results have been attained fore being pinned. Specimens stored in tubes with with a slightly modified version of the method de- 80% ethanol must be kept in darkness to prevent scribed by Wilkey (1962). The cut tip of abdomen is bleaching of body colours. after treatment in 10% KOH solution transferred to Study of the male genitalia is needed for a safe Essig’s Fluid (EAF) double stained with lignin pink determination of many species of Chloropidae. The and acid fuchsine for 1–2 hours and then trans- tip of the abdomen is removed and put in a 10% so- ferred to 96% alcohol. Stained parts are also stored lution of potassium hydroxide (KOH) and left there in glycerol. from 1–12 hours until the soft parts have been dis- Chloropid larvae, like those of other Diptera, pre- solved. The process may be accelerated to a few serve well in 80% ethanol. They are best killed minutes by heating the solution in a water bath. instantly in hot water beforehand, as this may After careful rinsing in destilled water to remove prevent contraction and retain their natural body all traces of KOH, the genital structures are ready shape. The process of making permanent slide for dissection and examination in alcohol or glyc- preparations of larvae is described by Nartshuk erol. For permanent storage, it is best to place the (1956), but today SEM techniques have substan- dissected parts in glycerol in a sealed microtupe or tially improved the study of the larvae.

Morphology, diagnostic characters and anatomy

Adults Head (Figs 8–17). The shape of the head is very dif- ferent between genera. In lateral view the “typical” head is rounded or square and the frons and vib- Adult chloropids of NW Europe a very small to rissal angle project beyond the eyes to the same medium sized (1.0–8.0 mm) “typical” flies. Habit- extent. In many cases, however, the frons or the ually, here are two main types. One type (Figs 1, 6, vibrissal angle (see below) projects far beyond the 7b), exemplified by Oscinella and Chlorops species, eyes. In species with a strongly produced frons as has a compact body with a large, broad head and in some Chloropinae the head appears rather trian- deepthorax,shortlegs,andashort,broadandflat- gular in lateral view. A strongly produced vibrissal tened abdomen and rather short, broad wings kept angle is more often seen in species of Oscinellinae. flat over the abdomen at rest. The other type, ex- The profile shape of the head is of great diagnostic emplified by Meromyza and Platycephala (Figs 5, value in many groups. A detailed study of the head 7a), has a slender body, a long and narrow head, and mouth parts of Chlorops pumilionis was made narrow thorax and a narrow, rather cylindrical ab- by Frew (1923b). domen. Especially the hind legs have in several As is typical for the chloropids have groups swollen femora and curved tibiae (Fig. 5). a well developed ptilinum.Theptilinal suture is Wingless (apterous) or short-winged (brachypter- sometimes rather incompletely closed. The frons ous) species or forms (Fig. 2) are relatively rare in is broad in both sexes and weakly sclerotized, so the North European fauna. The body colour varies that the head of newly emerged specimens tends from black over yellow, reddish or greenish, often to become deformed when dry-mounted. The ocel- with black or red stripes and marks and is shining lar triangle (frontal triangle) is usually well sclero- or more or less extensively dulled by greyish dust- tized and prominent (Figs 8, 10). Sometimes it is ing. so large that it covers the entire frons. The shape,

9 Figs 1–7b. Adult frit flies: 1: Calamoncosis glyceriae Nartshuk. 2: Elachiptera brevipennis (Meigen). 3: Gampsocera numerata (Heeger). 4: Lipara lucens Meigen. 5: Meromyza pratorum Meigen. 6: Oscinella maura (Fallén). 7a: Platycephala planifrons (Fabricius). 7b: Thau- matomyia notata (Meigen). Credit: Dmitry Gavryushin, 1–3, 7a; Tom Murray, 4; Jens Hyldstrup Larsen, 5; Klaas van der Veen, 6; Jarmo Holopainen, 7b.

10 Figs 8–17. Adult frit flies: 8: head, dorsal view of Pseudopachychaeta sp. 9: head, lateral view of Oscinella sp. 10: head, anterior view of Lipara sp. 11–14: head, lateral view and face and antennae, anterior view: 11, 12: Aphanotrigonum trilineatum (Meigen). 13, 14: Siphonella oscinina (Fallén). 15, 16: head, anterior view: 15: Polyodaspis rufijicornis (Macquart). 16: Lasiambia sp. 17: head, lateral view of Melanum laterale (Haliday) sp. (ant = antennomere; fl = flagellomere; setation:fr= frontal; if = interfrontal; oc = ocellar; orb = orbital; pooc = postocellar; pst = peristomal; vte = outer vertical; vti = inner vertical.)

11 colour, sculpture and pilosity of the ocellar trian- against a corresponding surface on the proepister- gle are of immense taxonomic value. Normally the num (cf. Chlorops pumilionis). The lower anterior ocellar triangle is without setulae except for the corner of the gena forms the vibrissal angle with ocellar pair, but sometimes there are one or two the vibrissal seta (vi). The vibrissal angle is more or rows along the side margins, or, rarely, setulae cover less projecting, right-angled or retreating. The area most of the surface, arising from small punctures. below the ptilinal suture and delimited laterally by The setae (bristles) of the head are sometimes quite the facial ridges is the face. Between the face and indistinct. Most of those mentioned below can usu- the anterior margin of the eye is the parafacial plate ally be distinguished though they are often small. (facial orbit, cheek). Above the insertion of the an- The frontal setae (fr) are more or less numerous tennae and separated from frons by the ptilinal su- on the frons. Along the margins of the ocellar tri- ture, is a small triangular sclerite, the lunule,that angle and curved up over it are the interfrontal se- mid-ventrally continues into the facial carina (fa- tae (if ). They can also be arranged on the surface cial ridge) between the antennae. The facial carina along the margins or be dispersed over the sur- is usually narrow, but broad in the genus Lipara. face. Along the eye margins there are one or more The facial margin is the lower limit of the face just recurved (reclinate), outcurved or procurved (pro- above the clypeus. clinate) orbital setae (orb). They cannot as a rule The antennae (Figs 18–24) comprise four parts: be classified as upper fronto-orbital setae (ors)or The scape (first antennal segment, ant1)isinmost lower fronto-orbital setae (ori) and they are here species very short. The pedicel (second antennal simply called orbital setae (orb). The orbital setae segment, ant2) is more variable in length. The fijirst are usually numerous (6–10) and rather small and flagellomere (third antennal segment, ant3), also even in size, but in some genera in both subfam- known as the postpedicel and though to repre- ilies one or two (rarely 3) setae are longer than sent a fusion of several flagellomeres (Stuckenberg, the remaining ones. On the ocellar triangle there 1999) shows great diversity: rounded, elongate, are three ocelli on an ocellar tubercle and two pro- trapezoid, etc. Finally, the arista is built of three clinate or reclinate, convergent or divergent ocel- distal flagellomeres, but the basal aristal segment is lar setae (oc). The direction of the ocellar setae very short and barely visible. The arista is very vari- has great taxonomic importance: in Oscinellinae able: slender or broad and flattened, nearly bare, they are straight or recurved and convergent; in pubescent or densely pubescent. Antennal char- Elachiptera they are more or less procurved but acters are of great diagnostic value in chloropids, convergent; and in Chloropinae and Rhodesielli- both for genera and for species. Andersson (1979) nae the ocellar setae are always procurved and di- noticed a distinct correlation in shape between the vergent. first flagellomere and the arista in different lineages On the vertex there are one outcurved outer ver- of the family. tical seta (vte) and one incurved inner vertical seta The proboscis (Figs 25–30) is usually soft and (vti) on each side. In the middle, behind the ocelli, short with round or oval labella,butratherfre- is a pair of postocellar setae (poc) (postvertical se- quently it is more sclerotized and lengthened, tae). The postocellar setae are usually convergent and with long and narrow labella angled back- or cruciate, except in some genera of Oscinellinae wards. A long proboscis is strongly correlated with (Lasiambia, Polyodaspis) that have straight posto- a strongly produced vibrissal angle. The structure cellar setae. Below the latter there is a well defined, of the proboscis was studied by Graham-Smith often dark coloured occipital fijield (occ) forming the (1930) in some non-Palearctic chloropids and by upper part of the occiput. Kulikova & Nartshuk (1991) in 25 chiefly Palearc- The compound eyes are large, circular or oval tic species from 20 genera. The labella have usually with a vertical, horizontal or diagonal long axis. All three pairs of pseudotracheae,butthereareonly facettes (ommatidia) are equal in size or the ante- two pairs in the slender labella of Siphonella oscin- rior and anterodorsal facettes are slightly enlarged. ina and up to 7 pairs in Lipara. Zaitzev (1992) gave The eyes are naked or covered with fine hairs. Be- a SEM picture of the labella of Siphonella oscinina low the eyes are the genae (jowls) that vary in width showing a rasping devise formed by pseudotra- from narrow to broad. The gena bears peristomalse- cheae. Finer or stronger setae are sometimes stand- tae (pst) and its broader hind part, the postgena,is ing between the pseudotracheae, e.g. in species of sometimes delimited by a low carina. The posterior Lasiambia (Tschirnhaus, 1992). Shcherbina (1970) surface of the postgena is sometimes flat and slides found on the labella of Meromyza saltatrix 86 sup-

12 Figs 18–24. Adult frit flies, antenna: 18: Hapleginella laevifrons (Haliday). 19: Epichlorops puncticollis (Zetterstedt). 20: Parectecephala longicornis (Fallén). 21: Oscinella capreolus (Haliday). 22: Elachiptera cornuta (Fallén). 23: Gampsocera numerata (Heeger). 24: Centori- soma elegantulum Becker. posed chemoreceptorial sensilla of two types dis- ous macrochaetae, that may be strong (setae)or tinguished by length. Elzinga & Broce (1986) made fine (setulae). The scutum can be bare or “hairy”, SEM studies of the labella of several species of with setulae evenly distributed or arranged in rows. Chloropidae and found that the pseudotracheae The rows of setae found on scutum are the dor- radiated individually from the prestomal opening socentrals (dc)andacrostichals (acr). Usually they at the base of the laella. They also described differ- are reduced and setulose except for the posterior ent types of rasping devices formed by the pseu- pair of dorsocentral before scutellum. Other scutal dotracheal rings and concluded that conventional setae are the anterior and posterior postalar setae prestomal teeth are not developed. The structure (apaand ppa)andprescutellar setae (psc). Species of proboscis indicates that adult chloropids feed with a better developed scutal chaetotaxy exist in on liquefied food. other parts of the world. The scutum is unicolorous, The palpi are of medium size, slightly upcurved, usually black, or striped with light ground colour cylindrical or flattened, with short setae. The palpi (yellow, green, brown) and five darker longitudi- are sometimes sexually dimorphic, being longer nal stripes (black, brown, red). The middle stripe and stouter in the males. (M1) usually ends distinctly anterior of the scutel- lum and can be divided by a narrow streak along Thorax. The thorax (Figs 31–34) is usually short and the acrostichal line. Laterally of M1 the inner lateral broad with a rather convex scutum (mesonotum), stripes (M2) continue from the postpronotal lobes but sometimes long and slender and flattened dor- to the dc. They are normally broadest in front. Lat- sally. The scutum is to varying extent shining or erally of M2 and behind the notopleural incisions mat (if covered in dustlike pruinescence), smooth are the outer lateral stripes (M3). In dark coloured or sculptured. On the thorax there are also numer- specimens of striped species M1 may extend to the

13 Figs 25–30. Adult frit flies, proboscis: 25: Oscinella frit (Linnaeus). 26: Siphunculina aenea (Macquart). 27: Siphonella oscinina (Fallén). 28: oral disc of Siphonella oscinina (Fallén). 29: Aphanotrigonum nigripes (Zetterstedt). 30: Meromyza nigriseta Fedoseeva.

14 Figs 31–34. Adult frit flies, thorax: 31: lateral view. 32: dorsal view. 33: dorsal view of Elachiptera cornuta (Fallén) with setulae in rows. 34: dorsal view with setulae evenly scattered. (a = anterior; acr = acrostichal; ap sc = apical scutellar; cal = callus; dc = dorsocentral; h = humeral; H = humeral spot; la sc = lateral scutellar; M1,2,3 = central, inner, and outer scutal stripes; npl = notopleural; p = posterior; pa = postalar; prpl = propleuron; psc = prescutellar; sc = scutellar; scp = scapular; sp = spiracle.)

15 scutellum or the dark stripes may be more or less near the humeral cross-vein (h) or as an indistinct confluent. weak fold along the first vein. The first three lon- The shape of the scutellum is quite variable: short gitudinal veins compose the radial vein (R1,R2+3, and semicircular to greatly prolonged and triangu- R4+5). The fourth vein is the medial vein (M). The lar, and dorsally convex or flattened. On the scutel- fifth vein is composed by a cubital vein and the lum there are setae along its margin. They can fijirst anal vein (CuA1). In the posterior basal part of be inserted on small or long, finger-like tubercles. the wing the veins are reduced or absent. The fijirst Apairofapical scutellar setae (ap sc) is usually well or anterior basal cell (br)isclosedbytheanterior developed while the lateral scutellar setae (la sc) cross-vein (r-m, ta). The second basal cell (bm)is are normally smaller but consisting of several pairs. fused with the discal cell (dm) because the cross- The postpronotal lobes (humeral calli) (h) form vein (bm-cu) between them is absent. Its former the anterodorsal corners of the thorax and are usu- place can usually be traced by a week flexure in the ally equipped with a dark postpronotal spot (H) middle of vein CuA1. The fused cell is terminated by in light coloured (yellow, brown, greenish) species the posterior cross-vein (dm-cu, tp). The anal veins and one (or more) postpronotal setae (h). The no- (A1,A2) are very reduced or absent. In most cases topleura (npl) form triangular depressions at the the alula is well developed and there is a small alar sides of the scutum. They have an anterior noto- calypter.Thehalteres are of medium size with a tri- pleural seta (anpl) and one or two (rarely more) angular knob coloured white, yellow or black. posterior notopleural setae (pnpl). A transverse su- Legs. The front, middle and hind legs are usually ture is not developed on scutum but indicated by simple and relatively short, composed of a coxa the notopleural incisions. (coxa1–coxa3), a trochanter,afemur (f1–f3), a tibia The sides of the thorax are made up by the pleu- (t1–t3) and a 5-segmented tarsus. The hind femora ral sclerites. These can be setose or bare, shining are in some species thickened with knobs or spines or dusted. The proepisternum (prep) (propleuron) below and correlated with this the hind tibiae in front has one or a few proepisternal setae (prep) are curved. In some species of Eribolus all femora below. A vertical proepisternal carina is very char- are enlarged, more strongly so in the males. The acteristic for chloropids but may sometimes be dif- legs are covered with fine setulae but distinct se- ficult to see without removing the head. Beneath tae are not present except for a ventral spur on the notopleuron is the anepisternum (anep)(meso- middle tibiae, sometimes even on hind tibiae. In pleuron), often with one or two anepisternal spots. many groups, but most commonly Oscinellinae, In other than completely black species the katepis- a so-called tibial organ (Fig. 45) is developed on ternum (kep) (sternopleuron) has a large katepis- the posterodorsal side of the hind tibiae in both ternal spot, of which the colour and dusting are sexes in shape of an elongate oval area densely co- of great importance. The sclerite above the hind vered with short and fine hairs. Its function is not coxa is the katepimeron (hypopleuron) and above known. The males of many species of Oscinellinae it and below the wing the anepimeron (pteropleu- also posses a femoral organ (Figs 46–61) on middle ron). They can both have taxonomically important femora consisting of a group or row of small warts spots. A precoxal bridge of T-form according to the each with a short or spine-like setula. Its function classification of Speight (1969) is developed be- is not known as well. The only species of Rhode- tween prosternum and the propleura. Below the siellinae (Aspistyla plumiger)foundinNWEurope scutellum the thorax is terminated by the medioter- possesses in both sexes femoral organs on middle gite (postnotum). Between the postpronotal lobe andhindfemora(Figs46,47). and anepisternum lies the mesothoracic anterior Abdomen (Figs 62, 63). The abdomen is ovoid, spiracle (sp1), and beneath the halter the posterior spiracle (sp ). oval or slender, cylindrical or flattened. The preab- 2 domen, which can be seen from above, is composed Wings (Figs 35–44). The costal vein (vein C) contin- of five segments. The tergites (T) 1 + 2, 3, 4, and 5 ues to the third or fourth vein and has a subcostal are well-developed, their lateral parts are usually break near at the end of the first vein. The costal turned down and partly covering the ventral side segments are denoted by mg1,mg2 etc. At the base of the abdomen, only leaving narrow membranous of vein C is the basicosta, and before that, nearest at strips between their margins and the narrow ster- the thorax, a triangular sclerite, the tegula.Thesub- nites. Only in Aphanotrigonum the tergites cover costal vein (Sc) is retained only for a short distance most of the ventral abdomen and have distinct

16 Figs 35–44. Adult frit flies, wing: 35: species of Oscinellinae. 36: Hapleginella laevifrons (Haliday). 37: Dicraeus sp. 38: Gampsocera nu- merata (Heeger). 39: Siphunculina aenea (Macquart). 40: Conioscinella zetterstedti Andersson. 41: Chlorops sp. 42: Camarota curvipennis (Latreille). 43: Diplotoxa sp. 44: Pseudopachychaeta sp. (40 after Brauns, 1939.) (mg1,2,3,4 = 1st to 4th costal segments.)

17 Figs 45–61. Adult frit flies, leg organs: 45: hind tibial organ of Oscinella frit (Linnaeus).46,47:femoralorganofmaleandfemaleAs- pistyla plumiger (Meigen).48–61:malefemoralorgan:48:Aphanotrigonum trilineatum (Meigen). 49: Calamoncosis minima (Strobl). 50: Eribolus nana (Zetterstedt). 51: (Meigen). 52: Dicraeus styriacus (Strobl). 53: Hapleginella laevifrons (Loew). 54: Incertella albipalpis (Meigen). 55: Oscinella frit (Linnaeus). 56a, 56b: Oscinimorpha minutissima (Strobl). 57: Oscinisoma cognatum (Meigen). 58: Elachiptera cornuta (Fallén). 59: Siphonella oscinina (Fallén). 60: Trachysiphonella scutellata (Roser). 61: Tricimba cincta (Meigen).

18 Figs 62–70. Adult frit flies, abdominal parts: 62: male abdomen, ventral view of Elachiptera sp. 63: female abdomen, ventral view of Oscinella sp. 64, 65: male abdomen, lateral view, with pregenital sclerite present (Oscinellinae) and absent (Chloropinae). 66: male abdominal vesiculae of Elachiptera brevipennis (Meigen). 67: male abdomen of Polyodaspis sulcicollis (Meigen) with vesicula, lateral view. 68–70: tip of male abdomen of with vesiculae, ventral view: 68: Lasiambia coxalis (Roser). 69: Thaumatomyia glabra (Meigen). 70: T. notata (Meigen). (68 after Hennig, 1941.) (ep = epandrium; hyp = hypandrium; psc = pregenital sclerite; sp = spiracle; s = sternite; t = tergite; v = vesicula.) ridges between their dorsal and ventral parts. In in this part of abdomen in some genera (Lasiambia, Elachiptera brevipennis tergites 1 + 2 are enlarged Polyodaspis, Thaumatomyia in NW Europe) (Figs and occupy nearly half of the abdomen. Sternites 66–70). The vesiculae in Thaumatomyia contain 1–5 (S1–S5) are usually small, but sometimes larger pheromon glands and also constitute a strong opti- or divided. Spiracles 1–5 (sp1–sp5) are placed in the cal signal (Kortba, 2009). membrane near lateral margins of the tergites. Hypopygium (Figs 71–82). The epandrium (ep) is In the male postabdomen there is only one scle- considered to represent tergite 9. In caudal view rite between tergite 5 and the epandrium. This scle- it is horseshoe-shaped, encircling the cerci in most rite is thought to represent the fused sternites 7–8. Oscinellinae or the movable forceps or surstyli (sur) It bears two approximated spiracles on either side in most Chloropinae. In Oscinellinae the movable (Fig. 64). In some Chloropinae is this sclerite is surstyli are usually simple convex sclerites. They very reduced or practically absent (Fig. 65). Ev- are more complex with processes of different shape ersible membranous vesiculae may be developed only in species of Gaurax. In most Chloropinae

19 Figs 71–75. Adult frit flies, male genitalia of Oscinellinae: 71: epandrium confluent fused cerci (Conioscinella sp.). 72: epandrium with separated narrow cerci (Dicraeus sp.). 73: epandrium with fused cerci (Lipara pullitarsis Doskočil & Chvála). 74: open hypandrium and phallic complex. 75: closed hypandrium and phallic complex. (73 after Chvála et al., 1974.) thesurstyliarequitecomplex,andsomegroups flat or more or less cup-shaped sclerite, the phal- built up by parts presently named the posterior, lapodemic sclerite (ph s) that usually reaches the median and anterior lobes of the surstyli. In some hypandrium anteriorly. The free end of the phallus Chloropinae, e.g. Lasiosina, the surstyli are firmly is the usually membranous distiphallus (dph). united with the epandrium and very likely im- The paired postgonites (pog) and pregonites (prg) movable. Sometimes the epandrium have addi- are of different shapes and standing in different tional processes, e.g. in Cetema. Encircled by the positions in different genera. They are weakly scle- epandrium is the supraanal lobe,thesubanal lobe, rotized except in most species of Meromyza (Fig. the anus and the cerci. The two first-mentioned 79). In Oscinellinae and most Chloropinae the are usually rather indistinct but their presence pre- and postgonites are arranged in a line, but is indicated by groups of small setae. The cerci in most small Oscinellinae the division between can be free or, especially in Chloropinae fused to- these structures is not visible. In some Chloropinae gether in a mesolobus (mes). A pair of sclerites is (Chlorops and related genera) the pregonites stand sometimes flanking the subanal lobe, but not in laterad of the postgonites. Rarely in Oscinellinae species from our region. The epandrium and its (Gaurax) the hypandrium and gonites are fused to- appendages form the caudal wall of the genital gether. In Chloropinae (Diplotoxa)thegonitesare capsule. When at rest the epandrium and its ap- seemingly fused together with phallus. The phal- pendages move downwards and forwards to cover lus is basally entered by a duct from the ejaculatory the genital pouch. The genital pouch is also delim- bulb with its minute ejaculatory apodeme (ej ap). ited by the semicircular hypandrium (hyp). Cau- Female postabdomen (Figs 83, 84). In the female dally the hypandrium articulates with the epan- postabdomen segments 6–8 are variously mod- drium and with ventral bases of the surstyli or a ified to form the ovipositor. Sternites 6–8 tend pair of ventral epandrial sclerites (ep s). The hypan- to be partly membranized and divided in pieces drium is in many Oscinellinae closed behind and and this sometimes even happens to tergites 6–8. forming a ring (Figs 74, 75). The inner copulatory The ovipositor is terminated by the cerci and the organs are the phallus and the gonites.Thebasal supranal and subanal plates. The cerci are usually end of the phallus, the basiphallus (bph) articu- soft and free but may be sclerotized and laterally lates with the posterior end of the phallapodeme. compressed or even joined into a flat, blade-like Anterior of the phallus the phallapodeme forms a structure.

20 Figs 76–82. Adult frit flies, male genitalia of Chloropinae: 76: posterior view. 77: ventral view, pre- and postgonites parallel (Chlorops sp.). 78: ventral view, pre- and postgonites in line (Cetema sp.). 79: hypandrium and phallic complex with sclerotized postgonites (Meromyza saltatrix (Linnaeus)). 80: epandrium and close-set surstyli (Thaumatomyia sp.). 81: epandrium and surstyli fused together (Lasiosina sp.). 82: epandrium with paralobes behind surstyli (Cetema sp.). (ep = epandrium; hyp = hypandrium; mes = mesolobus (fused cerci); ph s = phallapodemic sclerite; pog = postgonite; prg = pregonite; sur = surstylus.)

21 Figs 83–88. Adult frit flies male and female genitalia: 83: end of ovipositor, dorsal view (Cetema sp.). 84: end of ovipositor, lateral view. 85: inner male genitalia of Oscinellinae (Oscinella frit (Linnaeus)). 86: inner male genitalia of Chloropinae (Thaumatomyia notata (Meigen)). 87: inner genitalia with ripe eggs in all ovarian tubes (Lipara lucens Meigen). 88: ovarian tube with three ripe eggs (Dicraeus sp.). (ag = accessory gland; egt = egg tube; spt = spermatheca.)

Preimaginal stages Hennig (1943), Nartshuk (1956, 1962b, 1981, 1987), Nye (1958), Zhabinskaya (1963), Fedoseeva (1966), The life stages of Chloropidae consist of egg, three Chvála et al. (1974), Ismay (1998), Grochowska larval instars, pupa and imago. Viviparous species (1994, 2002, 2006, 2006a, 2007, 2008a, 2008b, are known from the tropics only. The preimagi- 2008c). All these papers include NW European nal stages of chloropids are poorly known com- species and may be useful. A general review of the pared to the imaginal stage. More or less detailed preimaginal stages is given by Ferrar (1987). descriptions of all the immature stages are only Unfortunately, it is not possible to give a diag- available for the economically important species nosis that will separate all larvae of Chloropidae Oscinella frit and Chlorops pumilionis. Publications from larvae of all other acalyptrate families. Like dealing with the immatures of other species also other schizophorous larvae chloropid larvae are exist. Some of them only with descriptions, oth- apodous, acephalic and of milky-white, pale-yellow ers include keys to genera or to species: Bala- or greenish colour. In family level keys to Diptera chowsky & Mesnil (1935), d’Aguillar (1943, 1953), larvae (e.g., Hennig, 1952; Tesky, 1981; Ferrar, 1987;

22 Figs 89–91. Frit flies, egg and egg structures: 89: Dicraeusfennicus Duda. 90a, 90b: egg outline and surface sculpture of Chloropsplanifrons (Loew). 91: part of egg with micropyle of Elachiptera tuberculifera (Corti), scale bar 0.05 mm (after Gaponov, 2000).

Smith, 1989; Smith & Ferrar, 2000) chloropid lar- producing Lipara species: up to 1.69 mm long vae are split into groups according to their biology in L. lucens (length:width ratio about 5:1); up to (plant feeders, predators of root aphids etc.) rather 1.26 mm long in L. pullitarsis (length:width ra- than based on morphological characters. tio 13:1) according to Chvála et al. (1974). Phy- tophagous species of the subfamily Chloropinae Eggs (Figs 89–91) have also rather big eggs: Meromyza 0.75 mm, Cetema 0.80 mm, Chlorops up to 1.5 mm, Lasiosina Eggs of Chloropidae are insufficiently studied. De- 0.90 mm, Platycephala 1.4–1.6 mm. scriptions and illustrations mainly exist for a few The colour of the eggs is usually white or yel- economically important species based on light mi- lowish, but pale brown in some Elachiptera and croscopic techniques. Most of the data were sum- Thaumatomyia, and blackish in Lipara pullitar- marized by Ferrar (1987). More recently Gaponov sis and L. rufijitarsis. The surface structure of the (2000, 2003) used SEM techniques in studies of the rigid, protective exochorion is complex as in many surface structure of eggs from 12 species in 9 genera other Diptera. It is usually formed by longitudi- of Chloropidae. nal parallel, sometimes sinuous ridges and grooves Eggs are elongate oval, cylindrical, straight to with or without transverse ribs between them. slightly curved, tapering at both poles or only to- The ridges often anastomose, especially at the an- wards the anterior pole, where the micropyle is lo- terior third of the egg. The surface between the cated. The upper side is usually convex, the side fac- ridges may be smooth, rough or with low trans- ing the substrate flattened or even slightly concave. verse ridges, forming a regular pattern between the Some eggs appear asymmetrical (Oscinella). Egg size varies from 0.24–1.7 mm in length and 0.15– longitudinal ones (Cetema, Platycephala). On top 0.39 mm in width. Index (width/length) varies of these patterns may be found rows of tubercles from 0.23–0.63. Carnivorous flies of the genus (Chlorops fasciatus) or fine pits (Elachiptera tuber- Thaumatomyia have the shortest eggs: 0.24 mm culifera, Thaumatomyia glabra). The exochorion in T. notate,0.27mminT. glabra. Saprophagous of some species has a cellular or polygonal struc- species of the genus Elachiptera have eggs 0.5– ture (some Chlorops). Sometimes the exochorion is 0.8 mm in length, eggs of Oscinella frit 0.6–0.8 mm relatively smooth, thin elastic with delicate striae long. The biggest eggs are known from the gall or small pits (Gaurax dubius). Specialized gall-

23 forming species of the genus Lipara have a very thin The slightly bilobate cephalic segment (Figs 93, and weakly sculptured chorion. 94) is the smallest segment of the larval body. From The micropyle at the anterior pole of the egg is it arise interiorly or laterally the short 2-segmented, usually formed on small conical tubercle in the usually unsclerotized antennae. The apical seg- middle of an apical depression (Gaurax, Elachip- ment is sometimes completely retracted within the tera, Conioscinella, Cetema, Chlorops, Meromyza)or basal antennal segment. Behind the antenna is a on a plug-like protrusion (Oscinella, Lasiosina). The sensory organ consisting of several papillae sur- micropylar zone changes in fertilized eggs from rounded by a more or less sclerotized ring being conical to rosette-shaped. An aeropylar zone con- completely closed or partly open anteriorly. Be- sisting of a system of rather large respiratory crypts hind this sensory organ lies another sensory organ has been found in some genera. It usually lies on consisting of 1–3 papillae named in some papers the anterior egg pole around the micropyle. The as frontal palpi. These two sensory organs, previ- number of crypts varies over a wide limit even ously identified as maxillary and frontal palpi re- within a genus. Eggs of Meromyza nigriventris and spectively, are innervated from the maxillary nerve, M. ornata have 5–8 crypts. Eggs of Gaurax has and both are now named as maxillary papillae. 12–16 crypts not arranged within aeropylar zone. Immediately posterior to the mouth opening is Eggs of Thaumatomyia glabra has 9–12 large crypts a small sclerite, the labial plate. Ventral surface with net structure inside. Some additional aeropy- of the cephalic segment is occupied by the facial lar crypts may lie in the rows of the main surface mask composed of oral (cephalic) ridges being hya- of the egg (Cetema). The apical area with crypts line cuticular thickenings. The structure of the fa- may be separated from the rest of chorion by a cial mask varies considerably between genera and circular ridge (Thaumatomyia). No aeropylar zone species. The oral ridges usually both furcate and was found in eggs of Elachiptera, Oscinella, Co- make anastomoses and thus develop cephalic cells nioscinella, Lipara and Lasiosina. that may have serrate or dentate margins. Singu- lar cephalic papillae are situated between the oral ridges. The oral ridge are used to channel food into Larvae (Figs 92–100, 956–1014) the mouth, but are absent in the carnivorous larvae of Thaumatomyia. Usually the third instar of larvae have the most The complex cephalopharyngeal skeleton (Figs fully differentiated species characteristics. There- 95–97) consists of paired sclerites. At the anterior fore, the following account applies to fully grown end two black mouth hooks (mandibles) may be final instar larvae. exposed from the mouth, while the remaining parts Larval body (Fig. 92) cylindrical, elongated, 2– of the cephalopharyngeal skeleton are contained 10 mm long (biggest in Lipara species) and 0.5– in the first segments of the body. The apical tooth 3.0 mm in diameter. Ratio length:breadth varies of the mandibles is particularly in phytophagous normally from 6:1 (in Chlorops species) to 10:1 (in larvae followed by one to several smaller acces- Oscinella, Meromyza and Cetema), only 4:1 in Li- sory teeth arranged in one or in two rows. Imme- para. Larvae are generally white or yellowish, but diately below the mandibles are a pair of small greenish in Meromyza. The larval cuticle is nor- crescent-shaped dentate sclerites, that in some mally colourless, except with black pigmentation species (Meromyza,someLasiosina)arefusedto- in both ends of Lipara larvae and anteriorly in lar- gether and forming arch. The dentate sclerites are vae of Platycephala planifrons. The larval body is absent in Chlorops. Some additional sclerites are more or less transparent depending on the amount situated near the posterior end of the mandible in of fat bodies with green or brown contents of the some species. The mandibles articulate with the gut and yellow to brown malpighian tubules of- hypopharyngeal (hypostomal, intermediate) scle- ten well visible through the cuticle. Body usually rites that are joined ventrally by a hypopharyngial distinctly tapering anteriorly and bluntly rounded arch. In ventral view hypopharyngeal sclerites form posteriorly, in many species with two caudal pro- a H-shaped sclerite. Hypopharyngeal sclerites ar- cesses bearing the posterior spiracles. The larval ticulate or fused with the tentoropharyngeal (pha- body of gall producing species as Lipara is more ryngeal) sclerites that at their posterior end are stocky and without caudal processes. The cephalic divided in dorsal and ventral cornua (wings). The and following 11 segments are demarcated by slight two ventral cornua are fused on each side with the constrictions of the body. pharynx. Ventral pharyngeal ridges are present in

24 Figs 92–100. Frit flies, larvae and larval structures: 92: larva of Oscinella sp., lateral view. 93: facial mask of 3rd instar larva of Polyodaspis rufijicornis (Macquart) with parallel oral ridges. 94: facial mask of 3rd instar larva of Calamoncosis glyceria Nartshuk with oral ridges forming cells. 95: cephalopharyngeal skeleton of 1st instar larva of Oscinella frit (Linnaeus). 96a, 96b: cephalopharyngeal skeleton of 3rd instar larva of Oscinella frit (Linnaeus). 97: cephalopharyngeal skeleton of 3rd instar larva of Meromyza sp. 98: anterior spiracle of fan type (Oscinella sp.). 99: anterior spiracle of small transverse rosette type (Meromyza sp.). 100: puparium of Oscinella sp. (cph = cephalopharyngeal sclerite; ds = dental sclerite; hys = hypostomal sclerite; md = mandibula; pst = parastomal sclerite.)

25 many species but absent in some phytophagous posterior margin of the thoracic segments but not species (Chlorops, Platycephala). The tentoropha- ventrally. The eight abdominal segments all have ryngeal sclerites are joined anterodorsally by a dor- ventral creeping welts that are usually covered with sal bridge (dorsal sclerite, dorsal arch) in most spicules.Somelarvae(Meromyza) have no spicules, species. A pair of narrow sclerites, the parastomal but many rows of fine striae instead. Some larvae bars (atrial rods) project from the anterior mar- (e.g. Platycephala planifrons) have spicular zones gin of the tentoropharyngeal sclerites above the on the dorsal side of the abdominal segments as hypopharyngeal sclerites. Sometimes (Meromyza well, and others (Lipara, Calamoncosis)arewith- species) they are partly fused with hypopharyn- out any spicules. The last abdominal segment may geal sclerites. The tentoropharyngeal sclerites are be rounded, incised or with more or less prominent usually less sclerotized than the mandibles and hy- processes for the posterior spiracles. On ventral popharyngeal sclerites, but the degree of scleroti- side of the last segment is the anal opening, either zation varies considerably between species. surrounded by spicules or with spicules present Mature larvae of Chloropidae are amphipneustic. only in front and behind it. The anterior spiracles are situated laterally on the Larvae of the first instar are small, less than 1 mm, first thoracic segment. Most larvae have fan-like and nearly transparent. They are metapneustic, anterior spiracles consisting of 5–12 buds (digitate anterior spiracles absent. Posterior spiracles have processes) (Figs 98, 99). Fewer larvae have anterior spiracles of a transverse rosette-type with a higher only two slits and usually with conspicuous inter- number of buds, e.g. 17 in Platycephala planifrons spiracular processes, even if these processes are (Fig. 912). The posterior spiracles on the eighth ab- absent in the mature larvae (e.g. in Lipara). The dominal segment (Figs 958, 962, 1001) are either cephalopharyngeal skeleton is not divided into hy- sessile or sitting on paired spiracular lobes (tubu- popharyngeal and tentoropharyngeal sclerites, and lar mountings, stigmophores), sometimes brown in the mandibles articulate either with long ante- colour. The shape and distance between the spirac- rior processes of the tentoropharyngeal sclerites or ular lobes are of diagnostic value. The posterior with an accessory pair of small rounded or square spiracles are surrounded by the peritreme and each sclerites (Fig. 95). Parastomal sclerites are some- consists of three elliptic slits set roughly at right times present, closely appressed to hypostomal angles to each other. The slits sometimes protrude sclerites (some Meromyza). The ridges of the facial from the spiracular plate (some Thaumatomyia). mask are less numerous and unbranched. Spicular Button inconspicuous. Between the spiracular slits zones on abdominal segments are sometimes bet- arise groups of four usually branched interspiracu- ter developed, e.g. in larvae of Oscinella. lar processes, but these may be unbranched, hair- Second instar larvae closely resemble the third like or rarely consist of three short spines (Cala- instar, but are smaller. They are amphipneustic, moncosis). The interspiracular processes are ab- but the anterior spiracles are relatively smaller and sent in Camarota curvipennis,someChlorops and knoblike with very short buds. The cephalopharyn- Lipara. The two main dorsal tracheal trunks are geal skeleton is also smaller and less sclerotized. sometimes visible through the larval cuticle. Some Puparia (Fig. 100) are shorter and thicker than larvae (e.g. Chlorops planifrons)havetheposte- the mature larvae, coloured dark or light brown, rior spiracular processes tapering and brownish. pale yellow, light greenish or almost transparent. In these cases the posterior spiracles are probably The anterior end is dorsoventrally flattened, the non-functional. surface often very wrinkled, especially at anterior The larval cuticle has thickenings or spicules of and posterior ends. The pupal spiracles remain in- variable size and form (Figs 959, 976, 980). The ar- ternal and do not break through the puparial wall rangement and size of the spicules vary between as respiratory horns. Nearly all larval characters genera and species and are important in larval are visible on the puparia except the arrangement identification. The three thoracic segments are of spicules on thoracic segments. Inside the pu- usually encircled by spicular zones consisting of several irregular rows of spicules or a series of over- parium is retained the cephalopharyngeal skeleton lapping striae (Cetema species). Spicular zones on and cephalic integument of the last instar larva. the second and/or third thoracic segments are ab- sent in some species. Larvae of Lipara may have a row of small sclerotized papillae dorsally on the

26 Anatomy of adults and larvae long to the polytrophic meroistic type, in which each oocyte is nursed by its own cluster of germ Anatomical studies of both adults and larvae of cells. Each ovary has 4–12 ovarioles, the number of Chloropidae are insufficient and fragmentary. ovarioles being correlated with body size. Female Oscinella (1.5–2.5 mm long) have 4–8 ovarioles, fe- Adults. Frey (1921) studied the skeleto-muscular male Lipara (6–8 mm long) 10–12 ovarioles. There anatomy of the head and mouth parts of adult is usually only one ripe egg in each ovariole, but in Chlorops speciosus, Platycephala planifrons, Sipho- Dicraeus usually two. The short oviducts from the nella oscinina and Thaumatomyia sp., and later ovaries join into an unpaired, rather long oviduct. (Frey, 1923) the skeletal morphology of the head The oviduct is proximally inserted by a pair of long and mouth-parts of Chlorops pumilionis.Kanmiya and thin, sometimes spiraled spermathecal ducts. (1981) illustrated indirect and direct flight muscles They arise from two unsclerotized spermathecae. of a Lipara sp. in a paper on substrate vibration as The basal oviduct also has a ventral receptacle and a means of communication. Ovtchinnikova (1989) receives a pair of rather long accessory glands. studied the musculature of the male genitalia in Li- Characters of the male reproductive system may para lucens. prove useful in taxonomic and phylogenetic stud- Inner reproductive organs of both sexes were in- ies. It is evident that tubular testes, distal fusion vestigated by Bordon (1971) in Oscinella pusilla,and of vas deferens, and absence of vas deferens may Nartshuk (1990, 1996, 1997c) studied the male in- be considered as derived states within the family. ner reproductive organs in 50 Palearctic species These characters are at terms with other apomor- from 21 genera. The male reproductive system con- phic characters of the subfamily Chloropinae and sists of two testes, each formed by one follicle as in the tribes Chloropini and Meromyzini. other Diptera (Figs 85, 86). The testes are brightly coloured yellow, brown or reddish. A small vesicle Larvae. Larval anatomy is better known than the is found on the anterior end of testes, opposite to anatomy of adults. Wandolleck (1899) described the insertion of vas deferens. Both vasa deferen- the larval anatomy of Platycephala planifrons and tia unite into a vas efferens with a basal swelling Frew (1923a) of Chlorops pumilionis and Meromyza named vesicula seminalis. Vas efferens gradually nigriventris. The larval alimentary canal of Os- narrows and becomes ductus ejaculatorius. Two cinella frit was illustrated by Shapiro (1961) and long tubular accessory glands insert at the level some details by Ryzhkova (1962) and Vilkova where the vasa deferentia join. (1963). The sclerotized larval mouth parts have Some differences in the structure of the male re- been considered above. productive system were found between the differ- The tracheal system mainly consists of a pair of ent subfamilies and genera of Chloropidae. These dorso-lateral longitudinal trunks uniting the ante- concern especially the testes and vasa deferentia rior and posterior tracheae. A series of ten dorsal that can be referred to different types. Testes: (1) transverse tracheal commissures (only the second globular in Conioscinella (Oscinellinae) and Neo- looped forwards) connect between the two main haplegis (Chloropinae); (2) ovoid, pyriform or con- longitudinal trunks. The first and tenth commis- ical in most Oscinellinae and Cryptonevra (Chlo- sures are ticker than the others. The first commis- ropinae) (Fig. 85); (3) elongated or nearly tubu- sure gives off anteriorly directed branches supply- lar in most Chloropinae (Fig. 86). Vasa deferen- ing the cephalopharyngeal complex, the remain- tia: (1) short, entering vesicula seminalis separately ing commissures are devoid of branches. The main in most Oscinellinae and some Chloropinae; (2) tracheal trunks also give off eleven lateral trunks long, entering vesicula together in Platycephala of which the anterior three are small and supply- and Meromyza (Chloropinae); (3) long, united be- ing the cephalopharyngeal complex. The remain- fore reaching vesicula in Trachysiphonella (Os- ing eight trunks pass downwards to supply the lat- cinellinae) and Neohaplegis (Chloropinae); (4) ab- eral body wall, bifurcate and unite with its posterior sent in some Chloropinae, especially in Chloropini. branches into a ventro-lateral trunk. Vesicula seminalis is present in Oscinellinae but The alimentary canal begins with the pharynx absent in most of Chloropinae. situated between the tentoropharyngeal sclerites. The female reproductive organs (Figs 87, 88, 111, The pharynx leads into the narrow, tubular oesoph- 112) consist of paired ovaries composed of sev- agus that leads into the proventriculus situated eral ovarioles. The ovaries, as in all Diptera, be- in the anterior part of the second abdominal seg-

27 ment. The proventriculus leads into the midgut Chloropspumilionis. The hind gut is slightly swollen that have several pouches (ventricular caeca) at and bulbous, lined with a well marked sclerotized the beginning. The midgut makes some big loopes intima. and eventually becomes very distended by its con- The salivary glands are a pair of simple tubular tents of dark brown colour. There is no peritrophic bodies lying on either side of the gut. The glands membrane in the midgut. There are two pairs of unite into a narrow salivary exit duct lined by a scle- malpighian tubules that unite at each side be- rotized intima. fore opening into the gut at the junction of mid- Nervous system. The brain is situated above the and hindguts. Shapiro (1961) illustrated the four oesophagus in the first abdominal segment. It is malpighian tubules in Oscinella frit as equal in connected by broad lateral commissures to a ven- length. Frew (1923a) found that the anterior tubules tral mass of fused thoracic and abdominal ganglia. are almost twice as long as the posterior ones in

Bionomics and ecology

Habitats and adult habits species of Eutropha and Conioscinella zetterstedti occur exclusively on sandy coastal beaches. Adults Most Chloropidae prefer open habitats such as of the southern European Trachysiphonella pori are grasslands and meadows, clearings and edges of associated with ant nests (Harkness & Ismay, 1976). forests, and all kinds of wetlands such as marshes, It remains to be found out, if this also is the case fens and bogs. The adults are mostly seen crawling for the two Trachysiphonella species occurring in on stems and leaves of plants. Apterous and short- northern Europe. winged species uncapable of flight are mostly seen Relatively few species of Chloropidae from the crawling or jumping on or near the ground. genera Gaurax, Gampsocera, Hapleginella, Aspisty- Adult chloropids may preferably feed on plant la, Tricimba,andLasiambia are forest inhabitants. secretions and honeydew on foliage and stems of Some species are known to occur in canopy plants. Some species also take up juice from fruits. (Thunes et al., 2004), many others occur on fo- Flowers (preferably Apiaceae, Rosaceae and Ra- liage or the trunks of trees closer to the ground. nunculaceae) are visited by relatively few species, Bährmann (2002) found 13 species of chloropids e.g. some Oscinella, Chlorops and Meromyza.Some on trunks of Salix and Populus in Germany, which Thaumatomyia visit flowers of Fabaceae. Siphun- made the family the most diverse after Phori- culina aenea is mainly caught on flowers. Kevan & dae and Hybotidae among 44 families of Diptera Baker (1983) listed Chloropidae among the Diptera . families with species visiting and pollinating flow- Some chloropid flies occasionally gather in huge ers. Some authors noted the attraction of newly numbers. Most well known are the huge autum- mowedlawnsandgreenstoadultOscinella frit. nal aggregations taking place indoors and involv- The attraction peaked about 1 hour after mowing ing Thaumatomyia notata together with Tricimba according to Falk (1982). A few species have been humeralis (Kotrba & Nartshuk, 2009). An outdoor observed feeding on various decomposing organic aggregation of Thaumatomyia glabra on Wisteria substrates in a liquefied state. Some non-Palearctic flowers in the USA was described by Sorensen species feed on wounds and eye secretions of mam- & Sorensen (1997). The non-Palearctic species mals. Apotropina gregalis formed aggregations in rock The chloropid fauna seems most diverse in mead- shelters and domestic dwellings in Namibia (Kirk- ows rich in grasses, sedges and reeds, that consti- Spriggs et al., 2001). tute the larval host plants for most species. Phrag- mites beds and moorland fens with Carex and other wetland monocots are important habitats for many Modes of larval feeding species. A few species (e.g. some Rhopalopterum, Pseudopachychaeta rufijiceps) are tyrphobionts, i.e. Larval habits and diets are very diverse in Chlorop- primarily found in raised peat bogs. Others, such as idae, especially in Oscinellinae. The subject was re-

28 viewed by Nartshuk (1972a, 1972b, 1987) and Ferrar 8. Two polyphagous species Polyodaspis rufijicor- (1987). Saprophagy, mycetophagy, necrophagy, co- nis and Tricimba cincta embrace several of the prophagy, phytophagy and zoophagy are all feeding above feeding modes (in more details under the behaviours documented from various chloropid corresponding species). larvae. Still, for most species we know practically nothing about the larval habits. Most of the investigated species have phytopha- Seasonal occurrence gous larvae, especially in the subfamily Chloro- pinae. These may be classified in four different groups: Only the phenology of Oscinella frit has been stud- ied in some detail (number of generations, adult 1. Inhabitants of shoots or stems of grasses (Poa- activity periods) by several authors in Scandinavia. ceae) and sedges (Cyperaceae) usually feeding For the remaining species information on phenol- on shoot and intercalary meristems of their ogy in this book is based on label data in the col- host plants: Oscinella, Conioscinella, Incertella, lections and the study by Nartshuk (1962a) in the Rhopalopterum, and most species of Chloro- Leningrad oblast of Russia with similar environ- pinae (Figs 101, 102). mental and climatic conditions. She recognized 10 2. Inhabitants of generative organs of and groups of Chloropidae with different live cycles and Cyperaceae feeding in particular on the deve- periods of adult activity based on own field obser- loping seeds: Dicraeus, Pseudopachychaeta (Figs vations. Less detailed data could be obtained from 103, 104). the labels of the examined material. Therefore, only 3. Gall-forming species: Lipara (the well known in- 2 groups concerning hibernation and 6 groups con- ducers of cigar galls on Common reed), but even cerning flight activity are distinguished here. Platycephala, some species of Chlorops (pumilio- The family includes univoltine, bivoltine and tri- nis, strigulus, riparius)andMeromyza nigriven- voltine species, but some species can be caught tris produce galls or gall-like swellings on their as adults all year without distinct generations. host plants (Figs 105–107). Monophagous species in particular tend to have a 4. Inquilines of gall-forming species: Calamonco- short flight period determined by the time when sis, Cryptonevra, Incertella zuercheri. their host plants are optimal targets for oviposi- tion and larval development. Species of Dicraeus Larvae of non-phytophagous (saprophagous and developing in seeds of Poaceae and species of Li- carnivorous) species of Chloropidae in northern para inducing galls in Phragmites australis are uni- Europe live in following kinds of substrates: voltine and have very short periods of adult activ- ity. Species of Dicraeus are active when their host 1. In wood under bark of trees infested by other grasses are flowering. Phragmitesaustralis, the food insects, mainly larvae of xylophagous beetles: plant of Lipara species, is only susceptible to infes- some Gaurax and Lasiambia. tation by Lipara over a very short period of time. 2. Cones of coniferous trees infested by other in- Some monophagous species of Chlorops (C. pumil- sects: Hapleginella laevifrons,someGaurax. ionis, C. limbatus, C. interruptus) are bivoltine, but 3. Mushrooms and bracket fungi: Gampsocera, the larvae of different generations feed on differ- Tricimba, Aspistyla,someGaurax. ent parts of their host plant. Oscinella frit makes 4. Mammalian excrements: Siphunculina aenea. 2–3 strongly overlapping generations per season 5. Shoots of monocots (Poaceae, Cyperaceae and in Sweden (Jonasson, 1975ff.; Lindblad, 1997). The Juncaceae), rarely dicots, attacked by other in- number of generations seems to depend on sum- sects, often phytophagous species of Chloropi- mer temperature and thus varies from one season dae: Aphanotrigonum, Elachiptera, Eribolus, In- to the next. The number of generations per season certella, Oscinisoma,someConioscinella, La- is impossible to find out for several species with siosina. apparently strongly overlapping generations. 6. Egg masses of spiders and short-horned grass- Two distinct groups of species can be distin- hoppers: Pseudogaurax venustus, Lasiambiapal- guished in relation to hibernation. One group over- posa, L. coxalis, Speccafrons halophila. wintersasadults:someElachiptera, Conioscinella, 7. Plant roots inhabited by root aphids: Thaumato- Aphanotrigonum, Tricimba, Eribolus, Oscinisoma, myia. some Thaumatomyia,andPseudopachychaeta. All

29 Figs 101–107. 101: shoot of wheat, damaged by frit fly Oscinella frit (Linnaeus). 102: stem and part of spike damaged by goud fly Chlorops pumilionis (Bjerkander). 103: egg of Dicraeus fennicus Duda on seed scale of Elytrigia repens.104:larvaofDicraeus fennicus within seed of Elytrigia repens.105:gallonRoegneria canina produced by Chlorops riparius Smirnov. 106: gall on common reed (Phragmites australis) produced by Lipara larvae. 107: photos of galls of Lipara spp.

30 Table 1. Larval life habits of the chloropid genera occurring in the area investigated. Subfamily/Genus Feeding habit(s) Place of development Subfamily Rhodesiellinae: Aspistyla mycetophagous or saprophagous fungi Subfamily Oscinellinae: Aphanotrigonum saprophagous shoots of Poaceae and Cyperaceae Calamoncosis phytophagous shoots of Poaceae, chloropid galls on Phragmites australis Conioscinella phytophagous and saprophagous shoots of Poaceae, galls of insects on oak trees Dicraeus phytophagous seeds of Poaceae Elachiptera phytosaprophagous shoots of Poaceae and other plants Eribolus phytosaprophagous shoots of Cyperaceae Gampsocera mycetophagous Myxomycetes Gaurax saprophagous (xylophagous) and necrophagous rotten wood, bark of trees and coniferous cones, bracket fungi, galls of Agrobacterum tumefacies Hapleginella saprophagous and necrophagous damaged coniferous cones Incertella phytosaprophagous and phytophagous shoots of Poaceae, Lipara galls on Phragmites australis Lasiambia saprophagous (xylophagous) and carnivorous rotten wood, sap of wood, egg pods of Orthoptera and Mantodea Lipara phytophagous gallformers on Phragmites australis Oscinella phytophagous shoots or seeds of Poaceae and Cyperaceae Oscinimorpha unknown unknown Oscinisoma phytosaprophagous shoots of Poaceae and Cyperaceae Polyodaspis polyphagous walnuts, stems of damaged plants, head of Asteraceae, leaves rolled by beetles Pseudogaurax carnivorous egg sacks of Aranea Rhopalopterum phytophagous or phytosaprophagous shoots of Carex and other Cyperaceae Siphonella ? carnivorous ? egg cocoons of Aranea, Coccoidea on grasses Siphunculina saprophagous or coprophagous mammalian faces, bird nests Speccafrons carnivorous egg cocons of Aranea Trachysiphonella unknown unknown Tricimba saprophagous, necrophagous and fungi, rotten shoots of plants, bird nests, dead mycetophagous mollusks Subfamily Chloropinae: Camarota phytophagous shoots of Poaceae Centorisoma phytophagous shoots of Poaceae Cetema phytophagous shoots of Poaceae Chlorops phytophagous shoots of Poaceae and Cyperaceae (Carex spp.) Chloropsina phytophagous or phytosaprophagous shoots of Poaceae and other plants Cryptonevra phytophagous or phytosaprophagous inquilines in Lipara galls on Phragmites Diplotoxa phytophagous or phytosaprophagous rhizome and stems of Eleocharis spp. Epichlorops unknown unknown Eurina phytophagous gallforming on Phragmites, Scirpus and other Cyperaceae Eutropha ? saprophagous plants and other substrates Lasiosina phytosaprophagous and ? phytophagous shoots of Poaceae and Cyperaceae Melanum unknown unknown Meromyza phytophagous shoots of Poaceae Neohaplegis phytophagous or phytosaprophagous shoots of Cyperaceae (Carex spp.) Parectecephala phytophagous shoots of Poaceae Platycephala phytophagous shoots of Phragmites Pseudopachychaeta phytophagous inflorescence of Cyperaceae (Eriophorum, Eleocharis) Thaumatomyia carnivorous rhizome of plants on root aphids Trichieurina ? phytophagous shoots of Cyperaceae (Carex spp.)

31 species of Oscinellinae belonging to this group Eribolusgracilis, E.hungaricus, Trachysiphonella have saprophagous larvae developing in rotting scutellata, Incertella antennata, Polyodaspis sul- shoots of Cyperaceae or Poaceae. Species of Chlo- cicollis, Microcercis trigonella. Most species of ropinae in this group have carnivorous (Thaumato- Dicraeus, developing in seeds of Poaceae, be- myia notata)orphytophagous(Pseudopachychaeta long to this group, because this period is time of rufijiceps) larvae. Some species overwintering as flowering for most grasses. All listed species are adults have been caught during the cold season: phytophagous. Eribolus nana on 10.xi and 25.ii, Oscinisoma cog- 4. Species with adult activity in late summer (July, natum on 1.xii and 6.i. Elachiptera tuberculifera August): many species of Chlorops (C. fijinitimus, was taken in a light trap in December in Norway C. geminatus, C. kirigaminensis, C. rufijinus, C. (Hågvar & Greve, 2003). Species of Elachiptera, centromaculatus), Aphanotrigonum cinctellum, Tricimba and Thaumatomyia notata were found in Conioscinella sordidella, C. zetterstedti and winter in bird’s nests or between scales of Picea Apistyla plumiger. cones. Mass occurrence of Thaumatomyia notata 5. Species with adult activity in autumn (August, inside houses is a late season event and probably September): Chlorops ringens, C. varsoviensis, a hibernation activity. Skuhravá (1980) stated that Paractecephala longicornis, Siphonella oscinina. Platycephala planifrons overwinters as adult in the 6. A special phenology is seen in Pseudopachy- Czech Rep., but in Fennoscandia this species has chaeta rufijiceps, a larval feeder on the inflores- not been caught after September. cences of Eriophorum vaginatum.Thespecies The second group of species overwinters as lar- has one generation per year. Newly emerged vae, or sometimes as praepupae or puparia. Most adults appear in June and stay active to the end of the phytophagous species belong to this group. of the season. They then hibernate as adults and Their larvae hibernate in the shoots, seeds or galls become active again in April and early May next of their host plants. It is not known if overwinter- season, at the flowering time of their host plant. ing larvae (or adults) at any time enter a state of diapause. It is presumed for hibernating adults of Thaumatomyia notata (Weidner, 1990). It is known Role in land ecosystems and species that the adults of some species (Dicraeus, Thau- matomyia notata) first emerge from their puparia of economic importance after these have been moistened by water. It is possible to divide NW European Chloropidae Chloropidae constitute a substantial proportion of in 6 groups by their flight activity pattern. species richness in grass ecosystems such as mead- ows, marches, waste lands, pastures, some agricul- 1. Species with adult activity during the entire tural fields and so on and play a significant role season from April or May to September or Oc- in the function of these ecosystems in the temper- tober. Many species belong to this group; they ate zone. Recent studies demonstrate that some hibernate either as adults or larvae. Some have species are abundant in woodlands as well, and are saprophagous larvae developing in rotting plant involved in the destruction of rotten wood, conifer- tissues or phytophagous larvae developing in ous cones and fungi (Komonen et al., 2004; Thunes young shoots: species of Aphanotrigonum, Eri- et al., 2004). A surprisingly high abundance and bolus, Conioscinella, Oscinella, Elachiptera, In- species diversity of Chloropidae were found in the certella, Rhopalopterum, Cetema, Chlorops, canopy of trees in a tropical rain forest in southeast- Meromyza, Epichlorops, Diplotoxa. ern Asia (Floren, 2003). 2. Species with adult activity in spring and early Larvae of most Chloropidae are either scavengers summer (May, June): Lipara lucens, L. pullitar- and phytophagous on species associated with the sis, Cryptonevra consimilis,andCalamoncosis monocot plants families Poaceae, Cyperaceae, Jun- aprica, all associated with Phragmites australis, caceae and a few others. The abundance of Chlo- and Dicraeus styriacus, Chlorops frontosus, C. ropidae in grassland habitats gives them an impor- strigulus. All listed species have phytophagous tant role as primary consumers of plant tissues and larvae. detritus feeders. Few Chloropidae have predaceous 3. Species with adult activity in the first half of larvae. Larvae of Pseudogaurax and Speccafrons summer (June, July): Conioscinella gallarum, C. and some Lasiambia feed on eggs of spiders and mimula, Dicraeus nitidus, D. tibialis, D. vagans, orthopterans. The larvae of Thaumatomyia feed ex-

32 clusively on aphids living on plant roots. The well subtile (Thomson), G. jaculator (Linnaeus) (Gaste- known mass occurrences of Thaumatomyia no- ruptiidae) and Gambrus carnifex (Gravenhorst) tata have also be observed in Scandinavia: Finland (Ichneumonidae) inhabited by Hylaeus pectoralis. (Haartman, 1950), Sweden (Lampa, 1893; T. Pape, The family Chloropidae has a great potential pers. comm.) and suggest that the species helps as bioindicator to monitor the condition of envi- in the control of subterranean aphids. Chloropi- ronments and ecosystems according to McGeoch dae obviously also play a significant role as a food (1998): chloropids are abundant and distributed resource for numerous parasitoid Hymenoptera over a wide range of terrestrial habitats; some and some parasitic nematodes and fungi. Adult species are very selective in their habitat require- and larval chloropids are also used as prey by vari- ments; spatial and temporal distributions predict- ous predatory Hymenoptera and Diptera, and birds ably ensure long term continuity; essential data on such as swallows, swifts and tits. Chloropids make biology is available for many species; the species 20% of the food of the Common Swift (Apus apus) are sampled easily by sweeping of low vegetation during the period of bringing up nestlings accord- and by various traps allowing quantitative results. ing to Parmenter & Owen (1954). Crabronid wasps On top of this adult chloropids are overall eas- belonging to Lindenius, Rhopalum and Crossocerus ily distinguished from other small acalyptrate flies use Chlorops fliesaslarvalprovision(Berland,1925; by structural (large ocellar triangle, weak chaeto- unpubl. obs.). Blue tits have been observed feed- taxy) and often colour characteristics (contrast- ing on Lipara larvae from small galls on Common ing dark pattern on a yellow or greenish back- reed during winter (Tischler, 1943; Mook, 1967). ground). Studies in Poland show that the pro- Great Spotted Woodpecker (Dendrocopus major L.) portion of Chloropidae among terrestrial arthro- opens large galls of Lipara lucens (Kramer, 1917). pods in grassland ecosystems increases markedly Old cigar galls of Lipara are used by some bees in habitats disturbed by man compared to natural and wasps for nesting (Giraud, 1863; Wagner, 1907; meadow habitats (Frydlewicz-Ciesielska, 1961; Ole- Elfing, 1951; Valkeila, 1956; Waitzbauer, 1969; Imhof, chowicz, 1971; Dabrowska-Prot, 1984, 1986, 1987). 1979; Wolf, 1988, 1991; Westrich, 1989; Dely- Nowakowski (1982) found out that 75% of all aca- Draskovits et al., 1994; Anonymous, 2000). Nests lyptrate flies sampled from urban lawns in Poland of following Hymenoptera were found: belonged to Chloropidae. Evidently, some species of Chloropidae even become more abundant in Apidae: Hylaeus pectoralis Förster, H. gracilicornis areas subjected to moderate pollution from both Morawitz, H. moricei (Friese), Megachile centun- sulphur- and fluorine-containing aerial emission cularis Linnaeus, M.versicolor Smith, Hoplitisleu- (Dabrowska-Prot, 1984; Bährmann, 1985; Zvereva, comelana (Kirby). 1993b; Kozlov & Zvereva, 1997). Zvereva (1993a) Crabronidae: Pemphredon rugifer (Dahlbom), P. underscored this tolerance to pollution in a study lethifer (Shuckard), P. inornata Say, P. wesmaeli on the insect fauna in a strongly disturbed area (Morawits), Pemphredon sp., Rhopalum clavipes near the plant “Severonikel” on the Kola Peninsula, (Linnaeus), Trypoxylon attenuatum F. Smith, T. where chloropids proved to be remarkably abun- fijigulus (Linnaeus), Passaloecus gracilis (Curtis), dant. P. clypealis Faeser, P. corniger Shuckard, P. singu- Kibík (2002) divided species of Chloropidae in laris (Dahlbom). three groups according to their ability to cope with Vespidae: Symmorphus bifaciatus (Linnaeus), S. anthropogenic deteriorations of ecosystems: (1) fuscipes (Herrich-Schäffer). anthropo-intolerant species (e.g., Incertella scot- Tenthridinidae: Brachythops flavens (Klug). Prob- ica, Pseudopachychaeta rufijiceps); (2) anthropo- ably the galls were incidentally used for overwin- neutral species (e.g., Aphanotrigonum trilineatum, tering. Chlorops pumilionis); and (3) anthropo-tolerant species (e.g. Elachiptera tuberculifera). Parasitoids and kleptoparasites of the listed bees Some of the species of Chloropidae have a great and wasps have also been reared from Lipara galls: economic importance as stem miners or grain feed- Pseudomalus auratus (Linnaeus) (Chrysididae), ers of cereals and fodder grasses. In particular the Campodorus variegatus (Jurine) (Ichneumonidae) Frit Fly (Oscinella frit) is regarded as a major pest and Macronychia sp. (Sarcophagidae) inhabited by species of cereals. It occurs in all Fennoscandia and Pemphredon rugifer and P. wesmaeli; Gasteruption Denmark (Znamenskaya, 1962). There are some nigrescens Schletterer, G. assectator (Linnaeus), G. closely related species (Oscinella pusilla, O. vastator

33 and O. nigerrima) that may also damage cereals, but yellow tip and base, and second aristal segment the taxonomy of these are still not fully clarified. about one-third as long as third segment. Another The Frit Fly (Oscinella frit) is a tiny black fly about is O. vastator that only differs from Oscinella pusilla 1–1.5 mm long with black arista, shining ocellar tri- in having the tibiae of front and middle legs never angle, clear wings and dusted scutum (Cover illus- entirely yellow and short brownish wings usually tration). It belongs to a complex of similar species not reaching the tip of the abdomen. or forms, but the “true” Frit fly is characterized by Several authors (Nartshuk, 1956; Ibbotson, 1961; having the posterior tibiae entirely black or nar- Le Berre & Chevin, 1961; Moreau, 1963; Vickerman, rowly yellowish at base and tip, front and mid- 1980; Savage, 1982; Nielsen, 1985) found differences dle tibiae usually black centrally and more widely in behaviour and host plant preferences between yellowish at base and tip, and second aristal seg- different species within the frit fly complex. Larvae ment is about half as long as third aristal segment. of O. frit develop in West Europe in young shoots of The species is polyphagous and injurious to bar- cereals and some grasses and within unripe grains ley, wheat, ray, and especially oats, and many of cereals, mainly oats, but also wheat and maize. fodder grasses. The larvae attack young shoots as In East Europe O. frit has not been found on unripe well as unripe grains. The species overwinters as grains of cereals except for maize. Nartshuk (1956) larva in seedlings of winter cereals or wild grasses found that O. pusilla prefers wheat and barley to and pupates the following spring. The puparia stay other cereals, while O. frit prefers wheat and oats. within the shoots. Adults of the first generation O. pusilla and O. vastator develop only in young emerge from the end of April to early in June. Fertil- shoots. Moreau (1963) found that O. pusilla prefers ized females then lay eggs either on the lower part barley and wheat to oats and never develops in un- (usually under the ligula) of seedlings of spring ce- ripe grains of cereals in France. reals or on the ground nearby the host plant. The In respect to fodder grasses, Oscinella frit and larvae eat a tunnel inwards and downwards in a O. vastator showthesamepreferenceforthean- spiral manner until they reach the center of the nual Lolium multiflorum and perennial L. perenne shoot, which results in destruction of the growth W Europe, while O. pusilla shows a preference for zone of the shoot. Usually one larva is found per shoot. Infested plants are easily recognized by the L. perenne (Jepson & Heard, 1959; Wetzel, 1967; dry central leaf (Fig. 101). Attacked seedlings ei- Vickerman, 1978a; Nielsson & Nielsson, 1984; Niel- ther die or react by producing new shoots result- son, 1985). Andersson (1956, 1967) reared O. pusilla ing in weakened plants and diminished production from stems of barley and wheat in Sweden, Rygg of grains. Injury is more pronounced in late-sown (1966a) from the same cereals in Norway, but also fields of spring cereals. Pupation takes place in the from rye, timothy and maize. Overall it seems that injured plants and the adults of the second gener- O. pusilla prefers grasses of the tribe Hordeae that ation emerge from June to August. Females of the include the cereals wheat and barley, while O. frit summer generation lay to some extent their eggs prefers grasses of the tribe Avenae that include oats on the developing ears of cereals where the larvae (Nartshuk, 1956; Chevin et al., 1974). feed on the developing grains. In years with a long Sampling with colour traps indicated that Os- warm summer adults of a third autumn generation cinella frit prefers blue coloured traps and O. pusilla may show up in August to September. Adults of the yellow ones. Some behavioural differences were summer and autumn generations complete the life also noted between the three species found most cycle by laying eggs on seedlings of wild grasses and often in agricultural fields. Adults of O. pusilla winter cereals. Larvae of frit flies are involved in the and O. vastator aremorestationary,preferringto transmission of barley yellow dwarf virus (Jess & live near ground in perennial vegetation. O. frit Mowat, 1986). Several chloropids, not the least Os- is a more mobile species and thus more capable cinella frit and allies, are causing “silvertop” injury of spreading to annual crops (Vickerman, 1980; in fodder grasses. This term describes a condition Nielsen & Nielsen, 1984; Nielsen, 1985). in which the stem is injured and the seed head dries The Frit fly problem has been investigated in de- off prematurely. tails in all Nordic countries: Other species of the Oscinella frit complex have been reported as cereal pests in NW Europe. One is Norway: Schøyen (1895, 1896, 1899, 1918a, 1918b, Oscinella pusilla that has entirely yellow front and 1919), Rygg (1966a, 1966b, 1967, 1968), Håbjørg middle tibiae, posterior tibiae black centrally with (1979).

34 Denmark: E. Rostrup (1893), S. Rostrup (1923), C. pumilionis is less important as a pest of cere- Anonymous (1928), Bagger & Thygesen (1971), als in the Nordic countries than Oscinella species, Jensen & Thygesen (1971), Nødegaard & Hansen but damage to cereals are reported from Finland (1972), Bagger & Nielsen (1975), Bejer & Esbjerg (Reuter, 1902b; Listo, 1926a; Kanervo & Vappula, (1978), Nielsen & Nielsen (1984a, 1984b), Nielsen 1962; Vappula, 1965) and Sweden (Bjerkander, 1789; (1985a, 1985b, 1994). Dahlbom, 1837; Larsson, 1885; Lampa, 1887, 1888, Sweden: Linnaeus (1750), Dahlbom (1837), Hed- 1891, 1892; Hedström, 1892; Wahlgren, 1918). ström (1884, 1892), Post (1884, 1885), Larsson Meromyza nigriventris has been reported to at- (1885), Lampa (1887, 1888, 1891, 1892), Aurivil- tack wheat, oats and barley in several Nordic coun- lius (1892), Trybom (1895), Nilsson-Ehle (1902, tries, but the species is of minor economic impor- 1906), Tedin (1902), Wahlberg (1918), Cunliffe tance. The larvae live under the uppermost leaf (1929, 1930), Anonymous (1935), Ahlberg (1937), sheath of cereals and cause “silvertop” disease (see Schwan (1939), Johansson (1943, 1960, 1962, above). It has been recorded under other names 1975), Ackerman (1951), Andersson & Ossian- from Finland (Reuter, 1902a, 1902b; Listo, 1926a; nilsson (1951), Andersson (1956, 1967), Anerud Vappula, 1965), Norway (Rygg, 1966a), and Den- (1957), Haegermark (1960, 1977), Larsson (1961, mark (Nielsen, 1994). 1978, 1979, 1984), Nitsh (1970), Jonasson (1975, Several other species of Chloropidae have some- 1976,1977a,1977b,1978,1980,1982a,1982b,1985a, times been reared from cereals and fodder grasses 1986, 1988a, 1988b, 1988c), Swenson (1976), Had- in the Scandinavian countries: Incertella albipalpis, berg (1978), Norlander (1978a, 1978b), Sigvald Aphanotrigonum trilineatum, Elachiptera cornuta, (1978), Curry, (1986), Lindblad (1988, 1991, 1993, E. tuberculifera, Tricimba cincta, Eribolus nana, La- 1997, 1999, 2001), Lindblad & Solbeck (1998), siosina herpini and Cetema elongatum (Johansson, Lindblad & Sigvald (1999). 1942, 1960; Kallio, 1950; Kolstad, 1965; Vappula, Finland: Reuter (1895, 1896, 1900), Listo (1926b), 1965; Rygg, 1967; Nielsen, 1994). Larvae of these Pohjakallio (1936), Kallio (1950), Jamailainen & species are scavengers and secondary invaders of Kanervo (1953), Hardh (1955), Tiittanen (1959a, plants with attacks from primary consumers of 1959b), Kanervo (1960a, 1960b), Kanervo & fresh plant tissues. Their activity period usually Vappula (1962), Vappula (1965), Ervio (1978), starts later than that of the phytophagous Oscinella. Markkula (1985), Kurppa (1990a, 1990b, 1991). One more species, Thaumatomyia notata merits Karelia and the Kola Peninsula: Kanervo (1942), attention. This fly sometimes enters buildings in Znamenskaya (1962). enormous swarms in the autumn, especially after a warm and dry summer (Figs 108–112) for over- Another cereal pest of some importance is the wintering. Haartman (1950) described this phe- Gout Fly, Chlorops pumilionis. This fly is 3–4 mm nomenon from Turku in 1948. Mass occurrences long and of yellow colour with scutal stripes and of T. notata inside buildings were also observed in a spot on katepisternum black. The broad ocellar Finland and Sweden near Stockholm in 1997. The triangle reaches to middle of frons and has a me- flies cannot, however, hibernate in heated build- dian groove. The triangle is black in dark speci- ings and will then dry out and die within some mens but with broad yellow margins in pale spec- weeks. T. notata can under these circumstances be imens. Antennae extensively dark coloured. Legs very disturbing by smearing ceilings and walls with yellow with tarsi and tips of front tibiae darkened. their excrements. Mass occurrences of this fly are C. pumilionis attacks mainly wheat but also bar- most likely to take place in buildings surrounded ley and couch grass (Elytrigia repens). The larvae by extensive lawns. It has also been reported that overwinter in a gall formed low down on the stem T. notata may choose the same building for this be- of its host plant. Pupation takes place within its havior several times in succession. The existing lit- gall in spring. Grasses and cereals attacked by lar- erature on this aggregation phenomenon is exten- vae of the winter generation of C. pumilionis are sive. A selection of the more important ones is as stunted and do not develop floral parts. Females follows: Waga (1848), Kiesenwetter (1857), Weyen- of the summer generation lay eggs on the upper berg (1871), Letzner (1873), Reh (1928), Hase (1929), leaves of their host plant. The larvae feed on de- Roszypal (1930), Uhlmann (1930), Lindner (1931), veloping ears and then move down the culm to Tzygankov (1931), Zuska (1966), Weidner (1990), pupate within a leaf sheath. Infested plants be- Nartshuk (2000b, 2000c) and Kotrba & Nartshuk come stunted with little or no seed development. (2009). Specimens of T. notata from Waga’s obser-

35 Figs 108–112. Thaumatomyia notata (Meigen): 108: map showing mass occurrences in Europe (white dots before 1990, black dots after 1990). 109: adult fly in autumn (abdomen inflated by fat body). 110: adult fly in spring and summer. 111: ovarium in autumn. 112: ovarium in summer filled with ripe eggs. (108 after Kotrba & Nartshuk, 2009; 109, 110 after Lindner, 1931.)

36 vationsinPolandin1848arestillkeeptinNHMW ly 50% of a sample of larvae of O. frit and O. vastator (unpubl. obs.). The earliest notes on these aggre- was infested by parasitoids. gations of flies date back to 1736, 1766 and 1805 in Also the parasitoids of Lipara and other chloro- England, 1807 in Germany, and 1812 in Poland, but pid species living as inquilines in Lipara galls on without specifying their identity with a scientific Common reed (Phragmites australis) are quite well name. Jenyns (1832) described the phenomenon in investigated: Giraud (1863), Blair (1932, 1934), England and for the first time proposed an iden- Erdös (1952, 1955, 1957, 1961), Valkeila (1959a, tity of the fly as Chlorops laeta. Subsequent au- 1959b), Mook (1961, 1967), De Bruyn (1987), Gro- thors agreed on the family assignment of the fly chowska (1989), Gromysz-Kałkowska & Grocho- involved in mass occurrences, but with absolute wska (1992) and Dely-Draskovits et al. (1993, 1994). no concensus on its specific identity. In various Records of parasitoids of other Chloropidae are combinations with the generic names Chlorops, rare (Nartshuk, 1962a; Kiauka & Nartshuk, 1972; Chloropisca and Thaumatomyia the species has Dyurich & Kostyukov, 1978; Dyurich, 1980; Norlan- been recorded by the following names: lineata der & Grijpma, 1991). Fabricius, nasuta Schrank, didyma Zetterstedt, ob- The reported parasitoids of Chloropidae are scurella Zetterstedt, minuta Zetterstedt, prodigiosa mainly reared from the puparia or from stems or Zenker, copiosa Schiner, laeta Meigen, hypostigma inflorescences containing chloropid larvae and/or Meigen, circumdata Meigen, ornata Meigen, notata puparia. Such plant parts, especially chloropid Meigen, nasuta Meigen, flavifrons Meigen, and tae- galls, may easily contain other insects and their niopus Meigen. Of these names prodigiosa Zenker, parasitoids. This aspect, as well as the possible oc- circumdata Meigen, ornata Meigen and copiosa currence of hyperparasitoids, suggests that the ex- Schiner are now regarded as synonyms of notata isting host records for parasitoids are frequently Meigen, the other ones are based on misidentifica- based on mistakes. tion. The behavior of the present species is alluded Most parasitoids of Chloropidae are found to in the names prodigiosa and copiosa and the among Hymenoptera (Table 2). Their development generic name Thaumatomyia means “the mysteri- takes place in the chloropid larvae followed by ous fly”. their emergence from the puparia. It has been ob- An analysis of the literature data on mass occur- served that the braconid wasp Polemochartus li- rences of T. notata in Europe for more than two parae oviposits in the eggs of Lipara found exposed centuries revealed that these occur at intervals of on shoots of Phragmites, but that development ac- 10–12 years. A hypotheses proposed by Nartshuk tually takes place in the larvae (Mook, 1961). Larvae (2000b, 2000c) suggests that the aggregations be- of Lipara lucens parasitized by P. liparae pupate in havior of T. notata coincide with the 11-year cycle of September instead of spring in the following year as the sun activity with its documented influence on they normally do (Varley & Butler, 1933; Chvála et the weather conditions. al., 1974). Stenomalina liparae (Pteromalidae) par- asitizes the young larvae of Lipara (Chvála et al., 1974; Athen & Tscharntke, 1999). The wingless bra- conid Chasmopodon apterus,thepteromalidTri- Parasitoids, predators and parasites chomalus statutus and a figitid wasp (Ganaspis sp.) lay their eggs in young larvae of their hosts, while Parasites and parasitoids of the Chloropidae are the pteromalid Spalangiafuscipes and the eulophid known in more detail for the agriculturally im- Pronotalia liparae oviposit in the host puparia. Tri- portant species of the Oscinella frit species com- chomalus cristatus lays several eggs in one larva of plex and Chlorops pumilionis. Several authors re- Oscinella, but only one parasitoid will survive. Most port on parasitoids reared from frit flies in Scan- parasitoids are solitary, but examples of gregarious dinavia (Lampa, 1888; Henriksen, 1918–1919; Grad- species are: Tetrastichus legionaries with up to 40 well, 1957; Nordlander, 1978a, 1978b; Nielsen, 1994), specimens emerging from a single puparium of Li- other parts of Europe (Kurdyumov, 1912; Meyer, para, Clytina giraudi with 3–15 specimens from one 1923; Imms, 1930, 1932; Riggert, 1935; Nikolskaya, puparium of a Cryptonevra or Calamoncosis sp., 1937; Dyurich, 1980; Moore, 1983) and in North Crataepiella platycephalae with 18–55 specimens America (Simmonds, 1952, 1953a, 1953b, 1954, from single puparia of Platycephala umbraculata, 1956). Species of the O. frit complex tend to be and Pronotalia liparae and Tetrastichus djuritshae strongly parasitized. Moore (1983) found that near- with 44–56 specimens from puparia of Lipara lu-

37 cens. Aggregations of Oscinella larvae and their hy- Broscus cephalotes, Pterostichus cupreus, P. punctu- menopterous parasitoids were observed in grass- latus, Carabus convexus) are listed as predators of lands in England and that the mortality among frit flies (Toldaev, 1972). Bembidion lampros feeds overwintering frit fly larvae differed between par- on eggs of Oscinella situated on the ground (Nord- asitized and unparasitized ones (Umoru, 1993c, lander, 1978a). Also predaceous mites of the genus 1994). Nordlander (1978a) noted that puparia of Os- Pergamasus (Parasitidae) eat chloropid eggs (Jep- cinella from shoots of cereals were parasitized up son & Southwood, 1958). Jones (1968a) noted that to 50% in Sweden, but puparia in grains of cereals the activity of larger carabid beetles may disturb were practically all unparasitized. females of Oscinella frit in laying eggs on shoots Eggs parasitoids of Chloropidae are not known. of oats. The plant bug Miris dolabratus (Miridae) Some testing of the potential of Trichogramma was once recorded as a predator of frit flies (Collin, evanescens (Trichogrammatidae) as a control agent 1918), but this seems questionable. Ochthera man- against Oscinella frit in Russia have been under- tis (Ephydridae) feed on adult chloropids (Hobby, taken in the laboratory and in field (Chernoponev- 1931). sky, 1939). This egg parasitoid did infest eggs of Adults of the Oscinella frit complex often carry the frit fly in the experiments, but the develop- red larvae of the mite Microtrombidium demeijerei ment of parasitoids took 28 days instead 9 days in (Oudemans) (Microtrombiidae) (Riggert, 1935). eggs of Lepidoptera. Spalangia drosophilae (Ptero- Riggert (1935) proposed a negative influence of malidae) is reported as a parasitoid of Oscinella this mite on adults of Oscinella frit similar to that frit in North America (Simmonds, 1952, 1953a, of parasitic Nematoda. Another microtrombiid 1953b, 1954, 1956). This species has never been mite species, Ettmuelleria caudatum was found on reared from frit flies in Europe, but some research the abdomen of Chlorops pumilionis as well as on was undertaken to find out the potential of this Sphaeroceridae (Smith, 1955). species as a control agent of O. frit in NW Russia Adults and larvae of the Oscinella frit complex and Latvia (Kovalchuk, 1968, 1972, 1977; Skaldere, are hosts of parasitic and saprophylic nematodes. 1976a, 1976b, 1979). It was shown later on that S. Young fertilized females of the obligate parasitic drosophilae in Europe primarily infests larvae of nematode Howardula oscinellae (Allantonemati- Sphaeroceridae and Drosophilidae living in dung, dae) penetrate and live in the larval haemocoel and only accidentally uses O. frit as a host (Mar- shakov, 1983). Marshakov (1984) presented a re- of Oscinella spp. The viviparous parasite is car- view of the genuine, casual and fake parasitoids of ried through the pupal stage and produces its lar- frit flies. vae in the body of the adult host fly. These larvae Cleigastra apicalis (Scathophagidae) is a well leave their host fly by entering its gut. Flies in- known predator of Lipara larvae (Wagner, 1907; fested by this nematode are affected in the develop- Theowald, 1961; Chvála et al., 1974; Askew & Shaw, ment of their inner reproductive organs (Goodey, 1979; De Bruyn, 1985, 1987). Other predators of Li- 1930a, 1930b; Filipjev, 1934; Wachek, 1955; Poinar, para larvae are birds (see chapter ‘Role in land 1975; Moore & Hunt, 1984; Slobodyanyuk, 1984). ecosystems and species of economic importance’). Other species of parasitic nematodes belonging Coenosia tigrina (Muscidae) is a predator of stem- to Mermithidae and Auguillulidae have been re- boring flies, including Oscinella frit and other Chlo- corded from adult Chlorops pumilionis and La- ropidae (Parmenter, 1968; Kuehne, 1991). Aelothrips siosina herpini (Tzygankov, 1930). A parasitic fasciatus, a species of thrips (Thysanoptera) has mermithid nematode, probably a Hexamermis sp., been recorded as an egg predator of Chloropidae has also been found in larvae of O. frit (Moore & (Znamensky, 1926; Tzygankov, 1930). Some unspe- Hunt,1984).EventhesaproxylicnematodePana- cialized predators may help control some of the grolaimus rigidus (Panagrolaimidae) causes some common agricultural pest species among Chlorop- mortality of Oscinella spp. larvae (Poinar, 1972, idae. Heidger & Nentwig (1989) noted that the spi- 1975). Usually only two or three, but up to six were der Dictyna arundinaceae (Linnaeus) builts its web found in the haemocoel of living larvae, and as preferably at the ears of wheat and catches almost many as 12 in a dead larva (Moore & Hunt, 1984, exclusively adults of Oscinella frit. Some Coccinell- 1987). Jepson & Southwood (1958) found that the idae are known to eat larvae of O. frit (Collin, 1918) poor state of overwintering larvae of Oscinella frit and several species of Carabidae (Bembidion lam- was caused by the presence of a yeast and a ciliate pros, B. quadrimaculatum, Pseudoophonus rufijipes, protozoan in their body cavity.

38 Table 2. Parasitoids of Chloropidae recorded from Europe. Parasitoids Chloropidae hosts Authors Proctotrupoidea Diapriidae Basalys (= Loxotropa) crassiclava Oscinella frit Chukanova & Kovalchuk, 1974; Dyurich, 1980; Talitskiy & Kieffer Kuslitskiy, 1990 ?Basalys tritoma (Thomson) Oscinella frit Meyer, 1923; Schander & Meyer, 1925; Vojnovskaja-Kriger, 1929; Imms, 1930, 1932; Riggert, 1935; Simmonds, 1952; Jones, 1965, 1968b; Fulmek, 1968; Chukanova & Kovalchuk, 1974; Nordlander, 1978b Basalys sp. Oscinella frit Chukanova & Kovalchuk, 1974 Trichopria aequata (Thomson) Oscinella frit Vojnovskaya-Kriger, 1929; Riggert, 1935; Simmonds, 1952; (= variipes Kieffer) Fulmek, 1968; Chukanova & Kovalchuk, 1974 Trichopria (= Tropidopria) Oscinella frit Fulmek, 1968 compressa Thomson T. verticillata Latreille Meromyza nigriventris Dyurich, 1980 M. pratorum Talitskiy & Kuslitskiy, 1990 Trichopria (= Ashmeadopria)sp. Oscinella frit Riggert, 1935; Simmonds, 1952; Fulmek, 1968; Chukanova &Kovalchuk,1974 Trichopria sp. Oscinella frit Chukanova & Kovalchuk, 1974 Platygastridae Paramisius sp. Oscinella frit Chukanova & Kovalchuk, 1974 Platygaster sp. Oscinella frit Chukanova & Kovalchuk, 1974; Moore, 1983 Ceraphronoidea Megaspilidae Dendrocerus chloropidarum Hapleginella sp. or Gaurax sp. Dessart, 1990 Dessart Cynipoidea Eucoilidae ?Eucoila sp. Oscinella frit Fulmek, 1968 Ganaspis mundata Förster Oscinella frit Nordlander, 1978b Ganaspis sp. Oscinella frit Ivatsik & Kovalchuk, 1975 cone inhabitant Chloropidae Chukasnova & Kovalchuk, 1974; Nordlander & Grijpma, 1991 Grybliographa (= Pseudeucoila) Oscinella frit Simmonds, 1952; Fulmek, 1968 (= Psichacra)sp. Hexacola fuscipes Meyer Oscinella frit Meyer, 1923; Riggert, 1935 H. hexatoma (Hartig) Oscinella frit Hedike, 1923; Meyer, 1923; Schander & Meyer, 1925; Imms, 1930; Simmonds, 1952; Kerrich & Quinlan, 1960; Fulmek, 1968; Chukanova & Kovalchuk, 1974; Nordlander, 1978b; Moore, 1983, 1984; Umoru, 1993b Hexacola sp. Oscinella frit Chukanova & Kovalchuk, 1974 Kleidotona psiloides Westul. Oscinella frit Chukanova & Kovalchuk, 1974 Odoutecoila sp. Oscinella frit Chukanova & Kovalchuk, 1974 Rhoptromeris heptoma (Hartig) Oscinella frit Baranov, 1912; Kurdjumov, 1912; Collin, 1918; Meyer, 1923; (= eucera Hartig) (= widhalmi Schander & Meyer, 1925; Vojnovskaja-Kriger, 1929; Kurdjumov) ?Imms, 1930, 1932; Riggert, 1935; Simmonds, 1952; Kerrich & Quinlan, 1960; Anonymous, 1963; Jones, 1965, 1968b; Fulmek, 1968; Nordlander, 1978a, 1978b; Dyurich, 1980; Howell, 1982; Talitskiy & Kuslitskiy, 1990; Umoru, 1993b; Nielsen, 1994 Rhoptromeris miris Belizin Oscinella frit Chao (Chien ming), 1959; Chukanova & Kovalchuk, 1974

39 Table 2 (Continued). Parasitoids Chloropidae hosts Authors Rhoptromeris strobigena Hapleginella laevifrons Nordlander & Grijpma, 1991 Nordlander & Grijpma Conioscinella gallarum? Gaurax niger? Gaurax ephippium? Rhoptromeris sp. Oscinella frit Chukanova & Kovalchuk, 1974 Chalcidoidea Pteromalidae Callitula bicolor Spinola Oscinella frit Imms, 1930, 1932; Riggert, 1935 Polyodaspis rufijicornis Nikolskaya, 1937; Goodliffe, 1942 Elachiptera cornuta Simmonds, 1952; Hemer, 1960 Chlorops pumilionis Fulmek, 1962, 1968; Zamfirov, 1962 Lasiosina herpini Jones, 1965, 1968b Meromyza nigriventris Chukanova & Kovalchuk, 1974; Dzhanokmen, 1978; Nordlander, 1978b; Dyurich, 1980; Talitskiy & Kuslitskiy, 1990 C. pyrrhogaster (Walker) Oscinella frit Goodliffe, 1942; Simmonds, 1952; Fulmek, 1968; Dzhanokmen, 1978 C. (= Micromelus) rufomaculatus Chlorops pumilionis; Meromyza Tzygankov, 1930 Nees (secondary parasitoid?) nigriventris Cyrtogaster clavicornis Walker Oscinella frit Chukanova & Kovalchuk, 1974 C. vulgaris Walker Coelinidea nigra Balachowsky & Mesnil, 1935; Riggert, 1935; Simmonds, (hyperparasitod) 1952; Hemer, 1960; Fulmek, 1962, 1968; Jones, 1968b; Chukanova & Kovalchuk, 1974; Dzhanokmen, 1978; Nordlander, 1978b Eupelmes artropurpureus Dalman Chlorops pumilionis; Lasiosina Dzhanokmen, 1978 herpini Eupteromalus hemipterus Walker Chlorops pumilionis Hemer, 1960 Oscinella frit Graham, 1969; Dzhanokmen, 1978 Halticoptera aenea Walker Oscinella frit Meyer, 1924; Mejer, 1929; Vojnovskaya-Kriger, 1929; Imms, 1930; Nikolskaya, 1937; Simmonds, 1952; Chao (Chien ming), 1959; Györfi, 1962; Chukanova & Kovalchuk, 1974; Ivatsik & Kovalchuk, 1975; Dyurich, 1980; Talitskiy & Kuslitskiy, 1990 H. circulus (Walker) (= fuscicornis Oscinella frit Cunliffe, 1921; Meyer, 1923; Vojnovskaja-Kriger, 1929; Walker) (= petiolata Thomson) Schander & Meyer, 1925 (= suilis Walker) Lasiosina herpini Imms, 1930, 1932; Riggert, 1935; Simmonds, 1952; Jones, 1965,1968b;Fulmek,1968;Ivatsik&Kovalchuk,1975; Nordlander, 1978a, 1978b; Dyurich, 1980; Howell, 1982; Moore, 1983, 1984; Talitskyi & Kuslitskiy, 1990; Umoru, 1993b; Nielsen, 1994 ?H. hieracii (Thomson) Chlorops pumilionis Fulmek, 1962, 1968 H. patellana (Dalman) Thomson Oscinella frit Fulmek, 1968 Homoporus (= Merisus) Oscinella frit Wilhelm, 1891; Collin, 1918; Meyer, 1923; Mejer, 1929 destructor (Say) (= intermedius Lindeman) Elachiptera cornuta Vojnovskaya-Kriger, 1929; Imms, 1930; Riggert, 1935; Simmonds, 1952; Fulmek, 1968; Chukanova & Kovalchuk, 1974; Dzhanokmem, 1978 H. (M.) febriculosus (Girault) Lasiosina herpini Nikolskaya, 1937 H. (M.) mordellistenae Crawford Lasiosina herpini Nikolskaya, 1937 Lamprotatus chrysochlorus Lipara lucens; L. rufijitarsis Séguy, 1934; Fulmek, 1968 Walker Merismus nitidus Walker Lipara pullitarsis galls Dely-Draskovits et al., 1993

40 Table 2 (Continued). Parasitoids Chloropidae hosts Authors Merisus sp. Oscinella frit Chukanova & Kovalchuk, 1974 Mesopolobus (Amblymerus)sp. Dicraeus ingratus; D. tibialis; D. Moiseev, 1950; Agafonova, 1962 agropyri; D. humeralis Pachyneuron concolor Förster Oscinella frit Chukanova & Kovalchuk, 1974 P. formosum Walker Lipara lucens Giraud, 1863; Séguy, 1934 L. rufijitarsis Fulmek, 1968 P. umbratum Delucchi Oscinella frit Dzhanokmen, 1978 Pachyneuron sp. Oscinella frit Chukanova & Kovalchuk, 1974 ?Polycystus oscinidis Kurdjumov Oscinella frit Mokrzecki, 1913; Collin, 1918; Meyer, 1923; Imms, 1930; Riggert, 1935; Simmonds, 1952; Fulmek, 1968 ?Pteromalus muscorum Walker Chlorops pumilionis Lampa, 1888 P. planiscuta Thomson Oscinella frit Mejer, 1929; Simmonds, 1952; Fulmek, 1968 P. puparum Linnaeus Oscinella frit Rörig, 1893; Collin, 1918; Meyer, 1923; Imms, 1930; Riggert, 1935; Simmonds, 1952; Fulmek, 1968 Pteromalus (Habrocytus) sp. Oscinella frit Ruschka & Fulmek, 1915 Chlorops pumilionis Meyer, 1923; Riggert, 1935; Simmonds, 1952; Fulmek, 1962, 1968; Chukanova & Kovalchuk, 1974 Racosina deplanata Bouček Polyodaspis rufijicornis Kiauka & Nartshuk, 1972 Cryptonevra flavitarsis Askew & Shaw, 1978 Elachiptera cornuta Dyurich, 1980; Abraham, 1983; Talitskiy & Kuslitskiy, 1990 Roptrocerus mirus (Walker) Chlorops pumilionis Fulmek, 1962, 1968 ?Semiotelus nigripes Lindeman Oscinella frit Wilhelm, 1891; Collin, 1918; Riggert, 1935; Simmonds, 1952; Hemer, 1960 Spalangia fuscipes Nees Oscinella frit Nikolskaya, 1937; Simmonds, 1952; Bouček, 1963; Fulmek, 1968;Kovalchuk,1969;Chukanova&Kovalchuk,1974; Ivatsik & Kovalchuk, 1975; Dzhanokmen, 1978; Nordlander, 1978b; Dyurich, 1980; Marshakov, 1983; Talitskiy & Kuslitskiy, 1990 S. nigra Latreille Oscinella frit Riggert, 1935; Fulmek, 1968 Spalangia sp. (?S. nigra) Oscinella frit Simmonds, 1952 Sphegigaster sp. Oscinella frit Chukanova & Kovalchuk, 1974 Stenomalina communis (Nees) Chlorops pumilionis Lampa, 1888; Ruschka & Fulmek, 1915 (= continua Walker) (= rugosa Thomson) (= laetus Ruschka) C. geminatus Kearns, 1931; Balachowsky & Mesnil, 1935 Meromyza pratorum Nikolskaya, 1937 M. nigriventris Fulmek, 1962, 1968; Graham, 1969; Dyurich, 1980; Talitskiy & Kuslitskiy, 1990 S. epistena Walker Chloropidae Dzhanokmen, 1978 S. illudens (Walker) Chlorops pumilionis Lampa, 1888 (= crassicornis Dalman) S. liparae (Giraud) Lipara galls Giraud, 1863; Wagner, 1907; Blair, 1932, 1944b L. lucens; L. similis Otten, 1940; Ruppolt, 1956–1957 L. rufijitarsis Gracham & Claridge, 1965 L. pullitarsis Mook, 1967, 1971; Fulmek, 1968; Waitzbauer, 1969; Chvála et al., 1974; Dzhanokmen, 1978; Skuhravý, 1981; Abraham & Carstensen, 1982; De Bruyn, 1987; Wolf, 1988; ?Gromysz-Kałkowska & Grochowska, 1992; Dely-Draskovits et al., 1993, 1994; Athen & Tscharntke, 1999 S. micans (Olivier) Oscinella frit Gureau, 1861; Porchinsky, 1881 Chlorops pumilionis Rörig, 1893; Schesterikov, 1910 Meromyza nigriventris Kurdjumov, 1912; Collin, 1918

41 Table 2 (Continued). Parasitoids Chloropidae hosts Authors Lasiosina herpini Frew, 1923a; Meyer, 1923; Imms, 1930; Tzygankov, 1930; Séguy, 1934; Balachowsky & Mesnil, 1935; Riggert, 1935; Nikolskaya, 1937; Goodliffe, 1939, 1942; Simmonds, 1952; Fulmek, 1962, 1968; Hemer, 1962; Gracham & Claridge, 1965; Chukanova & Kovalchuk, 1974; Dzhanokmen, 1978 S. muscarum Linnaeus Coelinidea nigra Wahlgren, 1918; Tzygankov, 1930; Balachowsky & Mesnil, (hyperparasitoid) 1935; Fulmek, 1962, 1968 Stenomalina sp. Chlorops pumilionis Fulmek, 1962, 1968; Dyurich, 1980; Talitskiy & Kuslitskiy, 1990 Trichomalopsis micropterus Oscinella frit Nikolskaya, 1937; Simmonds, 1952; Fulmek, 1968; Lindeman Chukanova & Kovalchuk, 1974; Dyurich, 1980; Talitskiy & Kuslitskiy, 1990 Trichomalopsis sp. Oscinella frit Chukanova & Kovalchuk, 1974 Trichomalus cristatus (Förster) Oscinella frit Baranov, 1912; Collin, 1918 Thaumatomyia notata Meyer, 1923; Schander & Meyer, 1925; Mejer, 1929; Vojnovskaja-Kriger, 1929; Riggert, 1935; Nikolskaya, 1937; Simmonds, 1952; Chao (Chien ming), 1959; Fulmek, 1968; Chukanova & Kovalchuk, 1974; Ivatsik & Kovalchuk, 1975 T. nanus (Walker) (= frontalis Oscinella frit Meyer, 1923; Riggert, 1935 Thomson) Chloropidae Fulmek, 1968; Dzhanokmen, 1978; Nordlander, 1978a, 1978b T. posticus Walker Oscinella frit Dzhanokmen, 1978; Dyurich, 1980 Chloropidae Talitskiy & Kuslitskiy, 1990 T. statutus (Förster) Oscinella frit Nartshuk, 1962a Pseudopachychaeta rufijiceps Graham, 1969 Chloropidae Dzhanokmen, 1978 Trichomalus sp. Oscinella frit Ivatsik & Kovalchuk, 1975 Eupelmidae Eupelmus atropurpureus Salman Chlorops pumilionis Tzygankov, 1930 Lasiosina herpini Nikolskaya, 1937 Meromyza nigriventris Fulmek, 1962, 1968 Eupelmes microzonus Förster Chlorops pumilionis Tzygankov, 1930 Lasiosina herpini Nykolskaya, 1937 Meromyza nigriventris Fulmek, 1962, 1968 Eupelmella vesicularis Retzius Chlorops pumilionis Tzygankov, 1930 Meromyza nigriventris Nikolskaya, 1937; Fulmek, 1962, 1968 Eurytomidae Eudecatoma fasciata (Thomson) Lipara lucens Fulmek, 1968 E. nartshukae Zerova Lipara lucens Zerova, 1977 E. palustris Erdös (= paludicola Eurina ducalis; Lipara lucens Zerova, 1978 Zerova) E. phragmiticola Zerova Lipara galls Zerova, 1978 Eurytoma lucidula Zerova Chlorops strigulus Zerova, 1995 ?Tetramesa phragmitis (Erdös) Lipara galls; preferable L. Erdös, 1957a; Dely-Draskovits et al., 1993 rufijitarsis Torymidae Torymus arundinis Walker Lipara similis Fulmek, 1968 Eulophidae Aprostocetus arundinis (Giraud) Lipara lucens Valkeila, 1959; Fulmek, 1968; Tryapitsyn & Kostyukov, 1978 A. ciliatus (Nees) Lipara lucens Fulmek, 1968 A. pausiris (Walker) Lipara lucens Wolf, 1988, 1991

42 Table 2 (Continued). Parasitoids Chloropidae hosts Authors Aprostocetus sp. L. lucens; L. similis; L. rufijitarsis Dely-Draskovits et al., 1994 Closterocerus lagus (Walker) Oscinella frit Howell, 1982; Moore, 1983, 1984; Nielsen, 1994 Digliphus sp. Oscinella frit Chukanova & Kovalchuk, 1974 Hemiptarsenus sp. Oscinella frit Chukanova & Kovalchuk, 1974 Horismeniu specularis Erdös Chloropidae Tryapitsyn & Kostjukov, 1978 Kostjukovius platycephalae Platycephala umbraculata Dyurich & Kostjukov, 1978; Dyurich, 1980; Talitskiy & Kostjukov Kuslitskiy, 1990 Mellitobia sp. Lipara lucens Abraham & Carstensen, 1982 L. pullitarsis Necremnus leucarthros Nees Oscinella frit Hemer, 1962 Neochrysocharis albipes Oscinella frit Simmonds, 1952; Fulmek, 1968 Kurdjumov N. immaculatus Kurdjumov Oscinella frit Kurdjumov, 1912; Meyer, 1923; Mejer, 1929; Imms, 1930; Riggert, 1935; Nikolskaya, 1937; Simmonds, 1952; Fulmek, 1968; Tryapitsyn & Kostjukov, 1978 Pediobius epigonus (Walker) Oscinella frit Tryapitsyn & Kostjukov, 1978 P. facialis (Giraud) Lipara lucens galls Giraud, 1863; Fulmek, 1968 L. similis Dely-Draskovits et al., 1994 (hyperparasitoid of Ephialtes arundinis) L. rufijitarsis P. metallicus (Nees) Oscinella frit Meyer, 1923; Imms, 1930; Simmonds, 1952; Fulmek, 1968 P. saulius Walker Lipara galls Viggiani, 1964 (hyperparasitoid of Exeristes arundinis) Pediobius (Pleurotropis)sp. Oscinella frit Chukanova & Kovalchuk, 1974 Pronotalia inflata Graham Lipara lucens Graham, 1991; Wolf, 1991 P. liparae Gradwell Lipara lucens Gradwell, 1957; Valkeila, 1959 P. valkeilai Gradwell Calamoncosis minima Gradwell, 1957; Valkeila, 1959 Tetrastichus atroceruleus (Nees) Lipara lucens Fulmek, 1968 T. djuritshae Kostjukov Chlorops novakii Kostjukov, 1979; Talitskiy & Kuslitskiy, 1990 T. legionarius (Giraud) Lipara lucens Giraud,1863;Henriksen,1918–1919 L. similis Otten, 1940; Erdös, 1955, 1956, 1957b L. rufijitarsis Ruppolt, 1956–1957 Cryptonevra flavitarsis Fulmek, 1968; Waitzbauer, 1969 Neohaplegis tarsata Chvála et al., 1974; Dyurich, 1980; Hudek et al., 1981; De Bruyn, 1987; Wolf, 1988; Talitskiy & Kuslitskiy, 1990; Plant, 1997 Tetrastichus sp. Lipara lucens Abraham & Carstensen, 1982 L. pullitarsis Dyurich, 1980 Chlorops geminatus Talitskiy & Kuslitskiy, 1990 C. novakii Tetrastichus sp. Oscinella frit Howell, 1982 Tetrastichus sp. Dicraeus ingratus Agafonova, 1962 D. tibialis Elasmidae Crataepiella liparae Gradwell Lipara lucens Fulmek, 1962; Tryapitzyn & Kostjukov, 1978 C. carlinarum Szelényi & Erdös Calamoncosis minima Gradwell, 1957; Valkeila, 1959 Aphelinidae Aphelinus flavipes Förster Oscinella frit Anonymous, 1963 ?Centrodora amoena Förster Lipara lucens Fulmek, 1968 Trichogrammatidae Trichogramma evanescens Oscinella frit Chernoponevkina, 1939 Westwood Trichogramma sp. Oscinella frit Fulmek, 1968

43 Table 2 (Continued). Parasitoids Chloropidae hosts Authors Signiphoridae Clytina giraudi Erdös Cryptonevra flavitarsis Erdös, 1957a, 1957b, 1958; Fulmek, 1968 C. diadema Tryapitsyn, 1978; Dyurich, 1980 Lipara lucens Talitskiy & Kuslitskiy, 1990 L. rufijitarsis Calamoncosis minima Neochaplegis tarsata Elachiptera cornuta Mymaridae Gonatocerus sulphureus Förster Oscinella frit Schander & Meyer, 1924; Simmonds, 1952; Fulmek, 1968 Ichneumonoidea Ichneumonidae Bathythrix decipiens Gravenhorst Lipara lucens Blair, 1932; Séguy, 1934; Fulmek, 1968 (?hyperparasitoid) ?Campodorus variegates (Jurine) Lipara lucens Fulmek, 1968 (= sanguicollis Gravenhorst) ?Diadegma chrysostictum Lipara lucens Starke, 1940 (Gravenhorst) D. fenestralis Holmgren Oscinella frit Chukanova & Kovalchuk, 1974 Endromopoda arundinator Lipara lucens Giraud,1863;Wagner,1907 (Fabricius) (= melanopyga Gravenhorst) L. rufijitarsis Blair, 1932; Hedwig, 1950, 1955 L. pullitarsis Aerts, 1956; Fulmek, 1962, 1968 L. similis Waitzbauer, 1969; Chvála et al., 1974 Platycephala planifrons Doskočil & Chvála, 1974; Imhof, 1979; Dyurich, 1980; Kasparjan, 1981; Grochowska, 1989; Talitskiy & Kuslitskiy, 1990 E. detrita (Holmgren) Lipara galls Wagner, 1907 L. lucens Henriksen, 1918–1919; Blair, 1932, 1944b L. similis Starke, 1940; Šedivý, 1963; Fulmek, 1968 L. rufijitarsis Waitzbauer, 1969; Raghi-Atri, 1980 E. phragmitidis (Perkins) Lipara lucens Perkins, 1957; Mook, 1967; Waitzbauer, 1969 L. similis Waitzbauer et al., 1973; Chvála et al., 1974 L. rufijitarsis Imhof, 1979; Fitton et al., 1988 Platycephala planifrons Skuhravý & Skuhravá, 1978; Dyurich, 1980; Hudec et al., 1981; Skuhravý, 1981; Talitskiy & Kuslitskiy, 1990; Dely-Draskovits et al., 1993; Reader, 2003 Endromopoda sp. (as Scambus sp.) Lipara galls, primarily L. De Bruyn, 1987 rufijitarsis Exeristes arundinis Kriechbaumer Lipara lucens Henriksen, 1918–1919; Seyrig, 1927 Platycephala planifrons Hellén, 1949; Valkeila, 1949; Fulmek, 1968; Waitzbauer, 1969; Waitzbauer et al., 1973; Kasparjan, 1981; Dely-Draskovits et al., 1994 Gelis liparae (Giraud) Lipara lucens Giraud, 1863 L. rufijitarsis Henriksen, 1918–1919; Fulmek, 1968 G. notabilis (Förster) Lipara lucens Henriksen, 1918–1919; Blair, 1932 Glypta sp. Oscinella frit Fulmek, 1968; Waitzbauer, 1969; Chukanova & Kovalchuk, 1974 Phrygaedeon troglodytes Giraud Lipara rufijitarsis Aerts, 1940 ?Pimpla turionellae Linnaeus Chlorops pumilionis; C. strigulus Fulmek, 1968 Promethes sulcator (Gravenhorst) Oscinella frit Anonymous, 1963 Scambus planatus (Hartig) Lipara lucens Fulmek, 1968

44 Table 2 (Continued). Parasitoids Chloropidae hosts Authors S. sagax (Hartig) Lipara lucens Fulmek, 1968 L. similis S. nigricans (Thomson) Lipara lucens Fulmek, 1968; Dely-Draskovits et al., 1974; Kasparyan, 1981 (= similis Bridgman) (var. habermehli Schmideknecht) S. nucum Ratzeburg Lipara lucens Starke, 1940 Braconidae ?Aleiodes rufijicornis Lipara lucens Starke, 1940; Hedwig, 1950; Fulmek, 1968 (Herrich-Schäffer) (= Rhogas dimidiatus Spinola) Baryproctus barypus (Marshall) Lipara similis Dyurich, 1980 L. rufijitarsis Talitskiy & Kuslitskiy, 1990; Dely-Draskovits et al., 1994 ?Blacus (B.) interstitialis Ruthe Oscinella frit Fischer, 1963; Haeselbarth, 1973; Achterberg, 1988; Koponen, 1991, 1993 Bracon discoideus Wesmael Chlorops pumilionis Fulmek, 1968 Bracon (B.) longicollis Wesmael Chlorops pumilionis Kearns, 1931; Fulmek, 1962, 1968 B. (B.) longulus Thomson Oscinella frit Ivatsik & Kovalchuk, 1975 Bracon (B.) minutator (Fabricius) Lipara lucens Henriksen, 1918–1919 (= abscissor Nees) Lipara galls Séguy, 1934; Fulmek, 1968 Bracon sp. Chlorops pumilionis Fulmek, 1962, 1968 Bracon (Glabrobracon) discoideus Chlorops pumilionis Fulmek, 1962, 1968 Westmael ?Cardiochiles saltator (Fabricius) Chlorops pumilionis Porchinsky,1881;Fulmek,1962,1968 var. brachialis Rondani Chasmodon apterus (Nees) Oscinella frit Cunliffe, 1921; Vojnovskaya-Kriger, 1929 Oscinella vastator; O. frit Riggert, 1935; Simmonds, 1952 Meromyza saltatrix Jones, 1965, 1969b; Fulmek, 1968 Lasiosina herpini Dyurich, 1980; Moore, 1983, 1984; Moore et al., 1986; Moore & Hunt, 1987; Moore & Ridout, 1987; Talitskiy & Kuslitskiy, 1990; Umoru, 1993a, 1993b Chorebus lar Morley Oscinella frit Dyurich, 1980 Chorebus sp. aff. ampliator (Nees) Oscinella frit Dyurich, 1980; Talitskiy & Kuslitskiy, 1990 Chorebus sp. Oscinella frit Chukanova & Kovalchuk, 1974 C. (Gyrocampa) pospelovi Oscinella frit Kurdjumov, 1912; Meyer, 1923; Schander & Meyer, 1925; Kurdjumov Riggert, 1935 Coelinidea nigra (Nees) Oscinella frit Goureau, 1861; Lampa, 1888 Chlorops pumilionis Marshall, 1891; Rörig, 1893 C. novakii; C. strigulus Wahlgren, 1918; Znamensky, 1923 C. laetus Tzygankov, 1930 Lasiosina herpini Kearns, 1931; Séguy, 1934 Meromyza pratorum Balachowsky & Mesnil, 1935 M. nigriventris Goodliffe, 1939, 1942; Fulmek, 1962, 1968; Anonymous, 1963; Dyurich, 1980; Talitskiy & Kuslitskiy, 1990 C. vidua (Curtis) Chlorops pumilionis Fulmek, 1962, 1968 ?Coeloides abdominalis Lipara lucens Fulmek, 1968 (Zetterstedt) ?C. melanostigma Strand Lipara lucens Starke, 1940; Hedwig, 1950; Fulmek, 1968 (= stigmaticus Hell) Dacnusa areolaris (Nees) Chlorops pumilionis Ruschka & Fulmek, 1915; Fulmek, 1962, 1968 Dacnusa sp. Chlorops pumilionis Fulmek, 1962, 1968 ?Gonatocerus sulphuripes Förster Oscinella frit Schander & Meyer, 1925 Microgaster sp. Oscinella frit Chukanova & Kovalchuk, 1974

45 Table 2 (Continued). Parasitoids Chloropidae hosts Authors Opius caesus Haliday Oscinella frit Anonymous, 1963 O. irregularis Wesmael Oscinella frit Anomymous, 1963 O. maculipes Wesmael Oscinella frit Simmonds, 1952; Fulmek, 1968; Dyurich, 1980; Talitskiy & Kuslitskiy, 1990 O. piceus Thomson Oscinella frit Anonymous, 1963 Opius sp. Oscinella frit Bhaattacharyya, 1957; Umoru, 1993b Phaenocarpa helophiliae Calamoncosis glyceriae Achterberg, 1998 Achterberg Polemochartus aboletus Papp Lipara similis Papp, 1992; Dely-Draskovits et al., 1993, 1994; Umoru, 1993a, 1993b; Nielsen, 1994 P. breviventris (Telenga) Lipara sp. Papp, 1992 P. liparae (Giraud) Lipara galls Giraud, 1863; Marshall, 1891 Preferable L. lucens Reynvaan & Docters van Leeuwen, 1906 L. pullitarsis Wagner, 1907; Blair, 1932, 1944b L. similis Seguy, 1934; Starke, 1940 L. rufijitarsis Erdös, 1955, 1957a, 1957b; Ruppolt, 1956, 1957; Mook, 1961, 1967;Fulmek,1968;Waitzbauer,1969;Waitzbaueretal., 1973; Chvála et al., 1974; Imhof, 1979; Skuhravý, 1981; Abraham & Carstensen, 1982; Maetô, 1983b; De Bruyn, 1987, 1994a, 1994b; Wolf, 1988; Dely-Draskovits et al., 1993, 1994 P. melas (Giraud) Lipara galls Giraud, 1863; Marshall, 1891 Preferable L. rufijitarsis Reynvaan & Docters van Leeuwen, 1906; Fulmek, 1968; Waitzbauer et al., 1973, 1974 Platycephala planifrons Chvála et al., 1974; Skuhravý, 1981; Maetô, 1983b; De Bruyn, 1987; Dely-Draskovits et al., 1993, 1994 Protodacnusa tristis (Nees) Oscinella frit Ruschka & Fulmek, 1915 Chlorops pumilionis Vojnovskaya-Kriger, 1929; Riggert, 1935; Simmonds, 1952; Fulmek, 1962, 1968 Sarops rea Nixon Cryptonevra sp. Maetô, 1983a C. flavitarsis Dyurich, 1980; Talitskiy & Kuslitskiy, 1990 Sigalphus caudatus (Curtis) Oscinella frit Curtis, 1860 ?Triaspis caudata (Nees) Oscinella frit Rörig, 1893; Collin, 1918 Chlorops pumilionis Meyer, 1923; Riggert, 1935; Simmonds, 1952; Fulmek, 1962, 1968 Triaspis obscurella Nees Oscinella frit Ivatsik & Kovalchuk, 1975 Aphidiidae ?Aphidius avenae Haliday Oscinella frit Cunliffe, 1921; Simmonds, 1952; Fulmek, 1968 (= granarius Marshall) ?Aphidius sp. Oscinella frit Chukanova & Kovalchuk, 1962

Notes. A question mark (?) in front of the name of a parasitoid indicates that this parasitoid probably came from some other insect host living in the same infected stem as the chloropid larva. Thename“Oscinella frit” refers to any species within O. frit complex. Only currently accepted names for the parasitoid species are used in the table, meaning that theser may differ substantially from those given in the original papers. The names of parasitoids were checked by Dr S.A. Belokobylsky (Braconidae), Dr A.I. Khalaim (Ichneu- monidae), Dr O.V. Kovalev (Cynipoidea), Dr M.A. Kozlov (Proctotrupoidea), Dr V.A. Tryapitzyn (Chalcidoidea) (all Zoological Institute Russian Academy of Sciences, S.-Petersburg, Russia), Dr Z.A. Efremova (Eulophidae) (Ulyanovsk, Russia), Dr K.A. Dzhanokmen (Alma- Ata, Kazakhstan) and Dr M.D. Zerova (Schmalhausen Institut of Zoology, Kiev, Ukraine).

46 Ryzhkova (1962) found in the salivary glands of bacteria found in frit flies are considered as and other organs of larvae of Oscinella frit and O. pathogenous. Of these, 26 species have been re- pusilla symbiotic bacteria which in her opinion corded from chloropid eggs, larvae, pupae and are involved in the digestion of plant tissues. Later, adults (Aleksandrov et al., 1969; Aleksandrov & Sasonov (1966) investigated in detail the relation- Fedosov, 1971; Gritsenko & Fedosov, 1972). Only ship between micro-organisms and larvae of Os- freshly laid eggs are sterile. The microflora of the cinella species. He found no microorganisms in the larvae is formed by bacteria occurring in plant tis- salivary glands of the larvae using both light and sues. Some bacteria are transmitted from larvae to electron microscope, but the bacteria Pseudomo- adults. Adults taken in the field contain up to 19 nas fluorescens, P. mycophaga and Chromobacter species of bacteria, adults hatching from puparia flavum were found in the larval gut and Bacterium in the laboratory up to 5 species (Gritsenko & Fe- ciliaerogenas on the larval body. He also found the dosov, 1972). following bacteria in plant tissues fed upon by the Two species of entomophagous fungi have been larvae: Mycobacterium globiforme, Pseudomonas found in England on two species of Chloropidae: fluorescens, P. herbicola, Bacterium aquatile acidi- Entomophthora americana on Lipara lucens and fijicans, B. candicans lactium. These epiphytic and E. culicis on Thaumatomyia glabra (Leatherdale, phytopatogenous bacteria are known as decom- 1966, 1970). posers of complex plant proteins. Another group

Distribution and zoogeography

The family Chloropidae is worldwide in distribu- Becker, Homalura Becker, Metopostigma Becker, tion. Some Palearctic species extend northwards to Paraeurina Duda, Phyladelphus Becker and Platy- the coast of the Arctic Ocean (Nartshuk, 2002b). cephalisca Nartshuk) out of 28 do not reach North Three species: Oscinella frit s.l., Cetema cereris and Europe. All missing genera except Homalura and Chlorops hypostigma are recorded from Faroe Is- Platycephalisca have a Mediterranean distribution. lands, one species: Oscinella frit from Iceland, and In comparison to the 203 species of chloropids no species from Greenland (Mazon, 1890; Mei- found in Fennoscandia and Denmark, the follow- jere, 1910; Becker, 1915; Nielsen & Tuxen, 1954; ing species numbers can be given for some Eu- Lyneborg, 1968; Lindroth et al., 1973; Messersmith, ropean countries: Great Britain – 175 species (Is- 1982; Ólafsson, 1991). Mazon (1890) mentioned also may, 1998), Germany – 208 (Wendt, 1999; Schu- Madiza palpora (sic!) Fln., now Lasiambia palposa mann, 2002), Belgium – 112 (De Bruyn & Tschirn- (Fallén) from Iceland, but this has no been con- haus, 1991), Netherlands – 140 (Beuk & Nartshuk, firmed. Recently 484 specimens of Chloropidae 2002), Poland – 170 (Nartshuk, 2002a), Latvia – were collected in Iceland (Prescher et al., 2005) 115 (Karpa, 2001), Lithuania – 97 (Nartshuk & that all proved to belong to O. frit (M. v. Tschirn- Pakalniškis, 2004), Hungary – 217 (Dely-Draskovits, haus, pers. comm.). 2001), Schwitzerland – 134 (Dely-Draskovits, 1998), The diversity of chloropids in terms of genera Czech and Slovak Republics – 151 (Roháček, 1997), and species increases southwards in Europe and Bulgaria – 138, all Balkan Peninsula – 180 (Bescho- remaining parts of the Palearctic Region. In Fenno- vski, 1985), Italy – 190 (Nartshuk, 1995), Spain – 91 scandia and Denmark are presently recorded 203 (De Bruyin & Báez, 2002; Nartshuk, 2004). species in 48 genera. This is nearly 50% of the Euro- The chloropid fauna of Fennoscandia and Den- pean species, but 80% of the genera. The subfamily mark is composed of species with very different Siphonellopsinae with two European genera is ab- total distribution ranges: (1) widespread Old World sent in North Europe. In Oscinellinae with 27 Euro- species found in the Palearctic, Afrotropical and/or pean genera only Aprometopis Becker is not found Oriental Regions – 5 (2.4%), Holarctic – 17 (8.3%), in North Europe. In the Chloropinae 8 genera (As- trans-Palearctic boreal – 67 (33.3%), south- suania Becker, Capnoptera Loew, Elachiptereicus Palearctic eremial (steppes and semideserts) – 3

47 (1.4%), Eurasian – 43 (21.1%), Euro-Mediterranean 1. This group includes pronouncedly southern 12 (5.9%), European (including Euro-Caucasian) – (nemoral) species extending no further north 55 (27.1%), and Mediterranean-Atlantic (coastal) – than Denmark and the southernmost provinces 1 species (0.5%). ofSweden(SK,HA,SM,Öl,GO).Itconsistsof1 The actual distribution of the different chloropid species of Rhodesiellinae, 13 species of Oscinel- species occurring in Fennoscandia and Denmark linae and 15 species of Chloropinae. Some of the is insufficiently known, especially in Norway, Kare- southern species have only been found on is- lia and the Kola Peninsula. Still, some general re- lands: Dicraeus raptus, Chlorops fijinitimus, C. pal- marks may be in place here. A preliminary anal- lidiventris, C. pannonicus, Chloropsina rohaceki, ysis of species distributions within the subfamily Neohaplegis glabra on Öland, D. ingratus on Chloropinae was given earlier (Nartshuk, 2002b). Bornholm and Gotland. This group mainly in- The area of present Fennoscandia and Denmark cludes species with Euro-Mediterranean, some was covered entirely with ice during the Riss glacial with coastal or eremial habitat preferences. period, and only the western part of Jutland (Den- Eutropha fulvifrons is a coastal, atlantic- mark) was ice free during the following Würm Mediterranean species that properly belongs in glacial period. Recolonization of this territory took this group despite a record from NB. place gradually during the retreat of the glacier, 12– 2. This group includes species distributed north- 10 000 years ago. The latitudinal extend of Fenno- wards to the 60°N latitude in Fennoscandia, scandia and Denmark is impressive (nearly 18 de- which approximately coincides with the north- grees from the southern border of Denmark to the ern limit of the boreo-nemoral zone: in Swe- Norwegion coast at the Arctic Ocean) and corre- den to VR and UP, in Finland to Al and Ka, in sponds to the latitudinal extend of the remaining Russia to Vib and southern Kr. A diverse group European territory. The main route of coloniza- consisting of 23 species of Oscinellinae and 20 tion of Fennoscandia came from the south through species of Chloropinae. This group mainly in- Denmark. cludes species with European, Euro-Caucasian As for many other insect groups it is evident and Euro-Mediterranean distributions. that most chloropid species occurring in the north- 3. This group includes species extending north- ern parts of Fennoscandia have Holarctic or trans- wards approximately to the 66°N latitude in Palearctic total distributions, whereas species con- Fennoscandia, which approximately coincides fined to Denmark and the southern parts of Fenno- with the northern limit of the middle boreal scandia have a stronger representation of primarily zone: in Sweden to LU and NB, in Finland to Lk. European and Euro-Mediterranean species. As in This is the largest group consisting of 19 species most other groups of organisms the species diver- of Oscinellinae and 38 species of Chloropinae. sity of Chloropidae decrease gradually to the north Some of the species, e.g. Trichieurina pubescens, in Fennoscandia. Thus, for most species the north- Meromyza sibirica and Neohaplegis tarsata,may ern border of their European distribution range well extend further north, but there is no data goes across Fennoscandia at some latitude. Only yet. This group includes the diverse fauna of 39 species, or 19.2% of the Scandinavian Chloropi- chloropids found associated with Common reed dae have been found north of the Arctic circle, and (Phragmites australis),aplantwithanorthern most of these have much wider distributions fur- range coinciding well with that of the present ther southwards in Fennoscandia and Denmark. group 3. There are surprisingly few examples of chloropid 4. This group includes the most widely distributed species with exclusively high boreal to subarc- species in Fennoscandia embracing the north- tic distributions in Fennoscandia: Conioscinella ern boreal zone, some of them (Oscinella frit, abiskoi, C. tornensis and C. messaurea and Aphan- O. ventricosi, Chlorops planifrons, C. scutellaris, otrigonum norrbotticum.TheConioscinella species C. varsoviensis, Melanum laterale, Pseudopachy- all belong to the C. frontella species complex, which chaeta rufijiceps, Thaumatomyia trifasciata) even suggests relatively recent speciation. subarctic tundra (Nartshuk, 2002b). The group Most chloropid species of Fennoscandia and is well represented by 19 species of Oscinellinae Denmark can be referred to one of the following and 17 species of Chloropinae. This group con- four groups determined by the position of their sists mostly of widely distributed Holarctic or northernmost distribution range: trans-Palearctic species

48 High latitude melanism. In several widely dis- rax, especially females, are often almost wholly tributed species of Chloropidae it is very evident black, and the usually discrete black scutal stripes that specimens from the northern boreal zone of seen in Chlorops speciosus and C. meigenii tend to Fennoscandia tend to be significantly more dark- widen and fuse together in northern specimens. coloured as normal. Northern specimens of Gau-

Taxonomy

Monophyly and relationships in the superfamily Chloropoidea based on the ob- servation by Sturtevant (1926) that the females of The Chloropidae is a well defined family belonging both families possess a ventral pocket-like sem- to the Schizophora, a specious higher taxon tradi- inal receptacle and desclerotized spermathecae. tionally divided in two major groups of families: Ca- A year later Hennig (1972) regretted this opinion, lyptratae and . Chloropids belong to because sclerotized spermathecae were found in the acalyptrate grade. The monophyly of the fam- Carnidae, a family he considered most closely re- ily is widely accepted. Apomorphies for the family lated to . Therefore, the absence of scle- after Andersson (1977): rotized spermathecae in Milichiidae and Chlorop- idae might be considered as a homoplasy. 1. First and second pairs of orbital bristles latero- Speight (1969) made a comprehensive compar- proclinate. ative study of the prosternum in Acalyptratae. He 2. Propleuron with vertical carinate ridge found that Chloropidae possess a well developed 3. “Tibial organ” present on hind tibia. prosternal bridge and proposed the inclusion of the 4. Vein Sc reduced in apical half. family in Drosophiloidea together with Drosophil- 5. Cross-vein bm-cu absent. idae (including Camillidae), Diastatidae, Canaci- 6. Vein CuA1 slightly sinuate near middle of penul- dae, Ephydridae, Tethinidae and Braulidae. Hen- timate sector nig (1972) argued that antennal morphology spoke 7. Cell cup absent. against uniting the Chloropidae and Milichiidae 8. Male tergite 6 absent. with Drosophilidae. However, Hackman & Väisä- nen(1985),basedoncharactersofwingveinC, Additional character states have been suggested followed Speight (1969) and included Chloropidae as chloropid apomorphies by earlier authors, e.g. in Drosophiloidea. Griffiths (1972), in a compar- Malloch (1934), Hennig (1958) and Griffiths (1972), ative study of male terminalia of cyclorrhaphous but they rather represent apomorphic states rela- Diptera, considered the Chloropidae most closely tive to the ground plan of all acalyptrate families. related to the families Acartophthalmidae, Milichi- The relationship of the Chloropidae to the nu- idae and Carnidae. Wheeler (1994c) supported a merous other acalyptrate families has for long sister group relationship between Chloropidae and been controversial. Several earlier authors placed Milichiidae and rejected a closer relationship be- Chloropidae as an isolated group of “higher” Aca- tween these families and the Risidae and Crypto- lyptratae (Malloch, 1948; Oldroyd, 1954; Hennig, chetidae. Brake (2000), in a cladistic analysis of 1958; Harrison, 1959). Hennig (1958) was uncer- Milichiidae, supported the idea of a sister group re- tain about the position of the Chloropidae. He sug- lationship between Chloropidae and Milichiidae. gested on one hand that the family might belong to She also presented a detailed review of previous ac- either Drosophiloidea or Milichioidea, but ended counts on the relationships between Milichiidae, up with erecting a superfamily Chloropoidea with Carnidae and Chloropidae. a single family Chloropidae. Later, Hennig (1969) It should be emphasized finally that the phylo- placed the family as a questionable member of the genetic relationships of Chloropidae are far from superfamily Milichioidea together with Milichi- being elucidated. The sister group of Chloropidae idae and Carnidae without specified evidence. may not be found without a detailed analysis of Hennig (1971) united Milichiidae and Chloropidae species of the big genus Apotropina Hendel (sub-

49 Table 3. Assignments of the family Chloropidae in different classifications of Diptera. Taxon Stone et al., 1965 Delfinado & Hardy, 1977 Steyskal, 1974* Griffiths, 1972* Suborder Cyclorrhapha Brachycera Cyclorrhapha Infraorder Division Schizophora Schizophora Sectio Acalyptratae Superfamily Chloropoidea Muscoidea Chloropoidea Prefamily Tephritoinea Included families CHLOROPIDAE CHLOROPIDAE, CHLOROPIDAE, Milichiidae, Acartophthalmidae, Carnidae Milichiidae, Carnidae Crosskey, 1980 Evenhuis, 1989 McAlpine, 1989 Suborder Cyclorrhapha Cyclorrhapha Brachycera Infraorder Division Schizophora Schizophora Acalyptratae Schizophora Sectio Acalyptratae Superfamily Drosophiloidea Ephydroidea Ephydroidea Prefamily Family group Included families CHLOROPIDAE, Camillidae, CHLOROPIDAE, Curtonotidae, CHLOROPIDAE, Camillidae, Ephydridae, Diastatidae, Drosophilidae, Ephydridae, Ephydridae, Diastatidae, Curtonotidae, Drosophilidae, Cryptochetidae, Tethinidae, Curtonotidae, Drosophilidae, Milichiidae, Carnidae, Tethinidae, Canacidae, Neurochaetidae Cryptochetidae, Tethinidae, Canacidae Canacidae

Yeats & Wiegmann, 1999 Pape & Thompson, 2010 Suborder Brachycera Cyclorrhapha Division Muscomorpha Schizophora Sectio Schizophora Acalyptratae Acalyptratae Superfamily Carnoidea Included families CHLOROPIDAE, Canacidae, CHLOROPIDAE, Carnidae, Cryptochetidae, Acartophthalmidae, Milichiidae, Tethinidae Australimyzidae, Braulidae, Canacidae, Carnidae, Cryptochetidae, Milichiidae

*Note. In the classifications of Griffiths, 1972 and Steyskal, 1974 a number of taxa between suborder and superfamily has been omitted.

family Siphonellopsinae) showing the most basic and M. saltatrix byLinnaeus(1758,1761),followed characters of the family. The genus is mainly trop- by M. umbelliferarum by Scopoli (1763), M. hordei, ical and out of scope of our work. Therefore we M. pumilionis and M. avenae by Bjerkander (1777, only summarize how the family has been classi- 1778, 1781), M. lineata, M. minuta, M. umbraculata fied in catalogues and manuals of Diptera from dif- and M. planifrons by Fabricius (1781, 1787, 1794, ferent zoogeographical regions published the last 1798), and M.nasuta by Schrank (1781). Five of these decades to show diverse opinions (Table 3). names are still valid and include some notorious pest species of cereals and cultured grasses that were first described from Sweden. Classification and nomenclature The oldest names for chloropid genera are Chlo- rops proposed by Meigen (1803) and Oscinis by The first species of the present family Chloropi- Latreille (1804). Fallén (1820) revised the Swedish dae described with valid binominal names were fauna of these flies and referred them to their own classified in the genus Musca Linnaeus: Musca frit family Oscinidae (as “Oscinides”) based on La-

50 treille’s name. It was established much later that which, in combination with the size of the group, Oscinis Latreille is a junior synonym of Chlorops makes morphology based cladistic analysis of the Meigen. For that reason Becker (1909, 1910) pro- family very difficult. He proposed a phylogeny for posed the genus Oscinella for Oscinis auct., nec the family (Andersson, 1979), but stressed that it Latreille, and replaced the family name Oscinidae was weakly supported and implied many instances with Chloropidae even if it is younger, as first pro- of homoplasy. posed by Rondani (1856) in the family Agromyzi- In the present work we divide the family Chlorop- dae as one of two chloropid “stirps” [groups] Chlo- idae in three subfamilies: Rhodesiellinae, Oscinelli- ropina and Oscinina. Schiner (1864) adopted the nae and Chloropinae. A fourth subfamily Siphonel- name Chloropina, but treated it as a subgroup of lopsinae is extralimital and is not discussed fur- the family Muscidae. The name Chloropidae is now ther here. Rhodesiellinae was earlier proposed as a firmly established and valid for the family accord- subfamily but without much argument (Nartshuk, ing to ICZN Art. 40.2. It is correctly cited with au- 1983). It is plesiomorphic in terms of Chloropinae thor and year as Chloropidae Rondani, 1856 (1820) by having costal vein extended to vein M, and ple- according ICZN Recommandation 40A. siomorphic in terms of Oscinellinae by having long, Lioy (1864) divided his family “Heteromyziti” in procurved and divergent ocellar setae. seven subfamilies, four of them based on generic Rhodesiellinae and Oscinellinae share the fol- names of present Chloropidae: Elachipterini, Si- lowing characters: phonellini, Meromyzini, and Chloropsini, but this classification was not followed by subsequent au- 1. Costal vein goes to M (plesiomorphy). thors. Becker (1910) rather followed Rondani by 2. Ocellar triangle large (not present in all Oscinel- dividing the family in two subfamilies: Oscinellinae linae) (apomorphic). and Chloropinae. Unfortunatly, the family-group 3. Scutellum elongated with apical setae inserted names Elachipterini Lioy, 1864 and Siphonellini on tubercles (only in a few species of Oscinelli- Lioy, 1864 have priority over Oscinellinae Becker, nae) (apomorphic). 1910. Therefore, an application to ICZN is required 4. Femoral organ present (apomorphic). to preserve the well established name Oscinellinae 5. Hypandrium closed (only in a few species of Os- for the subfamily. cinellinae). Duda (1930) divided Choropidae in three sub- Rhodesiellinae and Chloropinae share the fol- families: Oscinellinae, Chloropinae and Hippelati- lowing characters: nae. Later, after inclusion of exotic forms, he split the family in two subfamilies: Siphonellopsinae 1. Ocellar setae long, procurved and divergent and Chloropinae. The latter subfamily he split in (plesiomorphic). two tribes Chloropini and Oscinellini; the latter 2. Tibial organ absent (except in some Chloropi- tribe was then split in two superfamilies: Hippela- nae) (apomorphic). toidea and Oscinelloidea. Andersson (1977) pro- 3. Sclerites in postabdomen of male usually re- posed a new tribe Rhodesiellini in the subfamily duced (apomorphic). Oscinellinae and divided all remaining genera of 4. Cerci reduced or small and fused into mesolobus the Old World into several informal genus groups: (apomorphic). 9 in the tribe Oscinellini and 12 in the subfamily 5. Distiphallus membranous and long (apomor- Chloropinae. Kanmiya (1983) changed the con- phic). tentsofsomeofAndersson’sgenusgroupsand erected 4 new genus groups. Nartshuk (1983, 1987) As we see it, there are two possible cladistics sce- proposed to divide the family into 21 tribes. narios (Figs 113, 114): Andersson (1977, 1979) and Nartshuk (1987) pre- + sented some ideas on the phylogeny of Chlorop- Chloropinae is sister group to Rhodesiellinae Os- cinellinae. idae, but stressed that much further work was + needed. Andersson (1977, 1979) remarked that the Oscinellinae is sister group to Rhodesiellinae family Chloropidae has evolved relatively recently Chloropinae.

51 Figs 113, 114. Alternative ideas about relationships between subfamilies of Chloropidae (Ch = Chloropinae, Os = Oscinellinae, Rh = Rhodesiellinae, Si = Siphonellopsinae). 113: more weight given to length of costal vein (char. 18). 114: more weight given to direction of ocellar setae (char. 16). Characters: 1 – propleural ridge present; 2 – anal vein and anal cell absent; 3 – flexure present on vein CuA1; 4 – tibial organ present; 5 – only one costal break; 6 – three pairs of strong dorsocentral setae; 6a – one pair of strong dorsocentral setae; 7 – katepisternum with strong seta; 7a – katepisternum without seta; 8 – orbital setae three strong pairs; 8a – orbital setae weak and numerous; 9 – male postabdomen asymmetrical (sternites 6–8 discrete); 9a – male postabdomen symmetrical (sternites 6–8 fused); 10 – subcostal vein reaches costal vein separate from vein R1; 10a – subcostal vein distally merged with vein R1; 11 – one postpronotal seta; 11a – two postpronotal setae; 12 – epandrium without processes; 12a – epandrium with anteroventral processes; 13 – inner vertical seta shorter than outer vertical seta; 13a – inner vertical seta longer than outer vertical seta; 15 – male surstyli simple; 15a – male surstyli divided into three lobes; 16 – ocellar setae procurved and divergent; 16a – ocellar setae upright and convergent; 17 – male cerci discrete or fused; 17a – male cerci reduced; 18 – vein C extended to vein M1; 18a – vein C ending at vein R4+5; 19 – larvae saprophagous; 19a – larvae to some extent phytophagous. (Open squares = plesiomorphic states; filled squares = apomorphic states.)

Key to subfamilies

1 Three rather long, outcurved or procurved scutellum than to apical setae orb. vti stronger than vte. if 1–3. Two rather ...... Siphonellopsinae(notintheNordic strong h, one recurved and one incurved. One countries) strong seta on katepisternum. Lateral setae on – orb numerous (5–10) and short. vte stronger scutellum situated closer to anterior margin of than vti. if more numerous or if few then up-

52 right or recurved. Usually only 1 recurved h or 3 oc procurved and divergent. Male cerci absent. without distinct h. No seta on katepisternum, Anepisternum covered with setulae. Scutel- bare or with some setulae. Usually several lat- lum usually more or less triangular and flat- eral setae on scutellum, if only one then sit- tened. Apical scutellar setae usually arising uated closer to apical setae than to anterior from warts or long projections marginofscutellum...... 2 . . . . . Rhodesiellinae (only genus Aspistyla in 2 Costa reaches R4+5 (except some Dicraeus in Nordic countries) which 2nd costal sector is 3–4 times longer – oc upright or recurved, parallel, convergent or than 3rd costal sector) (Figs 35–40). Male cruciate. Male cerci usually present. Anepis- cerci apically free (Figs 71, 73) ...... 3 ternum bare or with setulae. Scutellum usu- – Costa not reaching R4+5 (Figs 41–44). Male ally rounded and convex. Apical scutellar se- cerci completely fused, forming a subrectan- tae rarely arising from warts . . . . Oscinellinae gular sclerite (Figs 76, 80, 81) . . . Chloropinae

Key to Fennoscandian genera

1 Wings fully developed, usually longer – Arista thin, without strong pubescence thanabdomen...... 2 (Figs18–20)...... 7 – Wings reduced in size (Figs 2, 40), 5 (4) Two long orb on each side of frons (Figs shorter than abdomen, or practically 270, 273). Scutum punctuated and absent...... 52 scutellum with tubercles at base of api- 2 (1) Costal vein reaching vein M (Figs 35– cal and subapical setae (Figs 276, 282, 39), if ending between apices of R4+5 284, 287) and M then vein R2+3 long and 2nd ...... Elachiptera Macquart (p. 90) costal sector 3 times as long as 3rd costal –Morethantwoorb on each side of frons sector(Fig.37)...... 3 being short and equal in size ...... 6 – Costal vein extending only to R4+5 or 6 (5) First flagellomere rounded and black; slightly beyond, never reaching vein M arista subapical, black or white (Fig. 21) (Figs 41–44). (Subfamily Chloropinae) ...... Oscinella Becker p. p. (p. 124) ...... 32 – First flagellomere angular on upper cor- 3 (2) Ocellar setae rather long, directed an- ner, predominantly yellow; arista apical teriorly and divergent (Fig. 115). Body (Fig. 23). Male wing with black spots black, shining. Arista distinctly pubes- (Figs 3, 38) cent. (Subfamily Rhodesiellinae) ...... Gampsocera Schiner (p. 100) ...... Aspistyla Duda (p. 59) 7 (4) Large flies, body 4–8 mm long, antennae – Ocellar setae small, upright, convergent separated by a wide dusted keel (Figs 4, or parallel. Body black or partly yellow, 9)...... Lipara Meigen (p. 118) shining or matt by dusting. (Subfamily Oscinellinae) ...... 4 – Small flies, body 1–4 mm long, anten- nae usually not separated, keel absent or 4 (3) Arista thick, cylindrical or flattened, or verynarrow(Figs13–15)...... 8 appearing thick due to long and dense 8 (7) Scutum with 3 or 5 deeply impressed pubescence, coloured black or white punctuate lines (Figs21–23)...... 5 ...... Tricimba Lioy (p. 149)

53 – Scutum without punctuate lines, but tance between crossveins longer than punctuate depressions may be present dm-cu ...... 16 along dorsocentral line on posterior part 16 (15) Pleura dull by dusting, if shining below ofscutum...... 9 then all body black including genae and 9(8) VeinR2+3 long, 2nd costal sector 3–4 face...... 17 times longer than 3rd costal sector (Fig. – Pleura shining, without dust except on 37)...... Dicraeus Loew (p. 85) upper part of anepisternum; if pleura dusted then proboscis long and genicu- –VeinR2+3 notsolong,2ndcostalsector no more than 2 times longer than 3rd late...... 18 costal sector (Figs 35, 36, 39) ...... 10 17 (16) Genae wider than first flagellomere (Fig. 215) 10 (9) Vein R2+3 very short, 2nd costal sector Colliniella Nartshuk & Andersson (p. 77) distinctly shorter than 3rd costal sector – Genae at most as wide as first flagellom- (Fig. 39) ere (Figs 121, 126, 134) ...... Siphunculina Rondani (p. 146) ...... Aphanotrigonum Duda (p. 61) –VeinR+ long, 2nd costal sector longer 2 3 18 (16) Vibrissal angle distinctly projecting be- than 3rd costal sector (Figs 35, 36) . . . 11 yond eye margin. Proboscis may be long 11 (10) Thorax dorsoventrally flattened . . . . . 12 and geniculate (Figs 460, 463, 469) – Thoraxnotflattened...... 14 ...... Oscinimorpha Lioy (p. 133) 12 (11) Scutum and scutellum with coarse mi- – Vibrissal angle not projecting beyond crosculpture. Apical setae of scutellum eye margin and proboscis short (Figs on small tubercles (Figs 473, 479) 347,360,424,530)...... 19 ...... Oscinisoma Lioy (p. 135) 19 (18) Frons except ocellar triangle with small – Scutum and scutellum smooth, without shining punctures. Eyes densely haired. microsculpture, at most slightly sculp- Scutellum with many marginal setae turedbeforescutellum...... 13 (Fig. 532) 13 (12) Scutum densely dusted and dull ...... Speccafrons Sabrosky (p. 148) ...... Eribolus Becker (p. 96) – Frons evenly haired, without punctures. – Scutum without or with very thin dust, Eyes sparsely short haired. Scutellum shining, sometimes with a green metal- with 1 apical and 2–3 subapical pairs of lic lustre setae...... 20 .....Rhopalopterum Duda p. p. (p. 141) 20 (19) Scutum with setulae dispersed ...... 21 14 (11) Ocellar triangle densely dusted or sub- – Scutum with setulae in rows shining through very thin dust ...... 15 ...... Incertella Sabrosky (p. 109) – Ocellar triangle without any dusting, 21 (20) Scutum densely grey-dusted, convex, if shining, sometimes with punctures and shining then setae and setulae yellow. setulaeonitssurface...... 22 Distance between crossveins shorter 15 (14) Body yellow with black or reddish stripes than basal sector of vein M1 (Fig. 34), if stripes fused and scutum ...... Conioscinella Duda p. p. (p. 78) black then postpronota and pleura part- – Scutum black, very thinly dusted, flat- ly yellow. Crossvein dm-cu distinctly tened. Distance between crossveins no oblique; distance between crossveins shorter than basal sector of vein M1 equals length of dm-cu (Fig. 543) (Fig. 514) ....Trachysiphonella Enderlein (p. 148) .....Rhopalopterum Duda p. p. (p. 141) – Body black or densely grey-dusted. 22 (14) Genae distinctly produced and anten- Crossvein dm-cu more upright and dis- nae separated by keel. Proboscis very

54 long and geniculate (Figs 13, 27) 28 (27) Scutellum rounded with remote apical ...... Siphonella Macquart (p. 145) setae (Figs 323, 325) – Genae barely produced. Proboscis ...... Gaurax Loew (p. 101) shorter. Female in doubtful cases with – Scutellum conical with apical setae ap- compressed, sclerotized cerci (Fig. 184) proximated (Fig. 493) ...... 23 ...... Pseudogaurax Malloch (p. 139) 23 (22) Postocellar setae (poc) parallel, frons 29 (27) Ocellar triangle reaching front margin of distinctly punctuate (Figs 371, 380, 484) frons (Fig. 431). Setulae on scutum few, ...... 24 arranged in single rows along acrostichal – poc convergent or cruciate, frons not and dorsocentral lines. Male cerci nearly punctuate, dull or covered with silky round or rather large (Figs 426, 429, 434, hairs...... 25 441, 445) 24 (23) Antennae fully separated by thin keel ...... Oscinella Becker p. p. (p. 124) (Figs 14, 486), wing cell br sometimes – Ocellar triangle shorter than frons (Figs widened (Fig. 487) 223, 346, 362). Setulae on scutum more ...... Polyodaspis Duda (p. 138) numerous and evenly dispersed, or at – Antennal keel only developed in upper least arranged in multiple rows . . . . . 30 part of face (Figs 15, 382) 30 (29) Body elongated, scutum flattened. Se- ...... Lasiambia Anonymous (p. 114) tulae on scutum dark and evenly dis- 25 (23) Ocellar triangle with setulae on its sur- persed. Body sometimes with light me- face (Figs 149, 159, 170, 177, 198, 212, 342) tallic luster ...... 26 .....Rhopalopterum Duda p. p. (p. 141) – Ocellar triangle without setulae on its – Body short, scutum convex. Setulae on surface, only on frons outside its side scutum either yellow or arranged in dou- margins...... 27 blerows...... 31 26 (25) Ocellar triangle with setulae arranged in 31 (30) Setulae on scutum yellow, abundant and a full length row along its side margins dispersed (Fig. 342) ...... Conioscinella Duda p. p. (p. 78) ...... Hapleginella Duda (p. 108) (one – Setulae on scutum black, arranged in species H. laevifrons) double rows – Ocellar triangle with setulae in 2–3 ir- ...... Microcercis Beschovski (p. 122) regular rows, arising from distinct small 32 (2) R + closely following and fused with R punctures (Figs 149, 159, 170, 177, 198, 2 3 1 and C (Figs 42, 566). Apical scutellar se- 212) . . . . Calamoncosis Enderlein (p. 68) tae placed on disc of scutellum (Fig. 565) 27 (25) Body yellow with black stripes on scu- ...... Camarota Meigen (p. 154) tum, or black, sometimes with yellow –R+ well separated from R and C (Fig. marks. Scutum shining, with dispersed, 2 3 1 41). Apical scutellar setae, if developed, rather long setulae. Eyes and arista with placed on margin of scutellum ...... 33 long pubescence. If body entirely black, then it is shining and arista is white. 33 (32) One to three orbital bristles distinctly longer than remaining ones (Figs 732, Wing cell R1 rather short and broad, 764, 918). Body always yellow with black broader than cell R2+3 (Fig. 324) . . . . 28 stripes that may become fused on scu- – Body uniformly black, setulae on scu- tum...... 34 tum short, sometimes arranged in rows. – Orbital bristles all about same size (Figs Eyes and dark arista with short pubes- 569, 598, 602). Body yellow with black cence...... 29 scutalstripesorwhollyblack...... 35

55 34 (33) Crossveins of wing approximated, their face of scutum smooth, evenly dusted separation not greater than length of ...... Diplotoxoides Andersson (p. 190) dm-cu (Fig. 44). Small species 41 (39) Ocellar triangle with one or more rows ....Pseudopachychaeta Strobl (p. 223) of setulae within side margins (Figs 705, – Crossveins more separated from each 930,939,944,948)...... 42 other than length of dm-cu (Fig. 766) – No setulae within side margins of ocel- Larger species lar triangle. Setulae present on frons out- ...... Lasiosina Becker (p. 196) side ocellar triangle (Figs 569, 752) 35 (33) Hind legs with femora greatly enlarged ...... 44 and tibiae usually curved (Figs 5, 904) 42 (41) Scutum entirely yellow with black or red ...... 36 longitudinal stripes (Fig. 7b). Scutellum – Hind legs without enlarged femora flattened, with apical setulae approxi- ...... 37 mated 36 (35) Veins R1 and R2+3 straight (Fig. 902)...... Thaumatomyia Zenker (p. 226) First flagellomere much longer than – Scutumentirelyblack...... 43 deep (Figs 900, 907). Frons strongly pro- 43 (42) First flagellomere rounded apically duced. Body large, reddish yellow (Fig...... Neohaplegis Beschovski (p. 218) 7a)...... Platycephala Fallén (p. 219) – First flagellomere acuminate apically –VeinsR1 and R2+3 curved towards C. (Figs 706, 707) First flagellomere barely longer than ...... Cryptonevra Lioy (p. 186) deep. Frons weakly produced. Body 44 (41) Arista shorter than remaining antenna, smaller, greenish nearly bare (Fig. 754). Body setulae white ...... Meromyza Meigen (p. 200) ...... Eutropha Loew (p. 194) 37 (36) Head triangular in side view (Figs 745, – Aristaaslongasorlongerthanremain- 953)...... 38 ing antenna. Body setulae usually black – Head rounded or rectangular in side ...... 45 view (Figs 570, 591, 599, 728) ...... 39 45 (44) First flagellomere longer than deep (Figs 38 (37) Body grey-dusted, densely covered with 20, 24, 570, 893, 894). Arista white long setae (Figs 952, 953). Anepister- ...... 46 num and anepimeron naked – First flagellomere only as long as deep...... Trichieurina Meigen (p. 230) Aristablackorwhite...... 47 – Body brownish-yellow with only short 46 (45) Scutum black, rugose setae. Anepisternum and anepimeron ...... Centorisoma Becker (p. 155) setulose ...... Eurina Meigen (p. 193) – Scutum yellow, smooth, with five dull 39 (37) Crossveins r-m and dm-cu approxi- black stripes, lateral stripes with a vel- mated, their separation not greater than vety black spot anteriorly length of dm-cu (Figs 43, 44, 729) . . . 40 ...... Parectecephala Becker (p. 218) – Crossveins r-m and dm-cu more sepa- 47 (45) Scutum yellow with black or red longitu- rated, their distance from each other dinalstripes...... 48 greater than length of dm-cu ...... 41 – Scutum black, at most postpronota and 40 (39) A row of setulae on ocellar triangle in- notopleurapartlyyellow...... 49 side its lateral margin (Fig. 728). Surface 48 (47) Genae linear (Figs 697, 700). Male gen- of scutum rugose and shining italia: pre- and postgonites in line (Figs ...... Diplotoxa Loew (p. 190) 699, 702) . . . Chloropsina Becker (p. 185) – A row of setulae on frons outside lateral – Genae wider, at least half as wide as margin of ocellar triangle (Fig. 732). Sur- first flagellomere (Figs 591, 595, 599,

56 603, 608). Male genitalia: postgonites 52 (1) oc setae procurved, divergent. Body fus- situated outside of pregonites (Fig. 77) cous yellow with black scutal stripes and ...... Chlorops Meigen p. p. (p. 159) black marks on pleura 49(47) Surfaceofscutumsmooth...... 50 ...... Lasiosina Becker p. p. (p. 196) – Surfaceofscutumrugose...... 51 – oc setae recurved or upright, convergent 50 (49) Vibrissal angle acute (Figs 17, 779) (Fig. 270). Body black with grey dust or ...... Melanum Becker (p. 200) thoraxredandabdomenblack.....53 – Vibrissal angle obtuse (Figs 616, 619, 623, 53 (52) Body black, grey-dusted. Scutellum with- 639).....Chlorops Meigen p. p. (p. 159) out warts at base of marginal setae (Figs 238, 239) 51 (49) Ocellar triangle reaching anterior mar- ...... Conioscinella Duda p. p. (p. 78) gin of frons (Fig. 737). Middle tibiae with small apical spur – Thorax red, abdomen black with tergites ...... Epichlorops Zetterstedt (p. 192) 1–2 extended. Scutellum with warts at base of marginal setae (as Fig. 276) – Ocellar triangle not reaching anterior ...... Elachiptera Lioy p. p. (p. 90) margin of frons. Middle tibiae with strong apical spur ...... Cetema Hendel (p. 155)

Preliminary key to genera for 3rd instar larvae

Larval descriptions exist for only a small number – Hind spiracles not trilobate, sessile or of genera and species. The present key to genera, born on posterior processes (Figs 454, intended for mature (3rd instar) larvae is there- 909,915,970,973)...... 5 fore very preliminarily. It can be used for puparia 3 (2) Facial mask with numerous nearly par- as well, as many larval characters remain intact on allel ridges. Anterior spiracles of rosette puparia: form of posterior end, spiracular zone of type with 10–12 buds (Fig. 958). abdominal segments, cephalopharyngeal skeleton Branched hairs on hind spiracles ab- and anterior spiracles. sent. All three thoracic segments with- out spicules, abdominal segments with 1 Larvae large, up to 8–12 mm, rather thick many spicules on creeping welts (Fig. and white. First or three first body seg- 959). Larvae in shoots of grasses ments and last segment dorsally sclero- ...... Camarota Meigen (p. 154) tized and blackish (Figs 398, 403). Pos- terior spiracles sessile. Larvae formings – Facial mask with few ridges or ridges galls on Phragmites australis (Figs 106, practicallyabsent(Fig.961)...... 4 107)...... Lipara Meigen (p. 118) 4 (3) Facial mask smooth. Cephalic segment – Larvae smaller and narrower, of white, large. Distal segment of antennae larger yellowish or greenish colour, without than basal one and round (Fig. 961). dorsal sclerotisations (Figs 92, 185). Pos- Three thoracic segments narrow and terior spiracles sessile or born on pos- long (Fig. 960). Larvae among roots of terior processes (Figs 454, 909, 815, 915, plants 963,970,973)...... 2 ...... Thaumatomyia Zenker (p. 226) 2 (1) Hind spiracles trilobate, sometimes – Facial mask simple, with maximally four more or less sclerotized (Figs 956, 957, crescent ridges. Larvae in shoots of 962)...... 3 grasses...... Conioscinella p. p. (p. 78)

57 5 (2) Hypopharyngial and tentoropharyngeal abdominal segments large and brown sclerites articulated. Mandibles more or (Figs 93, 964, 965). Larvae in different less slender, without or with accessory media...... Polyodaspis Duda (p. 138) teeth on ventral side (Figs 96, 456, 964, – Some spicules present above and/or be- 968, 971). Facial mask usually with com- lowanus...... 10 plex pattern of ridges. Hind spiracles 10 (9) Larvae in shoots of grasses and other born on posterior processes (Figs 717, plants, usually monocots 725,973,979)...... 6 ...... Aphanotrigonum Duda (p. 61), – Hypopharyngial and tentoropharyngeal Elachiptera Macquart (p. 90), Incertella sclerites fused (Figs 97, 911, 916, 997, Sabrosky (p. 109) 1003, 1010). Mandibles very massive, al- – Larvae in rotten wood, bracket fungi, ways with accessory teeth on ventral coniferous cones or in egg sacks of spi- side. Facial mask usually with more sim- ders...... 11 ple pattern. Hind spiracles sessile or 11 (10) Spicules on ventral side of abdominal born on posterior processes ...... 14 segments large, acute and wide basally, 6 (5) Mandible slender, without accessory arranged in two rows on ventral side of teeth on ventral side (Fig. 964). Facial 2nd to 8th abdominal segments (Fig. mask consists of several rows of parallel, 966). Larvae found in egg sacks of spi- oftendentateridges(Fig.93)...... 7 ders . . . . Pseudogaurax Malloch (p. 139) – Mandible rather massive and with ac- – Spicules on ventral side of abdominal cessory, sometimes very small teeth on segments smaller and apically obtuse, ventral surface. Facial mask contains arranged in several rows on ventral side ridges uniting into cells (Figs 94, 449, of 2nd to 8th abdominal segments (Figs 455)...... 12 969, 972). Larvae found in rotten wood, 7 (6) Anus surrounded by coarse spicules (Fig. bracket fungi, and coniferous cones 725)...... 8 ....Gaurax Loew (p. 101), Hapleginella Duda (p. 108) – Anal area with fewer spicules above and/or below anus, or without spicules 12 (6) First three abdominal spicular zones ...... 9 consist ventrally of confluent rows of 8 (7) Dental sclerite arched (as Fig. 97). 2nd finely serrate striae; on the following and 3rd thoracic segments without spic- abdominal segments these transverse ules. Creeping welts covered with coarse fields of striae are accompanied in front and thin spicules, but anterior row inter- by rows of small spicules (Fig. 976). Ab- rupted in the middle. Larvae in shoots of domen deeply divided posteriorly into grasses...... Lasiosina Becker (p. 196) two large spiracular lobes (Fig. 977) ...... Cetema Hendel (p. 155) – Dental sclerite small and of different form, or absent (Figs 712, 719). 2nd and – All abdominal spicular zones consist 3rd thoracic segments without spicules. ventrally of several transverse rows of Creeping welts covered with coarse and spicules of same or different size . . . . 13 thin spicules, anterior row usually entire 13 (12) Facial mask: frontal ridges between max- (Figs715,716,721,722).Larvaeinshoots illary papillae absent; genal ridges for- of Pragmites australis,ofteninLipara ming only 3–4 plain cells (Figs 449, 455). galls...... Cryptonevra Lioy (p. 186) Spicular fields or individual rows on 9 (7) Anal area and thoracic segments with- 2ndand3rdthoracicsegmentsusually out spicules. Spicules on ventral side of present, if absent then creeping welts on abdominal segments all armed with

58 coarse spicules (Figs 450–453) – Spicular zone of abdominal segments at ...... Oscinella Becker (p. 124) leasttosomeextentconsistingofwell – Facial mask more complex: about 4 rows defined spicules (Figs 909, 915) . . . . . 18 of conspicuous, transverse densely den- 17 (16) Antennae widely separated, situated an- tate frontal ridges between maxillary terolaterally on the head lateral to max- papillae; genal ridges forming more than illary papillae. Maxillary papillae com- 4 finely dentate cells (Fig. 978). Spicular pletely surrounded by a thick dark zones on 2nd and 3rd thoracic segments brown ring (Figs 989, 992, 996, 999, absent (Fig. 980) 1002). Dentate sclerite absent or very ...... Conioscinella Duda (p. 78) small. Caudal end of body variable. 14 (5) Spicular zones absent ventrally and dor- Larvae white or yellowish, in shoots of sally on abdominal segments, or only grasses...... Chlorops Meigen (p. 159) with a single row of spicules ...... 15 – Antennae situated in front of maxil- – Spicular zones present on both thoracic lary papillae. Maxillary papillae not sur- andabdominalsegments...... 16 rounded by thickened ring (Fig. 1008). 15 (14) Larvae slightly depressed dorsoventrally, Dentate sclerite arched (Figs 1010, 1011, up to 4 mm long, pale yellow, in seeds of 1014). Caudal end of body dorsally with a furrow (Fig. 1012). Larvae pale green grasses...... Dicraeus Loew (p. 85) ...... Meromyza Meigen (p. 200) – Larvae not depressed, white, in stems 18 (16) Posterior spiracles born on processes of Phragmites australis, both intact and from caudal end of body (Fig. 981). Lar- stems with Lipara galls or infected by vae yellowish, in inflorescences of Cyp- Platychephala planifrons; also found in eraceae other grasses ....Pseudopachychaeta Strobl (p. 223) ...... Calamoncosis Enderlein (p. 68) – Posterior spiracles sessile (Figs 909, 915). 16 (14) Spicular zone of abdominal segments Larvae white, stocky, living in shoots of consists ventrally of numerous homoge- Phragmites australis nous, often slightly confluent ridges ...... Platycephala Fallén (p. 219) without well defined spicules present (Figs987,1009,1013)...... 17

Subfamily RHODESIELLINAE

Genus Aspistyla Duda, 1933 lomere reniform, wider than long. Scutum shining, scutellum with apical and lateral setae on small tubercles. Tibial organ indistinct. Femoral organ Macrostyla Lioy, 1864: 1126 (preocc. Winnertz, formed by a group of small warts with short thin 1846). Type species: Chlorops plumiger Meigen, setae, presents in both sexes and in male on hind 1830 by monotypy. femora as well. Male genitalia: epandrium with Aspistyla Duda,1933:224,asreplacementname for Macrostyla Lioy, 1864. Type species: Chlorops terminal surstyli; pregonites with strong setae; dis- plumiger Meigen, 1830, automatic. tiphallus long and slender, curved. Black species with large naked eyes, haired arista and linear genae. Ocellar triangle large shining, Distribution. – Worldwide, most diverse in the Old reaching front margin of frons. Setae of head long, World tropics with many species in the Oriental ocellar setae procurved and divergent. First flagel- and Afrotropical Regions.

59 Figs 115–119. Aspistyla plumiger (Meigen): 115: head, dorsal view. 116: head, lateral view. 117: scutellum. 118: hypopygium, dorsal view. 119: hypandrium and phallic complex.

1. Aspistyla plumiger setae long, standing on small tubercles, apical se- (Meigen, 1830) tae longer than scutellum. Legs with femora black, tibiae and tarsi yellow. Distal segment of tarsi dark- Figs 46, 47, 115–119 ened. Halteres yellow. Body length nearly 2 mm.

Chloropsplumiger Meigen, 1830: 153; Meigen’s plate Distribution. – Rare. Denmark: EJ: Svejbæk, 229: 12 (Morge, 1976b). Lectotype in NHMW 19.vii.1964 (V. Koch), 1 female. Sweden: SK, ÖL. Fin- (Nartshuk, 1997a). land: Al. Not found in Norway or Russian parts of Fennoscandia. Body black, shining. Ocellar triangle large shin- Trans-Palearctic species, known from the British ing, reaching front margin of frons, with long se- Islands to the Far East of Russia. tulae along sides. Setae of head long, ocellar setae procurved and divergent. Eyes large, naked. Genae Biology. – Adults from end of June to September, in linear. First flagellomere wider than long with black broad-leaf woods and bushes. It has been reared long-haired arista. Palpi brown. Scutum and pleura from a mushroom (Xerocomus chrysenteron)in shining, scutellum triangular, apical and subapical Tatarstan, Russia (Khalidov, 1984; Yakovlev, 1994).

60 Subfamily OSCINELLINAE

Genus Aphanotrigonum Duda, 1932 num shining. Male genitalia with strongly ta- pering and straight surstyli (Figs 139–141) Aphanotrigonum Duda, 1932: 35. Type species: ...... A. norrbotticum sp. n. Chlorops trilineatus Meigen, 1830 by original – Head and legs partly yellow. Scutellum trape- designation. zoid with apical setae wider apart (Fig. 127). One posterior npl seta. Pleura mainly dusted. Small black, densely dusted species with mainly Male genitalia with surstyli of different shape dusted pleura. Ocellar triangle small and dusted. Genae with produced vibrissal angle. poc short, up- and directed mesad (Figs 123–125, 128, 129, right and convergent. Face with distinct but short 132,133)...... 6 carina. First flagellomere small and thick basally. 6 Hind tibiae with a small but distinct black npl 1 + 2. Tibial organ oval. Femoral organ consists spur. Prothoracic episternum shining ...... 7 of two rows of warts with short thin setulae. Male – Hind tibiae without a spur. Prothoracic epis- genitalia: hypandrium closed. ternum dull. Male genitalia Figs 132, 133 Distribution. – Holarctic, New Zealand, Oriental ...... A. inerme Collin Region. 7 All coxae and femora dark. Male genitalia Figs 128–130...... A. femorellum Collin Key to species of Aphanotrigonum – At least anterior and middle femora partly yel- low. Male genitalia Figs 123–125 Duda ...... A. cinctellum (Zetterstedt) 1 Wings shorter than abdomen. Male genitalia Fig.120...... A. brachypterum (Zetterstedt) 2. Aphanotrigonum brachypterum – Wingslongerthanabdomen...... 2 (Zetterstedt, 1848) 2 Scutellum with many marginal setae, apical Fig. 120 setae short and not much wider apart than poc Oscinis brachyptera Zetterstedt, 1848: 2660. Holo- setae(Figs135,143)...... 3 type in ZMLU (Andersson, 1966; Dely- – Scutellum with no more than two pairs of Draskovits, 1981). marginal setae, apical setae long and wider Aphanotrigonum trilineatum var. microptera Duda, apart than poc setae(Fig.122)...... 5 1932: 36. Lectotype male by present designa- 3 Three or five brown stripes on scutum and tion, and paralectotypes 4 males and 6 females scutellum. Male genitalia Figs 146–148 all from Denmark: NEZ: Tibirke, 19.x.1914 (W. Lundbeck), in ZMUC. Duda (1932) misspelled ...... A. trilineatum (Meigen) Tibirke as “Fibicke”. Syn. n. – Scutum evenly grey without brown stripes ...... 4 Body black, densely grey-dusted. Wings shortened, 4 Male genitalia: epandrium large; cerci not though extend nearly to the end of abdomen. Ocel- lar triangle dull, reaching halfway to front margin tapering, with several longish setae; surstyli of frons. Frons brownish yellow in front. Scutellum broad (Fig. 131) with 8–10 small marginal setae, apical setae ap- ...... A. hungaricum Dely-Draskovits proximated. Pleura dusted. Femora mainly black. – Male genitalia: epandrium small; cerci slightly Body length 1.7–1.8 mm. tapering, with one long seta; surstyli narrow Distribution. – Rare species. Denmark: NWJ: (Figs 136–138) ...... A. nigripes (Zetterstedt) Hansted Reservatet, 08.vi.1960 (ZMUC Exp.), 1 fe- 5 All body black except first flagellomere below. male; NEZ: Without locality (R.W. Schlick), 1 male; Scutellum rounded, apical setae rather close Tibirke, 19.x.1914 (Krüger), 5 males, 6 females; B: together. Two posterior npl setae. Anepister- Dueodde, 18.vi.1964 (L. Lyneborg, O. Martin & B.

61 Figs 120–125. Aphanotrigonum brachypterum (Zetterstedt): 120: hypopygium, dorsal view. A. cinctellum (Zetterstedt): 121: head, lateral view. 122: scutellum. 123: hypopygium, dorsal view. 124: hypopygium, lateral view. 125: male genitalia, ventral view.

Petersen), 1 female. Sweden: Sk: Björnstorp, 1 male Note. – Examination of a long series of specimens (holotype) (examined by Dely-Draskovits, 1981); from different localities revealed a sexual varia- Kullaberg, 16.viii.1966 (H. Andersson), 1 female. tion in colour of legs: femora dark in males, yellow European species also known from Germany and in females. Therefore we consider Oscinis cinctella Hungary. Zetterstedt and O. fasciella Zetterstedt as male and Note. – Andersson (1966) and Ismay (1980b) con- female of the same species. sidered A. brachypterum as a short-winged form of A. nigripes (Zetterstedt), but Dely-Draskovits Body mainly black, lightly grey-dusted. Frons yel- (1981), after examination of the type specimens, low anteriorly; ocellar triangle black with grey treated it as a separate species based on differences dust, reaching halfway to front margin of frons, be- in the male genitalia. hind reaching eye margin. Face and genae yellow, vibrissal angle produced. Palpi yellow. Antennae 3. Aphanotrigonum cinctellum yellow, first flagellomere darkened. Scutum grey, (Zetterstedt, 1848) scutellum short, rectangular. Apical setae longer Figs 121–125 than scutellum, wide apart. Pleura dusted. npl 1 + 1. Oscinis cinctella Zetterstedt, 1848: 2659. Holotype Base of abdomen yellow, remaining tergites with male in ZMLU (Andersson, 1966). black bands. Halteres yellow. Legs in female exten- Oscinis fasciella Zetterstedt, 1855: 4808. Holotype sively yellow, in male femora dark; hind tibiae with female in ZMLU (Andersson, 1966). Syn. n. small dark spur. Body length 1.0–1.3 mm.

62 Distribution. – Denmark: SJ, NEJ, F, NEZ. Sweden: puncture on sides of ocellar triangle. orb numer- SK, SM, Öl, GO, BO, SÖ. Norway: HEn: Isterfossen. ous and very short. Genae as wide as first flagel- Finland: Al, Ab, Om, Ob. Russia: Kr: Petrozavodsk. lomere, yellow in front, Vibrissal angle produced. Widely distributed in Palearctic, eastwards to First flagellomere dark brown, yellowish at base. Yakutia, East Siberia, southwards to Israel. Scutum and pleura dusted bluish grey. Scutum and scutellum punctate. Scutellum with approximated Biology. – Adults in July to early September in apical setae and 4–5 subapical setae on each side. dry habitats, mainly sandy beaches. Tschirnhaus npl 1 + 2; postalar seta rather long. Abdomen broad (1981) recorded as host plant Puccinella maritima and flat with lateral parts of tergites on ventral and Bährmann & Weipert (1989) reared the species side of abdomen, greyish brown. Male abdominal from Elytrigia and Puccinella. sternites 3 and 4 wider than long. Halteres yellow. Legs mainly black. Male genitalia: epandrium large, 4. Aphanotrigonum femorellum cerci broad, surstyli with thick setulae at base. Body Collin, 1946 length 2.0 mm. Figs 126–130 Distribution. – Denmark: WJ, F. Sweden: SK. Fin- land: Al: Sund (R. Frey); Ab: Nyström; N: Tvär- Aphanotrigonum femorella Collin, 1946: 127. Lec- minne. Not found in Norway or Russian parts of totype male (Dely-Draskovits, 1981), paralecto- Fennoscandia. types 3 males, 9 females in OXUM (Pont, 1995). Recorded from Europe and Afghanistan. Similar to A. cinctellum in having npl setae 1 + 1, apical scutellar setae wide apart, hind tibiae with 6. Aphanotrigonum inerme small dark spur. Main differences: first flagellom- ere larger and more extensively brownish on outer Collin, 1946 side, abdomen darker, pale hind margins on ter- Figs 132, 133 gites 3–5 narrower. Legs much darker, all femora dark, tibiae with dark band, tarsi brownish. Body Aphanotrigonum inerme Collin, 1946: 128. Lecto- length 1.5–1.75 mm. type male (Dely-Draskovits, 1981) and paralec- totypes 12 males and 9 females in OXUM (Pont, Distribution. – Rare species. Denmark: WJ: Skallin- 1995). gen, 17.vii.1982 (E. Bro Larsen); F: Langeland, Kelds- nor, 6.viii.1975 (Lyneborg, Martin & Michelsen). Similar to A. femorella, but without an anteroven- Sweden: SK: Foteviken, 25.viii.1960, coastal mead- tral spur on hind tibiae, prothoracic episternum dusted, abdomen without pale bands on hind mar- ow, 28 specimens (H. Andersson). Not found in + Norway, Finland and adjacent territories of Russia. gins of tergites except on female tergite 5. npl 1 1. Widely distributed but rare Palearctic species, Apical scutellar setae wide apart. Body length 1.5– known from Europe to Mongolia, and North Africa. 1.8 mm. Biology. – Adults in July and August in dry mead- Distribution. – Rare species. Denmark: WJ: Fanø, ows and on sandy beaches. Tschirnhaus (1981) re- 30.v.1919, 10.vii.1921 (T. Mortensen), 2 males, 2 fe- corded as host plants Puccinella maritima and Fes- males;Ringkøbing,24.vii.1951(K.O.Leth),1female; tuca rubra. NEZ: no further data (R.C. Stæger), 1 female. Swe- den: SK: Revinge, 26.vi.1985 (H. Andersson), 1 fe- male. Not found in Norway, Finland or Russian 5. Aphanotrinonum hungaricum parts of Fennoscandia. Dely-Draskovits, 1981 European species. Fig. 131 Biology. – Adults in May to July. In Bulgaria the species occurs on sand dunes with Phragmites aus- Aphanotrinonum hungaricum Dely-Draskovits, tralis (Beschovski, 1982). 1981: 131. Holotype in HNHM. Similar to A. nigripes. Body black, densely grey- dusted. Frons black, fuscous yellow in front. Ocellar 7. Aphanotrigonum nigripes triangle dull grey, reaching halfway to front mar- (Zetterstedt, 1848) gin of frons. A few tiny setulae arising from minute Figs 29, 134–138

63 Figs 126–133. Aphanotrigonum femorellum Collin: 126: head, lateral view. 127: scutellum. 128: hypopygium, dorsal view. 129: hypopy- gium, lateral view. 130: male genitalia, ventral view. A. hungaricum Dely-Draskovits: 131: hypopygium, dorsal view. A. inerme Collin: 132: hypopygium, dorsal view. 133: surstylus, lateral view. (131–133 after Dely-Draskovits, 1981.)

64 Oscinisnigripes Zetterstedt, 1848: 2652. Holotype in dusted. It is characterized by the wholly black ZMLU (Andersson, 1966). frons; antennae separated by very thin facial carina; pleura partly shining (contrary to all other species Body black, densely grey-dusted. Frons black, fus- of the genus); male genitalia with long surstyli. The cous yellow in front. Ocellar triangle dull grey, dark body colour, partly shining pleura and struc- reaching halfway to front margin of frons. A few ture of male genitalia provide the best diagnostic tiny setulae arising from minute punctures on characters of the new species. sides of ocellar triangle. orb numerous and very short. Genae as wide as first flagellomere, yellow in Description. – Bodyblack,darklydusted,exceptfor front, vibrissal angle produced. First flagellomere shining parts of pleura. Only face and ventral part dark brown, yellowish at base. Scutum and pleura of first flagellomere fuscous yellowish. Frons a little dusted bluish grey. Scutum and scutellum punc- longer than wide, dull. Ocellar triangle very small, tate. Scutellum with approximated apical setae and nearly reaching halfway to front margin of frons. 4–5 subapical setae on each side. npl 1 + 2and Setae of head very short. Genae distinctly produced a rather long postalar seta. Abdomen broad and beyond eye, wider than first flagellomere, brown- flat with lateral parts of tergites on ventral side ish with black lower margin. First flagellomere very of abdomen, greyish brown. Halteres yellow. Legs small, rounded, reddish below. Antennae separated mainly black. Body length 1.5–2.0 mm. by very thin facial carina. Proboscis rather long. Distribution. – Denmark: EJ, NWJ, NEZ, B. Sweden: Palpi darkened. Scutum nearly square with dis- north to NB and LU. Norway: Ø, AK, HEn, Bø, AAy, tinct puncture line in the middle and a pair of Ry. Finland: north to Ob and Ks. Russia: Vib, Kr. small depressions before scutellum. Scutellum flat, Eurasian species, known from Europe to Yakutia, rounded, apical setae not longer than scutellum, East Siberia and Mongolia. two small subapical setae on each side. Distance between apical setae not longer than 1.5 times Biology. – Adults from May to October, hibernate. distance between poc. Pleura partly shining, only Common in wet meadows, bogs and marshes. dusted on anepimeron and upper part of katepis- Larvae live in shoots of different grasses includ- ternum. npl 1 + 2. Halteres yellow. Abdomen and ing Calamagrostis epigeios, Leymus arenarius, legs entirely black. No spur on hind tibiae. Male Avena, Puccinella maritima (Uffen & Chandler, genitalia: cerci small and wide apart; surstyli rather 1978; Tschirnhaus, 1981). Collected on agricultural long, strongly tapering and nearly parallel; hypan- grasses in Denmark (Nielsen & Nielsen, 1984). drium closed, rather massive; 2 trichoid sensillae Bread from stems of wheat and oats in Bulgaria distinct in front of hypandrium. Body length 1.2– (Krusteva & Beschovski, 2000, 2001). Kibric et al. 1.25 mm. (1999) listed the species as tyrphophilous. Adults have been found in nest of Microtus sp. (Collin, Etymology. – The species is named after the type 1939). locality.

8. Aphanotrigonum norrbotticum 9. Aphanotrigonum trilineatum Nartshuk & Andersson, sp. nov. (Meigen, 1830) Figs 139–141 Figs 10, 11, 48, 142–148 Holotype, male: Sweden: Norrbotten: Högsön, Chlorops trilineatus Meigen, 1830: 162; Meigen’s Råneå, 01.vii.1972 (H. Andersson). Paratypes: 20 plate 210: 11 (Morge, 1976b). Lectotype in males, 31 females, same locality, 01, 06 and NHMW (Nartshuk, 1997a). 07.vii.1972 (H. Andersson); 2 males (in alcohol), Aphanotrigonum beschovskii Dely-Draskovits, 1978: Lule Lapmark: Messaure, 08–15 and 22–29.vii.1974 128. (K. Müller). Holotype and 49 paratypes in ZMLU, 4 Body black, densely bluish grey-dusted. Frons dark paratypes in ZISP. brown behind and yellowish in front, covered with Diffferential diagnosis. – In general appearance short pale setulae on minute black tubercles. Ocel- most similar to species of the Aphanotrigonum tri- lar triangle poorly demarcated at sides, nearly lineatum species group: genae with produced vib- reaching middle of frons, black, grey-dusted with rissal angle, ocellar triangle small, dusted, frons two rows of black setulae at sides. Genae slightly

65 Figs 134–141. Aphanotrigonum nigripes (Zetterstedt): 134: head, lateral view. 135: scutellum. 136: hypopygium, dorsal view. 137: hypopy- gium, lateral view. 138: male genitalia, ventral view. A. norrbotticum Nartshuk & Andersson: 139: hypopygium, dorsal view. 140: surstylus, lateral view. 141: hypandrium and phallic complex.

narrower than first flagellomere, vibrissal angle femora all black, tibiae black with both ends yel- produced. Antennae yellow, first flagellomere black lowish. Body length 1.5–2.0 mm. above. Head setae black, short and stout. Scutum densely covered with bluish grey dust, with 3–5 Distribution. – Very common. Denmark: EJ, WJ, brown longitudinal stripes along acrostichal and NEZ. Sweden: north to LU and NB. Norway: Ø, AK, dorsocentral lines and sometimes on sides of scu- HEn, HEs, Bø, VE, AAy. Finland: north to Li. Russia: tum and scutellum. Scutellum slightly wider than Kr, Lr. long with short stout apical setae and a pair of sub- Probably Holarctic, as Beaulier & Wheeler (2001) apical setae. Pleura dusted. npl 1(2) + 2. Legs with recorded the species with some reservation from

66 Figs 142–148. Aphanotrigonum trilineatum (Meigen): 142: head, dorsal view. 143: scutellum. 144: wing. 145: male abdomen, ventral and dorsal view. 146: hypopygium, dorsal view. 147: hypopygium, lateral view. 148: male genitalia, ventral view.

Canada. In the Palearctic Region a boreal, wide- It has been reared from stems of cereals (wheat, spread species ranging from the British Isles to ray, barley, oats) and foddergrass. Johanssen (1960) Japan. reared it from shoots of cereals in Sweden, Rygg (1966a) from seedlings of oats and wheat in Nor- Biology. – Adults from March to October on bogs and other wetlands, also in meadows and agri- way. The following wild grasses and sedges are also cultural fields. Hibernates as adults. Kallio (1950) known as host plants: Festuca pratensis, Calam- listed it as a pest of cereals in Finland. Collected on agrostis epigeios, Leymus arenarius, Poa pratensis agricultural fields in Denmark (Nielsen & Nielsen, and Eriophorum vaginatum (Kreiter, 1928; 1984). Hackman (1963) found adults in borrows of Nartshuk, 1972; Uffen & Chandler, 1978; Krusteva voles (Microtus, Clethrionomys) in Finland. & Beschovski, 2000, 2001).

67 Some larval structures were illustrated by Kreiter 3 Scutellum longer than wide (Figs 161, 201). (1928) and Nartshuk (1987). Wings and wing veins not whitish ...... 4 – Scutellumaslongaswideorshorter(Fig.172). Genus Calamoncosis Enderlein, 1911 Wings and veins M whitish ...... C. duinensis (Strobl) Calamoncosis Enderlein, 1911a: 235. Type species: 4 Scutellum long, triangular, basally flattened, Lipara rufijitarsis auct. nec Loew, 1858 (= Lipara with apical and several subapical setae on dis- minima Strobl, 1893) by original designation. tinct tubercles (Fig. 161). Male genitalia Figs Rhaphiopyga Nartshuk, 1971: 292. Type species: 152,163...... C. aspistylina Duda Calamoncosis glyceriae Nartshuk, 1958 by origi- – Scutellum shorter, convex at base, with api- nal designation. cal and subapical setae on very small tubercles Black species with hairy eyes, surface of ocellar (Fig. 201). Male genitalia Figs 203, 204 triangle shining with many fine setulae situated ...... C. minima (Strobl) in punctures. Face concave with facial carina be- 5 Only one posterior npl seta. Halteres and palpi ing narrowly triangular above and very thin be- black. Male genitalia Figs 152, 153 low. Genae usually narrower than first flagellom- ...... C. aprica (Meigen) ere, vibrissal angle more or less produced. poc upright, convergent. First flagellomere rounded – Two posterior npl setae. Halteres and palpi rectangular, wider than long; arista short. Scutum yellow or palpi more or less darkened . . . . . 6 entirely polished, convex, covered with small setu- 6 Vibrissal angle distinctly projecting beyond lae. Scutellum sometimes with scutellar setae on level of eyes (Fig. 178). Setae and setulae on + small or long tubercles. npl 1 1(2). Tibial organ thorax black. Male genitalia Figs 180–182. large, oval. Femoral organ with numerous warts Ovipositor laterally compressed and cerci scle- on a rounded area. Male genitalia: cerci basally fused, surstyli simple; postgonite relatively large rotized (Figs 183, 184) . . C. glyceriae Nartshuk and broad. – Vibrissal angle obtuse and not projected (Fig. The genus is divided into two subgenera: Calam- 213). Setae and setulae on thorax whitish. oncosis s. str. has female cerci simple and soft, co- Ovipositor not compressed and cerci not vered with numerous hairs; Rhaphiopyga Nartshuk, sclerotized...... C. oscinella (Becker) 1971 has female cerci laterally compressed, faintly sclerotized and sometimes elongated. Distribution. – Probably worldwide. Species key to known 3rd instar larvae of Calamoncosis Key to species of Calamoncosis Enderlein 1 Creeping welts ventrally on abdominal seg- 1 Ocellar triangle reaching 0.75–1.0 of distance ments without spicules. Larvae (Figs 207–211) to front margin of frons (Figs 159, 170, 192). 2–3 in galls of Lipara spp. on Phragmites posterior npl setae...... 2 ...... C. minima (Strobl) – Ocellar triangle shorter, at most reaching – Creeping welts ventrally on abdominal seg- slightly more than halfway to front margin of ments with one or two rows of small spicules frons (Figs 149, 177, 212), if longer then only 1 (Figs167,190)...... 2 posterior npl seta...... 5 2 Stigmal plate of hind spiracles with 3 short 2 Frons except ocellar triangle dull. Notopleura processes and without interstigmatic tufts shining. Male genitalia Figs 195–197 (Figs 164–169). Larvae in stems of Phalaris ...... C. halterata sp. n. arundinaceae ...... C. aspistylina Duda – Frons covered with silky hairs. Notopleura – Stigmal plate of hind spiracles with 4 pro- dusted...... 3 cesses and interstigmatic tufts; these tufts of-

68 ten broken in puparia, but indicated by scars stem of reed contained from 1 to 52 larvae. Pupa- fromtheirbases(Fig.189)...... 3 tion takes place inside the host plant. Adults from 3 Creeping welts ventrally on abdominal seg- a new generation were taken in traps from July 7 to August 11. Grochowska supposed that the species ments with two rows of spicules (Fig. 190). Lar- has two generations per year, but the host plant of vae living gregariously in unsheathed inflores- the summer generation is unknown. cences of Glyceria spp. Egg and larvae of three instars described by Gro- ...... C. glyceriae Nartshuk chowska (2002). – Creeping welts ventrally on abdominal seg- ments only with one short row of spicules (Fig. 158). Larvae in galls formed in stems of Phrag- 11. Calamoncosis (Calamoncosis) mites by Platycephala planifrons aspistylina ...... C. aprica (Meigen) Duda, 1935 Figs 159–169

10. Calamoncosis (Calamoncosis) Calamoncosis aspistylina Duda, 1935: 158. Holotype aprica in ZMHU (Wendt, 1975). (Meigen, 1830) Body black, shining. Frons longer than wide, co- Figs 149–158 vered with silky hairs. Ocellar triangle narrow, ex- tending nearly to front margin of frons, punctate Chlorops apricus Meigen, 1830: 158; Meigen’s plate along side margins, median part smooth, shining. 211: 4 (Morge, 1976b). Lectotype in NHMW Genae as wide as first flagellomere is long. First (Nartshuk, 1997a). flagellomere black, shorter than wide, basal anten- Black, shining except for dull frons. Ocellar trian- nal segments yellow. Scutum convex and punctate, gle short, extending 1/2 to 2/3 of distance to front evenly covered with black setulae. Scutellum elon- margin of frons, with many setulae. Genae a lit- gated. Thick marginal scutellar setae inserted on tle wider than first flagellomere. First flagellomere large tubercles, 3–4 pairs of subapical setae. Pleura shorter than high, black. Palpi black. Pleura shin- shining, anepisternum wrinkled. npl 1 + 2. Hal- ing. npl 1 + 1. Halteres dark. Female cerci simple. teres black. Female cerci simple. Body length 2.5– Body length up to 3 mm. 3.0 mm. Distribution. – Denmark: SJ, EJ, WJ, F, SZ. Sweden: Distribution. – Known only from southern Sweden: SK, SM, VR. Norway: AK, VE, AAy. Finland: Al, Ab, SK and BL. N, Sb. Not found in Russian parts of Fennoscandia. European species. European species. Biology. – Adults from late May to early of August Biology. – Adults from May to July in wet places. in wet meadows. Larvae phytophagous or phytosaprophagous, liv- Larvae phytophagous, live in stem of Phalaris ing as inquilines in stems of Phragmites australis arundinacea (Nartshuk, 1981; Sharipov, 1981). damaged by larvae of Platycephala planifrons or Larvae described by Nartshuk (1981). rarely by caterpillars of Archanara geminipuncta (Noctuidae) (Tschirnhaus, 1981; Grochowska, 2002). Uffen & Chandler (1978) recorded as host 12. Calamoncosis (Calamoncosis) plant foliage of Draba incana, but this seems to be duinensis doubtful. Grochowska (2002) studied the life cycle of the (Strobl, 1909) species in Poland. Adults appear in middle of May Figs 170–176 and lay eggs on stems of Phragmites australis at- Siphonella duinensis Strobl, 1909: 288. Syntypes 2 tacked by Platycephala planifrons. Most larvae of females in NMBA (Chvála, 2008). C. aprica were found among frass in existing lar- val feeding tunnels and had their alimentary tract Body black, shining. Ocellar triangle reaching front filled with some brown substance, but some larvae margin of frons, shining and coarsely punctured were also found feeding on fresh plant tissue. Each except on median glabrous area. Genae as wide

69 Figs 149–158. Calamoncosis aprica (Meigen): 149: head, dorsal view. 150: head, lateral view. 151: scutellum. 152: hypopygium, dorsal view. 153: hypandrium and phallic complex. 154: anterior end of 3rd instar larva. 155: cephalopharyngeal skeleton. 156: anterior spiracle. 157: posterior spiracle. 158: puparium. (154–158 after Grochowska, 2002.) as first flagellomere. Palpi yellow or slightly dark- Widespread Palearctic species, known from Eu- ened, projecting beyond facial margin. Basal seg- rope eastwards to Japan and in North Africa. ment of antennae yellow, first flagellomere dark Biology. – Adults from May to beginning of Septem- brown, broader than long. Scutellum short semicir- ber on Common reed (Phragmites australis). cular widely rounded apically. Apical setae shorter + Larvae phytophagous, live as inquilines in Lipara than scutellum. npl 1 2. Wing slightly whitish. galls on the folded leaves. Halteres with back macula on knob. Female cerci simple. Body length 2.0–2.5 mm. 13. Calamoncosis (Rhaphiopyga) Distribution. – Denmark: WJ, NWJ, F, NEZ. Sweden: glyceriae SK, HA, SM, VS. Finland: Ab, N. Russia: Vib. Not Nartshuk, 1958 found in Norway. Figs 1, 177–191

70 Figs 159–169. Calamoncosis aspistylina Duda: 159: head, dorsal view. 160: head, lateral view. 161: scutellum. 162: cerci and surstyli, dorsal view. 163: male genitalia, ventral view. 164: facial mask of 3rd instar larva. 165: mandibula. 166: tentoropharyngeal skeleton. 167: spicules on 6th abdominal segment. 168: puparium. 169: posterior end of puparium with spiracles.

71 Figs 170–176. Calamoncosis duinensis (Strobl): 170: head, dorsal view. 171: head, lateral view. 172: scutellum. 173: hypopygium, dorsal view. 174: surstylus, lateral view. 175: hypandrium and phallic complex. 176: tip of ovipositor, lateral view.

Calamoncosis glyceriae Nartshuk, 1958: 100. Holo- Distribution. – Denmark: EJ, NWJ, NEZ. Sweden: type in ZISP. SK,VS.Norway:Ø:Gansröd,Fredrikstad.Finland: Om. Russia: Vib, Kr. Body black, only antennae and parts of legs yel- Eurasian species. low. Frons longer than wide, dull. Ocellar triangle reaching halfway to front margin of frons. Frons Biology. – Adults are very abundant in wet places, covered with rather short and thin setulae. Genae near lakes and rivers, from May to September. brown, narrower than first flagellomere, slightly Larvae phytophagous, feed gregariously in un- sheathed inflorescences of Glyceriamaxima, G. flui- produced at vibrissal angle. Antennae yellow with tans, G. aquatica (Collin, 1946, as “C. laminiformis small round first flagellomere, arista shortly pubes- (Becker)”; Nartshuk, 1958). cent and shorter than frons. Palpi rather long, Larvae described by Ismay & Rotheray (1998). brown. Proboscis slightly elongated. Scutum and scutellum coarsely punctate, scutum shining, noto- pleura dusted, pleura shining. Apical scutellar setae 14. Calamoncosis (Calamoncosis) shorter than scutellum, with one or two pairs of halterata rather thin subapical setae. npl 1 + 2. Femora black Nartshuk & Andersson, sp. nov. except basally and at tips, tibiae yellow with a black Figs 192–197 band, tarsi yellow. Halteres yellow. Ovipositor later- ally compressed with sclerotized cerci. Body length Holotype, male: Sweden: Närke: Hjälmarsberg, Öre- 2.5 mm. bro, 26.vii.1941 (T. Nyholm). Paratype, female: Swe-

72 Figs 177–184. Calamoncosis glyceriae Nartshuk: 177: head, dorsal view. 178: head, lateral view. 179: scutellum. 180: hypopygium, dorsal view. 181: surstylus, lateral view. 182: hypandrium and phallic complex. 183: ovipositor, lateral view. 184: tip of ovipositor, lateral view.

73 Figs 185–191. Calamoncosis glyceriae Nartshuk, 3rd instar larva: 185: lateral view. 186: anterior end. 187: cephalopharyngeal skeleton. 188: anterior spiracle. 189: posterior spiracles. 190: spicules on 5th abdominal segment. 191: puparium. (After Ismay, 1998.)

den: Gotland: Hammars, Norrlanda, 12–18.vii.1979 Scutum longer than wide, finely punctated and (R. Danielsson). Holotype and paratype in ZMLU. covered with short dense setulae. Scutellum trian- gular, longer than wide at base, with very small tu- Diffferential diagnosis. –SimilartoC. oscinella bercles at base of apical and subapical setae. Scutel- Becker, but distinguished by longer and narrower lum coarser punctated than scutum and dusted. frons, longer ocellar triangle, shorter dark setu- Apical scutellar setae lost in male, in female only lae on frons, setae and setulae on thorax black one seta present shorter than scutellum; two or (not whitish), genae narrower, and scutellum more three pairs of rather thin subapical setae. Pleura coarsely punctated. shining, anepisternum rugose. npl 1 + 2. Wings Description. – Male, female. Body entirely black ex- longer than abdomen, slightly brownish. Cell br cept for yellow antennae and tarsi of all legs. Frons not widened. Second sector of costal vein 1.7 times longer than wide, dull. Ocellar triangle narrow, ex- longer than third, third costal sector 1.25 longer tending nearly to front margin of frons, on its sur- than fourth. Halteres with stem and knob black. face along side margins with two rows of setulae. Legs black except both ends of tibiae and all tarsi Frons covered with rather short and thin setulae, yellow. Body length 2.5 mm. orb fine, oc a little longer than orb, poc and vte nearly twice as long as oc. Genae punctuated, with slightly produced vibrissal angle, about as wide 15. Calamoncosis (Calamoncosis) as first flagellomere. Antennae yellow with small round first flagellomere, arista shortly pubescent minima and shorter than frons. Eyes large oval with vertical (Strob1, 1893) long axis, shortly pubescent. Palpi short, black. Figs 49, 198–211

74 Figs 192–197. Calamoncosis halterata Nartshuk & Andersson: 192: head, dorsal view. 193: head, lateral view. 194: scutellum. 195: hypopy- gium, dorsal view. 196: surstylus, dorsal view. 197: hypandrium and phallic complex.

Lipara minima Strobl, 1893: 225. Holotype male in Larvae phytophagous, feed on rolled young lives NMBA (Chvála, 2008). of Phragmites or live as inquilines in galls of Li- para species. Usually several larvae in a gall, feed- Body black, shining. Ocellar triangle extending ing on folded leaves above chamber of the Lipara nearly to front margin of frons, shining, punc- larva (Séguy, 1934; d’Aguillar, 1953; Valkeila, 1956; tured except on median glabrous area. Genae as Nartshuk, 1972a). wide as first flagellomere. Palpi yellow to dark Larvae described by d’Aguillar (1953) and Gro- brown, projecting beyond facial margin. Basal seg- chowska (2002). ment of antennae yellow, first flagellomere dark brown, broader than long. Scutum longer than wide. Scutellum elongate triangular, convex basally; 16. Calamoncosis (Calamoncosis) apical setae on small tubercles, shorter than scutel- oscinella lum. npl 1 + 2. Wings hyaline. Halteres with back macula on knob. Female cerci simple. Body length (Becker, 1910) 2.5–3.0 mm. Figs 212–214 Distribution. – Denmark: WJ, LFM, NEZ. Sweden: Lipara oscinella Becker, 1910: 145. Holotype proba- SK,SM,VG,NB,LU.Norway:AK,HEs,Bø,VE.Fin- bly in DEIM (Rohlfien & Edward, 1970). land: north to Om. Russia: Vib. Body black, shining with all setae and setulae Common and widespread, trans-Palearctic spe- white. Frons black, finely punctate and covered cies. with whitish setulae. Ocellar triangle shining, ex- Biology. – Adults from April to September on Com- tending slightly more than halfway to front margin mon reed (Phragmites australis). of frons. Eyes densely pubescent. Genae nearly as

75 Figs 198–211. Calamoncosis minima (Strobl): 198: head, dorsal view. 199: head, lateral view. 200: face and antennae, anterior view. 201: scutellum. 202: wing. 203: hypopygium, dorsal view. 204: male genitalia, ventral view. 205: apex of ovipositor. 206–211: 3rd instar larva: 206: cephalopharyngeal skeleton. 207: facial mask. 208: mandibula. 209: anterior spiracle. 210: posterior spiracle. 211: posterior end. (207 after d’Aguillar, 1943.)

76 Figs 212–214. Calamoncosis oscinella (Becker): 212: head, dorsal view. 213: head, lateral view. 214: scutellum. wide as first flagellomere, not produced beyond eye eye vs. not so produced; profile of head slightly con- margin. Antennae yellow, first flagellomere dark- cave vs. straight; arista pubescent vs. naked; genae ened above. Palpi yellow. Scutum shining, covered no wider than first flagellomere vs. wider than first with whitish setulae. Notopleura and anepister- flagellomere; femoral organ present vs. absent; ter- num in upper part thinly grey-dusted, remaining gites of abdomen ventrally bent and separated by parts of pleura shining. Scutellum convex, as wide ridge from dorsal parts vs. normally shaped. as long. Apical scutellar setae approximated and Duda’s species meijerei further differs from the short, two pairs shorter subapical ones. Abdomen genus Conioscinella by the dusted pleura, wide whitish pubescent. Femora black, tibiae yellow, genae, naked arista and eyes and absence of femo- hind tibiae with black band. Halteres yellow. Body ral organ. Therefore, we erect a new genus Colli- length 2.0–2.3 mm. niella for it. Distribution. – Rare species. Sweden: SK: Skäralid, Etymology. – The genus is named in honour of 09.vi.1965 (H. Andersson), 1 male. Norway: HEs: the famous British dipterist J.E. Collin, who con- Abborhøgda, Kongsvinger. Finland: N: Grankulla, tributed significantly to the knowledge of British 29.v.1918 (R. Frey), 1 female; Espoo (R. Frey), 1 fe- Chloropidae. Gender feminine. male. Not found in Denmark or Russian parts of Fennoscandia. Description. – Body densely dusted. Setae of head Known also from Germany. short and weak, orb 5–6 hardly different from the few indistinct fr. if along side margins of ocellar triangle. Ocellar triangle dusted, extending about Genus Colliniella Nartshuk & halfway to front margin of frons. Eyes oval with di- Andersson, gen. nov. agonal long axis, without hairs. Genae broad. Vib- rissal angle not projected. Facial margin not visible in side view. Facial carina triangular above, narrow Type species: Conioscinella meijerei Duda, 1933 by below, not reaching the facial margin. Arista naked. present designation. Proboscis not prolonged. Conioscinella meijerei was transferred to the genus Scutum dusted with few small dispersed setu- Aphanotrigonum together with some other species lae, ac and dc setulae not differentiated. Thoracic by Collin (1946) because of the dusted pleura. How- setae: 1 h,1+ 2 npl,1pa,1dc, all weak. Scutel- ever, Duda’s species species differs significantly lum short, distinctly convex. Two pairs of marginal from Chlorops trilineatus Meigen, the type species scutellar present. Legs slender, with a short spur of Aphanotrigonum, in other respects. The main on middle tibia. Tibial organ present, femoral or- differences between Aphanotrigonum trilineatum gan absent. Second and third costal sectors of same (Meigen) and Conioscinella meijerei Duda are: length. R2+3,R4+5 and M straight. Abdomen oval, Setae of head and thorax strong and black vs. dusted, tergites not bent on ventral side and not weak and whitish; vibrissal angle produced beyond divided into three parts. Male genitalia: cerci sep-

77 Figs 215–219. Colliniella meijerei (Duda): 215: head, lateral view. 216: scutellum. 217: last segments of tarsus. 218: hypopygium, dorsal view. 219: hypandrium and phallic complex. arated by a circular area; gonites long and narrow, yellow. Male genitalia: gonites narrow and long, not divided. projecting distinctly behind the base of phallus. Body length 1.5–2.0 mm. 17. Colliniella meijerei Distribution. – Denmark:NWJ,NEZ,B.Sweden:SK, (Duda, 1933), comb. n. HA, ÖL, GO. Russia: Vib. Not so far found in Norway Figs 215–219 and Finland, but expected there. The species is known from Europe and Mongolia. Conioscinella meijerei Duda,1933:61.Lectotypein Biology. – Adults in late of May to August on sandy ZMAN (Nartshuk, 2003). beaches with Leymus arenarius. Ardö (1957) reared Body black, covered with light grey dust. Frons yel- the species from leaves of Honckenya peploides, low in front, remaining parts of head black. Ocellar Nartshuk (1962a) from ears of Leymus arenarius on triangle grey-dusted, extending 3/4 of distance to the Karelian Isthmus. front margin of frons. Genae much wider than first flagellomere, vibrissal angle not produced. First flagellomere rounded or slightly angulate, dark- Genus Conioscinella Duda, 1929 ened to black with yellowish base. Genae more than 1/3 as wide as eye height. Palpi yellow. All se- Conioscinella Duda, 1929: 166, 169. Type species: tae of head white. Scutum grey-dusted, with white Oscinella soluta Becker, 1912 by designation of hardly visible setulae. Pleura mainly dusted. npl Sabrosky, 1941: 751. 1 + 2. Abdomen grey, with pale setulae. Femora mainly black, tibiae yellow, medially black; middle Black grey-dusted species with evenly arranged se- tibiae with spur, hind tibiae without spur. Halteres tulae on scutum. Ocellar triangle short, dusted.

78 Genae with weakly produced vibrissal angle. poc 7 Setae on head long, orb about 8. Male geni- upright, cruciate. First flagellomere rounded oval. talia: epandrium enlarged (Figs 235–237) + Pleura mainly shining. npl 1 2. Tibial organ oval...... C. tornensis sp. nov. Femoral organ consists of a row of warts. Male geni- talia: cerci and surstyli simple; hypandrium usually – Setae on head short, orb no more than 6. Male open, but sometimes closed. genitalia: epandrium normal sized (Figs 220– 222,228,229)...... 7 Distribution. –Worldwide. 7. Male genitalia: cerci triangular, wide apart and directed ventrally (Figs 220–222) Key to species of Conioscinella Duda ...... C. abiskoi sp. nov. – Male genitalia: cerci quadrate and directed 1 Wings much shorter than abdomen, some- mesad(Fig.228)...... C. livida Nartshuk times absent; even halteres sometimes absent Note. – A species group closely related to C. fron- (Fig. 40). Male genitalia Fig. 240 tella (Fallén) occurs in the northern part of distri- ...... C. zetterstedti Andersson bution area of the genus. These species are darker – Wings normally developed, longer than ab- coloured than C. frontella and have wider genae. domen,halteresalwayspresent...... 2 They are rather difficult to separate and exami- 2 Setae and setulae on head and sometimes on tation of the male genitalia is essential for reli- scutumyellowish...... 3 able identification. Three species are described here, but more species may show up in northern – Setae and setulae on head and on scutum Fennoscandia. black...... 4 3 Ocellar triangle and scutum shining, scutum densely covered with yellow setulae. Male 18. Conioscinella abiskoi genitaliaFig.227...... C. gallarum (Duda) Nartshuk & Andersson, sp. nov. – Ocellar triangle and scutum dusted and dull. Figs 220–222 Setae and setulae on scutum black, rarely yellow...... C. sordidella (Zetterstedt) Holotype, male: Sweden: Norrbotten: Tärendö, 22.vi.1983 (S.O. Ulefors). Paratypes: 1 female, Swe- 4 Genae not wider than front tibiae. Vibrissal den: Torne Lapmark: Vittangi, 16.vii.1955, on Carex angle not strongly projected beyond eye mar- (H. Andersson); 2 females, Torne Lapmark: Abisko, gin(Fig.224)...... 5 Betula-forest SW of marble quarry, 22.vii.1983, – Genae wider than fore tibiae, nearly equal 2500 m (H. Andersson). Holotype and paratypes in to width of first flagellomere. Vibrissal angle ZMLU. strongly projected beyond eye margin . . . . . 6 Diffferential diagnosis. – Similar to C. frontella,but 5 Femora dark in both sexes, middle tibiae darker coloured with more produced vibrissal an- broadly darkened in males, narrowly dark- gle. Male genitalia: cerci smaller and wider apart, ened in females. Frontal setae long. Male gen- hypandrium closed. italia Figs 225, 226 ...... C. frontella (Fallén) Description. – Male, female. Frons black, narrowly – Femora dark in males only, middle tibiae yel- yellow anteriorly. Ocellar triangle black, dusted, ex- low in both sexes. Frontal setae short. Male tending nearly to middle of frons. Basal antennal genitalia Figs 232, 233 . . . . . C. mimula Collin segments brown, reniform first flagellomere yel- 6 Genae as wide as first flagellomere, vibrissal low, darkened above. Genae narrower than first angle distinctly produced and proboscis long. flagellomere, vibrissal angle produced. Genae and Male genitalia Figs 230, 231 face yellow, except genae black along lower margin. Palpi yellow...... C. messaurea sp. nov. Thorax black, scutum dusted and evenly covered – Genae narrower than first flagellomere, vib- with setulae. Scutellum short. Distance between rissal angle less produced and proboscis apical setae nearly the same as between apical and shorter...... 7 nearest subapical setae. Pleura shining except on

79 Figs 220–226. Conioscinella abiskoi Nartshuk & Andersson: 220: hypopygium, dorsal view. 221: surstylus, lateral view. 222: hypandrium and phallic complex. C. frontella (Fallén): 223: head, dorsal view. 224: head, lateral view. 225: hypopygium, dorsal view. 226: hypandrium and phallic complex. upper part of anepisternum. npl 1 + 2. Postno- low, darkened above. Arista black. Eyes sparsely tum shining. Abdomen black. Legs black with tib- haired. Palpi yellow. Scutum and scutellum grey- iae at both ends and tarsi yellowish. Halteres yel- dusted and evenly covered with setulae. Scutellum low. Wings hyaline, venation unremarkable for the rounded, apical setae longer than scutellum. Pleura genus. Male genitalia: cerci small and rather wide only dusted above, shining on remaining surface. apart, surstyli narrow, hypandrium closed. npl 1 + 2. Abdomen brown, covered with black se- tulae. Sexual dimorphism in colour of legs: male legs mainly dark with knees, tibiae distally and tarsi 19. Conioscinella frontella except last segment yellowish; female legs paler (Fallén, 1820) with front coxae, all trochanteres, front and middle Figs 223–226, 978–980 tibiae and hind tibiae basally and distally yellowish. Body length nearly 2.0 mm. Oscinis frontella Fallén, 1820: 7. Lectotype in ZMLU (Andersson, 1963). Distribution. – Common species. Denmark: EJ, WJ, Body black, grey-dusted. Frons narrow, yellowish NEJ, LFM, NWZ, NEZ, B. Sweden: north to LU and in front. Ocellar triangle black grey-dusted, extend- NB.Norway:AK,HEs,HEn,Bø,AAy,VAy,Ry,HOi, ing 3/4 of frons. First flagellomere rounded, yel- STi, Nsi. Finland: north to Li. Russia: Vib, Kr, Lr.

80 Widely distributed Palearctic species, known bud of Populustremula in Finland and from Cirsium from Europe to Mongolia and Israel. head in Karelia (Russia) (specimens in collections). Biology. – Adults from middle of June to middle of August in meadows. Hackman (1963) found adults 21. Conioscinella livida in burrows of voles (Microtus, Clethrionomys)in Nartshuk, 1970 Finland. Figs 228, 229 Larvae phytophagous, living in shoots of many grasses: Elytrigia repens, Anthoxanthum odoratum, Conioscinella livida Nartshuk in Nartshuk et al., Phleum pratense, Dactylis glomerata, Deschampsia 1970: 418. Holotype in ZISP. caespitosa, D. flexuosa, Festuca ovina, Holcus lana- Body black, bluish grey-dusted. Frons narrow, yel- tus (Balachowsky & Mesnil, 1935; Nye, 1958, 1959; lowish in front. Ocellar triangle black, grey-dusted, Nartshuk, 1962a, 1972a; Karpova, 1972; Tschirn- extending halfway to front margin of frons, poorly haus, 1981, 1992; Bährmann & Weipert, 1989) and delimited from remaining frons. First flagellom- in oats and barley stems (Krusteva & Beschovski, ere rounded, yellow, darkened above. Arista black. 1998, 2000). A record from Silene dioica appears Eyes haired. Palpi yellow. Scutum and scutellum unreliable. Larvae hibernate in the basal tillers of bluish grey-dusted and evenly covered with setu- grasses and pupate the following spring. lae. Scutellum rounded, apical setae shorter than Larvae described by Balachowsky & Mesnil scutellum. Distance between apicals a little longer (1935) and Nye (1958). than distance between apical and subapical se- tae. Pleura dusted above and shining on remaining 20. Conioscinella gallarum surface. npl 1 + 2. Abdomen brown, covered with black setulae. Legs black. Halteres yellow. Body (Duda, 1933) length nearly 2.0 mm. Fig. 227 Distribution. – Finland: N: Espoo, Sammatti; Sb: Tropidoscinis gallarum Duda, 1933: 82. Syntypes in Varpaisjärvi. Total 3 males, 4 females, all taken in ZMHU (Wendt, 1975). June.Russia:Lr:Petsamo,1male.Notfoundinother Nordic countries. Body black, shining. Frons yellow anteriorly and European species, known from Estonia, St. Pe- gradually darkened posteriad. Ocellar triangle tersburg district and Germany (Bavaria) (Tschirn- black, shining, very sparsely dusted, nearly reach- haus & Schacht, 2000). ing front margin of frons. Antennae yellow, first flagellomere slightly wider than long, infuscate dor- Biology. –Adultsinbogs. sally. Genae about half as wide as first flagellom- ere, pale yellow and shining black behind. Scutum shining and evenly covered with short yellowish 22. Conioscinella messaurea setulae. Scutellum wider than long, dusted, apical Nartshuk & Andersson, sp. nov. setae longer than scutellum, a pair of subapical se- Figs 230, 231 tae shorter. Pleura shining. npl 1 + 2. All head and thoracic setae yellow. Halteres yellow. Legs entirely Holotype, male: Sweden: Lule Lapmark: Messaure, yellow. Body length 1.5–2.0 mm. 15–27.vii.1974 (K. Müller). Paratypes: 31 males, 57 females, same data as holotype except 17.vi– Distribution. – Denmark: NEZ. Sweden: SK, HA, 15.viii.1974. All specimens in alcohol. Holotype and NÄ,SÖ,VS.Norway:AK:Toien,Oslo,23.vi.1851 most paratypes in ZMLU, some paratypes in ZISP. (Siebke). Finland: Ab, N, St. Russia: Vib. Trans-Palearctic species, known from the British Diffferential diagnosis. – Similar to C. frontella,but Isles to Japan. differs in following characters: genae wider, nearly as wide as first flagellomere, vibrissal angle dis- Biology. – Adults from May to end of July in forests tinctly produced beyond eye margin and proboscis and bushes. Adults have been reared from galls longer. General colour darker, lower margin of of Andricus and Biorrhiza pallida (Hymenoptera, genae blackish, frons entirely dark without yellow Cynipidae) on oak, from cones of Larix decidua band in front, all femora and tibiae black in both (Duda, 1933; Nordlander & Grijpma, 1991), from sexes, only tarsi yellowish. Antennae yellow with

81 Figs 227–233. Conioscionella gallarum (Duda): 227: male genitalia, ventral view. C. livida Nartshuk: 228: hypopygium, dorsal view. 229: hypandrium and phallic complex. C. messaurea Nartshuk & Andersson: 230: hypopygium, dorsal view. 231: hypandrium and phallic complex. C. mimula Collin: 232: hypopygium, dorsal view. 233: hypandrium and phallic complex.

82 first flagellomere reniform, shorter than high, dark- Oscinis sordidella Zetterstedt, 1848: 2659. Holotype ened, but not black; arista black, short pubescent. in ZMLU (Andersson, 1966). Male genitalia: differences from C. frontella rather Conioscinella flavifrons Duda,1933:52,asvar.ofC. subtle, but cerci wider apart. frontella (Fallén) (Tschirnhaus, 1992). Description. – Male, female. Ocellar triangle dull, Black, grey-dusted with yellow setae and setulae extending halfway to front margin of frons. Frons on head and thorax. Frons yellow. Ocellar trian- covered with small black setulae. All setae of head gle black, grey-dusted, extending halfway to front small. orb numerous, no less than 7–8 on each margin of frons. First flagellomere rounded, yel- side of frons. Face and genae fuscous yellow. Genae low. Arista black. Eyes sparsely haired. Palpi yellow. distinctly produced, as wide as first flagellomere, Scutum and scutellum grey-dusted and evenly co- blackish along lower margin. Palpi fuscous yellow vered with pale setulae. Scutellum rounded, apical to blackish. Proboscis with elongated labella. setae longer than scutellum. Pleura dusted above Scutum and scutellum black, grey-dusted. Scu- and shining on remaining surface. npl 1 + 2. Ab- tum evenly covered with setulae. Scutellum domen brownish. Legs entirely yellow, including rounded behind, a little shorter than wide at base. coxae. Body length nearly 2.0 mm. Apical setae longer than scutellum, subapical setae small, distance between apical and subapical se- Distribution. – Denmark: LFM, NEZ, B. Sweden: tae equals 1/2 distance between apical setae. Pleura north to VS. Norway: AK, HEs (Thunes et al., 2004). shining except on upper part of anepisternum. npl Finland: Al, Ab, N. Russia: Vib, Kr. 1 + 2. All setulae and setae of head and thorax European species. black. Abdomen brown. Wing venation typical for Biology. – Adults from end of June to end of Au- the genus. Halteres yellow. Legs black, tarsi some- gust in dry meadows; found in canopies of Pinus times partly yellowish. Body length 1.75–2.0 mm. silvestris in Norway (Thunes et al., 2004). Larvae phytophagous, live in shoots of Dactylis 23. Conioscinella mimula glomerata, Lolium perenne, Leymus sp., Collin, 1946 sp., Anthoxanthum odoratum, Elytrigia repens (Kar- Figs 232, 233 pova, 1972; Tschirnhaus, 1981, 1992). Conioscinella mimula Collin, 1946: 129. Syntypes 2 males, 9 females in OXUM (Pont, 1995). 25. Conioscinella tornensis Very similar to C. frontella, but smaller, male legs Nartshuk & Andersson, sp. nov. with yellow front and middle tibiae, female legs en- Figs 235–237 tirely yellow. Frons in upper part dark, frontal and Holotype, male, Sweden: TO: Stordalen, 18.vii.1957 thoracic setae and setulae dark, first flagellomere (P.I. Persson). There is another male from the same yellow, brownish above. Body length 1.0–1.5 mm. locality and with the same date, but without head. Distribution. – Denmark: EJ, NEJ, F, LFM, NEZ, B. Holotype in ZMLU. Sweden:northtoJÄ.Finland:northtoTa.Russia: Vib. Not so far found in Norway. Diffferential diagnosis. –LargerthanC. frontella and European species. differs by darker body, longer orb and other se- tae, especially on femora and abdomen, darkened Biology. – Adults from June to middle of August in wings and larger epandrium. meadows. Larvae phytophagous, live in shoots of Anthoxan- Description. – Male. Frons black, only narrowly thum odoratum, Bromus sp., Lolium perenne,hiber- brown in front. Ocellar triangle extending a little nate in shoots of their host plant and pupate fol- more than halfway to front margin of frons, densely lowing spring (Nye, 1958, 1959; Tschirnhaus, 1981). dusted. orb numerous (c. 8) and rather long, longer Larvae described by Nye (1958). than vti and oc, but thinner. Eyes ovoid, covered with short hairs. Genae as wide as first flagellom- ere, fuscous yellow, black on lower margin, with 24. Conioscinella sordidella distinctly produced vibrissal angle. Antennae yel- (Zetterstedt, 1848) low, first flagellomere reniform; arista black, shortly Fig. 234 pubescent. Palpi yellow.

83 Figs 234–240. Conioscinella sordidella (Zetterstedt): 234: head, lateral view. C. tornensis Nartshuk & Andersson: 235: hypopygium, dorsal view. 236: surstylus, medial view. 237: hypandrium and phallic complex. C. zetterstedti Andersson: 238: head, lateral view. 239: scutellum. 240: hypandrium and phallic complex.

Scutum and scutellum black, grey-dusted, evenly Conioscinellazetterstedti Andersson, 1966: 72. Holo- covered with setulae. Scutellum semicircular. Api- type in ZMLU. cal setae longer than scutellum, two subapical se- tae on each side. Pleura shining except on upper Head black, front margin of frons, face and genae part of anepisternum. npl 1 + 2. Postnotum black, yellow. Ocellar triangle black, dusted, extending shining. 3/4 of frons. Basal antennal segments yellow, first Abdomen black, covered with rather long setae. flagellomere rounded, yellow and darkened above, Legs entirely black, including tarsi. Wing brownish especially on outer side. Scutum smooth and with venation usual for the genus. Halteres yel- dusted. Scutellum very short and varies in size. low. Male genitalia: epandrium large, with rounded It is proportionally larger in full-winged speci- cerci and slightly tapering surstyli; hypandrium mens. Apical scutellar setae wide apart. Wings vary open; postgonite with long hairs. Body length markedly in size. Venation reduced in short-winged 2.2 mm. specimens. Halteres yellow, smaller in short- winged specimens. Rare full-winged specimens oc- 26. Conioscinella zetterstedti cur: Finland: Ta: Hartola, 11.vi.1987 (Koponen), 1 Andersson, 1966 specimens; Ok: Paltamo, 03.vii.1991 (Koponen), 1 Figs 40, 238–240 specimen. Femora black, front and middle tibiae

84 yellow, especially in females, hind tibiae black ex- The genus is divided into several subgenera cept yellow at ends. Body length 1.25–1.5 mm. mainly based on differences in the male genitalia. The subgenus Dicraeus has slender male cerci in Distribution. – Denmark: EJ, WJ, NWJ, NEJ, NEZ. a lateral position near base of simple surstyli. The Sweden: SK, GO, UP, along coast of the Baltic Sea subgenus Oedesiella has similar male cerci, but to NB. Finland: N, Ta, Sa, Om, Ok, ObS, Li. Russia: surstyli are long, of complex shape and with dis- Vib, Kr. Not found in Norway. tinct setae. In the subgenus Paroedesiella male Holarctic species. cerci are minute and in median position or absent, Biology. – Adults xerobiontic, from middle of May and surstyli are short and broad with spine-like se- to end of September, preferable in coastal dune ar- tae. eas, but sometimes inland. Biology and variability in size of the wings and halteres have been stud- Distribution. – Probably worldwide, but not so far ied by Brauns (1937, 1938, 1959, as Conioscinella found in the Neotropical Region. brachyptera). Ardö (1957) found adults in dunes in dense tufts of Leymus and Ammophila. Flies are ex- Key to species of Dicraeus Loew cellent jumpers and run rapidly on plants and on the sand surface. It has also been found in mam- 1 Costal vein ending between apices of veins mals’ nests, peat-bogs, swamps, shore debris and R4+5 and M. Male genitalia Figs 243, 244 leaf litter (Mohrig, 1967; Wheeler, 1994b; Beaulieu ...... D. ingratus (Loew) & Wheeler, 2001). Adults were found in Sphagnum – Costal vein extending to end of vein M . . . . . 2 in February in the Netherlands (Pelt, 1926). Larvae have been found in rotting shoots of Am- 2 Crossvein dm-cu absent (Fig. 247). Male geni- mophila arenaria injured by Meromyza pratorum taliaFigs248,249...... D. raptus (Haliday) (Balachowsky & Mesnil, 1935). Tschirnhaus (1981) – Crossvein dm-cu present(Fig.37)...... 3 listed Elytrigia, Ammophila and Molinia coerulea as 3 Postpronota and pleura partly yellow, in dark host plants. Karpova (1972) reared the species from specimens yellow parts not distinct. Male gen- Agropyron cristatum. Beaulier & Wheeler (2001) found the species on Carex in Canada. italiaFigs266–268...... D. vagans (Meigen) – Thoraxentirelyblack...... 4 4 Pleura dusted; only one posterior npl setae. Genus Dicraeus Loew, 1873 Male genitalia Fig. 258 ...... D. tibialis (Macquart) Dicraeus Loew, 1873: 51. Type species: Dicraeus ob- – Pleura mainly shining, dusted only on upper = scurus Loew, 1873 ( Oscinis raptus Haliday, part of anepimeron and anepisternum; two 1838) by monotypy. posterior npl setae...... 5 Oedesiella Becker, 1910: 146. Type species: Oedesiella discolor Becker, 1910 by monotypy. 5 Genae and front margin of frons yellow . . . . 6 Paroedesiella Enderlein, 1936: 187. Unavailable. – Genae fuscous yellow or blackish, frons en- Post-1930 name proposed without designation tirely black. Genae distinctly wider than first ofatypespecies. flagellomere (Figs 245, 246) Paroedesiella Anonymous, 1937: 394. Type species: ...... D. nitidus (Wahlgren) Oscinella styriaca Strobl, 1898 by monotypy. Paroedesiella Sabrosky, 1941: 761. Type species: Os- 6 Ocellar triangle with thin layer of dust . . . . . 7 cinella styriaca Strobl, 1898 by original designa- – Ocellar triangle shining, without dusting . . . 8 tion. 7 Male genitalia Fig. 242 . . . . . D. fennicus Duda Black or yellow species with black stripes on scu- – Male genitalia Figs 250–252 tum, with short semicircular scutellum, rather ...... D. rossicus Stackelberg broad genae, long vein R + . poc upright and cruci- 2 3 8 Male genitalia Figs 254–256 ate. Ocellar triangle usually thinly dusted. First flag- ellomere rounded oval, arista shortly pubescent...... D. styriacus (Strobl) Tibial organ distinct, narrow.Femoral organ formed – Male genitalia Figs 259–261 by a group of warts with short stout spines...... D. tjederi sp. nov.

85 27. Dicraeus (Oedesiella) fennicus Distribution. – Only found in Denmark: B: Baste- Duda, 1933 mose, 26.vi.1964 (O. Martin & B.V.Petersen), 1 male, Figs 105, 106, 241, 242 1 female. Holarctic species, in Palearctic known from Eu- rope, southern Siberia, Israel. Dicraeus fennicus Duda,1933:67,asvar.ofDicraeus styriacus Strobl. Syntypes in NHMH. Biology. – Larvae phytophagous, feeding on de- Body black, thinly dusted. Ocellar triangle black, veloping seeds of Bromus inermis and B. erectus slightly dusted, extending nearly to front margin of (Agafonova, 1962; Nartshuk, 1967; Gołebiowska & frons. Face, frons except for hind part and genae Nawrot, 1970). yellow; genae wider than first flagellomere. Basal antennal segments yellow, first flagellomere yel- low, darkened above. Palpi yellow. Scutum thinly 29. Dicraeus (Paroedesiella) nitidus dusted, pleura shining except on upper part of Wahlgren, 1913 anepisternum. npl 1 + 2. Abdomen brown. Femora Figs 245, 246 and hind tibiae black, other tibiae and tarsi yellow. Body length 2.0–2.5 mm. Dicraeus nitidus Wahlgren, 1913: 52. Holotype fe- male in ZMLU. Distribution. – Rather common. Denmark: WJ, NEJ, Dicraeus napaeus Collin, 1946: 133. Holotype male NWZ, NEZ, B. Sweden: north to UP including GO, in OXUM (Pont, 1995). Syn. n. alsoNB.Norway:Ø,AK.Finland:Al,Ab,N,Ka,St, Ta, Om. Russia: Vib, Kr. Note. – Dicraeus nitidus Wahlgren was described Holarctic species, recorded in the Palearctic from on a female from Öland. We had the opportunity Europe to Yakutia, East Siberia and Mongolia. to examine males collected on Öland that coincide in all essential characters with the holotype of D. ni- Biology. – Adults from end of June to end of July, tidus. The male genitalia of these specimens agree rare in August. with a drawing of D. napaeus by Collin (1946). Larvae phytophagous, feeding on the unripe Body black, thinly grey-dusted. Ocellar trian- seeds of Elytrigia repens, Elymus pungens, E. pyc- gle black. Frons in front, face and genae blackish. nanthus and Elymus sp. (Parmenter, 1955; Nartshuk, Genae wider than first flagellomere, lower part of 1960, 1967; Tschirnhaus, 1981). Nielsen & Nielsen genae darker and punctate. Antennae nearly black. (1984) recorded this species on agricultural grasses Palpi brownish. Pleura mainly shining. npl 1 + 2. in Denmark, but this is probably because Elytrigia Legs black with yellowish ends of tibiae and yel- repens is a common weed everywhere. lowish brown tarsi. Abdomen black. Body length 2.0–2.3 mm. 28. Dicraeus (Dicraeus) ingratus Distribution. – Denmark: NWJ: Tved Plantage; B. (Loew, 1866) Sweden:northtoUP.Finland:Al.NotfoundinNor- Figs 243, 244 way or Russian parts of Fennoscandia. European species. Eutropha ingrata Loew, 1866: 26. Type material not Biology. – Adults in June to July in meadows. found in ZMHU (Wendt, 1975, 1978). Larvae phytophagous, but host plant previously Body black, thinly dusted. Frons fuscous yellow. unknown. We have examined specimens from Ocellar triangle black, dusted, extending to 4/5 of Öland reared from unripe seeds of Helictotrichon frons. Face and anterior part of genae fuscous yel- pratense. The distribution of the species in the area low, clypeus black. Genae as wide as or wider than investigated coincides well with the distribution of first flagellomere. Basal antennal segments brown, the host grass. first flagellomere nearly black. Palpi black. Scutum thinly dusted, pleura shining except on upper part of anepisternum. npl 1 + 2. Abdomen black. Legs 30. Dicraeus (Dicraeus) raptus mainly black. Wings clear, costal vein not reaching (Haliday, 1838) end of vein M. Body length 2.0–2.5 mm. Figs 247–249

86 Figs 241–252. Dicraeus fennicus Duda: 241: head, lateral view. 242: hypopygium, postero-lateral view. D. ingratus (Loew): 243: hypopy- gium, dorsal view. 244: hypopygium, lateral view. D. nitidus Wahlgren: 245: hypopygium, dorsal view. 246: surstylus, medial view. D. raptus (Haliday): 247: wing. 248: hypopygium, dorsal view. 249: hypopygium, postero-lateral view. D. rossicus Stackelberg: 250: hypopygium, dorsal view. 251: hypopygium, lateral view. 252: surstylus, medial view. (242 after Collin, 1946.)

87 Oscinis raptus Haliday, 1838: 187. Type in BMNH. Oscinis styriaca Strobl, 1898: 263. Syntype male in NMBA (Chvála, 2008). Body black, grey-dusted. Frons at front margin, Dicraeusvallaris Collin, 1946: 133. Syntypes 8 males, face and genae yellow. Ocellar triangle black, thinly 7 females in OXUM (Pont, 1995). Synonymy dusted and shining, extending to 2/3 of frons. An- established by J.W. Ismay and published by tennae and palpi yellow, first flagellomere dark- Nartshuk (2002a). ened above. Genae wider than first flagellomere. Scutum black, dusted, pleura mainly shining, Body black, grey-dusted. Frons in front, face and dusted above. npl 1 + 2. Abdomen brown with yel- genae in front yellow. Genae as wide as first flag- low base. Wing without crossvein dm-cu.Halteres ellomere. Antennae with basal segments yellow or yellow. Legs with all femora and hind tibiae black. brownish yellow, first flagellomere brownish. Palpi Body length 2.0 mm. brownish yellow. Pleura mainly shining with yellow + Distribution. – Only from Sweden: Öland: Köpinge anepimeral suture. npl 1 2. Legs black, with yel- alvar, 1 female, also recorded by Ringdahl (1947). lowish knees and tarsi. Abdomen brownish. Body European species. length 2.0–2.3 mm. Biology. – Larvae phytophagous, associated with Distribution. – Denmark: B. Sweden: SK, BL, ÖL, Bromus ramosus (Ismay, 1981d). GO,ÖG,UP,DR.Finland:Al,Ab,N,St.Russia:Vib. Not found in Norway. The distribution coincides well with the distribution of its host plant (map). 31. Dicraeus (Oedesiella) rossicus European species. Stackelberg, 1955 Biology. – Adults from June to beginning of July in Figs 250–252 meadows. Dicraeus rossicus Stackelberg, 1955: 332. Holotype Larvae phytophagous, feeding on developing in ZISP. seeds of Helictotrichon pubescens (Nartshuk, 1960, 1967) and probably Leymus arenarius. Similar to D. fennicus, but ocellar triangle and scu- tum more shining. Body black, frons narrowly yel- low in front. Face and genae yellow, occiput black. 33. Dicraeus (Dicraeus) tibialis Ocellar triangle subshining black, extending 4/5 (Macquart, 1835) length of frons. Genae wider than first flagellom- Figs 257, 258 ere. Palpi yellow. Scutum and scutellum thinly grey- dusted, subshining. Scutellum semicircular, apical Oscinis tibialis Macquart, 1835: 263. Types probably setae longer than scutellum. Pleura shining except in MHNL. on upper part of anepisternum. Sulcus between anepisternum and anepimerom yellow. npl 1 + 2. Body mainly black with face, genae, palpi, anterior Legs mainly black, front and middle tibiae yellow- part of frons, halteres and abdomen yellow. First ish. Halteres yellow. Body length 2.0 mm. flagellomere yellow basally. Ocellar triangle ex- tending 1/2 length of frons, covered with thin dust. Distribution. – Finland: Al: Jomala, 1 female; N: Genae slightly wider than first flagellomere. Scu- Helsinki, 2 females. Russia: Kr: Sonostroff (Suon- tum and pleura mainly dusted. npl 1 + 1. Femora saari), Nemetskiy Kuzov Is. Lr: Kandalaksha, 1 fe- black, anterior tibiae yellow to brownish. Abdomen male. The Finnish records are in need of verifica- yellow with lateral dark spots at base. Body length tion as based on females only. 1.8–2.0 mm. Trans-Palearctic species, known from Europe to Japan. Distribution. – Denmark: SJ, LFM, NEZ. Sweden: SK, SM. Finland: Al, N. Russia: Kr. Not found in Nor- Biology. – Larvae phytophagous, feeding on deve- way. loping seeds of Elymus caninus (Nartshuk, 1960, Holarctic species, found more recently in New 1967). Zealand (Ismay, 1991). Biology. – Adults in June to early July in meadows. 32. Dicraeus (Paroedesiella) styriacus Larvae phytophagous, feeding on developing (Strobl, 1898) seeds of Bromus inermis, B. erectus and Helictotri- Figs 52, 253–256 chon pubescens (Nartshuk, 1960, 1967; Agafonova,

88 Figs 253–261. Dicraeus styriacus (Strobl): 253: head, lateral view. 254: hypopygium, dorsal view. 255: surstylus, ventral view. 256: hypan- drium and phallic complex. D. tibialis (Macquart): 257: head, lateral view. 258: hypopygium, dorsal view. D. tjederi Nartshuk & Andersson: 259: hypopygium, dorsal view. 260, 261: surstylus, lateral and medial views.

89 1962) and probably other grasses. Falcoz (1921) partly yellow. In dark specimens postpronota with reared the species from a mole hole. a dark spot or all thorax black. Head black on oc- ciput and ocellar triangle, yellow in front. Ocellar triangle extending 3/4 length of frons. Antennae 34. Dicraeus (Paroedesiella) tjederi and palpi yellow. Genae wider than first flagellom- Nartshuk & Andersson, sp. nov. ere. npl 1 + 2. Abdomen yellow or more or less Figs 259–261 darkened. Legs mainly yellow. Body length nearly 2.0 mm. Holotype, male, Sweden: SK: Klammersbäck, Rau- lunda, 15.vi.1948 (B. Tjeder). Holotype in ZMLU. Distribution. – Denmark: common and wide- spread. Sweden: north to UP, also GO. Finland: Diffferential diagnosis. – Belongs to the subgenus Al, N. Russia: Vib: Bolshoi Beresovyi Is. near Pri- Paroedesiella, because of male genitalia: cerci ab- morsk. Not found in Norway. sent, surstyli short. It is most similar to D. styriacus European species. but differs in structure of the male genitalia: inci- sion of surstyli deeper in the new species, almost as Biology. – Adults from end of May to middle of July in D. vagans, and lower margin of surstylus without in meadows and at forest edges. a projection on anterior part as in D. vagans and D. Larvae phytophagous, feeding on developing styriacus. seeds of Arrhenatherum elatius (Nartshuk, 1960, Description. – Male. Head black except yellow on 1967). face, genae and narrowly on anterior part of frons. Ocellar triangle shining without any dust. Chaeto- taxy of head typical for the genus. Frons and face in Genus Elachiptera Macquart, 1835 the holotype rather deformed and antennae thus partly retracted. Antennae black. Genae not wider Elachiptera Macquart, 1835: 621. Type species: Chlo- than first flagellomere, with a row of black setulae rops brevipennis Meigen, 1830 by original desig- along lower margin. nation. Thorax entirely black, scutum thinly dusted and New generic synonyms published by Nartshuk & subshining except posterior part more densely Tschirnhaus (2012). dusted and dull like scutellum. Scutum evenly covered with black setulae. Scutellum triangular, Black or rarely yellow species with slightly flattened apical setae longer than scutellum, subapical se- scutum and scutellum and scutellar setae arising tae inserted a little nearer to base of scutellum from small warts. Scutum with rather coarse stel- than to apical setae. Pleura shining. npl 1 + 2. late microsculpture, evenly dusted or dust arranged Wings clear with brown veins. Costal vein reach- in lines along dorsocentral lines. First flagellom- ing end of M. Second costal sector 3.5 times longer ere oval or reniform, arista usually broadly thick- than third one. Third costal sector 1.5 times longer ened with dense, rather long hairs. Tibial organ short, oval. Femoral organ present as two rows or a than fourth. Veins R4+5 and M parallel. Cell br not widened. Halteres yellow. Legs with all femora group of warts. Wings sometimes reduced. In some black, except at tips, front and middle tibiae fus- species basal tergites fused. Male genitalia with cous yellow, hind tibiae with black band. Abdomen separate cerci, simple surstyli and broad triangu- dark brown. Body length 2 mm. lar postgonites. Distribution. – All regions except Australasia. 35. Dicraeus vagans (Meigen, 1830) Key to species of Elachiptera Macquart Figs 51, 262–268 1 Wings shortened, barely reaching end of ab- Chlorops vagans Meigen, 1830: 389. Type lost, in domen or shorter (Fig. 2). Body partly red- MNHN only a pin with Meigen’s name label as dish yellow, abdomen black. Syntergite 1 + 2 already noted by Becker (1902). of abdomen long, occupying basal half of ab- Body yellow with black scutum except for yellow domen. Male genitalia Figs 271, 272 postpronota and stripes along notopleura. Pleura ...... E. brevipennis (Meigen)

90 Figs 262–268. Dicraeus vagans (Meigen): 262: head, dorsal view. 263: head, lateral view. 264: wing. 265: scutellum. 266: hypopygium, ventral view. 267: surstylus, medial view. 268: hypandrium and phallic complex.

91 – Wings normally developed, usually longer occupying 1/2 length of abdomen. Wing length ei- thanabdomen...... 2 ther nearly as long as abdomen or only extending + 2 Scutum evenly thinly dusted or shining only to hind margin of syntergite 1 2, in this case with onsides...... 3 reduced venation. Body length 1.5–2.0 mm. – Scutum mainly shining, with two dusted Distribution. – Found only in Denmark: SJ, EJ, F, stripes and sometimes dusted before scutel- NEZ. lum(Fig.280)...... 4 European species. 3 Scutellum longer than wide with long tuber- Biology. – Adults in May to September in fens, cles at base of apical setae (Fig. 287). Arista costal marches and other wetlands with Carex, Scir- thick, flattened. Legs yellow pus and Typha. Adults crawl on plants near to the ...... E. tuberculifera (Corti) ground (Mohrig, 1967). Hibernates as adults. Tschirnhaus (1981) reared the species from – Scutellum not as long as wide with small tu- shoots of Dactylis glomerata. Some early authors bercles at base of apical setae (Fig. 284). Arista (Smith, 1856; Brauer, 1883) reported that eggs of thin, not flattened but densely pubescent. this species were found under the short wings of Legs mainly dark . . . . . E. scrobiculata (Strobl) the bug Nabis subapterus, but this record seems be 4 Apical scutellar setae nearly twice as wide doubtful. Vimmer (1906) discussed reduction of ve- apart as their distance to lateral setae (Fig. nation in brachypterous specimens. 282). Scutum dusted before scutellum 37. Elachiptera cornuta ...... E. diastema Collin – Apical scutellar setae close together, barely (Fallén, 1820) wider apart than their distance from lateral Figs 22, 33, 58, 273–279 setae (Fig. 276). Scutum not dusted before Oscinis cornuta Fallén, 1820: 6. Lectotype in ZMLU scutellum. Male genitalia Figs 277–279 (E. cor- (Andersson, 1963). nuta speciescomplex)...... 5 Body black, shining. Frons yellow, ocellar triangle 5 Arista thin, legs darkened black, smooth and shining, extending 2/3 of frons...... E. cornuta form A Two pairs of orb longer than remaining ones. Genae – Arista varying in thickness, legs yellow . . . . . 6 yellow with silvery dust, nearly as wide as first flag- ellomere. Antennae yellow, first flagellomere black 6 Arista thickened, aristal pubescence shorter at aristal base. Arista variable, widened and thickly than diameter of arista . . . . E. cornuta form B pubescent. Palpi yellow. Scutum and scutellum – Arista variable in thickness but aristal pubes- shining, coarsely punctate, scutum with two lon- cence not shorter than aristal diameter. gitudinal dusted stripes along dorsocentral lines...... E. cornuta (Fallén) Apical and subapical setae of scutellum on small tubercles. Distance between apical setae as dis- 36. Elachiptera brevipennis tance between apical and subapical setae. Pleura (Meigen, 1830) smooth, shining except on upper part. npl 1 + 1. Se- Figs 2, 66, 270–272 tae of head and abdomen yellow. Abdomen black, shining. Legs yellow or more or less darkened. Body Chlorops brevipennis Meigen, 1830: 159; Meigen’s length 2.0–3.0 mm. plates211:12;237:4;245:13(Morge,1976b).Lec- totype in NHMW (Nartshuk, 1978a). Distribution. – Throughhout Fennoscandia and Denmark, reaching the coast of the Arctic Ocean. Ground colour varies from yellowish red to dark Pan-Palearctic species. reddish brown, abdomen being darker than tho- rax. Ocellar triangle shining, reaching front margin Biology. – Common species in diverse types of of frons. Arista thickened and pubescent, black. habitats. Hibernates as adults in birds’ nests, Li- Two pairs of orb longer than remaining ones. Genae para galls, under scales of coniferous cones, and as wide as first flagellomere. Antennae brown. between the leaves of Typha (Tschirnhaus, 1992; Scutum densely punctate. Scutellum black with 4 Krivokhatsky & Nartshuk, 2001). small tubercles at base of scutellar setae. Abdomen Larvae saprophytophagous, secondary invaders, black, shining. Syntergite 1 + 2ofabdomenlong, develop in rotting tissue of different grasses: Phrag-

92 Figs 269–279. Elachiptera brevipennis (Meigen): 269: head, dorsal view. 270: head, lateral view. 271: hypopygium, dorsal view. 272: hypan- drium and phallic complex. E. cornuta (Fallén): 273: head, dorsal view. 274: head, lateral view. 275: wing. 276: scutellum. 277: hypopygium, dorsal view. 278: hypopygium, lateral view view. 279: hypandrium and phallic complex.

93 mites, Agrostis, Dactylis, Calamagrostis, Deschamp- Elachiptera diastema Collin, 1946: 147. Syntypes 9 sia, Festuca, etc., and all species of cereals. Also males and 6 females in OXUM, 1 syntype in the found in other mono- and dicotyledon plants: de- Manchester Museum (Pont, 1995). caying leaves of Typha latifolia and T. angustifolia, Similar to E. cornuta, but differs from it the follow- Bolboschoenus maritimus, Scirpus, Schoenoplectes ing characters: wings shorter, dusted stripes on scu- taberhaemontani, Triglochin maritimus, Atriplex tum joined behind by a prescutellar dusted band, triangularis, A. prostrata, Gladiolus communis, scutellum shorter and more truncate, apical setae Solanum fruticosum, Oenanthe crocata, Cannabis much farther apart, distance between apical setae sativa (Kreiter, 1927, 1928; Goidanich, 1928; Jepson nearly twice the distance between apical and sub- & Southwood, 1960a; Vilkova, 1962; Tschirnhaus, apical setae. Setulae of head and abdomen dark. 1981, 1992). Rygg (1966) reared it from seedlings of Body length 2.0–2.5 mm. wheat damaged by Phorbia fumigata (Anthomyi- idae) in Norway, and Johansson (1960) from ce- Distribution. – Denmark: common and wide- real shoots in Sweden. Mayer (1949) recorded the spread. Sweden: north to NB along the Baltic Coast. species as a pest of tomatoes, and Show (1955) as a Norway:Ø,HEs.Finland:N,Ka,Ta,Ok,Ob,Ks.Rus- pest of gladioli. It has been reared from a Schoeno- sia: Vib, Kr. plectes stem and from a stem of Bolboschoenus mar- Euro-Mediterranean species. itimus infested by Bactra robustana (Lepidoptera) together with Eribolus danicus.Thespecieshas Biology. – AdultsinMayandmorecommoninAu- no economic importance in cereal fields (Walton, gust and September in wet habitats. 1937; Vilkova, 1962). Larvae phytosaprophagous, developing in shoots Larvae were described by Vimmer (1926, 1931), of grasses (Dactylis glomerata and Festuca arund- Kreiter (1928), and Balachowsky & Mesnil (1935). inaceae (Sharipov, 1981; Tschirnhaus, 1981)). Col- Jepson & Southwood (1960a) gave distinguishing lected on agricultural grasses in Denmark (Nielsen characters of eggs and first instar larvae from those & Nielsen, 1984). Kibric et al. (1999) listed the of Oscinella frit. species as tyrphophilous. Note. – The species is very variable in size, colour of legs and thickness of arista. We found two deviat- 39. Elachiptera scrobiculata ing forms of the present species: (1) Form A is found (Strobl, 1901) in northern Fennoscandia and differs from normal Figs 283, 284 E. cornuta in having a smaller body, black legs and rather thin arista. (2) Form B differs from normal Oscinisscrobiculata Strobl, 1901: 233. Syntypes male E. cornuta in being larger and having a very broad and female not found in NMBA or the Klagen- arista and bright yellow legs; it occurs in southern furt Museum (Chvála, 2008). Scandinavia especially in small ponds on Typha. Elachiptera tenuiseta Frey, 1947: 69. Holotype in The taxonomic status of these forms is unsettled. NHMH, but Nartshuk in 1990 found only the pin They either represent distinct taxa or two extremes and labels without fly. within one variable species. There are some differ- Body black, shining. Frons dark brown to black. ences in the male genitalia between these forms, Ocellar triangle black, shining, extending 3/4 of but some specimens are not always separable from frons. Two pairs of orb longer than remaining pairs. the typical form of E. cornuta. Their status is unclar- Face and genae dark brown, genae as wide as ified. Examination of specimens from other parts of first flagellomere. First flagellomere black. Arista the vast range of E. cornuta is clearly needed to find rather thickly pubescent, black. Palpi dark. Scu- out is this is a species or species complex. Besides, tum dusted, scutellum wider than long, trape- there exist many names proposed as species, sub- zoid, coarsely punctate. Apical setae wide apart, species and forms that are tentatively considered on small tubercles. Pleura shining. npl 1 + 1. Ab- as synonyms of E. cornuta. domen black. Legs partly black. Halteres yellow. Body length 1.7–2.0 mm. 38. Elachiptera diastema Distribution. – Sweden: north to LU and NB. Nor- Collin, 1946 way: AAy: Jåmåsknutene, Froland. Finland: north to Figs 280–282 LkE. Russia: Vib. Not so far found in Denmark.

94 Figs 280–287. Elachiptera diastema Collin: 280: head and thorax, dorsal view. 281: head, lateral view. 282: scutellum. E. scrobiculata (Strobl): 283: head, lateral view. 284: scutellum. E. tuberculifera (Corti): 285: head, dorsal view. 286: head, lateral view. 287: scutellum.

Trans-Palearctic boreal species, known from 40. Elachiptera tuberculifera British Isles to Far East of Russia. Everywhere, but (Corti, 1909) not common. Figs 285–287 Biology. – Adults from May to October, proba- Crassiseta tuberculifera Corti, 1909: 132. Types not bly hibernating like other species of the genus, in seen, possibly in the Genova Museum. wet habitats with Carex. Apparently, the present species was found by Bleton & Fieuzet (1943) in Body black. Frons yellow, ocellar triangle black, stems of sorghum damaged by Atherigona (Musci- shining, extending nearly to front margin of frons. dae) in Morocco. Genae yellow, postgenae black. Antennae yellow,

95 first flagellomere darkened on apex. Arista black, as first flagellomere (Fig. 296). Ocellar trian- broadly thickened with short marginal hairs. Tho- gle extending almost to front margin of frons rax black, scutum dusted, postpronota shining. (Fig. 295). Male genitalia Figs 299, 300 Scutellum a little longer than wide, flattened and ...... E. nana (Zetterstedt) weakly shagreened, dusted, with 3 pairs of setae on finger like tubercles. Legs yellow. Body length 2.0– – Frons entirely black or slightly brownish in 2.5 mm. front. Genae in front distinctly narrower than first flagellomere (Figs 289, 291, 292, 302) Distribution. – Denmark: NEZ. Sweden: north to LU ...... 2 and NB. Norway: Ø, AK, HEs, On, Bv, VE, AAy, Ry. 2 Ocellar triangle shining except ocellar tuber- Finland: north to Om. Russia: Vib, Kr, Lr. Trans-Palearctic, polyzonal species. cle...... E. gracilior (Meijere) – Ocellartriangleentirelydull...... 3 Biology. – Adults in wetlands, meadows and moist places in forests, very common, taken in traps from 3 Genae in front nearly as wide as front tibiae March to September. Hibernating adults found in (Fig. 292), first flagellomere slightly reddish- birds’ nests (Tschirnhaus, 1992; Krivokhatsky & yellow at base. All tibiae yellow at both ends. Nartshuk, 2001). Taken in a light trap in December Femora not enlarged. Palpi dark. Male geni- in Norway (Hågvar & Greve, 2003). talia Figs 293, 294 . . . . . E. hungaricus Becker Larvae phytosaprophagous, developing in in- – Genae wider, nearly half width of first flagel- sect borrows with decaying plant tissue and frass. Found as secondary invader of grasses and all ce- lomere. Femora enlarged, more distinctly so in reals (Vilkova, 1962), but also in stems of other males(Figs303,304)...... 4 monocots: Liliaceae, Iridaceae, Typhaceae. It has 4 Larger, 1.5–2 mm. In males first flagellomere been reared from stems of several cereals (wheat, yellow, darkened at aristal base, palpi yellow. barley, rye), mostly infested by Phorbia fumigata In females first flagellomere dark, but yellow (Anthomyiidae) in Norway and Sweden (Johans- at base, palpi brown. Femora moderately en- son, 1960; Rygg, 1966). larged in males. Male genitalia Figs 305, 306 ...... E. slesvicensis Becker Genus Eribolus Becker, 1910 – Smaller, 1.0 mm. In males first flagellomere dark, reddish-yellow at base, palpi brown and all femora greatly enlarged. In females first Eribolus Becker, 1910: 127. Type species: Eribolus sudeticus Becker, 1910 (= Oscinis nana Zetter- flagellomere and palpi entirely black, femora stedt, 1838) by designation of Enderlein, 1911: slightly enlarged ...... E. danicus sp. nov. 206. Black, dusted or shining species with flattened scu- 41. Eribolus danicus tum. Ocellar triangle usually dusted. Vibrissal angle Nartshuk & Andersson, sp. nov. rounded. First flagellomere squarish or rounded. Figs 288–290 poc upright and cruciate. Setulae on scutum ar- ranged in lines. Pleura mainly shining. npl 1 + 1. Holotype male and paratype female: Denmark: Legs sometimes with thickened femora, more dis- NEZ: Amager, bread from stem of Bolboschoenus tinct in males. Tibial organ oval. Femoral organ a maritimus infested by Bactra robustana (Lepidop- group of slightly differentiated short setulae. Male tera), collected 04.vi.1921, fly emerged 05.vii.1921 genitalia: cerci and surstyli simple, hypandrium (Gusmann). Other paratypes: Sweden: SK: Alnarp, closed behind, postgonites enlarged. 20.vi.1960 (H. Andersson), 1 female; Lomma, 16.viii. 1959 (H. Andersson), 1 female; Silvåkra, 26.v.1982 Distribution. – Holarctic and Neotropical Regions. (C. Hansson), 1 female. Holotype and 1 paratype in ZMUC, 3 paratypes in ZMLU. Key to species of Eribolus Becker Diffferential diagnosis. – Most similar to E. hungari- cus Becker, but distinguished by smaller size, wider 1 Frons narrow but distinctly reddish-yellow an- genae, more distinctly enlarged femora, especially teriorly. Genae wide, almost as wide in front in males.

96 Description. – Male, female. Small, black. Frons 43. Eribolus hungaricus shorter than wide, dull. Ocellar triangle extend- Becker, 1910 ing 2/3 length of frons, dull, behind nearly as wide Figs 292–294 as frons. Three rather long and thin orb on each side of frons. poc and vte stronger and little longer Eribolus hungaricus Becker, 1910: 128. Syntype fe- than orb. First flagellomere large and round, arista male in ZMHU (Wendt, 1975). shortly pubescent, nearly as long as frons. Genae in front a little narrower than a half width of first flag- Small, very flattened species with black dull body. ellomere, distinctly widened posteriorly. Eyes short Frons black, brownish in front. Ocellar triangle dull oval. extending 3/4 of frons. Genae narrower than first Scutum flat, dusted. Scutellum short, triangular. flagellomere. First flagellomere black, yellowish at Apical setae thin, shorter than scutellum. Pleura base. Palpi black. Scutum flat, dull. Scutellum short shining. npl 1 + 1. Wings little brownish, longer with apical setae short and wide apart, inserted than abdomen. Second sector of costal vein nearly near to subapical setae. Femora black, tibiae yellow 1.5 longer than third sector which is 1.25 times with black band. Body length 1.25 mm. longer than fourth sector. Cell br not widened. Distribution. – Denmark: SJ, WJ, F, NEZ. Sweden: Veins R4+5 and M divergent apically. Halteres yel- SK, SM, ÖG. Norway: HOi. Finland: Al, Ab, N, Ta. low. Femora of all legs strongly enlarged in male, Russia: Vib. distinctly less so in female. Femora black, tibiae European species. black or a little paler at each end, tarsi yellowish. Body length 1 mm. Biology. – Adults in June to July in wet habitats with Phragmites, recorded as host plant by Tschirnhaus (1981). Ismay (1980a) pointed out that this species 42. Eribolus gracilior is common in reed beds in saline dunes in England. (Meijere, 1918) Fig. 291 44. Eribolus nana Oscinella gracilior Meijere, 1918: 140. Lectotype in (Zetterstedt, 1838) Figs 295–300 ZMAN (Nartshuk, 2003). Small, very flattened species, entirely black. Frons Oscinis nana Zetterstedt, 1838: 782. Lectotype in dull, ocellar triangle shining, with square dusted ZMLU (Andersson, 1966). spot on ocellar tubercle, extending halfway to front Body black, dull, narrow and flattened. Frons red- margin of frons. Antennae black, first flagellomere dish in front. Ocellar triangle dull, extending 3/5 rounded. Genae a little narrower than first flagel- of frons. Genae wider than first flagellomere. First lomere. Scutum flat, scutum and scutellum thinly flagellomere black, rounded. Palpi black. Scutum dusted and subshining. Pleura shining except on + flattened, grey-dusted. Scutellum shorter than long upper part of anepisternum. npl 1 1. Femora with short apical and subapical setae. Pleura shin- black, front tibiae yellow, other tibiae yellow with ing below. Femora black except yellow at apices, black band, tarsi yellow. Wings narrow, longer than tibiae yellow with black band. Body length 1.5– abdomen. Veins R4+5 and M divergent apically. 2.0 mm. Body length 1.0 mm. Distribution. – The most abundant and widespread Distribution. – Sweden: SK, HA, ÖL, VG, and also species of Eribolus. Denmark: EJ, NWJ, F, NEZ. Swe- further north in NB. Finland: Ta: Forssa, 26.vi.1964 den: north to LU and NB. Finland: north to Ks. Rus- (M. Loikaaanen), Ok: Paltamo, 10.vii.1981, 30.vi.1991 sia: Vib, Kr. Not recorded yet from Norway. (Koponen). Not so far found in Denmark, Norway Holarctic species, widely distributed in the Pale- or Russian parts of Fennoscandia. arctic. Eurasian species. Biology. –Adultshibernateandmaybecaught Biology. – Adults in May to August in bogs and from February to October in different kinds of wet- other wetlands, often swept from edges of open wa- lands: bogs, fens, marshes, especially on reeds (Is- ter without Phragmites (Ismay, 1980a). may, 1980a).

97 Figs 288–294. Eribolus danicus Nartshuk & Andersson: 288: head, dorsal view. 289: head, lateral view. 290: wing. E. gracilior (Meijere): 291: head, lateral view. E. hungaricus Becker: 292: head, lateral view. 293: hypopygium, dorsal view. 294: hypandrium and phallic complex.

Larvae often several together in shoots and stems 45. Eribolus slesvicensis of Carex, Scirpus, Sparganium and probably other Becker, 1910 hygrophilous monocots with borrows by other in- Figs 301–306 sects, often Cordilura spp. (Scathophagidae) (Val- ley et al., 1969; Nartshuk, 1972a; Uffen & Chandler, Eribolus slesvicensis Becker, 1910: 128. Syntypes (1 1978; Valley & Foote, 1996). Johansson (1943) re- male, 1 female) in ZMHU (Wendt, 1975). ported it from cereals in Sweden, but this needs to Eribolus crassipes Nartshuk,1972c:910.Holotypein be confirmed. Valley et al. (1969) found 10 larvae in ZISP. Syn. n. astemofCarex injured by Cordilura varicornis Cur- Small flattened species with black dull body and ran in North America. enlarged femora, especially in male. Frons black,

98 Figs 295–306. Eribolus nana (Zetterstedt): 295: head, dorsal view. 296: head, lateral view. 297: wing. 298: scutellum. 299: hypopygium, dorsal view. 300: hypandrium and phallic complex. E. slesvicensis Becker: 301: head, dorsal view. 302: head, lateral view. 303: male legs. 304: female legs. 305: hypopygium, dorsal view. 306: hypopygium, lateral view.

99 Figs 307–309. Gampsocera numerata (Heeger): 307: head, dorsal view. 308: head, lateral view. 309: cerci and surstyli, dorsal view. slightly brownish in front. Ocellar triangle black, with small cerci and simple surstyli. Phallic com- dull, extending 3/4 of frons. Genae nearly as wide as plex fused. first flagellomere. Antennae mainly yellow, but first flagellomere darkened above. Palpi yellow. Scutum Distribution. – Palearctic, Oriental and Neotropical dull, but pleura shining below. Scutellum small, Regions. apical setae short, rather wide apart. Femora en- larged. Femora and hind tibiae black with yellow 46. Gampsocera numerata ends, front and middle tibiae mainly yellow. Body length 1.5–2.0 mm. (Heeger, 1858) Figs 3, 23, 38, 307–309 Distribution. – Rare species. Sweden: SK: Lands- krona, 17.vii, 1 male; Skälderviken, 05.viii.1955 Chlorops numerata Heeger, 1858: 302. Lectotype in (Ringdahl), 2 males. Finland: Al: Sund (R. Frey); NHMW (Nartshuk, 1997b). Föglö, vii.1939 (A. Nordman); N: Tvärminne; Par- Frons, face and genae yellow. Frons broader than voo. Not found in Denmark, Norway or Russian long. Setae and setulae black. Ocellar triangle black, parts of Fennoscandia. shining, extending half of frons, yellow behind and Eurasian species, eastward to Mongolia. brown in anterior part. Antennae and palpi yel- Biology. – Adults in June and August in wetlands, low. Scapus very short. First flagellomere prolonged associated with reed beds (Phragmites australis), dorsodistally; arista short and black, apically tick- especially at lakes and sea shores. Tschirnhaus ened and densely haired. Genae with silvery dust, (1981) listed as host plants Carex and Bolboschoenus narrower in front than behind, with rounded vib- maritimus. Ismay (1980a) found the species in rissal angle and long pale vibrissa. Thorax yel- saline habitats in England. low with scutum black, shining, except on narrow stripes along notopleura. Postpronota with black spot. Scutellum black or brown, wider than long; apical setae black, as long as scutellum. Pleura Genus Gampsocera Schiner, 1862 shining, yellow with black marks or nearly entirely black. Abdomen brownish black. Legs mainly black Gampsocera Schiner, 1862: 431. Type species: Chlo- or with indistinct black marks. Wings hyaline in fe- rops numerata Heeger, 1858 by monotypy. male and with black marks in male. Body length 2.0–2.5 mm. Black species with obliquely prolonged first flagel- lomere and apically thickened arista. Ocellar trian- Distribution. – Rare. Sweden: UP: Uppsala, 27.x.1975 gle shining. poc upright, cruciate. Male wings with (L. Hedström), 1 female. Norway: AK, AAy. Finland: black marks. Tibial organ distinct, oval. Femoral or- Ta: Tavastia (Palmén), 1 male. Not found in Den- gan absent. Male genitalia: epandrium flattened, mark or Russian parts of Fennoscandia.

100 Eurasian species, known from Europe to Yakutia, (Figs 310, 311). Body mainly, sometimes en- East Siberia, but scarce everywhere. tirelyblack...... G. borealis (Duda) Biology. – Adults occur in forests. Larvae probably – Male genitalia: cerci shorter and wider apart, myxo-mycetophagous as once found under plas- surstyli shorter (Figs 315, 316). Colour of body modium of a slime mold. Heeger (1858) found lar- varied, but thorax normally yellow with black vae in a stem of Alcea rosea damaged by Apion stripes fused on anterior part of scutum, scu- curvirostris (Brentidae). He observed a pair in cop- tellum normally yellow, pleura with four black ula and a female laying eggs in the decaying stem spots...... G. dubius (Macquart) with excrements of Apion. 5 Apical part of wings slightly darkened espe- cially in male, best seen in fresh specimens. Male genitalia Figs 339, 340 Genus Gaurax Loew, 1863 ...... G. maculipennis (Zetterstedt) – Wings without darkening. Male genitalia dif- Gaurax Loew, 1863: 35. Type species: Gaurax fes- ferent...... 6 tivus Loew, 1863 by monotypy. 6 Ocellar triangle yellow, shining, only on ocel- Species black or yellow with black markings. Ocel- lar tubercle black. Hind tibiae with dark nar- lar triangle shining. Eyes usually distinctly haired. row band at base. Usually one dark spot on Genae narrow, usually with silvery dust. poc up- pleura, on anepisternum, but some females right, cruciate. First flagellomere oval or reniform. have 3 or 4 dark spots. Male genitalia: surstyli Arista usually densely pubescent, but not thick- with narrow process directed backwards (Figs ened. Tibial organ distinct, oval. Femoral organ ab- 326–328)...... G. fascipes Becker sent. Male genitalia: cerci usually long, surstyli of complex structure with long setae, phallic complex – Ocellar triangle mainly or entirely black, shin- fused. ing or dusted. Usually 4 dark spots on pleura, Species of Gaurax are very variable in body and or thorax entirely black. All legs entirely yel- size, and only examination of the male genitalia en- low, rarely tibial organ on hind tibiae sur- sures correct identification. roundedbydark,orentirelydark...... 7 Distribution. –Worldwide. 7 Body mainly black except reddish yellow at front margin of frons and on genae, or also scutellum, notopleura and hind part of post- Key to species of Gaurax Loew pronotayellow...... 8 – Ventral side of thorax yellow with black spots. 1 First flagellomere black with white pubes- Legsyellow...... 9 cence; arista white, slightly thickened. Male genitalia Figs 331–333 8 Body black except reddish yellow at front mar- ...... G. leucarista Nartshuk gin of frons and on genae. All legs darkened. – Antennaedifferent...... 2 Male genitalia Fig. 341 . . . . G. suecicus sp. nov. 2 First flagellomere rounded, nearly as wide as – Thorax black, with scutellum, a large spot cov- long(Figs313,334)...... 3 ering notopleura, hind part of postpronota – First flagellomere reniform, wider than long and upper part of anepisternum yellow. Legs (Fig.338)...... 5 yellow with black mark on all femora and mid and hind tibiae . . . . G. flavomaculatus (Duda) 3 First flagellomere large, nearly as wide as height of face (Fig. 334). Male genitalia Figs 9 Male genitalia: cerci tapering and parallel, 335–337...... G. macrocerus (Nartshuk) closetoeachother,surstyliwithacuteprocess directed medially ...... G. fungivorus sp. nov. – First flagellomere smaller, narrower than heightofface(Fig.313)...... 4 – Male genitalia: broader and wider apart, surstyli with some processes on lower margin 4 Male genitalia: cerci narrow, rather long; (Figs317–320)....G. ephippium (Zetterstedt) surstyli strongly elongated, widened apically

101 Note. – Flies of the genus Gaurax are not numerous 48. Gaurax dubius in the collections. Species identification is difficult (Macquart, 1835) because of considerable variation in colour within Figs 312–316, 971–973 species and between sexes. Therefore examination of the male genitalia is essential for reliable identi- Oscinis dubius Macquart, 1835: 604. Type probably fication. Additional species, including undescribed in MHNL. ones, may well occur in the investigated area. Body yellow with three black stripes on scutum and four black spots on pleura. Frons as long as 47. Gaurax borealis wide. Ocellar triangle short, extending half length (Duda, 1933) of frons, yellow with black spot on ocellar tuber- Figs 310, 311 cle, shining. First flagellomere nearly as wide as long, yellow or more or less darkened above. Genae Botanobia borealis Duda,1933:80,asvar.ofB. du- narrow with silvery dusting. Male genitalia: cerci bia (Macquart). Syntypes (1 male, 2 females) in rather short and wide apart, surstyli short. Body NHMH. length 1.3–1.5 mm. Usually scutum and scutellum black, covered with Distribution. – Sweden: SK, SM; SÖ: Tyresta Na- pale setulae, postpronota and notopleura yellow, ture Reserve, 11.vi–17.vii, 28.vii–15.viii.1999, 04.viii– but postpronota with black spot. Pleura yellow 24.ix.2000, in Malaise traps (Viklund, Wikars & with 4 large black spots, but females from north- Ahnlund), 1 male, 4 females; probably also LU: Mes- ern localities entirely black. Head yellow with black saure, but only females available. Norway: AK, HEs occiput. Frons as long as wide. Ocellar triangle (Komonen et al., 2004). Finland: Al, Ab, N, Li. Rus- short, extending half length of frons, black, shining. sia: Vib, Kr. Not found in Denmark and Norway. First flagellomere nearly as wide as long, more or Holarctic species. Occurs in forests. less darkened. Genae narrow with silvery dusting. Palpi yellow. Setae of head and thorax black. Ab- Biology. – Larvae primary fungivorous, develop in domen darkened dorsally. Legs yellow. Halteres yel- bracket fungi Piptoporus betulinus and Fomitopsis low. Male genitalia: cerci narrow and long, surstyli pinicola (Smith, 1964; Pielou, 1966; Pielou & Pielou, elongated. Body length 1.3–1.5 mm. 1968;Pielou&Verma,1968;Økland&Hågvar,1994; Distribution. – Rare. Sweden: SÖ: Tyresta Nature Komonen et al., 2004). Smith (1964) noted that Reserve, 28.v–15.vii.1999 (B. Viklund), 26.v– flies were reared from old dry bracket fungi. Adults 27.vii.2000 (Viklund, Wikars & Ahnlund), 1 male, also visit other fungi, Coriolus and Bjerkanderia (Uf- 1 female, Malaise traps. Finland: Ab, Ok, Ks, LkE. fen & Chandler, 1978). Larvae and puparia were Russia: Vib: Jukki, 12.vi.1932 (Stackelberg), 1 female; also found under bark of coniferous trees infested Kr: Paanajärvi, females? (specimens darkened), re- by the bark beetle Pityogenes chalcographus (Bog- corded by Frey (1933) as Conioscinella gallarum. danova, 1975; Nartshuk, 1987) and in cones of Picea Not found in Denmark and Norway. abies (Saksons, 1973; Skrzypczynska, 1982), but may be the latter records may concern G. ephippium. Note. – Described as a var. of dubia Macquart Bogdanova (1975) considered the species as ento- from Finland, but Duda (1933) did not specify mophagous. exact localities and number of specimens. Frey Larvae described by Smith (1964), some larval (1933), who published details about the Finnish structures illustrated by Nartshuk (1987). Chloropidae identified by O. Duda, listed the fol- lowing localities: Ab: Lojo (Krogerus); OK: Kajana (Wuoretanus); Ks: Kuusamo, Paanajärvi (Frey); 49. Gaurax ephippium LKem: Muonio (Palmén). We found in NHMH: 1 fe- male, Kajana (Wuoretanus), type No 8150; 1 female, (Zetterstedt, 1848), sp. rev. Figs 317–320 Lapponia (Palmén), type No 8151; and 1 male, Ku- usamo (Frey), type No 8149. The last specimen is Oscinis ephippium Zetterstedt, 1848: 2664. Holo- presently designated as lectotype. type in ZMLU (Andersson, 1966). Restored from Eurasian species, from Europe to Mongolia. synonymy with Gaurax dubius (Macquart, Biology. –Adultsinforests. 1835).

102 Figs 310–320. Gauraxborealis (Zetterstedt): 310: hypopygium, dorsal view.311: hypopygium, lateral view. G. dubius (Macquart): 312: head, dorsal view. 313: head, lateral view. 314: antenna. 315: hypopygium, lateral view. 316: hypopygium, dorsal view. G. ephippium (Zetterstedt): 317: hypopygium, dorsal view. 318: surstylus, medial side. 319: hypopygium, lateral view. 320: male cercus, dorsal view.

103 Gaurax strobilum Karps, 1981: 187. Holotype and 1 wide. Ocellar triangle short, extending half length paratype in ZISP, other paratypes in Institute of of frons, yellow shining, black spot only on ocel- Biology in Salaspils, Latvia. Syn. n. lar tubercle. First flagellomere reniform, wider than long. Genae narrow with silvery dusting. Posterior Body yellow with three black stripes on scutum and tibiae with small black ring in basal half. Male gen- four black spots on pleura. Scutal stripes fused in italia: surstyli with rather long narrow process and anterior part of scutum, shining, covered with pale incision in low angle. Body length 1.3–1.5 mm. setulae. Scutellum yellow, apical setae longer than scutellum. Frons as long as wide. Ocellar triangle Distribution. – Sweden:SK,ÖG,NÄ,SÖ,VS,HS. short, extending half length of frons, yellow with Norway: AK: Lyndøya, Oslo. Finland: N: Tvärminne black spot on ocellar tubercle, shining. First flagel- (R. Frey). Russia: Vib: Vyborg. Not found in Den- lomere reniform, wider than long, yellow or more mark. or less darkened above. Genae narrow with silvery Euro-Caucasian species. dusting. Palpi yellow. Male genitalia: cerci rather short and wide, surstyli short with patch of tick se- Biology. – Adults in woodland. tae on inner surface. Body length 1.3–1.5 mm. Larvae develop in bracket fungi (Piptoporus be- tulinus), rotting wood and cones of Larix and Picea Distribution. – Sweden:SK,ÖG,VG,BO,SÖ,NB. (Smith, 1964, 1967; Milišauskas, 1976; Skrzyp- Finland:Al:Saltvik(R.Frey);N:Ruotsinkylä,20– czyńska, 1978, 1982; Grebenshchikova & Naumov, 22.vi.1942 (Tahvonen), from cones of Picea,28 1979). It has also been reared from crown gall dis- specimens. Not found in Denmark, Norway or Rus- ease caused by Agrobacterium tumefacies (Tschirn- sian parts of Fennoscandia. haus, 1992). Collin (1939) recorded that it has been European species. bred from nests of blackbird and linnet in England and also found it in windows of his house. Biology. – Adults in June to July in woodland. It has Larvae described by Smith (1967). Some larval been reared from cones of Larix decidua and Picea structures illustrated by Nartshuk (1987). abies and from a twig of Pinus silvestris (Karps, 1981; Nordlander & Grijpma, 1991; and specimens in col- lections seen by us). 51. Gaurax flavomaculatus Note. – Oscinis ephippium Zetterstedt has for long (Duda, 1933) been considered as a synonym of Oscinis dubius Figs 329, 330 Macquart, 1835. Our re-examination of the male holotype in Coll. Zetterstedt in ZMLU revealed that Goniopsita flavomaculata Duda, 1933: 107. Holo- Zetterstedt’s species is not identical with the cur- type in ZMHU. rent interpretation of Gaurax dubius (Smith, 1964; Gaurax britannicus Deeming, 1980: 93. Holotype in Nartshuk, 1970). First flagellomere of G. ephippium BMNH. is reniform, not round as in G. dubius,andthemale Body black, shining. Head yellow, black on occiput, genitalia also differ, but coincide with those of G. ocellar triangle black. Eyes uniformly covered with strobilum Karps. short hairs. Antennae and clypeus black. Face and frons with exception of ocellar triangle and black 50. Gaurax fascipes parts with silvery sheen. Thorax black except yel- low on scutellum, notopleura, hind part of post- Becker, 1910 pronota and upper part of anepisternum. Legs yel- Figs 321–328 low with apical part of all femora and basal part of mid and hind tibia infuscate. Wings hyaline. Hal- Gaurax fascipes Becker, 1910: 116. Syntypes in tere yellow. Body length about 2 mm. HNHM and probably in ZMHU. Distribution. – Only Norway: AAy: Lillesand, S- Body yellow with three black partly fused stripes facing slope, viii–ix.2004, Malaise trap, 5 females. on scutum and one black spot on pleura. Some- European species. times stripes fused only in anterior part of scutum. Central stripe not reaching scutellum, but a black- Biology. – Adults in broad-leaf forest. The larvae ish spot may be present before scutellum. Scutum live under bark of trees, probably feeding on frass or around hind dc usually yellow. Frons as long as dead insects. In East Europe some specimens were

104 Figs 321–333. Gaurax fascipes (Meigen): 321: head, dorsal view. 322: head, lateral view. 323: thorax, dorsal view. 324: wing. 325: scutel- lum. 326: hypopygium, dorsal view. 327: hypopygium, lateral view. 328: hypandrium and phallic complex. G. flavomaculatus Duda: 329: head, dorsal view. 330: localities in Europe. G. leucarista Nartshuk: 331: hypopygium, lateral view. 332: hypopygium, dorsal view. 333: hypandrium and phallic complex.

105 reared from Quercus robur and Armeniaca vulgaris 53. Gaurax leucarista (Nartshuk, 2008). Nartshuk, 1962 Figs 332, 333 52. Gaurax fungivorus Gaurax leucarista Nartshuk, 1962a: 255. Holotype Nartshuk & Andersson, sp. nov. in ZISP. Holotype, male: Norway: HOy: Raudli, Os, 20.vi– Body black, shining. Frons square, with brown silky 16.vii.1991 ex Peptoporus betulinus (K.H. Thunes). hairs. orb 5, rather long. Ocellar triangle extend- Paratypes: 3 males and 1 female pinned, 18 males ing 2/3 length of frons, black, shining. Eyes bare. and 16 females in alcohol, same data as holotype Genae brown, in front half as wide as first flag- except 20.vi–23.vii.1991, 1 male. Sweden: SÖ: 5 km ellomere. Antennae black, first flagellomere with WSW Sköldinge, Flen, 10.iv.1967, ex Ganoderma ap- white pubescence. Arista densely white pubescent planum (I.Nordin),1male,2females;UP:Vallen- and seems to be thickened. Scutum and pleura tuna, 31.viii.1958 (K.J. Hedquist), 1 male. Holotype, shining, scutellum dusted. npl 1 + 1. Legs yellow all Swedish paratypes and 11 male and 8 female pa- with slight darkening on femora and hind tibia. ratypes from Norway in ZMLU; 2 paratypes from Halteres white. Body length 1.5 mm. Norway in ZISP; 16 paratypes in alcohol in ZMHU. Distribution. – Only known from Russia: Kr: Kon- Diagnosis. – Male genitalia with short surstyli sim- dopozhsky District: Kivach Nature Reserve, 1 male, ilar to G. macrocerus and G. maculipennis,butG. 18–20.vi.1989, Malaise trap in Populus tremula for- fungivorus differs from the former by a smaller first est (Polevoi). flagellomere and longer and narrower male cerci, Rare European species described from south- and from the latter by having clear wings and a ern part of Leningrad oblast, where it was caught small pointed process on surstyli. on young Populus tremula, and further found in Moscow Oblast and Karelia. Description. – Male, female. Head yellow with black occiput. Frons little longer than wide. Ocellar triangle black, thinly dusted, short, extending half 54. Gaurax macrocerus length of frons. Setae and setulae of head yellow or (Nartshuk, 1962) black. First flagellomere reniform, wider than long, Figs 334–337 entirely yellow or darkened on outer margin. Arista black with pale base. Eyes haired. Genae in front Botanobia macrocera Nartshuk, 1962a: 253. Holo- equal to width of first flagellomere, widened be- type in ZISP. hind. Palpi yellow. Scutum shining black, only narrow lateral mar- Headyellow.Fronsaslongaswide.Ocellartrian- gins yellow, evenly covered with rather long pale gle black, shining, extending half length of frons. setulae. Postpronota yellow with black spot. Scutel- Genae narrow with silvery dust. Antennae yellow, lum black, or in females yellow or with black sides first flagellomere in male large, nearly half vertical basally. In females scutal stripes not fused behind diameter of eye. Arista black. Scutum black covered leaving yellow spots around hind dc.Pleurayel- with pale dense setulae. Postpronota, notopleura + and scutellum yellow or scutellum black. Pleura low with four black, shining spots. npl 1 1. Setae + of thorax black. Wings not coloured. Halteres yel- yellow with four black spots. npl 1 1, a second low. Legs yellow, posterior tibiae darkened around smaller upper hind seta sometimes present. Legs sensory spot. Abdomen black dorsally, epandrium yellow. Abdomen black. Epandrium black with yel- and female cerci black. Male genitalia: cerci rather low surstyli. Body length 1.3–1.5 mm. long, surstyli short with acute process directed to Distribution. –Rarespecies.Sweden:HA,VG,SÖ. the middle and numerous setae on inner surface. Finland:Al:Saltvik,1male;Oa:Bergo,17–28.vi.1946 Body length 1.7–2.0 mm. (Hellén), 1 male; Ks: Kuusamo, 1 male. Not found in Note. – Larval development of the present new Denmark, Norway or Russian parts of Fennoscan- species takes place in bracket fungi. It is thus pos- dia, but described from locality 130 km south from sible, that existing records of other Gaurax species St. Petersburg. from bracket fungi refer to the present species, as European species. species of Gaurax are difficult to identify. Biology. – Adults in woodland.

106 Figs 334–341. Gaurax macrocerus (Nartshuk): 334: head, lateral view. 335: hypopygium, dorsal view. 336: hypopygium, lateral view. 337: hypandrium and phallic complex. G. maculipennis (Zetterstedt): 338: antenna. 339: hypopygium, dorsal view. 340: apex of abdomen, lateral view. G. suecicus Nartshuk & Andersson: 341: hypopygium, lateral view.

55. Gaurax maculipennis Distribution. – Sweden: SK (type area); SÖ: Svin- (Zetterstedt, 1848) mossen, Katrineholm, 12.iv.1966 (O. Elofson), 1 Figs 338–340 male, 3 females, reared from Polyporus fumetaria on Betula. Finland: Al, Ab, N, Ta, Kb. Not found in Denmark, Norway or Russian parts of Fennoscan- Oscinis maculipennis Zetterstedt, 1848: 2665. Holo- dia. type in ZMLU (Andersson, 1966). European boreal species. Head yellow with black occiput. Frons as long as Biology. – Adults in woodlands, was reared in Fin- wide. Ocellar triangle short, extending half length land (Lemland) from tree fungus (swamp, Corylus) of frons, yellow with black spot on ocellar tuber- (specimens in collection) and from Fomitopsis pini- cle, dusted. First flagellomere reniform, wider than cola (Komonen et al., 2004). long, yellow. Genae narrow with silvery dusting. Scutum with three black fused stripes. Postpronota 56. Gaurax suecicus and spots around hind dc yellow, postpronota with Nartshuk & Andersson, sp. nov. black spot. Scutellum blackish. Wings of male with Fig. 341 darkening in apical part, well seen in fresh speci- mens, but nearly disappeared in specimens in col- Holotype, male, Sweden: Lule Lappmark: Gällivare, lection. Legs yellow. Body length 1.3–1.5 mm. 13.vii.1955 (H. Andersson). Holotype in ZMLU.

107 Figs 342–345. Hapleginella laevifrons (Loew): 342: head, dorsal view. 343: head, lateral view. 344: hypopygium, dorsal view. 345: hypan- drium and phallic complex.

Diagnosis. – On entirely black thorax similar to G. Distribution. – Only known from the holotype from niger Czerny, but differs by partly yellow head and Lule Lappmark in Sweden. bare eyes. Genus Hapleginella Duda, 1933 Description. – Male. Head black with frons in front and genae reddish yellow. Ocellar triangle black, Hapleginella Duda, 1933: 77. Type species: Oscinis shining. Eyes bare. First flagellomere small, wider laevifrons Loew, 1858 by original designation. than long, reddish yellow, darkened above. Arista Black, shining species. Ocellar triangle large shin- black with short pubescence. Setulae on genae ing with a row of setulae on its surface along side pale, head setae brown. Palpi reddish yellow with margins. Frons faintly shining. Vibrissal angle right. pale setulae. poc upright, cruciate. First flagellomere rounded. Thorax all black, scutum shining, covered with Scutum evenly covered with very short setulae. Tib- long pale setulae, distinctly parted along acros- ial organ oval. Femoral organ consists of two rows tichal lines. Scutellum dusted, black in posterior of warts. Male genitalia: cerci and surstyli simple, half. Apical scutellar setae longer than scutellum, hypandrium open, phallapodemic sclerite bowl- two pairs of short subapical setae. Dorsal setae shaped. and setulae of thorax brown, ventral setulae pale. Distribution. – Holarctic. Pleura and postnotum black, shining. Legs slen- der, blackish. Wings not coloured, costal sectors as 57. Hapleginella laevifrons 55:60:40:20. R2+3 slightly procurved. R4+5 and M almost parallel. Halteres yellow. Abdomen black, (Loew, 1858) Figs 18, 36, 53, 342–345, 967–970 very thinly dusted. Epandrium large, cerci wide apart, surstyli with process directed behind. Body Oscinis laevifrons Loew, 1858: 70. Type material not length 1.5 mm. found ZMHU (Wendt, 1975, 1978).

108 Oscinella inquilina Hendel, 1933: 47. Lectotype in Black with densely or thinly dusted scutum. Frons NHMW (Nartshuk, 1997b). dull, ocellar triangle smooth, dusted. Front mar- gin of frons convex. Vibrissal angle right. poc up- Body black, shining. Ocellar triangle smooth shin- right, cruciate. First flagellomere rounded, wider ing with a row of setulae on its surface along side than long. Scutum covered with setulae arranged margins, reaching margin of frons. Frons faintly in distinct rows. Pleura mainly shining. npl 1 + 1 shining. First flagellomere yellow, strongly dark- or 1 + 2. Tibial organ oval. Femoral organ consists ened above. Genae half as wide as first flagellom- of two rows of warts with short, stout setulae. Male ere, yellowish with white dust. Palpi yellow. Scutum genitalia: cerci short, surstyli rather broad. Hypan- shining evenly covered with very short setulae. npl + drium closed, rarely open. 1 2. Scutellum shorter than wide at base, dusted. The species Incertella scotica, I. zuercheri and I. Apical scutellar setae longer than scutellum. Legs nigrifrons are quite similar to Oscinella and their yellow or femora and hind tibiae darkened. Body generic position is not clear. We leave them for the length 1.5–1.7 mm. moment in Incertella. Further study is much re- Distribution. – Sweden: north to LU. Norway: AK, quired to clarify the status and limits of the genus- HEs. Finland: north to Li. Russia: Vib, Kr, Lr. Not group taxa Incertella, Microcercis, Cyclocercula,and found in Denmark. Oscinella and probably Rhopalopterum. Trans-Palearctic boreal species known from the Distribution. – Holarctic. British Isles to Far East Russia. Biology. – Adults from March to July in woodlands. Key to species of Incertella Sabrosky The larvae develop in cones of different conif- erous trees (Pinus, Picea, Abies, Larix)infested 1 Frons anteriorly and genae yellow or yellow- by pyralid and tortricid moths or weevils: Dio- ish, yellow part on frons sometimes very nar- ryctria abietella, Retinia perangustana, Zeiraphera row. Body with rather thick grey dusting . . . 2 griseana, Clepsis retiferana, Pissodes validirostris (Hendel, 1933; Gaidene & Nartshuk, 1963; Stad- – Frons and genae entirely dark. Body with nitsky, 1969; Saksons, 1973; Gaidene, 1976; Mili- ratherthindusting,subshining...... 4 šauskas, 1976; Skrzypczyńka, 1978, 1987; Nordlan- 2 First flagellomere yellow, darkened only at der & Grijpma, 1991). Larvae can develop in other aristal base. Genae narrow, not wider than parts of coniferous trees. Nartshuk examined spec- front tibiae (Fig. 352). Front and middle tib- imens reared from rotting top buds of young seed- iaemainlyyellow...... I. antennata (Collin) lings of Pinus sibirica cultivated on the Karelian Isthmus. The larvae are saprophagous scavengers – Firstflagellomereentirelyblack...... 3 feeding on excrements, dead insects or rotting 3 Anterior half of frons and palpi yellow. Genae plant tissues in borrows of primary plant feeders wider than front tibiae (Fig. 347). All tibiae and do not cause primary damage of coniferous yellow at both ends. Male genitalia Figs 350, cones and seeds. Hibernates as larvae or/and pupae 351...... I. albipalpis (Meigen) (Gaidene & Nartshuk, 1963; Stadnizky, 1969; Kaidis & Georgevits, 1972; Skrzypczinska, 1974, 1978, 1986; – Frons very narrow yellow anteriorly, some- Rognes, 1983; Skrzypczinska et al., 1987; Nordlan- times nearly entirely black, palpi black. Genae der & Grijpma, 1991). A note by Duda (1932–1933) not wider than front tibiae (Fig. 360). Male that it has been reared from galls of Andricus termi- genitaliaFig.351...... I. kerteszi (Becker) nalis (Cynipidae) is questionable in the opinion of 4Wingcellbr widened in the middle. Ocellar Tschirnhaus (1992). triangle reaching front margin of frons. Male Larvae were described by Gaidene & Nartshuk genitaliaFig.367...... I. zuercheri (Duda) (1963). –Wingcellbr not widened in the middle. Ocel- lar triangle reaching front margin of frons or Genus Incertella Sabrosky, 1980 only half length of frons (Figs 353, 362) . . . . 5 Incertella Sabrosky, 1980: 420. Type species: Os- 5 Ocellar triangle reaching front margin of frons, cinella incertella Becker, 1912 by original desig- sometimes with shining spots near ocelli nation. (Fig. 362). Anterior notopleural seta weaker

109 than posterior. Male genitalia Figs 365, 364 (1966) reared it from seedlings of oats and wheat in ...... I. scotica (Collin) Norway. Kallio (1950) listed it as a pest of cereals in – Ocellar triangle extending half length of frons Finland. It has also been reared from a disposal tip and compost (Werner, 1997), no doubt accidentally (Fig.353)...... 6 with its host plants. 6 Wings darkened (Figs 356, 357). Male geni- Some larval structures illustrated by Kreiter talia Fig. 358 (1928)...... I. karteshensis Nartshuk & Przhiboro – Wings clear or slightly greyish 59. Incertella antennata ...... I. nigrifrons (Duda) (Collin, 1946) Fig. 352 58. Incertella albipalpis Tropidoscinis antennata Collin, 1946: 136. Syntypes (Meigen, 1830) 6 males and 3 females in OXUM (Pont, 1995). Figs 54, 346–351 Body black, grey-dusted. Frons in front, face and Chlorops albipalpis Meigen, 1830: 163; Meigen’s genae yellow. Ocellar triangle reaching middle of plate 211: 10 (Morge, 1976b). Lectotype in frons, dusted. Genae narrow, whitish yellow near NHMW (Nartshuk, 1997a). eye and dark along ventral margin with a single row of dark setulae. Palpi yellow. oc, orb and vti equal Body black, grey-dusted. Frons entirely yellow, oc- in length, vte a little longer. Antennae yellow with ciput and postgenae black grey-dusted. Ocellar tri- first flagellomere darkened especially on outer side. angle black densely grey-dusted, extending half Setulae on scutum arranged more or less in lines. length of frons. Antennae entirely black, first flag- Pleura shining, except on upper part of anepister- ellomere large and rounded. Genae pale yellow, num. All setae and setulae black. Wings slightly slightly narrower than first flagellomere, postgenae brownish below costa and at base. Halteres yellow. black. Palpi yellow. Scutum grey-dusted, setulae ar- Femora black, hind tibiae yellow with black band, ranged in line. Scutellum semicircular, apical setae other tibiae and tarsi except dark last segments yel- a little longer than scutellum, wide apart. Subapi- low. Body length 1.0–1.25 mm. cal setae nearly 1/2 as long as apical. Pleura mainly shining, dusted on upper part of anepisternum. npl Distribution. – Denmark: SJ: Sottrupskov, 1 + 1. Legs with all femora dark except both ends, 12.viii.1896 (W.Wüstnei), 1 female; NEZ: Sortemose, hind tibiae with dark band, other tibiae yellow or Ganløse Orned, 16.vi.1961 (Exp. Zool. Mus.), 1 fe- with dark band. Body length 1.5–1.8 mm. male.Sweden:SK,SM,VR.NotfoundinNorway, Finland or Russian parts of Fennoscandia. Distribution. – Denmark: common and widespread. European species. Sweden: throughout the country. Norway: Ø, AK, HEs, Bv, VE, VAy, TRy, TRi. Finland: north to Lk. Rus- Biology. – Not common, adults in June to August in sia: Vib, Kr, Lr. meadows. Trans-Palearctic polyzonal species, known from the British Isles to Japan. 60. Incertella karteshensis Biology. – Common species, adults from the end of Nartshuk & Przhiboro, 2009 May to October in diverse types of meadows, edges Figs 353–359 of forest, crop fields. Incertella kartashensis Nartshuk & Przhiboro, Larvae phytophagous or phytosaprophagous, de- 2009: 5. Types in ZISP. velop in shoots of diverse grasses and cereals (wheat, ray, barley, oats) often damaged by other Body black. Frons entirely black, anterior mar- insects (Kreiter, 1928). As host plants are listed gin straigth. Ocellar triangle 2/3 as long as frons, Lolium perenne, Poa spp., Phleum pratense, Festuca densely dusted and not well limited from frons pratense, F. rubra, Bromus inermis, Anthoxanthum surface. Eyes with sparse fine hairs. Genae nar- odoratum, Elytrigia repens, Calamagrostis epigeios row. Antennae entirely black, first flagellomere and Dactylis glomerata (Karpova, 1965; Tschirn- small, wider than long. Arista rather long, short haus, 1981; Nielsen, 1994). Found on agricultural pubescent. Palpi black. Scutum with dense irregu- grasses in Denmark (Nielsen & Nielsen, 1984). Rygg lar rows of tiny setae. Scutellum small semicircular.

110 Figs 346–352. Incertella albipalpis (Meigen): 346: head, dorsal view. 347: head, lateral view. 348: scutellum. 349: wing. 350: hypopygium, dorsal view. 351: hypandrium and phallic complex. I. antennata (Collin): 352: head, lateral view.

Pleura dusted above and shining below. npl 1 + 1. dusted, extending half length of frons. Antennae Abdomen black, shining. Wings longer than ab- entirely black, first flagellomere rounded. Genae domen, intensively darkened except for a lighter brownish, as wide as 1/2 of first flagellomere. Palpi area between veins M and CuA1.Cellbr not wid- black. Scutum grey-dusted, setulae arranged in ened. All femora, front and middle tibiae dark line. Scutellum as long as 1/2 basal width, rounded. brown to black, hind tibiae yellow with black band Apical scutellar setae little longer than scutellum. in the middle. Body length 1.8–2.2 mm. Subapical setae nearly 1/3 of apical. Pleura mainly shining, dusted on upper part of anepisternum. npl Distribution. – Found only in Russia: Kr: in the up- 1 + 1. Legs mainly black with tarsi yellowish. Body per, intertidal zone of the White Sea. length 1.5–1.8 mm. Distribution. – Denmark: WJ, NEJ, F, LFM, NEZ, B. 61. Incertella kerteszi Sweden: north to ÅS. Norway: HEs: Abborhøgda, (Becker, 1910) Kongsvinger. Finland: north to Ok. Russia: Vib, Kr, Figs 360, 361 Lr. Trans-Palearctic boreal species, known from the Oscinellakerteszi Becker, 1910: 163. Types in HNHM. British Isles to Kamchatka. Body black, grey-dusted. Frons black, narrowly red- Biology. – Adults from end of May to early October dish in front. Ocellar triangle black densely grey- in wet meadows, edges of forests.

111 Figs 353–361. Incertella kartashensis Nartshuk & Przhiboro: 353: head, dorsal view. 354: 1st flagellomere and arista. 355: fore, middle and hind legs. 356: male wing. 357: female wing. 358: hypopygium, dorsal view. 359: tip of ovipositor, dorsal. I. kerteszi (Becker):360:head, lateral view. 361: cerci and surstylus.

Larvae develop in shoots of Deschampsia caespi- as wide as first flagellomere. Antennae entirely tosa (Nartshuk, 1972a). black, first flagellomere small. Arista very short pubescent. Palpi black. Scutum thinly dusted and 62. Incertella nigrifrons subshining. Scutellum semicircular. Apical setae wide apart. Pleura dusted above and shining be- (Duda, 1933) low. npl 1 + 1. Abdomen black, shining. Wings Conioscinella nigrifrons Duda, 1933: 161. Syntypes in slightly greyish. Cell br not widened. Legs black, NHMH. tarsi yellowish-brown. Body length 1.0–1.2 mm. Smaller and darker than I. kerteszi.Bodyblack. Distribution. – Sweden:SK,HA,SM,ÖL,UP.Fin- Frons entirely black. Ocellar triangle 2/3 as long land: Ab, N, Ob. Russia: Lr. Not so far found in Nor- as frons, dusted and not well limited from frons way and Denmark. surface. Eyes short pubescent. Genae narrow, half Trans-Palearctic species.

112 Figs 362–367. Incertella scotica (Collin): 362: head, dorsal view. 363: head, lateral view. 364: hypopygium, dorsal view. 365: hypandrium and phallic complex. I. zuercheri (Duda): 366: head, lateral view. 367: cerci and surstyli, dorsal view. (362–365 after Ismay & Rotheray, 1998.)

63. Incertella scotica Biology. – Adults in June to beginning of August, on (Collin, 1946) bogs with Carex. Tyrphobiont species. Figs 362–365 64. Incertella zuercheri Tropidoscinis scotica Collin, 1946: 137. Lectotype fe- male and paralectotype female in OXUM (Is- (Duda, 1933) may, 1993; Pont, 1995). Figs 366, 367 Body black, shining, similar to species of the genus Tropidoscinis zuercheri Duda, 1933: 82. Lectotype in Oscinella. Ocellar triangle extends to front margin ZMAN (Nartshuk, 2003). of frons, covered with very thin dust and subshin- Similar to species of Oscinella. Body black, shining. ing, with a row of 6–7 inwardly directed setulae Ocellar triangle covered with very thin dust and just inside lateral margin and shining spots near subshining, reaching front margin of frons. Eyes ocelli. Genae slightly broader than anterior tibia. + more longer and densely haired than in Oscinella npl 1 1, posterior seta slightly longer and much species. Genae half as wide as first flagellomere. stouterthananterior.Femoralcombonmalemid- Antennae and palpi black. Scutum and scutellum dle femora a single row of setae. Halteres yellow. covered with thin dust and subshining. Apical setae Legs black with pale femora-tibial joints of anterior wide apart, longer than scutellum. Pleura mainly and middle legs and all tarsi. Male genitalia: cerci shining. npl 1 + 1. Wings longer than abdomen. Hal- nearly circular, surstyli broad basally with setose teres yellow. Femora black, hind femora slightly en- inner patch, narrowed apically, hypandrium open. larged. Front and middle tibiae black or yellowish Body length 1.0–1.3 mm. at both ends. Body length 1.0–1.3 mm. Distribution. – Denmark:B.Sweden:SK,HA,SM, Distribution. – Denmark: EJ: Without locality, UP.Finland:Ab,N,Ob,Ks.Russia:Lr:LuttoRiver. 20.viii.1956 (H. Andersson), 1 female. Sweden: SK, Not so far found in Norway. HA,SM,UP,NB.Norway:AK,VE.Finland:Ab,N,Ta, European species. Tb, Sb. Russia: Vib, Kr.

113 Eurasian species, known eastwards to Yakutia, domen (Fig. 68). Male genitalia Fig. 379 East Siberia...... L. coxalis (Roser) Biology. – Adults from middle of May to September – Pregenital part of abdomen before epandrium on Common reed (Phragmites australis). without membranous vesicules (Fig. 387). Larvae phytophagous, live as inquilines in Lipara Male genitalia Figs 385, 386 galls between folded leaves in upper part of galls...... L. palposa (Fallén) Tschirnhaus (1981) listed as host plant also Bol- 3 Genae more smooth with single row of short boschoenus maritimus and Glyceria maxima,but these records need to be confirmed. setulae along darkened lower margin (Fig. Egg, all larval stages and puparium are described 372). Male genitalia Figs 375–377 by Grochowska (2008b)...... L. brevibucca (Duda) – Genae with several rows of short setulae along Genus Lasiambia Anonymous, 1937 lower margin and genae not so smooth (Fig. 368). Male genitalia Figs 369, 370 Lasiambia Enderlein, 1936: 188. Unavailable. Post- ...... L. baliola Collin 1930 name proposed without designation of a type species. 65. Lasiambia baliola Lasiambia Anonymous, 1937: 393 (see Evenhuis et al., 2008: 17). Type species: Oscinella fycoperda (Collin, 1946) Becker, 1910 by original designation. Figs 368–370 Lasiambia Sabrosky, 1941: 754. Type species: Os- Fiebrigella baliola Collin, 1946: 60. Lectotype male cinella fycoperda Becker, 1910 by original desig- and 8 paralectotypes in OXUM (Pont, 1995; Is- nation. may, 2000). Black, shining species with coarsely punctate scu- Body black. Ocellar triangle shining, extending lit- tum. Vibrissal angle produced anterior to anterior tle more than half length of frons. Genae narrower margin of eye. poc upright and parallel. Genae be- than width of first flagellomere, widened back- low the eye is obliquely divided into a rugose and wards, dark, with setulae in more than a single shining posterior and ventral part and a silvery row. Palpi stout and brownish. Scutum with thin dusted anterior and dorsal part. Most setae are pubescence and rather shining. npl 1 + 2, dorsal slightly thickened. Facial carina separates entirely hind seta smaller than other ones. Scutellum punc- or partly the base of antennae. Tibial organ oval. tate, undusted on disk and shining. Tergite 5 of Femoral organ absent. Male genitalia: cerci basally abdomen twice length of tergite 4. The apex of ter- fused, surstyli broad ovate pointed. Hypandrium gite 5 with stronger setulae than in L. brevibucca. open. Femora black, tibiae almost entirely black. Wings Distribution. – Recorded from all regions, except brownish along costal half. Halteres yellow. Body Australasian. length 1.5–1.8 mm. Distribution. – Sweden: SK, SÖ, UP, NB. Finland: Al. Key to species of Lasiambia Not so far found in Denmark, Norway or Russian Anonymous, 1937 parts of Fennoscandia. European species. 1 Genae wider than first flagellomere and vib- Biology. – Adults in June to July and again in Sep- rissal angle distinctly produced beyond eye tember in woodland and even in parks in metro- margin(Figs378,381)...... 2 politan area; in London once found on rather large – Genae not wider than first flagellomere and excrescence on the trunk of a wych elm (Allen, vibrissal angle not so distinctly produced be- 1981a). Larvae develop in rotten wood, especially broad yondeyemargin(Figs368,372)...... 3 leaved wood. It has been reared from an ulcerous 2 Pregenital part of abdomen before epandrium elm tree, sappy chestnut bark, sappy excrescence with two membranous vesicules, one on each on trunk of a wych elm (Ulmus glabra) (Collin, side, in dry specimens situated within ab- 1946; Allen, 1981a; Godfrey, 1998).

114 Figs 368–377. Lasiambia baliola (Collin): 368: head, lateral view. 369: hypopygium, dorsal view. 370: hypandrium and phallic complex. L. brevibucca (Duda): 371: head, dorsal view. 372: head, lateral view. 373: scutellum. 374: wing. 375: hypopygium, dorsal view. 376: hypan- drium and phallic complex. 377: tip of male abdomen, ventral view. (368–370 after Ismay.)

115 66. Lasiambia brevibucca and produced from oral cavity. Scutum shining, (Duda, 1933) densely punctate. Scutellum rounded behind with Figs 372–377 apical setae shorter than scutellum, subapical se- tae nearly 1/2 length of apical. Pleura shining. npl Goniopsita brevibucca Duda, 1933: 107. Syntypes in 1 + 1. Legs mainly black, but coxae from bright yel- ZMHU, NHMH and HNHM (Wendt, 1975). low to little paler than remaining parts of legs, ends Body black, shining. Frons shining, ocellar trian- of femora and tibiae brownish. Male has two pairs of vesiculae of intersegmental membrane, between gle shining as frons, extending 3/5 length of frons. + Genae narrower than first flagellomere, yellow tergite 5 and synsternite 7 8. Usually these vesic- with whitish dust and with single row of setulae on ulae are inside of abdomen in dry specimens. Body narrowly darkened ventral margin. Genae slightly length 2.5 mm. broaderinfemaleandsetulaeinmorethanone Distribution. – Finland: Al. Russia: Kr. Not found in row. Palpi large, yellow. First flagellomere yellow Denmark, Sweden or Norway. darkened above. Scutum shining densely covered European species. with setulae. Scutellum dusted and slightly rugose. Pleura shining. npl 1 + 2, dorsal hind seta is much Biology. – Larvae are parasitic and develop in egg smaller. Tergite 5 of abdomen 1.5 times length of masses of Chrysochaon dispar Germar (Acrididae) tergite 4. Coxae yellow, femora black, tibiae dis- (Hennig, 1941) and possible of other Acrididae. tinctly pale at both ends. Halteres yellow. Body There are many records of larvae of P. palposa deve- length 1.5–2.0 mm. loping in egg pods of Acrididae, but some of these may as well refer to L. coxalis. Distribution. – Denmark: LFM, NWZ, NEZ. Sweden: Hennig (1941) gave drawing of tentoropharyngeal SK,SM,VG,UP.Norway:AK:Lindøya,Oslo.Finland: skeleton of larva. Al: Eckerö, 17.vii.1922 (R. Frey), 3 males, 2 females (syntypes); Ab. Not found in Norway or Fennoscan- 68. Lasiambia palposa dian districts of Russia. European species. (Fallén, 1820) Figs 380–387 Biology. – Adults in June to August in woodland associated with damaged trees and occurs even in Oscinis palposa Fallén, 1820: 6. Lectotype in NRMS parks in metropolitan area; in London once found (Andersson, 1963). on rather large excrescence on the trunk of a wych Very similar to L. coxalis, differs in structure of elm (Allen, 1981a). male postabdomen: no vesiculae of intersegmen- Larvae develop in rotten wood, sap flows of dam- tal membrane, between tergite 5 and synsternite aged trees and in rot-holes. It has been reared from 7 + 8. Body length 2.5 mm. sappy horse-chestnut (Aesculus hippocastanum) (Godfrey, 1988). Distribution. – Denmark: WJ, NWJ, NEJ, F, NWZ, B. Sweden:northtoVB.Norway:AK,HEs,Bø,VE,TEy, 67. Lasiambia coxalis AAy. Finland: north to ObS. Russia: Vib, Kr. Widely distributed Palearctic species, known (Roser, 1840) from Europe eastwards to Yakutia, East Siberia and Figs 68, 378, 379 Mongolia and southwards to Israel. It has been re- Chlorops coxalis Roser, 1840: 62. Lectotype in SMNS corded in Iceland by early authors (Mazon, 1890; (Nartshuk, 1994). Meijere, 1910), but was not mentioned by more re- Goniopsita oophila Hennig, 1941: 198 (Nartshuk, cent authors (Nielsen & Tuxen, 1954; Lindroth et 1994). Type specimens in DEIM. al., 1973; Messersmith, 1982; Ólafsson, 1991). Black, shining species. Frons shining, reddish yel- Biology. – Adults from June to the end of Septem- low in front and greatly punctate. Ocellar trian- ber in meadows, wood edges. gle 3/5 length of frons, black smooth shining. First Larvae carnivorous, live in and may feed on egg flagellomere rounded, reddish, black above. Genae pods of Acrididae. As host following species were wider than first flagellomere, whitish yellow along recorded: Chortippus biguttulus, C. albomargina- eye margin, brown below. Vibrissal angle produced. tus, C. apricarius, Stenobothrus eurasius, S. nigro- Palpi brown to black, large, especially in male maculatus, Omocestus viridulus (Frauenfeld, 1863;

116 Figs 378–387. Lasiambia coxalis (Roser): 378: head, lateral view. 379: hypopygium, dorsal view. L. palposa (Fallén): 380: head, dorsal view. 381: head, lateral view. 382: face and antennae, anterior view. 383: scutellum. 384: wing. 385: hypopygium, dorsal view. 386: hypandrium and phallic complex. 387: tip of male abdomen, ventral view.

117 Bezrukov, 1922; Chapinsky, 1923; Zakhvatkin, 1954; 2 Setulae on body yellowish. Male genitalia: Figs Nartshuk, 1972b; Tschirnhaus, 1992). It is possi- 391,392...... L. lucens Meigen ble that larvae may feed other food. Adults are – Setulae on body whitish. Male genitalia: Fig. often taken in autumn on fallen trunks of Be- tula in Leningrad oblast and Gusev (1928) found 396...... L. similis Schiner this species on trunks of Picea infested by Ips sex- 3 Male genitalia: cerci fused and far projecting dentatus (Coleoptera). It has been reared from (Fig. 73) . . . . . L. pullitarsis Doskočil & Chvála mushroom Leccinum scabrum in Tatarstan (Russia) – Male genitalia: cerci separated (Fig. 395) (Khalidov, 1984). Beschovski (1982) and Beschovski ...... L. rufijitarsis Loew & Georgiev (1993) recorded rearing this species from fruit of Ficus carica and from galls of Paran- threne tabaniformis (Lepidoptera, Aegeridae) in Species key to 3rd instar larvae of Greece, but maybe this record concerns some other Lipara Meigen species. Mature larvae of Lipara 6–10 mm long and 2–3 mm at widest diameter. Body milky white or yellowish, Genus Lipara Meigen, 1830 cylindrical or slightly curved. One or three thoracic segments sclerotized and black on dorsal side. Fa- Lipara Meigen, 1830: 1. Type species: Lipara lucens cial mask with ridges in cells. Antennae two seg- Meigen, 1830 by monotypy. mented, apical segment is situated inside of basal Large stout black species with hairy eyes, broad segment. Maxillar palpus surrounded by sclero- facial carina and convex densely haired scutum. tized half-ring. Cephalopharyngeal skeleton black, Head and thorax setae indistinct hairlike. Pleura Mandibles with acessory teeth in two rows, hypos- with coarsely wrinkled or granulate microsculp- tomal and tentoropharyngial sclerites fused. Small ture. Tibial organ formed by an oval group of warts. papillae and spicules are situated on thoracic and Femoral organ present. Male genitalia: cerci small, abdominal segments. Abdominal segments devoid sometimes fused, surstyli broad simple, postgo- of sclerotized spicules. Anterior spiracles of fan- nites haired. Adults have sound communication by type, posterior spiracles sessile on surface of 8th vibration, sound produced by males being specific, abdominal segment, with branched interspiracu- sound produced by females of different species are lar processes. more similar (Mook & Bruggemann, 1968; Kan- Lipara similis Schiner, 1854 (Figs 403–408) from miya, 1981). Assumed mechanoreceptor is situated central and western Europe is included in the key, on the mid coxa (Kanmiya, 1981). The genus in- as the species may show up in Denmark and south cludes the largest species of the family. Larvae de- Sweden. velop in stems of Common reed (Phragmites aus- tralis). The biology of the four species in Europe 1 Only one or three anterior segments of body was studied in detail by Grochowska (2011). black dorsally, posterior segment not black (Figs 398, 399, 409, 410, 415, 416) ...... 2 Distribution. – Holarctic, Australia. – Anterior and posterior segments of body black Key to species of Lipara Meigen (Figs403,404)...... L. similis Schiner 2 Three anterior segments black dorsally (Fig. Lipara similis Schiner, 1854 (Fig. 396) from central 399). Mandibles with accessory teeth in two and western Europe is included in the key, as the rows, 3 in each row (Fig. 402). Larva 8–10 mm species may show up in Denmark and south Swe- long milky white in cigar-like gall with hard den. shortened internodes ...... L. lucens Meigen 1 Setulae on scutum orientated uniformly back- – Only one anterior segment black dorsally or ward. Body smaller, up to 4–5 mm ...... 3 dorsally and ventrally (Figs 410, 416)...... 3 – Setulae on scutum serially directed toward 3 First thoracic segment of body black dor- each other producing longitudinal stripes (Fig. sally and ventrally (Fig. 416). There are small 390). Body larger, up to 8 mm. Male genitalia: spicules on the second segment of body (first cerci separated (Figs 391, 396) ...... 2 thoracic) ventrally and around anal opening

118 (Fig. 416). Mandibles with accessory teeth in by males being specific, sounds produced by fe- two rows, a tooth in each row (Figs 417, 418). males of different species are more similar (Mook Anterior spiracles 6–9 buds (Fig. 419). Larva & Bruggemann, 1968). Kanmiya (1990) described milky white flattened ventrally 8–9 mm long geographical variation in the vibratory courtship (Fig. 415) in slightly enlarged stem of Prag- signals of the species. Newly emerged female carry mature eggs and can mate immediately. Eggs are mites...... L. rufijitarsis Loew laid externally on the surface of the reed stems or – First segment of body black only dorsally (Fig. leaves. Larvae phytophagous, producing terminal 410). There are small spicules on the first seg- cigar- or spike-like gall on shoots of Phragmites aus- ment ventrally,on the second segment of body tralis. After hatching the larvae crawl to the top, (first thoracic) dorsally and ventrally and on enter under the edge of a leaf sheath and gnaw the third segment (the second thoracic) ven- their way down through the enwrapped leaves un- trally (Fig. 410). Mandibles usually with two til they reach the growth point. Larvae feed on the additional teeth in each row (Figs 411, 412). enwrapped leaves and later enter the gall cham- Anterior spiracles 4–5 buds (Fig. 413). Larva ber. At the end of summer the full grown larvae enter diapause. The following spring larvae pupate 6–9 mm long milky white (Fig. 409) in cigar- in the chamber. Galls rather thick and contain a likegall...... L. pullitarsis Doskočil & Chvála hardened gall chamber. Common reed growing on land with a diameter about 3–4 mm are preferred 69. Lipara lucens as host plants (Valkeila, 1956; Mook, 1961, 1971; The- owald, 1961; De Bruyn, 1987; Wolcsánszky et al., Meigen, 1830 1993). Females avoid to lay eggs on reed standing in Figs 388–394, 397–402 water. The cause of this behaviour is explained by Lipara lucens Meigen, 1830: 1; Meigen’s plate 210: 5 anatomical structure and biology of the reed show- (Morge, 1976b). Holotype in NHMW (Nartshuk, ing xerophilous and hygrophilous habit (Ambrus, 1997a). 1977). The meristem of infested plants stops func- tioning. Several histological changes are caused by Large robust species, the largest species of the modified metabolism of the tissues: sclerotisation, genus and of the family in the Palearctic region. development of lignine deposits, accumulation of Body black densely covered with long yellowish amylum in the ground tissue and clogging of the hairs, arranged on scutum in distinct longitudinal phloem or rhexigenous vessels (Reynvaan & Doc- stripes with hairs directed right and left. Antennae ters van Leeuwen, 1906; Wolcsánszky et al., 1993). with basal segments yellowish, first flagellomere Gromyssz-Kałkowska & Hubicka (1978) studied mainly blackish. Antennae divided by rather broad oxygen consumption during larval development facial carina. Genae wide. Pleura roughly wrinkled. of the species. Morphological and histochemical Legs darkened. Body length 5.5–8.0 mm. analysis of galls and effect of gall formation on the Distribution. – Denmark: SJ, EJ, WJ, NEJ, LFM, SZ, host plant were studied by De Bruyn et al. (1998), NEZ,B.Sweden:SK,SM,VG,UP,VS.Norway:AK, Vandevyvere & De Bruyn (1998). These authors de- Bø, some specimens in NHMW without exact local- scribe the process of cecidogenesis in detail. Gall ity and data. Finland: N, Ta, Sa, Kb. Russia: Vib. formation starts when the larvae feed outside the Euro-Mediterranean species. Sabrosky (1958) actual developing gall, on the enwrapped leaves. considered the species as introduced in North During gall growth internodes elongation is re- America. duced. Internally the pith parenchyma is destined to become the nutritive tissue. The tissue cylin- Biology. – Adults in May to June on Phragmites aus- der around the gall chamber widens up to three tralis, occurs predominantly on suboptimally de- times normal size. When gall is completed, the veloped reed growing on dry soil (Mook, 1971; Gro- larva gnaws through the growing point and enters chowska, 2011). Kontkanen (1939) described habi- the gall chamber, where it consumes the nutritive tats in Finland. tissues. The sclerenchymatization process starts There is only one generation per year. Overwin- now resulting in an extremely hardened gall. ter as larvae inside the gall. Adults emerge from Many species of Chloropidae, species of other the end of May until early June. Adults have sound families of Diptera and other groups of communication by vibration, sounds produced live as inquilines in folded leaves at upper part

119 Figs 388–402. Lipara lucens Meigen: 388: head, dorsal view. 389: head, lateral view. 390: thorax, dorsal view. 391: hypopygium, dorsal view. 392: hypandrium and phallic complex. 393: apex of ovipositor, dorsal view. 394: oscillogramma of vibration signals of male (A: specimen from Latvia; B: specimen from England; dotted vertical line, from 20 msec). L. rufijitarsis Loew: 395: outline of cerci. L. similis Schiner: 396: outline of cerci. L. lucens Meigen, 3rd instar larva: 397: facial mask. 398: lateral view. 399: anterior end. 400: anterior spiracle. 401: posterior spiracle. 402: cephalopharyngeal skeleton. (395, 396, 399 after Chvála et al., 1974.)

120 of Lipara galls. Chloropidae: Calamoncosis minima, 70. Lipara pullitarsis C. duinensis, Incertella zuercheri, Elachiptera cor- Doskočil & Chvála, 1971 nuta, Eribolus hungaricus, Oscinella angustipennis, Figs 73, 409–414 Cryptonevra diadema, C. flavitarsis, C. consimilis. Two more European species of Chloropidae Homa- Lipara pullitarsis Doskočil & Chvála, 1971: 102. lura tarsata Meigen and C. nigritarsis (Duda) are Holotype in DECU, paratypes in DECU, NMPC associated with Lipara galls. They are not known and BMNH. from Scandinavia, but may show up in southern part of Denmark. Records of Oscinella frit and O. Black, shining robust species, but distinctly smaller pusilla from galls (Giraud, 1863; Blair, 1932; The- than L. lucens. Scutum evenly and densely covered owald, 1961) probably concern Incertella zuercheri. with yellowish setulae not arranged in stripes. An- Some authors (Seguy, 1934; Wend, 1968) recorded tennae mainly black, basal segments sometimes Oscinella nitidissima from Lipara galls. brown. Facial carina broader than in L. lucens,half Cecidomyiidae: Giraudiella inclusa (Frauen- width of first flagellomere. Legs black with yellow feldt), Microlasioptera flexuosa (Winnertz), Lasiop- base of tarsi. Body length 3.5–5.0 mm. tera hungarica Möhn, L. arundinis Schiner, Asy- Distribution. – Denmark: NEZ. Sweden: SK, HA, napta phragmitis (Giraud). SM,SÖ,UP,DR.Norway:AK,Bø,VE,AAy.Finland: Anthomyzidae: Anthomyza collini Andersson, A. gracilis (Fallén). Ab, N. Russia: Vib. Agromyzidae: Phytomyza succisae Hering (The- Euro-Mediterranean species. owald, 1961). The last species is usually known as Biology. – Adults in May to June on Phragmites aus- a leaf-miner on Succisa pratensis. Therefore, the tralis. record from the Lipara galls is doubtful. Larvae phytophagous, produce terminal gall on Many bees and wasps make their nests in 2-years shoots of Phragmites australis. Galls are soft and do vacated galls (see chapter ‘Role in land ecosys- not contain a hardened gall chamber. Flies prefer- tems and species of economic importance’). Sev- able infest thicker stems of host plant than L. lucens eral species of mites (Acarina) were recorded with- (De Bruyn, 1987). Some species of Chloropidae and in galls of Lipara lucens and L. pullitarsis: Siteroptes other insects live as inquilines in folded leaves in (S.) avenae (Müller), S. (S.) reniformis Krantz (Pyg- upper part of the gall. All species of inquilines are mephoridae), Tarsonemuslacustris Schaarschmidt, listed under L. lucens. Stenotarsonemus phragmitidis (v. Schlechtendal) Larvae described by Chvála et al. (1974), egg, all and S. (Mahunkacus) gibber Suski (Tarsonemidae), larval stages, puparium and biology described by Urodiaspis tecta (Kramer) (Urodiaspidae), Tydeus Grochowska (2006, 2011). sp. (Tydeidae) (Imhof, 1979; Rach & Carstensen, 1981; Abraham & Carstensen, 1982). The bush- cricket Conocephalus dorsalis (Latreille) (Cono- 71. Lipara rufijitarsis cephalidae) is also associated with Lipara galls Loew, 1858 (Waitzbauer, 1969). Recently Reader (2003) de- Figs 395, 415–420 scribed interaction betweenL.lucensand Cryp- tonevra flavitarsis (on my opinion erroneously Lipara rufijitarsis Loew, 1858: 57. Lectotype in named as L. rufijitarsis in the paper) in the term of ZMHU, designated by Doskočil & Chvála (1971). intraguild kleptoparasitism. It is difficult to agree Wend (1975) listed the species in the Index, but with this point of view, as larvae of these species not in the main text of her paper. feed different part of plant and feeding of larvae Cryptonevra flavitarsis is not influenced on gall for- Black, shining robust species, but distinctly smaller mers. than L. lucens. Scutum evenly and densely covered For parasitoids, predators and other inhabitants with whitish setulae, which not arranged in stripes. in Lipara galls see the chapters ‘Parasitoids, preda- Antennae mainly yellow, basal segments some- tors and parasites’ and ‘Role in land ecosystems and times brown. Facial carina narrower than in L. pul- species of economic importance’. litarsis, in the middle scarcely one third as wide The most detailed descriptions of egg and larvae a first flagellomere, but distinctly widening above are done by Waitzbauer (1969), Chvála et al. (1974) and below. Legs black with bright yellow tarsi. Body and Beschovski (1995). length 3.5–5.0 mm.

121 Figs 403–414. Lipara similis Schiner, 3rd instar larva: 403: lateral view. 404: anterior end. 405: cephalopharyngeal skeleton. 406: mandibula. 407: anterior spiracle. 408: posterior spiracle. L. pullitarsis Doskočil & Chvála, 3rd instar larva: 409: lateral. 410: anterior end. 411: cephalopharyngeal skeleton. 412: mandibula. 413: anterior spiracle. 414: posterior spiracle. (403 after Grochowska, 2006a; 404, 410 after Chvála et al., 1974.)

Figs 415–420. Lipara rufijitarsis Loew, 3rd instar larva: 415: lateral view. 416: anterior end. 417: cephalopharyngeal skeleton. 418: mandibula. 419: anterior spiracle. 420: posterior spiracle. (416 after Chvála et al., 1974.)

Distribution. – Only Sweden: UP: Flottsund, galls Genus Microcercis Beschovski, 1978 collected 03.iii.1951, flies emerged 22.iv.1951 (S. Jo- hansson), 2 females and a puparium. Even if only Microcercis Beschovski,1978:28,assubgenusofOs- females are available, we are confident about the cinella Becker. Type species: Oscinella trigonella identification. Duda, 1933 by original designation. European species, known from Central Europe. Body black. Ocellar triangle shining, extending half Biology. – Larvae are phytophagous as in other length of frons. Genae narrow. Antennae and palpi species of the genus and produce a top gall on black. First flagellomere round. Scutum shining shoots of Phragmitesaustralis. Usually galls are nar- with setulae arranged more or less in rows. Scutel- rower than galls of L.lucens, but sometimes they are lum and hind part of scutum before scutellum dusted. Male genitalia: cerci small. Surstyli short indistinguishable from galls of L. lucens (Chvála et wide, not tapering. Hypandrium closed. al., 1974). Egg, all larval stages, puparium and biol- ogy described by Grochwska (2007a, 2011). Distribution. –Palearctic.

122 Note. – Beschovski (1978) described Microcercis as a subgenus of Oscinella, but later (Beschovski, 1981b) he treated it as a subgenus of Incertella Sabrosky, 1980. The type species of the genus In- certella, Conioscinella incerta Becker, is a non- Palearctic species that we have not examined and therefore we are not able to discuss the taxonomic status of the names Incertella and Microcercis and consider here the latter one as separate genus.

Key to species of Microcercis Beschovsky Figs 421–423. Microcercis trigonella (Duda): 421: hypopygium, dorsal view. 422: surstylus, lateral view. 423: hypandrium and phallic complex. 1 Scutum shining except part before scutellum. Acrostichal and dorsocentral setulae arranged in double rows. Male genitalia Figs 421, 422 ...... M. trigonella (Duda) 1984b), where it was listed as a valid species in Os- – Scutum dusted as in most species of Oscinella. cinella. Unfortunately only females are present in Acrostichal setae arranged in double rows the available material deposited in the Zoological ...... M. kroeberi (Duda) Museum, Hamburg. Without being able to exam- ine the male genitalia we provisionally assign kroe- beri to the genus Microcercis based on following 72. Microcercis kroeberi characters: frons dull and straight in front (not con- (Duda, 1935), comb. & stat. nov. cave as in Oscinella), ocellar triangle short and shin- ing, acrostichal setulae arranged in double rows, Oscinella kroeberi Duda, 1935: 158, as var. of O. frit npl setae 1 + 1(inOscinella usually 1 + 2). For a (Linnaeus). Syntypes in ZMHU. discussion about relationships between Microcer- cis and Incertella see under Microcercis. Body black. Frons dull, brownish in front. Ocel- lar triangle shining, extending half length of frons. Genae as narrow as front tibia. Antennae and palpi 73. Microcercis trigonella black. First flagellomere round. Scutum dusted, with setulae arranged in double rows along acros- (Duda, 1933) tichal lines. Scutellum dusted, apical setae longer Figs 421–423 than scutellum, wide apart, one very short subapi- cal seta on each side. Pleura mainly shining, except Oscinella trigonella Duda, 1933: 196. Syntypes in on upper part. npl 1 + 1. Legs black, except brown- HNHM. ish tarsi. Wings slightly brownish. Body length 1.0– Body black. Frons dull, brownish in front. Ocel- 1.5 mm. lar triangle shining, extending half length of frons. Distribution. – Denmark: SJ: Sønderborg, vi.1897 Genae as narrow as front tibia, brownish. Anten- (W. Wüstnei), 1 female; WJ: Tipperne, viii.1973 (E. nae and palpi black. First flagellomere round. Scu- Rald), 2 females; NEJ: Kringelrøn & Hornfiskrøn, tum mainly shining, with setulae arranged in dou- Læsø, 14.vii.1969 (Zool. Mus. Copenh. Exp.), 1 fe- ble rows along acrostichal and dorsocentral lines. male; F: Keldsnor, Langeland, 06.viii.1975 (Lyne- Scutellum and hind part of scutum before scutel- borg, Martin & Michelsen), 2 females. Russia: Kr: lum dusted. npl 1 + 1. Legs mainly black, but tibiae Kartesh, 21–25.vii.1993 (Sugonyaev), 1 female. yellowish at ends. Body length 1.0–1.5 mm. Note. – Duda (1935) described kroeberi from north- Distribution. – Denmark: SJ, EJ, WJ, LFM. Sweden: ern Germany as a variety of Oscinellafrit.Thistaxon north to TO. Norway: Bv. Finland: north to Le. Rus- has remained unrecognized in the literature except sia: Vib, Kr, Lr. in the Catalogue of Palaearctic Diptera (Nartshuk, Eurasian species.

123 Biology. – Adults from the end of May to early Au- Key to species of Oscinella Becker gust in wet meadows. Collin (1939) recorded numerous specimens bred by Mr. Basden from mole’s nest in June in England. 1 Scutum shining . . . . . O. nitidissima (Meigen) Larvae live in stems of Juncus efffusum, J. gerardii, – Scutum dusted, not shining, only postpronota Glyceria fluitans, Phalaris arundinaceae, Festuca maybeshining...... 2 arundinaceae and Deschampsia caespitosa (Ro- gochaya, 1960; Sharipov, 1981; Tschirnhaus, 1981). 2 Arista densely pubescent, enlarged, white or black (Fig. 21). Scutum densely dusted . . . . . 3 – Arista dark, rather thinly pubescent. Scutum Genus Oscinella Becker, 1909 lessdenselydusted...... 4 3 Arista white (Fig. 439) . . . . O. maura (Fallén) – Arista black. Male genitalia Figs 429, 430 Oscinella Becker, 1909: 120. Type species: Musca frit ...... O. capreolus (Haliday) Linnaeus, 1758 by designation of ICZN, 1978: 203 (Opinion 1100). 4 Postpronota shining, scutum dusted Pachychoeta Bezzi, 1895: 72 (preocc. Bigot, 1857)...... O. ventricosi Nartshuk Type species: Elachiptera aterrima Strobl, 1880 – Postpronota and scutum both dusted ...... 5 = ( Oscinis capreolus Haliday, 1838) by original 5 Last segment of all tarsi widened (Fig. 448) designation...... 6 Bezzi, 1906: 50, as replacement – Last segment of all tarsi not widened ...... 7 name for Pachychoeta Bezzi, 1895. Type species: Elachiptera aterrima Strobl, 1880 (= Oscinis 6 Legs more extensively dark, only extreme tips capreolus Haliday, 1838), automatic. of femora yellow, trochanter of front legs Pachychaetina Hendel, 1907: 98, as replacement black. Vibrissal angle rounded (Fig. 424). Ocel- name for Pachychoeta Bezzi, 1895. Type species: lar triangle dusted along sides Elachiptera aterrima Strobl, 1880 (= Oscinis ...... O. angularis Collin capreolus Haliday, 1838), automatic. – Legs less extensively dark, all femora with Note. – An application to the ICZN (Case 3576) broadly yellow tips, trochanter of front legs to conserve Oscinella Becker, 1909 by reversal of yellow. Vibrissal angle rather angular (Fig. precedence with Melanochaeta Bezzi, 1906 and 447). Ocellar triangle not dusted along sides Pachychaetina Hendel, 1907 has been made by ...... O. trochanterata Collin Tschirnhaus & Nartshuk (2012). 7 Wings long and narrow, c. 3 times longer than wide (Fig. 425). Male genitalia Figs 426, 427 Black species mainly with dusted or rarely shin- ing scutum. Frons dull, ocellar triangle large shin- ...... O. angustipennis Duda ing. Front margin of frons concave. Vibrissal an- – Wings wider, only about 2.5 times longer than gle right or slightly retreated. poc upright, cruci- wide(Fig.433)...... 8 ate. First flagellomere rounded, as long as wide. 8 Genae very narrow, not as wide as front tibiae Arista mainly slender dark with short pubescence ...... O. cariciphila Collin or rarely white with dense pubescence or enlarged with black pubescence. Scutum convex with setu- – Genae not so narrow, wider than front tibiae lae arranged in lines. Pleura mainly shining. Tibial ...... 9 organ oval. Femoral organ about 6–9 warts in a row. 9 Tibiae of front and middle legs entirely yellow, Male genitalia: cerci discrete and large, surstyli sim- tibiae of hind legs black on middle part. Male ple, hypandrium open. genitalia Figs 445, 446 . . . O. pusilla (Meigen) The genus is divided into two subgenera: Os- – Legs variably but more extensively dark, tib- cinella s. str. has large, more or less triangular male iae never entirely yellow. (O. frit (Linnaeus) cerci; Cyclocercula Beschovski, 1978 has nearly cir- speciescomplex)...... 10 cular male cerci. 10 Legs entirely black Distribution. –Worldwide...... O. nigerrima (Macquart)

124 – Legs not entirely black, at least part of front Musca hordei Bjerkander 1777; Sweden and middle tibiae yellow or yellowish . . . . . 11 Musca avenae Bjerkander 1781; Sweden Oscinis flavipes Olivier 1813; France 11 Genae distinctly wider than first flagellomere. Oscinis nigra Olivier 1813; France Wings darkened and usually not reaching end Chlorops fumipennis Meigen 1830; Germany ofabdomen...... O. vastator (Curtis) Oscinis brunnitarsis Macquart 1835; France – Genae not wider than first flagellomere . . . 12 Hydrellia rufijitarsis Meigen 1838; Germany 12 Hind tibiae entirely dark, front and middle tib- Chlorops nigrita Meigen 1838; Germany iae never entirely yellow. Genae wider, nearly Chlorops aenea Roser 1840; Germany as wide as first flagellomere. Wings clear and Chlorops distincta Roser 1840; Germany Chlorops nigrimana Roser 1840; Germany reaching end of abdomen. Male genitalia Figs Oscinis granarius Curtis 1846; England 434–436...... O. frit (Linnaeus) Oscinis frit var. nigripes Strobl 1894; Austria – Hind tibiae dark with yellowish base and tip, Oscinella frit var. plumiseta Duda 1933; Europe front and middle tibiae extensively or entirely Oscinella sziladyi Duda 1933; Bulgaria yellow. Genae distinctly narrower than first flagellomere. Wings a little darkened The applied literature dealing with frit flies as ...... O. hortensis Collin a pest in crops of cereals and fodder grasses in Europe, notably from England, Germany, France, Note. –“Oscinella frit” in the traditional, broad Denmark, Sweden and Norway, usually distin- sense is a complex of several very similar and vari- guishes between 3 species: Oscinella frit, O. pusilla able species, e.g. in respect to the width of genae and O. vastator, sometimes even O. hortensis.Di- and colour of legs, and thus very difficult to iden- agnostic characters used to distinguish the first tify.From this complex only species found regularly three species are given in the chapter ‘Role in land in crop fields are included into the key. ecosystems and species of economic importance.’ The Oscinella frit complex consists of the follow- The relative length of the second and third segment ing taxa either considered as distinct species or of arista helps to separate species occurring in agri- forms of doubtful taxonomical status, part of them cultural fields or reared from cereals and cultivated described exclusively or predominantly on larval grasses, but this character may not work in relation stages: to other species of the complex. An attempt to solve the taxonomy of the O. frit complex using caryolog- O. agropyri Mesnil, described from France on larval ical methods (Michailova & Beschovski, 1985) did stage in Balachowsky & Mesnil, 1935 not give unambiguous results. The chromosomes O. alopecuri Mesnil, described from France on lar- found were long and slender and difficult to study. val stage in Balachowsky & Mesnil, 1935 Extended breeding experiments with pure cultures O. festuceae Mesnil, described from France on lar- and molecular studies may help to settle the out- val stage in Balachowsky & Mesnil, 1935 standing taxonomic problems still pending within O. frit,formArrhanatherum, described on larval the Oscinella frit species complex. stage by Jepson & Nye (1957) and Nye (1958) O. frit,formAnthoxanthum, described on larval stage by Jepson & Nye (1957) and Nye (1958) Species key to known 3rd instar larvae O. exigua Collin, 1946, described from England as var. of O. frit (Linnaeus) of Oscinella O. hortensis Collin, 1946; England O. nigerrima (Macquart, 1835); France 1 3rd thoracic segment on ventral side covered O. phlei Nartshuk, 1956; European part of Russia with one or several rows of spicules (Figs 451, O. pusilla (Meigen, 1830); Germany 452,453,457,458)...... 2 O. vastator (Curtis, 1845); England – 3rd thoracic segment without ventral rows of spicules (Fig. 450) ...... O. angularis Collin Several other names are customarily listed as synonyms of O. frit (Linnaeus), even though the 2 Ventral creeping welts of abdominal segments type specimens no longer exist, or have never been with one or two rows of enlarged spicules (Figs adequately examined: 453,457,458)...... 3

125 – Ventral creeping welts of abdominal segments Biology. – Adults in June to July and in September without rows of enlarged spicules, if some in wet meadows and marshy habitats. large specules present, they do not form dis- Larvae phytophagous, live gregariously in shoots creterows(Figs451,452)...... 5 of Phalaris arundinaceae (Collin, 1946; Nartshuk, 1956; Svirina, 1984). The tissues of the host plant 3 Ventral creeping welts of abdominal segments abovethefeedingsiteoflarvaebecomebrownand with two rows of enlarged spicules (Fig. 458) the central leaf dries up...... O. ventricosi Nartshuk Larva described by Nartshuk (1956). – Ventral creeping welts of abdominal segments only with one row of enlarged spicules (Figs 453,457)...... 4 75. Oscinella (Cyclocercula) 4 On creeping welts of abdominal segments be- angustipennis hind of a row from larger spicules only a row (Duda, 1932) of smaller spicules presents (Fig. 457) Figs 425–427 ...... O. pusilla (Meigen) Oscinella angustipennis Duda, 1932: 87. Lectotype – On creeping welts of abdominal segments two in ZMAN (Nartshuk, 2003). rows from smaller spicules presents (Fig. 453) ...... O. nitidissima (Meigen) Body black, shining. Frons dull. Ocellar triangle smooth shining extending 3/4 length of frons. 5 On creeping welts of abdominal segments Genaehalfaswideasfirstflagellomere.Firstflag- small spicules form 3–4 rows (Fig. 452). Stig- ellomere rounded. Arista longer haired than in O. mophores closed, distance between them frit. Scutum and scutellum dusted. Legs black with shorter than a stigmophore. Larvae yellowish brownish tarsi. Wings long and narrow, 3 times ...... O. maura (Fallén) longer than wide, sometimes slightly darkened. – On creeping welts of abdominal segments Vein R2+3 distinctly concave. Halteres yellow. Body small spicules form only a row (Fig. 451)Lar- length 1.0–1.3 mm. vaewhite...... O. frit (Linnaeus) Distribution. – Found only in Russian parts of Fennoscandia: Vib: Jukki; Peri. 74. Oscinella (Oscinella) angularis European species. Collin, 1946 Biology. – Adult in June to July in wet meadows Figs 424, 449, 450 and Phragmites beds. It has been reared from Li- para galls in Netherlands (De Bruyn, 1985). Lar- Oscinella angularis Collin, 1946: 141. Syntypes: 13 vae live between the enwrapping leaves in upper males, 4 females in OXUM, 1 female in BMNH part of gall. Ismay (1981b) found the species on (Pont, 1995). Glyceria sp. on marsh. Uffen & Chandler (1978) re- Body black. Ocellar triangle extending nearly to corded Phalaris arundinaceae as food plant, but front margin of frons, shining, but covered with Drake (1988) recorded no correlation between the thin dust at tip and along sides. Genae with occurrence of this plant and the fly in Great Britain. roundedvibrissalangleandheadinsideview rather retreating. Scutum dusted. Legs black with knees and tip of tibiae yellowish. Middle and hind 76. Oscinella (Oscinella) capreolus tarsi yellow at base. The last segments of all tarsi (Haliday, 1838), comb. n. enlarged. Knob of halteres yellow, stem dark. Body Figs 21, 428–430 length 1.5–2.0 mm. Osciniscapreolus Haliday, 1838: 187. Described from Distribution. – Denmark: F: Humble, Langeland, ‘England’. Whereabouts of type material un- 20.vii.1964(O.Martin),1female.Sweden:SK,SM. known. Ismay et al. (2001) did not mention this Finland: N, Ta, Om. Russia: Vib. Not so far found in species when revising Haliday’s collection. Norway. European species, but found in North America, Differs from other Oscinella species by flattened probably introduced (Nartshuk, 2001). and densely pubescent black arista. Body black.

126 Figs 424–430. Oscinella angularis Collin: 424: head, lateral view. O. angustipennis Duda: 425: wing. 426: hypopygium, dorsal view. 427: hypopygium, lateral view. O. capreolus (Haliday): 428: wing. 429: hypopygium, dorsal view. 430: hypandrium and phallic complex. (426, 427 after Beschovski, 1978.)

Frons dull. Ocellar triangle smooth shining, reach- Bezzi, 1906, Pachychoeta Bezzi, 1895, and Pachy- ing front margin of frons. 4 pairs of orb,longer chaetina Hendel, 1907. As the generic name Mela- than remaining setulae of frons. First flagellomere nochaeta Bezzi, 1906 has priority over the widely rounded. Genae narrower than first flagellomere. used generic name Oscinella Becker, 1909, an ap- Scutum and scutellum dusted, scutellum as long peal has been made to the ICZN to conserve of the as wide at base with broadly rounded apex. Pleura generic name Oscinella Becker, 1909 and include mainly shining. npl 1 + 2. Wings brownish. Halteres it into Official List of Generic Names in Zoology dark. Body length nearly 2 mm. (Tschirnhaus & Nartshuk, 1912). Other Palearctic species currently listed within Distribution. – Rare species, found only in Den- the genus Melanochaeta are now transferred to mark: LFM: Resle, 28.v.1882 (H.J. Hansen), 1 female. the genus Lasiochaeta Corti, 1909 (type species: Euro-Caucasian species. Elachiptera pubescens Thalhammer), now restored Note. – Collin (1946) stated about Melanochaeta from synonymy with Melanochaeta.Themalegeni- capreola (Haliday) that it “except for its arista, talia of Lasiochaetapubescens are of the Elachiptera might easily pass as a species of Oscinella”. Tschirn- type with apically widened surstyli, see Andersson haus (1992) even expressed the opinion that Hali- (1977: fig. 31). day’s species rightly belongs to Oscinella.Examina- tion of the male genitalia from specimens kindly 77. Oscinella (Oscinella) cariciphila sent by Tschirnhaus to Nartshuk supports this opinion. Oscinis capreolus has male genitalia of Os- Collin, 1946 cinella type with rather big, nearly triangular cerci Oscinella cariciphila Collin, 1946: 141. Syntypes: 20 and simple gonites – not widened in apical part males and 8 females in OXUM (Pont, 1995). as in the Palearctic Melanochaeta pubescens (Thal- hammer). We thus propose the new combination Body black, shining. Ocellar triangle reaching front Oscinella capreola (Haliday), and accordingly treat margin of frons. Genae not wider then fore tibia. the following genus-group names, that have Osci- Scutum and scutellum dusted, pleura mainly shin- nis capreolus Haliday, 1838 as type species, in syn- ing. npl 1 + 2. Legs with front and middle femora onymy with Oscinella Becker, 1909: Melanochaeta very narrowly yellow at tip, front and middle tibiae

127 entirely yellow, hind tibiae black and only narrowly countries (see chapter ‘Role in land ecosystem and fuscous yellow about base and tip. Wings rather species of economic importance’). More than 60 short and broad. Halteres yellow with brownish species of wild grasses are recorded as host plants, stem. Body length 1.5–2.0 mm. but notably the species does not develop in shoots of Dactylis glomerata (Vickerman, 1978a). However, Distribution. – Sweden: SK, HA, GO, NÄ, UP, VR, the uncertain taxonomy within the O. frit species DR,HS.Norway:Ø,AK,HEs,VE,STi.Finland:N. complex suggests that some records refer to other, Russia: Vib; Kr: Kondopozhsky District: Kon, Bukol- closely related species. Usually polyphagous her- nikov Is., 24.vi.2003 (Polevoi), 1 male; Kartesh, 16– bivorous species exhibit a distinct preference for 29.vii.1966 (Tanasijtshuk), 2 females. Not so far one or a few host species. Nartshuk (1956) assumed found in Denmark. that O. frit prefers wild grasses of the tribe Avenae. Eurasian species. Vickerman (1978b) found out that the survival of O. Biology. – Adults in June in wet habitats with Carex, frit larvae was lower on barley than on either oats or probably larvae live in shoots of Carex. wheat. The females laid most eggs on oats, Lolium perenne and Festuca rubra, but relatively few eggs on barley in laboratory experiments (Vickerman, 78. Oscinella (Oscinella) frit 1978a). Larvae feeding on different grasses may dif- (Linnaeus, 1758) fer in the arrangement of spicules on ventral side of Figs 25, 55, 85, 95, 96, 103, 431–437, 451 the body. Jepson & Nye (1957, 1959) and Nye (1958) described such differences in larvae from Arrhen- Musca frit Linnaeus, 1758: 598. atherum and Anthoxanthum, respectively. It is not clear if these larvae belong to different species or Body black. Frons dull with smooth shining ocellar exhibit variation within O. frit under influence of triangle reaching front margin of frons. First flagel- the different host plants. It has been shown at least lomere rounded. Second aristal segment 0.5 times that no cross-breading takes place between O. frit as long as third aristal segment. Warts in femoral and O. pusilla (Le Berre & Chevin, 1963; Agapova, comb situated close each other and rarely fused. 1966). Le Berre & Chevin (1961) demonstrated in Genae nearly as wide as first flagellomere or wider. experiments that colour of tibiae and arrangement Scutum and scutellum dusted. Pleura mainly shin- + of spicules on thoracic and abdominal segments of ing. npl 1 2. Legs mainly black or front and middle larvae are characters dependent on both tempera- tibiae partly yellow. Body length 1.5–2.0 mm. ture and host plant. O. frit has been reared from dis- Distribution. – One of the commonest species of posal dumps and compost (Werner, 1997), without the family. Widely distributed species (or group of doubt from larvae occurring in host plant remains. closely related species), known from all of Fenno- Larvae described by many authors, but in most scandia to the coast of Arctic Ocean. The species detail by Steel (1931, all instars), Balachowsky & has been found in all zoogeographical regions of Mesnil (1935), Riggert (1935), Nartshuk (1956, all the World. instars), Jepson & Nye (1957), Nye (1958), Jepson & Southwood (1960a, first instar). Biology. – Adults in May to September in meadows, wood edges, crop fields. Two or three generation peryearinNordiccountries.Overwinteraslarvae 79. Oscinella (Oscinella) hortensis within tillers of grasses. Adults have been found on Collin, 1946 xerothermic locality in November and December Fig. 438 in Germany (Thuringia) (Bährmann, 1996). Adults are very vagile, found up to 300 m above ground Oscinella hortensis Collin, 1946: 142. Syntypes: 44 (Johnson et al., 1962). Thunes et al. (2004) found males and 14 females in OXUM (Pont, 1995). adult in canopies of Pinus silvestris in Norway. Larvae phytophagous, develop in shoots and Similar to O. frit, but colour of legs paler. Collin seeds of cereals: wheat, barley, rye, oats and in wrote that in colour of legs and width of genae this shoots of many grasses. The species damages cere- species is intermediate between O. cariciphila and als and fodder grasses in all countries where it oc- O. pusilla. Shining ocellar triangle almost reaching curs, and there exist numerous publications on the front margin of frons. First flagellomere small as in biology and control of this species from all Nordic O. pusilla but genae narrower though not so narrow

128 as in O. cariciphila. Arista pubescence shorter. Scu- down inside a folded leaf (Mashek, 1955). It has tum more finely punctuated and less dusted than been found on agricultural grasses in Denmark in O. pusilla. Femora black, front and middle tibiae (Nielsen & Nielsen, 1984). yellow with black ring about middle. Hind tibiae Larvae described by Balachowsky & Mesnil darker. Male genitalia: surstyli narrower than in O. (1935), Nartshuk (1956), and Nye (1958). pusilla. Body length 1.0–1.5 mm. Distribution. – Probably in all Nordic countries, but the species is difficult distinguish from other 81. Oscinella (Oscinella) nigerrima species of O. frit group. (Macquart, 1835) Biology. – Host plants Lolium perenne, Deschamp- sia flexuosa, Poa sp. (Nielsen, 1994). Two genera- Oscinis nigerrima Macquart, 1835: 603. Types prob- tions per year in England (Vickerman, 1980). ably in MHNL. Note. – Nartshuk (1997a) studied a type specimen Similar to O. frit, but legs entirely black. Systematic of Oscinis vindicata Meigen, 1830, which appeared position of the species is not clear, possible it is only identical to Oscinella hortensis, but made no formal a dark coloured spring form of O. frit. synonymisation, as the taxonomy of this group is Distribution. – Not fully clarified as difficult to sep- insufficiently studied. arate from other closely related species, but evi- dently widespread in Denmark and Fennoscandia. 80. Oscinella (Oscinella) maura Biology. – The species develops on Festuca ovina (Fallén, 1820) (Nielsen, 1994) and Arrhenatherum elatius (Vick- Figs 6, 439, 452 erman, 1978a). Vickerman (1978b) established that larvae of the species did not survive on oats or bar- Oscinis maura Fallén, 1820: 6. Lectotype in ZMLU ley, but a few larvae survived on wheat. The first (Andersson, 1966). species of the genus to appear in spring. Two gen- Black species with densely white pubescent arista, erations per year in England (Vickerman, 1980). more densely punctured and dull scutum than in other species of the genus. Wings darkened and halteres often darkened. Legs usually black, but tib- 82. Oscinella (Oscinella) nitidissima iae sometimes partly yellow (Ismay, 1981c). Kario- = + (Meigen, 1838) type 2n 6 0 X. Morphology of chromosomes Figs440,453,454 different from other species of the O. frit complex (Michailova & Beschovski, 1985). Body length 1.7– 2.3 mm. Chlorops nitidissimus Meigen, 1838: 388. Lectotype female in MNHN (Nartshuk, 2006). Distribution. – Common species. Denmark: from all districts except WJ. Sweden: north to DR. Nor- Body black. Similar to O. frit, but scutum not dusted, way:AK,On,Bø,VE,Ry,HOi.Finland:northtoOa glabrous and shining. Frons dull. Ocellar triangle and Sa. Russia: Vib, Kr. smooth shining reaching front margin of frons. Known from Europe, Siberia, the Caucasus and Antennae black. First flagellomere round. Genae Iran. nearly as wide as first flagellomere. Scutellum dusted. Apical scutellar setae longer than scutel- Biology. – Adults from end of May to early of Sep- lum. Pleura shining. npl 1 + 2. Legs mainly black, tember, in meadows, wood edges, fields of fodder but ends of tibiae and tarsi yellowish. Body length grasses, 2–3 generations per year, but Vickerman 1.5–1.8 mm. (1980) observed only one generation in England. Larvae phytophagous, live in shoots of Dactylis Distribution. – Denmark: widespread. Sweden: glomerata (Séguy, 1934; Mashek, 1955; Nartshuk, north to LU. Norway: Bø: Akerøy, Hvaler, 6.vi.1992; 1956; Jepson & Nye, 1957; Nye, 1959), but Wetzel HOy: Apeltunv., Bergen, 02.viii.1936, 1 male. Fin- (1967) listed also Bromus inermis Leyss. Females lay land: north to Ob and Ks. Russia: Vib, Kr, Lr. eggs on the inside of a folded leaf of grass. Newly Holarctic species, widely distributed in Palearc- hatched larvae feed on their egg chorion and moves tic.

129 Figs 431–440. Oscinella frit (Linnaeus): 431: head, dorsal view. 432: head, lateral view. 433: wing. 434: hypopygium, ventral view. 435: hypopygium, lateral view. 436: hypandrium and phallic complex. 437: apex of ovipositor, ventral view. O. hortensis Collin: 438: head, lateral view. O. maura (Fallén): 439: head, lateral view. O. nitidissima (Meigen): 440: head, lateral view.

Biology. – Adults from end of May to end of August cus lanatus, Alopecurus geniculatus (Jepson & Nye, on diverse habitats: meadows, wood edges, crop 1957;Nye,1958,1959;Rogochaya,1960;Wetzel, fields. Found on agricultural grasses in Denmark 1967; Mowat, 1974, 1975). Some authors reared (Nielsen & Nielsen, 1984). Two generations per year the species from galls of Lipara lucens (Kramer, in England (Vickerman, 1978). 1917), but probably this represents misidentified In- Larvae phytophagous, develop in shoots of Poa- certella zuercheri. In Canada was recorded unusual cea, preferable in different species of Agrostis damage of oats caused by larvae of this species (Nartshuk, 1956; Vickerman, 1978a), but some au- (Comeau & Pelletier, 1977). Vickerman (1978b) es- thors listed also Anthoxanthum odoratum, Dactylis tablished that larvae of the species did not survive glomerata, Festuca arundinaceae, F. pratensis, F. on either oats, wheat or barley. rubra, Lolium multiflorum, L. perenne, Phleum Larvae described by Nartshuk (1956, as O. pratense, Poa annua, P. pratensis, P. trivialis, Hol- agrostis), and Nye (1958).

130 83. Oscinella (Cyclocercula) Very similar to O. frit, but differs in colour of legs: norrbottica front and middle tibiae entirely yellow, hind tib- iae yellow with a black band in the middle. Second Nartshuk & Andersson, sp. nov. aristal segment 0.3 times as long as third aristal seg- Figs 441–444 ment. Genae as wide as first flagellomere. Rows of Holotype, male, Sweden: Norrbotten: Jupukka, Pa- warts of femoral comb in male wider apart than jala, 28.vii.1955 (H. Andersson). Paratypes: Norr- in O. frit and distance between warts longer. Body botten: Jupukka, Pajala, 27–31.vii.1955 (H. Anders- length 1.5–2.0 mm. son), 2 males, 5 females; Kaunisjärvi, 08.vii.1951 (Ander, Ardö, Berden & Dahl), 1 female; Norrbot- Distribution. – Denmark: widespread. Sweden: ten, Kengis, 29–30.vii.1955 (H. Andersson), 1 male, north to TO. Norway: Ø, AK, HEn, Bø, HO, SFi, NT, 1 female. Holotype and 8 paratypes in ZMLU, 2 pa- Fn. Finland: north to Li. Russia: Vib, Kr, Lr. ratypes in ZISP. Widely distributed in the Palearctic Region. Diffferential diagnosis. – Beschovski (1978) de- Biology. – Adults occur in diverse habitats includ- scribed Cyclocercula as a subgenus of Oscinella ing crop fields. Becker with two included species O. (C.) angus- Larvae phytophagous, live in shoots and more tipennis and O. (C.) nartshukiana Beschovski, the rarely in seeds of cereals, but usually avoid oats, and latter is type species of the subgenus. The rounded in shoots of many wild grasses, preferable from the shapeofthemalecerciisthemaindefiningchar- tribe Hordeae (Nartshuk, 1956). O. pusilla and O. frit acter that also refers the present species to Cyclo- are not cross-breeding (Le Berre & Chevin, 1963; cercula.ItisdistinguishedfromO. (C.) angustipen- Agapova, 1966). A well known serious pest of cere- nis by shorter wings and body and from O. (C.) als in many European countries. nartshukiana by rounded first flagellomere with- Larvae described by Balachowsky & Mesnil (1935, out an angular upper corner. as O. grossa), Nartshuk (1956), Jepson & Nye (1957), Description. – Male, female. All body black. Frons and Nye (1958). square, dull. Ocellar triangle shining, nearly reach- ing front margin of frons. Genae narrow, no more Note. – Becker (1902) who studied Meigen’s types than half as wide as first flagellomere. Vibrissal an- in Vienna and Paris wrote “1 zerstörte Type in Paris. gle rounded. First flagellomere round, arista with 1 Exemplar in Wien...” (p. 319). Nartshuk (1997a) rather long pubescence. examined the specimen in NHMW and found that Scutum, postpronota and scutellum dusted. Se- it coincides with O. cariciphila Collin, has narrow tulae on scutum arranged in rows. Scutellum semi- genae equal of diameter of front tibiae, all femora circular, apical setae longer than scutellum, thick black, the anterior and middle tibiae and all tarsi at base and twice as far apart than their distance yellow, the posterior tibiae black with a yellow to small subapical setae. Pleura shining except on ends. According to Meigen’s description, all tibiae upper part. npl 1 + 2. Wings brownish. Cell br not and tarsi yellow (no Oscinella specieswithsuchcol- widened. Second and third costal sectors subequal. oration of legs in European fauna), but in Meigen’s Veins R4+5 and M divergent distally. Halteres yel- figure (Morge, 1976b) the base of hind tibiae are low with dark stem. Legs black. Male genitalia: cerci slightly darkened, as in the specimen in NHMW. rounded; surstyli broad; hypandrium open. Nartshuk examined the specimen in MNHN as Etymology. – The species is named after the type well. It is a male without head, part of legs and area, the Swedish province of Norrbotten. a wing. Front legs absent, middle tibiae all yel- low and hind leg black except tarsus. Scutum very densely dusted, more than in other species of the 84. Oscinella (Oscinella) pusilla genus. (Meigen, 1830) The name Oscinella pusilla Meigen is widely used Figs 445, 446, 455–457 in the applied literature. In order to conserve this important name it is necessary to request the ICZN Chlorops pusillus Meigen, 1830: 157; Meigen’s plate to reject the existing syntypes and designate a neo- 216: 17 (Morge, 1976b). Type specimen in type of Chlorops pusillus Meigen in accordance NHMW (Nartshuk, 1997a). with the accepted usage.

131 Figs 441–448. Oscinella norrbottica Nartshuk & Andersson: 441: part of hypopygium, dorsal view. 442: cerci. 443: cercus and surstylus, latero-dorsal view. 444: hypandrium and phallic complex. O. pusilla (Meigen): 445: hypopygium, ventral view. 446: hypopygium, lateral view. O. trochanterata Collin: 447: head, lateral view. 448: male middle tarsus. (448 after Collin, 1946.)

85. Oscinella (Oscinella) Biology. – Adults in June and August, probably two trochanterata generations per year. Occur in wet meadows, edges of ponds and marshy habitats, more rare than O. an- Collin, 1946 gularis, developing in the same plant-host. Figs 447, 448 Larvae phytophagous, develop in unshelled in- florescence of Phalaris arundinaceae (Collin, 1946; Oscinella trochanterata Collin, 1946: 140. Syntypes Nartshuk, 1956). The tissues of the infested plant 16 males and 12 females in OXUM, 1 male and 1 turns brown above the feeding site of the larvae. female, in BMNH (Pont, 1995). Wetzel (1967) listed as food plant also Bromus in- ermis. Body black. Ocellar triangle shining, extending Larvae described by Balachowsky & Mesnil (1935, 2/3–3/4 length of frons. Genae with rounded vib- as O. gracilior), and Nye (1958). rissal angle and head in profile rather retreating. Scutum densely dusted. The last segments of all tarsi enlarged and fourth segment short and deeply 86. Oscinella (Oscinella) vastator excised on upper apical margin. Trochanters of (Curtis, 1845) front legs conspicuously yellow. All femora broadly yellow at tip, hind tibiae yellow at both ends. Stem Oscinis vastator Curtis, 1845: 493. and knob of halteres yellow. Body length 2.5– Body black. Similar to O. frit, but larger up to 2.8 mm. 3.0 mm and genae wider than first flagellomere, middle tibiae yellow on both ends, but hind tibiae Distribution. – Denmark: SJ, NWJ, NEJ, NWZ, never so distinctly yellow at base and tip as in O. NEZ, B. Sweden: SK, BL, HA, ÖL. Russia: Vib. Not pusilla. Nye (1958) supposed that O. vastator and found in Norway and Finland. O. festucae Mesnil might be conspecific on base of European species. structures of larvae. Body length 2.3–3.0 mm.

132 Distribution. – Probably everywhere, but individ- short dusted. Face with facial carina. First flagel- ual specimens very difficult to distinguish from lomere rounded. Pleura mainly shining. npl 1 + 2. other closely related species of the genus. Found Tibial organ narrow oval. Femoral organ forming in meadows and agricultural fields. It is possible one row of warts. Male genitalia: cerci discrete, and that the species occurs in all Nordic countries. surstyli simple, hypandrium open. According to many authors the species damages different cereals. Vickerman (1978a) established, Distribution. –Palearctic. that O. vastator is the most polyphagous species and its preferred hosts are Lolium multiflorum, L. perenne and Festuca pratensis,butfemaleslayno Key to species of Oscinimorpha Lioy eggs on cereals in the laboratory. Nielsen (1994) found this species in Denmark on Lolium perenne, 1 All setae and setulae whitish. Pleura dull. Vib- L. multiflorum, Poa spp., mainly P. trivialis,butnot rissal angle strongly produced and proboscis on wheat. Other authors listed following grasses as rather long and geniculate (Figs 460, 461) host plants: Agrostistenuis, Bromusinermis, Festuca ...... O. albisetosa Duda pratensis, F. rubra, Alopecurus pratensis, Phleum – All setae and setulae black. Pleura partly shin- pratense, Arrhenatherum elatius, Lolium perenne, ing. Vibrissal angle less produced and pro- L. multiflorum (Jepson & Heard, 1959; Nye, 1959; boscis not so long and not geniculate ...... 2 Wetzel, 1967). Larvae described by Jepson & Nye (1957), Nye 2 Genae nearly a half as wide as first flagellom- (1958), and as O. festucae by Nartshuk (1956). ere, vibrissal angle more produced (Fig. 469). Body entirely black, including head and legs 87. Oscinella (Oscinella) ventricosi ...... O. sordidissima (Strobl) Nartshuk, 1956 – Genae distinctly narrower, not wider than Fig. 458 front tibia, vibrissal angle less produced (Fig. 463). Head and legs partly yellow. Male geni- Oscinella ventricosi Nartshuk, 1956: 873. Holotype talia Figs 466, 467. . . .O. minutissima (Strobl) in ZISP. Body black. Similar to O. frit. Ocellar triangle shin- 88. Oscinimorpha albisetosa ing and reaching front margin of frons. Genae as wide as first flagellomere. Scutum dusted, but post- (Duda, 1932) pronota shining without dusting. Body length 1.5– Figs 459–461 2.3 mm. Conioscinella albisetosa Duda, 1932: 47. Syntypes in Distribution. – Sweden: north to TO. Finland: Ta. ZMHU (Wendt, 1975). Probably more widespread in Fennoscandia, but confused with other species of the O. frit complex. Small grey-dusted species. Head yellow with black Eurasian species, eastwards to Yakutia, East Sibe- occiput. Ocellar triangle small black grey-dusted ria; occurs preferably in the north. and extending halfway to front margin of frons. Antennae yellow, first flagellomere small rounded. Biology. – Adults in July to August in rich mead- Genae as wide as first flagellomere, with strongly ows. Larvae phytophagous, develop in shoots of produced vibrissal angle. Proboscis thin long, palpi Alopecurus spp. (Nartshuk, 1956). yellow, long narrow. Scutum and scutellum light Larva described by Nartshuk (1956). grey-dusted. Scutellum half as long as wide at base. Apical setae shorter than scutellum. Pleura dusted. npl 1 + 2. All setae and setulae yellow. Abdomen Genus Oscinimorpha Lioy, 1864 yellowish at base. Legs yellow or femora and tibiae partly darkened. Wings whitish with yellow veins. Oscinimorpha Lioy, 1864: 1126. Type species: Oscinis Halteres yellow. Body length 1.5–1.7 mm. obliqua Macquart, 1835 by monotypy. Distribution. – Scarce. Denmark: SJ: Rømø; NWJ: Black grey-dusted species with produced vibrissal Struer; F: Langeland. Sweden: SK, ÖL. Not found in angle and prolonged proboscis. Ocellar triangle Norway, Finland or Russian parts of Fennoscandia.

133 Figs 449–458. Oscinella spp., 3rd instar larva and puparium: O. angularis Collin: 449: facial mask. 450: spicules on ventral side. O. frit (Linnaeus): 451: spicules on ventral side. O. maura (Fallén): 452: spicules on ventral side. O. nitidissima (Meigen): 453: spicules on ventral side. 454: posterior end of puparium. O. pusilla (Meigen): 455: facial mask. 456: mandibula. 457: spicules on ventral side. O. ventricosi Nartshuk: 458: spicules on ventral side.

134 Eurasian species, reaching north to S Sweden in and setulae black. Legs nearly entirely black. Body Europe, but the species goes much further north in length 1.5–2.0 mm. continental East Siberia, near Yakutsk. Distribution. – Sweden:SK,ÖL,GO,UP,DR.Fin- Biology. – Adults in June to August in dry meadows land: N. Russia: Vib. Not so far found in Denmark and sandy habitats. Larvae probably phytophagous, or Norway. as it has been reared from Elytrigia and Pucclinella Eurasian species. (Bährmann & Weipert, 1989). Biology. – Adults in June to August in meadows. In Great Britain the species has a restricted distri- 89. Oscinimorpha minutissima bution (Ismay, 1994b) and the conservation status (Strobl, 1900) of notable (Falk, 1991). It has been found in Scot- land on flat area behind the narrow fore dunes with Figs 56, 462–467 patches of short turf (Cole, 1996). Siphonella minutissima Strobl, 1900: 631. Syntypes 2 males in NMBA (Chvála, 2008). Genus Oscinisoma Lioy, 1864 Black grey-dusted species. Frons dull, narrow yel- lowish in front. Ocellar triangle black grey-dusted, extending 2/3 length of frons. Antennae yellow Oscinisoma Lioy, 1864: 1125. Type species: Chlorops with first flagellomere darkened. Genae narrower vitripennis Meigen, 1830 (= Chlorops cognatus than first flagellomere, distinctly produced. Pro- Meigen, 1830) by designation of Coquillett, 1910: boscis little elongated. Palpi yellow. Scutum and 582. scutellum dusted, pleura mainly shining. Scutel- Black species with flattened granulate scutum. lum semicircular, apical setae as long as scutellum Head much wider than long and deep. Ocellar tri- or longer. All setae and setulae black. Legs with angle shining or dusted. Vibrissal angle right, not femora and hind tibiae mainly black, remaining produced. poc upright, cruciate. First flagellom- parts yellow. Body length 1.0–1.5 mm. ere rounded, wider than long. Scutum markedly Distribution. – Denmark: SJ, NWJ, NEJ, F, SZ. Swe- rugose, evenly covered with dense setulae aris- den: north to TO. Norway: AK, HEn, VAy, Ry. Fin- ing from small prominences. Pleura mainly shin- land:northtoOm.Russia:Vib;Lr:KhibinyMts, ing. npl 1 + 2. Apical scutellar setae arising from 16.viii.1928 (Cheburova). small tubercles in most species. Tibial organ dis- Euro-Mediterranean species. tinct oval. Femoral organ composed of group of warts. Male genitalia with large cerci and slender Biology. – Adults from June to middle of August on surstyli. Hypandrium closed, postgonites with me- diverse meadows, wood edges. dian directed projection. As host plants Salvia sclarea, Secale, Oryza and Silene dioica were listed (Nartshuk, 1972a; Tschirn- Distribution. – Holarctic and Oriental. haus, 1981). Key to species of Oscinisoma Lioy 90. Oscinimorpha sordidissima (Strobl, 1893) 1 Ocellar triangle dusted and subshining. Male Figs 468–470 genitalia Figs 480, 481 ...... O. germanica (Duda) Siphonella sordidissima Strobl, 1893: 252. Syntype female in NMBA (Chvála, 2008). – Ocellar triangle not dusted and shining . . . . 2 2 Postpronota shining. Wings wider and shorter. Body black, dark grey-dusted. Frons black, ocel- Male genitalia Figs 475, 476 lar triangle black, dusted, extending half length of ...... O. cognatum (Meigen) frons. Antennae and palpi black. Genae narrower than first flagellomere, with produced vibrissal an- – Postpronota dusted. Wings narrower and gle. Scutum and scutellum finely punctuate and longer. Male genitalia Figs 482, 483 dusted. Pleura shining below. npl 1 + 2. All setae ...... O. gilvipes (Loew)

135 Figs 459–470. Oscinimorpha albisetosa (Duda): 459: head, dorsal view. 460: head, lateral view. 461: face and antennae, anterior view. O. minitissima (Strobl): 462: head, dorsal view. 463: head lateral view. 464: scutellum. 465: wing. 466: hypopygium, dorsal view. 467: hypandrium and phallic complex. O. sordidissima (Strobl): 468: head, dorsal view. 469: head lateral view. 470: wing.

91. Oscinisoma cognatum tum and scutellum flattened, coarsely punctuate, (Meigen, 1830) thinly dusted. Postpronota shining. Apical scutel- Figs 57, 471–476 lar setae on very small tubercles rather wide apart. Legs and halteres yellow. Body length 1.5–2.0 mm. Chloropscognatus Meigen, 1830: 154; Meigen’s plate 216: 16 (Morge, 1976b). Lectotype in NHMW Distribution. – Denmark:SJ,F,SZ,NEZ.Sweden: (Nartshuk, 1997a). north to VR and UP,and further in NB. Norway: VAy: Straumen,Farsund.Finland:Al,Ab,N,Ta,Sa,Om. Body black, shining. Body and wings are shorter Russia: Vib, Kr. than in O. gilvipes. Frons in front and face yel- The commonest species of the genus. Trans- low. Ocellar triangle black, shining, extending 3/4 Palearctic boreal species. length of frons. First flagellomere varies in the colour, from entirely yellow to darkened above. Biology. – Adults from May to October, common on Genae as wide as first flagellomere, yellow with bogs, fens, marshes and other wetlands. Flies live whitish dusting but black below. Palpi yellow. Scu- near the ground in deep litter, hibernate as adults.

136 Figs 471–483. Oscinisoma cognatum (Meigen): 471: head, dorsal view. 472: head, lateral view. 473: scutellum. 474: wing. 475: hypopygium, dorsal view. 476: hypandrium and phallic complex. O. germanicum (Loew): 477: head, dorsal view. 478: head, lateral view. 479: scutellum. 480: hypopygium, dorsal view. 481: hypandrium and phallic complex. O. gilvipes (Loew): 482: hypopygium, dorsal view. 483: hypandrium and phallic complex. (482, 483 after Ismay, 1976.)

It has been reared from shoots of Glyceria aquat- microsculpture and distinctly dusted. Postpronota ica, G. maxima, Phalaris arundinaceae, Festuca dusted. Colour of legs is strongly variable from en- arundinaceae, Carex sp., Scirpus, Sparganium, Ty- tirely yellow to darkened femora and tibiae. The pha (Rogochaya, 1960; Nartshuk, 1972a; Sharipov, male genitalia are larger than in the other species 1981; Tschirnhaus, 1992). of the genus. Distribution. – Rare. Found only in Sweden: SK: 92. Oscinisoma germanicum Skäralid, 20.vi.1941 (T. Nyholm), 1 male; SM: Rödsle, (Duda, 1932) Oskarshamn, 22.vii.1983 (R. Danielsson), 1 male. Figs 477–481 European species. Biology. – Collin (1939) reported that Mr. Basden Discogastrella germanica Duda, 1932: 42. Holotype found the species alive in one of his artificial decoy in ZMHU (Wendt, 1975). nests. Frons dusted, yellow with yellow setae and setulae. Ocellar triangle black, dusted in contrast to other species of the genus, extending half length of frons. 93. Oscinisoma gilvipes First flagellomere entirely yellow. Scutum not so (Loew, 1858) flat as in other species of the genus, with weaker Figs 482, 483

137 Oscinis gilvipes Loew, 1858: 71. Syntype in ZMHU apical setae approximated (Fig. 487b). npl 1 + (Wendt, 1975). 2(3). Vibrissal angle not strongly produced Frons dusted, black behind and yellow in ante- and proboscis shorter. Male genitalia Figs 488, rior third. Ocellar triangle black, shining, extending 489...... P. rufijicornis (Macquart) 2/3–3/4 down frons. orb about10,small,but2– –Wingcellbr not widened in the middle. Scu- 3 may be longer. Eyes with pale short hairs. First tellum with apical setae wide apart and no flagellomere wider than long, in male yellow with more than two lateral (subapical) on each apically darkened part, in female darkened on the side. Vibrissal angle strongly produced and upper half. Arista black. Genae in front as wide proboscis longer (Fig. 490). npl 1 + 1. Male as first flagellomere, wider behind, yellow in front genitalia Figs 491, 492 and black behind. Palpi yellow. Scutum longer than wide, shining, rugose, flattened in hind 3/4. Post- ...... P. sulcicollis (Meigen) pronota and notopleura dusted. npl 1 + 1. Most part of pleura shining. Scutellum black, wider than long, rugose. Subapical setae on small tubercles in some 94. Polyodaspis rufijicornis specimens. Wings longer than in O. cognata,longer (Macquart, 1835) than abdomen in some specimens. Halteres yellow. Figs 14, 93, 484–489, 964, 965 Body length 1.5–1.7 mm. Siphonella rufijicornis Macquart, 1835: 585. Types Distribution. – Rare. Found only in Sweden: SK, SM, probably in MHNL. UP. European species. Body black. Frons brown covered with brownish setulae on distinct punctures. Ocellar triangle in- Biology. – Adults from May to the end of September distinct, shining, minutely shagreened, extending in wet habitats, probably hibernating as adults. It has been reared from Glyceria maxima beyond 2/3 length of frons. poc, vte and vti sube- (Tschirnhaus, 1992). qual and rather stout. Antennae mainly yellow with first flagellomere rounded and darkened on dorsal margin. Arista black with very short pubescence. Genae a little wider than first flagellomere, dark Genus Polyodaspis Duda, 1933 brown, distinctly rugose, anterior part covered with more or less silvery dust. Vibrissal angle strongly Polyodaspis Duda, 1933: 61. Replacement name produced. Facial carina well developed, triangular for Macrothorax Lioy, 1864 (preocc. Jaennicke, above, very thin below, separating the deep anten- 1867). Type species: Siphonella rufijicornis Mac- nal foveae. Proboscis elongated. Palpi yellow with quart, 1835, automatic. black setulae and one strong apical seta. Black species with well developed facial carina, dis- Scutum subshining, densely shagreened. Scutel- tinctly produced vibrissal angle, shining frons and lum rounded apically, weakly flattened and sha- ocellar triangle. poc strong upright parallel. Genae greened with a pair of approximated apical and divided by oblique carina into a lower posterior 6–8 pair of short stout subapical setae on small tu- darker part with coarse sculpture and numerous bercles. Pleura shining, wrinkled and covered with + setulae and a smooth upper anterior part. Tib- small setulae. npl 1 2(3). Halteres with large black ial organ small oval. Femoral organ absent. Male knob. Legs mainly black. Abdomen black with genitalia: cerci simple, surstyli broad, hypandrium short black setulae. In wing cell br widened. Male open. genitalia: epandrium small with squarish strongly sclerotized cerci and broad surstyli pointed at tip. Distribution. – Palearctic, Oriental and Afrotropi- Body length 2.5–3.0 mm. cal Regions. Distribution. – Uncommon, except in Finland. Den- mark:?SJ:Nolocality[Crepis Litocha](W.Wüst- Key to species of Polyodaspis Duda nei). Norway: AK: Rakeie, Oslo; Kirkeby, Maridalen (bread from flowerhead of Cirsium helenioides). 1Wingcellbr widened in the middle (Fig. 487). Finland: N, Ta, Sa, Oa, Sb. Russia: Kr; Lr: Kandalak- Scutellum with numerous marginal setae and sha; Kola; Yläluostari. Not found in Sweden.

138 Widely distributed in the Palearctic Region, more Proboscis long with narrow labella. Palpi from yel- abundant in the Asian part of the Palearctic Region, lowish to black, as long as front tibia. Scutum recorded from the Oriental Region as well. and scutellum shining, slightly flattened. Scutel- lum with apical setae long and not approximated, Biology. – Adults in dry habitats. subapical setae thin and short. Pleura with dis- Larvae feed on different substances, usually en- tinct microsculpture. npl 1 + 1. Male abdomen with riched proteins, and exhibit very wide ecological membranous vesicula between 5 tergite and syns- plasticity in relation to the choice of food sub- ternite 7 + 8, seen only on macerated abdomen. strates and environmental conditions. They may Legs mainly black. Body length 1.5–1.8 mm. be saprophagous, phytophagous, necrophagous or parasitic (Kiauka & Nartshuk, 1972; Nartshuk, Distribution. – Uncommon. Denmark: B: Bagå, 1972b). Flies were reared from inflorescence of 16.vi and 19.vii.1964, 2 males. Sweden: SK, SM, GO, Asteraceae: Knautia arvensis together with larvae SÖ, UP. Norway: AK: Hovedøya and Lindøya, Oslo of Larinus sp., Cirsium spp. together with Fjord. Finland: Al, Ab, N. Not found in Russian parts caterpillars of Microlepidoptera and larvae of of Fennoscandia. Tephritidae, walnut fruits, both intact and infested Widespread in W Palearctic, eastwards to Yaku- by other insects, large heads of artichoke (Cy- tia, East Siberia and Mongolia. nara scolymus)togetherwithlarvaeofApion car- Biology. – Adults in May to July in dry habitats. duorum,stemsofCannabis sativa and Onopordon Larvae as in P. rufijicornis exhibit wide ecologi- acanthium together with Lixus pollinosus,stemsof cal plasticity in relation to the choice of food sub- maize usually damaged by other insects, from dead strates and environmental conditions. The adults and living caterpillars of Pyrausta nubilalis living in were reared from stem of broomrape (Orobanche stem of maize, from tachinid fly living as parasite of speciosa D.C.) damaged by other insects (Martelli, Pyrausta caterpillar, from Laspeyresia caterpillars 1935). Meijere (1938) suggested that the larvae are on apple and plum, from rolling leaves with lar- predators, but Martelli (1935) regarded them as vae of the beetle Byctiscus rugosus on aspen, from saprophytophagous. Apoderus coryli on river birch, found as ectopara- Some larval structures illustrated by Nartshuk site on larvae of Hoplocampa (Hymenoptera, Ten- (1987). thredinidae) on pear and plum trees, from galls of Paranthrene tabaniformis (Lepidoptera, Sesiidae) (Perris, 1830, 1839, 1873, 1876; Frauenfeld, 1863; Genus Pseudogaurax Malloch, 1915 Schiner, 1864; Perris & Laboulbéne, 1871; Vukaso- vič, 1926; Goidanich, 1928; Falcoz, 1930; Sachtleben, Pseudogaurax Malloch, 1915: 159. Type species: 1930, 1939; Martelli, 1933a, 1933b; Rogochaya, 1960; Gaurax anchora Loew, 1866 by original desig- Nartshuk & Kiauka, 1963; Beschovski & Georgiev, nation. 1993). The species is often introduced with dam- aged walnuts fruits. Yellow species with black markings, long triangular Larvae described by Kiauka (1974). scutellum with approximate apical scutellars. Ocel- lar triangle shining. First flagellomere obliquely prolonged with thickened pubescent arista. Tibial 95. Polyodaspis sulcicollis organ distinct, narrow oval. Femoral organ absent. (Meigen, 1838) Male genitalia: cerci fused basally, surstyli simple. Figs 67, 490–492 Distribution. –Worldwide. Chlorops sulcicollis Meigen, 1838: 387. Lectotype fe- male in MNHN (Nartshuk, 2006). 96. Pseudogaurax venustus Body black, shining. Frons shining brownish and (Czerny, 1906) punctuate. Ocellar triangle extending half length Fig. 493 of frons with a row setulae on lateral margins. An- Gaurax venustus Czerny, 1906: 299. Lectotype in tennae from yellow to black, first flagellomere a lit- NHMW (Nartshuk, 1997b). tle angulated. Arista black with short pubescence. Genae shining dark brown, black on below mar- Body yellow with black, shining longitudinal stripes gin, vibrissal angle acute and distinctly produced. on scutum. Frons narrow, longer than wide, yel-

139 Figs 484–492. Polyodaspis rufijicornis (Macquart): 484: head, dorsal view. 485: head, lateral view. 486: face and antennae, anterior view. 487: wing. 487b: scutellum. 488: hypopygium, dorsal view. 489: hypandrium and phallic complex. P.sulcicollis (Meigen): 490: head, lateral view. 491: hypopygium, dorsal view. 492: hypandrium and phallic complex.

140 low with yellow setae and setulae. Ocellar tri- dusted than scutum. Male genitalia Figs 515, angle yellow with black ocellar tubercle, smooth 516...... R. tomentosum sp. nov. shining, extending half length of frons. Occiput 2 Genae wider than front tibia. Scutum with with small black spot. Eyes large and dense and dusting at sides and behind, covered with pale long pubescent. Antennae yellow, first flagellom- ere wider than long. Arista short, basally thick setulae. Legs entirely darkened, only basitarsi and yellow, apically thin and darkened. Genae nar- of all legs yellowish row, less than half width of first flagellomere with ...... R. atricillum (Zetterstedt) rounded vibrissal angle. Scutum with 3 black, shin- – Genae narrow, near linear (Fig. 498). Legs yel- ing stripes covered with yellow setulae. Scutellum lowordarkened...... 3 elongate triangular, yellow, apical setae longer than 3 Legsentirelyyellow...... 4 scutellum and approximated. Pleura smooth shin- ing with black marks. npl 1 + 2. Legs and halteres – Legs more or less dark, at most all femora yellow. Body length 2.0–2.5 mm. black...... 5 4 Body with distinct greenish lustre, especially Distribution. – Rare. Only Russia: Kr: Kivach Nature on abdomen. Scutum not flattened before Reserve, Malaise trap, 1 female (Nartshuk, 1999). scutellum, covered with brown setulae. Male European species, known from Austria and Hun- genitalia Figs 494–496 gary...... R. anthracinum (Meigen) Biology. – Life habits of this species unknown, – Body black, without metallic luster. Scutum other species of the genus are carnivorous, larvae flattened before scutellum. Male genitalia Figs develop in egg sack of Aranea. 500–502...... R. atricorne (Zetterstedt) 5 Wings brownish (Fig. 504). Legs including all Genus Rhopalopterum Duda, 1929 femoraandmosttibiaeblack...... 6 – Wings transparent not brownish. Front tibiae Rhopalopterum Duda, 1929: 167. Type species: Os- yellow, hind tibiae and sometimes middle tib- cinella limitata Becker, 1912 by monotypy. iae with black band. Postpronota and noto- pleural depression dusted. Male genitalia Figs Black, shining species with elongated thorax and 508–510...... R. fasciola (Meigen) relatively long wings. Frons silky shining, ocellar triangle large shining or in thin dust. Genae usu- 6 Ocellar triangle with small black metallic shin- ally linear, vibrissal angle straight. poc upright, cru- ing half-moon-shaped spot before front ocel- ciate. First flagellomere rounded, wider than long. lus. Scutum dusted at sides. No hairy spot Scutum rather flat, sometimes distinctly flattened on inner surface of epandrium. Hypandrium behind, shining or thinly dusted and evenly co- open. Male genitalia Figs 505–507 vered with setulae. Thorax sometimes with metal- ....R. brunneipenne Beschovski & Lansbury lic green lustre. Pleura smooth, mainly shining. npl – Ocellar triangle shining without metallic spot. 1 + 1. Tibial organ oval. Femoral organ composed Postpronota and greater part of notopleural of two indistinct rows of warts. Male genitalia: cerci and surstyli simple, hypandrium open, postgonites depression shining without dust. Hairy spot usually with sclerotized projection. present on inner surface of epandrium. Hy- pandrium closed. Male genitalia Figs 511–513 Distribution. – Holarctic and Neotropical Regions...... R. femorale Collin

Key to species of Rhopalopterum Duda 97. Rhopalopterum anthracinum 1 Scutum and ocellar triangle shining, without (Meigen, 1830) Figs 494–496 dusting, scutellum covered with thin dust ...... 2 Chlorops anthracinus Meigen, 1830: 156; Meigen’s – Scutum and ocellar triangle subshining, co- plate 216: 14 (Morge, 1976b). Lectotype in vered with rather thin dust, scutellum ticker NHMW (Nartshuk, 1997a).

141 Figs 493–502. Pseudogaurax venustus Czerny: 493: thorax, dorsal view. Rhopalopterum anthracinum (Meigen): 494: hypopygium, dorsal view. 495: surstylus, lateral view. 496: hypandrium and phallic complex. R. atricorne (Zetterstedt): 497: head, dorsal view. 498: head, lateral view. 499: wing. 500: hypopygium, dorsal view. 501: surstylus, lateral view. 502: hypandrium and phallic complex.

Body elongated, black, shining with green lustre, 98. Rhopalopterum atricillum especially on abdomen. Frons black, silky shining, (Zetterstedt, 1838) ocellar triangle shining, extending 3/4 length of frons or longer, genae linear. Antennae and palpi black. Scutum shining, evenly covered with brown Oscinis atricilla Zetterstedt, 1838: 781. Lectotype in setulae. Notopleural depression dusted. Scutellum ZMLU (Andersson, 1966). rounded, dusted. Legs entirely and halteres yellow. Body black, elongated. Frons dull, ocellar triangle Body length 1.8–2.0 mm. shining, reaching 3/4 length of frons. Antennae and palpi black. Genae wider than front tibia. Scu- Distribution. – Denmark:F,NWZ,NEZ,B.Sweden: tum shining, evenly covered with pale setulae, but north to HS. Finland: north to Ob. Russia: Vib, Kr. extensively dusted behind and especially at sides. Not recorded yet in Norway. Legs entirely darkened, only basitarsi of all legs yel- Eurasian boreal species. lowish. Halteres yellow. Body length 1.5–1.8 mm. Distribution. – Sweden: north to TO. Norway: HOi, Biology. – Adults in June to August in wetlands Nsy, TRi. Finland: north to Li. Russia: Kr, Lr. Not so with Carex spp., which are known as host plants far found in Denmark. (Nartshuk, 1972a). Tyrphophilous species. Eurasian boreal species.

142 Biology. – Adults from June to early September in 101. Rhopalopterum fasciola wetlands with Carex spp., which probably repre- (Meigen, 1830) sent the host plants. Tyrphobiont species. Figs 508–510 Chlorops fasciolus Meigen, 1830: 154; Meigen’s plate 99. Rhopalopterum atricorne 237: 9 (Morge, 1976b). Lectotype in NHMW (Zetterstedt, 1838) (Nartshuk, 1997a). Figs 497–502 Body elongated, black, shining. Frons dull, ocel- Oscinis atricornis Zetterstedt, 1838: 780. Lectotype lar triangle shining, extending 3/4 length of frons, in ZMLU (Andersson, 1966). genae almost linear. Antennae black. Postpronota Lioscinella platythorax Nartshuk, 1958: 99. and notopleural depression dusted. Legs with fem- (Nartshuk & Andersson, 2003). ora black, hind tibiae and sometimes middle with black band. Body length 1.8–2.0 mm. Body elongated, black, shining. Frons dull black, velvety; ocellar triangle shining, extending 3/4 Distribution. – Denmark: EJ, NEZ, B. Sweden: north length of frons, genae linear, but a little wider than toHS,alsoNB.Finland:Ab,Ta,Kb,Om.Russia:Vib, in R. anthracinum. Antennae and palpi black. Scu- Kr. Not so far found in Norway. tum shining, evenly covered with brown setulae, Biology. – Adults in June to August in wetlands with distinctly flattened in hind part. Postpronota and Carex. Probably species of Carex are host plants. notopleural depression dusted. Scutellum rounded dusted. Legs entirely yellow. Halteres yellow. Body length 1.8–2.0 mm. 102. Rhopalopterum femorale Distribution. – Sweden: north to TO. Finland: Ab, N, Collin, 1946 Ta, Sb. Russia: Vib, Kr. Not so far found in Norway Figs 511–513 and Denmark. Lioscinella femoralis Collin, 1946: 145. Lectotype Eurasian boreal species. male and 6 males, 8 females paralectotypes Biology. – Adults in June to August in wetlands with in OXUM (Beschovski & Lansbury, 1987; Pont, Carex spp. Probably species of Carex are host plants 1995). (Nartshuk, 1972a). The species is more abundant Body elongated, black, shining. Frons dull, ocel- than R. anthracinum. lar triangle shining, extending 3/4 length of frons, genae almost linear. Antennae black. Postpronota 100. Rhopalopterum brunneipenne and greater part of notopleural depression shin- ing, without dust. Wings faintly brownish. Legs Beschovski & Lansbury, 1987 with femora black, all tibiae with black band. Body Figs 503–507 length 1.8–2.0 mm. Rhopalopterum brunneipenne Beschovski & Lans- Distribution. – Sweden: north to TO. Finland: north bury, 1987: 95. Holotype male in OXUM (Pont, to Lk. Russia: Vib, Kr. Not so far found in Norway 1995). and Denmark. Eurasian boreal species, known eastwards to Frons a little longer than wide. Ocellar triangle Mongolia. extending 2/3 length of frons, shining with small black metallic shining half-moon-shaped spot. Biology. – Adults in June to August in wetlands with Genae linear. First flagellomere rounded, palpi Carex spp. Probably species of Carex are host plants black. Scutum before scutellum, notopleura, up- (Nartshuk, 1972a). Tyrphophilous species. per part of anepisternum and scutellum dusted. Postpronota shining. Legs mainly black. Wings dis- tinctly brownish, especially in the anterior half. 103. Rhopalopterum tomentosum Body length 2.0 mm. Nartshuk & Andersson, sp. nov. Figs 514–516 Distribution. – Finland: Ab, Sb. Russia: Kr. Described from England. Elsewhere only known Holotype male, Sweden: SK: Åsljunga, 05.vi.1960 from NW Russia and Lithuania. (H. Andersson). Paratypes 6 males and 6 females

143 Figs 503–510. Rhopalopterum brunneipenne Beschovski & Lansbury: 503: head, dorsal view. 504: wing. 505: hypopygium, dorsal view. 506: hypandrium and phallapodeme, ventral view. 507: postgonite. R. fasciola (Meigen): 508: hypopygium, dorsal view. 509: hypopygium, ventral view. 510: hypandrium and phallic complex. (503, 504, 506, 507 after Beschovski & Lansbury, 1987.)

Figs 511–516. Rhopalopterum femorale (Collin): 511: hypopygium, dorsal view. 512: surstylus, lateral view. 513: hypandrium and phallic complex. R. tomentosum Nartshuk & Andersson: 514: wing. 515: hypopygium, dorsal view. 516: hypandrium and phallic complex.

144 with same data. Holotype and 10 paratypes in shining. poc upright convergent. Scutellum rather ZMLU, 2 paratypes in ZISP. long triangular, apical setae on small tubercles. npl 1 + 2. Tibial organ distinct oval. Femoral organ con- Diffferential diagnosis. – Differs from other species of the genus by thin dust on ocellar triangle and sists of a dense group of long setulae. Male geni- thorax and structure of epandrium. talia: cerci close together, surstyli simple. Hypan- drium closed. Description. – Male, female. Body black, subshin- ing with rather thin dust. Head wider than thorax. Distribution. – Probably worldwide, but not known Frons as long as wide, narrowed anteriorly, dull. in the Neotropical and Australasian Regions. Front margin of frons straight. Ocellar triangle ex- tending 2/3 length of frons, subshining, covered with thin dust as in Incertella zuercheri.Headsetae 104. Siphonella oscinina rather long, 5 thin orb, vte little longer than vti and (Fallén, 1820) poc. Genae a little wider than anterior tibiae and Figs12,13,27,28,59,517–521 widened posteriad. Vibrissal angle rounded with short vi. Antennae black, first flagellomere small Madiza oscinina Fallén, 1820: 6. Lectotype in ZMLU and rounded, arista with rather long pubescence. (Andersson, 1963). Palpi short black. Scutum longer than wide, covered with thin dust. Body black, shining. Ocellar triangle narrow, ex- Scutellum short, semicircular, thicker dusted than tending half length of frons. orb numerous. Face scutum. Apical scutellar setae longer than scutel- with narrowly triangular carina above, which after lum. Two pairs of subapical setae. Pleura shin- narrowing below and widening again reaches fa- ing. npl 1 + 1,butsomefemalehavesecondthin cial margin. Vibrissal angle strongly produced, pro- posterior seta. Wings brownish. Costal sectors as boscis long. Palpi long cylindrical yellowish. First 30:25:18:10. Halteres yellow. Male genitalia: cerci flagellomere rounded, wider than long, rather thick more or less transversal. Hypandrium open. Body below, yellow, darkened above. Scutum coarsely length 1 mm. punctate, evenly covered with numerous short se- tulae arising from stellate granulae. Scutellum with Note. – The new species is provisionally placed in the same microsculpture. Pleura smooth shining, the genus Rhopalopterum, even though it shares only anepisternum wrinkled and dusted above. some characters in dusting of ocellar triangle and Halteres brown. Body length 2.0–3.0 mm. scutum with black Incertella species. It differs from species of Incertella by the longer and less con- Distribution. – Denmark: SJ, EJ, WJ, NEJ, LFM, NEZ. vex scutum. Also the shape of the hypandrium and Sweden: north to TO. Norway: Ø, AK, HEs, Bø, VE, epandrium is more like in other species assigned to VAy,Ry,SFi,STi,Fn.Finland:northtoLi.Russia:Vib, the genus Rhopalopterum. The new species belongs Kr, Lr. to a complex of small, entirely black species which Holarctic species. are rather difficult to distinguish. There exist some other, probably new species within this group in Biology. – Adults autumnal, from the end of July to our material. We refrain however from describing September. more species at this moment, as more material is Data on bionomics of the species controversial. needed to assess character variation and and habi- Coquillett (1898) reported that from egg cocoon of tat preferences. Aranea (species not named) were reared 4 spec- imens of S. oscinina. Nartshuk (1962a) proposed that larvae feed on coccids Greenisca glyceriae Genus Siphonella Macquart, 1835 (Green), living on Agrostis vulgaris.NotebyDuda (1932–1933) that Schwangart (1906) reported par- asitism of larvae in egg cocoon of Aranea virgata Siphonella Macquart, 1835: 584. Type species: Hahn is not correct. Schwangart not exactly named Madiza oscinina Fallén, 1820 by original desig- the species, but considered it as species related to S. nation. oscinina. Probably Schwangart’s note refers to Lasi- Black, shining species with strongly produced vib- ambia coxalis or L. palposa. Adults were found on rissal angle, distinct high facial carina and genic- Common stinkhorn, Phallus impudicis in Finland ulate proboscis with long labella. Ocellar triangle (Luther, 1946; Smith, 1956).

145 Figs 517–521. Siphonella oscinina (Fallén): 517: head, dorsal view. 518: scutellum. 519: wing. 520: hypopygium, ventral view. 521: hypan- drium and phallic complex.

Genus Siphunculina Rondani, 1856 – Body grey pollinose, all setae and setulae of body whitish. Male genitalia Figs 527–529 Siphunculina Rondani, 1856: 128. Type species: Si- ...... S. nidicola Nartshuk phunculina brevinervis Rondani, 1856 (= Si- phonella aenea Macquart, 1835) by original des- ignation. 105. Siphunculina aenea Black, shining species or grey pollinose with short (Macquart, 1835) Figs 26, 39, 522–526 vein R2+3. Ocellar triangle shining or dusted. Vib- rissal angle produced. poc upright convergent. Face Siphonella aenea Macquart, 1835: 585. Types prob- with carina completely dividing antennal foveae. ably in MHNL. Tibial organ small oval. Femoral organ absent. The Diastata albinervis Roser, 1840. Lectotype in SMNS genus has been associated with dung and birds (Nartshuk, 1994). nest. Body black. Frons black and evenly dusted, head se- Distribution. –Worldwide. tae black and short. Ocellar triangle black, shining, reaching margin of frons, with a shallow median furrow. Antennae yellow, first flagellomere more Key to species of Siphunculina or less darkened, wider than long, thick basally. Rondani Facial carina broadly triangular between anten- nae, then narrowing and again broadening before 1 Body shiny black, all setae and setulae of body facial margin. Genae slightly narrower than first black. Male genitalia Figs 525, 526 flagellomere. Vibrissal angle weakly produced. Scu- ...... S. aenea (Macquart) tum shining with fine setiferous punctuations, de-

146 Figs 522–529. Siphunculina aenea (Macquart): 522: head, dorsal view. 523: head, lateral view. 524: scutellum. 525: hypopygium, dorsal view. 526: hypandrium and phallic complex. S. nidicola Nartshuk: 527: hypopygium, dorsal view. 528: posterior end of abdomen, lateral view. 529: male genitalia, ventral view. pressed in hind half. Pleura black, shining. Scutel- Body black grey pollinose. Frons black, ocellar tri- lum rather short, apical and two pairs of subapical angle shining, dusted on ocellar tubercle only, ex- setae very short and stout. Femora black, tibiae yel- tending 2/3–3/4 length of frons, with a shallow me- low with black band. Halteres black. Body length dian furrow. Head setae and setulae whitish. An- 1.5–2.0 mm. tennae and palpi yellow. Arista short pubescent, nearly bare. Genae slightly narrower than first flag- Distribution. – Sweden: HA, NÄ, DR. Norway: AK: ellomere. Vibrissal angle weakly produced. Scutum Konglungen, Asker, 4.vi.1997 (G.E.E. Söli). Finland: dull and with dispersed short whitish setulae, se- northtoOm.Russia:Vib.NotfoundinDenmark. tae of thorax whitish. Pleura black, shining except Widely distributed species in the Palearctic Re- on upper part of anepisternum. Notopleural setae gion eastwards to Japan. 1 + 1. Scutellum trapezoid, apical setae little shorter Biology. – Adults from May to August, mainly in than scutellum. subapical setae very close to api- cals. Legs yellow, femora and tibiae of hind legs woodland, often on flowers. Also found in a swal- slightly darkened. Femora black, tibiae yellow with low’s nest (Krivokhatsky & Nartshuk, 2001). black band. Halteres brown. Body length 2.0 mm. It has been reared from faeces of wild brown bear and cow dung in Japan (Kanmiya, 1982, 1989). Distribution. – Not yet found in Denmark and Fennoscandia, but expected in Finland. Eurasian species described from Russia: Lenin- 106. Siphunculina nidicola grad Oblast and Mongolia. Nartshuk, 1971 Biology. – In Mongolia bread from a kite’s (Milvus) Figs 527–529 nest. Siphunculina nidicola Nartshuk, 1971: 290.

147 Genus Speccafrons Sabrosky, 1980

Speccafrons Sabrosky, 1980: 424. Type species: Os- cinella mallochi Sabrosky, 1938 by original des- ignation. Body black, shining or dusted. Eyes densely pubes- cent. Frons with numerous small bare spots scat- tered over the finely tomentose surface. Vibrissal angle rounded, not projected in advance of eye. First flagellomere suborbicular. npl 1 + 2. Tibial organ present. Femoral organ consists of a row of warts bearing short spines. Halteres brown- ish black. Male genitalia with large semiglobose surstyli, cerci discrete. Distribution. –Holarctic.

107. Speccafrons halophila (Duda, 1933) Figs 530–534 Conioscinella halophila Duda, 1933: 53. Syntypes in NHMW, HNHM and NHMH. Figs 530–534. Speccafrons halophila (Duda): 530: head, lateral Body black grey-dusted. Frons dull, brownish black view. 531: antenna. 532: scutellum. 533: hypopygium, dorsal behind and reddish yellow in front. Setulae of view. 534: hypandrium and phallic complex. frons arise from shining punctures. Ocellar tri- angle small, extending half length of frons, black grey-dusted. Eyes densely haired. Antennae yel- Collin (1946) found it identical with the present low brownish, first flagellomere wider than long. species. Genae as wide as first flagellomere or narrower. Palpi rather thick and long, yellowish. Scutum with setulae not arranged in rows. Scutellum semicircu- Genus Trachysiphonella Enderlein, lar, with numerous setae on hind margin. Pleura 1936 mainly shining. npl 1 + 2. Legs mainly black, but front and middle tibiae yellow at ends. Body length 1.8–2.0 mm. Trachysiphonella Enderlein, 1936: 187. Type species: Oscinis pumilio auct. nec Zetterstedt, 1848 Distribution. – Denmark: SZ, NEZ, B. Sweden: north (= Chlorops scutellatus Roser, 1840) by mono- to LU. Finland: Ab, N, Sb. Russia: Vib, Kr. Not so far typy. found in Norway. Yellow species with 3–5 black or red stripes on scu- Trans-Palearctic species, known from the British tum and black marks on pleura similar in colour to Isles to Russian Far East. those of Chlorops species. Frons dull with dusted Biology. – Adults in June to September, in meadows small ocellar triangle. Vibrissal angle produced. and wetlands with Phragmites australis. Proboscis prolonged. Face with distinct narrow ca- Larvae carnivorous, developing in egg sacks of rina. npl 1 + 2. Tibial organ distinct. Femoral organ the spiders Larinioides sericatus and Singa nitidula present as oval group of warts. Male genitalia: cerci (Schwangart, 1906; Pereleshina, 1928, as “Coni- and surstyli simple, hypandrium open, postgonites oscinella frontella”; Séguy, 1934 and Vachon, 1952, with sclerotized projections. as “Oscinisoma frontellum”; Nartshuk, 1972b). Distribution. – The Palearctic Region. Schwangart (1906) determined the flies reared from eggssacksasaspeciesnearSiphonella oscinina,but

148 Key to species of Trachysiphonella 109. Trachysiphonella scutellata Enderlein (Roser, 1840) Figs 60, 539–546 1 Body larger, reaching 2.0 mm. Stripes on scu- tum black sometimes fused. Pleura with four Chlorops scutellatus Roser, 1840: 63. Syntypes in black marks. Male genitalia Figs 544–546 SMNS (Andersson, 1966; Nartshuk, 1994)...... T. scutellata (Roser) Body yellow with black marks. Frons yellow and – Body smaller, reaching 1.5 mm. Stripes on scu- ocellar triangle very small, extending no more tum black or red. Pleura only with one black than 1/3 length of frons, black and dusted. Anten- or red mark. Male genitalia Fig. 538 nae and palpi yellow, first flagellomere darkened above. Genae distinctly produced and proboscis ...... T. rufijiceps (Macquart) elongated. Scutum yellow with black longitudinal stripes, sometimes fused in dark coloured speci- 108. Trachysiphonella rufijiceps mens. Scutellum yellow or darkened. Pleura with 4 black spots. Legs mainly yellow. Halteres yellow. (Macquart, 1835) Body length 1.8–2.0 mm. Figs 535–538 Distribution. – Denmark: LFM, NEZ. Sweden: north Siphonella rufijiceps Macquart, 1835: 586. Types to LU. Norway: AK: Oslofjord: Lindøya; Hovedøya; probably in MHNL. Baerum; Oksenøya; Storøykilen. Finland: north to Chlorops pygmaeus Meigen, 1838: 385, syn. n. Lec- Lk. Russia: Vib; Kr: 3 km SW Konchozero, totype, male, in MNHN (Nartshuk, 2006). 15.vii.2002 (Polevoi); Kr: Kivach, 04.vii.2001 and 03.ix.2002 (Polevoi). Body yellow with black to red marks. Frons yellow Eurasian species known from Europe eastwards and ocellar triangle very small, extending no more to Yakutia, East Siberia and Mongolia and south- than 1/3 length of frons, blackish and dusted. An- wards to Israel. tennae and palpi yellow. Genae produced to dif- ferent degree and proboscis elongated correspond- Biology. – Adults in June to July in meadows. ingly. Scutum yellow with black to red longitudi- nal stripes. Scutellum yellow. Pleura with only one black spot. Legs and halteres yellow. Body length Genus Tricimba Lioy, 1864 1.5 mm. Distribution. – Denmark: SJ, NEZ. Sweden: north to Tricimba Lioy, 1864: 1125. Type species: Oscinis li- VS. Finland: Al, Ab, N. Russia: Vib. Not so far found neella Fallén, 1820: 8 by designation Enderlein, in Norway. 1911: 207. Eurasian species. Nartshukiella Beschovski, 1981c: 119, as subgenus of Tricimba Lioy, 1864. Type species: Chlorops Biology. – Adults in June to August in meadows. cincta Meigen, 1830 by original designation. Note. – The present species is very variable in Black densely dusted species with 3–5 longitu- colour, projection of genae and correlated length dinal punctuate lines on scutum. Ocellar trian- of proboscis, but we were unable to find distinct gle dusted. Vibrissal angle produced. poc upright differences in the structure of male genitalia sup- convergent. First flagellomere rounded. npl usu- porting a subdivision into several species. There- ally 1 + 2. Scutellum sometimes thick with convex fore, we only recognized two species in our large side margins, and scutellar apicals stout. Tibial or- Palearctic material based on size, number of se- gan oval. Femoral organ consists of two rows of tulae on scutellum and distance between apical warts or absent. Male genitalia: cerci separated or setae on scutellum (Figs 537, 542): the larger Tra- more or less fused, hypandrium closed or open, chysiphonella scutellata (Roser) and the smaller T. pre- and postgonites indistinctly separated. Female rufijiceps (Macquart, 1835). Oscinis flavella Zetterst- with ovipositor membranous. edt, 1848, and Siphonella diplotoxoides Strobl, 1893 European species of the genus Tricimba are re- are both considered as likely synonyms of T. rufiji- ferred to 2 subgenera: Tricimba s. str. having mar- ceps (Macquart, 1835). ginal scutellar setae short, located at the same

149 Figs 535–546. Trachysiphonella rufijiceps (Macquart): 535: head, dorsal view. 536: head, lateral view. 537: scutellum. 538: hypopygium, dorsal view. T. scutellata (Roser): 539: head, dorsal view. 540: head, lateral view. 541: face and antennae, anterior view. 542: scutellum. 543: wing. 544: hypopygium, dorsal view. 545: surstylus, medial view. 546: hypandrium and phallic complex.

level as the lower surface of scutellum with tuber- Key to species of Tricimba Lioy cles at the base, cell br widened, male cerci fused at base, and hypandrium open; and Nartshukiella Beschovski having marginal scutellar setae located at the same level as upper surface of scutellum with 1 Ocellar triangle with shining spot before an- no tubercles at the base, cell br not widened, male terior ocellus. Postpronota yellow. Male geni- cerci separated, hypandrium closed. A third sub- talia Figs 552, 553 . . . T. (N.) humeralis (Loew) genus, Schumaniella Beschovski, 1981, is confined – Ocellar triangle entirely dull without shining to North Africa and the Near East. spot. Postpronota coloured as remaining parts Distribution. –Worldwide. ofthorax...... 2

150 2 Three faintly impressed punctuate lines on and Clethrionomys in Finland. Hibernate as adults. scutum. Scutellum flat and apical setae not It has been recorded many times indoors in the au- stout, scutellum often yellow apically (Fig. tumn together with Thaumatomyia notata.Adults 548). Male genitalia Figs 549, 550 were recorded also from birds’ nests (Hicks, 1953; ...... T. (N.) cincta (Meigen) Krivokhatsky & Nartshuk, 2001). The larvae are exceptionally diverse in feeding – Five deeply impressed punctuate lines on scu- habits. They have been reared from stems of cereals tum. Scutellum and apical setae stout, scutel- infested by other insects by many authors, includ- lum coloured as remaining parts of thorax ing seedlings of wheat in Norway (Rygg, 1966), from (Figs 556, 557). Cell br widened in the mid- rotting spathes of skunk cabbage (Symplocarpus dle (Fig. 558). Male genitalia Figs 560, 561 foetidus), from various mushrooms (Boletaceae, ...... T. (T.) lineella (Fallén) Agaricaceae, Russulaceae), from rotting bark of Alnus, berries of Sambucus racemosa,deadmol- lusks and meat baits (Brown, 1956; Myhályi, 1965; 110. Tricimba (Nartshukiella) cincta Dely-Draskovits, 1972; Krivosheina, 1974; Schatz- (Meigen, 1830) man, 1977; Grimaldi & Jennike, 1983; Krivosheina Figs 61, 547–550 et al., 1986; Tschirnhaus, 1992; Yakovlev, 1994). Anterior and posterior spiracles of larvae are fig- Chlorops cinctus Meigen, 1830: 162; Meigen’s plate ured by Krivosheina (2005). 227:12(Morge,1976b).LectotypeinNHMW (Nartshuk, 1997a). Note. – A lectotype designation for Chlorops cinc- Tricimba fungicola Dely-Draskovits, 1983: 349 tus Meigen by Dely-Draskovits (1983) is invalid, (Nartshuk, 1997a). because she designated a specimen that cannot belong to the syntypes series, as labeled “Madrid Body black grey-dusted. Frons mainly yellow, in- 7. August”. Meigen stated “Im September auf der fuscate behind. Ocellar triangle black grey-dusted, Blüthe der Möhren selten”. According to Pont extending halfway to front margin of frons. An- (1986) this indicates specimens collected in Stol- tennae yellow, first flagellomere orbicular, slightly berg near Achen, Germany. darkened apically. Head setae from yellow to black. Genae yellow, narrower than first flagellomere. Vib- rissal angle weakly produced. Palpi yellow. Scu- 111. Tricimba (Nartshukiella) tum densely grey-dusted with 3 longitudinal sulci humeralis of punctuations along acrostichal and dorsocen- tral lines. Scutellum somewhat conical, slightly (Loew, 1858) wider than long, distinctly flattened, usually with Figs 551–553 yellow tip, but specimens from N Fennoscandia have no yellow tip. Apical scutellar setae close-set, Oscinis humeralis Loew, 1858: 59. Syntypes in shorter than scutellum, three pairs of subapical se- ZMHU (Wendt, 1975). tae. Scutellar setae from yellow to black. Pleura Body black grey-dusted. Frons mainly yellow with mainly shining. npl 1 + 2. Legs in males mainly yellow setae and setulae. Ocellar triangle black yellow, in females femora darkened. Body length grey-dusted, extending half length of frons with 1.5–2.0 mm. shining spot in apical part between front ocellus Distribution. – Denmark: from all districts except and tip of triangle. Antennae yellow, first flagellom- SJ, WJ and SZ. Sweden: north to TO. Norway: AK, ere orbicular, slightly darkened apically. Arista yel- HEs,Bø,AAy,STi.Finland:northtoLi.Russia:Vib, low at base. Genae yellow, as wide as first flagellom- Kr, Lr. ere. Vibrissal angle weakly produced. Palpi yellow. Holarctic species. Scutum black with postpronota yellow, densely grey-dusted with 3 longitudinal sulci of punctu- Biology. – Adults from June to September in mead- ations along acrostichal and dorsocentral lines. ows, in forests, from canopies of Pinus silvestris in Scutellum slightly wider than long with yellow tip. Norway (Thunes et al., 2004), and in agricultural Apical scutellar setae as long as scutellum. Pleura fields in Denmark (Nielsen & Nielsen, 1984). Hack- shining below. npl 1 + 2. Abdomen yellow at base. man (1963) found adults in borrows of Microtus Legs mainly yellow, but hind femora and tibiae

151 Figs 547–553. Tricimba cincta (Meigen): 547: head, lateral view. 548: scutellum. 549: hypopygium, dorsal view. 550: hypandrium and phallic complex. T. humeralis (Loew): 551: scutellum. 552: hypopygium, dorsal view. 553: hypandrium and phallic complex.

with black band. Halteres yellow. Body length 1.5– Body black grey-dusted, except shining lower part 2.0 mm. of pleura. Frons yellow in front, infuscate behind. Ocellar triangle thick grey-dusted, nearly extend- Distribution. – Rare. Sweden: SK: Alnarp, 26.vi.1960 ing half length of frons. Antennae yellow, first flag- (H. Andersson), 1 male. Norway: 7 specimens in ellomere orbicular, yellow with small dark spot at NHMW labelled “Norway”. aristal base. Arista dark. Head setae short brown. Widely distributed species in the southern Pale- Genae yellow, about half as wide as first flagellom- arctic and Afrotropical Regions. ere. Vibrissal angle weakly produced. Palpi yellow. Biology. – Hibernate as adults. In Germany re- Scutum densely grey-dusted with 5 longitudinal corded many times inside buildings together with sulci of punctuations along acrostichal and dor- Thaumatomyia notata Meigen (Tschirnhaus, 1992). socentral lines and laterally from the latter ones. Also found in birds’ nests (Turdus philomelos and Scutellum somewhat conical, slightly wider than Hirundo rustica) (Zuska, 1966; Krivokhatsky & long, very thick, with scutellar setae arising on Nartshuk, 2001). Some specimens were reared from lower surface. Apical scutellar setae shorter than a disposal tip (Werner, 1997). scutellum, two pairs of subapical setae, all stout. Pleura mainly shining. npl 1 + 2. Legs mainly yel- low, middle and posterior femora and tibiae with 112. Tricimba (Tricimba) lineella dark strikes. Body length 1.1–1.5 mm. (Fallén, 1820) Figs 554–561 Distribution. – Denmark: SJ, NEZ. Sweden: north to UP. Norway: AK: Hovedøya, Oslofjord. Finland: Al. Oscinis lineella Fallén, 1820: 8. Lectotype in NRMS Holarctic species, known in the Palearctic Region (Andersson, 1963). from Europe to Japan and Israel.

152 Figs 554–561. Tricimba lineella (Fallén): 554: head, dorsal view. 555: head, lateral view. 556: scutellum, dorsal view. 557: scutellum, lateral view. 558: wing. 559: apical end of middle femur with appendage (piece of lateral wall of femur removed). 560: hypopygium, dorsal view. 561: hypandrium and phallic complex.

Biology. – Adults from the end of April to August, ocybe bresadolae, Russula pectinata, Xerocomus mainly in forests, often on flowers, but also in parks chrysenteron, Leccinum calopus, Paxillus involutus, in cities (Allen, 1981b). Adults hibernate in old Lycoperdon gemmatum, and probably Langerman- birds’ nests in America (Malloch, 1934, as T. spinig- nia gigantea (Falcoz, 1930; Dely-Draskowits, 1972; era). Allen, 1981b; Krivosheina et al., 1986; Yakovlev, The larvae develop in many different fungi: Can- 1994). Also from rotting spathes of skunk cabbage tharellus cibarius, Calocybe gambosa, Collybia Symplocarpus foetidus (Araceae) in North America dryophila, Amanita verna, Cortinarius uraceus, In- (Brown, 1956).

153 Figs 562–568. Camarota curvipennis Latreille: 562: head, dorsal view. 563: head, lateral view. 564: thorax, lateral view. 565: scutellum. 566: wing. 567: hypopygium, dorsal view. 568: male genitalia, ventral view.

Subfamily CHLOROPINAE

Genus Camarota Meigen, 1830 113. Camarota curvipennis (Latreille, 1805) Figs 42, 562–568, 956–959 Camarota Meigen, 1830: 7. Type species: Camarota flavitarsis Meigen, 1830 (= Oscinis curvipennis Oscinis curvipennis Latreille, 1805: 383. Syntype, 1 Latreille, 1805) by monotypy. specimen, in MNHN (Ismay, 1999). Black species with triangular head, thick arista, Body black with partly yellow head and legs. Head large ocellar triangle and curved wings with thick longer than wide with very short setae. Eyes oval veins, vein R2+3 fused with R1 and costa. Frons dis- with horizontal long axis. Antennae mainly black, tinctly projected, face with narrow carina. In eyes but first flagellomere with yellow spot around arista anterior ommatidia larger than posterior. First flag- insertion. Arista broad lanceolate black. First flag- ellomere trapezoid. Tibial organ present. Male gen- ellomere 1.5 times longer than wide. Genae broad italia: mesolobus short wide, middle lobe of sursty- with black spot. Ocellar triangle black reaching lus with hook-like apical end. Gonites divided into margin of frons, with a shallow depression in front post- and pregonites. of ocellar tubercle. oc short. vte and vti indistinct. Palpi yellow. Scutum black subshining with coarse Distribution. – Palearctic and Afrotropical Regions. microsculpture, covered with pale setulae. Scutel-

154 lum rounded wider than long. Apical setae short Tibial organ present. Intra-epandrial sclerite well convergent, situated on the middle of the disk. developed. Mesolobus large. Surstyli with distinct Pleura mainly dusted and smooth. Anepisternum upper lobe and hook-like middle lobe. Gonites di- with shining wrinkled area below. Katepisternum vided into pre- and postgonites. shining at middle and in front below, remaining part silvery dusted. Wings brownish, convex, and Distribution. – Palearctic Region, most species are folded over the abdomen at rest and give the fly found in eastern part. a beetle-like appearance. Halteres black. Legs with femora black, tibiae darkened in the middle. Tibial 114. Centorisoma elegantulum organ narrow oval. Body length 2.5–3.0 mm. Becker, 1910 Distribution. – Nartshuk have seen two females in Figs 24, 569–575 MNHN labeled “Denmark”. We have not seen other material from the territory investigated. Duda Centorisoma elegantulum Becker, 1910: 107. Syn- (1933) mentioned specimens from Finland and types in HNHM. Sweden, but Nartshuk did not find specimens in Body black with partly yellow head and legs. Ocel- NHMH and NRMS, and there are no specimens in lar triangle large, behind broad as frons, rounded ZMLU. off at apex, reaching margin of frons, black, shin- There is only one species of the genus in the ing. Head setae small. First flagellomere 1.5 times Palearctic Region. The species is known from all as long as wide. Arista thick white haired. Genae Europe except for the northern parts, and from as wide as first flagellomere, with black spot. Face the Caucasus, southern part of Palearctic Asia and with black spot. Scutum with coarse granulate North Africa. sculpture, dusted behind. Scutellum dusted. Api- Biology. – Larvae phytophagous, develop in shoots cal scutellar setae short and rather far apart. Legs and ears of different cereals: barley,oats, rye, wheat, yellow, middle tibiae with rather long black spur. but also on Elytrigria repens and possible other Halteres yellow. Body length 2.0–2.5 mm. grasses. Four or five larvae may live in a single Distribution. – Only from Vib: Bolshoy Beresovy Is., stem. Infestation of young shoots results in arrested Vyborg (Nartshuk, 1984a). growth and no development of ears. The species Eurasian species, known from Europe and West is considered as a pest of cereals in some coun- Siberia eastwards to the Baikal Lake. tries of southern Europe (Italy, France, former Yu- goslavia, Bulgaria) (Rondani, 1873; Marchal, 1894, Biology. – Rather scarce. Adults in dry meadows 1897; Osten-Sacken, 1896; Balachowsky & Mes- and on sandy beaches, often caught on Elytrigia nil, 1935; Lyubenov, 1960; Grujičić, 1961; Zamfirov, repens. 1962; Krusteva & Beschovski, 1998, 2000). Larvae Larvae develop in shoots of Elytrigia repens (Kar- of Elachiptera cornuta have been reported living to- pova, 1972; Nartshuk, 1972a). gether with the present species. Larvae described by Marchal (1884), Balacho- wsky & Mesnil (1935), and description repeated by Genus Cetema Hendel, 1907 Nye (1958). Cetema Hendel, 1907: 98, replacement name for Centor Loew, 1866 (preocc. Schoenherr, 1848). Genus Centorisoma Becker, 1910 Type species: Oscinis cereris Fallén, 1820, auto- matic. Centorisoma Becker, 1910: 106. Type species: Cen- Black species with partly yellow head, pleura, scu- torisomaelegantulum Becker, 1910 by monotypy. tellum and legs. Body elongated. Ocellar triangle Body elongated, usually black with coarsely sculp- large, shining. First flagellomere rounded squarish, tured scutum and scutellum, more rarely thorax as long as wide. Arista white or black. Scutum mod- yellow and scutum smooth. First flagellomere more erately convex, coarsely sculptured. Middle tib- or less prolonged with thick usually white arista. iae with rather long spur. Front and middle tibiae Ocellar triangle large smooth shining. Face and in males sometimes with long hairs. Tibial organ genae with central dark shining sclerotized area. present. Pregenital sclerite in male well developed.

155 Figs 569–575. Centorisoma elegantulum Becker: 569: head, dorsal view. 570: head lateral view. 571: wing. 572: hypopygium, dorsal view. 573: male genitalia, lateral view. 574: male genitalia, ventral view. 575: ovipositor, dorsal view.

Male genitalia: All European species have an epan- – Anterior and middle tibiae of male with long drium with posterior lateral corners prolonged and hairs(asinFig.577)...... 3 tapering. Middle lobe of surstyli serrate at apex; 3 Male genitalia: processes of epandrium pregonites very long, postgonites with short hook at posterior end. In females sternum 5 very large. straight, long and tapering (Fig. 587). Pre- genital sclerite yellowish Distribution. –Holarctic...... C. neglectum Tonnoir – Male genitalia: processes of epandrium curved mesad (Figs 585, 586). Pregenital sclerite black Key to species of Cetema Hendel ...... C. myopinum (Loew)

1 Arista white. Anterior and middle tibiae of male with long hairs (Fig. 577). Male genitalia 115. Cetema cereris Figs578,579...... C. cereris (Fallén) (Fallén, 1820) – Arista black. Anterior and middle tibiae of Figs 576–580 malewithorwithoutlonghairs...... 2 2 Anterior and middle tibiae of male without Oscinis cereris Fallén, 1820: 3. Lectotype in ZMLU long hairs. Epandrium of male genitalia with (Andersson, 1963). straight processes (Figs 581–584) Frons almost as long as wide, ocellar triangle black, ...... C. elongatum (Meigen) shining. Genae half as wide as first flagellomere or

156 a little wider than that. First flagellomere slightly Biology. – Adults in June to August in meadows and longer than wide, mainly yellow with infuscate dor- edges of forests. Univoltine species, overwinters as sal margin. Arista white. Scutum black shagreened, larva. postpronota yellow with black spot. Pleura yellow Larvae phytophagous, develop in shoots of Poa- with black spots on all parts. Legs almost yellow ceae. Host plants: Dactylis glomerata, Holcus lana- except front tarsus and 2–3 last tarsal segments of tus, Elytrigia repens, Lolium perenne, L. multiflo- other legs. Front and middle tibiae and their ba- rum, Hordeum murinum, Agrostis tenuis, A. alba, sitarsi with long white hairs in males and black spur A. stolonifera, A. canina, Poa annua, P. pratensis, on tip. Terminal segment of male abdomen weakly P. nemoralis, P. trivialis, Phleum pratense (Bela- curved ventrad and clubbed in lateral view. Tergites chowsky & Mesnil, 1935; Nye, 1958; Karpova, 1972; 4 and 5 longer than tergite 3, synsternite 7 + 8 Nartshuk,1972a;Mowat,1975;Tschirnhaus,1981, rather large and sclerotized in male. Sternite 5 of fe- 1992; Nielsen, 1994). male abdomen enlarged. Body length 3.0–3.5 mm. Larvae described by Kreiter (1928) and Bala- chowsky & Mesnil (1935). A photo of the anterior Distribution. – Rather common species. Denmark: end of the larva is given by Sol (1974). found in most districts. Sweden: everywhere north- wards to LY. Norway: AK, HEs, HEn, Os, Bø, Bv, VE, AAy,HOy,HOi, Nnø. Finland: northwards to Li. Rus- 117. Cetema myopinum sia: Vib, Kr, Lr. (Loew, 1866) Trans-Palearctic species in boreal and temper- Figs 585, 586 ate zones, extending from Great Britain to Sakhalin (Russia) and Japan. Séguy (1934) recorded the Centor myopinus Loew, 1866: 9. Type material not species from 2000 m in Norway. found in ZMHU (Wendt, 1975, 1978). Biology. – Adults in June to August, in meadows, Body stout and in male hind end very large and forest edges and clearings. Johansson (1960) re- curved ventrally. Frons yellow, ocellar triangle shin- marked briefly on a habitat in Sweden. ing black, extending 2/3 length of frons. Arista Larvae phytophagous, develop in shoots of Poa- black. In male front and middle tibiae with long cea: Poa, Alopecurus, Festuca, Agrostis, Secale white hairs. In female the 5th sternite of abdomen (Balachowsky & Mesnil, 1935; Nye, 1958; Nartshuk, large. Setulae of abdomen pale. Body length 2.5– 1972a; Tschirnhaus, 1981, 1992). 3.0 mm. Distribution. – Denmark: SJ, WJ, ?NEZ. Sweden: 116. Cetema elongatum northwards to JÄ. Norway: HEs, Bø, Bv. Finland: northwards to Om. Russia: Vib, Kr. (Meigen, 1830) Eurasian species, known from Great Britain to Figs 581–584 the Baikal Lake and northern Mongolia. ChloropselongatusMeigen, 1830: 151; Meigen’s plate Biology. – Adults in June to August in meadows. 211: 2 (Morge, 1976b). Lectotype in NHMW Larvae phytophagous, develop in shoots of many (Nartshuk, 1997a). grasses, but prefer species of Agrostis (Nartshuk, Cetema obliqua Beschovski, 1984: 213. Holotype in 1962a). Hodson (1926) recorded the species from ZMBA. spring oats. He found several larvae living together Cetema simile Ismay, 1985: 35. Holotype in OXUM. in one shoot; they also pupated in the same shoot. Smaller with narrower body than in preceding Newly emerged adults copulated in their cage, but species. Front and middle male legs without long no eggs or larvae were found on oats shoots pre- white hairs. Arista black. Setulae of abdomen dark. sented to them. Nye (1958) considered this attack Male genitalia: epandrium small, slightly laterally due to larval migration from a newly ploughed compressed. Body length 2.0–2.5 mm. grassfield. Distribution. – Denmark: found in all districts. Swe- den: northwards to LY. Norway: northwards to Nsy. 118. Cetema neglectum Finland: northwards to Om. Russia: Vib, Kr. Tonnoir, 1921 European species. Figs 587, 588, 974–977

157 Figs 576–588. Cetema cereris (Fallén): 576: wing. 577: middle femur and tibia. 578: end of abdomen, lateral view. 579: hypopygium, dorsal view. 580: tip of ovipositor, lateral view. C. elongatum (Meigen): 581: end of abdomen, lateral view. 582: hypopygium, dorsal view. 583a, 583b: hypopygium, lateral view, two variants. 584: hypandrium and phallic complex. C. myopinum (Loew): 585: end of abdomen, lateral view. 586: hypopygium, dorsal view. C. neglectum Tonnoir: 587: end of abdomen, lateral view. 588: hypandrium and phallic complex.

158 Cetema neglectum Tonnoir, 1921: 131. Syntypes in has modified male front tarsi with shortened ba- MHNB. sitarsi and asymmetrical claws and a sclerotized aedeagus Similar to C. cereris, but arista black and epan- drium in male yellowish. Male front and middle Distribution. –Worldwide. tibiae with long white hairs. Setulae of abdomen pale. Pregenital sclerite yellowish. Body length 2.8– 3.0 mm. Key to species of Chlorops Meigen Distribution. – Denmark: from all districts except NWJ and B. Sweden: northwards to ÅN. Norway: AK:Landøya;Bø:Verksøya,Hurum.Finland:only 1 Scutum entirely black, postpronota in southern part: Al, Ab, N, Sa. Russia: Vib. black or yellow with black spots. Scutel- European species. lum sometimes black or darkened . . . . 2 Biology. – Adults in June to August in meadows. – Scutum yellow with 3–5 black or red- Univoltine species. dish stripes. Postpronota usually yellow Larvae phytophagous, develop in shoots of Poa- with black spot. Scutellum usually yel- ceae. Host plants: Lolium perenne, Festuca praten- low, very rarely with dark middle . . . . . 4 sis, Poa trivialis, P. pratensis, Agrostis stolonifera, Elymus pycnanthus. Larvae hibernate and pupate 2 (1) Basal segments of antennae yellow or the following spring. Hodson (1926) found larvae fuscous yellow. Genae wider than first in tillers of spring oats that he identified with C. flagellomere, often with black setulae. myopinum, but Collin (1966) indicated that they Yellow parts of body usually darkened belong to the present species. Probably this attack ...... 3 was due to larval migration from a newly ploughed – Antennae entirely black. Genae at most grassfield nearby (Nye, 1958). as wide as first flagellomere. Yellow parts Larvae described by Nye (1958). of body sharply contrasted with black parts. Ocellar triangle with acute apex and not reaching front margin of frons Genus Chlorops Meigen, 1830 (Fig. 673). Scutum smooth and polished. Male genitalia Figs 675, 876 Chlorops Meigen, 1830: 278. Type species: Musca ...... C. scutellaris (Zetterstedt) pumilionis Bjerkander, 1778 by designation of 3 (2) Ocellar triangle black with strongly ICZN, 1955: 423. Opinion 348. Name No 868 in demarcated outline, sometimes with Official List of Names in Zoology. windows of lighter colour outside ocelli, Yellow species usually with black or rarely with but these tend to disappear in dark spec- yellow-red stripes on scutum and spots on pleura. imens (Fig. 679). Male genitalia: outer Ocellar triangle differs in shape, usually shining, gonites short (Figs 681, 682) sometimes with ridges along lateral margins or sul- ...... C. speciosus Meigen (dark form) cus in the middle. Antennae yellow to black. First flagellomere rounded, as long as or a little longer – Border of ocellar triangle not so strongly than wide. Arista slender, black or white. Palpi outlined, the tip often yellowish, but no short or long and thick, usually yellow, rarely en- light windows on the sides of ocelli. Male tirely black or black on tip. Scutum smooth, dusted genitalia: outer gonites long (Figs 636, or shining. Stripes black or red, sometimes partly 637).....C. meigenii Loew (dark form) or completely fused. Tibial organ absent. Male gen- 4 (1) Palpi long and thick, sometimes black italia: middle lobe of surstylus with large granulate ontip...... 5 or hook like apex; lower lobe long hairy; pregonites and postgonites situated parallel to each another, – Palpi not long, yellow, sometimes black and often equal in length. on tip or entirely black, in that case A species group within Chlorops,oftenrecog- arista white and markedly pubescent nized as subgenus Sclerophallus Beschovski, 1978, ...... 9

159 5 (4) Stripes on scutum dusted. Palpi greatly – Ocellar triangle yellow or black, often darkened in both sexes (Fig. 640). Male black spot occupies only central part genitalia Figs 641, 642 of triangle leaving sides yellow. If small ...... C. nigripalpis (Duda) black spots are on ocellar tubercle and – Stripes on scutum more or less shining. on triangle then surface of triangle not Palpi yellow, darkened at tip in one sex, smooth but wrinkled along sides . . . . 12 only in male or only in female ...... 6 10 (9) Antennae and palpi black. Arista white, 6 (5) Genae not wider than first flagellomere. slightly thickened by dense pubescence. Ocellar triangle with more or less dis- Black ocellar and apical spots on ocel- tinct groove in apical part. Palpi, espe- lar triangle connected by dark line. Male cially in male, darkened at tip genitalia Figs 592, 593 ...... C. palpatus Smirnov ...... C. anthracophagoides Strobl – Basal antennal segments and palpi al- – Genae wider than first flagellomere. ways yellow. Arista black, grey or whitish Ocellar triangle without groove in api- and slender with short pubescence cal part. Palpi yellow or darkened only ...... 11 infemale...... 7 11 (10) First flagellomere yellow. Stripes on scu- 7 (6) Spot on katepisternum yellow ...... 8 tum usually red, but sometimes black, – Spot on katepisternum black. Abdomen grey-dusted. Male genitalia Figs 666, 667 yellow with distinct narrow black bands ...... C. rufijinus (Zetterstedt) on front margins of tergites 2–5 and – First flagellomere black. Stripes on scu- sometimes with black median line. First tum black, grey-dusted flagellomere from yellow to almost ...... C. planifrons Loew forma black. Stripes on scutum shining or sub- triangularis Becker shining, only thinly pollinose. Male gen- 12 (9) Genae narrower, subequal to width of italia Figs 671, 672 . . . C. scalaris Meigen first flagellomere or narrower (Figs 619, 8 (7) Ocellar triangle mainly yellow with small 639,693)...... 13 dark spot only on ocellar tubercle and – Genae broader, equal to width of first around it. Stripes on scutum shining. flagellomere or usually distinctly wider Abdomen predominantly yellow. Palpi (Fig.623)...... 17 yellow in both sexes, palpi of female longer and thicker, than in C. ringens 13 (12) Male front metatarsus long, 5 times ...... C. centromaculatus (Duda) longer than thick, front claws symmet- rical. Stripes on scutum shining and – Ocellar triangle predominantly black, abdomen predominantly yellow with yellow only along side margins and black setulae or stripes on scutum sometimes with yellow window spots. dusted and abdomen dark dorsally Stripes on scutum subshining, thinly ...... 14 pollinose. Abdomen yellow with black – Male front metatarsus short, no more bands, sometimes connected by median than 3 times longer than thick, front line. Palpi in female darkened at tip. claws asymmetrical. Stripes on scutum Male genitalia Figs 663, 664 dusted. Abdomen blackish or in male ...... C. ringens Loew pale yellow with white setae or dark- 9 (4) Ocellar triangle smooth, shining, yellow ened with black setulae, in female red- with two small black spots, one on ocel- dishbrown...... 15 lar tubercle and another on apex of ocel- 14 (13) Stripes on scutum black, shining. ab- lar triangle, sometimes black spots are domen predominantly yellow with black connectedbydarkline...... 10 setulae. Basal segments of antennae yel-

160 low. Ocellar triangle only with central inantly black. Outside margin of inner black spot, leaving sides yellow. Male lateral stripes on scutum with triangu- genitalia Figs 620, 621 lar projection behind postpronota (Fig...... C. hypostigma Meigen 665)...... C. rossicus Smirnov – Stripes on scutum black, dusted, ab- – First flagellomere yellow or darkened on domen dark dorsally. Antennae and upper and outer sides. Spot on katepis- ocellar triangle entirely black ternum yellow or brownish on upper ...... C. bjerkanderi sp. n. part. Black spot on ocellar triangle not 15 (13) Anepisternum with some small setulae, occupying apical part, side margins and which are usually black in female and hind corners of triangle. Outside mar- white in male. Ocellar triangle with yel- gin of inner lateral stripes on scutum lowsides...... 16 more or less even, without projection – Anepisternum bare without setulae. (Fig. 635). Male genitalia Figs 636, 637 Ocellar triangle entirely black. Palpi yel- ...... C. meigenii Loew low. Basal antennal segments yellow. 20 (18) Stripes on scutum shining. Ocellar tri- First flagellomere rather large, black. angle entirely black or with fuscous yel- Borders of ocellar triangle sharply out- low window spots laterally from central lined. Abdomen blackish (Figs 631, 632) line, but side margins and hind corners ...... C. limbatus Meigen always black. Borders of ocellar trian- 16 (15) Setulae on scutum blackish. Ocellar tri- gle sharply outlined. Genae with black angle mainly black, side margins and hairs. Body larger, 3.5–6.0 mm. Male hinter corners brownish or sometimes genitalia Figs 681, 682 yellow. Antennae black or brownish be- ...... C. speciosus Meigen low. Palpi black or yellow. Abdomen red- – Stripes on scutum subshining with thin dish brown in female and darkened dor- dusting. Ocellar triangle black or yellow sally in male. Male genitalia Figs 694, along side margins and on hind corners, 695...... C. varsoviensis Becker usually without light window spots. Bor- – Setulae on scutum whitish. Ocellar tri- der of black parts not so sharply out- angle widely yellow at sides. First flag- lined. Genae without black hairs. Body ellomere yellow below. Palpi yellow. Ab- smaller3.0–3.5mm...... 21 domen yellow with small black spots on 21 (20) Abdomen brown or black dorsally. Basal the second tergite. Male genitalia Fig. segments of antennae brownish. Ocellar 647...... C. pallidiventris (Duda) triangle entirely black or narrowly yel- 17 (12) Stripes on scutum black, shining, with- low along side margins and on hind cor- out dusting or subshining and thinly ners. Black stripes on scutum wider. Fe- dusted. Spot on katepisternum black, male cerci triangular if adjoined (dorsal shining or reddish yellow without view). Male genitalia Fig. 691 dusting...... 18 ...... C. troglodytes (Zetterstedt) – Stripes on scutum black densely grey- – Abdomen dorsally yellow with black dusted. Spot on katepisternum grey- bands. Ocellar triangle broadly yellow dusted, shining black or reddish yellow along side margins and on hinter cor- ...... 22 ners. Basal segments of antennae yellow. 18 (15) Wings weakly darkened, greyish or Black stripes on scutum narrower. Fe- brownish...... 19 male cerci truncate if adjoined (dorsal – Wingsuncoloured...... 20 view). Male genitalia as Fig. 597 19 (18) First flagellomere and spot on katepis- ...... C. calceatus Meigen ternum black. Ocellar triangle predom-

161 22 (17) Pubescence on first flagellomere longer – First flagellomere as long as wide. Palpi than aristal pubescence (Fig. 599). Ocel- and middle of scutellum black or brown- lar triangle entirely black to mainly yel- ish. Spot on katepisternum black, grey- low. Anepisternum often with some dusted. Male genitalia Figs 688, 690 blackorwhitesetulae...... 23 ...... C. strigulus (Fabricius) – First flagellomere with shorter pubes- 28 (26) Basal segments of antennae yellow. Cen- cence. Anepisternum with some white tral scutal stripe (M1)extendingthrough or black setulae or without setulae scutellum. Scutal stripes M2 and M3 ...... 25 connected only in front (Fig. 604). Body 23 (22) Anepisternum without setulae. First smaller, 3.0–3.5 mm. Male genitalia Figs flagellomere yellow to brownish, Palpi 605,606...... C. fijinitimus Becker from yellow to black. Abdomen with – Basal segments of antennae black. Both wide brown bands on tergites. Male gen- stripes on each side of scutum fused. italia Fig. 600 ...... C. fasciatus Meigen Body larger, 4.0–4.5 mm. Male genitalia – Anepisternum with setulae ...... 24 Figs610,611...... C. frontosus Becker 24 (23) Antennae and palpi black. Ocellar trian- 29 (25) Spot on katepisternum black and dens- gle mainly black. Scutellum often dark- elygrey,dusted...... 30 ened. Male genitalia Figs 589, 590 – Spot on katepisternum shining or sub- ...... C. adjunctus Becker shining black without dusting, or red- – Antennae and palpi yellow, first flagel- dishyellow...... 31 lomere slightly darkened on upper side. 30 (29) Anepisternum with some black setu- Ocellar triangle yellow with darkening lae. Basal segment of antennae yellow, on corners and along sides to brownish, first flagellomere yellow with darkened with small groove from anterior ocellus upper side. Front metatarsus and tip to apex. Scutellum yellow of front tibiae yellow. Ocellar triangle ...... C. dasycerus Loew mainly yellow with black central line 25 (22) Arista white, slender or thickened with and without median groove before front dense pubescence. Ocellar triangle wrin- ocellus. Male genitalia Fig. 628 kled on apex and along sides. Scutellum ...... C. laetus Meigen often darkened. Palpus sometimes black – Anepisternum bare, without setulae...... 26 Basal segment of antennae black or – Arista black, slender with short pubes- strongly darkened, first flagellomere cence. Ocellar triangle black or partly black, very rarely fuscous yellow at base yellow, smooth, not wrinkled. Palpi on inner side. Front metatarsus and tip yellow...... 29 of front tibiae blackish. Ocellar trian- 26 (25) Antennae predominantly yellow, first gle entirely black to almost yellow with flagellomere darkened on upper side black median line and black ocellar tu- ...... 27 bercle, in front of front ocellus with a – Antennae entirely or only first flagellom- groove that transforms into a low ridge ereblack...... 28 in apical part of the triangle. Male geni- 27 (26) First flagellomere slightly longer than talia Figs 661, 662 wide. Palpi and scutellum yellow. Spot ...... C. pumilionis (Bjerkander) on katepisternum yellow. On scutum 31 (29) First flagellomere black or at most stripes M and M connected in front 2 3 slightly brownish at base on inner side (Fig. 650). Male genitalia Figs 651, 652 ...... 32 ...... C. pannonicus Strobl

162 – First flagellomere yellow or at most dark- – Headashighaslong.Aristawhite.Pal- ened only on upper and outer sides pus usually black at tip. Black spot on ...... 38 ocellar triangle variable, usually with 32(31) Spotonkatepisternumblack...... 33 yellow window spots, sometimes re- – Spot on katepisternum reddish yellow duced to small spots near ocelli and me- ...... 36 dianline...... C. zernyi (Duda) 33 (32) Ocellar triangle wrinkled on sides or also 38 (31) Ocellar triangle large, smooth, shining, inapicalpart...... 34 reaching front margin of frons, without – Ocellar triangle smooth, at most with distinct linear part (Fig. 622). Stripes central groove before of front ocellus on scutum often partly reddish brown...... 35 Male genitalia Fig. 624 34 (33) Ocellar triangle yellow, shining with ...... C. interruptus Meigen small dark spot on ocellar tubercle. Black – Ocellar triangle with more or less ex- stripes on scutum narrow, divided by pressed linear part, smooth or wrinkled wide yellow ones. Scutellum yellow or swelled along lateral sides, often with ...... C. pallifrons (Duda) whitish spots in front of ocellar tubercle – Ocellar triangle black, shining. Black ...... 39 stripes on scutum wide, divided by nar- 39 (38) Ocellar triangle wrinkled at apex and row yellow ones. Scutellum often along side margins (Figs 612, 625) . . . 40 darkened . . . C. obscurellus (Zetterstedt) – Ocellar triangle smooth, not wrinkled 35 (33) Ocellar triangle large with short linear ...... 41 part which is no more than 1/4–1/5 40 (39) Ocellar triangle dull, short, extending length of frons. Postnotum evenly 0.5 length of frons and continues to an- dusted. Female cerci long and narrow, terior margin of front as a line (Fig. 625). apart in dry specimens Lateral sides of ocellar triangle swelled...... C. planifrons Loew (typical form) Black spots situated on ocellar tubercle, – Ocellar triangle shorter, linear part on side margins and on apex. Palpi short longer, nearly 1/2 length of frons. Dusted and thick. Male genitalia as Figs 626, 627 parts on postnotum divided by shining ...... C. kirigaminensis Kanmiya part. Female cerci short – Ocellar triangle shining, long, linear part ...... C. riparius Smirnov notsodistinct(Fig.612).Medianpartof 36 (32) Ocellar triangle large with short linear ocellar triangle whitish. Male genitalia part no more than 1/4–1/5 length of as Figs 613, 614 . . . C. geminatus Meigen frons. Female cerci long and narrow and 41 (39) Ocellar triangle more or less evenly nar- apart in dry specimens. Male genitalia rowing to apex, blackish. Anepisternum Figs 655–657 with some yellow setulae. Body smaller, C. planifrons Loew (intermediate form) 3–4 mm. Male genitalia Fig. 601 – Ocellar triangle with linear part nearly ...... C. fijiguratus (Zetterstedt) 1/2 length of frons. Female cerci short – Ocellar triangle sharply narrowed in ...... 37 middle of frons with long linear part, of- 37 (36) Head higher than long (Fig. 677). Genae ten with whitish spots before ocelli (Fig. broad. Arista black. Black spot on ocellar 615). No setulae on anepisternum. Body triangle small, usually with two window larger, 4–6 mm. Male genitalia Fig. 617 spots near ocelli. Palpi yellow. Male gen- ...... C. gracilis Meigen italiaFig.678...... C. serenus Loew

163 Species key to regionally known 3rd – Lobes of anal segment small, not brown or instar larvae of Chlorops Meigen tapering (Fig. 998), but posterior spiracles maybebrown(Fig.1001)...... 6 1 Posterior spiracles situated on distinct 6 (5) Posterior spiracles not coloured, rather lobes of anal segment, sometimes lobes wide apart on edges of anal segment (Fig. rather small or brownish coloured (Figs 998). Spicular zones rather narrow. Larvae 990, 993–995, 998, 1000, 1004). Body of lar- 5.5–7.5 mm long, in shoots of cereals and vae cylindrical elongated (Figs 988, 991) Elytrigia repens ...... 2 ...... C. pumilionis (Bjerkander) – Posterior spiracles situated on surface of – Posterior spiracles coloured (Fig. 1001), sit- anal segment (Fig. 987). Body of larvae uated on edges of anal segment or on inner nearly oval and slightly flattened (Fig. 986). sides of incision of anal segment (Figs 1000, Larvae 4–5 mm long, in galls on Brachy- 1004)...... 7 podium and Elytrigia 7 (6) Posterior spiracles (Fig. 1001) somewhat ...... C. strigulus Fabricius cap-shaped with 3–4 projections and 2 (1) Lobes of anal segment longer than wide brown. Anus small, not more than 1/6 of (Fig.993)...... 3 width of anal segment (Fig. 1000). Larvae – Lobes of anal segment not longer than 5–6 mm long, in shoots of Elytrigia repens wide, sometimes very small, brownish ...... C. interruptus Meigen coloured (Figs 998, 1000) ...... 5 – Posterior spiracles situated on inner side of 3 (2) Posterior spiracles situated on the end of incision of anal segment; anus wide, occu- lobes (Figs 993, 994). Maxillary brown pies 2/3–3/4 of width of anal segment (Fig. papillae close to each another (Fig. 992). 1004). Larvae up to 10 mm long, in shoots Anus wide, occupies all width of segment of Deschampsia caespitosa ...... 4 ...... C. speciosus Meigen – Posterior spiracles situated on the inner surface of lobes (Fig. 990) and without in- 119. Chlorops adjunctus terspiracular processes. Maxillary brown papillae wide apart (Fig. 989). Larvae up to Becker, 1910 Figs 589, 590 10 mm long, in shoots of large species of Carex...... C. frontosus Meigen Chlorops adjunctus Becker, 1910: 90. Syntypes in 4 (3) Maxillary palpi (Fig. 992) large, brown ring HNHM. surrounding them narrow. Posterior spir- Ocellar triangle and antennae black. First flagel- acles with short interspiracular processes. lomere with white pubescence. Palpi blackish. Scu- Larvae up to 10 mm long, in shoots of tal stripes broad, nearly confluent, black densely Phalaris arundinaceae dusted. Central stripe continues onto scutellum ...... C. limbatus Meigen and scutellum darkened. Anepisternum with sev- eral black setulae. All pleural spots black. Ab- – Brown ring surrounding maxillary palpi domen dorsally black and with distinct setulae. wide, especially on inner side. Posterior Body length 3.0–4.0 mm. spiracles without interspiracular processes. Larvae 8–10 mm long, in shoots of Carex Distribution. – Rare species. Only from Sweden: SK: ...... C. zernyi (Duda) Vitemölla, 25.vii.1919 (E. Wahlgren), 1 male. European species. 5 (2) Lobes of anal segment brown and tapering, posterior spiracles probably not functional (Fig. 995). Larvae 7–10 mm long, in shoots 120. Chlorops anthracophagoides of large species of Carex Strobl, 1901 ...... C. planifrons Loew Figs 591–593

164 Chlorops anthracophagoides Strobl, 1901: 231. Holo- Etymology. – The species is named in honor of the type female in NMBA (Chvála, 2008). Swedish scientist C. Bjerkander, who described the first species of the genus. Ocellar triangle yellow with black side margins and two black spots, one on ocellar tubercle and an- other on tip. Both black spots connected by black 122. Chlorops calceatus line, as result forming two yellow windows on ocellar triangle. Antennae black, first flagellomere Meigen, 1830 rather large. Arista thick white. Genae narrower Fig. 597 than first flagellomere. Palpi black. Scutal stripes broad black, dusted, scutum before scutellum dark- Chloropscalceatus Meigen, 1830: 146; Meigen’s plate ened. Katepisternum with black spot. Legs yellow. 160:2(Morge,1976a).Lectotype,1female,in Body length 2.0–2.5 mm. MNHN (Nartshuk, 2006). Distribution. – Rare species. Denmark: Sandager Ocellar triangle extending half length of frons, (“Sandacker”), Rinkenæs, 24.vi.1896 (W.Wüstnei), 1 smooth shining black, but hind corners yellow. An- male. Sweden: NB: Eriksberg (in NHMH, Nartshuk, tennae with basal segments yellow, first flagellom- 1998). Russia: Vib, Kr. Not known yet from Norway ere black. Arista black. Genae wider than first flag- and Finland. ellomere. Palpi small yellow. Scutal stripes black, Euro-Caucasian species. thinly dusted and subshining. Spot on katepister- num black, shining. Abdomen yellow with narrow black bands on tergites and central black line. Legs 121. Chlorops bjerkanderi yellow with black last tarsal segments. Cerci of fe- Nartshuk & Andersson, sp. nov. male black. Body length 2.5 mm. Figs 594–596 Distribution. – Very common. Denmark: from all Holotype, female, Sweden: SK: Rövarekulan, Rols- districts except NWJ and SZ. Sweden: northwards berga, 01.ix.1974 (H. Andersson). Holotype in to LY. Norway: northwards to STi. Finland: north- ZMLU. wards to ObS. Russia: Vib, Kr. Eurasian species. Diagnosis. – A new species characterized by the following combination of characters not seen in Biology. – Adults in June to early August, in mead- any other Palearctic species: ocellar triangle and ows and forest edges. antennae entirely black, genae narrow, scutal Larvae phytophagous, living on Festuca rubra stripes narrow, dusted, spot on katepisternum (Tschirnhaus, 1981) and possibly other grasses. black, shining. Description. – Body yellow with black marks. Frons 123. Chlorops centromaculatus longer than wide, yellow, covered with short black (Duda, 1933) setulae. All head setae black, rather short, vte, vti and poc equal in size. Ocellar triangle entirely Oscinis centromaculatus Duda, 1933: 61. Syntypes in black, shining, extending 2/3 length of frons. Oc- NHMW, HNHM and probably other museums. ciput with wide black stripe leaving narrow yellow stripes along eyes margin. Antennae entirely black. Ocellar triangle extending 3/4 length of frons and Palpi yellow. continues as line to front margin of frons, mainly Scutum longer than wide, ratio length:width yellow, black only on ocellar tubercle and here of- as 3.0:2.3. Scutal stripes narrow, dusted, central ten with yellow windows. Genae wider than first one not extending to scutellum. Scutellum yellow. flagellomere. Basal segments of antennae yellow, Pleura with small black, shining mark on anepis- first flagellomere black, arista black. Palpi rather ternum, black, shining spot on katepisternum and enlarged yellow. Scutal stripes black, shining, cen- black, dusted spot on katepimeron. Postnotum tral stripe short. Spot on katepisternum reddish- black, dusted. Abdomen brown dorsally. Wings yellow. Abdomen mainly yellow with very narrow transparent, venation usual for the genus. Halteres black bands on tergites. Legs yellow, last tarsal seg- yellow. Legs yellow with darkened tarsi. Cerci of ment darkened. Cerci of female yellow. Body length ovipositor wide black. Body length 3.5 mm. 3.0 mm.

165 Figs 589–600. Chlorops adjunctus Becker: 589: hypopygium, ventral view. 590: hypandrium and phallic complex. C. anthracophacoides Strobl: 591: head, lateral view. 592: hypopygium, dorsal view. 593: hypandrium and phallic complex. C. bjerkanderi Nartshuk & Andersson: 594: head, dorsal view. 595: head, lateral view (antenna omitted). 596: antenna. C. calceatus Meigen: 597: male genitalia, ventral view. C. fasciatus Meigen: 598: head, dorsal view. 599: head, lateral view. 600: male genitalia, ventral view.

166 Distribution. – Rare species. Sweden: NÄ: Kvist- pleural spots black, shining. Abdomen dorsally bro, Mullhyttan, 30.vii.1941 (T. Nyholt), 1 male; with black bands on tergites. Body length 3.5– UP: (Boheman), 1 female; Forsmark, 18.vii.1975 (R. 4.0 mm. Lenneborg), 1 male. Finland: N: Helsinki, 02.viii.1976, 23.viii.1977, 21.ix.1979 (M. Koponen); Distribution. – Not yet found in Denmark and Ta: Karkola, 15.viii.1974 (M. Koponen). Russia: Vib: Fennoscandia, but expected in SE Fennoscandia. Lake Pionerskoye, Polarny, 07.viii.1998 (Przhiboro), Eurasian species. 1 male, 1 female. Not recorded from Denmark and Norway. European species. 126. Chlorops fijiguratus Biology. – Probably autumnal species, adults from (Zetterstedt, 1848) late July to September. Fig. 601 Oscinis fijiguratus Zetterstedt, 1848: 2631. Holotype 124. Chlorops dasycerus in ZMLU (Andersson, 1966). Chlorops zonulatus Wahlgren, 1913: 52. Synonymy Loew, 1866 by Duda (1933), confirmed by Nartshuk & An- dersson (2002). Lectotype in ZMLU (Nartshuk Chlorops dasycerus Loew, 1866: 46. Type material & Andersson, 2002). not found ZMHU (Wendt, 1975, 1978). Ocellar triangle extending half length of frons and Ocellar triangle black with pale windows. First flag- continues as line to front margin of frons, mainly ellomere yellow to brownish, with longish white yellow, black on ocellar tubercle and with more pubescence. Arista black. Palpi yellow or blackish. or less distinct black median line. Genae distinctly Genae wider than first flagellomere. Scutal stripes wider than small first flagellomere. Antennae yel- black and densely dusted. Scutellum darkened at low, first flagellomere sometimes darkened above, base. Anepisternum with several black setulae. All arista black. Palpi yellow. Scutal stripes black, pleural spots black. Abdomen dorsally black or dusted, rather short, not reaching scutellum. Spot brown. Body length 3.5–4.0 mm. on katepisternum yellow or reddish-yellow. Ab- Distribution. – Rare species. Denmark: NEJ: Rød- domen yellow, often with very narrow black bands stensklit, Skagen (Bræstrup), 1 female. Sweden: SK: on tergites and black central line. Legs yellow, last Vitemölla, 15.ix.1959 (H. Andersson), 1 female. Not tarsal segment darkened. Cerci of female yellow. found in Norway, Finland or Fennoscandian dis- Body length 2.5–3.0 mm. tricts of Russia. Euro-Caucasian species. Distribution. – Sweden: SK, Vittskövle, female holotype of Oscinis fijiguratus. ÖL: Borgholms Alvar, 22.vi.1910, type specimens of C. zonulatus 125. Chlorops fasciatus Wahlgren. Russia: Kr: Paanajärvi (Frey), 1 male. Not found in Denmark or Norway. Meigen, 1830 Eurasian species. Figs 598–600 Biology. – Unknown. A record by Kryshtal (1947) Chlorops fasciatus Meigen, 1830: 143; Meigen’s plate that larvae damaged stems and ears of Elytrigia 276: 14 (Morge, 1976b). Types not found (Becker, repens probably refers to C. novakii Strobl. 1902). Ocellar triangle black with yellow side margins and posterior corners. First flagellomere from yel- 127. Chlorops fijinitimus low to brownish, in white long pubescence. Arista Becker, 1910 black. Palpi yellow or blackish. Genae wider than Figs 602–606 first flagellomere. Scutal stripes rather narrow and densely dusted, central stripe sometimes contin- Chlorops fijinitimus Becker, 1910: 96. Syntypes in ued onto scutellum. Scutellum yellow or with black ZMHU and DEIM (Rohlfien & Ewald, 1970; stripe. Anepisternum without black setulae. All Wendt, 1975).

167 Figs 601–611. Chlorops fijiguratus (Zetterstedt): 601: male genitalia, ventral view. C. fijinitimus Becker: 602: head, dorsal view. 603: head, lateral view. 604: thorax, dorsal view. 605: hypopygium, ventral view. 606: hypandrium and phallic complex. C. frontosus Meigen: 607: head, dorsal view. 608: head, lateral view. 609: antenna. 610: hypopygium, dorsal view. 611: male genitalia, ventral view.

168 Ocellar triangle extending nearly to front margin Larvae phytophagous, develop in shoots of large of frons, black, with yellow hind corners, wrinkled species of Carex (Séguy, 1934; d’Aguillar, 1943). along side margins, sometimes with yellow marks Larvae described by d’Aguillar (1943). lateral to ocelli. Genae distinctly wider than first flagellomere. Basal segments of antennae fuscous yellow, first flagellomere black, slightly longer than 129. Chlorops geminatus wide. Arista whitish. Palpi basally yellow, apically Meigen, 1830 blackish. Scutal stripes black, dusted, sublateral Figs 612–614 stripes connected with lateral in front. Scutum be- fore scutellum darkened and this diffuse spot oc- Chlorops geminatus Meigen, 1830: 141; Meigen’s cupies scutellum. Spot on katepisternum reddish- plates 159: 10; 211: 19 (Morge, 1976b). Lectotype, yellow, often with black upper part. Abdomen yel- female, in MNHN (Nartshuk, 2006). Specimen low with black bands on tergites and black central in NHMW does not belong to the type series line. Legs mainly yellow, femora partly black, last (Nartshuk, 1997a). tarsal segment darkened. Cerci of female yellow. Ocellar triangle extending nearly to front margin Body length 2.5–3.0 mm. of frons, yellow, shining, wrinkled along side mar- Distribution. – Rare species, known only from Swe- gins. Ocellar tubercle and tip of triangle black and den: ÖL: Byxelkrok, 05.viii.1958 (B.H. Hansen), 2 sometimes central black line connects these black males, 1 female; Halltorps Hage, 22.vii.1978 (H. An- spots. Genae 1.5 times wider than first flagellom- dersson),1male,1female. ere, yellow. Antennae entirely yellow, first flagel- The species is known from Europe, the Caucasus lomere sometimes darkened above. Arista black and Central Asia. or brown. Palpi rather thick yellow. Scutal stripes black, dusted, rather short. Spot on katepisternum Biology. – Adults in July to August, in dry meadows. yellow. Abdomen yellow with narrow black bands on tergites. Legs yellow. Body length 3.5–4.0 mm. 128. Chlorops frontosus Distribution. – Rare. Denmark: WJ: Tipperne, Meigen, 1830 Ringkøbing Fjord (E. Rald). Sweden: ÖL: Figs 607–611, 988–990 Mörbylånga, 09.vii.1917 (Tullgren), 1 female; GO: Hammars, Norrlanda, 12–18.viii.1979 (R. Daniels- Chloropsfrontosa Meigen, 1830: 148; Meigen’s plates son), 1 male, 1 female; ÖG: Kastad, Örberga, 213: 10; 276: 15 (Morge, 1976b). Lectotype in 20.vii.1979 (H.W. Waldem), 1 female. Russia: Kr: NHMW (Nartshuk, 1997a). Ruskeala, 28.vii.1949 (Nyland), 1 male. Not found in Norway or Finland. Ocellar triangle extending nearly to front margin Eurasian species. of frons, black, wrinkled along side margins and with central sulcus. Genae distinctly wider than Biology. – Adults in July to August in meadows. first flagellomere, yellow with black lower mar- It has been reared from Hierochloe odorata (Kar- gin. Antennae black, first flagellomere longer than pova, 1972). wide. Aristal pubescence thick and white. Palpi black. Scutal stripes wide black, dusted, outer lat- eral stripes fused with inner lateral ones. Scutellum 130. Chlorops gracilis blackish. Spot on katepisternum black, dusted. Ab- Meigen, 1830 domen dorsally black. Legs mainly black, coxae and Figs 615–617 tibiae partly yellow. Body length 3.0–3.5 mm. Chlorops gracilis Meigen, 1830: 140; Meigen’s plate Distribution. – Scarce. Denmark: SJ: Gråsten (W. 160: 1 (Morge, 1976a). Holotype, specimen with- Wüstnei); NEZ: Høed Skov. Sweden: SK, SÖ, UP. out abdomen and hind legs, in MNHN Russia: Kr: Kondopozhsky Distr.: Malyi Lednikovski (Nartshuk, 2006). Is., 24.vi.2003 (Polevoi), 1 female. European species. Ocellar triangle extending nearly half length of frons, yellow, with median slightly darkened sulcus, Biology. – Adults in May to June, on fens and black on ocellar tubercle and sometimes with yel- marshes with Carex. low windows on this black spot. Genae 1.5 times

169 Figs 612–624. Chlorops geminatus Meigen: 612: head, dorsal view. 613: hypopygium, dorsal view. 614: male genitalia, ventral view. C. gracilis Meigen: 615: head, dorsal view. 616: head lateral view. 617: hypandrium and phallic complex. C. hypostigma Meigen: 618: head, dorsal view. 619: head, lateral view. 620: hypopygium, ventral view. 621: hypandrium and phallic complex. C. interruptus Meigen: 622: head, dorsal view. 623: head, lateral view. 624: male genitalia, ventral view.

170 wider than first flagellomere. Antennae entirely Biology. – Adults from late May to August, in mead- yellow. Arista rather long haired, black, but basally ows, edges of forests. Univoltine species, overwin- yellow. Palpi yellow. Scutal stripes black, densely tering as third instar larva within shoots of grasses. dusted, not extended to scutellum. Spot on katepis- Larvae phytophagous, develop in shoots of ternum yellow. Abdomen yellow shining, some- Dactylis glomerata, Poa sp., P. trivialis, Festuca sp. times with narrow black stripes on tergites. Legs (Séguy, 1934; Mowat, 1975; Uffen & Chandler, 1978; yellow. Body length 4.5–5.0 mm. Tschirnhaus, 1981). Distribution. – Rather common in dry habitats. Sweden: northwards to JÄ. Finland: only in south- ern regions: Ab, N, St, Ta, Sa, Tb, Kb. Russia: Vib; 132. Chlorops interruptus Kr: Kondopozhsky Distr.: Shoksha, 13.vii.2004 Meigen, 1830 (Polevoi), 1 male. Not so far recorded from Den- Figs 622–624, 999–1001 mark and Norway. Eurasian species. Chlorops interruptus Meigen, 1830: 145; Meigen’s Biology. – Adults in June to July in dry meadows. plate 213: 9 (Morge, 1976b). Lectotype in NHMW One generation per year. Hibernates as larvae in (Nartshuk, 1997a). shoots of its host plant. Ocellar triangle large, reaching front margin of Larvae phytophagous, develop in shoots of Cala- frons, yellow shining, medially with furrow and magrostis epigeios (Kröber, 1910; Nartshuk, 1962a; black line from ocellar tubercle to front end. Genae Uffen & Chandler, 1978; Dubbert et al., 1998). Lar- nearly 1.5 times wider than first flagellomere. An- vae of other Diptera, mostly chloropids, live in tennae entirely yellow, arista black. Palpi yellow. shootsinfestedbythisspecies:Aphanotrigonum Scutal stripes black, dusted, rather narrow and not trilineatum, A. nigripes, Lasiosina herpini,and extended to scutellum, sometimes stripes partly Asteia concinna Meigen (Asteiidae). brownish-red. Spot on katepisternum yellow. Ab- domen yellow. Legs yellow, last tarsal segments 131. Chlorops hypostigma darkened. Body length 2.5–3.5 mm. Meigen, 1830 Distribution. – Only from Sweden: UP: Hässelby, Figs 618–621 Börje, 03.ix.1965 (Kurs). Euro-Mediterranean species. Chlorops hypostigma Meigen, 1830: 141; Meigen’s plates159:12;211:20;213:12(Morge,1976b).Lec- Biology. – Two generations per year in Central Eu- totype male, and paralectotypes 2 males and 1 rope. female in MNHN (Nartshuk, 2006). Larvae phytophagous, develop in shoots of Elyt- rigia and ?Phragmites (Séguy, 1934; Balachowsky & Ocellar triangle extending nearly half length of frons, black with rather wide yellow side margins, Mesnil, 1935; Uffen & Chandler, 1978). The record sometimes with small yellow window before front of Phragmites as host plant needs verification. ocellus. Genae slightly narrower than first flagel- Larvae described by Balachowsky & Mesnil lomere. Basal segments of antennae fuscous yellow, (1935). first flagellomere black. Arista black. Palpi yellow. Scutal stripes black, shining, rather narrow and not extend scutellum. Spot on katepisternum black, 133. Chlorops kirigaminensis shining. Abdomen yellow or sometimes with nar- Kanmiya, 1978 row black stripes on tergites. Legs yellow, last tarsal Figs 625–627 segments blackish. Body length 2.0–2.5 mm. Distribution. – Common. Denmark: widespread. Chlorops kirigaminensis Kanmiya, 1978: 55. Holo- Sweden: northwards to LY. Norway: AK, HEs, VAy, type and paratypes in Entomological Labora- Ri,HOi,MRy.Finland:northwardstoObS.Russia: tory Faculty of Agriculture Kyushu University Vib, Kr. (Kanmiya, 1978). Paratype, female in USNM, Euro-Caucasian species. Washington, D.C.

171 “Chlorops zonulatus Wahlgren”; Smirnov, 1958; with 5 black densely grey-dusted stripes. Scutel- Smirnov & Fedoseeva, 1976; Nartshuk, 1998. lum yellow. Spot on katepisternum yellow, anepis- Misidentifications. ternum with some black setulae. Abdomen yellow with narrow black bands on tergites and black me- Frons wider than long. Ocellar triangle short, ex- dian line. Body length 3.5–4.0 mm. tending to middle of frons and then continues to front margin as a line. Ocellar triangle shining, cen- Distribution. – Not common. Sweden: SM, ÖL, ÖG, trally yellow and with black spots on ocellar tuber- SÖ, UP. Norway: AK, HOy. Finland: Sa. Russia: Vib, cle, in front of ocelli, on side margins and on apex. Kr. Not found in Denmark. Surface of triangle between black spots whitish- The species is known from Europe and Kazakh- yellow and sulcate medially and laterally. Genae as stan. wide as first flagellomere. First flagellomere yellow, darkened at base of arista, arista black. Palpi yellow. Biology. – Autumnal species, adults in August to Scutum with 5 black densely grey-dusted stripes September. covered with yellowish setae anteriorly and black setulae posteriorly. Scutellum yellow with 6 setae. Spot on katepisternum yellow, spot on anepister- 135. Chlorops limbatus num black. Postnotum black, dusted. Body length Meigen, 1830 3.5–4.0 mm Figs 629–632, 991–993 Distribution. – Sweden: LU: lake NW of upper course of river Vietasjokk, 14.vii.1961 (Dahm, Chlorops limbatus Meigen, 1830: 150; Meigen’s Landin & Löfqvist), 1 male. More common in Fin- plates 211: 13; 213: 11 (Morge, 1976b). Lectotype land:Al,Ab,N,St,Ta,Sa,Oa,Om.Russia:Vib;Kr: in NHMW (Nartshuk, 1997a). Paanajärvi. Not found in Denmark and Norway. Chlorops discicornis Zetterstedt, 1848: 2625, non Widely distributed in Asian part of the Palearctic Loew, 1866: 66 (Nartshuk & Andersson, 2002). eastwards to Japan and in East Europe, but not re- Chlorops fulviceps Roser, 1840: 62 (Nartshuk, 1994). corded in West Europe except for Sweden and Fin- Chlorops brevimanus Loew, 1866 (Nartshuk, 1997b). land. An eastern Eurasian species. Frons wider than long. Ocellar triangle extending Biology. – Adults in July to August in meadows. half length of frons, black smooth with little raised side margins. Genae narrow, about 2/3 width of Note. – The species was described from Japan first flagellomere. First flagellomere black, rather (Kanmiya, 1978), then synonymized with C. gemi- large, slightly broader than long. Arista black. Palpi natus by Kanmiya (1983) but subsequently restored rather stout yellow. Scutum with 5 black densely from synonymy by Nartshuk (2002a). grey-dusted rather wide stripes. Scutellum yellow. Spot on katepisternum black. Abdomen yellow with black bands on tergites or sometimes black- 134. Chlorops laetus ish dorsally. Legs yellow with medially black hind Meigen, 1830 tibiae and darkened tarsi. Front basitarsus of male Fig. 628 short and claws of front legs asymmetrical. Male genitalia with sclerotized aedeagus. Body length Chlorops laetus Meigen, 1830: 143; Meigen’s plate 2.5–3.0 mm. 211: 15 (Morge, 1976b). Type specimens not found in Coll. Winthen in NHMW, probably de- Distribution. – Rather common. Denmark: from all stroyed. districts except WJ and LFM. Sweden: northwards to UP and VR, and further north in NB. Norway: Frons wider than long. Ocellar triangle narrow, ex- MRy, NTi. Finland: northwards to ObN. Russia: Vib, tending to middle of frons followed by linear exten- Kr. sion to front margin of frons. Ocellar triangle with Trans-Palearctic species, known from the British yellow windows and black spots in front of ocelli, Isles to Japan. on side margins and on apex. Genae 1.5 times as wide as first flagellomere. Basal segments of anten- Biology. – Adults in May to September, in wet nae fuscous yellow. First flagellomere basally yel- meadows and on banks of rivers and lakes. Two low, black above. Arista black. Palpi yellow. Scutum generations per year.

172 Figs 625–638. Chlorops kirigaminensis Kanmiya: 625: head, dorsal view. 626: hypopygium, dorsal view. 627: male genitalia, ventral view. C. laetus Meigen: 628: male genitalia, ventral view. C. limbatus Meigen:629:head,dorsalview.630:head,lateralview.631:hypopygium, ventral view. 632: hypandrium and phallic complex. C. meigenii Loew: 633: head, dorsal view. 634: head, lateral view. 635: thorax, dorsal view. 636: hypopygium, ventral view. 637: apex of surstyli. 638: hypandrium and phallic complex.

173 Larvae phytophagous, develop in shoots of Oscinis nigripalpis Duda, 1933: 186. Holotype in Phalaris arundinacea, larvae of the summer gen- NHMH, currently on loan in ZMLU. eration on exposed panicles. The leaves above the Chlorops crassipalpis Smirnov, 1958: 1166. Holotype stem-feeding larvae dry out and turn brown. Larvae in ZMMU (Nartshuk & Andersson, 2002). hibernate in basal and aerial shoots and pupate the following spring (Nartshuk, 1962a; Sharipov, 1981). Frons longer than wide. Ocellar triangle extending Records of other host grasses belonging to Alopecu- 2/3 length of frons followed by linear extension to rus and Phleum pratense (Wetzel, 1967; Tschirn- front margin of frons, predominantly yellow with haus, 1981) need verification. median black spot, occupying ocellar tubercle and Larvae described by Balachowsky & Mesnil narrowingtoapex.Genae1.5timesaswideasfirst (1935), Oettingen (1935) and Nye (1958). flagellomere. Basal antennal segments fuscous yel- low. First flagellomere and arista black. Palpi yellow basally, black in apical half. Scutum with 5 black, 136. Chlorops meigenii dusted stripes not extended to scutellum. Scutel- Loew, 1860 lum yellow. Spot on katepisternum black, anepis- Figs 633–638 ternum with small black spot. Abdomen yellow Chlorops meigenii Loew,1860:43.Thenameisin- with black bands on tergites. Legs mainly yellow, cluded in Official List of Names in Zoology, last tarsal segments darkened. Body length 3.0 mm. Opinion 2040 (Case 3190), 2003: 116. Distribution. – Rare species. Sweden: NB: Högsön, Ocellar triangle predominantly black, shining, Råneå, 31.vii.1978 (H. Andersson), 1 female; reaching front margin of frons and yellowish at Jupukka, Rajala, 28.vii.1955 (H. Andersson), apex. Genae as wide as first flagellomere or a lit- 5 males. Finland: N: Helsinki (Ahnger), 1 female. tle wider. Basal antennal segments fuscous yel- Russia: Kr: female holotype of Oscinis nigripalpis. low. First flagellomere small rounded, mainly dark The species is known from Europe and Kazakh- brown or yellow with darkening dorsally. Arista stan. black. Palpi small yellow. Scutum with 5 black, shining stripes, central one with indistinct outline posteriorly. Inner lateral stripes without projection 138. Chlorops obscurellus on outer margin. Stripes nearly fused in some dark specimens collected in northern Fennoscandia. (Zetterstedt, 1838) Spot on katepisternum reddish brown or blackish Figs 644–646 dorsally. Abdomen yellow with black bands. Wings brownish. Legs mainly yellow, but femora and hind Oscinis obscurellus Zetterstedt, 1838: 779. Holotype tibiae partly black, last tarsal segments darkened. in ZMLU (Andersson, 1966). Body length 3.0–3.5 mm. Frons as long as wide. Ocellar triangle large, behind Distribution. – One of the most common species of touching eyes, extending nearly to front margin of the genus. Denmark: SJ, F, LFM, NWZ, NEZ. Swe- frons, black, except hind corners, smooth and shin- den: northwards to TO. Norway: Ø, AK, HEs, AAy, ing, wrinkled on apex. Genae a little wider than HOy, HOi, SFy, SFi, Nsy. Finland: northwards to Li. first flagellomere. Basal antennal segments brown, Russia: Vib, Kr, Lr. first flagellomere and arista black. Palpi yellow.Scu- Trans-Palearctic species ranging from Europe to tum with 5 black, dusted stripes, three central ones Kamchatka. very broad and extended to scutellum. Scutellum Biology. – AdultsinMaytoAugust,inwetmeadows yellow, basally black. Spot on katepisternum black, and forest clearings. Adults were found in a swal- shining. Abdomen brown to black dorsally with low’s nest (Hicks, 1959). narrow yellow side margins. Legs yellow, femora Probable host plants are some species of Calama- and hind tibiae partly black, last tarsal segments grostis growing in wet places, other than C. epigeios. darkened. Body length 2.0–3.0 mm. Distribution. – Denmark: SJ, LFM, NEZ. Sweden: 137. Chlorops nigripalpis northwards to NB. Finland: Ta, Oa, Kb, Om. Russia: (Duda, 1933) Vib. Not found in Norway. Figs 639–642 Eurasian species.

174 Figs 639–652. Chlorops nigripalpis (Duda): 639: head, dorsal view. 640: head, lateral view. 641: hypopygium, ventral view. 642: hypan- drium and phallic complex. C. obscurellus (Zetterstedt): 643: head, dorsal view. 644: head, lateral view. 645: hypopygium, ventral view. 646: hypandrium and phallic complex. C. pallidiventris (Duda): 647: male genitalia, ventral view. C. pannonicus Strobl: 648: head, dorsal view. 649: head, lateral view. 650: thorax, dorsal view. 651: hypopygium, ventral view. 652: hypandrium and phallic complex.

175 Biology. – Adults in June to August, in wet mead- Abdomen yellow with narrow black bands on ter- ows and swamps. gites. Legs yellow, last tarsal segments darkened. Larvae phytophagous, develop in shoots of Carex Body length 3.0–3.5 mm. riparia without causing external changes (De Bruyn, 1989). Distribution. – Rare. Only from Sweden: VS: Skäret, Valskog, 11.vii.1989 (H. Andersson), 1 female; VR: Storfors, 28.vi.1988 (H. Andersson), 2 females. 139. Chlorops pallidiventris European species. (Duda, 1933) Fig. 647 141. Chlorops palpatus Oscinis pallidiventris Duda, 1933: 164. Syntype, 1 fe- Smirnov, 1958 male, in ZMHU (Wendt, 1975). Chlorops palpatus Smirnov, 1958: 1169, 1174. Syn- Frons as long as wide or a little longer. Ocellar tri- types probably in ZMMU. angle short, extending nearly 1/3 length of frons followed by linear extension to front margin of Frons longer than wide. Ocellar triangle extending frons. Black spot on ocellar triangle occupies cen- 2/3–3/4 length of frons followed by linear exten- tral part; side margins narrowly and hind corners sion to front margin of frons, shallow in the middle. widely yellow. Genae narrower than first flagel- Black spot on ocellar triangle occupies central part, lomere. Antennae yellow, first flagellomere black side margins and hind corners yellow. Basal seg- above, arista black. Palpi yellow. Scutum with 5 ments of antennae yellow, first flagellomere black, black, dusted rather narrow stripes not extended arista black. Palpi rather long and broad, brown- to scutellum. Setulae of scutum yellow. Scutellum ish to black. Scutum with 5 black, shining stripes, yellow, basally blackish. Spot on katepisternum central stripes not extended to scutellum. Scutel- black, shining. Anepisternum with some setulae. lum yellow. Spot on katepisternum shining, reddish Abdomen whitish-yellow covered with yellow se- yellow, with brown stripe above. Abdomen yellow, tulae. Legs yellow. Basitarsus of front legs in male shining, covered with whitish setulae. Legs yellow. short, only 2 times as long as broad and claws of Body length 2.0–3.0 mm. front legs asymmetrical. Body length 2.0–2.5 mm. Distribution. – Rare. Sweden: HA: Årnilt, Enslöv, Distribution. – Rare. Only from Sweden: ÖL: Hall- 04.viii.1972 (H. Andersson), 2 males, 1 female; UP: torps Hage, 19–23.v.1976 (H. Andersson & R. (Boheman), 1 specimen. Norway: VE: Sandefjord Danielsson), 1 male. (Siebke). European species, known elsewhere only from Eurasian species, known from European part of Hungary and Bulgaria. Russia.

140. Chlorops pallifrons 142. Chlorops pannonicus Strobl, 1909 Strobl, 1893 Figs 648–652 Chlorops pallifrons Strobl, 1909: 287. Syntypes 2 males, 4 females in NMBA (Chvála, 2008). Chlorops pannonica Strobl, 1893: 228. Syntypes 5 males, 5 females in NMBA (Chvála, 2008). Frons wider than long. Ocellar triangle a little Chlorops balticus Wahlgren, 1913: 51. Holotype in longer than half length of frons, yellow, sulcate ZMLU (Duda, 1933; Nartshuk & Andersson, along side margins, continuing as a yellow line 2002). to front margin of frons. Ocellar tubercle black. Genae 1.5 times wider than first flagellomere. Basal Frons longer than wide. Ocellar triangle extending antennal segments yellow, first flagellomere and nearly to front margin of frons with median sul- arista black. Palpi yellow. Scutum with 5 black, cus and often wrinkled along side margins. Black dusted stripes, central one extended to scutellum. spot on ocellar triangle occupies nearly all sur- Scutellum yellow. Spot on katepisternum black. face except hind corners. Genae a little wider than

176 first flagellomere. Antennae yellow, first flagellom- on katepisternum black or yellow, but intermedi- ere black above, a little longer than wide. Arista ate colour also occurs: yellow with black above. whitish. Palpi small yellow. Scutum with 5 black, Abdomen yellow with narrow black bands on ter- dusted stripes, outer lateral stripes connected with gites and often with median black line. Legs yellow, inner lateral ones in front and fused with them last tarsal segments darkened. Cerci of female nar- behind. Scutellum yellow, basally at sides black- row and long, in dry specimens divergent. Body ish. Spot on katepisternum from yellow to black, length 2.5–3.0 mm. The light coloured form was covered with thin dust and subshining. Abdomen described as C. triangularis Becker, 1910: 97, which brown or blackish. Legs yellow, femora partly black Duda (1933) considered as variety of C. planifrons. and last tarsal segments darkened. Body length Distribution. – Very common and widespread. Den- 2.0–2.5 mm. mark: SJ, EJ, NEJ, NEZ. Sweden: northwards to TO. Distribution. – Rare. Only from Sweden: ÖL: female Norway: HEn. Finland: north to Li. Russia: Vib, Kr, holotype of Chlorops balticus and another female Lr. from July 1916 also collected by Wahlgren; GO: one Trans-Palearctic, widely distributed polyzonal female from collection of Boheman (NRMS). Holo- species, known from the British Isles to the Far East type of Chlorops balticus was collected on the alvar of Russia, in Europe southwards to Spain. (Wahlgren, 1913). Biology. – Adults from middle June to middle Au- Xerophilous species widely distributed in the gust,onfensandmarsheswithlargetussocksof Palearctic steppe zone from Hungary to the Far Carex. One generation per year. East, with local extensions northwards in dry, sandy Larvae phytophagous, develop in shoots of Carex, habitats. Finding places nearest to Gotland and e.g. C. inflata, C. rostrata (Nartshuk, 1962a; Zhabin- Öland in Estonia: Islands of Kõinastu and Saare- skaya, 1963). Larvae of the first instar hibernate in maa (Nartshuk, 1998). shoots of its host plant and continue feeding the Note. – Duda (1933: 189) considered Chlorops balti- following spring. Larvae of the summer generation cus Wahlgren as a synonym of C. pannonicus. make a furrow on stem that begins immediately Nartshuk (1984b) listed the species as a separate below inflorescence and gradually widens down- species. Now, after examination of holotype of wards. At some point is an interruption of the fur- Chlorops balticus, we accept this synonymy. row below which it suddenly increases in width, evidently an indication of larval molting. Larvae pupate in the basal part of the Carex stem. Some larvae leave their first stem in order to continue 143. Chlorops planifrons feeding on another stem (Nartshuk, 1962a). Loew, 1866 Larvae described by Zhabinskaya (1963). Figs 653–657, 995

Chlorops planifrons Loew, 1866: 55. Syntypes in 144. Chlorops pumilionis ZMHU (Wendt, 1975). (Bjerkander, 1778) Chlorops triangularis Becker, 1910: 97, as var. of C. Figs 104, 658–662, 996–998 planifrons Loew. Syntypes in ZMHU (Wendt, 1975). Musca pumilionis Bjerkander, 1778: 240. Types not found; see under Meromyza saltatrix. Body yellow or sometimes in specimens collected Chlorops pumilionis gotlandica Holmgren, 1887: 75. in autumn reddish. Frons longer than wide. Ocel- Types not found (Nartshuk & Andersson, 2002). lar triangle large, extending 4/5 length of frons, smooth shining. Black spot on ocellar triangle Body yellow or sometimes in specimens collected rather variable: occupies all triangle or divided into in autumn reddish. Frons as long as wide. Ocellar two spots, one on ocellar tubercle and another triangle extending slightly more than half length rhomboid in form at the apex of triangle. Genae a of frons and continues as line to front margin of little wider than first flagellomere. Basal segments frons. Its surface black or brown, smooth shining of antennae yellow, first flagellomere rather large with median sulcus before front ocellus and me- black. Arista slightly whitish. Palpi yellow. Scutum dian ridges in apical part. In light coloured spec- with 5 black, dusted stripes. Scutellum yellow. Spot imens only ocellar tubercle, sulcus and ridge are

177 Figs 653–662. Chlorops planifrons Loew: 653: head, dorsal view. 654: head, lateral view. 655: hypopygium, ventral view. 656: surstyli and mesolobus. 657: hypandrium and phallic complex. C. pumilionis (Bjerkander): 658: head, dorsal view. 659: head, and part of thorax, lateral view. 660: wing. 661: hypopygium, ventral view. 662: hypandrium and phallic complex.

178 black. Genae wider than first flagellomere. Basal yellow windows. Genae wider than small first flag- segments of antennae brown or black, first flag- ellomere. Basal segments of antennae yellow or ellomere and arista black. Palpi yellow. Scutum brownish, first flagellomere black or brownish be- with 5 wide black, dusted stripes. Scutellum yellow. low. Arista black. Palpi long yellow and black api- Spot on katepisternum black, dusted. Abdomen cally in females. Scutum with 5 black thinly dusted yellow with black bands on tergites or blackish. stripes. Scutellum yellow or rarely with black spot. Legs mainly yellow, apical part of tibiae and ba- Spot on katepisternum yellow. Abdomen yellow sitarsus or all tarsus of front legs black, last tarsal with narrow black bands on tergites. Legs yellow, segments of other legs darkened. Body length 2.5– hind tibiae with black band. Two last tarsal seg- 3.5 mm. ments darkened. Body length 2.5–3.5 mm. Distribution. – Denmark: from all districts. Sweden: Distribution. – Denmark: NEZ, B. Sweden: north to north to ÅN. Norway: Ø, AK, HEs, Os, Bv, Bø, VAi, DR. Norway: AK. Finland: Ab, N, Ka, Sa. Russia: Vib, HOi. Finland: north to Om. Russia: Vib, Kr. Kr. Eurasian temperate species, widespread from Eurasian temperate species, known from south- Europe to Iran, Israel, West Siberia and Mongolia. ernSwedenandFinlandsouthwardstoBulgaria Biology. – Adults from end of May to early Septem- and eastwards to Yakutia, East Siberia. ber, in two generations. Adults of the second gener- Biology. – Adults in meadows. Autumnal species ation were found in a swallow’s nest (Krivokhatsky occurring in August and September. & Nartshuk, 2001). Well known pest of cultivated cereals, especially wheat, occationally barley and rye, but not oats. Larvae of the winter generation develop in shoots 146. Chlorops riparius of cereals causing thickening of the attacked shoots Smirnov, 1958 (“gout swelling”), while larvae of the summer gen- Fig. 107 eration feed on the exposed, developing ear and stem of cereals (Porchinsky, 1881). Plant reactions Chlorops riparius Smirnov, 1958: 1167, 1174, as sub- to the larval attacks were studied specially by species of C. nigripalpis (Duda). Syntypes prob- Goidanich (1956). Many species of wild grasses ably in ZMMU. have been recorded as possible host plants of the Goats fly by different authors, but most likely only Frons wider as long. Ocellar triangle extending Elytrigia repens and Aegylops sp. are genuine host nearly half length of frons or little longer followed plants. Injuries on cereals caused by Chlorops by linear extension to front margin of frons, with pumilionis have been reported from several Nordic sulcus from front ocellus to apex. Black spots on countries (Lampa, 1888; Johansson, 1960; Rygg, ocellar triangle rather narrow, side margins of tri- 1966a), but the species appears to be of little eco- angle widely yellow. Genae wider than small first nomic importance. flagellomere. Basal segments of antennae yellow Larval descriptions are given by many authors, or brownish, first flagellomere black or brownish notably by Frew (1923a), Balachowsky & Mesnil below. Arista black, distinctly pubescent. Palpi yel- (1935), Nye (1958) and Zhabinskaya (1963). low short and rather stout. Scutum with 5 black densely dusted stripes, central stripe not extending to scutellum. Scutellum yellow. Spot on katepister- 145. Chlorops ringens num black and dusted. Abdomen yellow or with Loew, 1866 very narrow black bands on tergites. Legs yellow, Figs 663, 664 two last tarsal segments darkened. Body length 3.0–4.0 mm. Chlorops ringens Loew, 1866: 46, 60. Syntypes in Distribution. – Sweden:VG,UP.Norway:SFi:Vass- ZMHU (Wendt, 1975). bygda, Aurland, 16.viii.1941 (N. Knaben); Loen, Fronsaswideaslong.Ocellartriangleextending Stryn, 02.viii.1942 (N. Knaben), 1 male. Not found in 2/3 length of frons followed by linear extension Denmark, Finland or Russian parts of Fennoscan- to front margin of frons. Black spots on ocellar dia. triangle rather small triangular and usually with Eurasian species.

179 Biology. – Species occurring sporadically in August, 148. Chlorops rufijinus near small streams in forest, probably one genera- (Zetterstedt, 1848) tion per year. Figs 666–668 Larvae phytophagous, live in shoots of Elymus caninus, hibernate as larvae of first or second in- Oscinis rufijinus Zetterstedt, 1848: 2628. Holotype in star. The mature larvae produce following spring a ZMLU (Andersson, 1966). gall-like swelling near the ground on young shoots Oscinis citrinellus Zetterstedt, 1848: 2630 (Nartshuk (Nartshuk, 1962a). Some inquilines live within the & Andersson, 2002). galls: Elachiptera cornuta and Asteia concinna Oscinis bipunctus Duda, 1933: 173. Meigen (Asteiidae) (Nartshuk, 1962a). Oscinis bipunctus hungaricus Duda, 1933: 173. Oscinis bipunctus fennicus Duda, 1933: 173. Note. –Typesofhungaricus not found in ZMHU 147. Chlorops rossicus (information of A. Soós in Zuska, 1960). Zuska Smirnov, 1955 (1960) designated a lectotype, 1 female, of bipunc- Fig. 665 tus in NHMH (N 8159). Species very variable in colour of scutal stripes, spot Chlorops rossicus Smirnov, 1955: 334. Holotype on katepisternum and first flagellomere. Frons as probably in ZMMU. wide as long or a little wider. Ocellar triangle ex- Frons usually wider than long with black upright tending 2/3–3/4 length of frons and continues as setulae. Ocellar triangle black, extending half line to front margin of frons, yellow with two black length of frons or little longer followed by linear spots: one on ocellar tubercle and another rhom- extension to front margin of frons. Black spot on boid spot on apex, both spots connected by black ocellar triangle occupies nearly all triangle except line, side margins of triangle narrowly black as well. narrow yellow lines along side margins and hind Genae as wide as first flagellomere. Antennae yel- angles, black middle line goes from front ocellus low or first flagellomere darkened above. Arista to apex of triangle. Genae with some black setu- whitish with yellow basal part. Palpi yellow. Scu- lae, wider than first flagellomere. Basal segments tum with 5 red, partly black or entirely black, thinly of antennae yellow or brownish, first flagellomere dusted stripes. Scutellum yellow. Spot on katepis- and arista black. Palpi yellow. Scutum with 5 wide ternum from yellow to black, dusted. Abdomen black, shining stripes and often blackish before yellow with narrow black bands on tergites. Legs scutellum. Inner lateral stripes widened in front yellow, last tarsal segments darkened. Body length and outer margin with triangular incision opposite 2.5–3.0 mm. notopleura. Scutellum yellow. Spot on katepister- Distribution. – Sweden: northwards to NB. Norway: num black, shining. Wings brownish, costal vein AK,HOi.Finland:Al,Ta.Russia:Vib,Kr.Notre- extending beyond tip of R4+5. Abdomen blackish corded from Denmark. dorsally. Legs yellow, or with black stripes on hind Widespread trans-Palearctic species, known tibiae. Two last tarsal segments darkened. Body southwards to Bulgaria and eastwards to Mongolia length 2.5–3.5 mm. and Japan, but everywhere rare. Distribution. – Denmark: NEJ: Danzigmand, Læsø, Biology. – Adults in July to August in meadows. 12.vii.1969 (ZMUC-Exp.), 1 female; NEZ: Lyngby Mose,03.vii.1964(O.Martin),1male,2females, 149. Chlorops scalaris 21.vi.1972 (K. Schnack), 1 female. Sweden: 1 female. Sweden: SK: Revingehed, 06.vi.1993 (R. Daniels- Meigen, 1830 son), 1 female; SÖ: Trehörningen, Huddinge, Figs 669–672 21.vi.1960 and 03.vii.1965 (L. Hedström), 2 females Chlorops scalaris Meigen, 1830: 145; Meigen’s plate in ZMUU. Norway: SFy, STy, TRy. More common in 159: 11 (Morge, 1976a). Lectotype and some par- Finland: N, Ka, St, Ta, Sb, Li. Russia: Vib, Lr. alectotypes in NHMW (Nartshuk, 1997a), other European species. paralectotypes in MNHN. Biology. – Adults in June and July, in meadows and Frons wider than long. Ocellar triangle extend- edges and clearings in forests. ing 2/3 length of frons followed by linear exten-

180 Figs 663–672. Chlorops ringens Loew: 663: hypopygium, latero-ventral view. 664: hypandrium and phallic complex. C. rossicus Smirnov: 665: thorax, dorsal view. C. rufijinus (Zetterstedt): 666: hypopygium, ventral view. 667: surstylus. 668: hypandrium and phallic complex. C. scalaris Meigen: 669: head, dorsal view. 670: head, lateral view. 671: hypopygium, ventral view. 672: hypandrium and phallic complex.

sion to front margin of frons, yellow with small Distribution. – Denmark: common and wide- median black spot; side margins, apex of triangle spread. Sweden: northwards to LU and NB. Norway: and windows near ocelli yellow. Genae wider than Ø,AK,HEs,HEn,Bø,VE,VAy,Ry,HOi,SFi,TRy,Fv, first flagellomere. Basal segments of antennae yel- Fn. Finland: northwards to ObS. Russia: Vib, Kr. Eurasian polyzonal species, ranging southwards low, small first flagellomere yellow darkened above, to Italy and Bulgaria and eastwards to East Siberia rarely blackish. Arista black. Palpi long yellow. Scu- and Mongolia. tum with 5 black thinly dusted stripes. Scutellum Biology. – Adults in June and July, rather common yellow. Spot on katepisternum black, shining. Ab- everywhere in meadows. domen yellow with narrow black bands on tergites. Larvae phytophagous, host plants Holcuslanatus, Legs yellow, last tarsal segments darkened. Body Poa pratensis (Tschirnhaus, 1981, 1992) and proba- length 2.5–3.5 mm. bly other grasses.

181 150. Chlorops scutellaris (M. Koponen), 1 female; Apalholm, Lemland, 12– (Zetterstedt, 1838) 24.viii.1954 (H. Lindberg), 1 male, 1 female. Not Figs 673–676 found in Russian parts of Fennoscandia. Euro-Mediterranean species, known from mid- Oscinis scutellaris Zetterstedt, 1838: 779. Lectotype dle and southern Europe, Israel and Algeria. in ZMLU (Andersson, 1966). Biology. – Adults in meadows in July and again in Chlorops laevicollis Becker, 1910: 89. Holotype in August to September, likely in two generations. ZISP (Nartshuk, 1998). Oscinis freyi Duda, 1933: 199. Lectotype in NHMH (Frey, 1944; Nartshuk, 1998). 152. Chlorops speciosus Rather dark species. Frons brown, wider than long. Meigen, 1830 Ocellar triangle nearly reaching front margin of Figs 679–683, 1002–1004 frons, entirely black, shining. Genae narrower than first flagellomere. Antennae entirely black, includ- Chloropsspeciosus Meigen, 1830: 146; Meigen’s plate ing arista. Palpi black. Scutum entirely black, shin- 159: 1 (Morge, 1976a). Lectotype in NHMW ing. Scutellum and pleura darkened. Spot on (Nartshuk, 1997a). katepisternum black, shining. Abdomen brown to black. Legs mainly black. Body length 2.0–2.5 mm. Frons wider than long. Ocellar triangle extending 2/3–3/4 length of frons, black or brown smooth Distribution. – Sweden northwards to LY. Norway: shining with very distinct borders. Genae distinctly Fn:TanaBru.Finland:northtoLi.Russia:Vib,Lr. wider than first flagellomere with many black setu- Not found in Denmark. lae. Basal segments of antennae dark, first flagel- Northern Eurasian species, more likely of Sibe- lomere black. Arista black. Palpi small yellow. Scu- rian origin, known from North Europe eastwards to tum with 3 broad black, shining stripes (in dark East Siberia. specimens from northern Fennoscandia stripes Biology. – AdultsinJulyinwetmeadowsandfens. fused), covered with black setulae. Scutellum yel- low or darkened. Spot on katepisternum black, shining. Abdomen yellow with black bands on ter- 151. Chlorops serenus gites or blackish. Legs partly black. Body length 3.5–4.5 mm, some females up to 6.0 mm. Loew, 1866 Figs 677, 678 Distribution. – The most common species of Chlo- rops occurring throughout Denmark and Chlorops serenus Loew, 1866: 62. Syntypes in ZMHU Fennoscandia. (Wendt, 1975). Eurasian species, found commonly in Europe Frons wider than long. Ocellar triangle extending southwards to Spain and Bulgaria, ranging east- nearly to front margin of frons, yellow with small wards to Altai, West Siberia. median black spot, side margins, apex of trian- Biology. – Adults in June to August in wet mead- gle and two windows near ocelli yellow. Genae ows. distinctly wider than first flagellomere. Basal seg- Larvae phytophagous, live in stems of Deschamp- ments of antennae yellow, first flagellomere black. sia caespitosa and hibernate basally in the stem of Arista black. Palpi yellow. Scutum with 5 black, their host plant. dusted stripes, covered with yellow setulae. Scutel- Larvae described by Balachowsky & Mesnil lum yellow. Spot on katepisternum yellow, rarely (1935), Nye, (1958) and Zhabinskaya (1963), de- blackish above, shining. Abdomen yellow with nar- scriptions of different authors differ in some de- row black bands on tergites and median black line. tails. Legs yellow, last tarsal segments darkened. Body length 3.0–3.5 mm. Distribution. – Scarce. Denmark: SJ, F, LFM, NEZ, B. 153. Chlorops strigulus Southern part of Sweden: SK, HA, and also UP. (Fabricius, 1794) Norway: AK, AAy. Finland: Al: Kökar, 11.vi.1978 Figs 684–690, 986, 987

182 Figs 673–683. Chlorops scutellaris (Zetterstedt): 673: head, dorsal view. 674: head, lateral view. 675: hypopygium, dorsal view. 676: hy- pandrium and phallic complex. C. serenus Loew: 677: head, lateral view. 678: hypandrium and phallic complex. C. speciosus Meigen: 679: head, dorsal view. 680: head, lateral view. 681: mesolobus and surstyli. 682: male genitalia, ventral view. 683: apex of ovipositor, ventral view.

Musca strigula Fabricius, 1794: 334. Type in MNHN Distribution. – Rare. Denmark: EJ, SZ, NEZ, B. Swe- (Zimsen, 1964). den: UP: Hässelby, Börje, 09.v.1959 (L. Hedström), 1 male. Not found in Norway, Finland or Russian Frons wider than long. Ocellar triangle reaching parts of Fennoscandia. front margin of frons, black dull and sulcate along Eurasian species. side margins and on apex. Genae distinctly wider than first flagellomere, yellow with black lower Biology. – Adults from end of May to June, in dry margin. Basal segments of antennae brown, first meadows. flagellomere longer than wide and black. Arista Phytophagous larvae produce top galls on shoots thick white pubescent with basal segment dark. of Brachypodium sylvaticum and B. retusum (Séguy, Palpi black. Scutum with 5 black, dusted stripes, 1934; Balachowsky & Mesnil, 1935; Nye, 1958). of which 3 central stripes are broad and some- Baudyš (1912) described galls on Elytrigia repens times fused before scutellum. Scutellum darkened. that in his opinion were produced by C. strigulus, Anepisternum with many black setulae. Spot on but probably the gall former was C. novakii Strobl, katepisternum black, dusted. Abdomen dorsally a species not found in the Nordic countries. brown to black with narrow yellow bands on ter- Larvae described by Rübsaamen (1895), Bala- gites. Legs mainly black. Body length 2.5–3.5 mm. chowsky & Mesnil (1935) and Nye (1958).

183 Figs 684–695. Chlorops strigulus (Fabricius): 684: head, dorsal view. 685: head, lateral view. 686: antenna. 687: wing. 688: hypopygium, ventral view. 689: hypandrium and phallic complex. 690: posterior end of male abdomen, lateral view. C. troglodytes (Zetterstedt): 691: male genitalia, ventral view. C. varsoviensis Becker: 692: head, dorsal view. 693: head, lateral view. 694: hypopygium, ventral view. 695: hypandrium and phallic complex.

154. Chlorops troglodytes northwards to NTi. Finland: northwards to Ks. Rus- (Zetterstedt, 1848) sia: Vib, Kr, Lr. Fig. 691 Eurasian polyzonal species, known from Great Britain eastwards to Yakutia, East Siberia and Mon- Oscinis troglodytes Zetterstedt, 1848: 2626. Lecto- golia. type in ZMLU (Andersson, 1966). Biology. – Adults in June to August in meadows. Frons wider than long. Ocellar triangle extending half length of frons, entirely black, shining. Genae wider than first flagellomere. Basal segments of an- 155. Chlorops varsoviensis tennae brown, first flagellomere and arista black. Becker, 1910 Palpi small yellow. Scutum with 5 black thinly Figs 692–695 dusted and slightly shining stripes. Scutellum yel- low. Spot on katepisternum black, shining. Ab- Chlorops varsoviensis Becker, 1910: 92. Syntype, 1 domen dorsally brown to black with narrow yel- male, in ZMHU (Wendt, 1975). low bands on tergites. Cerci of ovipositor black. Legs yellow with darkened apical tarsomeres. Body Ocellar triangle medially with a large poplar leaf- length 2.0–2.5 mm. shaped black spot. Genae narrower than width of first flagellomere. Antennae usually black or first Distribution. – Rather common. Denmark: WJ, flagellomere reddish ventrobasally. Palpi blackish. NWJ, LFM. Sweden: northwards to NB. Norway: Scutum with 5 black stripes, sparsely grey-dusted.

184 Pleura with black spots. Anepisternum with setu- Yellow species with black stripes on scutum and lae, white in males and black in females. Abdomen black spots on pleura. Head broader than tho- reddish-brown in females. Legs almost yellow with rax. Eyes large, genae linear and postgenae nar- apical tarsomeres darkened. In male front basitar- row. Ocellar triangle smooth shining with convex susshort,about2.0–2.5timesaslongaswideand sides and reaching front margin of frons, usually front inner claw larger and more curved than outer with a row of setulae on its surface along side mar- one. Body length 3.0–3.5 mm. gins. First flagellomere orbicular or slightly longer than wide. Scutum shining or rarely dusted. There Distribution. – Sweden: northwards to TO. Norway: are band-like sclerites along anus. Male genitalia: Fi:Gargia,Alta.Finland:northwardstoLi.Russia: mesolobus small; surstyli closely alligned to epan- Vib, Kr. Lr. Not so far recorded from Denmark. drium; pre- and postgonites situated in line. Tibial Trans-Palearctic species, known from France to organ present. Japan, southwards to Spain, North Italy and Bul- garia, eastwards to Mongolia. Distribution. – Probably worldwide. Biology. – Adults in July and August in wet habitats with Carex. 157. Chloropsina distinguenda Larvae phytophagous develop in stems of Carex. (Frey, 1909) Figs 696–699 156. Chlorops zernyi Chlorops distinguendus Frey, 1909: 11. Holotype in (Duda, 1933) NHMH.

Oscinis zernyi Duda, 1933: 201. Holotype female in Head wider than thorax. Frons narrowed in front. NHMW. Ocellar triangle large, smooth black, shining, reach- ing front margin of frons. vte setae on yellow spots. Body yellow. Frons as long as wide, setae and se- Genae linear, not wider than palpus. Antennae tulae black. Ocellar triangle large, reaching front including arista and palpi black. Scutum nearly margin of frons, but narrow in distal third, smooth square, black, shining, scutal stripes nearly fused. shining, mainly black, but yellow along sides, hind Scutellum yellow. Pleura with several black, shining angles and on tip, wrinkled on hind angles. Genae marks. Abdomen black dorsally and yellow ven- a little wider than first flagellomere. Parafacialia as trally. Legs mainly yellow with darkened femora wide as gena. Basal segments of antennae brown, and tarsi. Body length 2.3 mm. first flagellomere black. Arista white, thickly pubes- cent. Palpi large, blackish on tip. Scutum with 5 Distribution. – Very rare. Sweden SK: Hässleholm, black, dusted stripes, covered with yellow setulae, 21.vii.1981 (H. Andersson). Finland: Al: Tinström, not extended to scutellum. Scutellum yellow. Spot Sund(R.Frey),femaleholotype;Ab:Jurmo,Kor- on katepisternum yellow-brownish, shining. Ab- po, 04.viii.1960 (Meinander); N: Linna, Helsinki (R. domen yellow with narrow black bands on tergites, Frey), 1 female; Oa: Berga, 17.vi.1946 (Hellén). covered with black setulae. Legs yellow. Cerci of fe- Elsewhere only recorded from the Czech Repub- male yellow. Body length 4.0–4.5 mm. lic. Distribution. – Russia: Vib: Beloostrov, 15.viii.1956 (Nartshuk), 2 females. 158. Chloropsina rohaceki Eurasian species. Nartshuk, 2000 Figs 700–702 Biology. – Usually in Carex beds. Chloropsina rohaceki Nartshuk, 2000a: 256. Holo- type in ZMOC, paratypes in ZISP and ZMLU. Genus Chloropsina Becker, 1911 Head wider than thorax. Frons longer than wide. Ocellar triangle large, smooth black, shining, reach- Chloropsina Becker, 1911: 51. Type species: Chlo- ing front margin of frons. vte setae on yellow spots. ropsina oculata Becker, 1911 by designation of Genae narrower than palpus. First flagellomere yel- Malloch, 1931: 76. low, darkened dorsally. Arista pale. Palpi yellow.

185 Figs 696–702. Chloropsina distinguenda (Frey): 696: head, dorsal view. 697: head, lateral view. 698: hypopygium, dorsal view. 699: male genitalia, ventral view. C. rohaceki Nartshuk: 700: head, dorsal view. 701: hypopygium, dorsal view. 702: male genitalia, ventral view.

Scutum nearly square, with 5 black, shining stripes, Black species, only frons, antennae and legs partly median stripe reaching scutellum as black spot yellow. Head setae well developed, vte longer than with indistinct margins. Scutellum yellow with vti. Ocellar triangle large, broad behind, almost black side margins. Pleura with several black, shin- reaching front margin of frons, with reticulate mi- ing marks. Abdomen black dorsally and yellow ven- crosculpture, not dusted. Face with carina, trian- trally. Legs entirely yellow. Body length 2 mm. gular between antennae and narrow below. First flagellomere about as long as wide with pointed up- Distribution. – Only Sweden: BL: Kristianopel, per corner. Scutum black with faint granulate mi- 08.vii.1987 (R. Danielsson). crosculpture, dusted and evenly covered with nu- European species, known elsewhere from Slo- merous setulae. Femoral and tibial organs present. vakia. Male genitalia: middle lobe of surstylus granu- late. No sclerites lateral to anus. Postgonites short, connected with long pregonites. Distiphallus long Genus Cryptonevra Lioy, 1864 complex with coronated sclerites. Contrary to Neo- haplegis testes in male tubular as usual in Chloropi- nae, not round. Cryptonevra Lioy, 1864: 1125. Type species: Chlorops flavitarsis Meigen, 1830 by monotypy. Distribution. – Palearctic and Oriental Regions.

186 Key to species of Cryptonevra Lioy Similar to C. flavitarsis, but differs in having paler front tibiae and all short pubescence of head, tho- rax, abdomen and legs pale. Only vte setae on head; 1 Frons yellow in front. Male genitalia Fig. 711 thoracic and scutellar setae black. Frons slightly ...... C. diadema (Meigen) longer than wide, black (rarely with reddish-brown – Fronsentirelyblack...... 2 anterior margin). Ocellar triangle black, nearly 2 Setulae on head, scutum, abdomen and legs reaching front margin of frons. Its surface minutely black. Scutum shining. Front tibiae black. punctuate and subshining. Basal segments of an- Male genitalia Figs 709, 710 tennae black or rarely reddish brown. First flagel- lomere slightly longer than wide, distinctly angu- ...... C. flavitarsis (Meigen) lated dorsodistally,black. Arista black and minutely – Setulae on head, scutum, abdomen and legs pubescent basally, distally glabrous. Scutum black, pale. Scutum dull. Front tibiae pale. Male gen- minutely shagreened, dusted on notopleura and italia Figs 703, 704 . . . . . C. consimilis (Collin) before scutellum, evenly clothed with pale setu- lae. Scutellum grey-dusted. Pleura mainly shin- ing. Wings slightly whitish. Veins R4+5 and M di- Species key to known 3rd instar larvae vergent distally. Abdomen basally brownish yel- low, remaining part black grey-dusted and covered of Cryptonevra with white setulae. Femora black except for yellow knees, tibiae and tarsi yellow. Male with tarsal seg- Mature larvae white cylindrical 3–5 mm long. Fa- ment 4 of middle legs with black setae on ventral cial mask with linear branched ridges. Hypostomal surface. Body length 2.5–2.8 mm. and tentoropharyngeal sclerites separate. Mandi- bles with small accessory teeth in two rows. Body Distribution. – Known only from southern Sweden: with numerous lines of spicules of different size SK: Krankesjön, 30.iv.1986, 11.v.1986 (B. Stille); SM: on locomotory creeping welts on ventral side of Kalmar, 07.v and 10.v.1986 (B. Stille). body. Anal opening surrounded by spicules. Ante- European species. rior spiracles of fan type with 4–9 buds. Posterior Biology. – The larvae live as inquilines in galls of spiracles on stigmophores and with branched in- Lipara similis on Phragmites australis. They feed in- terspiracular processes. side the hollow of the reed stem, and before pupa- tion most of them pass down below the L. similis 1 Anterior lines of spicules on ventral locomo- larvae (Collin, 1932). tory creeping welts of abdominal segments continuous (Figs 721, 722). Anterior spiracles with 6–8 buds (Fig. 720) 160. Cryptonevra diadema ...... C. diadema (Meigen) (Meigen, 1830) – Anterior lines of spicules on ventral locomo- Figs 711, 719–726 tory creeping welts of abdominal segments Chlorops diadema Meigen, 1830: 158; Meigen’s plate not continuous, separated laterally (Figs 715, 211: 3 (Morge, 1976b). Syntype, a female in 716). Anterior spiracles with 4–9 buds (Fig. NHMW (Ismay, 1994a). 713)...... C. flavitarsis (Meigen) Frons slightly longer than wide, broadly reddish yellow on anterior part and black posterad. Ocel- lar triangle predominantly black, reddish brown on 159. Cryptonevra consimilis apex and nearly reaching front margin of frons. Its (Collin, 1932) surface minutely punctuate and subshining. Basal Figs 703, 704 segments of antennae reddish brown. First flagel- lomere slightly longer than wide, distinctly angu- Haplegisconsimilis Collin, 1932: 113. Lectotype male lated dorsodistally, basally and ventrally reddish and 1 male, 1 female paralectotypes in OXUM, brown to yellow, black dorsally. Arista black and 3 males, 2 females paralectotypes in BMNH, all minutely pubescent basally, distally glabrous. Scu- bread from reed stem inhabited by Liparasimilis tum black, minutely shagreened, dusted on noto- (Ismay, 1994a; Pont, 1995). pleura and before scutellum, evenly clothed with

187 Figs 703–711. Cryptonevra consimilis (Collin): 703: hypopygium, dorsal view. 704: hypandrium and phallic complex. C. flavitarsis (Meigen): 705: head, dorsal view. 706: head, lateral view. 707: antenna. 708: wing. 709: hypopygium, ventral view. 710: hypandrium and phallic complex. C. diadema (Meigen): 711: male genitalia, ventral view. (703, 704 after Ismay, 1994.) yellowish setulae. Scutellum grey-dusted. Pleura ThelarvaeliveinshootsofPhragmites australis, mainly shining. Wings slightly whitish. Veins R4+5 usually as inquilines in Lipara galls. Egg, all lar- and M divergent distally. Femora black except for val stages and pupurium described by Grichowska yellow knees, anterior tibiae yellow, all tarsi yellow. (2008c). Male with tarsal segments 2–4 of middle legs with black setae on ventral surface. Abdomen basally brownish yellow, remaining part black grey-dusted. 161. Cryptonevra flavitarsis Body length 2.5–2.8 mm. (Meigen, 1830) Figs 705–710, 712–718 Distribution. – Denmark:SJ,WJ,NWJ,NEJ,F,NEZ. Sweden: SK, BL, HA, SM, VG, UP. Finland: north to Chlorops flavitarsis Meigen, 1830: 161; Meigen’s Om along the Baltic coast. Russia: only Vib. Not re- plate 211: 6 (Morge, 1976b). Lectotype in NHMW corded from Norway. (Nartshuk, 1997a). Trans-Palearctic species, known from Europe to Japan, in southern part of Eurasia and in North Frons slightly longer than wide, black. Ocellar tri- Africa. angle black dull, nearly reaching front margin of frons. Genae nearly as wide as first flagellomere. Biology. – Adults in May to July. First flagellomere slightly longer than wide, dis-

188 Figs 712–726. Cryptonevra flavitarsis (Meigen), 3rd instar larva: 712: cephalopharyngeal sclerite. 713: anterior spiracle. 714: posterior spiracle. 715: spicules on 2nd abdominal segment. 716: spicules on 7th abdominal segment. 717: posterior end. 718: puparium. Cryptonevra diadema (Meigen), 3rd instar larva: 719: cephalopharyngeal sclerite. 720: anterior spiracle. 721: spicules on 2nd abdominal segment. 722: spicules on 7th abdominal segment. 723: posterior spiracle. 724: mandibula. 725: posterior end. 726: puparium. (726 after Grochowska, 2008b.)

189 tinctly angulated dorsodistally, black. Arista black fused into a rigid aedegal complex which articu- and minutely pubescent. Scutum black, minutely lates with hypandrium and probably functions as shagreened and shining, thinly dusted on noto- “pseudophallus”. pleura and before scutellum, evenly clothed with blackish setulae. Scutellum grey-dusted. Pleura Distribution. – Holarctic, Australia. mainly shining. Wings slightly greyish. Veins R4+5 and M divergent distally. Femora black except for 162. Diplotoxa messoria yellow knees, anterior tibiae black, tarsi yellow or brown. Male with tarsal segment 4 (rarely also (Fallén, 1820) 3) of middle legs with black setae on ventral sur- Figs 727–731 face. Abdomen basally brownish yellow, remaining Oscinis messoria Fallén, 1820: 5. Lectotype in ZMLU part black grey-dusted, abdominal setae dark. Body (Andersson, 1963). length 1.5–2.5 mm. Frons as long as wide, yellow. Ocellar triangle black, Distribution. – Common. Denmark: from all dis- shining, extending nearly to front margin of frons, tricts except SZ and NWZ. Sweden: north to JÄ. Nor- behind nearly touching eyes. Genae and face yel- way: Ø, AK, VE. Finland: north to Om. Russia: Vib, low, genae a little narrower than first flagellom- Kr. ere. Antennae black, first flagellomere round, arista Trans-Palearctic species, known from the British white. Palpi yellow. Scutum with 3 wide, sometimes Isles eastwards to China and southwards to North nearly fused black, dusted stripes, which extend Africa. to scutellum. Pleura yellow with black spots on Biology. – Adults from the end of April to beginning anepisternum, katepisternum and katepimeron. of August on Common reed (Phragmites australis). Scutellum brown to black, apical setae shorter than The larvae develop as inquilines in Lipara galls scutellum. Abdomen dark brown to black, shining. (Giraud, 1863), repeated by many authors. Recently Femora and hind tibiae black, front and middle Reader (2003) characterized the activity of the tibiae fuscous yellow, tarsi darkened. Body length Cryptonevra flavitarsis larvae in the galls of Lipara 2.5–3.0 mm. lucens (wrongly given as L.rufijitarsis?) as “intraguild Distribution. – Denmark: from most districts. Swe- kleptoparasitism”. This seems questionable, how- den: north to NB along coast of Baltic Sea. Norway: ever, as the larvae of these two species feed on AK: Oslo (Siebke). Finland: north to ObS. Russia: different parts of the plant, and it does not seem Vib, Kr. that the presence of Cryptonevra larvae affects the Holarctic species, in the Palearctic from the Brit- well-being of the gall-inducing L. lucens larvae. ish Isles to the Far East of Russia. Egg, all larval stages and pupurium described by Grichowska (2008a). Biology. – Adults from middle of June to the be- ginning of September, on marshy habitats with Eleocharis. Some specimens were reared from this plant Genus Diplotoxa Loew, 1863 moved into laboratory (Nartshuk, 1962a). The spe- cies develops in rotting stems of Eleocharis macro- Diplotoxa Loew, 1863: 54. Type species: Diplotoxa stachya in North America (Valley et al., 1969). versicolor Loew, 1863 by monotypy. Tschirnhaus (1981) listed as host plants Eleocharis palustris and E. uniglumis. Other Nearctic species Species of Chlorops-like general shape. Body mainly of Diplotoxa are also associated with species of yellow with black nearly confluent scutal stripes, Eleocharis (Wearsch & Foote, 1994). irregular transversely striated microsculpture on scutum. Ocellar triangle large, almost reaching front margin of frons, shining. Anterior ommatidia larger than posterior ones. The wings have veins Genus Diplotoxoides Andersson, 1977 R2+3 and R4+5 strongly procurved, R4+5 and M divergent and crossveins approximated. Middle Diplotoxoides Andersson, 1977: 150, as subgenus tibiae with slightly curved spur. Tibial organ ab- of Diplotoxa Loew. Type species: Diplotoxa dal- sent. Male genitalia: phallus, pre- and postgonites matina Strobl, 1900 by original designation.

190 Figs 727–731. Diplotoxa messoria (Fallén): 727: head, dorsal view. 728: head, lateral view. 729: wing. 730: hypopygium, ventral view. 731: hypandrium and phallic complex.

Figs 732–736. Diplotoxoides dalmatina (Strobl): 732: head, dorsal view. 733: head, lateral view. 734: wing. 735: male genitalia, ventral view. 736: hypandrium and phallic complex.

Body yellow with black stripes and spots. Head 163. Diplotoxoides dalmatina as high as long. Anterior ommatidia larger than (Strobl, 1900) posterior ones. Ocellar triangle large smooth shin- Figs 732–736 ing. Head setae well developed. First flagellomere rounded. Scutum smooth, dusted. The wings have Diplotoxa dalmatina Strobl, 1900: 628. Syntypes veins R2+3 and R4+5 procurved, R4+5 and M di- not found in NMBA or the Sarajevo Museum vergent and crossveins approximated. Middle tib- (Chvála, 2008). iae with slightly curved spur. Tibial organ absent. Male genitalia: phallus, pre- and postgonites fused Frons yellow, as long as wide in the middle and nar- into aedegal complex which articulates with hy- rowed in front. orb 6–7 short recurved. poc shorter pandrium. than oc. Ocellar triangle black, shining, extend- ing nearly to front margin of frons, behind nearly Distribution. – Not clear, Palearctic? touching eyes. Genae and face yellow, genae a little

191 Figs 737–743. Epichlorops puncticollis (Zetterstedt): 737: head, dorsal view. 738: head, lateral view. 739: antenna. 740: wing. 741: head and thorax, dorsal view. 742: hypopygium, ventral view. 743: hypandrium and phallic complex. narrower than first flagellomere, narrowed in front. Pleura smooth yellow. Wings of Chlorops-type. Tib- Antennae black, first flagellomere round, arista ial organ absent. Male genitalia: pre- and postgo- black. Palpi yellow. Scutum with wide, usually nites situated parallel to each other. fused black thinly dusted subshining stripes, which Distribution. – Holarctic Region. extend to scutellum. Scutellum often darkened, apical setae longer than scutellum, wide apart. Pleura yellow with black spots on anepisternum, 164. Epichlorops puncticollis katepisternum and katepimeron. Abdomen dark (Zetterstedt, 1848) brown. Coxae yellow, femora and tibiae mainly Figs 19, 737–743 black, tarsi darkened. Body length 1.5 mm. Distribution. – Rare species. Denmark: NWJ: Thy Oscinis puncticollis Zetterstedt, 1848: 2636. Lecto- National Park. Sweden: SK. Finland: AB: Lohja; N: type in ZMLU (Andersson, 1966). Hanko, Helsinki; Om: Nykarleby, Pietarsaari. Rus- Frons as long as wide, yellow.Ocellar triangle reach- sia: Kr: Bolshie Gory. Not recorded yet in Norway. ing front margin of frons, black smooth shining, Trans-Palearctic species, found from British Isles anterior apex obtuse. Genae slightly wider than to Far East of Russia. first flagellomere. Antennae and arista black. Palpi Biology. – Adults from June to August, in wetlands. yellow. Scutum longer than wide, black except for lateral margins, undusted with strong stellate mi- crosculpture and evenly clothed with black setulae. Postpronota yellow with black spot. Scutellum yel- Genus Epichlorops Becker, 1910 low, apical setae rather approximated. Pleura yel- low with several black spots. Abdomen completely Epichlorops Becker, 1910: 77. Type species: Oscinis black dorsally except for extreme lateral margins. puncticollis Zetterstedt, 1848 by original desig- Legs mainly yellow, hind tibiae medially with black nation. band and all tarsi black, sometimes legs extensively darkened. Body length 2.6–3.5 mm. Body black with partly yellow head, pleura and legs. Head as long as high with short setae and setu- Distribution. – Common species. Denmark: wide- lae. Ocellar triangle very large, behind almost as spread, but absent from northern districts of Jut- broad as frons, reaching front margin of frons in a land. Sweden: north to NB. Norway: VE: Tønsberg rather broad tip. First flagellomere rounded. Scutel- (Siebke). Finland: north to Om. Russia: Vib, Kr. lum rounded, black with stellate microsculpture. Holarctic species.

192 Biology. – Adults from June to the beginning of Sep- jected. Ocellar triangle reaching front margin of tember in wet meadows and marshes with Carex. frons, with median black sulcus and coarse black In North America E. exilis (Coquillett) was reared punctuations. Genae broad and punctuate. Anten- from Carex crinita (Rogers et al., 1991). nae from brown to black, scapus long triangular, first flagellomere 1.5 times longer than wide. Scu- tum longer than wide, with four black punctu- Genus Eurina Meigen, 1830 ate stripes. Scutellum rounded, apical setae very short. Pleura densely dusted. All thoracic setae short. Wings grey brownish. Halteres fuscous yel- Eurina Meigen, 1830: 3. Type species: Eurina lurida low. Body length 6.0 mm. Meigen, 1830 by designation of Westwood, 1840: 147. Distribution. – Only from Denmark: SJ: Sønderborg, Body brown dull elongated. Head longer than high vi.1900 (W. Wüstnei), 1 male, 1 female; Sottrupskov, with frons strongly projecting, about as long as eye. 12 and 15.vi.1893, 08.vi.1895, 04.vi.1898 (W. Wüst- Face retreating. Eyes small rounded. Ocellar trian- nei), 2 males, 4 females. gle large, reaching front margin of frons, with longi- Euro-Mediterranean species. tudinal groove, surface covered with small setulae. Head setae very small. Antennae prolonged, first 166. Eurina lurida flagellomere trapezoid, longer than wide. Scutum flattened, evenly covered with short setulae. Scutel- Meigen, 1830 lum rounded flat. Tibial organ absent. Male gen- Figs 749–751 italia: pre-, postgonites, phallapodeme, and phal- Eurina lurida Meigen, 1830: 5; Meigen’s plate 146: lapodemic sclerites fused into a very broad disc-like 11 (Morge, 1976a). Lectotype male in MNHN structure. (Nartshuk, 2006). Distribution. – Probably worldwide. Body brown dull. Head longer than high with frons strongly projecting, longer than wide. Face re- treating. Eyes small rounded, bare. Ocellar tri- Key to species of Eurina Meigen angle large, straight-sided, reaching front margin of frons, with a conspicuous central groove and 1 Body dull grey, scutum with numerous lack sulci along side margins. Ocellar tubercle black. shining spots at base of setulae. Ocellar tri- Genae as wide as eye, covered with small setulae. angle wide with central sulcus and numer- Antennae prolonged, first flagellomere trapezoid, ous punctures at base of setulae, shining along longer than wide, brown to black, arista black. Scu- central sulcus and dusted on sides (Fig. 744). tum flattened with 3 black, dusted longitudinal Male genitalia Figs 747, 748 . . . . E. calva Egger stripes, evenly covered with short setulae. Scutel- – Body reddish with distinct black stripes on lum rounded flat with short apical setae. Anepis- ternum covered with many short black setulae. scutum and spots at base of setulae. Ocellar Abdomen reddish-brown dorsally, yellow ventrally. triangle shorter with central sulcus and nu- Wings slightly brownish. Legs yellow. Body length merous punctures at base of setulae, shining 6.0–6.5 mm. onwholesurface(Fig.749).MalegenitaliaFigs 750,751...... E. lurida Meigen Distribution. – Denmark: LFM: Ulfshale, 27– 29.v.1983 (Andersen & Michelsen), 2 females. Swe- den: SK: Klagshamn, 25.v.1981 (H. Andersson), 2 165. Eurina calva males; BO: Bro, 17.vii.1945 (E. Wieslander), 1 male. Egger, 1862 Norway: AK: Konglungen, Asker, 04.vi.1997 (G.E.E. Figs 744–748 Söli). Not found in Finland or Russian parts of Fennoscandia. Eurina calva Egger, 1862: 778. Lectotype in NHMW Euro-Mediterranean species. (Nartshuk, 1997b). Biology. – Adults on Common reed (Phragmites Body brown covered with thick greyish dust. Head australis) and other plants in coastal, saline habi- nearly triangular in side view. Frons strongly pro- tats (Cole, 1980; Allen, 1982; Alexander, 1983).

193 Figs 744–751. Eurina calva Eggers: 744: head, dorsal view. 745: head, lateral view. 746: wing. 747: male genitalia, ventral view. 748: hypopy- gium, dorsal view. E. lurida Meigen: 749: head, dorsal view. 750: male genitalia, ventral view. 751: hypopygium, dorsal view.

The larvae phytophagous, but data on host plant shorter than antennae. Scutum moderately con- controversial: Séguy (1934) mentioned Phragmites vex, smooth. Scutellum rounded. All setae and se- australis as host plant. Tschirnhaus (1981) records tulae white. Wings of Chlorops-type. Middle tib- Bolboschoenus maritimus. In Israel larvae of this iae with spur. Tibial organ present. Male genitalia: species develop on Eleocharis dulcis without pro- surstyli with pointed hook-like middle lobe; prego- ducing top galls (Kaplan et al., 1986), but Ismay nites straight, postgonites short curved. (1999), based on pers. comm. from A. Freidberg, listed Bolboschoenus maritimus as host plant but Distribution. – Palearctic, Afrotropical and Aus- failed to find galls on that plant in England. tralasian Regions.

Genus Eutropha Loew, 1866 Key to species of Eutropha Loew

Eutropha Loew, 1866: 26. Type species: Chlorops ful- 1 Pleura and scutellum yellow. Scutum with vifrons Haliday, 1833 by designation of Becker, grey-dusted stripes fused in some specimens 1910: 59...... E. variegata Loew Head as long as high with short setae. Ocellar tri- – Thorax and abdomen black, shining. Only angle smooth shining. Genae very wide. First flag- sides of scutum a little dusted. Male genitalia ellomere large rounded squarish. Arista very short, Fig.756...... E. fulvifrons (Haliday)

194 Not so far recorded in Russian parts of Fennoscan- dia. Biology. – Adults from May to October on sand dunes along sea coast, preferable at the fore dune on the Cakile-Salsola vegetation (Ardö, 1957). Séguy (1934) found puparia in sand on sea coast. Tschirnhaus (1981) proposed Ammophila arenaria as host plant. The adults of both sexes visited baits with pyrrolizid alkaloid and ingested these sec- ondary plant chemicals (Boppré & Pitkin, 1998). Larvae of the tropical species E. lindneri Sabrosky were found in a dead seal on a sea shore in Africa (Kirk-Spriggs et al., 2001)

168. Eutropha variegata Loew, 1866

Eutropha variegata Loew, 1866: 28. Type material not found in ZMHU (Wendt, 1975, 1978). Figs 752–756. Eutropha fulvifrons (Haliday): 752: head, dorsal Frons as long as wide, yellow with white setae view. 753: head, lateral view. 754: antenna. 755: wing. 756: male and setulae. Ocellar triangle extending nearly to genitalia, ventral view. front margin of frons, yellow shining with me- dian sulcus. Ocellar tubercle black. Face and genae yellow. Genae in male as wide as first flagellom- 167. Eutropha fulvifrons ere, in female 1.2 times wider than first flagel- (Haliday, 1833) lomere. Antennae yellow in male and black in fe- Figs 752–756 male. First flagellomere large, especially in male, with upper corner. Arista shorter than antennae, Chlorops fulvifrons Haliday, 1833: 172. Lectotype in black or whitish, nearly bare. Palpi yellow. Scu- NMID (Ismay et al., 2001). tum yellow with 5 blackish densely grey-dusted and white pubescent stripes. Scutellum yellow. Body almost entirely shining brownish-black with Pleura with black marks. Setae of thorax white. Ab- white setae and setulae. Frons longer than wide, domen brown, white pubescent. Halteres yellow. yellow in front and brown behind, or entirely Body length 2.0–2.5 mm. brown, shining with white setae and setulae. Ocel- lar triangle extending 2/3 length of frons followed Distribution. – Rare species. Norway: Ø: Larkollen, by linear extension to front margin of frons, black, 06.vii.1955 (Ardö), 1 female. Finland: Ab: Vihti; N: shining. Face and genae yellow, the latter with Tvärminne, Hanko. Russia: Vib. Not so far found in black below margin. Genae 1.2–2 times as wide as Denmark and Sweden. first flagellomere. Antennae black or first flagel- Palearctic species, known from Europe to Central lomere yellowish below. First flagellomere round Asia. or slightly angulated. Arista shorter than antennae, Biology. – Adults in July and August on sand dunes black, nearly bare. Palpi yellow. Scutum entirely along sea coasts. Flies were found under leaves of black, shining, scutellum from yellow to black. Se- Honckenya peploides. tae and setulae of thorax white. Abdomen black, It is known that some tropical species of the shining, white pubescent. Legs mainly black. Hal- genus are also associated with sea coast and de- teres yellow. Body length 2.0–2.5 mm. velop in mammalian carrion (Kirk-Spriggs et al., Distribution. – Denmark: WJ, NWJ, NEJ, NWZ, NEZ. 2001). Sweden:SK,HA,ÖL,GO,NB.Norway:Ø,AK,VE,Ry. Finland: N: Hanko, 25.vii.1927 (Krogerus), 1 male.

195 Genus Lasiosina Becker, 1910 5 Male genitalia: surstyli shorter and their tip dentate and directed to the middle (Figs 767– Lasiosina Becker, 1910: 73. Type species: Chlorops 771)...... L. herpini (Guérin-Méneville) cinctipes Meigen, 1830 by original designation. – Male genitalia: surstyli longer and directed straight (Figs 761, 762) Body yellow with black stripes and spots. Head as ...... L. danielssoni Nartshuk & Andersson long as high with well developed setae. 2–3 orb setae long. Ocellar triangle narrow, extending half length of frons followed by linear extension to front Species key to known 3rd instar larvae margin of frons, often with black rhomboid spot at tip, dusted but with shining areas at sides be- of Lasiosina Becker hind and on apical black spot. There is sexual di- 1 Body length 3 mm. Mandible with 1 triangu- morphism in colour in some species: antennae and lar middle tooth; dental sclerite more or less palpi in males yellow, in females black. First flagel- lomere rounded. Scutum yellow with black stripes, arched (Fig. 984). On ventral side of abdomen dusted. Wings of Chlorops-type. Tibial organ ab- creeping welts consist of 4–5 incomplete rows sent. Male genitalia: surstyli fused with epandrium, of spicules. In front of perianal opening a row granulate at apex. Postgonites apically curved up- of backward directed spicules (Fig. 985) wards...... L. brevisurstylata Dely-Draskovits – Body length 4 mm. Mandible with 3–4 trape- Distribution. –Worldwide. zoid middle teeth; dental sclerite more or less conical or semicircular (Fig. 982). On ven- Key to species of Lasiosina Becker tral side of abdomen creeping welts consist of 5–8 complete rows of spicules, usually uni- formly situated. In front of perianal opening 1 Ocellar triangle without shining rhomboid no spicules (Fig. 983) spot apically. Wings usually shorter than ab- ...... L. herpini (Guérin-Méneville) domen. Male genitalia Fig. 774 ...... L. parvipennis Duda 169. Lasiosina albipila – Ocellar triangle with shining rhomboid spot apically (Fig. 764). Wings normal, longer than (Loew, 1866) Figs 757–759 abdomen...... 2 2 Only one posterior npl seta. First flagellomere Diplotoxa albipila Loew, 1866: 37. Type material not and palpi in male yellow, in female black. Male found in ZMHU (Wendt, 1975, 1978). genitalia Fig. 760 Head broader than deep or long. Frons wider than ...... L. brevisurstylata Dely-Draskovits long, yellow with black setae and setulae. Ocel- – Two posterior npl setae...... 3 lar triangle extending half length of frons followed 3 Antennae black and palpi yellow in both sexes by linear extension to front margin of frons, yel- or only in males. Abdomen with white or black low with two black spots, one on ocellar tuber- cle and another rhomboid spot on apex. Genae setulae...... 4 wider than first flagellomere. Antennae black, first – Antennae and palpi yellow in male and black flagellomere rounded. Palpi yellow. Scutum with 3 in female. Abdomen with black setulae . . . . 5 black densely grey-dusted stripes, scutellum yel- 4 Abdomen with white setulae, only some setu- low. Pleura with several black marks. Spots on lae black. Male genitalia Figs 757–759 anepisternum and katepisternum shining. Abdo- ...... L. albipila (Loew) men brown, covered with white setulae, but some – Abdomen with black setulae. Male genitalia black setulae present. Legs partly black. Body Figs 772, 773, 775–777 length 2.5–3.0 mm...... L. intermedia Dely-Draskovits, Distribution. – Rare species. Sweden: SK: Ivö klack, L. subnigripes Dely-Draskovits Ivön, 19.v.1998 (R. Danielsson), 1 female; SM; UP.

196 Figs 757–763. Lasiosina albipila (Loew): 757: hypopygium, dorsal view. 758: hypopygium, lateral view. 759: male genitalia, ventral view. L. brevisurstylata Dely-Draskovits: 760: hypopygium, dorsal view. L. danielssoni Nartshuk & Andersson: 761: hypopygium, dorsal view. 762: hypopygium, lateral view. 763: hypandrium and phallic complex (part of hypandrium omitted).

Finland: N: Helsinki, 1 male. Russia: Vib. Not so far shining on upper part and dusted below. npl 1 + 1. found in Denmark and Norway. Abdomen blackish dorsally, covered with black se- European species. tulae. Legs yellow. Body length 2.0–2.6 mm. Biology. – Adults in meadows. It was found in Distribution. – Found only in Finland: St: Eura, Switzerland that adults feed on secreta from chrys- 16.vi.1982 (Koponen), 1 female. omelid larvae of Agelastica alni (Pschorn-Walcher, Eurasian species known eastwards to Yakutia, 1956). East Siberia. Biology. – In Bulgaria reared from shoots of winter 170. Lasiosina brevisurstylata wheat, oats and spring barley together with L. her- pini (Beschovski & Krusteva, 1995, 1997; Krusteva & Dely-Draskovits, 1977 Beschovski, 1998, 2000, 2001). Figs 760, 984, 985 Pupa described by Beschovski & Krusteva (1997). Lasiosinabrevisurstylata Dely-Draskovits, 1977: 267. Holotype and paratypes in HNHM. 171. Lasiosina danielssoni Frons as wide as long, yellow with black setae Nartshuk & Andersson, sp. nov. and setulae. Ocellar triangle nearly 1/3 as wide as Figs 761–763 frons, yellow shining, extending little more than half length of frons with shining black rhomboid Holotype male, Sweden: GO: Lummelunda bruk, spot on apex. Three pairs of long orb. vte longer 30.viii.1992 (R. Danielsson). Holotype in ZMLU. than vti. Genae wider than first flagellomere. In Diagnosis. – New species belongs to L. herpini male antennae and palpi yellow, first flagellomere group (= cinctipes auct.) and is similar to L. cinc- little darkened above. In female first flagellomere tipes in colour of frons, antennae, palpi and legs. and palpi black. Scutum with 3 black grey-dusted It differs by structure of male genitalia with longer stripes, narrowed behind, scutellum yellow. Pleura surstyli with straight ends not curved towards mid- with several black marks. Spot on katepisternum dle and elongated hypandrium.

197 Description. – Male. Body yellow with black grey- Genae wider than first flagellomere. First flagel- dusted stripes on scutum and four spots on pleura. lomere rounded, yellow, darkened around inser- Frons as long as wide, little narrowed anteriorly. tion of arista in male and entirely black in female. Base of ocellar triangle occupies 1/3 of frontal Palpi yellow in male and black in female. Scutum width. Ocellar triangle extending little more than with 3 black densely grey-dusted stripes, scutellum half length of frons followed by linear extension to yellow. Pleura with several black marks. Spot on front margin of frons. Ocellar tubercle black, dull, katepisternum black, completely dusted or shining another black, shining rhomboid spot situated on at anterior and posterior ends in some specimens. apex of triangle. Lateral sides narrow and line be- Abdomen covered with black setulae. Legs yellow. tween anterior ocellus and tip of ocellar triangle Body length 2.5–3.0 mm. brownish. Two long orb. vte, vti and oc nearly equal, poc little shorter. Frons covered with short black Distribution. – The most common species of the setulae. Eyes bare, strongly oblique. Genae wider genus.Sweden:SK,SM,ÖL,GO,ÖG,UP.Norway: than first flagellomere. Antennae yellow, first flag- AK: (Rygg, 1966a, as L. cinctipes). Finland: Ab, N, St. ellomere round, with small dark spot at aristal base. Russia: Vib, Kr. Not so far from Denmark. Arista black. Trans-Palearctic species. Scutum little longer than wide, yellow, with three Biology. – Adults in July to September in dry mead- wide black, dusted stripes. Central stripe narrowed ows and cereal fields. behind and not extended to scutellum. Postpronota The larvae develop in shoots of different grasses, with small brownish spot. Scutellum yellow, api- including cereals: rye, wheat, oats, barley as sec- cal setae longer than scutellum. Pleura with black, ondary invaders, often in the same shoots as Chlo- shining spot on anepisternum and entirely dusted rops pumilionis (Tzygankov, 1929, 1930; Balachow- spot on katepisternum. Remaining two spots black, sky & Mesnil, 1935; Goodliffe, 1939; Nye, 1958; + dusted. npl 1 2. Abdomen brownish dorsally. Legs Krusteva & Beschovski, 2000, 2001). It has been re- yellow with little darkened tarsi, especially on ante- corded as a pest of cereals in Finland (Kallio, 1950) rior legs. Wings transparent with venation typical and in Sweden (Johansson, 1960). Rygg (1966a) for the genus. Halteres yellow. All setae and setulae reared it from barley shoots in Norway. As host of head and thorax black. Male genitalia: surstyli plants were listed grasses from the genera Cala- rather long and little curved in lateral view; dis- magrostis, Agropyron, Bromus, Festuca, Glyceria, tally dentate on inner side but not curved towards Hordeum (Nartshuk, 1972a; Tschirnhaus, 1981). middle. Hypandrium elongated, pregonites long, Larvae described by Kreiter (1928), Tzygankov postgonites with two setae. Body length 2.5 mm. (1929, 1930), Balachowsky & Mesnil (1935), Oet- Female unknown. tingen (1935), Nye (1958), Beschovski & Krusteva Etymology. – The species is named in honor of R. (1997). The species has been misidentified as L. Danielsson who found the only specimen known cinctipes in nearly all applied publications and de- so far. scriptions of larvae.

172. Lasiosina herpini 173. Lasiosina intermedia (Guérin-Méneville, 1843) Dely-Draskovits, 1977 Figs 764–771, 982, 983 Figs 772, 773 Chlorops herpini Guérin-Méneville, 1843: 30. Syn- Lasiosina intermedia Dely-Draskovits, 1977: 269. types in MNHN. Holotype and paratypes in HNHM. Lasiosina cinctipes auct., nec Meigen, 1830 Head broader than deep or long. Frons wider than (Nartshuk, 1997a). long, yellow with black setae and setulae. Ocel- Headbroaderthandeeporlong.Fronswiderthan lar triangle extending half length of frons followed long, yellow with black setae and setulae. Ocel- by linear extension to front margin of frons, yel- lar triangle extending half length of frons followed low with two black spots, one on ocellar tubercle by linear extension to front margin of frons, yel- and another rhomboid shining spot on apex. Genae low with two black spots, one on ocellar tuber- nearly as wide as first flagellomere. First flagellom- cle and another rhomboid shining spot on apex. ere rounded, extensively darkened in both sexes,

198 Figs 764–777. Lasiosina herpini (Guérin-Méneville): 764: head, dorsal view. 765: head, lateral view. 766: wing. 767: hypopygium, dorsal view. 768: hypopygium, ventral view. 769: hypopygium, lateral view. 770: hypandrium and phallic complex. 771: postgonite and dis- tiphallus. L. intermedia Dely-Draskovits: 772: hypopygium, dorsal view. 773: hypandrium and phallic complex. L. parvipennis Duda: 774: hypopygium, lateral view. L. subnigripes Dely-Draskovits: 775: hypopygium, dorsal view. 776: hypopygium, lateral view. 777: hypandrium and phallic complex. (772, 773 after Ismay, 1991.)

but in male sometimes yellow at base. Palpi in male Biology. – Adults in May, and again in July and Au- from yellow to brown, in female black. Scutum gust in meadows. with 3 black densely grey-dusted stripes, scutel- lum yellow. Pleura with several black marks. Spot on katepisternum black below and yellow above, 174. Lasiosina parvipennis mainly shining, dusted only on lower angle. Ab- Duda, 1933 domen covered with black setulae. Legs extensively Fig. 774 darkened. Body length 2.5–3.0 mm. Lasiosina parvipennis Duda, 1933: 140. Lectotype in NHMH (Nartshuk, 1999). Distribution. – Found only in southern Sweden: SK, HA, ÖL, GO. Frons longer than wide, yellow, with black se- European species. tae and setulae. Ocellar triangle brown dull, main

199 part extending half length of frons, apical part produced acutely. Proboscis somewhat prolonged. linear. Face and genae yellow, genae as wide as First flagellomere rounded. Scutum rather con- first flagellomere. First flagellomere, arista and vex, smooth, dusted. Scutellum rounded. Wings of palpi black. Scutum yellow with black nearly fused Chlorops-type. Tibial organ absent. Female cerci densely dusted stripes. Scutellum yellow or dark- very short. Male genitalia: epandrium flattened ened. Wings variable in length: shorter or equal of posteriorly with a strong marginal carina. Surstyli abdomen. Legs with femora black, tibiae and tarsi apically black and densely sclerotized. Middle lobe yellow. Halteres yellow. Body length 1.5 mm. of surstyli rectangular. Postgonites very long, pre- gonites indistinct. Distribution. – Rare species. Found only in Finland: Ab,St,Ta,Sa,Kb,OmandRussia:Vib,Kr. Distribution. – Palearctic Region and New Zealand. Eurasian species, known from North Europe to East Siberia and Mongolia. 176. Melanum laterale Biology. – Adults in May, and again in September (Haliday, 1833) on bogs and marshes. Probably two generations per Figs 17, 778–782 year. Chlorops lateralis Haliday, 1833: 172. Lectotype in 175. Lasiosina subnigripes NMID (Ismay et al., 2001). Dely-Draskovits, 1977 Frons yellow, black pubescent. Ocellar triangle Figs 775–777 black, shining, dusted on ocellar tubercle. Ocel- lar triangle extending nearly to front margin of Lasiosina subnigripes Dely-Draskovits, 1977: 274. frons with acute apex. Face and genae yellow, genae Holotype and paratype in HNHM. as wide as first flagellomere, with strongly pro- jected and acute vibrissal angle. Antennae black, Head broader than deep or long. Frons as wide first flagellomere as long as wide. Palpi yellow. Scu- as long, yellow with black setae and setulae. Ocel- tum black, dusted covered with short black setulae. lar triangle extending half length of frons followed Scutellum yellow. Pleura yellow with usual black by linear extension to front margin of frons, yel- marks. Spot on katepisternum dusted. Abdomen low with two black spots, one on ocellar tuber- dorsally black with yellow lateral margins. Legs cle and another rhomboid shining spot on apex. mainly yellow. Body length 2.5–3.0 mm. Genae narrower than first flagellomere. In male first flagellomere rounded, extensively darkened, Distribution. – Denmark: from all districts. Sweden: palpi yellow. (Female unknown). Scutum with 3 along Baltic coast north to NB. Norway: AK, On, VA, black densely grey-dusted stripes, central one nar- Ry,HOi,STy,STi.Finland:northtoObS.Russia:Vib; rowed behind. Scutellum yellow. Pleura with com- Kr; Lr: Ponoj, 1 female. mon black marks. Spot on katepisternum black, Trans-Palearctic species, known from the British shining. npl 1 + 2. Abdomen covered with black se- Isles to Japan. tulae. Legs yellow. Body length 2.5 mm. Biology. – Adults from June to September in wet- Distribution. – Rare species. Found only in Sweden: lands, especially on coastal salt marshes, but also UP:Hässelby,Böje,20.v.1969,20.v.1970,08.v.1972 on shores of lakes and banks of rivers inland. (Kur), 5 specimens. The larvae probably phytophagous. Tschirnhaus European species. (1972) recorded Juncus gerardi as host plant.

Genus Melanum Becker, 1910 Genus Meromyza Meigen, 1830

Melanum Becker, 1910: 50. Type species: Chlorops Meromyza Meigen, 1830: 163. Type species: Musca lateralis Haliday, 1833 by original designation. saltatrix Linnaeus, 1761 by designation of Mac- quart, 1835: 589. Head as long as high, yellow with black. Ocellar tri- angle large, behind almost as broad as frons, with Body light green or yellowish green with dusted acute tip, smooth shining, black. Vibrissal angle black or red stripes and maculae. Head as long

200 Figs 778–782. Melanum laterale (Haliday): 778: head, dorsal view. 779: head, lateral view. 780: hypopygium, dorsal view. 781: end of male abdomen, lateral view. 782: hypandrium and phallic complex. as high or longer, with short setae. Ommatidia in some species on the borders of the Oriental and anterior part of eye larger than in posterior. Ocel- Neotropical Regions. lar triangle extending 1/2 or 2/3 length of frons, pale with black ocellar tubercle, smooth or with transverse ridges, especially in apical part. Face Key to species of Meromyza Meigen concave with indistinct carina. Scutum moderately (males only) convex, usually smooth and dusted, with black or red stripes. Scutellum rounded. Wings with veins 1 Palpi pale, at most only the tip a little R2+3 and R4+5 strongly procurved. Veins R4+5 and darkened...... 2 M distinctly divergent. Legs stout, hind femora – Palpi mainly black, at least on apical half strongly thickened, ventrally with two rows of black ...... 8 spines. Hind tibiae correspondingly curved. Tibial 2 (1) Abdomen with a central dark stripe and organ absent. Male genitalia: postgonites usually large, strongly sclerotized and black, rarely small two dark lateral spots at the base, no lat- and nearly transparent. Structure of male geni- eral dark marking on tergites 3–5. Surstyli talia, especially shape and colour of postgonites rather long and expanded medially with and surstyli are the only reliable characters to dis- long hairs (Figs 808, 829, 846) ...... 3 tinguish species of the genus. – Abdomen with three dark stripes or Larvae phytophagous, develop in shoots of Poa- nearly black dorsally. Surstyli different ceae, including cereals...... 5 3 (2) Aedeagus swollen at base, basally nearly Distribution. – Predominantly Holarctic, but one twice as broad as at middle. Lower mar- species is known in the Afrotropcal Region and gin of anterior process of postgonite

201 strongly convex medially, the tip of ante- 9 (8) Frontal triangle 1.5 times longer than rior process turned a little laterally. Frons broad at base with concave side margins, distinctly produced, hypopleural marks and wrinkled surface. Postgonite more yellow to reddish. Genitalia Figs 846–850 massive, posterior process long. Genitalia ...... M. pratorum Meigen Figs 798–801. Body larger, 3.5–4.0 mm – Aedeagus not swollen at base, basally not ...... M. elbergi Fedoseeva broader than at middle. Lower margin – Frontal triangle nearly equilateral, with of anterior process of postgonite more straight side margins. Postgonite small. straight and not turned laterally. Frons Genitalia Figs 871–875. Body smaller 2.5– less produced and hypopleural mark 3.0mm...... M. triangulina Fedoseeva black...... 4 10 (8) Anterior process of postgonite elongated 4 (3) Lower margin of anterior process of go- andpointedatthetip...... 11 nite a little convex and the tip less – Anterior process of postgonite broad at pointed. Genitalia Figs 808–811 the tip, sometimes depressed and turned ...... M. ingrica Nartshuk laterally, postgonite more massive . . . . 16 – Lower margin of anterior process of post- 11 (10) Posterior process of postgonite long and gonite more straight and the tip pointed. directed posteriorly, basal part of aedea- Genitalia Figs 829–833 gus swollen and nearly square. Central ...... M. ornata (Wiedemann) mesonotal stripe usually ending before 5 (2) Hind femora more swollen, 4 times as scutellum. Genitalia Figs 793–797 wide as hind tibia. Genitalia Figs 852– ...... M. curvinervis (Zetterstedt) 855...... M. rohdendorfiji Fedoseeva – Posterior process of postgonite shorter – Hind femora less swollen and only 2.5– and directed downwards or a little ante- 3 times wider than hind tibia. Genitalia riorly, sometimes hardly separated from different...... 6 anterior process. Aedeagus not swollen 6 (5) Central scutal stripe does not extend to basally...... 12 thescutellum...... 7 12 (11) Postgonite scarcely darkened, nearly – Central scutal stripe extending over the transparent or a little brown, posterior scutellum. Surstyli short and straight. process small, sometimes hardly sepa- Genitalia Figs 812–815 rated from anterior process. Hind femora ...... M. mosquensis Fedoseeva more swollen, nearly 3 times wider than 7 (6) Surstyli long and expanded at tip later- hindtibia...... 13 ally. Postgonites more massive with small – Postgonite more strongly sclerotized dark tooth at tip. Upper margin of anterior brown or almost black, posterior process process of postgonites slightly convex. well developed. Hind femora usually less Genitalia Figs 882–885 swollen, not more than 2–2.5 times wider ...... M. zimzerla Nartshuk thanhindtibia...... 15 – Surstyli short, not expanded laterally. 13 (12) Surstyli rather narrow and has a group of Postgonites with acute tip. Upper mar- black thickened bristles at the base. Gen- gin of anterior process of postgonites italia Figs 816–821 strongly convex. Genitalia Figs 789–792 ...... M. nigriseta Fedoseeva ...... M. bohemica Fedoseeva – Surstyli moderately broad, sometimes 8 (1) Frontal triangle with its boundary lin- long but without black thickened bristles early darkened. Genae with black hairs ...... 14 ...... 9 14 (13) Surstyli longer than epandrium. Ante- – Frontal triangle not darkened laterally. rior process of postgonite more elongated Genaewithoutblackhairs...... 10 and directed downwards. Central scutal

202 stripe ends before scutellum. Genitalia Species key to known 3rd instar larvae Figs 866–870 . . . . . M. sibirica Fedoseeva of Meromyza Meigen – Surstylus shorter than epandrium. Ante- rior process of postgonite shorter and di- rected anteriorly. Central scutal stripe ex- 1 Distal segment of antennae brownish and re- tending to scutellum. Black spot on ocel- tractedwithinbasalsegment...... 2 lar triangle elongated anteriorly. Geni- – Antennaenotcoloured...... 3 talia Figs 840–844 . . . M. pluriseta Péterfi 2 Anterior spiracles of transversal rosette type 15(12) Surstyli rather short and broad; postgo- with 10–12 buds on two lobes (Fig. 1005). Spic- nite small. Genitalia Figs 835–839 ular zones on abdominal segments consist of ...... M. palposa Fedoseeva numerous lines on anterior half of segment – Surstyli narrower, expanded medially; and a few lines on hind margin segments (Fig. postgonite longer. Genitalia Figs 822–826 1006). In addition wings of dentate sclerite ...... M. nigriventris Macquart point laterally instead of anterolaterally. Lar- 16 (10) Postgonite massive, posterior process of vae light green, in shoots of Calamagrostis postgonite divided into two parts, ante- epigeios and Ammophila arenaria rior process at tip straight, not pointed ...... M. pratorum Meigen or depressed. Ocellar triangle short (Fig. – Anterior spiracles of small rosette type with 783). Genitalia Figs 785–788 7–9 buds (Fig. 1007). Spicular zones on ab- ...... M. bifurcata sp. nov. dominal segments consist of 20–30 lines on – Posterior process of postgonite not di- anterior half of segment in a broadly trian- vided, tip of anterior process more or gular field plus a few lines on hind margin less pointed or depressed. Ocellar trian- (Fig. 1009). Wings of dentate sclerite point an- glelonger...... 17 terolaterally. Posterior spiracles not borne on 17 (16) Scutal stripes mainly or partly red. Tip lobes and close together. Larvae blue-green, in of anterior process of postgonites not shoots of Dactylis glomerata depressed...... 18 ...... M. variegata Meigen – Scutal stripes black. Surstyli expanded 3 Two pairs of additional teeth on inner side of medially. Postgonite massive with tip of mandible...... 4 anterior process depressed. Aedeagus – Three pairs of additional teeth on inner side of broadened basally. Genitalia Figs 859– mandible(Fig.1011)...... 6 865...... M. saltatrix (Linnaeus) 4 Anteriorspiracleswith7–8buds...... 5 18 (17) Hind femora less swollen, 2.5–3.0 times – Anterior spiracles with 9–10 buds. Spicular thicker than tibia. Surstyli long, expanded zones of abdominal segments triangular, oc- a little laterally. Postgonite narrower. cupy only anterior part of segment, on poste- Aedeagus with bifid process basally. Gen- rior part only 2–3 short lines present. Inter- italia Figs 876–881 spiracular processes branched 2 or 3 times...... M. variegata Meigen Larvae greenish yellow, 6–7 mm long, in – Hind femora strongly swollen, 4 times shoots of Poa pratensis, Festuca rubra, Agrostis thicker than tibia. Surstyli not expanded vulgaris ...... M. mosquensis Fedoseeva laterally. Postgonite massive with con- 5 Interspiracular processes on hind spiracles cave tip and small tooth-like projection. not branched. Spicular zones semicircular, Base of aedeagus without bifid process. occupy only anterior part of abdominal seg- Genitalia Figs 802–807 ments, on posterior part of segment only 2–3 ...... M. femorata Macquart short lines present. Larvae greenish yellow, 5 mm long, in shoots of Dactylisglomerata and Festuca rubra ...... M. triangulina Fedoseeva

203 – Interspiracular processes on hind spiracles long axis of eye. Frons produced beyond eye and forked at tips. Larvae greenish 6–7 mm long, parafacial plates equal half width of genae. Genae in shoots of Lolium perenne as wide as first flagellomere is long, with yellow se- ...... M. bohemica Fedoseeva tulae. Frons longer than wide, covered with black setulae. Only vte distinct among head setae. Ocel- 6 Anterior spiracles with 11–12 buds. Parastomal lar triangle short, extending little more than half bar with lanceolar widening. Larvae bright length of frons, yellow shining, with black, dusted green, 7–10 mm long, in shoots of Deschamp- ocellar tubercle and small brown points along side sia caespitosa ...... M. ornata (Wiedemann) margins. Antennae yellow, first flagellomere little – Anterior spiracles with 7–10 (11) buds...... 7 longer than deep, slightly darkened above. Arista 7 Spicular zones on ventral side of all abdomi- brown, nearly bare. Occiput with brown stripe and two brownish spots laterally. Palpi black in apical nal segments occupy only anterior part of seg- part. ments, no additional spicular lines at poste- Scutum longer than wide with three wide black, rior margin, but some striation present on dor- dusted stripes, central one extending over scu- sal side. Larvae green, 7–8 mm long, in shoots tellum. Pleura with black, shining spot on anepi- wheat, rye, barley, Elytrigia repens, Phleum meron, black, dusted spot on katepimeron and red- pratense ...... M. nigriventris Macquart dish with black upper angles on katepisternum. – Short additional spicular lines present on pos- Legs mainly yellow, front femora with black stripe. terior part of segments besides main area on Hind femora 2.5 times thicker than tibiae. Wings transparent with venation typical for the genus. anterior part of all segments. Spicular zones Abdomen with three black spots on tergites. Male short, not occupying half of segments . . . . . 8 genitalia: surstyli distinctly angulated on inner 8 Larvae greenish-yellow, anterior spiracles with side. Postgonites black, hind process large triangu- 8–9 buds. Dentate sclerite long, broadly flat- lar, front process bifidly divided, one twig longer tened. Larvae 6–7 mm long, in shoots of Poa than the other. Aedeagus with enlarged base. Body pratensis, Festuca rubra length 3.0 mm...... M. saltatrix (Linnaeus) 178. Meromyza bohemica – Larvae bright green, anterior spiracles with 9– 10 (11) buds. Dentate sclerite short, sharply Fedoseeva, 1962 curved. Larvae 7–8 mm long, in shoots of Elyt- Figs 789–792 rigia repens, Roegneria ramosum, Hierochloe Meromyza bohemica Fedoseeva, 1962: 472. Holo- odorata ...... M. nigriseta Fedoseeva type in ZISP. Ground colour yellow. Ocellar triangle light, shin- 177. Meromyza bifurcata ing, with slightly darkened side margins. First flag- ellomere longer than wide, yellow darkened above. Nartshuk & Andersson, sp. nov. Palpi narrow, yellow, rarely slightly darkened at tip. Figs 783–788 Genae with some black setulae in a row. Stripes of scutum reddish centrally, dusted, outer mar- Holotype male, Sweden: SK: Kullaberg, 02.vii.1974 gin of inner lateral stripes black. Central stripe (H. Andersson). Holotype in ZMLU. not extending over scutellum. Pleura with reddish Diagnosis. – New species belongs to the saltatrix marks, except oval black mark on anepisternum. group of species with black palpi, black stripes on Abdomen with central brown to black stripe and scutum and scutellum, three black stripes on ab- spots on 2–5 tergites. Hind femora 2.5–3.0 times domen, moderately thickened hind femora and wider than tibiae. Body length 2.5–3.0 mm. strongly sclerotized massive postgonites. It differs Distribution. – Found only in Denmark: EJ, LFM, from all known species by bifidly divided front pro- NWZ and southern Sweden: SK, HA, SM, ÖL, GO. cess of postgonites and also rather short ocellar European species. triangle. Biology. – Adults in July to September in meadows. Description. – Ground colour of body light yel- Larvae develop in shoots of Lolium perenne (Fe- low. Head longer than deep with almost horizontal doseeva, 1974).

204 Figs 783–797. Meromyza bifurcata Nartshuk & Andersson sp. n.: 783: head, dorsal view. 784: head, lateral view. 785: hypopygium, dorsal view. 786: postgonite. 787: postgonite, anterior view. 788: basiphallus. M. bohemica Fedoseeva: 789: hypopygium, dorsal view. 790: hy- popygium, lateral view. 791: male genitalia, ventral view. 792: postgonite. M. curvinervis (Zetterstedt): 793: hypopygium, dorsal view. 794: hypopygium, lateral view. 795: postgonite. 796: basiphallus. 797: male genitalia, latero-ventral view.

Eggs and larvae described by Hubicka (1967, as widely black on apical half. Stripes of scutum black, M. lolii Hubicka). densely grey-dusted. Central stripe ends before scutellum, but sometimes before scutellum a black 179. Meromyza curvinervis mark with indistinct borders. Sometimes central (Zetterstedt, 1848) stripe partly reddish and scutellum with black Figs 793–797 mark. Spot on katepisternum black and mainly Oscinis curvinervis Zetterstedt, 1848: 2620. Holo- dull. Abdomen with only central reddish brown to type in ZMLU (Andersson, 1966). black stripe. Hind femora 2.0–2.5 times wider than Ground colour yellow. Frons longer than wide. tibiae. Phallus swollen at base. Body length 2.0– Ocellar triangle extending 2/3 length of frons. Palpi 3.0 mm.

205 Distribution. – Scarce, only in southern Fennoscan- femora strongly swollen, nearly 4 times as thick as dia. Sweden: SK, ÖL, GO. Finland: Ab, Ka. Russia: tibiae. Body length 3.5–4.0 mm. Vib. Not so far found in Norway and Denmark. Distribution. – Southern species. Denmark: from all Eurasian species known from the British Isles to districts except NWJ and SZ. Sweden: SK, BL, SM, Mongolia. ÖL. Norway: AK, Ø. Biology. – Adults in July and August in meadows. European species. Tschirnhaus (1981) proposed as host plants Am- Biology. – Adults from the end of June to Septem- mophila arenaria and Corynephorus canescens. ber in meadows. Larvae phytophagous, develop in shoots of 180. Meromyza elbergi Dactylis glomerata. Fedoseeva, 1979 Figs 798–801 182. Meromyza ingrica Meromyza elbergi Fedoseeva,1979:118.Holotypein Nartshuk, 1992 ZISP. Figs 808–811 Ground colour fuscous yellow. Ocellar triangle Meromyza ingrica Nartshuk, 1992: 125. Holotype wrinkled, shining, with distinctly concave, slightly and 1 paratype in ZISP, other paratypes in darkened side margins. First flagellomere longer NHMH. than wide, yellow with darkening above. Palpi nar- Ground colour greenish. Body elongated, tip of ab- row, black in distal half. Genae 1.5 times wider than domen reaching well behind tip of hind femora. wide of first flagellomere, with some black setulae Head longer than deep. Frons produced in front as in irregular rows. Stripes of scutum black, or cen- long as breadth of first flagellomere. Ocellar trian- tral stripe reddish, dusted. Central stripe extending gle longer than wide at base, with wrinkled surface over scutellum. Pleura with reddish spots, except and black on ocellar tubercle only. Palpi greenish. black marks on anepisternum and katepimeron. Stripes of scutum black, densely grey-dusted. Cen- Abdomen with three brown to black stripes. Hind tral stripe reaching scutellum and scutellum with femora about 3 times as thick as tibiae. Body length black mark. Katepimeron with black or dark brown 3.5–4.0 mm. mark. Abdomen with only a central black stripe. Distribution. – Sweden: GO, NÄ, UP. Norway: AK. Hind femora moderately swollen, about 2.5 times Finland: Al. Not found in Denmark or Fennoscan- as thick as tibiae. Hypandrium covered with long dian districts of Russia. hairs. Body length to 4.0 mm. Central and eastern European species. Distribution. – Sweden: HS, JÄ, NB, LY. Norway: Biology. – Adults in July and August in dry mead- HOy, HOi. Finland: north to Ob. Russia: Vib. Not ows. found in Denmark. Known till now only from the Fennoscandian districts listed above. Possibly more widely dis- 181. Meromyza femorata tributed, but not distinguished from other species Macquart, 1835 of the M. pratorum species group. Figs 802–807 Biology. – Adults in July to August in wet meadows. Meromyza femorata Macquart, 1835: 589. Types probably in MHNL. 183. Meromyza mosquensis Ground colour yellowish. Frons produced in front Fedoseeva, 1960 no more than 1/2 breadth of first flagellomere. Figs 812–815 Ocellar triangle shining, rugose in apical part and Meromyza mosquensis Fedoseeva, 1960: 457. Syn- black on ocellar tubercle only. Palpi black on api- types in ZISP. cal half. Stripes of scutum red, grey-dusted, central stripe reaching scutellum and scutellum with red Ground colour greenish-yellow. Frons produced in mark. Pleura with red marks, except small black front no more than 1/2 breadth of first flagellom- mark on the anepisternum. Abdomen with central ere. Face more upright than in most Meromyza black stripe and small spots on 3–5 tergites. Hind species. Ocellar triangle with wrinkled rhomboid

206 Figs 798–811. Meromyza elbergi Fedoseeva: 798: hypopygium, dorsal view. 799: hypopygium, lateral view. 800: postgonite. 801: basiphal- lus. M. femorata Macquart: 802: hypopygium, dorsal view. 803: hypopygium, lateral view. 804: postgonite. 805: male genitalia, ventral view. 806: basiphallus, ventral view. 807: basiphallus, lateral view. M. ingrica Nartshuk: 808: hypopygium, dorsal view. 809: hypopygium, lateral view. 810: postgonite. 811: basiphallus.

207 dark coloured apical part and black ocellar tuber- Host plants: Elytrigia repens, E. tenerum, Ley- cle. Palpi yellow. Stripes on scutum rather wide mus ramosus, Hierochloe odorata, Agrostis vulgaris, and sometimes touching in front, black densely rarely larvae found in shoots of summer wheat and grey-dusted or central one reddish brown. Central winter rye (Karpova, 1965, 1972; Nartshuk, 1972a; stripe reaching scutellum and scutellum with dark Fedoseeva, 2003). brown mark. Pleura with brown to black marks. Ab- Larvae of third instar described by Fedoseeva domen short, dorsally shining, with 3 black stripes, (1966). in dark specimens entirely black dorsally. Hind femora moderately swollen, 2.5 times as thick as 185. Meromyza nigriventris tibiae. Body length 3.0–3.5 mm. Macquart, 1835 Distribution. – Denmark: SJ, WJ, F, LFM, NEZ, B. Figs 822–826, 1010–1012 Sweden: SK, BL, HA, SM ÖL, GO. Norway: Ø, AK. Meromyza nigriventris Macquart, 1835: 590. Type Finland: Al, Ab, N, Ta, Sa. Russia: Vib, Kr, Lr. probably in MHNL. European species, but not in the southernmost Meromyza basalis Roser, 1840: 63, Type 1 male in parts. SMNS (Ismay, 1981a; Nartshuk, 1994). Biology. – Adults in June to August in meadows. Meromyza cerealium Reuter, 1902a: 84. Type not Host plants: Poa pratensis, Agrostis vulgaris, Fes- found in NHMH (Nartshuk, 1992). Mesnil (1931) tuca rubra, F. pratensis, Elytrigia repens (Nartshuk, considered M. cerealium as a summer form of M. 1962a; Fedoseeva, 1966, 2003; Karpova, 1972). Hi- nigriventris (as M. saltatrix). bernates as larvae of third instar. Meromyza rostrata Hubicka, 1966: 16. Holotype in Larvae of all three instars are described by Fe- ZIWP. doseeva (1966). The species is very variable in colour, showing two distinct colour forms: 184. Meromyza nigriseta Summer form. Ground colour yellow. Frons pro- Fedoseeva, 1960 duced in front no more than 1/2 breadth of first Figs 30, 816–821 flagellomere. Ocellar triangle no longer than wide at base, yellow, wrinkled, black spot on ocellar tu- Meromyza nigriseta Fedoseeva,1960:456.Syntype bercle small. Palpi black in apical part and a lit- in ZISP. tle swollen. Stripes of scutum entirely reddish or Ground colour greenish-yellow. Frons produced in central stripe behind and outer marginal stripes front no more than 1/2 breadth of first flagellom- brown to black. Central stripe reaches scutellum ere. Ocellar triangle slightly wrinkled in apical part and scutellum with red or brown mark. Pleura with and with rather large black spot on ocellar tubercle. brownish marks, except small black spot on anepis- Palpi black slightly thick in apical part. Stripes of ternum. Abdomen with 3 reddish to brown stripes. scutum rather narrow, black densely grey-dusted or Hind femora moderately swollen, no more than 3 central one reddish brown. Central stripe reaches times as thick as tibiae. scutellum and scutellum with black mark. Ab- Spring form dark coloured. Ocellar triangle en- domen with 3 black stripes or only one central tirely black, or with large square black spot at base stripe and with spots on tergites 3–5. Hind femora and occiput black. Scutal stripes wide and black moderately swollen, no more than 3 times as thick and sometimes fused, leaving scutum black with as tibiae. Epandrium covered with black setae and small marginal yellow marks. Abdomen black dor- surstyli with thickened black setae in several rows sally, only hind margins of tergites narrowly yellow. on basal part. Body length 3.5–4.0 mm. Legs partly black. Body length 3.0–3.5 mm. Distribution. – Denmark:NEJ,F,NEZ,B.Sweden: northtoDR.Norway:AK.Finland:Ab,N,Sa.Russia: Distribution. – Denmark: SJ, WJ, NWJ, NEJ, LFM, Vib. NWZ, NEZ. Sweden: along Baltic coast north to NB. Eurasian species, known from the British Isles to Norway:AK,VE.Finland:northtoTb.Russia:Vib, Mongolia. Kr. Holarctic species, in the Palearctic widely dis- Biology. – Adults from the end of June to beginning tributed from the British Isles to Japan, in the of August, in meadows. Nearctic confined to western North America.

208 Figs 812–826. Meromyza mosquensis Fedoseeva: 812: hypopygium, dorsal view. 813: hypopygium, lateral view. 814: postgonite. 815: basiphallus. M. nigriseta Fedoseeva: 816: hypopygium, dorsal view. 817: hypopygium, lateral view. 818: postgonite. 819: male genitalia, ventral view. 820: basiphallus. 821: base of basiphallus. M. nigriventris Macquart: 822: hypopygium, dorsal view. 823: hypopygium, lateral view. 824: postgonite. 825: male genitalia, ventral view. 826: base of basiphallus.

209 Biology. – Adults from May to June and August to Eurasian species, ranging from the British Isles to September, two generations per year. Yakutia, East Siberia and Mongolia. Larvae develop in different cereals: wheat, bar- ley, rye and many wild grasses: Aegelops sp., Elytri- Biology. – Adults from end of June to August in wet gia repens, E. tenerum, Agropyron cristatum, Lolium meadows. perenne, Arrhanatherum elatius, Phleum pratense, Larvae live in shoots of Deschampsia caespitosa. Hierochloe odorata, Calamagrostis epigeios (Kar- Larvae of the first instar hibernate and in spring move into another plant to continue development. pova, 1972; Fedoseeva, 2003). Hibernates as larvae Larvae of all three instars are described by Fe- of the third instar within shoots of grasses. Lar- doseeva (1966, as M. sororcula). vae damage shoots and ears of cereals. This species (not M. saltatrix) is a pest of cereals (Fedoseeva, 1969; Fedoseeva & Perfiljeva, 1988; Krusteva, 1995). 187. Meromyza palposa Reared from cereals (wheat, barley) in Sweden (Jo- Fedoseeva, 1960 hanssen, 1960), Finland (Reuter, 1902a, as M. cere- Figs 835–839 alium) and Norway (Rygg, 1966a, as M. saltatrix var. Meromyza palposa Fedoseeva, 1960: 452. Syntype variegata and M. pratorum var. decora). Some spec- in ZISP. imens were reared from compost (Werner, 1997). Without doubt larvae occurred there with grass re- Ground colour yellow. Frons slightly produced in mains. front. Ocellar triangle as long as wide at base or Larvae were described by Kreiter (1929), Tzy- little longer, with indistinct apical part, with large gankov (1930), Balachowsky & Mesnil (1935), Nishi- black spot on ocellar tubercle. First flagellomere jima (1960), usually under name M. saltatrix.Full as wide as long. Palpi black, rather broad. Genae descriptions of all three larval instars are given by with pale setulae. Stripes of scutum black, densely Fedoseeva (1966). The species is named as M. salta- grey-dusted or central stripe brownish in the mid- trix in most applied publications (misidentifica- dle. Central stripe reaches scutellum and scutel- tion). lum with black or brown mark. Abdomen with 3 wide black stripes. Hind femora slightly swollen, 186. Meromyza ornata no more than in 2 times as thick as tibiae. Surstyli broad, densely covered with short setulae, gonites (Wiedemann, 1817) rather small. Body length to 2.5–3.5 mm. Figs 827–834 Distribution. – Denmark: WJ, NWJ, NEJ, B. Sweden: Oscinis ornata Wiedemann, 1817: 86. Holotype in north to NB along Baltic coast. Norway: present, NHMW (Nartshuk, 1997b). but district uncertain. Finland: north to Obs along Meromyza sororcula Fedoseeva,1962:470.Holo- Baltic coast. Russia: Vib. type in ZISP. European species. Ground colour greenish. Body elongated, head Biology. – Adults in June to August in dry meadows. longer than deep. Frons produced in front nearly as Larvae develop in shoots of Nardus stricta (Fe- long as breadth of first flagellomere. Ocellar trian- doseeva, 2003). gle longer than wide at base, with wrinkled surface and black on ocellar tubercle only. Occiput black. 188. Meromyza pluriseta Head setae and setulae predominantly black. Palpi greenish. Stripes of scutum wide, black, densely Péterfi, 1961 grey-dusted. Central stripe continuing over scutel- Figs 840–844 lum. Katepisternum with pale spot, katepimeron Meromyza pluriseta Péterfi, 1961: 167. Type deposi- with blackish mark. Abdomen with only central tory unknown. black stripe. Hind femora moderately swollen, 2.5 times as thick as tibiae. Epandrium covered with Ground colour fuscous yellow. Frons produced in long pale setulae. Body length to 4.0–4.5 mm. front as long as 1/2 breadth of first flagellomere. Ocellar triangle as long as wide at base or a little Distribution. – Denmark: widespread. Sweden: longer, with dark wrinkled rhomboid spot in api- north to LU. Norway: AK: Enebakk, Oslo; VE: Tøns- cal part, which connects with extended large black berg. Finland: north to ObN. Russia: Vib, Kr. spot of ocellar tubercle in dark specimens. In light

210 Figs 827–839. Meromyza ornata (Wiedemann): 827: head, dorsal view. 828: head, lateral view. 829: hypopygium, dorsal view. 830: hy- popygium, lateral view. 831: postgonite. 832: postgonite, another specimen. 833: male genitalia, ventral view. 834: basiphallus. M. palposa Fedoseeva: 835: hypopygium, dorsal view. 836: hypopygium, lateral view. 837: postgonite. 838: male genitalia, ventral view. 839: basiphal- lus.

211 Figs 840–851. Meromyza pluriseta Fedoseeva: 840: hypopygium, dorsal view. 841: hypopygium, lateral view. 842: postgonite. 843: postgo- nite, another specimen. 844: basiphallus. M. pratorum Meigen: 845: head, dorsal view. 846: hypopygium, dorsal view. 847: hypopygium, lateral view. 848: postgonite. 849: postgonite, another view. 850: male genitalia, ventral view. 851: basiphallus.

212 coloured specimens black spot on ocellar tubercle Biology. – Adults in June to August in dry meadows small. Palpi black except yellow at base. Stripes of and sandy, coastal habitats. scutum wide black, densely grey-dusted or lateral Larvae develop in shoots of Calamagrostis epi- stripes brownish in front. Central stripe reaching geios, C. neglecta, Ammophila arenaria and Hie- scutellum and scutellum with black or brown mark. rochloe odorata (Nartshuk, 1962a; Karpova, 1972; Pleura with large black marks. Abdomen with 3 Huiskes, 1979). One generation per year. Females wide black stripes, sometimes partly fused. Hind lay eggs on leaves. Larvae damage upper meristem femora 3 times as thick as tibiae. Front femora and central leaves. darkened. Male genitalia: surstyli broad, densely Larvae described by Balachowsky & Mesnil covered with short setulae. Gonites small and light. (1935). Body length 3.5–4.0 mm. 190. Meromyza rohdendorfiji Distribution. – Denmark: NWJ, NWZ, NEZ, B. Swe- den: SK, ÖL. Norway: On: Dovre. Finland: north to Fedoseeva, 1974 ObS. Russia: Vib, Kr, Lr. Figs 852–856 Eurasian species ranging from the British Isles to Meromyza rohdendorfiji Fedoseeva, 1974: 916. Holo- Yakutia, East Siberia and Mongolia. type in ZISP. Biology. – Adults in June to August in meadows. Ground colour brownish yellow. Frons produced in Larvae develop in shoots of Hierochloe odorata, front no more than 1/2 breadth of first flagellomere. Elytrigia repens (Karpova, 1972; Fedoseeva, 2003) Ocellar triangle longer than wide at base, shining, and probably other grasses. wrinkled in apical part and black on ocellar tuber- cle only.Palpi yellow, slightly widened at tip. Stripes of scutum densely grey-dusted, central one reddish 189. Meromyza pratorum brown, inner and outer lateral stripes black. Cen- Meigen, 1830 tral stripe reaching scutellum and scutellum with Figs 5, 845–851, 1005, 1006 black or brown mark. Pleura with red marks, ex- ceptblackmarkonanepisternum.Abdomenwith Meromyza pratorum Meigen, 1830: 165; Meigen’s central black stripe and small spots on tergites 2–5. plate 162: 11 (Morge, 1976a). Lectotype female Hind femora strongly swollen, nearly 4 times as and paralectotype female in MNHN (Nartshuk, thick as tibiae. Body length 3.5–4.0 mm. 2006). Distribution. – Denmark: widespread. Sweden: north to UP. Norway: Ø: Larkollen (Siebke); VAy: Ground colour greenish. Body elongated, tip of ab- Lindesnes, Spangereid. Finland: Ab: Dragsfjärd, domen reaching well behind tip of hind femora. Nagu. Head longer than deep. Frons produced in front as European species. long as breadth of first flagellomere. Ocellar trian- gle longer than wide at base, with wrinkled surface Biology. – Adults in July to August, in meadows. and black on ocellar tubercle only. Palpi greenish. Stripes of scutum black, densely grey-dusted. Cen- 191. Meromyza saltatrix tral stripe reaching scutellum and scutellum with (Linnaeus, 1761) black mark. Katepimeron with yellow mark. Ab- Figs 79, 857–865, 1013, 1014 domen with only central black stripe. Hind femora moderately swollen, no more than 2.5 times as Musca saltatrix Linnaeus, 1761: 555. The only speci- thick as tibiae. Hypandrium covered with long men in Coll. Linnaeus, Linnean Society, London, hairs. Phallus swollen at base. Body length 4.0– under Musca saltatrix is a female of Chlorops 6.0 mm. pumilionis (Bjerkander), but Ismay (1981a) found it questionable that this is an original Distribution. – Denmark: widespread. Sweden: specimen, as it disagrees with Linnaeus’ 1761- north to NB. Norway: Ø, AK, VE, Fn. Finland: north description saying “Femora postica crassa”, a to ObN. Russia: Vib, Kr. character omitted in the 1767-edition of his Sys- Holarctic species: widely distributed from the tema Naturae. An application should be made British Isles to Japan, and also in western North to the ICZN for permission to designate of a neo- America. type of Musca saltatrix in order to retain current

213 Figs 852–865. Meromyza rohdendorfiji Fedoseeva: 852: hypopygium, dorsal view. 853: hypopygium, lateral view. 854: male genitalia, ven- tral view. 855: postgonite. 856: basiphallus. M. saltatrix (Linnaeus): 857: head, dorsal view. 858: head, lateral view. 859: hypopygium, dorsal view. 860: hypopygium, lateral view. 861: hypandrium and phallic complex, ventral view. 862: hypandrium and phallic complex, lateral view. 863: postgonite. 864: postgonite, lateral view. 865: basiphallus.

214 usage of the genus Meromyza that has Musca genitalia: postgonites light-coloured, elongated and saltatrix Linnaeus as type species. curved. Body length 3.0–3.5 mm. Ground colour greenish. Frons produced in front by Distribution. – Only Sweden: 1 male without ex- no more than half width of first flagellomere. Ocel- act locality. Finland: Sb: Maaninka (Tuovilankas), 1 lar triangle longer than wide at base, with wrin- male. kled surface and black on ocellar tubercle only. Oc- The species is rather common in Palearctic Asia, ciput darkened. Head setae and setulae black. Palpi but rare in Europe. A few specimens are known blackened in apical half. Stripes of scutum black, from Austria, Germany, Poland, Estonia, Lithuania densely grey-dusted. Central stripe reaching scutel- and European part of Russia. lum and scutellum with black mark. Pleural marks reddish to black, spot on katepisternum dusted. 193. Meromyza triangulina Abdomen with three black stripes. Hind femora 3 times as thick as tibiae. Male genitalia: postgo- Fedoseeva, 1960 Figs 871–875 nites black, large triangular with tip of posterior process depressed. Phallus swollen at base. Body Meromyza triangulina Fedoseeva, 1960: 457. Syn- length 3.5–4.0 mm. type in ZISP. Distribution. – Very common species occurring Ground colour yellowish or light greenish. Frons throughout Denmark and Fennoscandia. produced in front less than 1/2 breadth of first flag- Holarctic: widely distributed trans-Palearctic and ellomere. First flagellomere little longer than deep, western Nearctic species. darkened above. Ocellar triangle as long as wide at base or little longer, with large black spot on Biology. – Adults from June to August, in meadows. ocellar tubercle and darkened side margins. Palpi Larvae phytophagous. As host plants were re- black an apical half, narrow. Genae with black se- corded Poa pratensis, Elytrigia repens and Festuca tulae on lower margin. Stripes of scutum black, rubra (Fedoseeva, 1966, 2003). Hibernates as larvae densely grey-dusted, central stripe narrowed be- of third instar. Larvae of all three instars described hind and reaching scutellum, and scutellum with by Fedoseeva (1966). black mark. Abdomen usually with 3 black stripes, Earlier authors frequently recorded the present rarely only one stripe present. Hind femora rather species as a pest of cereals, but these records are all thin, nearly 2 times as thick as tibiae. Male geni- based on misidentifications of M. nigriventris (Fe- talia: postgonites rather small, black. Body length doseeva, 1969; Fedoseeva & Perfiljeva, 1988). 2.5–3.0 mm. Distribution. – Denmark: all districts except SJ and 192. Meromyza sibirica EJ. Sweden: north to NB along Baltic coast. Norway: Fedoseeva, 1961 AK: Ryenberg, 22.vi.1850 (Siebke). Finland: Al, Ab, Figs 866–870 N, St. Russia: Vib, Kr. Widespread in Europe. Meromyza sibirica Fedoseeva, 1961: 706. Paratypes in ZISP. Biology. – Adults in June to July, and again in Sep- tember in meadows, probably two generations. Ground colour yellow. Frons slightly produced in Larvae live in shoots of Dactylis glomerata, Fes- front. Ocellar triangle little longer than wide at tuca ovina, F. rubra (Fedoseeva, 1966, 2003). Hiber- base, yellow, wrinkled at tip. Black spot on ocel- nates as larvae of the third instar. lar tubercle small. First flagellomere a little longer Larvae of the third instar described by Fedoseeva than deep, darkened above. Palpi black and slightly (1966) based on puparium. widened on apical half. Stripes of scutum wide, black or dark brown laterally, densely grey-dusted. Central stripe narrowed behind and does not 194. Meromyza variegata reach scutellum, and scutellum yellow, sometimes Meigen, 1830 with narrow fuscous yellow stripe. Abdomen usu- Figs 876–881, 1007–1009 ally with one black stripe, but sometimes with some lateral spots on tergites. Hind femora rather Meromyza variegata Meigen, 1830: 165. Syntype in thick, more than 3 times as thick as tibiae. Male MNHN.

215 Figs 866–875. Meromyza sibirica Fedoseeva: 866: hypopygium, dorsal view. 867: hypopygium, lateral view. 868: male genitalia, ventral view. 869: postgonite. 870: basiphallus. M. triangulina Fedoseeva: 871: hypopygium, dorsal view. 872: hypopygium, lateral view. 873: post- gonite. 874: postgonite, another specimen. 875: base of basiphallus.

Meromyza lidiae Nartshuk, 1992: 126. Holotype Distribution. – Denmark: SJ, NWJ, F, LFM, NEZ. and some paratypes in ZISP, other paratypes in Sweden: SK. Finland: Ab, N. Russia: Vib. Not found NHMH (Nartshuk, 2006). in Norway. European species distributed southwards to Bul- Ground colour yellowish. Frons produced in front garia. no more than 1/2 breadth of first flagellomere. Ocellar triangle shining, rugose in apical part and Biology. – Adults in July to August in dry meadows. black on ocellar tubercle only. Palpi black on api- Proposed host plant Phleum phleoides. cal part. Stripes of scutum black, densely grey- dusted or central one reddish brown. Central stripe 195. Meromyza zimzerla ends before scutellum and scutellum yellow. Pleura with brown to black marks. Abdomen with cen- Nartshuk, 1992 Figs 882–885 tral black stripe and small spots on tergites 3–5. Hind femora moderately swollen, 2.5 times as thick Meromyza zimzerla Nartshuk, 1992: 135. Holotype as tibiae. Aedeagus with bifurcate process at base. and some paratypes in ZISP, remaining paraty- Body length 3.5–4.0 mm. pes in NHMH.

216 Figs 876–885. Meromyza variegata Meigen: 876: hypopygium, dorsal view. 877: hypopygium, lateral view. 878: postgonite. 879: basiphal- lus, lateral view. 880: basiphallus, latero-ventral view. 881: basiphallus, ventral view. M. zimzerla Nartshuk: 882: hypopygium, dorsal view. 883: hypopygium, lateral view. 884: postgonite. 885: base of basiphallus.

Ground colour yellow. Frons produced in front no rather large and black. Surstyli long. Body length more than half width of first flagellomere. Ocellar 3.5–4.0 mm. triangle shining wrinkled in apical part, ocellar tu- Distribution. – Denmark: WJ, NEJ. Sweden: SK, SM, bercle with black spot. Palpi yellow. Stripes of scu- ÖL,NÄ.Norway:AK:Drøbak(Siebke).Finland:Al, tum black to reddish brown, densely grey-dusted. N, Ka. Not found in Russian parts of Fennoscandia. Central stripe not reaching scutellum, and scutel- European species. lum yellow. Pleura with red mark on katepisternum Biology. – Adults in July to August, in dry meadows and brown mark on katepimeron. Abdomen usu- with Phleum phleoides, probably its host plant. ally with 3 black or brown stripes. Hind femora 3 times as thick as tibiae. Male genitalia: postgonites

217 Genus Neohaplegis Beschovski, 1981 a broadly rounded apex. Genae narrower than first flagellomere. Scutum dull, covered posteriorly with Neohaplegis Beschovski, 1981a: 51. Type species: Os- grey dust not or just reaching level of anterior npl cinis tarsata Fallén, 1820 by original designa- setae. Legs black with yellow to black trochanters. tion. Body length 2.0–2.5 mm. Species with entirely black body. Ocellar triangle Distribution. – Denmark: NEZ: without further large and shining. vte and vti setae subequal in size. data (R.C. Stæger), 1 male, 1 female; Utterslev Mose, First flagellomere of antennae rounded. Tibial or- 17.vi.1906 (W. Lundbeck), 1 female. Sweden: north gan present. Male genitalia: surstyli short, granu- to NB. Finland: north to Om: Pietarsaari. Russia: lated; anus flanked by a pair setulose sclerites; dis- Vib, Kr. Not so far found in Norway. tiphallus simple; testes globular, not tubular as in Trans-Palearctic species. Cryptonevra and other genera of Chloropinae. Biology. – Adults in June to July in wetlands with Distribution. – Palearctic Region. Carex. Larvae live in shoots of Carex (Nartshuk, 1972a). Key to species of Neohaplegis 1 Ocellar triangle smooth and shining, with Genus Parectecephala Becker, 1910 straight side margins and acute apex. Legs Parectecephala Becker, 1910: 105. Type species: Os- black with brown to black trochanters. Male cinis longicornis Fallén, 1820 by designation of genitalia Figs 886–889 . . . . . N. glabra (Duda) Duda, 1933: 208. – Ocellar triangle subshining, with convex side margins and a rounded apex. Legs black with Body of Chlorops-type with elongated first flag- ellomere and velvety black spot in anterior part yellow to black trochanters. Male genitalia of inner lateral stripes. Head higher than long. Figs890,891...... N. tarsata (Fallén) Ocellar triangle large, reaching front margin of frons. First flagellomere squarish, longer than wide. 196. Neohaplegis glabra Arista white pubescent. Scutum yellow with black (Duda, 1933) stripes, smoothly dusted. Scutellum rounded, con- Figs 886–889 vex. Wings of Chlorops-type. Tibial organ absent. Male genitalia: middle lobe of surstyli with small, Haplegis glabra Duda, 1933: 131, as var. of H. tarsata flat apical part; pre- and postgonites parallel to (Fallén). Lectotype and paratype in ZMHU (Is- may, 1994a). each other. Body black. Frons dull. Ocellar triangle smooth Distribution. –Worldwide. and shining, with straight side margins and acute apex. Genae narrower than first flagellomere. Scu- 198. Parectecephala longicornis tum subshining, rather densely setulose, covered with brown dust posteriorly extending beyond an- (Fallén, 1820) terior npl setae. Legs black with brown to black Figs 20, 892–898 trochanters. Body length 2.0–2.5 mm. Oscinis longicornis Fallén, 1820: 4, as var. of O. lin- Distribution. – Found only in Sweden: ÖL. eata (Fabricius). Lectotype in ZMLU (Anders- Distributed in Europe and Kasakhstan. son, 1963, 1966). Andersson referred the name longicornis to Zetterstedt and accordingly desig- 197. Neohaplegis tarsata nated a lectotype from Zetterstedt’s collection. All specimens from Fallén’s collection belong to (Fallén, 1820) other species (Andersson, 1963: 104). However, Figs 890, 891 current usage refers the name to Fallén. Oscinis tarsata Fallén, 1820: 5. Lectotype in ZMLU Frons shorter than wide, yellow along eye margins (Andersson, 1963). and brown near ocellar triangle. Ocellar triangle Body black. Frons dull. Ocellar triangle subshining, reaching front margin of frons, narrowed anteri- with convex side margins and anteriorly ending in orly, yellow or brown with large black shiny spot.

218 Figs 886–891. Neohaplegis glabra (Duda): 886: hypopygium, dorsal view. 887: hypopygium, lateral view. 888: hypopygium, ventral view. 889: hypandrium and phallic complex. N. tarsata (Meigen): 890: hypopygium, ventral view. 891: male genitalia, ventral view. (886–889 after Ismay, 1994; 890 after Beschovski, 1981.)

Face and genae yellow. Genae as wide as first flag- ifrons Fabricius, 1798) by designation of Curtis, ellomere. First flagellomere 1.5 times longer than 1839: 725. wide, yellow, darkened above. Arista white, thick- ened. Palpi black. Thorax yellow or slightly reddish. Body brown, elongated, hind femora thickened. Scutum with 5 black, grey-dusted stripes. Lateral Head longer than high. Frons strongly produced stripes anteriorly with round velvety spot. Scutel- for at least half length of eyes. Eyes with horizontal lum yellow with median black stripe. Pleura with long axis. Ocellar triangle large, occupying almost black marks, spot on katepisternum sometimes entire frons, broadly reaching and semicircular at reddish. Anepisternum with setulae below. Ab- front margin of frons. Surface of frons covered with domen brown. Legs mainly yellow, femora with numerous setulae. Face sloping backwards, more black stripes. Body length 2.5–3.0 mm. or less keel-shaped. Antennae prolonged, first flag- ellomere longer than wide, sometimes narrowed in Distribution. – Denmark: widespread. Sweden: SK, apical part. Scutum flat, shining with brown or red HA, ÖL. Not found in Norway, Finland or Russian stripes and covered with numerous setulae. Scutel- parts of Fennoscandia. Central and South Europe, Near East and Asia. lum oval, flat, apical setae present or absent. Pleura extensively haired. Middle tibiae with 1–3 apical Biology. – Adults from the end of July to Septem- spurs. Hind femora thickened, ventrally in apical ber in dry, sandy meadows. Tschirnhaus (1981) sug- part with a ridge covered with granules. Hind tib- gested Corynephorus canescens, Carex arenaria or iae correspondingly curved. Male genitalia: middle Ammophila arenaria as possible host plants. lobe of surstyli flat, squarish or triangular; prego- nites slender, postgonites short.

Genus Platycephala Fallén, 1820 Distribution. – Palearctic and Oriental Regions.

Platycephala Fallén, 1820: 2. Type species: Platy- cephala culmorum Fallén, 1820 (= Musca plan-

219 Figs 892–898. Parectecephala longicornis (Fallén): 892: head, dorsal view. 893: head, lateral view. 894: antenna. 895: wing. 896: hypopy- gium, ventral view. 897: hypandrium and phallic complex. 898: apex of ovipositor, dorsal view.

Key to species of Platycephala Fallén Species key to known 3rd instar larvae of Platycephala 1 First flagellomere longer and much narrower at tip than at base (Fig. 907). Frons without Larvae 6–8 mm long, body cylindrical, milky white, punctures. Male genitalia Fig. 908 with or without a small sclerotization on the first ...... P. umbraculata (Fabricius) thoracic segment. Cephalopharyngeal skeleton black, hypostomal and tentoropharyngeal seg- – First flagellomere shorter and slightly nar- ments fused. Mandibles with accessory teeth. An- rower at tip than at base (Fig. 900). Frons with terior spiracles transverse, of rosette type. Poste- black punctures. Male genitalia Fig. 905 rior spiracles sessile on surface of 8th abdominal ...... P. planifrons (Fabricius) segment, with interspiracular branched processes.

220 Body with numerous spicules dorsally or dorsally Grochowska, 1986, 1994, 1995). Only one genera- and ventrally. tion per year. Grochowska (1986, 1994) described the life-cycle of the species in Poland, Skuhravý 1 Abdominal segments dorsally and ventrally & Skuhravá (1978) reported than the species Hi- with several rows of sclerotized spicules (Fig. bernates as adults in galls of Lipara in former 909). Spicules different in size, some of them Czechoslovakia. Larvae of Calamoncosis aprica live large and hook like (Fig. 914). Mandibles with as inquilines in stems of reed infested by P. plani- 3 large accessory teeth in each row (Fig. 911) frons (Grochowska, 2002)...... P. planifrons (Fabricius) Some larval parts described by Wandolleck (1899) and Nartshuk (1987), egg, all larval instars – Abdominal segments only dorsally with sev- and pupa by Grochowska (1994). eral rows of sclerotized spicules (Fig. 915). Spicules uniformly small (Fig. 917). Mandibles 200. Platycephala umbraculata with small accessory teeth only (Fig. 916) ...... P. umbraculata (Fabricius) (Fabricius, 1794) Figs 906–908, 915–917 199. Platycephala planifrons Musca umbraculata Fabricius,1794:348.Typelost, in Coll. Fabricius, ZMUC, only a name label (Fabricius, 1798) (Zimsen, 1964). Figs 7a, 899–905, 909–914 Platycephala nigra Meigen, 1830: 26 (Nartshuk, 1997a). Musca planifrons Fabricius, 1798: 565. Type lost, in Coll. Fabricius, ZMUC, only a name label (Zim- Bodyyellowishbrown.Fronsaslongaswide, sen, 1964). slightly produced anteriorly. Face with a median carina. Ocellar triangle large, occupying nearly en- Body yellowish brown. Frons longer than wide, pro- tire frons, but side margins slightly incurved, with duced anterior to eyes for 1/2 length of eye. Face shallow median sulcus, otherwise smooth with fine with a median carina. Ocellar triangle large, oc- punctures each inserted by a small setula. First flag- cupying entire frons, semicircular anteriorly, with ellomere 3 times longer than deep, narrowed at median sulcus and covered with black punctures apex. Apical segment of arista distinctly pubescent, each inserted by a small setula. First flagellom- white. Genae roughly wrinkled, behind as wide ere about 2 times longer than deep, rounded api- as eye. vi absent. Scutum shining, with brownish cally. Apical segment of arista distinctly pubescent, stripes, roughly punctured. Thoracic setae longer white. Genae roughly wrinkled, behind as wide than in P. planifrons. Scutellum rounded behind as eye. vi setae absent. Scutum with three dark with apical setae nearly a half as long as scutellum, brown stripes, roughly punctured and separated by subapical setae absent. Pleura with pale punctures. pale unpunctured stripes along dorsocentral lines. Abdomen flattened, long and narrow. Body length Scutellum nearly squarish, apical setae absent. 4–6 mm. Pleura with coarse black punctures and pale setu- lae. Abdomen flattened, long and narrow, dusted Distribution. – Denmark: SJ, EJ, WJ, NEJ, F, LFM, and with pale setulae. Body length 6–8 mm. NEZ.Sweden:SK,BL,HA,UP.Norway:Ø,VE.Fin- land: Al, Ab, N, Ta, Om. Not found in the Russian Distribution. – Denmark: all districts except NWJ parts of Fennoscandia. andB.Sweden:northtoHS.Norway:Ø,AK,VE, Trans-Palearctic species, known from the British AAy. Finland: north to ObS. Russia: Vib, Kr. Isles to Far East of Russia. More abundant in Asian Eurasian species. part of range. Biology. – Adults from June to September in beds Biology. – Adults in July to August. Less common of Common reed. One generation per year. and more demanding than P. planifrons,inbedsof Larvae phytophagous, living in stems of Com- Common reed growing in relatively dry habitats. mon reed (Phragmites australis). They induce little Larvae phytophagous, living in young shoots of visible stem galls that make the central leaf wither Phragmites australis at or slightly above ground and shorten two internodes of the infested stem level (Nartshuk, 1987; Grochowska, 1998). (Wandolleck, 1899; Skuhravý & Skuhravá, 1978; Some larval parts illustrated by Nartshuk (1987).

221 Figs 899–908. Platycephala planifrons (Fabricius): 899: head, dorsal view. 900: head, lateral view. 901: antenna. 902: wing. 903: thorax, lateral view. 904: hind femur and tibia. 905: male genitalia, ventral view. P. umbraculata (Fabricius): 906: head, dorsal view. 907: head, lateral view. 908: male genitalia, ventral view.

222 Figs 909–917. Platycephala planifrons (Fabricius), 3rd instar larva: 909: lateral view. 910: facial mask. 911: cephalopharyngeal skeleton. 912: anterior spiracle. 913: posterior spiracle. 914: spicules on larval body. P. umbraculata (Fabricius), 3rd instar larva: 915: lateral view. 916: cephalopharyngeal skeleton. 917: spicules on larval body.

Genus Pseudopachychaeta Fig. 927. Ovipositor Figs 928, 929 Strobl, 1902 ...... P. rufijiceps (Zetterstedt) – Scutum yellow with 3 black stripes distinctly Pseudopachychaeta Strobl, 1902: 500, as subgenus separated. Ocellar triangle yellow or darkened of Diplotoxa Loew, 1863. Type species: Diplotoxa (Pseudopachychaeta) pachycera Strobl, 1902 by ...... 2 monotypy. 2 Female cerci narrower, soft and densely pubescent (Figs 921, 922). Male genitalia Fig. Small species with procurved veins R2+3 and R4+5 920...... P. approximatonervis (Zetterstedt) and crossveins close together. Head higher behind than in front. Ocellar triangle shining or thinly – Female cerci deeper, compressed, strongly dusted. oc very strong, procurved. vte twice as long sclerotized and without pubescence (Figs 925, as vti. poc short, cruciate. 2–3 very strong, recurved 926). Male genitalia Figs 923, 924 orb. Genae widest behind, at middle not as broad ...... P. oscinina (Fallén) as first flagellomere. First flagellomere rounded. Arista thickened or slender. Scutum dusted, yellow 201. Pseudopachychaeta with black, sometimes very wide and fused stripes. Tibial organ absent. Male genitalia: surstyli fused approximatonervis with epandrium, coarsely granulate; post- and pre- (Zetterstedt, 1848) gonites fused. Figs 918–922 Distribution. –Worldwide. Oscinis approximatonervis Zetterstedt, 1848: 2622. Key to species of Pseudopachychaeta Lectotype in ZMLU (Andersson, 1966). Strobl Frons as long as wide, yellow, black haired with two long orb. Ocellar triangle reaching half length 1 Scutum with dusted black stripes fused. Ocel- of frons followed by linear extension to front mar- lar triangle entirely darkened Male genitalia gin of frons, yellow or brownish, thinly dusted. Face

223 and genae yellow, genae narrowed in front. All se- Known from Europe southwards to Spain tae on genae of females usually pale. Antennae (Nartshuk, 2004) and eastwards to Kazakhstan and brown, first flagellomere darkened, arista black. Yakutia, East Siberia. Scutum yellow with 3 black, dusted stripes, median stripe not reaching scutellum. Scutellum yellow. Biology. – Adults from the end of May to July, found Abdomen yellow to brown, dusted. Legs mainly in wet habitats. yellow. Female cerci cylindrical, covered in rather Larvae phytophagous, living in inflorescences of long, dense pubescence. Eleocharis spp. (Nartshuk, 1964). Distribution. – Denmark: NWJ, LFM, NEZ. Sweden: north to TO. Finland: north to Le. Russia: Vib, Kr. 203. Pseudopachychaeta rufijiceps Not so far found in Norway. (Zetterstedt, 1838) Holarctic species. Figs 927–929, 981 Biology. – Adults in May to June and again in Oscinis rufijiceps Zetterstedt, 1838: 780. Lectotype in September, probably overwintering. In freshwater ZMLU (Andersson, 1966). marches and other wet habitats. Fronsaslongaswide,yellow,blackhairedwithtwo Larvae phytophagous, living in inflorescences of long orb. Ocellar triangle reaching half length of various Cyperacea. Different species of Eriopho- frons followed by linear extension to front margin rum other than E.vaginatum according to Nartshuk of frons, brownish to black, dusted. Face and genae (1962a). Tschirnhaus (1981) listed as host plant yellow to brownish. Genae narrowed in front. An- Schoenoplectus tabernaemontani. The larvae are in tennae brown, first flagellomere darkened, arista North America (Valley et al., 1969) reported from black. Scutum yellow with 3 black, dusted fused inflorescences of Eleocharis obtusa and E. smalli stripes, median stripe reaching scutellum. In dark feeding on unripe seeds. coloured specimens scutal stripes more or less con- fluent with yellowish patches only on notopleura 202. Pseudopachychaeta oscinina and anterior part of postpronota. Scutellum yellow (Fallén, 1823) to brownish. Abdomen and legs darkened. Female Figs 923–926 cerci cylindrical, covered with setulae.

Phytomyza oscinina Fallén, 1823: 4. Holotype in Distribution. – The most abundant species of the ZMLU (Nartshuk & Andersson, 2002). genus, especially to the north. Sweden: north to TO. Lasiosinaheleocharis Nartshuk, 1964: 321 (Nartshuk Norway: Bv, HOy, HOi, Fn. Finland: north to Li. Rus- & Andersson, 2002). sia: Vib, Kr. Lr. Not from Denmark. Trans-Palearctic arcto-boreal species, ranging Frons as long as wide, yellow, black haired with from northern parts of the British Islands to the two long orb. Ocellar triangle reaching half length Chukotsky Peninsula, Far East Russia. of frons followed by linear extension to front mar- gin of frons, yellow or slightly brownish, thinly Biology. – Overwintered adults appear in spring, dusted. Face and genae yellow, genae narrowed in new generation active from July to September front. Part of setae on genae dark. Antennae brown, (Nartshuk, 1962a). Tyrphobiont, mainly in raised first flagellomere strongly darkened, arista black. bogs. Scutum yellow with 3 wide black, dusted stripes, Females lay eggs in early spring on inflores- median stripe not reaching scutellum. Scutellum cences of Eriophorum vaginatum and other species yellow. Pleura darkened, spot on katepisternum of Eriophorum (e.g. E. polystachyum, E. latifolium, and postnotum black, dusted. Abdomen brownish. but not on the delicate E. gracilis). Larvae phy- Legs mainly yellow. Female cerci shining, blade- tophagous, living 4–6 larvae together in one inflo- like, laterally compressed, almost without pubes- rescence, feeding on the developing seeds. They cence. pupate within the inflorescence. Tentoropharyngeal sclerites and pupa are illus- Distribution. – Sweden:SK,VG,UP,NB.Finland:Ta. trated by Nartshuk (1987). Russia: Vib. Not so far found in Denmark and Nor- way.

224 Figs 918–929. Pseudopachychaeta approximatonervis (Zetterstedt): 918: head, dorsal view. 919: head, lateral view. 920: male genitalia, ventral view. 921: ovipositor, lateral view. 922: apex of ovipositor, lateral view. P. oscinina (Fallén): 923: hypopygium, dorsal view. 924: male genitalia, ventral view. 925: ovipositor, lateral view. 926: apex of ovipositor, lateral view. P. rufijiceps (Zetterstedt): 927: hypopygium, dorsal view. 928: apex of ovipositor, lateral view. 929: ovipositor, lateral view.

225 Genus Thaumatomyia Zenker, 1833 with arched membranous vesicula seen best in macerated abdomen (Fig. 69) Thaumatomyia Zenker, 1833: 344. Type species: ...... T. glabra (Meigen) Thaumatomyiaprodigiosa Zenker, 1833 (= Chlo- – Scutum and scutellum distinctly haired. Scu- rops notatus Meigen, 1830) by monotypy. tum with black or red stripes, middle stripe Chlorops-like yellow species with black or red usually ending before scutellum. Male ab- stripes on scutum, flattened scutellum with ap- domen without membranous vesicula . . . . . 4 proximated apical scutellar setae. Frons somewhat 4 Body yellow with black or red stripes on scu- projecting, as broad as long. Ocellar triangle large tum. Male genitalia Figs 946, 947 with straight or convex sides, reaching or almost ...... T. rufa (Macquart) reaching front margin of frons, with one or more rows of setulae. First flagellomere rounded, usu- – Body darkened with black stripes. Male geni- ally longer than wide. Scutum rather convex with talia Figs 937, 938 . . . T. hallandica Andersson 5 black or red stripes, smooth and shining, al- most bare or covered with short setulae. Scutellum 204. Thaumatomyia glabra rounded or somewhat triangular, flat, bare or with (Meigen, 1830) numerous setulae. Apical scutellar setae close to- Figs 69, 930–936, 960–962 gether. Tibial organ present. Middle lobe of surstyli with nail- or hook-shaped apex; pre- and postgo- Chlorops glabra Meigen, 1830: 149; Meigen’s plates nites situated in line. Males of some species have 162: 4, 211: 18 (Morge, 1976b). Lectotype in eversible vesicles in the intersegmental membrane NHMW (Nartshuk, 1997a). of the postabdomen that become inflated during Frons slightly longer than wide. Ocellar triangle the premating period. These vesicles probably emit large, mainly black, linearly yellow at side mar- short-range sex pheromones. gins and posterolateral corners. Genae very wide. Distribution. –Worldwide. First flagellomere about 1.2 times as long as wide, distally brown, basally yellow. Only vte setae dis- tinct on head. Scutum glabrous with three broad Key to species of Thaumatomyia black stripes, all reaching scutellum. Setulae cover- Zenker ing scutum minute. Scutellum flat, nearly bare, api- cal setae black, approximated. Abdomen black dor- 1 Genae narrow, no more than half width of first sally. Male postabdomen with large eversible vesi- flagellomere; ocellar triangle with two rows of cles. Body length 2.3–2.6 mm. setulae inside lateral margins (Figs 939, 940). Male abdomen with two lateral membranous Distribution. – Common species. Denmark: found vesicules seen usually in macerated abdomen in all districts except SZ. Sweden: north to TO. Nor- (Fig.70)...... T. notata (Meigen) way:Ø,AK,HEs,Bv,TEy,VAy,Ry,HOy,HOi,SFy,STi, NTi. Finland: north to Le. Russia: Vib, Kr, Lr. – Genae as wide as or wider than first flagellom- Holarctic species. ere; ocellar triangle with one row of setulae inside lateral margins (Figs 930, 931, 944, 945, Biology. – Adults from June to September, abun- 948,949)...... 2 dant everywhere, found in a variety of habitats including freshwater marshes, very often on flower- 2 Ocellar triangle carinate in apical part; genae ing Medicago (Fabaceae). Aggregating males have very wide, wider than first flagellomere (Figs been observed on the inflorescence of Wisteria 948, 949). Male genitalia Figs 950, 951 sinensis in California, with single males occupy- ...... T. trifasciata (Zetterstedt) ing an individual blossom (Sorensen & Sorensen, – Ocellar triangle without carina in apical part, 1997). Steyskal (1945) described the behavior of genae not wider than first flagellomere (Figs a North American subspecies of this species (T. glabra var. bistriata Walker). He observed that 930,931,944,945)...... 3 males with exposed terminal vesicles would walk 3 Scutum and scutellum with sparse setulae. about on sugar maple leaves fluttering their wings Scutum with black, shining stripes, middle and swelling their vesicles. On approaching an- stripe reaching scutellum. Male abdomen other (male?) individual they would go infight us-

226 Figs 930–938. Thaumatomyia glabra (Meigen): 930: head, dorsal view. 931: head, lateral view. 932: antenna. 933: wing. 934: male gen- italia, ventral view. 935: hypandrium and phallic complex. 936: right surstylus. T. hallandica Andersson: 937: mesolobus and surstylus. 938: hypandrium and phallic complex. ing their front legs, raised to an angle of 45°, and 205. Thaumatomyia hallandica eventually one of them would be pushed over the Andersson, 1966 edge of the leaf and forced to escape. It was found Figs 937, 938 that the males were attracted by methyl anthrani- late (Landolt et al., 2000). Sabrosky (1935) men- Thaumatomyia hallandica Andersson, 1966: 63. tioned that this species was taken in a light trap, Holotype in ZMLU. although chloropids are seldom taken at light. Body generally infuscate. Frons slightly longer than Larvae carnivorous, living among roots of plants wide, dull. Ocellar triangle large, almost reach- feeding on root aphids, especially Pemphigidae, ing front margin of frons, shining, brownish black, as reported by Parker (1918) and in many subse- with straight side margins. Genae a little wider quent publications, see Nartshuk (1972b). There than first flagellomere. First flagellomere about 1.2 is one record of the species from a spider egg sac times as long as wide, black or basally yellow. Arista brown, shortly pubescent. vte strong. Palpi yellow (Sabrosky, 1943). It has been reared from compost to brown. Scutum glabrous with 5 broad black (Werner, 1997) that no doubt contained larvae im- stripes, middle one ending at middle of scutum. ported with plant material. Scutellum flat, apical setae black, approximated. Larvae described by Balachowsky & Mesnil Abdomen brown dorsally. Male without postab- (1935), puparium figured by Nartshuk (1987). dominal vesicles. Body length 2.0–2.6 mm.

227 Figs 939–947. Thaumatomyia notata (Meigen): 939: head, dorsal view. 940: head, lateral view. 941: wing. 942: right surstylus, ventral view. 943: hypandrium and phallic complex. T. rufa (Macquart): 944: head, dorsal view. 945: head, lateral view. 946: hypopygium, right half, ventral view. 947: hypandrium and phallic complex.

Distribution. – Denmark: found in all districts ex- ferent width, with 2–3 irregular rows of setulae cept SZ and NWZ. Sweden: north to LU. Norway: along side margins. Genae narrow, nearly 1/3 of north to Nnø. Finland: north to Le. Russia: Vib, Kr. first flagellomere. First flagellomere nearly orbic- Eurasian species, ranging from the British Isles to ular or sometimes very slightly wider than long, Mongolia and Central Asia. almost entirely black or brownish basally. vte and Biology. – Adults from June to August in dry mead- vti setae both developed and rather strong. Scu- ows and sand dunes. tum with 5 black, rarely partly reddish, stripes Larvae carnivorous, living among roots of plants not reaching scutellum. Scutellum flat, apical se- feeding on root aphids. tae black, approximated. Pleura with black or red spot on katepisternum. Male with pair of eversible 206. Thaumatomyia notata postabdominal vesicles. Body length 2.5–3.0 mm (Meigen, 1830) Figs 7b, 70, 110–114, 939–943 Distribution. – Very common. Denmark: SJ, LFM, NEZ. Sweden: north to NB along the Baltic coast. Chlorops notata Meigen, 1830: 144; Meigen’s plate Norway: AK, Os, HOi. Finland: north to Om. Russia: 211: 17 (Morge, 1976). Lectotype in NHMW Vib, Kr. (Nartshuk, 1997a). Widespread species, recorded from the Palearc- Fronsnearlyaslongaswide.Ocellartrianglefrom tic, Afrotropical and Oriental Regions. Palearctic black to yellow with median black stripes of dif- distribution from the British Isles to Japan.

228 Biology. – Adults from May to October, hiber- nate, found in a variety of habitats, often visit flowers. In Norway found by canopy fogging of Pinus silvestris (Thunes et al., 2004). In Switzer- land the fly has been observed by Pschorn-Walcher (1956) feeding on secretions from the larvae of Agelastica alni (Chrysomelidae). Adults aggregate in holes and crevices such as bird’s nests and be- tween scales of Picea cones for overwintering (Kri- vokhatsky & Nartshuk, 2001). Occationally, several millions of specimens enter houses in the autumn for overwintering. This has repeatedly been re- corded from various parts of Europe including the Nordic countries: Finland (Haartman, 1950) and Sweden (T. Pape, pers. comm.). Lampa (1893) de- scribed a swarm covering a fir tree of 15.5 m height. For more details about occurrences, see the chapter ‘Role in land ecosystems and species of economic importance’. Larvae carnivorous, living in the ground among roots of plants feeding on root aphids (Mesnil, 1933; Balachowsky & Mesnil, 1935; Aszél, 1942; Yarkulov, 1971). Reared from compost (Werner, 1997), un- Figs 948–951. Thaumatomyia trifasciata (Zetterstedt): 948: doubtedly from larvae imported with roots of head, dorsal view. 949: head, lateral view. 950: surstylus. 951: hy- plants. pandrium and phallic complex. Larvae described by Balachowsky & Mesnil (1935), some larval structures illustrated by Biology. – Adults from middle of June to August, in Nartshuk (1987). meadows. Larvae carnivorous, living in the ground among roots of plants where feeding on root 207. Thaumatomyia rufa aphids. It has been reared from compost (Werner, 1997). (Macquart, 1835) Anterior spiracles of larvae illustrated by Figs 944–947 Nartshuk (1987). Chlorops rufa Macquart, 1835: 593. Type probably 208. Thaumatomyia trifasciata in MHNL. (Zetterstedt, 1848) Frons slightly longer than wide. Ocellar triangle en- Figs 948–951 tirely brownish yellow to blackish, reaching front margin of frons, with one row of setulae along Oscinis trifasciata Zetterstedt 1848: 2609. Lecto- slightly concave side margins. Genae nearly as wide type in ZMLU (Andersson, 1966). as first flagellomere. First flagellomere distinctly Frons slightly longer than wide. Ocellar triangle longer than wide, mainly yellow, darkened dor- usually black, extending nearly to front margin sally. Head only with distinct vte setae. Scutum of frons, with one row of setulae along side mar- with 5 stripes not reaching scutellum. Stripes black gins and on anterior half of central ridge. Genae or sometimes reddish brown. Spot on anepister- 1.5–2.0 times wider than first flagellomere is deep. num black, remaining pleural spots usually reddish First flagellomere slightly longer than deep, mainly brown. Male without postabdominal vesicles. Body yellow, darkened dorsally. vte setae absent. Scu- length 2.0–2.5 mm. tum with 5 black stripes, central one not reach- ing scutellum. Pleural spots black. Male without Distribution. – Denmark: SJ, WJ, F, LFM, NEZ, B. postabdominal vesicles. Body length 3.5–4.0 mm. Sweden: north to TO. Norway: O, VE, HOi. Finland: north to Om. Russia: Vib, Kr. Distribution. – Widespread in Denmark and Fen- Trans-Palearctic species, known from the British noscandia. Isles to Japan and southwards to North Africa. Holarctic species.

229 Figs 952–955. Trichieurina pubescens (Meigen): 952: head, dorsal view. 953: head, lateral view. 954: hypopygium, dorsal view. 955: male genitalia, ventral view.

Biology. – Adults from June to August, found in wet 209. Trichieurina pubescens places such as bogs, marshes, swamps, fens and lit- (Meigen, 1830) toral zone of lakes. In Norway found by canopy fog- Figs 952–955 ging of Pinus silvestris (Thunes et al., 2004). Larvae as in other species of the genus carnivo- Eurina pubescens Meigen, 1830: 4; Meigen’s plate rous, feeding on root aphids among roots of plants. 209: 4 (Morge, 1976b). Lectotype in NHMW Meijere (1950) found larvae among roots of Ci- (Nartshuk, 1997a). chorium intybus (Asteraceae) in the Netherlands. Densely grey-dusted species, body covered with A. Przhiboro (pers. comm.) reared the species from longhairs.Fronsprojectingbeyondeyesaslong plant roots growing at the littoral zone of the White as length of an eye. Ocellar triangle reaching front Sea. margin of frons, dusted, with a longitudinal median groove and long hairs along sides. Second antennal Genus Trichieurina Duda, 1933 segment as long as wide, first flagellomere longer than wide; arista dark, basally whitish. Scutum dark Trichieurina Duda, 1933: 126. Type species: Eurina grey-dusted with double central line brown, co- pubescens Meigen, 1830 by original designation. vered with long hairs. Pleura, abdomen and legs completely dusted. Greyish species with strongly produced frons. Body covered with very thick dust and extremely long Distribution. – Found only in Finland: Kb: Ham- slender hairs. Head triangular in side view. Eyes maslahti, Pyhäselkä, 15.vi.1935, 04.vi.1938, small. Genae very wide. Ocellar triangle reaching 07.vii.1939 (Kontkanen), 2 males, 5 females. front margin of frons, covered with long hairs. First The species is known from Central and East Eu- flagellomere longer than wide, arista nearly bare. rope and Kazakhstan. Tibial organ absent. Male genitalia: middle lobe Biology. – Kontkanen (1935) described in detail the of surstyli serrate, postgonites and pregonites dis- habitat in Finland. The species occurs in swamps tinct, the latter transversally directed. with sedges. Carex spp. are likely host plants.

230 Figs 956–966. Camarota curvipennis (Latreille), 3rd instar larva: 956: posterior end. 957: anterior spiracle. 958: posterior spiracle. 959: spicular zones. Thaumatomyia glabra (Meigen), 3rd instar larva: 960: mature larva. 961: anterior end. 962: posterior spiracle. Platycephala planifrons (Fabricius), 3rd instar larva: 963: posterior end. Polyodaspis rufijicornis (Macquart), 3rd instar larva: 964: cephalopharyngeal skeleton. 965: spicular zones. Pseudogaurax chiyokae Kanmiya, puparium: 966: dorsal (left) and ventral (right). (956–962 after Bala- chowsky & Mésnil, 1935; 964, 965 after Kiauka, 1974; 966 after Kanmiya, 1972.)

231 Figs 967–977. Hapleginella laevifrons (Loew), 3rd instar larva: 967: anterior end of mature larva, lateral view. 968: cephalopharyngeal skeleton. 969: posterior end of mature larva, lateral view. 970: posterior end of puparium, ventral view. Gaurax dubius (Macquart), 3rd instar larva: 971: cephalopharyngeal skeleton. 972: spicules on 7th abdominal segment. 973: posterior end of puparium, ventral view. Cetema neglectum Tonnoir, 3rd instar larva: 974: facial mask. 975: cephalopharyngeal skeleton. 976: spicular zones. 977: posterior end. (971, 972 after Smith, 1964; 974–977 after Nye, 1958.)

232 Figs 978–985. Conioscinella frontella (Fallén), 3rd instar larva: 978: facial mask. 979: cephalopharyngeal skeleton. 980: spicular zones. Pseudopachychaeta rufijiceps (Zetterstedt): 981: posterior end of puparium. Lasiosina herpini (Guérin-Méneville): 982: mandibula, 3rd instar larva. 983: posterior end of puparium. L. brevisurstylata Dely-Draskovits: 984: mandibula, 3rd instar larva. 985: posterior end of puparium. (978–980 after Nye, 1958; 982–985 after Beschowsky & Krasteva, 1995.)

Figs 986–995. Chlorops strigulus (Fabricius), 3rd instar larva: 986: dorsal view. 987: posterior end. C. frontosus Meigen, 3rd instar larva: 988: dorsal view. 989: facial mask. 990: posterior end. C. limbatus Meigen, 3rd instar larva: 991: dorsal view. 992: facial mask. 993: pos- terior end. C. zernyi Duda, 3rd instar larva: 994: posterior end. C. planifrons Loew, 3rd instar larva: 995: posterior end. (986–993 after Balachowsky & Mésnil, 1935; 994 after Zhabinskaya, 1963.)

233 Figs 996–1004. Chlorops pumilionis (Bjerkander), 3rd instar larva: 996: facial mask. 997: cephalopharyngeal skeleton. 998: posterior end. C. interruptus Meigen, 3rd instar larva: 999: facial mask. 1000: posterior end. 1001: posterior spiracle. C. speciosus Meigen, 3rd instar larva: 1002: facial mask. 1003: cephalopharyngeal skeleton. 1004: posterior end. (996–998, 1002–1004 after Nye, 1958; 999–1001 after Balachowsky & Mésnil, 1935.)

Figs 1005–1014. Meromyza pratorum Meigen, 3rd instar larva: 1005: anterior spiracle. 1006: spicular zones. M. variegata Meigen, 3rd instar larva: 1007: anterior spiracle. 1008: facial mask. 1009: spicular zones. M. nigriventris Macquart, 3rd instar larva: 1010: cephalopha- ryngeal skeleton. 1011: mandibulae. 1012: posterior end. M. saltatrix (Linnaeus), 3rd instar larva: 1013: mature larva. 1014: facial mask. (1005, 1006, 1013, 1014 after Balachowsky & Mésnil, 1935; 1007, 1008 after Nye, 1958; 1009, 1010 after Fedoseeva, 1966; 1011, 1012 after Tsygankov, 1930.)

234 Abbreviations aed: aedeagus C: costal vein ane: anepisternum CuA1: fifth longitudinal vein ant1,ant2,ant3: first–third antennal segment M: medial, fourth longitudinal vein apog: anterior process of postgonite R1: first radial, first longitudinal vein ag: accessory gland R2+3:second+ third radial, second longitudinal bph: basiphallus vein c: cercus R4+5:fourth+ fifth radial, third longitudinal vein cph: cephalopharyngeal sclerite Sc: subcostal vein cx1,cx2,cx3: first–third coxa ta: r-m crossvein dph: distiphallus tp: dm-cu crossvein ds: dental sclerite egt: egg tube ej ap: ejaculatory apodeme acr: acrostical setae ep: epandrium anpl: anterior notopleural seta ep s: ventral epandrial sclerite apa: anterior postalar seta ap sc: apical scutellar setae f1,f2,f3:first–thirdfemur fc: facial carina dc:dorsocentralsetae fl1: first flagellomere, postpedicel fr:frontalsetulae fp: frontal palpi h:humeralseta hyp: hypandrium if : interfrontal setulae hys: hypostomal sclerite la sc: lateral scutellar setae kepml: ketepimerom oc: ocellar setae m: mesonotum orb:orbitalsetae md: mandilula (mouth hook) of larva pnpl: posterior notopleural setae mes: mesolobus, fused cerci ppa: posterior postalar setae mg1,mg2,mg3,mg4: first–fourth costal segment pra: prealar seta mn: metanotum prpl: propleural seta mp: maxillary palpi prs: presutural setae mt1,mt2,mt3: first–third metatarsus psc: prescutellar setae npl: notopleuron pst: peristomal setulae ph: phallapodeme pooc: postocellar, postvertical setae ph s: phallapodemic sclerite sa: supraalar seta pl: paralobes sc: scutellar setulae pog: postgonite scp: scapular seta pph: phallic paraphyses str: sensillum trichodeum ppog: posterior process of postgonite vi: vibrissa prg: pregonite prpl: propleuron vte:outerverticalsetae psc: pregenital sclerite vti: inner vertical setae pst: parastomal sclerite sp: spiracle ANPL: anepimeral spot spt: spermatheca H: humeral spot sur: surstylus KEMSP: katepimeral spot t1,t2,t3:first–thirdtibia KESSP: katepisternal spot ts: testes M1: central scutal (mesonotal) stripe v: vesicula M2: inner lateral scutal (mesonotal) stripe

235 M3: outer lateral scutal (mesonotal) stripe PTPL: pteropleural spot MN: metanotal spot S, s: sternite PRPL: propleural spot T, t: tergite

236 Catalogue 238 olnel meijerei Colliniella rohaceki Choropsina distinguenda Chloropsina zernyi Chlorops varsoviensis Chlorops troglodytes Chlorops strigulus Chorops speciosus Chlorops serenus Chlorops scutellaris Chlorops scalaris Chlorops rufijinus Chlorops rossicus Chlorops riparius Chlorops ringens Chlorops pumilionis Chlorops planifrons Chlorops pannonicus Chlorops palpatus Chlorops pallifrons Chlorops pallidiventris Chlorops obscurellus Chlorops nigripalpis Chlorops meigenii Chlorops limbatus Chlorops laetus Chlorops kirigaminensis Chlorops interruptus Chlorops hypostigma Chlorops gracilis Chlorops geminatus Chlorops frontosus Chlorops fijinitimus Chlorops fijiguratus Chlorops dasycerus Chlorops centromaculatus Chlorops calceatus Chlorops bjerkanderi Chlorops anthracophagoides Chlorops adjunctus Chlorops neglectum Cetema myopinum Cetema elongatum Cetema cereris Cetema elegantulum Centorisoma curvipennis Camarota oscinella Calamoncosis minima Calamoncosis halterata Calamoncosis glyceriae Calamoncosis duinensis Calamoncosis aspistylina Calamoncosis aprica Calamoncosis plumiger Aspistyla trilineatum Aphanotrigonum Andersson & Nartshuk norrbotticum Aphanotrigonum nigripes Aphanotrigonum inerme Aphanotrigonum hungaricum Aphanotrigonum femorellum Aphanotrigonum cinctellum Aphanotrigonum brachypterum Aphanotrigonum (Fallén) Meigen (Duda) (Zetterstedt) Meigen Loew Loew Meigen Smirnov Smirnov (Fabricius) Loew Meigen Smirnov Becker (Zetterstedt) Tonnoir (Duda) Meigen Meigen Meigen (Meigen) Strobl (Loew) (Meigen) Loew Becker (Zetterstedt) Loew Meigen (Bjerkander) (Duda) (Zetterstedt) Nartshuk (Zetterstedt) Meigen athk&Andersson & Nartshuk (Meigen) Strobl Meigen Becker (Latreille) (Strobl) (Duda) (Becker) Nartshuk athk&Andersson & Nartshuk (Strobl) Kanmiya Collin Duda (Frey) Becker (Zetterstedt) (Duda) (Zetterstedt) Strobl (Meigen) Collin Dely-Draskovits (Zetterstedt) P? P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P Germany P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P Great Britain P P P P P P P P P P P P P P P P P P SJ P P P P P P P P P P P P P P P P EJ P P P P P P P P P P P P P P P P P WJ P P P P P P P P P P P NWJ Denmark P P P P P P P P P P P P P NEJ P P P P P P P P P P P P P P P F P P P P P P P P P P P P P LFM P P P P P P P P P SZ P P P P P P P NWZ P? P? P P P P P P P P P P P P P P P P P P P P P P P P P P NEZ P P P P P P P P P P P P P P B P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P SK P P P P P P P P P P P P P P BL P P P P P P P P P P P P P P P P P P P P P P P HA P P P P P P P P P P P P P P P P P P P P P P P P P SM P P P P P P P P P P P P P P P P P P P P P P P P P P ÖL P P P P P P P P P P P P P P P P P P P P GO P P GS P P P P P P P P P P P P P P P P P P ÖG P P P P P P P P P P VG P P P P P P P P P P P P P P BO P P P DS P P P P P P P P P P P P P P P P NÄ P P P P P P P P P P P P P P P P P P P SÖ P P P P P P P P P P P P P P P P P P P P P P P P P P

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239 240 aisn brevisurstylata Lasiosina albipila Lasiosina palposa Lasiambia coxalis Lasiambia brevibucca Lasiambia baliola Lasiambia zuercheri Incertella scotica Incertella nigrifrons Incertella kerteszi Incertella karteshensis Incertella antennata Incertella albipalpis Incertella laevifrons Hapleginella suecicus Gaurax maculipennis Gaurax macrocerus Gaurax leucarista Gaurax fungivorus Gaurax flavomaculatus Gaurax fascipes Gaurax ephippium Gaurax dubius Gaurax borealis Gaurax numerata Gampsocera variegata Eutropha fulvifrons Eutropha lurida Eurina calva Eurina slesvicensis Eribolus nana Eribolus hungaricus Eribolus gracilior Eribolus danicus Eribolus puncticollis Epichlorops tuberculifera Elachiptera scrobiculata Elachiptera diastema Elachiptera cornuta Elachiptera brevipennis Elachiptera dalmatina Diplotoxoides messoria Diplotoxa vagans Dicraeus tjederi Dicraeus tibialis Dicraeus styriacus Dicraeus rossicus Dicraeus raptus Dicraeus nitidus Dicraeus ingratus Dicraeus fennicus Dicraeus flavitarsis Cryptonevra diadema Cryptonevra consimilis Cryptonevra zetterstedti Conioscinella tornensis Conioscinella sordidella Conioscinella mimula Conioscinella messaurea Conioscinella livida Conioscinella gallarum Conioscinella frontella Conioscinella abiskoi Conioscinella Egger Meigen (Zetterstedt) (Macquart) athk&Andersson & Nartshuk Haliday Duda Becker Macquart Wahlgren athk&Andersson & Nartshuk (Meigen) athk&Andersson & Nartshuk (Collin) Stackelberg (Mejiere) (Becker) (Loew) Duda Loew (Roser) (Collin) Nartshuk (Strobl) athk&Andersson & Nartshuk (Zetterstedt) (Fallén) (Haliday) (Zetterstedt) Loew (Duda) Nartshuk (Duda) Nartshuk (Meigen) (Fallén) Becker Becker (Collin) athk&Andersson & Nartshuk Collin (Zetterstedt) Collin (Meigen) (Duda) (Fallén) athk&Przhiboro & Nartshuk (Meigen) Collin (Heeger) (Loew) athk&Andersson & Nartshuk (Duda) (Meigen) (Duda) (Zetterstedt) (Zetterstedt) (Strobl) (Strobl) Andersson & Nartshuk (Corti) Dely-Draskovits Andersson P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P Germany P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P Great Britain P P P P P P P P P P P P P SJ P P P P P P P P P P P P EJ P P P P P P P P P P P P P WJ P P P P P P P P P P P P P

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241 242 ecosi&Lansbury & Beschovski brunneipenne Rhopalopterum atricorne Rhopalopterum atricillum Rhopalopterum anthracinum Rhopalopterum rufijiceps Pseudopachychaeta oscinina Pseudopachychaeta (Zetterstedt) approximatonervis Pseudopachychaeta venustus Pseudogaurax sulcicollis Polyodaspis rufijicornis Polyodaspis umbraculata Platycephala planifrons Platycephala longicornis Parectecephala gilvipes Oscinisoma germanicum Oscinisoma cognatum Oscinisoma sordidissima Oscinimorpha minutissima Oscinimorpha albisetosa Oscinimorpha ventricosi Oscinella trochanterata Oscinella pusilla Oscinella norrbottica Oscinella nitidissima Oscinella maura Oscinella frit Oscinella cariciphila Oscinella capreolus Oscinella angustipennis Oscinella angularis Oscinella tarsata Neohaplegis glabra Neohaplegis trigonella Microcercis kroeberi Microcercis zimzerla Meromyza variegata Meromyza triangulina Meromyza sibirica Meromyza saltatrix Meromyza rohdendorfiji Meromyza pratorum Meromyza pluriseta Meromyza palposa Meromyza ornata Meromyza nigriventris Meromyza nigriseta Meromyza mosquensis Meromyza ingrica Meromyza femorata Meromyza elbergi Meromyza curvinervis Meromyza bohemica Meromyza bifurcata Meromyza laterale Melanum rufijitarsis Lipara pullitarsis Lipara lucens Lipara subnigripes Lasiosina parvipennis Lasiosina intermedia Lasiosina herpini Lasiosina danielssoni Lasiosina .lat. s. Meigen Loew (Meigen) (Fallén) okčl&Chvála & Doskočil (Guérin-Méneville) (Haliday) Fedoseeva (Wiedemann) Nartshuk Fedoseeva Fedoseeva Nartshuk (Duda) Collin (Haliday) (Linnaeus) (Loew) Nartshuk Fedoseeva Péterfi (Fallén) (Duda) athk&Andersson & Nartshuk Macquart Collin Meigen Meigen (Meigen) Fedoseeva athk&Andersson & Nartshuk Dely-Draskovits athk&Andersson & Nartshuk Dely-Draskovits (Duda) Duda (Meigen) (Macquart) (Zetterstedt) (Meigen) Fedoseeva Macquart Fedoseeva Fedoseeva (Czerny) Collin (Fabricius) Duda (Duda) (Duda) (Zetterstedt) (Zetterstedt) (Fabricius) (Zetterstedt) (Strobl) (Strobl) (Zetterstedt) (Fallén) (Meigen) P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P Germany P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P Great Britain P? P P P P P P P P P P P P P P P P P P P P P SJ P P P P P P P P P P P P P P P P EJ P P P P P P P P P P P P P P P P P P P WJ P P P P P P P P P P P P P P P P P

NWJ Denmark P P P P P P P P P P P P P P P P P P P P P NEJ P P P P P P P P P P P P P P P P P P P P P F P P P P P P P P P P P P P P P P P P P P P P LFM P P P P P P P P P P SZ P P P P P P P P P P P P P P P P NWZ P P P P P P P P P P P P P P P P P P P P P P P P P P NEZ P P P P P P P P P P P P P P P P P P P P B P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P SK P P P P P P P P P P P P P P P P P P BL P P P P P P P P P P P P P P P P P P P P P P P P P P P P P HA P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P SM P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P ÖL P P P P P P P P P P P P P P P P P P P P P P P P P P P P GO P P P P P GS P P P P P P P P P P P P P P P ÖG P P P P P P P P P VG P P P P P P P P BO P P DS P P P P P P P P P NÄ P P P P P P P P P P P P P P P P SÖ P P P P P P P P P P P P P P P P P P P P P P P P P P P

UP Sweden P P P P P P P VS P P P P P P P P VR P P P P P P P P P P P P P P P P P P P P P DR P GA P P P P P P P HS P ME P HR P P P P P P P P P JÄ P P P P P ÅN P P P P VB P P P P P P P P P P P P P P P P P P P P P NB P P P P P P P ÅS P P P P P P P P P LY P PI P P P P P P P P LU P P P P P P P P P P TO Norway Finland Russia Ø+AK HE (s+n) O(s+n) B(ø+v) VE TE (y+i) AA (y+i) VA (y+i) R(y+i) HO (y+i) SF (y+i) MR (y+i) ST (y+i) NT (y+i) Ns (y+i) Nn (ø+n) TR (y+i) F(v+i) F(n+ø) Al Ab N Ka St Ta Sa Oa Tb Sb Kb Om Ok ObS ObN Ks LkW LkE Le Li Vib Kr Lr

P P P P P P

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P P P P P P P P P P P P P P P

P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P

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P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P

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243 NB: 244 and (Macquart) rcmalineella Tricimba humeralis Tricimba cincta Tricimba pubescens Trichieurina scutellata Trachysiphonella rufijiceps Trachysiphonella trifasciata Thaumatomyia rufa Thaumatomyia notata Thaumatomyia hallandica Thaumatomyia glabra Thaumatomyia halophila Speccafrons aenea Siphunculina oscinina Siphonella Andersson & Nartshuk tomentosum Rhopalopterum femorale Rhopalopterum fasciola Rhopalopterum siel frit Oscinella (Meigen) (Fallén) (Fallén) (Loew) (Meigen) (Macquart) .vastator C. (Duda) (Meigen) (Meigen) .lt ersnsacmlxo pce iha nete taxonomy: unsettled an with species of complex a represents lat. s. (Meigen) (Meigen) (Collin) (Macquart) (Zetterstedt) Andersson (Roser) (Curtis). P P P P P P P P P P P P P P P P Germany P P P P P P P P P P P P P P P Great Britain P P P P P P SJ P P P P P P EJ P P P P P WJ P P P NWJ Denmark P P P P NEJ P P P P P F P P P P P P P LFM P SZ P P P NWZ P P P P P P P P P P P P NEZ P P P P P P P B P P P P P P P P P P P P P P P SK P P P P P P P BL P P P P P P P P P P P HA P P P P P P P P P P P P SM P P P P P P P P P P ÖL P P P P P P P P GO GS P P P P P P P ÖG siel frit Oscinella P P P VG P P P P P P BO P DS P P P P P P P NÄ P P P P P P SÖ (Linnaeus), P P P P P P P P P P UP Sweden P? P P P P P VS P P P P VR P P P P P P P P P P P DR

.hortensis C. GA P P HS ME P P HR P P P JÄ P P P Collin, ÅN P P VB P P P P P P P NB .nigerrima C. P P ÅS P P P P LY P P PI P P P P P P P P LU P P P P P P TO Norway Finland Russia Ø+AK HE (s+n) O(s+n) B(ø+v) VE TE (y+i) AA (y+i) VA (y+i) R(y+i) HO (y+i) SF (y+i) MR (y+i) ST (y+i) NT (y+i) Ns (y+i) Nn (ø+n) TR (y+i) F(v+i) F(n+ø) Al Ab N Ka St Ta Sa Oa Tb Sb Kb Om Ok ObS ObN Ks LkW LkE Le Li Vib Kr Lr P P P P P P P P P P P P P P

P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P P? P P

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257 Howell, J., 1982. Parasitism and host-plant relationships International Commission on Zoological Nomenclature, in the dipterous stem borer complex of ryegrass. – Pes- 1984. Opinion 1264. Oscinis plumigera Loew, 1860 (In- ticide Science, 13: 367–372. secta, Diptera): suppression by use of the plenary pow- Hubicka, J., 1966. New species of the genus Meromyza Mg. ers. – Bulletin of Zoological Nomenclature, 41: 8–9. (Diptera: Chloropidae) from Poland. – Annales Uni- Ismay, J.W., 1975. Femoral combs – a secondary sexual versitatis M. Curie-Skłodowska, Sec. C: Biologia, 21 (2): character in British Oscinellinae (Diptera: Chloropi- 13–26. dae). – Journal of Entomology, 44: 21–25. Hubicka, J., 1967. A new species of the genus Meromyza Ismay, J.W., 1976. A revision of Oscinisoma (Diptera, Mg. (Diptera: Chloropidae) from Poland. – Bulletin de Chloropidae) in Britain. – Entomologist’s Gazette, 27: l’Académie Polonaise des Sciences, Ser. Biol., 15 (12): 107–112. 753–756. Ismay, J.W., 1980a. Recent records of Eribolus (Dipt., Hudec, K., J. Pelikán, V. Pokorny, B. Rychonovsky, V. Chloropidae). – Entomologist’s Monthly Magazine, Skuhravy & M. Skuhravá, 1981. Invertebrates and ver- 115: 96. tebrates attacking common reed stands (Phragmites Ismay, J.W., 1980b. Two brachypterous Chloropidae communis) in Czechoslovakia. 113 pp. Studie ČSAV, (Dipt.) new to Britain. – Entomologist’s Monthly Mag- Praha. azine, 115 (1979): 225–227. Huiskes, A.H.L., 1979. Damage to marram grass Am- Ismay, J.W., 1981a. British Meromyza (Dipt., Chloropidae). mophila arenaria by larvae of Meromyza pratorum – Entomologist’s Monthly Magazine, 116 (1980): 177– (Diptera). – Holarctic Ecology, 2 (3): 182–185. 197. Hwang, Y.-S. & M.S. Mulla, 1973. Attractants for synan- Ismay, J.W., 1981b. Oscinella angustipennis Duda (Dipt., thropic flies. 1. Some physico-chemical factors as re- Chloropidae) in Oxfordshire and Norfolk. – Entomol- latedtothepotencyofHippelates eye gnat attractants. ogist’s Monthly Magazine, 117: 2. – Journal of Economic Entomology, 66: 1055–1058. Ismay, J.W., 1981c. Oscinella maura (Fall.) (Dipt., Chlorop- Hwang, Y.-S., M.S. Mulla & H. Axelrod, 1976. Attractants idae) with pale tibiae. – Entomologist’s Monthly Mag- for synanthropic flies. Identification of attractants azine, 117: 2. and coattractants for Hippelates eye gnats (Diptera, Ismay, J.W., 1981d. Dicraeus raptus (Hal.) (Dipt., Chlorop- Chloropidae). – Journal of Agricultural and Food idae) associated with Bromus ramosus Huds. – Ento- Chemistry, 24: 164–169. mologist’s Monthly Magazine, 117: 34. Ibbotson, A., 1958. The behaviour of frit-fly in Northum- Ismay, J.W., 1985. The identity of Cetema elongata berland. – Annals of Applied Biology, 46: 474–479. (Meigen) (Dipt., Chloropidae). – Entomologist’s Ibbotson, A., 1989. Observations on the oviposition be- Monthly Magazine, 121: 35–38. haviour of frit fly (Oscinella frit L. Dipt., Chloropidae). Ismay, J.W., 1991. Dicraeus tibialis (Macquart) (Diptera: – Entomologia Experimentalis et Applicata, 3 (1): 84– Chloropidae) new to New Zealand. – New Zealand En- 92. tomologist, 14: 1–24. Imhof, G., 1979. Arthropod communities connected with Ismay, J.W., 1993. Rediscovery of Incertellascotica (Collin) Phragmites. – Monographiae Biologicae, 37: 389–397. (Dipt., Chloropidae) in Scotland. – Entomologist’s Imms, A.D., 1930. Observations on some parasites of Os- Monthly Magazine, 129: 49–54. cinella frit Linn. Part 1. – Parasitology, 22 (1): 11–36. Ismay, J.W., 1994a. A revision of the British Neohaplegis Imms, A.D., 1932. Observations on some parasites of Os- Beschovski and Cryptonevra Lioy (Dipt., Chloropidae). cinella frit Linn. Part 2. – Parasitology, 24 (3): 440–447. – Entomologist’s Monthly Magazine, 130: 1–18. International Commission on Zoological Nomenclature, Ismay, J.W., 1994b. Oscinimorpha sordissima (Strobl) 1955. Opinion 348. Supression, under the Plenary Pow- (Diptera: Chloropidae) in Oxfordshire. – Entomolo- ers, of the generic name Titania Meigen, 1800, for gist’s Monthly Magazine, 130: 199. the purpose of validating the generic name Chlorops Ismay, J.W., 1998. Chloropidae. In P. Chandler (ed): Meigen, 1803 (Class Insecta, Order Diptera). – Opin- Checklists of insects of the British Isles (New Series). ions and Declarations rendered by the International Part 1: Diptera. – Handbooks for the Identification of Commission on Zoological Nomenclature, 10: 421– British Insects, 12: 145–149. 436. Ismay, J.W., 1999. The British and Irish genera of International Commission on Zoological Nomencla- Chloropinae (Dipt., Chloropidae). – Entomologist’s ture, 1978. Opinion 1100. Designation of Musca frit Monthly Magazine, 135: 1–37. Linnaeus, 1758, as type-species of Oscinella Becker, Ismay, J.W., 2000. The British species of Lasiambia 1909 (Diptera, Chloropidae). – Bulletin of Zoological Sabrosky (Diptera, Chloropidae). – Dipterists Digest, 7 Nomenclature, 34: 203–204. (2): 59–70.

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264 Meijere, J.C.H. de, 1950. Achtste supplement op de Moore, D., 1983. Hymenopterous parasitoids of stem- Nieuwe Naamlijst van Nederlandsche Diptera van boring Diptera (e.g. Oscinella frit (L.)) in perennial rye- 1898 (Tweede supplement op mijne naamlijst van grass (Lolium perenne) in Britain. – Bulletin of Ento- 1939). – Tijdschrift voor Entomologie, 92 (1949): 1–14. mological Research, 73: 601–608. Mejer, N.F., 1929. [Hymenoptera reared from injurious in- Moore, D., 1984. Factors affecting parasitism of stem- sects in Russia.] – Izvestia po Prikladnoy Entomologii, borer larvae in Italian ryegrass (Lolium multiflorum). 4 (1): 231–248. (In Russian.) – Entomologia Experimentalis et Applicata, 36: 247– Mellini, E., 1954. Pterophorus microdaetylus Hbn. 251. (Lep., Pterophoridae) nella biocenosi di Eupatorium Moore, D., R.O. Clements & M.S. Ribout, 1986. Effects of cannabium. – Bollettino dell’Istituto di Entomologia posture establishment and renovation techniques on dell Università degli Studi di Bologna, 20: 375–387. the Hymenopterous parasitoids of Oscinella frit L. and Mesnil, L., 1931. Récoltes des Muscides nuisibles aux other stem-boring Diptera in ryegrass. – Journal of Ap- céréales. – Revue Pathologie végetale et d’Entomologie plied Ecology, 23 (3): 871–881. agricole de France, 18 (8–9): 266–274. Moore, D. & D.J. Hunt, 1984. Nematodes associated Mesnil, L., 1933. Sur deux Chloropides considérés à with dipterous stem-boring larvae (Chloropidae and tort comme nuisibles. – Revue Pathologie végetale et Opomyzidae) in ryegrass. – Annals of Applied Biology, d’Entomologie agricole de France, 20 (1): 3–7. 104: 291–296. Messersmith, D.H., 1982. A report on a collection Moore, D. & D.J. Hunt, 1987. The role of parasitic nema- of Diptera from Iceland and Greenland. – Fauna todes and hymenopterous parasitoids in causing pre- Norvegica, Serie B, 29 (1): 36–39. mature death of Oscinella spp. larvae. – Annals of Ap- Meyer, R., 1923. Die parasitischen Hymenopteren der plied Biology, 110: 209–211. Fritfliege. – Zeitschrift zur angewandte Entomologie, Moore, D. & M.S. Ridout, 1987. Avoidance of super- 9: 111–120. parasitism of stem-boring larvae by Chasmodon Michailova, P.V. & V.L. Beschovski, 1985. Karyotaxonom- apterus (Hym.: Braconidae). – Entomophaga, 32: ical studies on genus Oscinella (Diptera, Chloropidae). 299–302. – Acta Zoologica Bulgarica, 29: 25–36. Moore, D., M.S. Ridout & R.O. Clements, 1988. Mortal- Mihályi, F., 1965. Rearing flies from faeces and meat ity of Oscinella spp. due to parasitism in insecticide infected under natural condition. – Acta Zoologica treated and untreated ryegrass reseeds. – Journal of Academiae Scientiarum Hungaricae, 11: 153–164. Applied Entomology, 105: 154–159. Milišauskas, Z., 1976. Lietuvos TSR paprastosios egles˙ Moreau, J.P., 1963. Contribution à l’étude éthologiquede (Picea abies Karst.) kankorežiu˙ ˛ entomofauna. – Acta deux espèce d’oscinies, Oscinella frit L. et O. pusilla Entomologica Lituanica, 3: 37–47. (In Lithuanian with Mg. (Diptera, Chloropidae). – Comptes Rendus de English and Russian summaries.) l’Académie de Sciences, Paris, 256: 1831–1833. Mohrig, W., 1967. Beitrag zur Ökologie und Verbreitung Moreau, J.P.,1976. Distribution of adult insects in relation brachypterer Dipteren in norddeutschen Biotopen. – to the host-plants. – Symposia Biologica Hungarica, 16: Deutsche Entomologische Zeitschrift, 14 (1/2): 169– 181–185. 184. Morge, G., 1976a. Dipteren-Farbtafeln nach den bisher Mokrzecki, S.A., 1913. [Report on injurious insects and nich veröffentlichten Original-Handzeichnungen diseases of plants in the government of Taurida during Meigens: “Johann Wilhelm Meigen: Abbildung der the year 1912.] 23 pp. Simferopol, USSR. (In Russian.) europaeischen zweiflügeligen Insecten, nach der Mook, J.H., 1961. Observations on the oviposition be- Natur.” Pars II. – Beiträge zur Entomologie, 26 (1): 441, haviour of Polemon liparae Gir. (Hymenoptera, Bra- 80 pls. conidae). – Arch. Neerl. Zoologie, 14 (3): 423–430. Morge, G., 1976b. Dipteren-Farbtafeln nach den bisher Mook, J.H., 1967. Habitat selection by Lipara lucens nich veröffentlichten Original-Handzeichnungen Mg. (Diptera, Chloropidae) and its survival value. – Meigens: “Johann Wilhelm Meigen: Abbildung der Archives Néerlandaises de Zoologie, 17: 469–549. europaeischen zweiflügeligen Insecten, nach der Mook, J.H., 1971. Influence of environment on some in- Natur.” Pars III. – Beiträge zur Entomologie, 26 (2): 54, sects attacking common reed (Phragmites communis 145 pls. Trin.). – Hidrobiologia, Bucureşti, 12: 305–312 Morge, G., 1984. Diptera collectionis P.Gabriel Strobl – 13. Mook, J.H. & C.G. Bruggemann, 1968. Acustical commu- (Typen-Designierung der Exemplare der sogenannten nication by Lipara lucens (Diptera, Chloropidae). – En- ‘Typensammlung’). – Beiträge zur Entomologie, 34 (2): tomologia Experimentalis et Applicata, 11: 397–402. 319–335.

265 Musolff, E.W., 1959. Die Beobachtung der Verteilung von Nartshuk, E.P., 1972c. [Chloropidae (Diptera) from the Fritfliegen über Getreide unter Verwendung von Farb- Mongolian People’s Republic.] – Nasekomye Mongolii, schalen. – Nachrichtenblatt des Deutsche Pflanzen- Leningrad, 1: 905–932. (In Russian.) schutzdienst, Stuttgart, 11 (11): 164–165. Nartshuk, E.P., 1975. [A convenient method of fluid Myrsina, P.A., M.A. Sybchev, P.S. Pogrebnyak & A.A. mount keeping in collections.] – Entomologicheskoe Svishchuk, 1977. [Physiologically active substances.] – Obozrenie, 54 (3): 676–677. (In Russian.) Naukova Dumka, Kiev, 9: 68–70. (In Russian.) Nartshuk, E.P., 1978a. [Map 18. Range of Diplotoxa mes- Nartshuk, E.P., 1956. [Oscinella-Arten (Diptera, Chlorop- soria Fall.] In K.B. Gorodkov (ed.): Provisional Atlas of idae) des europäischen Teils der UdSSR und ihre the Insects of the European Part of U.S.S.R., 1: Maps 1– Nährpflanzen.] – Entomologicheskoe Obozrenie, 35: 20. (In Russian.) 856–822. (In Russian with German summary.) Nartshuk, E.P., 1978b. [Map 19. Range of Cetema cereris Nartshuk, E.P., 1958. [Two new Palaearctic species of Fall.] In K.B. Gorodkov (ed.): Provisional Atlas of the the family Chloropidae (Diptera).] – Trudy Zoologich- Insects of the European Part of U.S.S.R., 1: Maps 1–20. eskogo Instituta Akademii Nauk SSSR, 24: 99–102. (In (In Russian.) Russian.) Nartshuk, E.P., 1980. Eine neue Gaurax-Art (Diptera, Nartshuk, E.P., 1960. [On the biology of some species of Chloropidae) aus Polen. – Annales Zoologici, the genus Dicraeus (Diptera, Chloropidae).] – Ento- Warszawa, 35: 243–246. mologicheskoe Obozrenie, 39: 585–593. (In Russian Nartshuk, E.P., 1981. [A first record of the chloropid fly with English summary.) Calamoncosis aspistylina Duda (Diptera, Chloropidae) from the USSR and description of its larva.] – Entomo- Nartshuk, E.P., 1962a. [On the fauna and ecology logicheskoe Obozrenie, 60 (2): 422–426. (In Russian.) of Chloropidae (Diptera) of the Leningrad region.] [English translation: Entomological Review, Washing- – Trudy Zoologicheskogo Instituta Akademii Nauk ton, 60: 151–155.] SSSR, 31: 250–275. (In Russian.) Nartshuk, E.P., 1983. [Classification of the superfam- Nartshuk, E.P., 1962b. [A review of the Palaearctic species ily Chloropoidea (Diptera, Cyclorrhapha).] – Entomo- of the genus Calamoncosis End. (Diptera, Chloropi- logicheskoe Obozrenie, 62: 638–648. [English transla- dae).] – Entomologicheskoe Obozrenie, 41: 457–469. tion: Entomological Review, Washington, 62: 180–193.] (In Russian.) Nartshuk, E.P., 1984a. [Flies of the Island of Bol’shoi Nartshuk, E.P., 1964. [A new genus and new species of Berezovyi associated with grasses, with description of Chloropidae (Diptera) from Kazakhstan.] – Trudy Zo- a new species of the family Opomyzidae (Diptera).] ologicheskogo Instituta Akademii Nauk SSSR, 34: 302– – Trudy Zoologicheskogo Instituta Akademii Nauk 324. (In Russian.) SSSR, Leningrad, 123: 51–59. (In Russian.) Nartshuk, E.P., 1967. [Chloropid flies of the genus Di- Nartshuk, E.P., 1984b. Family Chloropidae. In Á. Soós craeus (Diptera, Chloropidae).] – Entomologicheskoe & L. Papp (eds): Catalogue of Palaearctic Diptera. 10 Obozrenie, 46: 415–438. (In Russian.) (Clusiidae–Chloropidae): 222–298. Nartshuk, E.P., 1970. [Revision of type-specimens of Nartshuk, E.P., 1987. [Chloropid flies (Diptera: Chloropidae (Diptera) described by Th. Becker and O. Chloropoidea): their classification, evolution and Duda from the collection of the Zoological Institute, connections with plants.] – Trudy Zoologicheskogo Academy of Sciences of the USSR.] – Entomologich- Instituta Akademii Nauk SSSR, 136: 1–268. (In eskoe Obozrenie, 49: 459–470. (In Russian.) Russian.) Nartshuk, E.P., 1971. Ergebnisse der zoologischen Nartshuk, E.P., 1988. [Chloropid-flies of the genus Forschungen von Dr. Z. Kasab in der Mongolei. 252. Meromyza Mg. (Diptera, Chloropidae) from Finland.] Chloropidae, Part I. Oscinellinae (Diptera). – Annales In V.V. Zlobin (ed.): Svyazi entomofaun severnoi Historico-Naturales Musei Nationalis Hungarici, 63: Evropy i Sibiri. [The connections between entomo- 275–299. fauna of the North Europe and Siberia.] – Leningrad: Nartshuk, E.P., 1972a. [Phytophagy in grass flies (Diptera, 112–117. (In Russian.) Chloropidae) and its evolution.] – Reports from the Nartshuk, E.P., 1990. Structure of male internal reproduc- 23rd meeting to the memory of N.A. Kholodkovsky in tive system of Chloropidae (Diptera). – The Second 1970, Leningrad: 3–49. (In Russian.) International Congress of Dipterology, Bratislava. Ab- Nartshuk, E.P., 1972b. [Predatism and parasitism in the stract Volume: 161. evolution of chloropid flies (Diptera, Chloropidae).] Nartshuk, E.P., 1992. Revision of the species of Meromyza – Zoologichesky Zhurnal, 51: 1342–1352. (In Russian Meigen (Diptera, Chloropidae) from Finland. – Ento- with English summary.) mologica Fennica, 3 (3): 121–138.

266 Nartshuk, E.P., 1994. Revision of the Chloropidae de- Nartshuk, E.P., 2002a. Additions and corrections to scribed by von Roser from Württemberg with lecto- Chloropidae (Diptera) of Poland. – Annales Zoologici, type designations (Diptera). – Zoosystematica Rossica, Warszawa, 52 (2): 319–325. 3: 153–158. Nartshuk, E.P., 2002b. Zoogeographical aspects of the Nartshuk, E.P., 1995. Famiglia Chloropidae. In A. Minelli North European fauna of Chloropinae (Diptera, Cy- et al. (eds): Checklist delle Specie della fauna Italiana, clorrhapha Chloropidae). – Proceedings of the Zoolog- 75: 15–20. ical Institute of Russian Academia of Sciences, S. Pe- Nartshuk, E.P., 1996. The reproductive system of grass- tersburg, Annual Report, 296: 87–94. flies (Diptera, Chloropidae). – The VI International Nartshuk, E.P., 2003. The type specimens of Palaearc- Conference on Regulation of Insect Reproduction, tic Chloropidae (Diptera: Cyclorrhapha) described by Česke Budějovice: 35–36. Van der Wulp and O. Duda in the Zoological Museum Nartshuk, E.P., 1997a. A revision of the Chloropidae in Amsterdam. – International Journal of Dipterologi- (Insecta: Diptera) described by J.W. Meigen from cal Research, 14 (1): 29–33. the Winthem’s collection. – Annalen des Naturhis- Nartshuk, E.P., 2004. Notes on the knowledge of Chlorop- torischen Museums in Wien, 99B: 387–406. idae (Diptera, Muscomorpha) of Spain, including de- Nartshuk, E.P., 1997b. The type specimens of Palaearctic scription of two new species. – Studia Dipterologica, Chloropidae (Insecta: Diptera) in the Natural History 10 (2) (2003): 653–664. Museum in Vienna. – Annalen des Naturhistorischen Nartshuk, E.P., 2006. [Parasites of grass flies (Diptera, Museums in Wien, 99B: 407–416. Chloropidae) from the order Hymenoptera in the Ho- Nartshuk, E.P., 1997c. [Comparative morphology of in- larctic Region.] – Entomologicheskoe Obozrenie, 85 ner reproductive system in Chloropid flies (Diptera, (2): 414–440. (In Russian.) [English translation: Ento- Chloropidae) and perspective of use of the charac- mological Review, Washington, 86 (5): 576–597.] ters of its structure in systematics and phylogenetics.] Nartshuk, E.P., 2006. A revision of the Meigen’s collec- – Entomologicheskoe Obozrenie, 76 (1): 36–50. (In tion of Chloropidae (Diptera) in the Museum National Russian.) [English translation: Entomological Review, d’Histoire Naturelle in Paris (France). – Zoosystemat- Washington, 77 (1): 31–42.] ica Rossica, 5 (1): 173–184. Nartshuk, E.P., 1998. A revision of grassflies of the tribe Nartshuk, E.P., 2008. Gaurax flavomaculatus (Duda, Chloropini (Diptera, Chloropidae) of Finland, Estonia 1933) (Diptera, Chloropidae) found in Norway – with and North-West Russia. – Entomologica Fennica, 9 (3): data on distribution and the mode of life of the species. 153–183. – Norwegian Journal of Entomology, 55: 233–234. Nartshuk, E.P., 1999. Grassflies of the subfamily Nartshuk, E.P. & H. Andersson, 2002. 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Periodicity of outbreaks of preda- nica, 20 (1): 4–8. tory fly Thaumatomyia notata MG. (Diptera, Chloropi- Nartshuk, E.P., E.S. Smirnov & L.I. Feedoseeva, 1970. dae) and its possible reasons. – Entomological Review, [Fam. Chloropidae – grass flies.] In G.Ya. Bei-Bienko Washington, 80: 911–918. (ed.): [Keys to the insects of the European part of the Nartshuk, E.P., 2000c. Outbreaks of carnivorous fly Thau- USSR.] 5 (Diptera and fleas) (2): 399–439. (In Rus- matomyia notata Meigen (Diptera: Chloropidae) and sian.) [English translation: Smithsonian Institution Li- their periodicity. – Trudy Zoologicheskovo Instituta braries & National Science Foundation, Washington, Rossiyskoi Akaddemii Nauk, 286: 93–100. DC, 1988: 667–731.] Nartshuk, E.P., 2001. Oscinella angularis Collin (Diptera: Nartshuk, E.P. & M. von Tschirnhaus, 2012. 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267 Nielsen, L.B., 1985a. Fritfluen – lille flue, stor virkning. – Nowakowski, J.T., 1982. Influence of urban pressure on Naturens Verden, 1985 (1): 24–29. (In Danish.) communities of Diptera-Acalypterata. In:M.Luniak Nielsen, L.B., 1985b. Differential flight behaviour of Os- (ed.): Zwierzata w swiecie zurbanizowanym. [Animals cinella spp. (Diptera, Chloropidae). – Zeitschrift für in the urban environment.] – Ossolineum, Warszawa: angewandte Entomologie, 100 (4): 392–398. 91–102. Nielsen, L.B., 1994. Distribution and population densities Nye, I.W.B., 1958. The external morphology of some of the of stem boring fly larvae (Diptera) and their parasitoids dipterous larvae living in the Gramineae of Britain. – in Danish farmland. – Journal of Applied Entomology, Transaction of the Royal Entomological Society of Lon- 117 (4): 321–331. don, 110 (15): 411–487. Nielsen, L.B. & B.O. Nielsen, 1984a. Oscinella frit (L.) and Nye, I.W.B., 1959. The distribution of shoot-fly larvae O. pusilla (Mg.) 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271 Shapinsky, D.V., 1923. [Biological observations on the frit fly. – Archives of Phytopathology and Plant Protec- Acrididae of the Tcheliabinsk district, Orenburg tion, 29: 111–118. province.] – Izvestia Moskowskogo Entomologich- Skaldere, S.K., 1976a. [Hibernation of Spalangiadrosophi- eskoo Obshchestva, 2: 26–29. (In Russian.) lae – a parasitoid of frit fly in Latvia.] – Puti Vne- Shapiro, I.D., 1961. [Pecularities of interrelations between dreniya Progressivnykh Metodov Zhashchity Rasteniy thefrit-fly(Oscinella frit L.) and injured plants and v Selskokhosaistvennoe Proisvodstvo, Riga: 181–184. their significance for corn resistance to this pest.] – En- (In Russian.) tomologicheskoe Obozrenie, 50 (4): 794–806. (In Rus- Skaldere, S.K., 1976b. [Effectiveness of Spalangia in con- sian.) trol of frit fly on winter rye.] – Puti Vnedreniya Progres- Sharipov, Kh.G., 1981. [Insects damage Phalaroides arun- sivnykh Metodov Zhashchity Rasteniy v Selskokho- dinaceae and Festuca arundinaceae in the Leningrad saistvennoe Proisvodstvo, Riga: 185–189. (In Russian.) district.] – Nauchnye Trudy Leningradskogo Selsko- Skaldere, S.K., 1979. [Spalangia – acclimatized parasitoid khosaistvennogo Instituta, 405: 20–25. (In Russian.) of frit fly in Latvia.] – Puti Dalneishego Sovershen- Shcherbina, V.P., 1970. [Sensory organs of the labella of stvovaniya Zhashchity Rasteniy v Belorussii i Respub- some species of grass flies.] – Nauka-zashchite Ras- likakh Pribaltiki, Minsk, 2: 97–98. (In Russian.) tenii, Voronezh: 32–38. (In Russian.) Skrzypcznska, M., 1978. [A review of the insects found Show, M.W., 1955. Elachiptera cornuta damaging gladioli. in the cones of the European larch Larix decidua Mill. – Plant Pathology, London, 4 (3): 110. and Polish larch L. polonica Rac. in Poland.] – Pol- Siebke, H., 1866a. Entomologisk Reise i Romsdals Amt skie Pismo Entomologiczne, 48 (4): 543–563. (In Pol- i Sommaren 1864. – Nytt Magasin for Naturviden- ish with English summary.) skapene, 14: 375–388. Skrzypczńska, M., 1982. The entomofauna of the cones of Siebke, H., 1866b. Entomologiske undersøgelser, fore- spruce, Picea abies (L.) Karst. in Poland. – Zeitschrift tagne i Sommaren 1865. – Nytt Magasin for Naturvi- für angewandte Entomologie, 92: 21–32. denskapene, 14: 389–420. Skrzypczńska, M., 1984. Insects of cones and seeds of the Siebke, H., 1872. Bidrag til Norges Insektfauna: beretning European larch Larix decidua Mill. and Polish larch, L. om en i Østerdalen foretagen Reise i 1870. 64 pp. J. polonica Rac. in Poland. In H.O. Yates (ed.): Proceed- Dahl, Christiania. ings of the cone and seed insects working party con- Siebke, H., 1877. Enumeratio insectorum norvegicorum. ference. – Ashville, NC: 6–14. Fasculum IV. Catalogum dipterorum continentem. Skrzypczńska, M., 1987. Beitrag zur Kenntnis der Ento- xiv + 255 pp. A.W. Broegger, Christianiæ. mofauna der Fichtenzapfen, Picea abies (L.) in Nor- Sigvald, R., 1978. Inventering over angrepp av fritfluga i wegen. – Anzeiger für Schädlingskunde und Pflanzen- havre 1977. – Växtskyddsrapporter, Uppsala, Jordbruk, Umweltsschutz, 60: 25–27. 52–55: 244. (English summary.) Skuhravý, V., 1980. Diptera attacking common reed in Simmonds, F.J., 1952. Parasites of the frit-fly Oscinella frit Czechoslovakia (Phragmitis communis). – Acta Uni- (L.) in eastern North America. – Bulletin of Entomo- versitatis Carolinae, Biologica, 5–6 (1977): 417–418. logical Research, 43: 503–542. Skuhravý, V. (ed.), 1981. Invertebrates and vertebrates at- Simmonds, F.J., 1953a. Inter-relationship of the parasites tacking common reed stands (Phragmites australis)in of the frit-fly Oscinella frit (L.) in eastern North Amer- Czechoslovakia. – Studie ČSAV, 1: 1–113. ica. – Bulletin of Entomological Research, 44: 387–393. Skuhravý, V. & M. Skuhravá, 1978. Das Schadbild Simmonds, F.J., 1953b. Observations on the biology and und Entwicklung der Art Platycephala planifrons mass-rearing of Spalangia drosophilae Ashm. (Hy- (F.) (Diptera, Chloropidae) an Schilf Phragmites menoptera, Spalangidae), a parasite of the frit-fly Os- communis. – Dipterologica Bohemoslovaca, 1: 311–318. cinella frit (L.). – Bulletin of Entomological Research, Slobodyanyuk, O.V., 1984. [Entomopathogenic nema- 44: 773–778. todes of Diptera. Order Tylenchida.] 198 pp. “Nauka”, Simmonds, F.J., 1954. Host finding and selection by Moscow. (In Russian.) Spalangia drosophilae Ashm. – Bulletin of Entomolog- Smirnov, E.S., 1955. [A new species of Chlorops Mg. ical Research, 45: 527–537. (Diptera).] – Zoologichesky Zhurnal, 34 (2): 334–335. Simmonds, F.J., 1956. Superparasitism by Spalangia (In Russian.) drosophilae Ashm. – Bulletin of Entomological Re- Smirnov, E.S., 1958. [Übersicht der Chlorops-Arten search, 47: 361–376. (Diptera, Chloropidae) aus der Umgegend von Sinelnikov, E.A. & A.E. Tzyplenkov, 1994. Mosaic viruses Moskau.] – Zoologichesky Zhurnal, 37 (8): 1157–1174. influence the susceptibility of barley to net blotch and (In Russian.)

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276 Wheeler, T.A., 1994a. A revision of the genus Epichlorops Zaitzev, V.F., 1992. [Evolutionary pathways of morpho- Becker (Diptera: Chloropidae). – Entomologica Scan- functional adaptations.] 188 pp. Nauka, S. Petersburg. dinavica, 25: 393–414. (In Russian.) Wheeler, T.A., 1994b. Conioscinella zetterstedti Anders- Zamfirov, Z., 1962. [New species of parasites in Bulgaria.] son (Diptera: Chloropidae), a chloropid fly with poly- – Selskostopanska Nauka, 1 (11): 1203–1206. (In Bulgar- morphic wing reduction, new to the Nearctic and Cen- ian.) tral Palaearctic regions. – Canadian Entomologist, 126: Zenker, J.C., 1833. [Eine neue Art Fliegen.] – Notizen 1377–1381. aus dem Gebiete der Natur- und Heilkunde, Erfurt, Wheeler, T.A., 1994c. Sister group relationships and the Weimar, 35: 344. groundplan of the Chloropidae and Milichiidae. – Zerova, M.D., 1977. [New species of the genus Eurytoma Third International Congress of Dipterology, Guelph. Illiger (Hymenoptera, Eurytomidae) from Mongolia Abstracts: 243. and USSR.] – Nasekomye Mongolii, 5: 487–495. (In Wiedemann, C.R.W., 1817. Neue Zweiflügler (Diptera Russian.) Linn.) aus der Gegend um Kiel. – Zoologisches Mag- Zetterstedt, J.W., 1837. Conspectus familiarum, generum azin, 1 (1): 61–86. et specierum Dipterorum, in fauna Insectorum Lap- Wilbur, D.A. & R. Fritz, 1939. Use of shoebox emergence ponica descriptorum. – Isis. Leipzig, 21 (1): 28–67. cages in the collecting of insects inhabiting grasses. – Zetterstedt, J.W., 1838. Sectio tertia. Diptera. In Insecta Journal of Economic Entomology, 32: 571–573. Lapponica (1838–1840). – Lipsiae [= Leipzig]: 477– Wilhelm, H., 1891. Die Haferfliege, Oscinis pusilla,und 868. die Mittel zu ihrer Bekämpfung. 41 pp. Gustav Fock, Zetterstedt, J.W., 1848. Diptera Scandinaviae disposita et Leipzig. descripta. – Lundae [= Lund], 7: 2581–2934. Wilkey, R.E., 1962. A simplified technigue for clearing, Zetterstedt, J.W., 1852. Diptera Scandinaviae disposita et staining and permanently mounting small Arthro- descripta. – Lundae [= Lund], 11: 4091–4545. poda. – Annals of the Entomological Society of Amer- Zetterstedt, J.W., 1855. Diptera Scandinaviae disposita ica, 55 (5): 606. et descripta, seu supplementum tertium, continens Wolcsánszky, E.S., Á. Dely-Draskovits & T. Vásárhelyi, addenda, corrigenda and emendanda tomis undecim 1993. Nádban fejlődő rovarok hatása a nádhajtás prioribus. Lundae [= Lund], 12: 1–xx + 4547–4942. (Phragmites communis) szöveti felépítésére. [The im- pact of insects on the tissues of reed (Phragmites com- Zetterstedt, J.W., 1860. Diptera Scandinaviae disposita et munis).] – Botanikai Közlemények, 80 (2): 141–150. (In descripta, seu ultimus, continens addenda, corrigenda Polish.) and emendanda tomis prioribus, una cum indice al- Wolf, H., 1988. Bewohner von Schilfgallen in den phabetico novarum specierum hujus and praeceden- Naturschutzgebieten “Am Berger Hang” und tis tomi, atque generico omnium tomorum. Lundae = + “Enkheimer Ried” in Frankfurt am Main (Insecta: [ Lund], 14: i–iv 6191–6609. Diptera, Hymenoptera). – Hessische faunistische Zhabinskaja, M.I., 1963. [Larvae of certain species of Briefe, 8 (1): 16–18. the genus Chlorops Mg. (Diptera, Chloropidae) of the Wolf, H., 1991. Bewohner von Schilfgallen in Langen (Hes- European part of the USSR.] – Entomologichskoe sen). – Hessische faunistische Briefe, 11 (2): 31–33. Obozrenie, 42: 924–934. (In Russian.) [English trans- Wright, R., 1966. Primary odours and insect attractation. lation: Entomological Review, Washington, 42: 498– – Canadian Entomologist, 98 (10): 1083–1093. 503.] Yarkulov, F., 1971. [On the biology of the predaceous flies Zhang, Q.-H. & J.R. Aldrich, 2004. Attraction of scaveng- Thaumatomyia sulcifrons Beck. and Th. notata Meig. ing chloropid and milichiid flies (Diptera) to metatho- (Chloropidae, Diptera).] – Zoologichesky Zhurnal, 50: racic scent gland compounds of plant bugs (Het- 1252–1254. (In Russian with English summary.) eroptera: Miridae). – Environmental Entomology, 33 Yakovlev, E.B., 1994. [Palaearctic Diptera associated with (1): 12–20. fungi and myxomycetes.] 127 pp. Petrozavodsk. (In Zimsen, E., 1964. The type-material of I.C. Fabricius. Russian with English summary.) 656 pp. Munksgaard, Copenhagen. Yeats, D.K. & B.M. Wiegmann, 1999. Congruence and Znamenskaya, M.K., 1941. [Pests of cereals and futter controversy: toward a higher-level phylogeny of the grasses in Khibines.] – Vestnik Zashchity Rasteniy, Diptera. – Annual Review of Entomology, 44: 397–428. Leningrad, 1: 42–44. (In Russian.) Zacher, F., 1920. Über einen für Deutschland neuen Ger- Znamenskaya, M.K., 1962. [Review of pests of cultivated stenschädling, Lasiosina cinctipes Meig. – Zeitschrift plants in the Murman district.] – Entomologicheskoe für Angewandte Entomologie, 6: 175. Obozrenie, 41 (2): 310–321. (In Russian.)

277 Zuska, J., 1960. Bemerkungen über einige Paläarctischen glish translation: Entomological Review, Washington, Arten der Gattung Chlorops Meigen und Beschrei- 73: 45–57.] bung einer neuen Art aus Tschechoslowakei (Diptera, Zvereva, E.L., 1993b. Population densities of some flies in Chloropidae). – Časopis Československé Společnosti an air pollution gradient near Monchegorsk. In M.V. Entomologické, 57 (4): 387–397. Kozlov, E. Haukioja & V.T. Yarmishko (eds): Aerial pol- Zvereva, E.L., 1993a. [Effect of industrial pollution on fly lution in Kola Peninsula. – Proceedings of interna- communities (Diptera, Brachycera).] – Entomologich- tional workshop, April 14–16, 1992, St. Petersburg: 371– eskoe Obozrenie, 72 (3): 558–569. (In Russian.) [En- 373.

278 Taxonomic index Includes the main entry for each taxon of Chloropidae treated in the systematic part. Synonyms are in italics

abiskoi Nartshuk & Andersson 79 calva Egger 193 adjunctus Becker 164 Camarota Meigen 154 aenea Meigen 146 capreolus Haliday 126 albinervis Roser 146 cariciphila Collin 127 albipalpis Meigen 110 Centorisoma Becker 155 albipila Loew 196 centromaculatus Duda 165 albisetosa Duda 133 cerealium Reuter 208 angularis Collin 126 cereris Fallén 156 angustipennis Duda 126 Cetema Hendel 155 antennata Collin 110 Chloropinae 154 anthracinum Meigen 141 Chlorops Meigen 159 anthracophagoides Strobl 164 Chloropsina Becker 185 Aphanotrigonum Duda 61 cincta Meigen 151 aprica Meigen 69 cinctellum Zetterstedt 62 approximatonervis Zetterstedt 223 cinctipes auct. 198 Aspistyla Duda 59 citrinellus Zetterstedt 180 aspistylina Duda 69 cognatum Meigen 136 atricillum Zetterstedt 142 Colliniella Nartshuk & Andersson 77 atricorne Zetterstedt 143 Conioscinella Duda 78 consimilis Collin 187 baliola Collin 114 cornuta Fallén 92 balticus Wahlgren 176 coxalis Roser 116 basalis Roser 208 crassipalpis Smirnov 174 beschovskii Dely-Draskovits 66 crassipes Nartshuk 98 bifurcata Nartshuk & Andersson 204 Cryptonevra Lioy 186 bipunctus Duda 180 curvinervis Zetterstedt 205 bjerkanderi Nartshuk & Andersson 165 curvipennis Latreille 154 bohemica Fedoseeva 204 Cyclocercula Beschovski 124 borealis Duda 102 brachypterum Zetterstedt 61 dalmatina Strobl 191 brevibucca Duda 116 danicus Nartshuk & Andersson 96 brevimanus Loew 172 danielssoni Nartshuk & Andersson 197 brevipennis Meigen 92 dasycerus Loew 167 brevisurstylata Dely-Draskovits 197 diadema Meigen 187 britannicus Deeming 104 diastema Collin 94 brunneipenne Beshovski & Lansbury 143 Dicraeus Loew 85 Diplotoxa Loew 190 Calamoncosis Enderlein 68 Diplotoxoides Andersson 190 calceatus Meigen 165 discicornis Zetterstedt 172

279 distinguenda Frey 185 Hapleginella Duda 108 dubius Macquart 102 heleocharis Nartshuk 224 duinensis Strobl 69 herpini Guérin-Méneville 198 hortensis Collin 128 Elachiptera Macquart 90 humeralis Loew 151 elbergi Fedoseeva 206 hungaricum Dely-Draskovits 63 elegantulum Becker 155 hungaricus Becker 97 elongatum Meigen 157 hungaricus Duda 180 ephippium Zetterstedt 102 hypostigma Meigen 171 Epichlorops Becker 192 Eribolus Becker 96 Incertella Sabrosky 109 Eurina Meigen 193 inerme Collin 63 Eutropha Loew 194 ingratus Loew 86 ingrica Nartshuk 206 fasciatus Meigen 167 inquilina Hendel 109 fasciella Zetterstedt 61 intermedia Dely-Draskovits 198 fasciola Meigen 143 interruptus Meigen 171 fascipes Becker 104 femorale Collin 143 karteshensis Nartshuk & Przhiboro 110 femorata Macquart 206 kerteszi Becker 111 femorellum Collin 63 kirigaminensis Kanmiya 171 fennicus Duda 86 kroeberi Duda 123 fennicus Duda 180 figuratus Zetterstedt 167 laetus Meigen 172 finitimus Becker 167 laevicollis Becker 182 flavifrons Duda 83 laevifrons Loew 108 flavitarsis Meigen 188 Lasiambia Anonymous 114 flavomaculatus Duda 104 Lasiambia Enderlein 114 freyi Duda 182 Lasiambia Sabrosky 114 frit Linnaeus 128 Lasiosina Becker 196 frontella Fallén 80 laterale Haliday 200 frontosus Meigen 169 leucarista Nartshuk 106 fulviceps Roser 172 limbatus Meigen 172 fulvifrons Haliday 195 lineella Fallén 152 fungicola Dely-Draskovits 151 Lipara Meigen 118 fungivorus Nartshuk & Andersson 106 livida Nartshuk 81 longicornis Zetterstedt 218 gallarum Duda 81 lucens Meigen 119 Gampsocera Schiner 100 lurida Meigen 193 Gaurax Loew 101 geminatus Meigen 169 macrocerus Nartshuk 106 germanicum Duda 137 Macrostyla Lioy 59 gilvipes Loew 137 maculipennis Zetterstedt 107 glabra Duda 218 maura Fallén 129 glabra Meigen 226 meigenii Loew 174 glyceriae Nartshuk 70 meijerei Duda 78 gotlandica Holmgren 177 Melanochaeta Bezzi 124 gracilior Mejiere 97 Melanum Becker 200 gracilis Meigen 169 Meromyza Meigen 200 messaurea Nartshuk & Andersson 81 hallandica Andersson 227 messoria Zetterstedt 190 halophila Duda 148 Microcercis Beschovski 122 halterata Nartshuk & Andersson 72 microptera Duda 61

280 mimula Collin 83 planifrons Loew 177 minima Strobl 74 Platycephala Fallén 219 minutissima Strobl 135 platythorax Nartshuk 143 mosquensis Fedoseeva 206 plumiger Meigen 60 myopinum Loew 157 pluriseta Péterfi 210 Polyodaspis Duda 138 nana Zetterstedt 97 pratorum Meigen 213 napaeus Collin 86 Pseudogaurax Malloch 139 Nartshukiella Beschovski 149 Pseudopachychaeta Strobl 223 neglectum Tonnoir 157 pubescens Meigen 230 Neohaplegis Beschovski 218 pullitarsis Doskočil & Chvála 121 nidicola Nartshuk 147 pumilionis Bjerkander 177 nigerrima Macquart 129 puncticollis Zetterstedt 192 nigra Meigen 221 pusilla Meigen 131 nigrifrons Duda 112 pygmaeus Meigen 149 nigripalpis Duda 174 nigripes Zetterstedt 63 raptus Haliday 86 nigriseta Fedoseeva 208 Rhaphiopyga Nartshuk 68 nigriventris Macquart 208 Rhodesiellinae 59 nitidissima Meigen 129 Rhopalopterum Duda 141 nitidus Wahlgren 86 ringens Loew 179 norrbottica Nartshuk & Andersson 131 riparius Smirnov 179 norrbotticum Nartshuk & Andersson 65 rohaceki Nartshuk 185 notata Meigen 228 rohdendorfi Fedoseeva 213 numerata Heeger 100 rossicus Smirnov 180 rossicus Stackelberg 88 obliqua Beschovski 157 rostrata Hubicka 208 obscurellus Zetterstedt 174 rufa Macquart 229 Oedesiella Becker 85 ruficeps Macquart 149 oophila Hennig 116 ruficeps Zetterstedt 224 ornata Wiedemann 210 ruficornis Macquart 138 oscinella Becker 75 rufinus Zetterstedt 180 Oscinella Becker 124 rufitarsis Loew 121 Oscinellinae 61 Oscinimorpha Lioy 133 saltatrix Linnaeus 213 oscinina Fallén 145 scalaris Meigen 180 oscinina Fallén 224 scotica Collin 113 Oscinisoma Lioy 135 scrobiculata Strobl 94 scutellaris Zetterstedt 182 Pachychaetina Hendel 124 scutellata Roser 149 Pachychoeta Bezzi 124 serenus Loew 182 pallidiventris Duda 176 sibirica Fedoseeva 215 pallifrons Strobl 176 simile Ismay 157 palpatus Smirnov 176 similis Schiner 118 palposa Fallén 116 Siphonella Macquart 145 palposa Fedoseeva 210 Siphunculina Rondani 146 pannonicus Strobl 176 slesvicensis Becker 98 Parectecephala Becker 218 sordidella Zetterstedt 83 Paroedesiella Anonymous 85 sordidissima Strobl 135 Paroedesiella Enderlein 85 sororcula Fedoseeva 210 Paroedesiella Sabrosky 85 Speccafrons Sabrosky 148 parvipennis Duda 199 speciosus Meigen 182 planifrons Fabricius 221 strigulus Fabricius 182

281 strobilum Karps 104 troglodytes Zetterstedt 184 styriacus Strobl 88 tuberculifera Corti 95 subnigripes Dely-Draskovits 200 suecicus Nartshuk & Andersson 107 umbraculata Fabricius 221 sulcicollis Meigen 139 vagans Meigen 90 tarsata Fallén 218 vallaris Collin 88 tenuiseta Frey 94 variegata Loew 195 Thaumatomyia Zenker 226 variegata Meigen 215 tibialis Macquart 88 varsoviensis Becker 184 tjederi Nartshuk & Andersson 90 vastator Curtis 132 tomentosum Nartshuk & Andersson 143 ventricosi Nartshuk 133 tornensis Nartshuk & Andersson 83 venustus Czerny 139 Trachysiphonella Enderlein 148 triangularis Becker 177 zernyi Duda 185 triangulina Fedoseeva 215 zetterstedti Andersson 84 Trichieurina Duda 230 zimzerla Nartshuk 216 Tricimba Lioy 149 zonulatus auct. 172 trifasciata Zetterstedt 229 zonulatus Wahlgren 167 trigonella Duda 123 zuercheri Duda 113 trilineatum Meigen 65 trochanterata Collin 132

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