no The frit flies (Chloropidae, Diptera) of Fennoscandia and Denmark [43/1, 1 ed.] 9789004167100, 9789004190665, 2013006443

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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: Oscinella 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 fly–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

This publication has been typeset in the multilingual “Brill” typeface. With over 5,100 characters covering Latin, IPA, Greek, and Cyrillic, this typeface is especially suitable for use in the humanities. For more information, please see www.brill.com/brill-typeface. ISSN 0106-8377 ISBN 978-90-04-16710-0 (hardback) ISBN 978-90-04-19066-5 (e-book) Copyright 2013 by Koninklijke Brill NV, Leiden, The Netherlands. Koninklijke Brill NV incorporates the imprints Brill, Global Oriental, Hotei Publishing, IDC Publishers and Martinus Nijhoff Publishers. All rights reserved. No part of this publication may be reproduced, translated, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without prior written permission from the publisher. Authorization to photocopy items for internal or personal use is granted by Koninklijke Brill NV provided that the appropriate fees are paid directly to The Copyright Clearance Center, 222 Rosewood Drive, Suite 910, Danvers, MA 01923, USA. Fees are subject to change. This book is printed on acid-free paper.

Contents Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . List of new taxa, synonyms and nomenclatural changes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Material and acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Review of previous studies of Chloropidae in Fennoscandia and Denmark . . . . . . . . . . . . . . . . . . . . . . . . . Methods of collecting, preparing and preserving . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Morphology, diagnostic characters and anatomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adults . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Preimaginal stages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eggs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Larvae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anatomy of adults and larvae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bionomics and ecology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Habitats and adult habits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Modes of larval feeding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Seasonal occurrence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Role in land ecosystems and species of economic importance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parasitoids, predators and parasites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Distribution and zoogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Monophyly and relationships . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Classification and nomenclature . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Key to subfamilies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Key to Fennoscandian genera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Preliminary key to genera for 3rd instar larvae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Subfamily Rhodesiellinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Aspistyla Duda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Subfamily Oscinellinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Aphanotrigonum Duda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Calamoncosis Enderlein . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Colliniella Nartshuk & Andersson, gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Conioscinella Duda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Dicraeus Loew . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Elachiptera Macquart . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Eribolus Becker . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Gampsocera Schiner . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Gaurax Loew . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Hapleginella Duda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Incertella Sabrosky . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Lasiambia Anonymous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Lipara Meigen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Microcercis Beschovski . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Oscinella Becker . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Oscinimorpha Lioy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Oscinisoma Lioy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Polyodaspis Duda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v

1 1 2 3 4 7 9 9 22 23 24 27 28 28 28 29 32 37 47 49 49 50 52 53 57 59 59 61 61 68 77 78 85 90 96 100 101 108 109 114 118 122 124 133 135 138

Genus Pseudogaurax Malloch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Rhopalopterum Duda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Siphonella Macquart . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Siphunculina Rondani . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Speccafrons Sabrosky . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Trachysiphonella Enderlein . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Tricimba Lioy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Subfamily Chloropinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Camarota Meigen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Centorisoma Becker . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Cetema Hendel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Chlorops Meigen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Chloropsina Becker . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Cryptonevra Lioy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Diplotoxa Loew . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Diplotoxoides Andersson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Epichlorops Becker . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Eurina Meigen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Eutropha Loew . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Lasiosina Becker . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Melanum Becker . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Meromyza Meigen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Neohaplegis Beschovski . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Parectecephala Becker . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Platycephala Fallén . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Pseudopachychaeta Strobl . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Thaumatomyia Zenker . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Trichieurina Duda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Abbreviations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Catalogue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Literature . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Taxonomic index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

vi

139 141 145 146 148 148 149 154 154 155 155 159 185 186 190 190 192 193 194 196 200 200 218 218 219 223 226 230 235 237 247 279

Abstract This volume deals with North European species of the family Chloropidae, known as frit flies or chloropid flies, a large family of acalyptrate flies including some notarial, economically pests of cereals and fodder grasses. The taxonomy, biology and faunistics of all NW European species are revised. Keys are given to subfamilies, genera and species for the adults, for the larvae to generic level as far as possible, and to specific level for a few genera. Brief descriptions of the adult flies are given for all genera and species. Species distributions in Fennoscandia and Denmark and elsewhere are briefly outlined and further tabulated in a catalogue. The known biology and ecology is summarized in general chapters and for each species. Illustrations are given of the male genitalia and also of other characters of diagnostic importance.

From Fennoscandia and Denmark are known 48 genera and 209 species of frit flies: Rhodesiellinae 1 species, Oscinellinae 111 species, Chloropinae 97 species. Among them 119 species are known from Denmark, 97 from Norway, 189 from Sweden, 144 from Finland, and 115 from included provinces of Russia (Karelian Isthmus, Karelia, and the Kola Peninsula). One new genus and 11 new species are described, and some new synonyms and nomenclatural changes are proposed. Some other new species, new synonyms, and overlooked specific names resulting from this revision have been published earlier (Nartshuk, 1992, 1998, 1999, 2002a; Nartshuk & Andersson, 2002; Nartshuk & Przhiboro, 2009; Nartshuk & Tschirnhaus, 2012).

List of new taxa, synonyms and nomenclatural changes New taxa: Colliniella gen. nov.; type species: Conioscinella meijerei Duda, 1933. Aphanotrigonum norrbotticum sp. nov. from North Sweden. Calamoncosis halterata sp. nov. from Sweden. Chlorops bjerkanderi sp. nov. from Sweden. Conioscinella abiskoi sp. nov. from North Sweden. Conioscinella messaurea sp. nov. from North Sweden. Conioscinella tornensis sp. nov. from North Sweden. Dicraeus tjederi sp. nov. from southern Sweden. Eribolus danicus sp. nov. from Denmark and Sweden. Gaurax norvegicus sp. nov. from Norway and Sweden. Gaurax suecicus sp. nov. from Sweden. Lasiosina danielssoni sp. nov. from Gotland, Sweden. Meromyza bifurcata sp. nov. from Sweden.

Rhopalopterum tomentosum sp. nov. from North Sweden. Restored from synonymy: Oscinis ephippium Zetterstedt, 1848. New synonyms: Aphanotrigonum brachypterum (Zetterstedt, 1848) = Aphanotrigonum trilineatum var. micropterum Duda, 1932. Aphanotrigonum cinctellum (Zetterstedt, 1848) = Oscinis fasciella Zetterstedt, 1855. Dicraeus nitidus Wahlgren, 1913 = Dicraeus napaeus Collin, 1946. Eribolus slesvicensis Becker, 1910 = Eribolus crassipes Nartshuk, 1972. Gaurax ephippium (Zetterstedt, 1848) = Gaurax strobilum Karps, 1981. Trachysiphonella rufijiceps (Macquart, 1835) = Chlorops pygmaeus Meigen, 1838 = Oscinis flavella Zetterstedt, 1848 = Siphonella diplotoxoides Strobl, 1893. 1

Aphanotrigonum trilineatum var. micropterum Duda, 1932. Gaurax borealis Duda, 1933.

New combination and status: Microcercis kroeberi (Duda, 1933). Lectotype designations:

Introduction The purpose of the present work is to present a revision of the species of Chloropidae or “frit flies” in a broad sense, also known as chloropid flies or grass flies, occurring in NW Europe with special emphasis on Denmark, Sweden, Norway, Finland and the adjacent Russian provinces of the Fennoscandian Peninsula. It provides keys to subfamilies, genera and species, descriptions of adult and larval morphology, biology, and local and Palearctic distribution. Chloropidae is a large family of small acalyptrate flies usually 2.0–4.0 mm in length, exceptionally 5.0–8.0 mm. They are morphologically and ecologically very diverse. The flies may be found in practically all kinds of terrestrial habitats. Greatest diversity and abundance of species is usually found in open landscapes ranging from sandy coasts, over dry to humid grasslands to bogs, fens, marshes and other types of wetlands. Fewer chloropids are specialized forest inhabitants. The common name of these flies contains in many languages the word “frit”: frit flies, Fritfliegen, fritflugor, fritfluer. This word was used by C. Linnaeus, when he described the first species of the family as Musca frit (now known as Oscinella frit), a serious pest of cereals in many European countries. The very first studies on chloropid flies came from Swedish entomologists: C. Linnaeus (1707– 1778), C. Bjerkander (1735–1795), C.F. Fallén (1764– 1830), J.W. Zetterstedt (1785–1874) and dealt with the local fauna. Their studies proved particularly important because they both contained descriptions of new species and contributed information on the biology and damage to agricultural crops caused by these flies. Many European and Palearctic species of Chloropidae were first described on basis of specimens originating from Sweden. Very many chloropids in Europe have their northernmost distributional limit running through Sweden and the neighboring countries Norway and Finland. Most frit flies, especially those belonging to the subfamily Chloropinae, have phytophagous larvae. In spite of their small size, their abundance gives them an important role in grass ecosystems as 2

first level consumers. Their role as pests of cereals and cultivated grasses should also not be neglected. Increased abundance of certain species of Chloropidae is used as a bioindicator that cereals and grasses are under environmental stress, e.g. from draught or industrial pollution. The studies made in preparation of the present book had some spin-off in terms of descriptions of new species from Fennoscandia and Denmark, new synonymies, finds of overlooked names, and a proposal for conservation of a specific name (Nartshuk, 1992, 1998, 1999, 2002a, 2004; Nartshuk & Andersson, 2002). Still, not all taxonomical problems concerning Scandinavian Chloropidae have been settled with the present book. Especially the taxonomic problems within Oscinella may not be fully elucidated on basis of morphology alone. The morphological terminology follows that proposed in the Manual of Nearctic Diptera (McAlpine, 1981). The treatment of each species begins with the valid name followed by the original combination followed by synonyms established after the Catalogue of Palearctic Diptera, vol. 10 (Soós & Papp (eds), 1984) and listed chronologically with their bibliographic references. Then follows a diagnosis and description to ensure correct differentiation from related species. These are supplemented with illustrations, mainly line drawings, of male genitalia and sometimes other body parts for nearly all the species. A few species cannot be safely identified without examination of the male and female genitalia. Each description is followed by an outline of the known distribution in Denmark and the Fennoscandian countries, and worldwide. Then follows a brief outline of bionomics in terms of phenology, habitat preferences and larval host plants. That piece of information, especially about host plants, is often based on extralimital observations in the absence of local data. A Catalogue giving the known distribution by faunistic province for each species in Denmark and Fennoscandia is provided in the format practiced in previous volumes of the series.

Altogether 209 species of Chloropidae in 48 genera have been found in Fennoscandia and Denmark, but a few more species can be expected. A lot remains however to be found out about the bionomics of Scandinavian frit flies, including such basics as the larval host plants of many phytophagous species. An examination of the arthropod community of a rain forest canopy (Floren, 2003) indicated that the Chloropidae were the most abundant fam-

ily of flies. Similar studies of the canopy fauna of Diptera in Scandinavia has only just started (Thunes et al., 2004). The authors hope that this volume of Fauna Entomologica Scandinavica will make the identification of NW European species of Chloropidae easier and thus promote the general knowledge about this important family of flies.

Material and acknowledgements The present work is based on examination of more than 100 000 specimens of Chloropidae from different collections in Scandinavian countries and material collected by the authors: H. Andersson mainly in Sweden from 1954; E.P. Nartshuk on the Karelian Isthmus (1956–2000) and in Finland (1995). Most material from Sweden is deposited in the Zoological museum of Lund University and consists of two main parts: the collections of C.F. Fallén, J.W. Zetterstedt, E. Wahlgren and O. Ringdahl with numerous type-specimens for their new species, and the material collected by H. Andersson from different parts of Sweden. Material from other Swedish collections were also examined, first of all the collection of the Naturhistoriska Riksmuseet in Stockholm, including old material of C.H. Boheman and P.F. Wahlberg. Further, a collection at the Zoological Museum of Göteborg University and two collections in Uppsala, deposited respectively in the Zoological Museum of the Uppsala University and in the Agriculture University. E.P. Nartshuk visited the Zoological Museum of the Helsinki University four times (1984, 1985, 1990, 2004) to study the large collection of chloropids from Finland, Karelia and the Kola Peninsula, including material of E.J. Bonsdoff, J. Sahlberg, R. Tuomikosky and types of O. Duda, R. Frey and F. Reuter. H. Andersson also borrowed types and interesting species from the Helsinki Museum. The collections of the Zoological Museum (now Natural History Museum of Denmark), University of Copenhagen with material from Denmark, including material of W.H.C.F. Wüstnei, R.C. Stæger, R.W. Schlick, W. Lundbeck, and L. Lyneborg, and also from other Scandinavian countries were examined by both authors (H. Andersson in 1996, E.P. Nartshuk in 2004).

Chloropidae from Norway were borrowed from the Zoological Museum of Oslo University and the Zoological Museum of Bergen University, including the collection of H. Siebke. The Chloropidae collections in the Oslo Museum were also studied by E.P. Nartshuk on a visit in 2008. Material from the Karelian Isthmus, including the collection of A.A. Stackelberg, Karelia and the Kola Peninsula deposited in the Zoological Institute of Russian Academy in St. Petersburg were studied by E.P. Nartshuk as well. Additional material from Finland was received from Dr M. Koponen, from Norway from Dr L. Greve Jensen, and K. Thunes, from Karelia from Dr A. Polevoi and Dr A. Przhiboro, from Sweden from Dr B. Vicklund. All records listed are based on specimens determined by the authors. Distribution records from literature are not used, except the type localities of species described from the territory investigated. For the loan of material and/or for information the authors thank Lita Greve Jensen, Karl Thunes, Pekka Vilkamaa, Bernard Lindeberg, Gunilla Ståhls, Leif Lyneborg, Verner Michelsen, Michael von Tschirnhaus, Alexei Polevoi, Andrei Przhiboro, John W. Ismay, Walter Hackman, Martti Koponen, Bert Vicklund. Hans Silverberg (Helsinki) located many collecting sites and interpreted old spellings of locality names in Finnish material. Our sincere thanks to Thomas Pape and Roy Danielsson for constant support and help, especially during visits of E.P. Nartshuk to Stockholm, Lund and Copenhagen, and to Vladimir I. Gusarov who organized the visit of E.P. Nartshuk to Oslo in 2008 to study Chloropidae collections from Norway. We are indebted to S. Riebe (Botanical Museum of the Lund University) for advice on biotic regions 3

and vegetation zones of northern Fennoscandia, and Anders Peltz (Klassiska och semitiska institutionen, Lund University) for advise on the Latin name of Musca frit. Visits by E.P. Nartshuk to Helsinki (1984, 1990), Stockholm and Lund (1997, 1999, 2002) were supported by the Russian Academy of Sciences and Svenska institutet (2004). Financial support from Russian foundation of Basic Researcn (No. 11-0800186, 13-04-00639). My sincere thanks go to V. Michelsen for his extensive editorial work on my MS. Types of many species were examined by E.P. Nartshuk in Deutsche Entomologische Institut, Eberswalde [now Müncheberg] (1993), Museum für Naturkunde an der Humboldt Universität, Berlin (1993), Staatliches Museum für Naturkunde, Stuttgart (1993), Hope Entomological Collection, University Museum, Oxford (1998), Zoölogisch Museum, Universiteit van Amsterdam (2001), Naturhistorisches Museum, Vienna (1996), Hungarian Natural History Museum, Budapest (1968, 1972, 1976, 1986) and Muséum National d’Histoire Naturelle, Paris by H. Andersson in 1960 and E.P. Nartshuk in 2001. The following abbreviations are used for the location of type specimens and other material mentioned in the text. BMNH – The Natural History Museum, London, England. DECU – Department of Entomology, Charles University, Prague, Czech Republic. DEIM – Deutsche Entomologische Institut, Müncheberg, Germany. HNHM – Hungarian Natural History Museum, Budapest, Hungary. MHNB – Museé d’Histoire Naturelle, Bruxelles, Belgium. MHNL – Musée d’Histoire Naturelle, Lille, France.

MNHN – Muséum d’Histoire Naturelle, Paris, France. NHMH – Natural History Museum, Helsinki, Finland. NHMW – Naturhistorisches Museum, Vienna, Austria. NMBA – Naturhistorisches Museum der Benediktiner-Abbei, Admont, Austria. NMID – National Museum of Ireland, Dublin, Ireland. NMPC – National Museum, Natural History, Prague, Czech Republic. NRMS – Naturhistoriska Riksmuseet, Stockholm, Sweden. OXUM – Hope Entomological Collections, University Museum, Oxford, England. SMNS – Staatliches Museum für Naturkunde, Stuttgart, Germany. ZISP – Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia. ZIWP – Zoological Institute, Polish Academy of Sciences, Warzaw, Poland. ZMAN – Zoologisches Museum, Universiteit van Amsterdam, The Netherlands. ZMBA – Zoological Museum, Bulgarian Academy of Sciences, Sofia, Bulgaria. ZMHU – Museum für Naturkunde an der Humboldt Universität, Berlin, Germany. ZMLU – Zoological Museum, Lund University, Lund, Sweden. ZMMU – Zoological Museum, Moscow University, Moscow, Russia. ZMOC – Zoological Museum, Opava, Czech Republic. ZMUC – Natural History Museum, University of Copenhagen, Denmark. ZMUU – Zoological Museum, Uppsala University, Sweden.

Review of previous studies of Chloropidae in Fennoscandia and Denmark Sweden C. Linnaeus published in 1750 the first data on Scandinavian Chloropidae in a paper on insects injurious to cereals. Under the name “korn-flugan” 4

Linnaeus described a fly which damaged grains of oats in Sweden. He formally described this fly as Musca frit in the 10th edition of his Systema naturae from 1758. The word “frit” is maintained in the vernacular names of the family Chloropidae

in many languages: frit flies, Fritfliegen, fritfluer, fritflugor. Two opinions have been expressed on the origin of this word. One opinion, repeated in many agricultural books, suggests with reference to C. Linnaeus that the word frit originates from Swedish language and means “light grain” (Schaufuss, 1893; Schander & Meyer, 1925). Schaufuss (1893) wrote “Frit bedeutet im Schwedischen ‘Leichte Waare’, der Name weist also auf die von der SommerGeneration angefressenen Körner hin” (p. 82). Indeed grains damaged by larvae of frit flies are unripe and therefore light. The other, better substantiated opinion expressed by Jablonowski (1911) suggests an old Latin origin of this word and refers to M. Terentius Varro. This is supported by the Oxford Latin Dictionary (Glare (ed.), 1968–1982) that gives the following explanation of the word “frit”: “A tiny particle, perh. undeveloped grain, at the top of an ear of corn; (as a type of minuteness) a ‘grain’ ” (p. 736) and cites two works using the word: Mostellaria by T. Maccius Plautus (d. 184 B.C.) and Res Rusticae by M. Terentius Varro (116–27 B.C.). Another species of Chloropidae Musca saltatrix was described by Linnaeus (1761) in Fauna svecica. Bjerkander (1777, 1778a, 1778b, 1779, 1781, 1789) described 3 species with biological observations on another harmful species, now known as Chlorops pumilionis (Bjerkander). The identity of the latter species was discussed by Lampa (1888) and Wahlberg (1918) and Bjerkander’s name of this well known pest of wheat and barley has been placed on the Offfijicial list of specifijic names in zoology (Name No. 868). The first monograph of Swedish Chloropidae was published in 1820 by C.F. Fallén. He listed 20 species in his family Oscinides. Only 16 species belong to Chloropidae in the modern sense, but these include 1 genus and 12 species proposed as new to science. Andersson (1963) revised Fallén’s species with lectotype designations, and 1 genus and 10 species are still valid. One more species of Chloropidae was described by Fallén (1823) in his Phytomyzides. J.W. Zetterstedt dealt with Chloropidae from northern Sweden in his Insecta Lapponica from 1838, and from all parts of Scandinavia in several volumes of his Diptera Scandinaviae (1848, 1852, 1855, 1860). He carried out an enormous amount of faunistic and taxonomic research. All 6 species of Chloropidae described by Zetterstedt (1838) are still valid. 28 species of Chloropidae were described as new in different volumes of Diptera Scandinaviae based on material from Sweden, Nor-

way, Denmark and Finland. Andersson (1966) revised all Zetterstedt’s species with lectotype designations. 15 species described by Zetterstedt are still valid. Only a few new taxa of Chloropidae were described from Sweden after Zetterstedt. Holmgren (1887) described Chlorops taeniopus gotlandica and its bionomics and parasites from Gotland, but this subspecific name has been omitted in most subsequent literature on Chloropidae. Wahlgren (1913) described 3 new species from Öland. Andersson (1966) described two new species in his revision of Zetterstedt’s collections. Records of chloropids new to Sweden and of new data on distribution were published in faunistic papers by Boheman (1852, 1864, 1868), Wallengren (1867), Roth (1896, 1897), Wahlgren (1915) and Poppius et al. (1916). Wahlgren (1919) summed up existing data and gave a review and a key to 19 genera and 65 species of Swedish Chloropidae in Svensk insektfauna. In succeeding years it was first of all O. Ringdahl who in numerous publications (1921, 1931, 1939, 1941, 1947, 1950, 1951, 1954, 1959a, 1959b, 1960) added species and distribution data to Wahlgren’s list of Swedish Chloropidae. Other contributions to the knowledge about Swedish Chloropidae are given in papers by Jansson (1922), Lindroth (1943), Hedström (1964), Lundberg (1964), Andersson & Danielsson (1980) and Nartshuk (2002a). New synonyms based on study of types specimens of Fallén, Zetterstedt and others were published by Nartshuk & Andersson (2002). Hedström (1994) stated that 100 (actually 101) species of Chloropidae were known from Sweden but estimated that the total number of species might rather be 150. The foregoing papers about Chloropidae contains mainly taxonomic and faunistic data with brief notes on habitats and phenology for some species. Ardö (1957) on the contrary carried out a study with emphasis on the ecology of Diptera occurring in marine shore dunes in Sweden, Norway and Denmark. He recognized 49 species of Chloropidae occurring in the dune ecosystems investigated. Ardö specifically studied the influence of some ecological factors on three species of Chloropidae being stenotopic inhabitants of marine shores: Colliniella meijerei, Conioscinella zetterstedti (as C. brachyptera) and Eutropha fulvifrons. Insects, including two chloropid species, associated with the polypore Fomitopsis pinicola across Fennoscandia were studied by Komonen et al. (2004). 5

Chloropid species injurious to cereals and agricultural grasses in Sweden have been most thoroughly studied. Applied aspects connected with Oscinella frit, O. pusilla and Chlorops pumilionis are treated in numerous papers. Linnaeus (1750) and Bjerkander (1777, 1778a, 1778b, 1779, 1781, 1789) were the first to describe the damage to cereals caused by Oscinella frit and Chlorops pumilionis, respectively. Both pest species were considered in Dahlbom’s (1837) review of harmful Scandinavian insects. Numerous papers dealing with these species were published later on, some of which are listed in the chapter ‘Role in land ecosystems and species of economic importance’. The whole complex of harmful species of Chloropidae in Sweden was analyzed by Johansson (1960).

Diptera from marine dune ecosystems in Scandinavia records several species of Chloropidae from Denmark. Lyneborg (1965, 1968) gave lists of Chloropidae from the Hansted Reservation, NW Jutland and, based on identifications by H. Andersson, from the Faroes. The first list includes 32 species in 18 genera and the second 3 species in 3 genera. Petersen (2001) published a check list of Danish Chloropidae that even mentions species expected to be present based on known distributions. Chloropidae as agricultural pests and applied problems with frit fly in Denmark were considered in numerous publications, some of them listed in the chapter ‘Role in land ecosystems and species of economic importance’.

Finland Norway Relatively little information is available on Norwegian Chloropidae. The first records were published by Zetterstedt (1838, 1848). A first list of Norwegian Chloropidae was published by Siebke (1872). Later on Siebke (1877) treated the Chloropidae in his Enumeratio insectorum norvegicum which included 35 species belonging to 15 genera as presently defined. New faunistic data on Norwegian Chloropidae were published later on by Bidenkap (1901) from northern provinces, Strand (1903), Kolstad (1965), Greve (1983, 1999), Skrzypcznska (1987) and Økland & Hågvar (1994). Some species from Norway were also recorded by Ardö (1957) in his research on marine dune ecosystems. Rygg (1966a) studied species of Chloropidae livings as larval pests on cereals and agricultural grasses in Norway and added some species new to the country. The arthropod canopy community of Scots pine (Pinus silvestris) in Norway was studied by Thunes et al. (2004) resulting in 2 chloropid species new to the Norvegian fauna. Nartshuk (2008) published a paper on Gaurax flavomaculatus (Duda) in Norway.

Denmark Chloropidae from Denmark were also recorded for the first time by Zetterstedt (1848) based on material collected by R.C. Stæger. Some meager faunistic records are represented in papers by Kemner (1937), Klefbeck (1951), both investigating the entomofauna of Anholt Is. Ardö’s (1957) study on 6

Some species of Chloropidae from Finland were recorded by Zetterstedt (1848, 1860) based mainly on material of J.E. Bonsdorff. Bonsdorff (1861) listed under “Oscinides” only species of the present genera Platycephala, Meromyza, Oscinis and Madiza. Chydenius & Furuhjelm (1858–1859) recorded some species. Reuter (1902a) described one species of Meromyza. Frey studied Finnish Chloropidae in some detail, resulting in the description of 3 new species (1908a, 1909, 1947), and many new faunistic records (1908a, 1908b, 1944, 1948, 1950). Becker (1910) recorded Diplotoxoides dalmatina from Finland. The rich collection of Finnish Chloropidae was identified by O. Duda and based on his results Frey (1933) published a list of 82 species in 30 genera. All earlier records of Finnish Chloropidae were critically reviewed in this work. Duda further used the material from the Finnish collection in his monograph on Chloropidae in E. Lindner’s Die Fliegen der Paläarktischen Region (1932– 1933). The monograph contains descriptions of 3 new species and several new records of Chloropidae from Finland. A list of Finnish Chloropidae was published again by Frey (1941) in Enumeratio Insectorum Fenniae. New faunistic data were also published by Lindberg & Saris (1952) and Hedström (1964). Some ecological observations together with new faunistic records were done by R. Krogerus (1932, 1960), Kontkanen (1935, 1939), H. Krogerus (1948), Luther (1946), Tiensuu (1948), and Hackman (1963, 1968). Gall producing Lipara and accompanying species were studied by Rable (1951) and Valkeila (1956, 1959a). Haartman (1950) recorded mass occurrences Thaumatomyia notata

indoors. A check list of the insects of Finland, including Chloropidae, was regularly updated by W.E. Hellén from 1926 to 1971. Kallio (1950) studied species of Chloropidae associated with cereals. Vappula (1965), in a monograph on insect pests of cultivated plants in Finland, treated 8 species of Chloropidae that to various extent damage cereals and cultivated grasses in Finland. Hackman (1980) published a check list of Finnish Diptera with Chloropidae represented by 91 species in 36 genera. Nartshuk (1988, 1992) revised the Finnish species of Meromyza and added some species new to the list. Later on she revised the remaining genera of the subfamily Chloropinae from Finland (1998, 1999). A new check list of Finnish Chloropidae by Kaharpää is available from the website http://www.iki.fi/kahanpaa/diptera/list/. Publications on Oscinella frit, Chlorops pumilionis and other harmful species of frit flies in Finland are listed in the chapter ‘Role in land ecosystems and species of economic importance’.

North-West European Russia The Chloropidae of the Russian Karelia, Kola Peninsula and Karelian Isthmus have never been studied specifically. The first records of Chloropidae from the Karelian Isthmus were given by Osten-Sacken

(1858). Chydenius & Furuhjelm (1858–1859) listed 11 species from Karelia. Frey (1933a) published a short list (5 species) found near Sosnovo (Metsäpirtti) on the Karelian Isthmus and in subsequent papers (1933b, 1941) recorded some species from all these territories. Duda (1933) described a new species of Chlorops from Karelia. Chloropidae from the Karelian Isthmus were recorded as a part of studies of the Diptera fauna of the St. Petersburg Province by Stackelberg (1921, 1958, 1965). Additional species and biological data were published by Nartshuk (1962a, 1978a, 1978b, 1992). Nartshuk (1984) studied Chloropidae of the Bol’shoi Berezovyi Is. near Primorsk. Nartshuk & Przhiboro (2009) published some new data on Karelian Chloropidae. Znamenskaya (1941, 1962) recorded Oscinella frit from the Kola Peninsula. Zvereva (1993a, 1993b) and Kozlov & Zvereva (1997) studied the impact of the industrial pollution in the Kola Peninsula on the insect fauna, including Chloropidae. A list of Chloropidae is given in the latter paper, but some records seem doubtful though. Nartshuk & Przhiboro (2009) described a new species of Incertella Sabrosky from the coast of the White Sea in Karelia. As a result of the Scandinavian tradition for taxonomic study of insects, no less than 49 valid species of Chloropidae have their type localities in Fennoscandia.

Methods of collecting, preparing and preserving Most Chloropidae are very small to small flies. The adults of most species are found in the herb stratum, especially among grasses, sedges and reeds. Some species visit flowers, especially Apiaceae. Species of Gaurax, Hapleginella and Aspistyla may be found crawling on foliage of trees or bushes. In general no special techniques are required to collect Chloropidae. Sweeping is a very productive method of Chloropidae collecting. Herbage in meadows, beaches, swamps, fens, bogs and marshes, hedges and clearings in forests, agricultural fields, and also low twigs of trees and shrubs should be examined. The flies are most easily picked up from the net by use of an aspirator. Sweeping of wet vegetation should be avoided. Using a pouter or vacuum sampler is an effective

method to collect small flies living in very low vegetation. Arnold et al. (1973) described a portable vacuum sampler powered by a small two-stroke petrol engine. Malaise traps are probably advantageous over other automated methods of collecting flying insects including Chloropidae by day. Window traps may also be useful, but are less effective. Chloropidae and most other Diptera are not very active at night. Accordingly, light traps normally give poor results. Different kinds of pitfall traps are very effective for wingless or brachypterous chloropids, but even collect fully winged species living near the ground. Pitfall traps are especially recommended to use in habitats with low and dense vegetation. Leponte (1956) collected many 7

Chloropidae from the canopy of trees by using a modified umbrella. Use of brightly coloured pan traps or Moericke traps is a productive way to collect adult Chloropidae (Lauva & Shutele, 1976; Barták, 1997; Kibík et al., 1999). Some authors found that white and yellow traps are the most effective in attracting Oscinella and other chloropids (Koch, 1960; Tešic, 1971; Ibbotson, 1989), others blue (Ibbotson, 1958; Musolff, 1959; Mayer, 1961; De Bruyn, 1986) or violet (Oschmann, 1974, 1979). It is thus possible that different species prefer different colours. Moreau (1963) found that O. frit prefers blue colour and O. pusilla yellow. Tschirnhaus (1981), who used coloured pan traps to collect Chloropidae, considered white pans to be the most efficient. Sticky traps are also used to collect frit flies in agricultural fields. Ibbotson (1958) used panes of glass coated with an adhesive. Kharchenko & Burov (1990) received a good result with coloured sticky traps to study the flight dynamics of flies in corn fields. Nielsen (1985) used blue adhesive traps to study flight behavior of Oscinella spp. Roháček et al. (1998) found that sweeping was the most productive method to collect Chloropidae. They compared sweeping, yellow pan traps and Malaise traps as alternative methods of collecting flies in a peatbog. Comparison of different methods of collecting has been made in more detail for Agromyzidae (Scheirs et al., 1997) than for Chloropidae. It is known that various natural as well as artificial odourous substances attract chloropid flies, but chemical baits are nevertheless rarely used for collecting purposes. Both sexes of species of Oscinella, Conioscinella, Olcella, Hippelates, Siphonella and females of Tricimba are attracted to 2,4hexadienyl butyrate, 2,4-hexadienyl propionate, heptyl and hexenyl butyrate, coproic and valeric acids (Fluno et al., 1972; Rogoff et al., 1973; Aldrich & Barros, 1996; Hibbard et al., 1997; Zwang & Aldrich, 2004) and Hippelates also to trimetil amine ammonia acetic acid (Hwang et al., 1976), oleic and linoleic acids (Hwang & Mulla, 1971). Males of Thaumatomyia glabra are attracted to methyl anthranilate (Landolt et al., 2000; James, 2005). Species belonging to Lasiochaeta, Oscinella, Chlorops and Eutropha came to baits of pyrrolizine alkaloid and ingested these secondary plant chemicals (Boppre & Pitkin, 1988). Only females of Tricimba cincta seem attracted to trans-5 dodecenol (Myrsina et al., 1977; Sybchev & Myrsina, 1977; Verba et al., 1991). Different caproic acids and ethyl caproate attract some Chloropidae (Jantz 8

& Beroza, 1967). Octyl butanoate is attractive to Siphonella sp. (Sugawara & Muto, 1974). Fifteen chloropid species, among them kleptoparasitic flies of the genus Olcella (predominantly females), were found attracted to alkenas: (E)-2-Octenal, (E)-Hexenal and (E)-2-Decenal (Aldrich & Barros, 1995). Adult frit flies are attracted to oils from sprouted seeds of cereals (Kryshtal et al., 1973). Zhang & Aldrich (2004) studied the attraction of female scavenging Chloropidae to hexyl butyrate and (E)-2-hexeny butyrate, both scent gland compounds of plant bugs (Miridae). Their list of species includes species of the genera Conioscinella, Lasiochaeta, Oscinella, Siphonella, and Chlorops, all occurring in Europe. Insecticidal knock-down fogging is now extensively used to collect flies and other insects in the forest canopy. The method proved a fairly effective method for Chloropidae in tropical rainforest (Floren, 2003). Also canopy fogging of pine trees has been practiced with interesting results in Norway (Thunes et al., 2004). We predict that this method is most efficient for species of Gaurax, Hapleginella and Lasiambia. Using cap traps in the canopy of trees in Warsaw parks (Poland) gave a new species of the genus Gaurax (Nartshuk, 1980). Wilbur & Fritz (1939), Southwood & Jepson (1962), Emden (1963) and Clements (1980, 1982) described several methods for collecting invertebrates from grasslands, where most Chloropidae live, including methods for collecting frit fly larvae and rearing adults from grasses. Chloropidae occurring in leaf litter, debris from birds’ nests, mammals’ borrows etc. are preferably extracted by placing the substrates in a box or dark plastic bag allowing exit to a light transparent vessel or bag. Alternative methods of collecting Chloropidae from grasslands, the preferred habitat of most Chloropidae, such as emergence traps and vacuum suckers are described by Clements (1980, 1982). Larvae of many phytophagous chloropids can be found by dissecting stems of grasses, sedges, reeds and other plants. Mature larvae should be allowed to stay within their host plant for pupation. Most flies emerge from puparia in about two weeks, but some species (Lipara, Dicraeus) hibernate and appear only the following spring. Adults of Dicraeus preferably emerge after the puparia have been in contact with water. Some authors (Nye, 1959; Clements, 1982) applied irritants, others (Goodhue & Emden, 1960) X-rays to accelerate the emergence of flies, but these methods appear to be of limited value.

If Chloropidae are collected dry it is preferable to pin them when still soft after having been killed. Double mounting is usually used for small Chloropidae. Taxonomic characters are easier to observe in dry-mounted flies than in fluid-preserved specimens. Flies that have dried out may be glued onto cardboard triangles. Alternatively, dry flies must be softened in a high humidity relaxing chamber before being pinned. Specimens stored in tubes with 80% ethanol must be kept in darkness to prevent bleaching of body colours. Study of the male genitalia is needed for a safe determination of many species of Chloropidae. The tip of the abdomen is removed and put in a 10% solution of potassium hydroxide (KOH) and left there from 1–12 hours until the soft parts have been dissolved. The process may be accelerated to a few minutes by heating the solution in a water bath. After careful rinsing in destilled water to remove all traces of KOH, the genital structures are ready for dissection and examination in alcohol or glycerol. For permanent storage, it is best to place the dissected parts in glycerol in a sealed microtupe or

microvial with a stopper that can be mounted on the same pin as the fly. This method was described by H. Andersson (1972) and Nartshuk (1975). Some light-coloured species of Chloropinae have weakly sclerotized male genitalia, especially the postgonites and aedeagus. Some methods of staining are recommended to better examine transparent parts. Fairly good results have been attained with a slightly modified version of the method described by Wilkey (1962). The cut tip of abdomen is after treatment in 10% KOH solution transferred to Essig’s Fluid (EAF) double stained with lignin pink and acid fuchsine for 1–2 hours and then transferred to 96% alcohol. Stained parts are also stored in glycerol. Chloropid larvae, like those of other Diptera, preserve well in 80% ethanol. They are best killed instantly in hot water beforehand, as this may prevent contraction and retain their natural body shape. The process of making permanent slide preparations of larvae is described by Nartshuk (1956), but today SEM techniques have substantially improved the study of the larvae.

Morphology, diagnostic characters and anatomy Adults Adult chloropids of NW Europe a very small to medium sized (1.0–8.0 mm) “typical” flies. Habitually, here are two main types. One type (Figs 1, 6, 7b), exemplified by Oscinella and Chlorops species, has a compact body with a large, broad head and deep thorax, short legs, and a short, broad and flattened abdomen and rather short, broad wings kept flat over the abdomen at rest. The other type, exemplified by Meromyza and Platycephala (Figs 5, 7a), has a slender body, a long and narrow head, narrow thorax and a narrow, rather cylindrical abdomen. Especially the hind legs have in several groups swollen femora and curved tibiae (Fig. 5). Wingless (apterous) or short-winged (brachypterous) species or forms (Fig. 2) are relatively rare in the North European fauna. The body colour varies from black over yellow, reddish or greenish, often with black or red stripes and marks and is shining or more or less extensively dulled by greyish dusting.

Head (Figs 8–17). The shape of the head is very different between genera. In lateral view the “typical” head is rounded or square and the frons and vibrissal angle project beyond the eyes to the same extent. In many cases, however, the frons or the vibrissal angle (see below) projects far beyond the eyes. In species with a strongly produced frons as in some Chloropinae the head appears rather triangular in lateral view. A strongly produced vibrissal angle is more often seen in species of Oscinellinae. The profile shape of the head is of great diagnostic value in many groups. A detailed study of the head and mouth parts of Chlorops pumilionis was made by Frew (1923b). As is typical for Schizophora the chloropids have a well developed ptilinum. The ptilinal suture is sometimes rather incompletely closed. The frons is broad in both sexes and weakly sclerotized, so that the head of newly emerged specimens tends to become deformed when dry-mounted. The ocellar triangle (frontal triangle) is usually well sclerotized and prominent (Figs 8, 10). Sometimes it is 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: Thaumatomyia 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 triangle are of immense taxonomic value. Normally the ocellar triangle is without setulae except for the ocellar pair, but sometimes there are one or two rows along the side margins, or, rarely, setulae cover most of the surface, arising from small punctures. The setae (bristles) of the head are sometimes quite indistinct. Most of those mentioned below can usually be distinguished though they are often small. The frontal setae (fr) are more or less numerous on the frons. Along the margins of the ocellar triangle and curved up over it are the interfrontal setae (if ). They can also be arranged on the surface along the margins or be dispersed over the surface. Along the eye margins there are one or more recurved (reclinate), outcurved or procurved (proclinate) orbital setae (orb). They cannot as a rule be classified as upper fronto-orbital setae (ors) or lower fronto-orbital setae (ori) and they are here simply called orbital setae (orb). The orbital setae are usually numerous (6–10) and rather small and even in size, but in some genera in both subfamilies one or two (rarely 3) setae are longer than the remaining ones. On the ocellar triangle there are three ocelli on an ocellar tubercle and two proclinate or reclinate, convergent or divergent ocellar setae (oc). The direction of the ocellar setae has great taxonomic importance: in Oscinellinae they are straight or recurved and convergent; in Elachiptera they are more or less procurved but convergent; and in Chloropinae and Rhodesiellinae the ocellar setae are always procurved and divergent. On the vertex there are one outcurved outer vertical seta (vte) and one incurved inner vertical seta (vti) on each side. In the middle, behind the ocelli, is a pair of postocellar setae (poc) (postvertical setae). The postocellar setae are usually convergent or cruciate, except in some genera of Oscinellinae (Lasiambia, Polyodaspis) that have straight postocellar setae. Below the latter there is a well defined, often dark coloured occipital fijield (occ) forming the upper part of the occiput. The compound eyes are large, circular or oval with a vertical, horizontal or diagonal long axis. All facettes (ommatidia) are equal in size or the anterior and anterodorsal facettes are slightly enlarged. The eyes are naked or covered with fine hairs. Below the eyes are the genae (jowls) that vary in width from narrow to broad. The gena bears peristomal setae (pst) and its broader hind part, the postgena, is sometimes delimited by a low carina. The posterior surface of the postgena is sometimes flat and slides 12

against a corresponding surface on the proepisternum (cf. Chlorops pumilionis). The lower anterior corner of the gena forms the vibrissal angle with the vibrissal seta (vi). The vibrissal angle is more or less projecting, right-angled or retreating. The area below the ptilinal suture and delimited laterally by the facial ridges is the face. Between the face and the anterior margin of the eye is the parafacial plate (facial orbit, cheek). Above the insertion of the antennae and separated from frons by the ptilinal suture, is a small triangular sclerite, the lunule, that mid-ventrally continues into the facial carina (facial ridge) between the antennae. The facial carina is usually narrow, but broad in the genus Lipara. The facial margin is the lower limit of the face just above the clypeus. The antennae (Figs 18–24) comprise four parts: The scape (first antennal segment, ant1 ) is in most species very short. The pedicel (second antennal segment, ant2 ) is more variable in length. The fijirst flagellomere (third antennal segment, ant3 ), also known as the postpedicel and though to represent a fusion of several flagellomeres (Stuckenberg, 1999) shows great diversity: rounded, elongate, trapezoid, etc. Finally, the arista is built of three distal flagellomeres, but the basal aristal segment is very short and barely visible. The arista is very variable: slender or broad and flattened, nearly bare, pubescent or densely pubescent. Antennal characters are of great diagnostic value in chloropids, both for genera and for species. Andersson (1979) noticed a distinct correlation in shape between the first flagellomere and the arista in different lineages of the family. The proboscis (Figs 25–30) is usually soft and short with round or oval labella, but rather frequently it is more sclerotized and lengthened, and with long and narrow labella angled backwards. A long proboscis is strongly correlated with a strongly produced vibrissal angle. The structure of the proboscis was studied by Graham-Smith (1930) in some non-Palearctic chloropids and by Kulikova & Nartshuk (1991) in 25 chiefly Palearctic species from 20 genera. The labella have usually three pairs of pseudotracheae, but there are only two pairs in the slender labella of Siphonella oscinina and up to 7 pairs in Lipara. Zaitzev (1992) gave a SEM picture of the labella of Siphonella oscinina showing a rasping devise formed by pseudotracheae. Finer or stronger setae are sometimes standing between the pseudotracheae, e.g. in species of Lasiambia (Tschirnhaus, 1992). Shcherbina (1970) found on the labella of Meromyza saltatrix 86 sup-

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: Centorisoma elegantulum Becker.

posed chemoreceptorial sensilla of two types distinguished by length. Elzinga & Broce (1986) made SEM studies of the labella of several species of Chloropidae and found that the pseudotracheae radiated individually from the prestomal opening at the base of the laella. They also described different types of rasping devices formed by the pseudotracheal rings and concluded that conventional prestomal teeth are not developed. The structure of proboscis indicates that adult chloropids feed on liquefied food. The palpi are of medium size, slightly upcurved, cylindrical or flattened, with short setae. The palpi are sometimes sexually dimorphic, being longer and stouter in the males. Thorax. The thorax (Figs 31–34) is usually short and broad with a rather convex scutum (mesonotum), but sometimes long and slender and flattened dorsally. The scutum is to varying extent shining or mat (if covered in dustlike pruinescence), smooth or sculptured. On the thorax there are also numer-

ous macrochaetae, that may be strong (setae) or fine (setulae). The scutum can be bare or “hairy”, with setulae evenly distributed or arranged in rows. The rows of setae found on scutum are the dorsocentrals (dc) and acrostichals (acr). Usually they are reduced and setulose except for the posterior pair of dorsocentral before scutellum. Other scutal setae are the anterior and posterior postalar setae (a pa and p pa) and prescutellar setae (psc). Species with a better developed scutal chaetotaxy exist in other parts of the world. The scutum is unicolorous, usually black, or striped with light ground colour (yellow, green, brown) and five darker longitudinal stripes (black, brown, red). The middle stripe (M1) usually ends distinctly anterior of the scutellum and can be divided by a narrow streak along the acrostichal line. Laterally of M1 the inner lateral stripes (M2 ) continue from the postpronotal lobes to the dc. They are normally broadest in front. Laterally of M2 and behind the notopleural incisions are the outer lateral stripes (M3 ). In dark coloured 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 confluent. The shape of the scutellum is quite variable: short and semicircular to greatly prolonged and triangular, and dorsally convex or flattened. On the scutellum there are setae along its margin. They can be inserted on small or long, finger-like tubercles. A pair of apical scutellar setae (ap sc) is usually well developed while the lateral scutellar setae (la sc) are normally smaller but consisting of several pairs. The postpronotal lobes (humeral calli) (h) form the anterodorsal corners of the thorax and are usually equipped with a dark postpronotal spot (H) in light coloured (yellow, brown, greenish) species and one (or more) postpronotal setae (h). The notopleura (npl) form triangular depressions at the sides of the scutum. They have an anterior notopleural seta (a npl) and one or two (rarely more) posterior notopleural setae (p npl). A transverse suture is not developed on scutum but indicated by the notopleural incisions. The sides of the thorax are made up by the pleural sclerites. These can be setose or bare, shining or dusted. The proepisternum (prep) (propleuron) in front has one or a few proepisternal setae (prep) below. A vertical proepisternal carina is very characteristic for chloropids but may sometimes be difficult to see without removing the head. Beneath the notopleuron is the anepisternum (anep) (mesopleuron), often with one or two anepisternal spots. In other than completely black species the katepisternum (kep) (sternopleuron) has a large katepisternal spot, of which the colour and dusting are of great importance. The sclerite above the hind coxa is the katepimeron (hypopleuron) and above it and below the wing the anepimeron (pteropleuron). They can both have taxonomically important spots. A precoxal bridge of T-form according to the classification of Speight (1969) is developed between prosternum and the propleura. Below the scutellum the thorax is terminated by the mediotergite (postnotum). Between the postpronotal lobe and anepisternum lies the mesothoracic anterior spiracle (sp1 ), and beneath the halter the posterior spiracle (sp2 ). Wings (Figs 35–44). The costal vein (vein C) continues to the third or fourth vein and has a subcostal break near at the end of the first vein. The costal segments are denoted by mg1 , mg2 etc. At the base of vein C is the basicosta, and before that, nearest at the thorax, a triangular sclerite, the tegula. The subcostal vein (Sc) is retained only for a short distance 16

near the humeral cross-vein (h) or as an indistinct weak fold along the first vein. The first three longitudinal veins compose the radial vein (R1 , R2+3 , R4+5 ). The fourth vein is the medial vein (M). The fifth vein is composed by a cubital vein and the fijirst anal vein (CuA1 ). In the posterior basal part of the wing the veins are reduced or absent. The fijirst or anterior basal cell (br) is closed by the anterior cross-vein (r-m, ta). The second basal cell (bm) is fused with the discal cell (dm) because the crossvein (bm-cu) between them is absent. Its former place can usually be traced by a week flexure in the middle of vein CuA1 . The fused cell is terminated by the posterior cross-vein (dm-cu, tp). The anal veins (A1 , A2 ) are very reduced or absent. In most cases the alula is well developed and there is a small alar calypter. The halteres are of medium size with a triangular knob coloured white, yellow or black. Legs. The front, middle and hind legs are usually simple and relatively short, composed of a coxa (coxa1 –coxa3 ), a trochanter, a femur (f1 –f3 ), a tibia (t1 –t3 ) and a 5-segmented tarsus. The hind femora are in some species thickened with knobs or spines below and correlated with this the hind tibiae are curved. In some species of Eribolus all femora are enlarged, more strongly so in the males. The legs are covered with fine setulae but distinct setae are not present except for a ventral spur on middle tibiae, sometimes even on hind tibiae. In many groups, but most commonly Oscinellinae, a so-called tibial organ (Fig. 45) is developed on the posterodorsal side of the hind tibiae in both sexes in shape of an elongate oval area densely covered with short and fine hairs. Its function is not known. The males of many species of Oscinellinae also posses a femoral organ (Figs 46–61) on middle femora consisting of a group or row of small warts each with a short or spine-like setula. Its function is not known as well. The only species of Rhodesiellinae (Aspistyla plumiger) found in NW Europe possesses in both sexes femoral organs on middle and hind femora (Figs 46, 47). Abdomen (Figs 62, 63). The abdomen is ovoid, oval or slender, cylindrical or flattened. The preabdomen, which can be seen from above, is composed of five segments. The tergites (T) 1 + 2, 3, 4, and 5 are well-developed, their lateral parts are usually turned down and partly covering the ventral side of the abdomen, only leaving narrow membranous strips between their margins and the narrow sternites. Only in Aphanotrigonum the tergites cover most of the ventral abdomen and have distinct

Figs 35–44. Adult frit flies, wing: 35: species of Oscinellinae. 36: Hapleginella laevifrons (Haliday). 37: Dicraeus sp. 38: Gampsocera numerata (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.)

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Figs 45–61. Adult frit flies, leg organs: 45: hind tibial organ of Oscinella frit (Linnaeus). 46, 47: femoral organ of male and female Aspistyla plumiger (Meigen). 48–61: male femoral organ: 48: Aphanotrigonum trilineatum (Meigen). 49: Calamoncosis minima (Strobl). 50: Eribolus nana (Zetterstedt). 51: Dicraeus vagans (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).

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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 Elachiptera brevipennis tergites 1 + 2 are enlarged and occupy nearly half of the abdomen. Sternites 1–5 (S1 –S5 ) are usually small, but sometimes larger or divided. Spiracles 1–5 (sp1 –sp5 ) are placed in the membrane near lateral margins of the tergites. In the male postabdomen there is only one sclerite between tergite 5 and the epandrium. This sclerite is thought to represent the fused sternites 7–8. It bears two approximated spiracles on either side (Fig. 64). In some Chloropinae is this sclerite is very reduced or practically absent (Fig. 65). Eversible membranous vesiculae may be developed

in this part of abdomen in some genera (Lasiambia, Polyodaspis, Thaumatomyia in NW Europe) (Figs 66–70). The vesiculae in Thaumatomyia contain pheromon glands and also constitute a strong optical signal (Kortba, 2009). Hypopygium (Figs 71–82). The epandrium (ep) is considered to represent tergite 9. In caudal view it is horseshoe-shaped, encircling the cerci in most Oscinellinae or the movable forceps or surstyli (sur) in most Chloropinae. In Oscinellinae the movable surstyli are usually simple convex sclerites. They are more complex with processes of different shape 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.)

the surstyli are quite complex, and some groups built up by parts presently named the posterior, median and anterior lobes of the surstyli. In some Chloropinae, e.g. Lasiosina, the surstyli are firmly united with the epandrium and very likely immovable. Sometimes the epandrium have additional processes, e.g. in Cetema. Encircled by the epandrium is the supraanal lobe, the subanal lobe, the anus and the cerci. The two first-mentioned are usually rather indistinct but their presence is indicated by groups of small setae. The cerci can be free or, especially in Chloropinae fused together in a mesolobus (mes). A pair of sclerites is sometimes flanking the subanal lobe, but not in species from our region. The epandrium and its appendages form the caudal wall of the genital capsule. When at rest the epandrium and its appendages move downwards and forwards to cover the genital pouch. The genital pouch is also delimited by the semicircular hypandrium (hyp). Caudally the hypandrium articulates with the epandrium and with ventral bases of the surstyli or a pair of ventral epandrial sclerites (ep s). The hypandrium is in many Oscinellinae closed behind and forming a ring (Figs 74, 75). The inner copulatory organs are the phallus and the gonites. The basal end of the phallus, the basiphallus (bph) articulates with the posterior end of the phallapodeme. Anterior of the phallus the phallapodeme forms a 20

flat or more or less cup-shaped sclerite, the phallapodemic sclerite (ph s) that usually reaches the hypandrium anteriorly. The free end of the phallus is the usually membranous distiphallus (dph). The paired postgonites (pog) and pregonites (prg) are of different shapes and standing in different positions in different genera. They are weakly sclerotized except in most species of Meromyza (Fig. 79). In Oscinellinae and most Chloropinae the pre- and postgonites are arranged in a line, but in most small Oscinellinae the division between these structures is not visible. In some Chloropinae (Chlorops and related genera) the pregonites stand laterad of the postgonites. Rarely in Oscinellinae (Gaurax) the hypandrium and gonites are fused together. In Chloropinae (Diplotoxa) the gonites are seemingly fused together with phallus. The phallus is basally entered by a duct from the ejaculatory bulb with its minute ejaculatory apodeme (ej ap). Female postabdomen (Figs 83, 84). In the female postabdomen segments 6–8 are variously modified to form the ovipositor. Sternites 6–8 tend to be partly membranized and divided in pieces and this sometimes even happens to tergites 6–8. The ovipositor is terminated by the cerci and the supranal and subanal plates. The cerci are usually soft and free but may be sclerotized and laterally compressed or even joined into a flat, blade-like structure.

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.)

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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 The life stages of Chloropidae consist of egg, three larval instars, pupa and imago. Viviparous species are known from the tropics only. The preimaginal stages of chloropids are poorly known compared to the imaginal stage. More or less detailed descriptions of all the immature stages are only available for the economically important species Oscinella frit and Chlorops pumilionis. Publications dealing with the immatures of other species also exist. Some of them only with descriptions, others include keys to genera or to species: Balachowsky & Mesnil (1935), d’Aguillar (1943, 1953), 22

Hennig (1943), Nartshuk (1956, 1962b, 1981, 1987), Nye (1958), Zhabinskaya (1963), Fedoseeva (1966), Chvála et al. (1974), Ismay (1998), Grochowska (1994, 2002, 2006, 2006a, 2007, 2008a, 2008b, 2008c). All these papers include NW European species and may be useful. A general review of the preimaginal stages is given by Ferrar (1987). Unfortunately, it is not possible to give a diagnosis that will separate all larvae of Chloropidae from larvae of all other acalyptrate families. Like other schizophorous larvae chloropid larvae are apodous, acephalic and of milky-white, pale-yellow or greenish colour. In family level keys to Diptera larvae (e.g., Hennig, 1952; Tesky, 1981; Ferrar, 1987;

Figs 89–91. Frit flies, egg and egg structures: 89: Dicraeus fennicus Duda. 90a, 90b: egg outline and surface sculpture of Chlorops planifrons (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 larvae are split into groups according to their biology (plant feeders, predators of root aphids etc.) rather than based on morphological characters.

Eggs (Figs 89–91) Eggs of Chloropidae are insufficiently studied. Descriptions and illustrations mainly exist for a few economically important species based on light microscopic techniques. Most of the data were summarized by Ferrar (1987). More recently Gaponov (2000, 2003) used SEM techniques in studies of the surface structure of eggs from 12 species in 9 genera of Chloropidae. Eggs are elongate oval, cylindrical, straight to slightly curved, tapering at both poles or only towards the anterior pole, where the micropyle is located. The upper side is usually convex, the side facing the substrate flattened or even slightly concave. Some eggs appear asymmetrical (Oscinella). Egg size varies from 0.24–1.7 mm in length and 0.15– 0.39 mm in width. Index (width/length) varies from 0.23–0.63. Carnivorous flies of the genus Thaumatomyia have the shortest eggs: 0.24 mm in T. notate, 0.27 mm in T. glabra. Saprophagous species of the genus Elachiptera have eggs 0.5– 0.8 mm in length, eggs of Oscinella frit 0.6–0.8 mm long. The biggest eggs are known from the gall

producing Lipara species: up to 1.69 mm long in L. lucens (length:width ratio about 5:1); up to 1.26 mm long in L. pullitarsis (length:width ratio 13:1) according to Chvála et al. (1974). Phytophagous species of the subfamily Chloropinae have also rather big eggs: Meromyza 0.75 mm, Cetema 0.80 mm, Chlorops up to 1.5 mm, Lasiosina 0.90 mm, Platycephala 1.4–1.6 mm. The colour of the eggs is usually white or yellowish, but pale brown in some Elachiptera and Thaumatomyia, and blackish in Lipara pullitarsis and L. rufijitarsis. The surface structure of the rigid, protective exochorion is complex as in many other Diptera. It is usually formed by longitudinal parallel, sometimes sinuous ridges and grooves with or without transverse ribs between them. The ridges often anastomose, especially at the anterior third of the egg. The surface between the ridges may be smooth, rough or with low transverse ridges, forming a regular pattern between the longitudinal ones (Cetema, Platycephala). On top of these patterns may be found rows of tubercles (Chlorops fasciatus) or fine pits (Elachiptera tuberculifera, Thaumatomyia glabra). The exochorion of some species has a cellular or polygonal structure (some Chlorops). Sometimes the exochorion is relatively smooth, thin elastic with delicate striae or small pits (Gaurax dubius). Specialized gall23

forming species of the genus Lipara have a very thin and weakly sculptured chorion. The micropyle at the anterior pole of the egg is usually formed on small conical tubercle in the middle of an apical depression (Gaurax, Elachiptera, Conioscinella, Cetema, Chlorops, Meromyza) or on a plug-like protrusion (Oscinella, Lasiosina). The micropylar zone changes in fertilized eggs from conical to rosette-shaped. An aeropylar zone consisting of a system of rather large respiratory crypts has been found in some genera. It usually lies on the anterior egg pole around the micropyle. The number of crypts varies over a wide limit even within a genus. Eggs of Meromyza nigriventris and M. ornata have 5–8 crypts. Eggs of Gaurax has 12–16 crypts not arranged within aeropylar zone. Eggs of Thaumatomyia glabra has 9–12 large crypts with net structure inside. Some additional aeropylar crypts may lie in the rows of the main surface of the egg (Cetema). The apical area with crypts may be separated from the rest of chorion by a circular ridge (Thaumatomyia). No aeropylar zone was found in eggs of Elachiptera, Oscinella, Conioscinella, Lipara and Lasiosina.

Larvae (Figs 92–100, 956–1014) Usually the third instar of larvae have the most fully differentiated species characteristics. Therefore, the following account applies to fully grown final instar larvae. Larval body (Fig. 92) cylindrical, elongated, 2– 10 mm long (biggest in Lipara species) and 0.5– 3.0 mm in diameter. Ratio length:breadth varies normally from 6:1 (in Chlorops species) to 10:1 (in Oscinella, Meromyza and Cetema), only 4:1 in Lipara. Larvae are generally white or yellowish, but greenish in Meromyza. The larval cuticle is normally colourless, except with black pigmentation in both ends of Lipara larvae and anteriorly in larvae of Platycephala planifrons. The larval body is more or less transparent depending on the amount of fat bodies with green or brown contents of the gut and yellow to brown malpighian tubules often well visible through the cuticle. Body usually distinctly tapering anteriorly and bluntly rounded posteriorly, in many species with two caudal processes bearing the posterior spiracles. The larval body of gall producing species as Lipara is more stocky and without caudal processes. The cephalic and following 11 segments are demarcated by slight constrictions of the body. 24

The slightly bilobate cephalic segment (Figs 93, 94) is the smallest segment of the larval body. From it arise interiorly or laterally the short 2-segmented, usually unsclerotized antennae. The apical segment is sometimes completely retracted within the basal antennal segment. Behind the antenna is a sensory organ consisting of several papillae surrounded by a more or less sclerotized ring being completely closed or partly open anteriorly. Behind this sensory organ lies another sensory organ consisting of 1–3 papillae named in some papers as frontal palpi. These two sensory organs, previously identified as maxillary and frontal palpi respectively, are innervated from the maxillary nerve, and both are now named as maxillary papillae. Immediately posterior to the mouth opening is a small sclerite, the labial plate. Ventral surface of the cephalic segment is occupied by the facial mask composed of oral (cephalic) ridges being hyaline cuticular thickenings. The structure of the facial mask varies considerably between genera and species. The oral ridges usually both furcate and make anastomoses and thus develop cephalic cells that may have serrate or dentate margins. Singular cephalic papillae are situated between the oral ridges. The oral ridge are used to channel food into the mouth, but are absent in the carnivorous larvae of Thaumatomyia. The complex cephalopharyngeal skeleton (Figs 95–97) consists of paired sclerites. At the anterior end two black mouth hooks (mandibles) may be exposed from the mouth, while the remaining parts of the cephalopharyngeal skeleton are contained in the first segments of the body. The apical tooth of the mandibles is particularly in phytophagous larvae followed by one to several smaller accessory teeth arranged in one or in two rows. Immediately below the mandibles are a pair of small crescent-shaped dentate sclerites, that in some species (Meromyza, some Lasiosina) are fused together and forming arch. The dentate sclerites are absent in Chlorops. Some additional sclerites are situated near the posterior end of the mandible in some species. The mandibles articulate with the hypopharyngeal (hypostomal, intermediate) sclerites that are joined ventrally by a hypopharyngial arch. In ventral view hypopharyngeal sclerites form a H-shaped sclerite. Hypopharyngeal sclerites articulate or fused with the tentoropharyngeal (pharyngeal) sclerites that at their posterior end are divided in dorsal and ventral cornua (wings). The two ventral cornua are fused on each side with the pharynx. Ventral pharyngeal ridges are present in

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 species (Chlorops, Platycephala). The tentoropharyngeal sclerites are joined anterodorsally by a dorsal bridge (dorsal sclerite, dorsal arch) in most species. A pair of narrow sclerites, the parastomal bars (atrial rods) project from the anterior margin of the tentoropharyngeal sclerites above the hypopharyngeal sclerites. Sometimes (Meromyza species) they are partly fused with hypopharyngeal sclerites. The tentoropharyngeal sclerites are usually less sclerotized than the mandibles and hypopharyngeal sclerites, but the degree of sclerotization varies considerably between species. Mature larvae of Chloropidae are amphipneustic. The anterior spiracles are situated laterally on the first thoracic segment. Most larvae have fan-like anterior spiracles consisting of 5–12 buds (digitate processes) (Figs 98, 99). Fewer larvae have anterior spiracles of a transverse rosette-type with a higher number of buds, e.g. 17 in Platycephala planifrons (Fig. 912). The posterior spiracles on the eighth abdominal segment (Figs 958, 962, 1001) are either sessile or sitting on paired spiracular lobes (tubular mountings, stigmophores), sometimes brown in colour. The shape and distance between the spiracular lobes are of diagnostic value. The posterior spiracles are surrounded by the peritreme and each consists of three elliptic slits set roughly at right angles to each other. The slits sometimes protrude from the spiracular plate (some Thaumatomyia). Button inconspicuous. Between the spiracular slits arise groups of four usually branched interspiracular processes, but these may be unbranched, hairlike or rarely consist of three short spines (Calamoncosis). The interspiracular processes are absent in Camarota curvipennis, some Chlorops and Lipara. The two main dorsal tracheal trunks are sometimes visible through the larval cuticle. Some larvae (e.g. Chlorops planifrons) have the posterior spiracular processes tapering and brownish. In these cases the posterior spiracles are probably non-functional. The larval cuticle has thickenings or spicules of variable size and form (Figs 959, 976, 980). The arrangement and size of the spicules vary between genera and species and are important in larval identification. The three thoracic segments are usually encircled by spicular zones consisting of several irregular rows of spicules or a series of overlapping striae (Cetema species). Spicular zones on the second and/or third thoracic segments are absent in some species. Larvae of Lipara may have a row of small sclerotized papillae dorsally on the 26

posterior margin of the thoracic segments but not ventrally. The eight abdominal segments all have ventral creeping welts that are usually covered with spicules. Some larvae (Meromyza) have no spicules, but many rows of fine striae instead. Some larvae (e.g. Platycephala planifrons) have spicular zones on the dorsal side of the abdominal segments as well, and others (Lipara, Calamoncosis) are without any spicules. The last abdominal segment may be rounded, incised or with more or less prominent processes for the posterior spiracles. On ventral side of the last segment is the anal opening, either surrounded by spicules or with spicules present only in front and behind it. Larvae of the first instar are small, less than 1 mm, and nearly transparent. They are metapneustic, anterior spiracles absent. Posterior spiracles have only two slits and usually with conspicuous interspiracular processes, even if these processes are absent in the mature larvae (e.g. in Lipara). The cephalopharyngeal skeleton is not divided into hypopharyngeal and tentoropharyngeal sclerites, and the mandibles articulate either with long anterior processes of the tentoropharyngeal sclerites or with an accessory pair of small rounded or square sclerites (Fig. 95). Parastomal sclerites are sometimes present, closely appressed to hypostomal sclerites (some Meromyza). The ridges of the facial mask are less numerous and unbranched. Spicular zones on abdominal segments are sometimes better developed, e.g. in larvae of Oscinella. Second instar larvae closely resemble the third instar, but are smaller. They are amphipneustic, but the anterior spiracles are relatively smaller and knoblike with very short buds. The cephalopharyngeal skeleton is also smaller and less sclerotized. Puparia (Fig. 100) are shorter and thicker than the mature larvae, coloured dark or light brown, pale yellow, light greenish or almost transparent. The anterior end is dorsoventrally flattened, the surface often very wrinkled, especially at anterior and posterior ends. The pupal spiracles remain internal and do not break through the puparial wall as respiratory horns. Nearly all larval characters are visible on the puparia except the arrangement of spicules on thoracic segments. Inside the puparium is retained the cephalopharyngeal skeleton and cephalic integument of the last instar larva.

Anatomy of adults and larvae Anatomical studies of both adults and larvae of Chloropidae are insufficient and fragmentary. Adults. Frey (1921) studied the skeleto-muscular anatomy of the head and mouth parts of adult Chlorops speciosus, Platycephala planifrons, Siphonella oscinina and Thaumatomyia sp., and later (Frey, 1923) the skeletal morphology of the head and mouth-parts of Chlorops pumilionis. Kanmiya (1981) illustrated indirect and direct flight muscles of a Lipara sp. in a paper on substrate vibration as a means of communication. Ovtchinnikova (1989) studied the musculature of the male genitalia in Lipara lucens. Inner reproductive organs of both sexes were investigated by Bordon (1971) in Oscinella pusilla, and Nartshuk (1990, 1996, 1997c) studied the male inner reproductive organs in 50 Palearctic species from 21 genera. The male reproductive system consists of two testes, each formed by one follicle as in other Diptera (Figs 85, 86). The testes are brightly coloured yellow, brown or reddish. A small vesicle is found on the anterior end of testes, opposite to the insertion of vas deferens. Both vasa deferentia unite into a vas efferens with a basal swelling named vesicula seminalis. Vas efferens gradually narrows and becomes ductus ejaculatorius. Two long tubular accessory glands insert at the level where the vasa deferentia join. Some differences in the structure of the male reproductive system were found between the different subfamilies and genera of Chloropidae. These concern especially the testes and vasa deferentia that can be referred to different types. Testes: (1) globular in Conioscinella (Oscinellinae) and Neohaplegis (Chloropinae); (2) ovoid, pyriform or conical in most Oscinellinae and Cryptonevra (Chloropinae) (Fig. 85); (3) elongated or nearly tubular in most Chloropinae (Fig. 86). Vasa deferentia: (1) short, entering vesicula seminalis separately in most Oscinellinae and some Chloropinae; (2) long, entering vesicula together in Platycephala and Meromyza (Chloropinae); (3) long, united before reaching vesicula in Trachysiphonella (Oscinellinae) and Neohaplegis (Chloropinae); (4) absent in some Chloropinae, especially in Chloropini. Vesicula seminalis is present in Oscinellinae but absent in most of Chloropinae. The female reproductive organs (Figs 87, 88, 111, 112) consist of paired ovaries composed of several ovarioles. The ovaries, as in all Diptera, be-

long to the polytrophic meroistic type, in which each oocyte is nursed by its own cluster of germ cells. Each ovary has 4–12 ovarioles, the number of ovarioles being correlated with body size. Female Oscinella (1.5–2.5 mm long) have 4–8 ovarioles, female Lipara (6–8 mm long) 10–12 ovarioles. There is usually only one ripe egg in each ovariole, but in Dicraeus usually two. The short oviducts from the ovaries join into an unpaired, rather long oviduct. The oviduct is proximally inserted by a pair of long and thin, sometimes spiraled spermathecal ducts. They arise from two unsclerotized spermathecae. The basal oviduct also has a ventral receptacle and receives a pair of rather long accessory glands. Characters of the male reproductive system may prove useful in taxonomic and phylogenetic studies. It is evident that tubular testes, distal fusion of vas deferens, and absence of vas deferens may be considered as derived states within the family. These characters are at terms with other apomorphic characters of the subfamily Chloropinae and the tribes Chloropini and Meromyzini. Larvae. Larval anatomy is better known than the anatomy of adults. Wandolleck (1899) described the larval anatomy of Platycephala planifrons and Frew (1923a) of Chlorops pumilionis and Meromyza nigriventris. The larval alimentary canal of Oscinella frit was illustrated by Shapiro (1961) and some details by Ryzhkova (1962) and Vilkova (1963). The sclerotized larval mouth parts have been considered above. The tracheal system mainly consists of a pair of dorso-lateral longitudinal trunks uniting the anterior and posterior tracheae. A series of ten dorsal transverse tracheal commissures (only the second looped forwards) connect between the two main longitudinal trunks. The first and tenth commissures are ticker than the others. The first commissure gives off anteriorly directed branches supplying the cephalopharyngeal complex, the remaining commissures are devoid of branches. The main tracheal trunks also give off eleven lateral trunks of which the anterior three are small and supplying the cephalopharyngeal complex. The remaining eight trunks pass downwards to supply the lateral body wall, bifurcate and unite with its posterior branches into a ventro-lateral trunk. The alimentary canal begins with the pharynx situated between the tentoropharyngeal sclerites. The pharynx leads into the narrow, tubular oesophagus that leads into the proventriculus situated in the anterior part of the second abdominal seg27

ment. The proventriculus leads into the midgut that have several pouches (ventricular caeca) at the beginning. The midgut makes some big loopes and eventually becomes very distended by its contents of dark brown colour. There is no peritrophic membrane in the midgut. There are two pairs of malpighian tubules that unite at each side before opening into the gut at the junction of midand hindguts. Shapiro (1961) illustrated the four malpighian tubules in Oscinella frit as equal in length. Frew (1923a) found that the anterior tubules are almost twice as long as the posterior ones in

Chlorops pumilionis. The hind gut is slightly swollen and bulbous, lined with a well marked sclerotized intima. The salivary glands are a pair of simple tubular bodies lying on either side of the gut. The glands unite into a narrow salivary exit duct lined by a sclerotized intima. Nervous system. The brain is situated above the oesophagus in the first abdominal segment. It is connected by broad lateral commissures to a ventral mass of fused thoracic and abdominal ganglia.

Bionomics and ecology Habitats and adult habits Most Chloropidae prefer open habitats such as grasslands and meadows, clearings and edges of forests, and all kinds of wetlands such as marshes, fens and bogs. The adults are mostly seen crawling on stems and leaves of plants. Apterous and shortwinged species uncapable of flight are mostly seen crawling or jumping on or near the ground. Adult chloropids may preferably feed on plant secretions and honeydew on foliage and stems of plants. Some species also take up juice from fruits. Flowers (preferably Apiaceae, Rosaceae and Ranunculaceae) are visited by relatively few species, e.g. some Oscinella, Chlorops and Meromyza. Some Thaumatomyia visit flowers of Fabaceae. Siphunculina aenea is mainly caught on flowers. Kevan & Baker (1983) listed Chloropidae among the Diptera families with species visiting and pollinating flowers. Some authors noted the attraction of newly mowed lawns and greens to adult Oscinella frit. The attraction peaked about 1 hour after mowing according to Falk (1982). A few species have been observed feeding on various decomposing organic substrates in a liquefied state. Some non-Palearctic species feed on wounds and eye secretions of mammals. The chloropid fauna seems most diverse in meadows rich in grasses, sedges and reeds, that constitute the larval host plants for most species. Phragmites beds and moorland fens with Carex and other wetland monocots are important habitats for many species. A few species (e.g. some Rhopalopterum, Pseudopachychaeta rufijiceps) are tyrphobionts, i.e. primarily found in raised peat bogs. Others, such as 28

species of Eutropha and Conioscinella zetterstedti occur exclusively on sandy coastal beaches. Adults of the southern European Trachysiphonella pori are associated with ant nests (Harkness & Ismay, 1976). It remains to be found out, if this also is the case for the two Trachysiphonella species occurring in northern Europe. Relatively few species of Chloropidae from the genera Gaurax, Gampsocera, Hapleginella, Aspistyla, Tricimba, and Lasiambia are forest inhabitants. Some species are known to occur in canopy (Thunes et al., 2004), many others occur on foliage or the trunks of trees closer to the ground. Bährmann (2002) found 13 species of chloropids on trunks of Salix and Populus in Germany, which made the family the most diverse after Phoridae and Hybotidae among 44 families of Diptera Brachycera. Some chloropid flies occasionally gather in huge numbers. Most well known are the huge autumnal aggregations taking place indoors and involving Thaumatomyia notata together with Tricimba humeralis (Kotrba & Nartshuk, 2009). An outdoor aggregation of Thaumatomyia glabra on Wisteria flowers in the USA was described by Sorensen & Sorensen (1997). The non-Palearctic species Apotropina gregalis formed aggregations in rock shelters and domestic dwellings in Namibia (KirkSpriggs et al., 2001).

Modes of larval feeding Larval habits and diets are very diverse in Chloropidae, especially in Oscinellinae. The subject was re-

viewed by Nartshuk (1972a, 1972b, 1987) and Ferrar (1987). Saprophagy, mycetophagy, necrophagy, coprophagy, phytophagy and zoophagy are all feeding behaviours documented from various chloropid larvae. Still, for most species we know practically nothing about the larval habits. Most of the investigated species have phytophagous larvae, especially in the subfamily Chloropinae. These may be classified in four different groups: 1. Inhabitants of shoots or stems of grasses (Poaceae) and sedges (Cyperaceae) usually feeding on shoot and intercalary meristems of their host plants: Oscinella, Conioscinella, Incertella, Rhopalopterum, and most species of Chloropinae (Figs 101, 102). 2. Inhabitants of generative organs of Poaceae and Cyperaceae feeding in particular on the developing seeds: Dicraeus, Pseudopachychaeta (Figs 103, 104). 3. Gall-forming species: Lipara (the well known inducers of cigar galls on Common reed), but even Platycephala, some species of Chlorops (pumilionis, strigulus, riparius) and Meromyza nigriventris produce galls or gall-like swellings on their host plants (Figs 105–107). 4. Inquilines of gall-forming species: Calamoncosis, Cryptonevra, Incertella zuercheri. Larvae of non-phytophagous (saprophagous and carnivorous) species of Chloropidae in northern Europe live in following kinds of substrates: 1. In wood under bark of trees infested by other insects, mainly larvae of xylophagous beetles: some Gaurax and Lasiambia. 2. Cones of coniferous trees infested by other insects: Hapleginella laevifrons, some Gaurax. 3. Mushrooms and bracket fungi: Gampsocera, Tricimba, Aspistyla, some Gaurax. 4. Mammalian excrements: Siphunculina aenea. 5. Shoots of monocots (Poaceae, Cyperaceae and Juncaceae), rarely dicots, attacked by other insects, often phytophagous species of Chloropidae: Aphanotrigonum, Elachiptera, Eribolus, Incertella, Oscinisoma, some Conioscinella, Lasiosina. 6. Egg masses of spiders and short-horned grasshoppers: Pseudogaurax venustus, Lasiambia palposa, L. coxalis, Speccafrons halophila. 7. Plant roots inhabited by root aphids: Thaumatomyia.

8. Two polyphagous species Polyodaspis rufijicornis and Tricimba cincta embrace several of the above feeding modes (in more details under the corresponding species).

Seasonal occurrence Only the phenology of Oscinella frit has been studied in some detail (number of generations, adult activity periods) by several authors in Scandinavia. For the remaining species information on phenology in this book is based on label data in the collections and the study by Nartshuk (1962a) in the Leningrad oblast of Russia with similar environmental and climatic conditions. She recognized 10 groups of Chloropidae with different live cycles and periods of adult activity based on own field observations. Less detailed data could be obtained from the labels of the examined material. Therefore, only 2 groups concerning hibernation and 6 groups concerning flight activity are distinguished here. The family includes univoltine, bivoltine and trivoltine species, but some species can be caught as adults all year without distinct generations. Monophagous species in particular tend to have a short flight period determined by the time when their host plants are optimal targets for oviposition and larval development. Species of Dicraeus developing in seeds of Poaceae and species of Lipara inducing galls in Phragmites australis are univoltine and have very short periods of adult activity. Species of Dicraeus are active when their host grasses are flowering. Phragmites australis, the food plant of Lipara species, is only susceptible to infestation by Lipara over a very short period of time. Some monophagous species of Chlorops (C. pumilionis, C. limbatus, C. interruptus) are bivoltine, but the larvae of different generations feed on different parts of their host plant. Oscinella frit makes 2–3 strongly overlapping generations per season in Sweden (Jonasson, 1975ff.; Lindblad, 1997). The number of generations seems to depend on summer temperature and thus varies from one season to the next. The number of generations per season is impossible to find out for several species with apparently strongly overlapping generations. Two distinct groups of species can be distinguished in relation to hibernation. One group overwinters as adults: some Elachiptera, Conioscinella, Aphanotrigonum, Tricimba, Eribolus, Oscinisoma, some Thaumatomyia, and Pseudopachychaeta. 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: larva of Dicraeus fennicus within seed of Elytrigia repens. 105: gall on Roegneria canina produced by Chlorops riparius Smirnov. 106: gall on common reed (Phragmites australis) produced by Lipara larvae. 107: photos of galls of Lipara spp.

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Table 1. Larval life habits of the chloropid genera occurring in the area investigated. Subfamily/Genus Subfamily Rhodesiellinae: Aspistyla Subfamily Oscinellinae: Aphanotrigonum Calamoncosis

Feeding habit(s)

Place of development

mycetophagous or saprophagous

fungi

saprophagous phytophagous

shoots of Poaceae and Cyperaceae shoots of Poaceae, chloropid galls on Phragmites australis shoots of Poaceae, galls of insects on oak trees seeds of Poaceae shoots of Poaceae and other plants shoots of Cyperaceae Myxomycetes rotten wood, bark of trees and coniferous cones, bracket fungi, galls of Agrobacterum tumefacies damaged coniferous cones shoots of Poaceae, Lipara galls on Phragmites australis rotten wood, sap of wood, egg pods of Orthoptera and Mantodea gallformers on Phragmites australis shoots or seeds of Poaceae and Cyperaceae unknown shoots of Poaceae and Cyperaceae walnuts, stems of damaged plants, head of Asteraceae, leaves rolled by beetles egg sacks of Aranea shoots of Carex and other Cyperaceae ? egg cocoons of Aranea, Coccoidea on grasses mammalian faces, bird nests egg cocons of Aranea unknown fungi, rotten shoots of plants, bird nests, dead mollusks

Conioscinella Dicraeus Elachiptera Eribolus Gampsocera Gaurax

phytophagous and saprophagous phytophagous phytosaprophagous phytosaprophagous mycetophagous saprophagous (xylophagous) and necrophagous

Hapleginella Incertella

saprophagous and necrophagous phytosaprophagous and phytophagous

Lasiambia

saprophagous (xylophagous) and carnivorous

Lipara Oscinella Oscinimorpha Oscinisoma Polyodaspis

phytophagous phytophagous unknown phytosaprophagous polyphagous

Pseudogaurax Rhopalopterum Siphonella Siphunculina Speccafrons Trachysiphonella Tricimba

carnivorous phytophagous or phytosaprophagous ? carnivorous saprophagous or coprophagous carnivorous unknown saprophagous, necrophagous and mycetophagous

Subfamily Chloropinae: Camarota Centorisoma Cetema Chlorops Chloropsina Cryptonevra Diplotoxa Epichlorops Eurina

phytophagous phytophagous phytophagous phytophagous phytophagous or phytosaprophagous phytophagous or phytosaprophagous phytophagous or phytosaprophagous unknown phytophagous

Eutropha Lasiosina Melanum Meromyza Neohaplegis Parectecephala Platycephala Pseudopachychaeta

? saprophagous phytosaprophagous and ? phytophagous unknown phytophagous phytophagous or phytosaprophagous phytophagous phytophagous phytophagous

Thaumatomyia Trichieurina

carnivorous ? phytophagous

shoots of Poaceae shoots of Poaceae shoots of Poaceae shoots of Poaceae and Cyperaceae (Carex spp.) shoots of Poaceae and other plants inquilines in Lipara galls on Phragmites rhizome and stems of Eleocharis spp. unknown gallforming on Phragmites, Scirpus and other Cyperaceae plants and other substrates shoots of Poaceae and Cyperaceae unknown shoots of Poaceae shoots of Cyperaceae (Carex spp.) shoots of Poaceae shoots of Phragmites inflorescence of Cyperaceae (Eriophorum, Eleocharis) rhizome of plants on root aphids shoots of Cyperaceae (Carex spp.)

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species of Oscinellinae belonging to this group have saprophagous larvae developing in rotting shoots of Cyperaceae or Poaceae. Species of Chloropinae in this group have carnivorous (Thaumatomyia notata) or phytophagous (Pseudopachychaeta rufijiceps) larvae. Some species overwintering as adults have been caught during the cold season: Eribolus nana on 10.xi and 25.ii, Oscinisoma cognatum on 1.xii and 6.i. Elachiptera tuberculifera was taken in a light trap in December in Norway (Hågvar & Greve, 2003). Species of Elachiptera, Tricimba and Thaumatomyia notata were found in winter in bird’s nests or between scales of Picea cones. Mass occurrence of Thaumatomyia notata inside houses is a late season event and probably a hibernation activity. Skuhravá (1980) stated that Platycephala planifrons overwinters as adult in the Czech Rep., but in Fennoscandia this species has not been caught after September. The second group of species overwinters as larvae, or sometimes as praepupae or puparia. Most of the phytophagous species belong to this group. Their larvae hibernate in the shoots, seeds or galls of their host plants. It is not known if overwintering 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 that the adults of some species (Dicraeus, Thaumatomyia notata) first emerge from their puparia after these have been moistened by water. It is possible to divide NW European Chloropidae in 6 groups by their flight activity pattern. 1. Species with adult activity during the entire season from April or May to September or October. Many species belong to this group; they hibernate either as adults or larvae. Some have saprophagous larvae developing in rotting plant tissues or phytophagous larvae developing in young shoots: species of Aphanotrigonum, Eribolus, Conioscinella, Oscinella, Elachiptera, Incertella, Rhopalopterum, Cetema, Chlorops, Meromyza, Epichlorops, Diplotoxa. 2. Species with adult activity in spring and early summer (May, June): Lipara lucens, L. pullitarsis, Cryptonevra consimilis, and Calamoncosis aprica, all associated with Phragmites australis, and Dicraeus styriacus, Chlorops frontosus, C. strigulus. All listed species have phytophagous larvae. 3. Species with adult activity in the first half of summer (June, July): Conioscinella gallarum, C. mimula, Dicraeus nitidus, D. tibialis, D. vagans, 32

Eribolus gracilis, E. hungaricus, Trachysiphonella scutellata, Incertella antennata, Polyodaspis sulcicollis, Microcercis trigonella. Most species of Dicraeus, developing in seeds of Poaceae, belong to this group, because this period is time of flowering for most grasses. All listed species are phytophagous. 4. Species with adult activity in late summer (July, August): many species of Chlorops (C. fijinitimus, C. geminatus, C. kirigaminensis, C. rufijinus, C. centromaculatus), Aphanotrigonum cinctellum, Conioscinella sordidella, C. zetterstedti and Apistyla plumiger. 5. Species with adult activity in autumn (August, September): Chlorops ringens, C. varsoviensis, Paractecephala longicornis, Siphonella oscinina. 6. A special phenology is seen in Pseudopachychaeta rufijiceps, a larval feeder on the inflorescences of Eriophorum vaginatum. The species has one generation per year. Newly emerged adults appear in June and stay active to the end of the season. They then hibernate as adults and become active again in April and early May next season, at the flowering time of their host plant.

Role in land ecosystems and species of economic importance Chloropidae constitute a substantial proportion of species richness in grass ecosystems such as meadows, marches, waste lands, pastures, some agricultural fields and so on and play a significant role in the function of these ecosystems in the temperate zone. Recent studies demonstrate that some species are abundant in woodlands as well, and are involved in the destruction of rotten wood, coniferous cones and fungi (Komonen et al., 2004; Thunes et al., 2004). A surprisingly high abundance and species diversity of Chloropidae were found in the canopy of trees in a tropical rain forest in southeastern Asia (Floren, 2003). Larvae of most Chloropidae are either scavengers and phytophagous on species associated with the monocot plants families Poaceae, Cyperaceae, Juncaceae and a few others. The abundance of Chloropidae in grassland habitats gives them an important role as primary consumers of plant tissues and detritus feeders. Few Chloropidae have predaceous larvae. Larvae of Pseudogaurax and Speccafrons and some Lasiambia feed on eggs of spiders and orthopterans. The larvae of Thaumatomyia feed ex-

clusively on aphids living on plant roots. The well known mass occurrences of Thaumatomyia notata have also be observed in Scandinavia: Finland (Haartman, 1950), Sweden (Lampa, 1893; T. Pape, pers. comm.) and suggest that the species helps in the control of subterranean aphids. Chloropidae obviously also play a significant role as a food resource for numerous parasitoid Hymenoptera and some parasitic nematodes and fungi. Adult and larval chloropids are also used as prey by various predatory Hymenoptera and Diptera, and birds such as swallows, swifts and tits. Chloropids make 20% of the food of the Common Swift (Apus apus) during the period of bringing up nestlings according to Parmenter & Owen (1954). Crabronid wasps belonging to Lindenius, Rhopalum and Crossocerus use Chlorops flies as larval provision (Berland, 1925; unpubl. obs.). Blue tits have been observed feeding on Lipara larvae from small galls on Common reed during winter (Tischler, 1943; Mook, 1967). Great Spotted Woodpecker (Dendrocopus major L.) opens large galls of Lipara lucens (Kramer, 1917). Old cigar galls of Lipara are used by some bees and wasps for nesting (Giraud, 1863; Wagner, 1907; Elfing, 1951; Valkeila, 1956; Waitzbauer, 1969; Imhof, 1979; Wolf, 1988, 1991; Westrich, 1989; DelyDraskovits et al., 1994; Anonymous, 2000). Nests of following Hymenoptera were found: Apidae: Hylaeus pectoralis Förster, H. gracilicornis Morawitz, H. moricei (Friese), Megachile centuncularis Linnaeus, M. versicolor Smith, Hoplitis leucomelana (Kirby). Crabronidae: Pemphredon rugifer (Dahlbom), P. lethifer (Shuckard), P. inornata Say, P. wesmaeli (Morawits), Pemphredon sp., Rhopalum clavipes (Linnaeus), Trypoxylon attenuatum F. Smith, T. fijigulus (Linnaeus), Passaloecus gracilis (Curtis), P. clypealis Faeser, P. corniger Shuckard, P. singularis (Dahlbom). Vespidae: Symmorphus bifaciatus (Linnaeus), S. fuscipes (Herrich-Schäffer). Tenthridinidae: Brachythops flavens (Klug). Probably the galls were incidentally used for overwintering. Parasitoids and kleptoparasites of the listed bees and wasps have also been reared from Lipara galls: Pseudomalus auratus (Linnaeus) (Chrysididae), Campodorus variegatus (Jurine) (Ichneumonidae) and Macronychia sp. (Sarcophagidae) inhabited by Pemphredon rugifer and P. wesmaeli; Gasteruption nigrescens Schletterer, G. assectator (Linnaeus), G.

subtile (Thomson), G. jaculator (Linnaeus) (Gasteruptiidae) and Gambrus carnifex (Gravenhorst) (Ichneumonidae) inhabited by Hylaeus pectoralis. The family Chloropidae has a great potential as bioindicator to monitor the condition of environments and ecosystems according to McGeoch (1998): chloropids are abundant and distributed over a wide range of terrestrial habitats; some species are very selective in their habitat requirements; spatial and temporal distributions predictably ensure long term continuity; essential data on biology is available for many species; the species are sampled easily by sweeping of low vegetation and by various traps allowing quantitative results. On top of this adult chloropids are overall easily distinguished from other small acalyptrate flies by structural (large ocellar triangle, weak chaetotaxy) and often colour characteristics (contrasting dark pattern on a yellow or greenish background). Studies in Poland show that the proportion of Chloropidae among terrestrial arthropods in grassland ecosystems increases markedly in habitats disturbed by man compared to natural meadow habitats (Frydlewicz-Ciesielska, 1961; Olechowicz, 1971; Dabrowska-Prot, 1984, 1986, 1987). Nowakowski (1982) found out that 75% of all acalyptrate flies sampled from urban lawns in Poland belonged to Chloropidae. Evidently, some species of Chloropidae even become more abundant in areas subjected to moderate pollution from both sulphur- and fluorine-containing aerial emission (Dabrowska-Prot, 1984; Bährmann, 1985; Zvereva, 1993b; Kozlov & Zvereva, 1997). Zvereva (1993a) underscored this tolerance to pollution in a study on the insect fauna in a strongly disturbed area near the plant “Severonikel” on the Kola Peninsula, where chloropids proved to be remarkably abundant. Kibík (2002) divided species of Chloropidae in three groups according to their ability to cope with anthropogenic deteriorations of ecosystems: (1) anthropo-intolerant species (e.g., Incertella scotica, Pseudopachychaeta rufijiceps); (2) anthroponeutral species (e.g., Aphanotrigonum trilineatum, Chlorops pumilionis); and (3) anthropo-tolerant species (e.g. Elachiptera tuberculifera). Some of the species of Chloropidae have a great economic importance as stem miners or grain feeders of cereals and fodder grasses. In particular the Frit Fly (Oscinella frit) is regarded as a major pest species of cereals. It occurs in all Fennoscandia and Denmark (Znamenskaya, 1962). There are some closely related species (Oscinella pusilla, O. vastator 33

and O. nigerrima) that may also damage cereals, but the taxonomy of these are still not fully clarified. The Frit Fly (Oscinella frit) is a tiny black fly about 1–1.5 mm long with black arista, shining ocellar triangle, clear wings and dusted scutum (Cover illustration). It belongs to a complex of similar species or forms, but the “true” Frit fly is characterized by having the posterior tibiae entirely black or narrowly yellowish at base and tip, front and middle tibiae usually black centrally and more widely yellowish at base and tip, and second aristal segment is about half as long as third aristal segment. The species is polyphagous and injurious to barley, wheat, ray, maize and especially oats, and many fodder grasses. The larvae attack young shoots as well as unripe grains. The species overwinters as larva in seedlings of winter cereals or wild grasses and pupates the following spring. The puparia stay within the shoots. Adults of the first generation emerge from the end of April to early in June. Fertilized females then lay eggs either on the lower part (usually under the ligula) of seedlings of spring cereals or on the ground nearby the host plant. The larvae eat a tunnel inwards and downwards in a spiral manner until they reach the center of the shoot, which results in destruction of the growth zone of the shoot. Usually one larva is found per shoot. Infested plants are easily recognized by the dry central leaf (Fig. 101). Attacked seedlings either die or react by producing new shoots resulting in weakened plants and diminished production of grains. Injury is more pronounced in late-sown fields of spring cereals. Pupation takes place in the injured plants and the adults of the second generation emerge from June to August. Females of the summer generation lay to some extent their eggs on the developing ears of cereals where the larvae feed on the developing grains. In years with a long warm summer adults of a third autumn generation may show up in August to September. Adults of the summer and autumn generations complete the life cycle by laying eggs on seedlings of wild grasses and winter cereals. Larvae of frit flies are involved in the transmission of barley yellow dwarf virus (Jess & Mowat, 1986). Several chloropids, not the least Oscinella frit and allies, are causing “silvertop” injury in fodder grasses. This term describes a condition in which the stem is injured and the seed head dries off prematurely. Other species of the Oscinella frit complex have been reported as cereal pests in NW Europe. One is Oscinella pusilla that has entirely yellow front and middle tibiae, posterior tibiae black centrally with 34

yellow tip and base, and second aristal segment about one-third as long as third segment. Another is O. vastator that only differs from Oscinella pusilla in having the tibiae of front and middle legs never entirely yellow and short brownish wings usually not reaching the tip of the abdomen. Several authors (Nartshuk, 1956; Ibbotson, 1961; Le Berre & Chevin, 1961; Moreau, 1963; Vickerman, 1980; Savage, 1982; Nielsen, 1985) found differences in behaviour and host plant preferences between different species within the frit fly complex. Larvae of O. frit develop in West Europe in young shoots of cereals and some grasses and within unripe grains of cereals, mainly oats, but also wheat and maize. In East Europe O. frit has not been found on unripe grains of cereals except for maize. Nartshuk (1956) found that O. pusilla prefers wheat and barley to other cereals, while O. frit prefers wheat and oats. O. pusilla and O. vastator develop only in young shoots. Moreau (1963) found that O. pusilla prefers barley and wheat to oats and never develops in unripe grains of cereals in France. In respect to fodder grasses, Oscinella frit and O. vastator show the same preference for the annual Lolium multiflorum and perennial L. perenne W Europe, while O. pusilla shows a preference for L. perenne (Jepson & Heard, 1959; Wetzel, 1967; Vickerman, 1978a; Nielsson & Nielsson, 1984; Nielson, 1985). Andersson (1956, 1967) reared O. pusilla from stems of barley and wheat in Sweden, Rygg (1966a) from the same cereals in Norway, but also from rye, timothy and maize. Overall it seems that O. pusilla prefers grasses of the tribe Hordeae that include the cereals wheat and barley, while O. frit prefers grasses of the tribe Avenae that include oats (Nartshuk, 1956; Chevin et al., 1974). Sampling with colour traps indicated that Oscinella frit prefers blue coloured traps and O. pusilla yellow ones. Some behavioural differences were also noted between the three species found most often in agricultural fields. Adults of O. pusilla and O. vastator are more stationary, preferring to live near ground in perennial vegetation. O. frit is a more mobile species and thus more capable of spreading to annual crops (Vickerman, 1980; Nielsen & Nielsen, 1984; Nielsen, 1985). The Frit fly problem has been investigated in details in all Nordic countries: Norway: Schøyen (1895, 1896, 1899, 1918a, 1918b, 1919), Rygg (1966a, 1966b, 1967, 1968), Håbjørg (1979).

Denmark: E. Rostrup (1893), S. Rostrup (1923), Anonymous (1928), Bagger & Thygesen (1971), Jensen & Thygesen (1971), Nødegaard & Hansen (1972), Bagger & Nielsen (1975), Bejer & Esbjerg (1978), Nielsen & Nielsen (1984a, 1984b), Nielsen (1985a, 1985b, 1994). Sweden: Linnaeus (1750), Dahlbom (1837), Hedström (1884, 1892), Post (1884, 1885), Larsson (1885), Lampa (1887, 1888, 1891, 1892), Aurivillius (1892), Trybom (1895), Nilsson-Ehle (1902, 1906), Tedin (1902), Wahlberg (1918), Cunliffe (1929, 1930), Anonymous (1935), Ahlberg (1937), Schwan (1939), Johansson (1943, 1960, 1962, 1975), Ackerman (1951), Andersson & Ossiannilsson (1951), Andersson (1956, 1967), Anerud (1957), Haegermark (1960, 1977), Larsson (1961, 1978, 1979, 1984), Nitsh (1970), Jonasson (1975, 1976, 1977a, 1977b, 1978, 1980, 1982a, 1982b, 1985a, 1986, 1988a, 1988b, 1988c), Swenson (1976), Hadberg (1978), Norlander (1978a, 1978b), Sigvald (1978), Curry, (1986), Lindblad (1988, 1991, 1993, 1997, 1999, 2001), Lindblad & Solbeck (1998), Lindblad & Sigvald (1999). Finland: Reuter (1895, 1896, 1900), Listo (1926b), Pohjakallio (1936), Kallio (1950), Jamailainen & Kanervo (1953), Hardh (1955), Tiittanen (1959a, 1959b), Kanervo (1960a, 1960b), Kanervo & Vappula (1962), Vappula (1965), Ervio (1978), Markkula (1985), Kurppa (1990a, 1990b, 1991). Karelia and the Kola Peninsula: Kanervo (1942), Znamenskaya (1962). Another cereal pest of some importance is the Gout Fly, Chlorops pumilionis. This fly is 3–4 mm long and of yellow colour with scutal stripes and a spot on katepisternum black. The broad ocellar triangle reaches to middle of frons and has a median groove. The triangle is black in dark specimens but with broad yellow margins in pale specimens. Antennae extensively dark coloured. Legs yellow with tarsi and tips of front tibiae darkened. C. pumilionis attacks mainly wheat but also barley and couch grass (Elytrigia repens). The larvae overwinter in a gall formed low down on the stem of its host plant. Pupation takes place within its gall in spring. Grasses and cereals attacked by larvae of the winter generation of C. pumilionis are stunted and do not develop floral parts. Females of the summer generation lay eggs on the upper leaves of their host plant. The larvae feed on developing ears and then move down the culm to pupate within a leaf sheath. Infested plants become stunted with little or no seed development.

C. pumilionis is less important as a pest of cereals in the Nordic countries than Oscinella species, but damage to cereals are reported from Finland (Reuter, 1902b; Listo, 1926a; Kanervo & Vappula, 1962; Vappula, 1965) and Sweden (Bjerkander, 1789; Dahlbom, 1837; Larsson, 1885; Lampa, 1887, 1888, 1891, 1892; Hedström, 1892; Wahlgren, 1918). Meromyza nigriventris has been reported to attack wheat, oats and barley in several Nordic countries, but the species is of minor economic importance. The larvae live under the uppermost leaf sheath of cereals and cause “silvertop” disease (see above). It has been recorded under other names from Finland (Reuter, 1902a, 1902b; Listo, 1926a; Vappula, 1965), Norway (Rygg, 1966a), and Denmark (Nielsen, 1994). Several other species of Chloropidae have sometimes been reared from cereals and fodder grasses in the Scandinavian countries: Incertella albipalpis, Aphanotrigonum trilineatum, Elachiptera cornuta, E. tuberculifera, Tricimba cincta, Eribolus nana, Lasiosina herpini and Cetema elongatum (Johansson, 1942, 1960; Kallio, 1950; Kolstad, 1965; Vappula, 1965; Rygg, 1967; Nielsen, 1994). Larvae of these species are scavengers and secondary invaders of plants with attacks from primary consumers of fresh plant tissues. Their activity period usually starts later than that of the phytophagous Oscinella. One more species, Thaumatomyia notata merits attention. This fly sometimes enters buildings in enormous swarms in the autumn, especially after a warm and dry summer (Figs 108–112) for overwintering. Haartman (1950) described this phenomenon from Turku in 1948. Mass occurrences of T. notata inside buildings were also observed in Finland and Sweden near Stockholm in 1997. The flies cannot, however, hibernate in heated buildings and will then dry out and die within some weeks. T. notata can under these circumstances be very disturbing by smearing ceilings and walls with their excrements. Mass occurrences of this fly are most likely to take place in buildings surrounded by extensive lawns. It has also been reported that T. notata may choose the same building for this behavior several times in succession. The existing literature on this aggregation phenomenon is extensive. A selection of the more important ones is as follows: Waga (1848), Kiesenwetter (1857), Weyenberg (1871), Letzner (1873), Reh (1928), Hase (1929), Roszypal (1930), Uhlmann (1930), Lindner (1931), Tzygankov (1931), Zuska (1966), Weidner (1990), Nartshuk (2000b, 2000c) and Kotrba & Nartshuk (2009). Specimens of T. notata from Waga’s obser35

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

vations in Poland in 1848 are still keept in NHMW (unpubl. obs.). The earliest notes on these aggregations of flies date back to 1736, 1766 and 1805 in England, 1807 in Germany, and 1812 in Poland, but without specifying their identity with a scientific name. Jenyns (1832) described the phenomenon in England and for the first time proposed an identity of the fly as Chlorops laeta. Subsequent authors agreed on the family assignment of the fly involved in mass occurrences, but with absolute no concensus on its specific identity. In various combinations with the generic names Chlorops, Chloropisca and Thaumatomyia the species has been recorded by the following names: lineata Fabricius, nasuta Schrank, didyma Zetterstedt, obscurella Zetterstedt, minuta Zetterstedt, prodigiosa Zenker, copiosa Schiner, laeta Meigen, hypostigma Meigen, circumdata Meigen, ornata Meigen, notata Meigen, nasuta Meigen, flavifrons Meigen, and taeniopus Meigen. Of these names prodigiosa Zenker, circumdata Meigen, ornata Meigen and copiosa Schiner are now regarded as synonyms of notata Meigen, the other ones are based on misidentification. The behavior of the present species is alluded to in the names prodigiosa and copiosa and the generic name Thaumatomyia means “the mysterious fly”. An analysis of the literature data on mass occurrences of T. notata in Europe for more than two centuries revealed that these occur at intervals of 10–12 years. A hypotheses proposed by Nartshuk (2000b, 2000c) suggests that the aggregations behavior of T. notata coincide with the 11-year cycle of the sun activity with its documented influence on the weather conditions.

Parasitoids, predators and parasites Parasites and parasitoids of the Chloropidae are known in more detail for the agriculturally important species of the Oscinella frit species complex and Chlorops pumilionis. Several authors report on parasitoids reared from frit flies in Scandinavia (Lampa, 1888; Henriksen, 1918–1919; Gradwell, 1957; Nordlander, 1978a, 1978b; Nielsen, 1994), other parts of Europe (Kurdyumov, 1912; Meyer, 1923; Imms, 1930, 1932; Riggert, 1935; Nikolskaya, 1937; Dyurich, 1980; Moore, 1983) and in North America (Simmonds, 1952, 1953a, 1953b, 1954, 1956). Species of the O. frit complex tend to be strongly parasitized. Moore (1983) found that near-

ly 50% of a sample of larvae of O. frit and O. vastator was infested by parasitoids. Also the parasitoids of Lipara and other chloropid species living as inquilines in Lipara galls on Common reed (Phragmites australis) are quite well investigated: Giraud (1863), Blair (1932, 1934), Erdös (1952, 1955, 1957, 1961), Valkeila (1959a, 1959b), Mook (1961, 1967), De Bruyn (1987), Grochowska (1989), Gromysz-Kałkowska & Grochowska (1992) and Dely-Draskovits et al. (1993, 1994). Records of parasitoids of other Chloropidae are rare (Nartshuk, 1962a; Kiauka & Nartshuk, 1972; Dyurich & Kostyukov, 1978; Dyurich, 1980; Norlander & Grijpma, 1991). The reported parasitoids of Chloropidae are mainly reared from the puparia or from stems or inflorescences containing chloropid larvae and/or puparia. Such plant parts, especially chloropid galls, may easily contain other insects and their parasitoids. This aspect, as well as the possible occurrence of hyperparasitoids, suggests that the existing host records for parasitoids are frequently based on mistakes. Most parasitoids of Chloropidae are found among Hymenoptera (Table 2). Their development takes place in the chloropid larvae followed by their emergence from the puparia. It has been observed that the braconid wasp Polemochartus liparae oviposits in the eggs of Lipara found exposed on shoots of Phragmites, but that development actually takes place in the larvae (Mook, 1961). Larvae of Lipara lucens parasitized by P. liparae pupate in September instead of spring in the following year as they normally do (Varley & Butler, 1933; Chvála et al., 1974). Stenomalina liparae (Pteromalidae) parasitizes the young larvae of Lipara (Chvála et al., 1974; Athen & Tscharntke, 1999). The wingless braconid Chasmopodon apterus, the pteromalid Trichomalus statutus and a figitid wasp (Ganaspis sp.) lay their eggs in young larvae of their hosts, while the pteromalid Spalangia fuscipes and the eulophid Pronotalia liparae oviposit in the host puparia. Trichomalus cristatus lays several eggs in one larva of Oscinella, but only one parasitoid will survive. Most parasitoids are solitary, but examples of gregarious species are: Tetrastichus legionaries with up to 40 specimens emerging from a single puparium of Lipara, Clytina giraudi with 3–15 specimens from one puparium of a Cryptonevra or Calamoncosis sp., Crataepiella platycephalae with 18–55 specimens from single puparia of Platycephala umbraculata, and Pronotalia liparae and Tetrastichus djuritshae with 44–56 specimens from puparia of Lipara lu37

cens. Aggregations of Oscinella larvae and their hymenopterous parasitoids were observed in grasslands in England and that the mortality among overwintering frit fly larvae differed between parasitized and unparasitized ones (Umoru, 1993c, 1994). Nordlander (1978a) noted that puparia of Oscinella from shoots of cereals were parasitized up to 50% in Sweden, but puparia in grains of cereals were practically all unparasitized. Eggs parasitoids of Chloropidae are not known. Some testing of the potential of Trichogramma evanescens (Trichogrammatidae) as a control agent against Oscinella frit in Russia have been undertaken in the laboratory and in field (Chernoponevsky, 1939). This egg parasitoid did infest eggs of the frit fly in the experiments, but the development of parasitoids took 28 days instead 9 days in eggs of Lepidoptera. Spalangia drosophilae (Pteromalidae) is reported as a parasitoid of Oscinella frit in North America (Simmonds, 1952, 1953a, 1953b, 1954, 1956). This species has never been reared from frit flies in Europe, but some research was undertaken to find out the potential of this species as a control agent of O. frit in NW Russia and Latvia (Kovalchuk, 1968, 1972, 1977; Skaldere, 1976a, 1976b, 1979). It was shown later on that S. drosophilae in Europe primarily infests larvae of Sphaeroceridae and Drosophilidae living in dung, and only accidentally uses O. frit as a host (Marshakov, 1983). Marshakov (1984) presented a review of the genuine, casual and fake parasitoids of frit flies. Cleigastra apicalis (Scathophagidae) is a well known predator of Lipara larvae (Wagner, 1907; Theowald, 1961; Chvála et al., 1974; Askew & Shaw, 1979; De Bruyn, 1985, 1987). Other predators of Lipara larvae are birds (see chapter ‘Role in land ecosystems and species of economic importance’). Coenosia tigrina (Muscidae) is a predator of stemboring flies, including Oscinella frit and other Chloropidae (Parmenter, 1968; Kuehne, 1991). Aelothrips fasciatus, a species of thrips (Thysanoptera) has been recorded as an egg predator of Chloropidae (Znamensky, 1926; Tzygankov, 1930). Some unspecialized predators may help control some of the common agricultural pest species among Chloropidae. Heidger & Nentwig (1989) noted that the spider Dictyna arundinaceae (Linnaeus) builts its web preferably at the ears of wheat and catches almost exclusively adults of Oscinella frit. Some Coccinellidae are known to eat larvae of O. frit (Collin, 1918) and several species of Carabidae (Bembidion lampros, B. quadrimaculatum, Pseudoophonus rufijipes, 38

Broscus cephalotes, Pterostichus cupreus, P. punctulatus, Carabus convexus) are listed as predators of frit flies (Toldaev, 1972). Bembidion lampros feeds on eggs of Oscinella situated on the ground (Nordlander, 1978a). Also predaceous mites of the genus Pergamasus (Parasitidae) eat chloropid eggs (Jepson & Southwood, 1958). Jones (1968a) noted that the activity of larger carabid beetles may disturb females of Oscinella frit in laying eggs on shoots of oats. The plant bug Miris dolabratus (Miridae) was once recorded as a predator of frit flies (Collin, 1918), but this seems questionable. Ochthera mantis (Ephydridae) feed on adult chloropids (Hobby, 1931). Adults of the Oscinella frit complex often carry red larvae of the mite Microtrombidium demeijerei (Oudemans) (Microtrombiidae) (Riggert, 1935). Riggert (1935) proposed a negative influence of this mite on adults of Oscinella frit similar to that of parasitic Nematoda. Another microtrombiid mite species, Ettmuelleria caudatum was found on the abdomen of Chlorops pumilionis as well as on Sphaeroceridae (Smith, 1955). Adults and larvae of the Oscinella frit complex are hosts of parasitic and saprophylic nematodes. Young fertilized females of the obligate parasitic nematode Howardula oscinellae (Allantonematidae) penetrate and live in the larval haemocoel of Oscinella spp. The viviparous parasite is carried through the pupal stage and produces its larvae in the body of the adult host fly. These larvae leave their host fly by entering its gut. Flies infested by this nematode are affected in the development of their inner reproductive organs (Goodey, 1930a, 1930b; Filipjev, 1934; Wachek, 1955; Poinar, 1975; Moore & Hunt, 1984; Slobodyanyuk, 1984). Other species of parasitic nematodes belonging to Mermithidae and Auguillulidae have been recorded from adult Chlorops pumilionis and Lasiosina herpini (Tzygankov, 1930). A parasitic mermithid nematode, probably a Hexamermis sp., has also been found in larvae of O. frit (Moore & Hunt, 1984). Even the saproxylic nematode Panagrolaimus rigidus (Panagrolaimidae) causes some mortality of Oscinella spp. larvae (Poinar, 1972, 1975). Usually only two or three, but up to six were found in the haemocoel of living larvae, and as many as 12 in a dead larva (Moore & Hunt, 1984, 1987). Jepson & Southwood (1958) found that the poor state of overwintering larvae of Oscinella frit was caused by the presence of a yeast and a ciliate protozoan in their body cavity.

Table 2. Parasitoids of Chloropidae recorded from Europe. Parasitoids Proctotrupoidea Diapriidae Basalys (= Loxotropa) crassiclava Kieffer ?Basalys tritoma (Thomson)

Basalys sp. Trichopria aequata (Thomson) (= variipes Kieffer) Trichopria (= Tropidopria) compressa Thomson T. verticillata Latreille Trichopria (= Ashmeadopria) sp. Trichopria sp. Platygastridae Paramisius sp. Platygaster sp. Ceraphronoidea Megaspilidae Dendrocerus chloropidarum Dessart Cynipoidea Eucoilidae ?Eucoila sp. Ganaspis mundata Förster Ganaspis sp.

Chloropidae hosts

Authors

Oscinella frit

Chukanova & Kovalchuk, 1974; Dyurich, 1980; Talitskiy & Kuslitskiy, 1990 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 Chukanova & Kovalchuk, 1974 Vojnovskaya-Kriger, 1929; Riggert, 1935; Simmonds, 1952; Fulmek, 1968; Chukanova & Kovalchuk, 1974 Fulmek, 1968

Oscinella frit

Oscinella frit Oscinella frit Oscinella frit

Oscinella frit

Dyurich, 1980 Talitskiy & Kuslitskiy, 1990 Riggert, 1935; Simmonds, 1952; Fulmek, 1968; Chukanova & Kovalchuk, 1974 Chukanova & Kovalchuk, 1974

Oscinella frit Oscinella frit

Chukanova & Kovalchuk, 1974 Chukanova & Kovalchuk, 1974; Moore, 1983

Hapleginella sp. or Gaurax sp.

Dessart, 1990

Oscinella frit Oscinella frit Oscinella frit cone inhabitant Chloropidae

Fulmek, 1968 Nordlander, 1978b Ivatsik & Kovalchuk, 1975 Chukasnova & Kovalchuk, 1974; Nordlander & Grijpma, 1991 Simmonds, 1952; Fulmek, 1968

Meromyza nigriventris M. pratorum Oscinella frit

Grybliographa (= Pseudeucoila) (= Psichacra) sp. Hexacola fuscipes Meyer H. hexatoma (Hartig)

Oscinella frit

Hexacola sp. Kleidotona psiloides Westul. Odoutecoila sp. Rhoptromeris heptoma (Hartig) (= eucera Hartig) (= widhalmi Kurdjumov)

Oscinella frit Oscinella frit Oscinella frit Oscinella frit

Rhoptromeris miris Belizin

Oscinella frit

Oscinella frit Oscinella frit

Meyer, 1923; Riggert, 1935 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 Chukanova & Kovalchuk, 1974 Chukanova & Kovalchuk, 1974 Chukanova & Kovalchuk, 1974 Baranov, 1912; Kurdjumov, 1912; Collin, 1918; Meyer, 1923; Schander & Meyer, 1925; Vojnovskaja-Kriger, 1929; ?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 Chao (Chien ming), 1959; Chukanova & Kovalchuk, 1974

39

Table 2 (Continued). Parasitoids Rhoptromeris strobigena Nordlander & Grijpma

Rhoptromeris sp. Chalcidoidea Pteromalidae Callitula bicolor Spinola

Chloropidae hosts Hapleginella laevifrons Conioscinella gallarum? Gaurax niger? Gaurax ephippium? Oscinella frit

Authors Nordlander & Grijpma, 1991

Oscinella frit Polyodaspis rufijicornis Elachiptera cornuta Chlorops pumilionis Lasiosina herpini Meromyza nigriventris

Imms, 1930, 1932; Riggert, 1935 Nikolskaya, 1937; Goodliffe, 1942 Simmonds, 1952; Hemer, 1960 Fulmek, 1962, 1968; Zamfirov, 1962 Jones, 1965, 1968b Chukanova & Kovalchuk, 1974; Dzhanokmen, 1978; Nordlander, 1978b; Dyurich, 1980; Talitskiy & Kuslitskiy, 1990 Goodliffe, 1942; Simmonds, 1952; Fulmek, 1968; Dzhanokmen, 1978 Tzygankov, 1930

C. pyrrhogaster (Walker)

Oscinella frit

C. (= Micromelus) rufomaculatus Nees (secondary parasitoid?) Cyrtogaster clavicornis Walker C. vulgaris Walker (hyperparasitod)

Chlorops pumilionis; Meromyza nigriventris Oscinella frit Coelinidea nigra

Eupelmes artropurpureus Dalman

Chlorops pumilionis; Lasiosina herpini Chlorops pumilionis Oscinella frit Oscinella frit

Eupteromalus hemipterus Walker Halticoptera aenea Walker

H. circulus (Walker) (= fuscicornis Walker) (= petiolata Thomson) (= suilis Walker)

Oscinella frit

Lasiosina herpini

?H. hieracii (Thomson) H. patellana (Dalman) Thomson Homoporus (= Merisus) destructor (Say) (= intermedius Lindeman)

Chlorops pumilionis Oscinella frit Oscinella frit

40

Chukanova & Kovalchuk, 1974 Balachowsky & Mesnil, 1935; Riggert, 1935; Simmonds, 1952; Hemer, 1960; Fulmek, 1962, 1968; Jones, 1968b; Chukanova & Kovalchuk, 1974; Dzhanokmen, 1978; Nordlander, 1978b Dzhanokmen, 1978 Hemer, 1960 Graham, 1969; Dzhanokmen, 1978 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 Cunliffe, 1921; Meyer, 1923; Vojnovskaja-Kriger, 1929; Schander & Meyer, 1925 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 Fulmek, 1962, 1968 Fulmek, 1968 Wilhelm, 1891; Collin, 1918; Meyer, 1923; Mejer, 1929

Lasiosina herpini Lasiosina herpini Lipara lucens; L. rufijitarsis

Vojnovskaya-Kriger, 1929; Imms, 1930; Riggert, 1935; Simmonds, 1952; Fulmek, 1968; Chukanova & Kovalchuk, 1974; Dzhanokmem, 1978 Nikolskaya, 1937 Nikolskaya, 1937 Séguy, 1934; Fulmek, 1968

Lipara pullitarsis galls

Dely-Draskovits et al., 1993

Elachiptera cornuta

H. (M.) febriculosus (Girault) H. (M.) mordellistenae Crawford Lamprotatus chrysochlorus Walker Merismus nitidus Walker

Chukanova & Kovalchuk, 1974

Table 2 (Continued). Parasitoids Merisus sp. Mesopolobus (Amblymerus) sp.

P. umbratum Delucchi Pachyneuron sp. ?Polycystus oscinidis Kurdjumov

Chloropidae hosts Oscinella frit Dicraeus ingratus; D. tibialis; D. agropyri; D. humeralis Oscinella frit Lipara lucens L. rufijitarsis Oscinella frit Oscinella frit Oscinella frit

?Pteromalus muscorum Walker P. planiscuta Thomson P. puparum Linnaeus

Chlorops pumilionis Oscinella frit Oscinella frit

Pteromalus (Habrocytus) sp.

Oscinella frit Chlorops pumilionis

Racosina deplanata Bouček

Roptrocerus mirus (Walker) ?Semiotelus nigripes Lindeman

Polyodaspis rufijicornis Cryptonevra flavitarsis Elachiptera cornuta Chlorops pumilionis Oscinella frit

Spalangia fuscipes Nees

Oscinella frit

S. nigra Latreille Spalangia sp. (?S. nigra) Sphegigaster sp. Stenomalina communis (Nees) (= continua Walker) (= rugosa Thomson) (= laetus Ruschka)

Oscinella frit Oscinella frit Oscinella frit Chlorops pumilionis

Pachyneuron concolor Förster P. formosum Walker

C. geminatus Meromyza pratorum M. nigriventris S. epistena Walker S. illudens (Walker) (= crassicornis Dalman) S. liparae (Giraud)

S. micans (Olivier)

Chloropidae Chlorops pumilionis Lipara galls L. lucens; L. similis L. rufijitarsis L. pullitarsis

Oscinella frit Chlorops pumilionis Meromyza nigriventris

Authors Chukanova & Kovalchuk, 1974 Moiseev, 1950; Agafonova, 1962 Chukanova & Kovalchuk, 1974 Giraud, 1863; Séguy, 1934 Fulmek, 1968 Dzhanokmen, 1978 Chukanova & Kovalchuk, 1974 Mokrzecki, 1913; Collin, 1918; Meyer, 1923; Imms, 1930; Riggert, 1935; Simmonds, 1952; Fulmek, 1968 Lampa, 1888 Mejer, 1929; Simmonds, 1952; Fulmek, 1968 Rörig, 1893; Collin, 1918; Meyer, 1923; Imms, 1930; Riggert, 1935; Simmonds, 1952; Fulmek, 1968 Ruschka & Fulmek, 1915 Meyer, 1923; Riggert, 1935; Simmonds, 1952; Fulmek, 1962, 1968; Chukanova & Kovalchuk, 1974 Kiauka & Nartshuk, 1972 Askew & Shaw, 1978 Dyurich, 1980; Abraham, 1983; Talitskiy & Kuslitskiy, 1990 Fulmek, 1962, 1968 Wilhelm, 1891; Collin, 1918; Riggert, 1935; Simmonds, 1952; Hemer, 1960 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 Riggert, 1935; Fulmek, 1968 Simmonds, 1952 Chukanova & Kovalchuk, 1974 Lampa, 1888; Ruschka & Fulmek, 1915

Kearns, 1931; Balachowsky & Mesnil, 1935 Nikolskaya, 1937 Fulmek, 1962, 1968; Graham, 1969; Dyurich, 1980; Talitskiy & Kuslitskiy, 1990 Dzhanokmen, 1978 Lampa, 1888 Giraud, 1863; Wagner, 1907; Blair, 1932, 1944b Otten, 1940; Ruppolt, 1956–1957 Gracham & Claridge, 1965 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 Gureau, 1861; Porchinsky, 1881 Rörig, 1893; Schesterikov, 1910 Kurdjumov, 1912; Collin, 1918

41

Table 2 (Continued). Parasitoids

Chloropidae hosts Lasiosina herpini

S. muscarum Linnaeus (hyperparasitoid) Stenomalina sp.

Coelinidea nigra

Trichomalopsis micropterus Lindeman

Oscinella frit

Trichomalopsis sp. Trichomalus cristatus (Förster)

Oscinella frit Oscinella frit Thaumatomyia notata

T. nanus (Walker) (= frontalis Thomson)

Oscinella frit

Chlorops pumilionis

Chloropidae T. posticus Walker T. statutus (Förster)

Trichomalus sp. Eupelmidae Eupelmus atropurpureus Salman

Eupelmes microzonus Förster

Eupelmella vesicularis Retzius Eurytomidae Eudecatoma fasciata (Thomson) E. nartshukae Zerova E. palustris Erdös (= paludicola Zerova) E. phragmiticola Zerova Eurytoma lucidula Zerova ?Tetramesa phragmitis (Erdös) Torymidae Torymus arundinis Walker Eulophidae Aprostocetus arundinis (Giraud) A. ciliatus (Nees) A. pausiris (Walker)

42

Authors 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 Wahlgren, 1918; Tzygankov, 1930; Balachowsky & Mesnil, 1935; Fulmek, 1962, 1968 Fulmek, 1962, 1968; Dyurich, 1980; Talitskiy & Kuslitskiy, 1990 Nikolskaya, 1937; Simmonds, 1952; Fulmek, 1968; Chukanova & Kovalchuk, 1974; Dyurich, 1980; Talitskiy & Kuslitskiy, 1990 Chukanova & Kovalchuk, 1974 Baranov, 1912; Collin, 1918 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 Meyer, 1923; Riggert, 1935

Oscinella frit Chloropidae Oscinella frit Pseudopachychaeta rufijiceps Chloropidae Oscinella frit

Fulmek, 1968; Dzhanokmen, 1978; Nordlander, 1978a, 1978b Dzhanokmen, 1978; Dyurich, 1980 Talitskiy & Kuslitskiy, 1990 Nartshuk, 1962a Graham, 1969 Dzhanokmen, 1978 Ivatsik & Kovalchuk, 1975

Chlorops pumilionis Lasiosina herpini Meromyza nigriventris Chlorops pumilionis Lasiosina herpini Meromyza nigriventris Chlorops pumilionis Meromyza nigriventris

Tzygankov, 1930 Nikolskaya, 1937 Fulmek, 1962, 1968 Tzygankov, 1930 Nykolskaya, 1937 Fulmek, 1962, 1968 Tzygankov, 1930 Nikolskaya, 1937; Fulmek, 1962, 1968

Lipara lucens Lipara lucens Eurina ducalis; Lipara lucens

Fulmek, 1968 Zerova, 1977 Zerova, 1978

Lipara galls Chlorops strigulus Lipara galls; preferable L. rufijitarsis

Zerova, 1978 Zerova, 1995 Erdös, 1957a; Dely-Draskovits et al., 1993

Lipara similis

Fulmek, 1968

Lipara lucens

Valkeila, 1959; Fulmek, 1968; Tryapitsyn & Kostyukov, 1978 Fulmek, 1968 Wolf, 1988, 1991

Lipara lucens Lipara lucens

Table 2 (Continued). Parasitoids Aprostocetus sp. Closterocerus lagus (Walker) Digliphus sp. Hemiptarsenus sp. Horismeniu specularis Erdös Kostjukovius platycephalae Kostjukov Mellitobia sp. Necremnus leucarthros Nees Neochrysocharis albipes Kurdjumov N. immaculatus Kurdjumov

Pediobius epigonus (Walker) P. facialis (Giraud)

P. metallicus (Nees) P. saulius Walker Pediobius (Pleurotropis) sp. Pronotalia inflata Graham P. liparae Gradwell P. valkeilai Gradwell Tetrastichus atroceruleus (Nees) T. djuritshae Kostjukov T. legionarius (Giraud)

Tetrastichus sp.

Tetrastichus sp. Tetrastichus sp. Elasmidae Crataepiella liparae Gradwell C. carlinarum Szelényi & Erdös Aphelinidae Aphelinus flavipes Förster ?Centrodora amoena Förster Trichogrammatidae Trichogramma evanescens Westwood Trichogramma sp.

Chloropidae hosts L. lucens; L. similis; L. rufijitarsis Oscinella frit Oscinella frit Oscinella frit Chloropidae Platycephala umbraculata Lipara lucens L. pullitarsis Oscinella frit Oscinella frit Oscinella frit

Oscinella frit Lipara lucens galls L. similis L. rufijitarsis Oscinella frit Lipara galls Oscinella frit Lipara lucens Lipara lucens Calamoncosis minima Lipara lucens Chlorops novakii Lipara lucens L. similis L. rufijitarsis Cryptonevra flavitarsis Neohaplegis tarsata

Lipara lucens L. pullitarsis Chlorops geminatus C. novakii Oscinella frit Dicraeus ingratus D. tibialis

Authors Dely-Draskovits et al., 1994 Howell, 1982; Moore, 1983, 1984; Nielsen, 1994 Chukanova & Kovalchuk, 1974 Chukanova & Kovalchuk, 1974 Tryapitsyn & Kostjukov, 1978 Dyurich & Kostjukov, 1978; Dyurich, 1980; Talitskiy & Kuslitskiy, 1990 Abraham & Carstensen, 1982 Hemer, 1962 Simmonds, 1952; Fulmek, 1968 Kurdjumov, 1912; Meyer, 1923; Mejer, 1929; Imms, 1930; Riggert, 1935; Nikolskaya, 1937; Simmonds, 1952; Fulmek, 1968; Tryapitsyn & Kostjukov, 1978 Tryapitsyn & Kostjukov, 1978 Giraud, 1863; Fulmek, 1968 Dely-Draskovits et al., 1994 (hyperparasitoid of Ephialtes arundinis) Meyer, 1923; Imms, 1930; Simmonds, 1952; Fulmek, 1968 Viggiani, 1964 (hyperparasitoid of Exeristes arundinis) Chukanova & Kovalchuk, 1974 Graham, 1991; Wolf, 1991 Gradwell, 1957; Valkeila, 1959 Gradwell, 1957; Valkeila, 1959 Fulmek, 1968 Kostjukov, 1979; Talitskiy & Kuslitskiy, 1990 Giraud, 1863; Henriksen, 1918–1919 Otten, 1940; Erdös, 1955, 1956, 1957b Ruppolt, 1956–1957 Fulmek, 1968; Waitzbauer, 1969 Chvála et al., 1974; Dyurich, 1980; Hudek et al., 1981; De Bruyn, 1987; Wolf, 1988; Talitskiy & Kuslitskiy, 1990; Plant, 1997 Abraham & Carstensen, 1982 Dyurich, 1980 Talitskiy & Kuslitskiy, 1990 Howell, 1982 Agafonova, 1962

Lipara lucens Calamoncosis minima

Fulmek, 1962; Tryapitzyn & Kostjukov, 1978 Gradwell, 1957; Valkeila, 1959

Oscinella frit Lipara lucens

Anonymous, 1963 Fulmek, 1968

Oscinella frit

Chernoponevkina, 1939

Oscinella frit

Fulmek, 1968

43

Table 2 (Continued). Parasitoids Signiphoridae Clytina giraudi Erdös

Mymaridae Gonatocerus sulphureus Förster Ichneumonoidea Ichneumonidae Bathythrix decipiens Gravenhorst (?hyperparasitoid) ?Campodorus variegates (Jurine) (= sanguicollis Gravenhorst) ?Diadegma chrysostictum (Gravenhorst) D. fenestralis Holmgren Endromopoda arundinator (Fabricius) (= melanopyga Gravenhorst)

E. detrita (Holmgren)

E. phragmitidis (Perkins)

Endromopoda sp. (as Scambus sp.) Exeristes arundinis Kriechbaumer

Gelis liparae (Giraud)

Chloropidae hosts

Authors

Cryptonevra flavitarsis C. diadema Lipara lucens L. rufijitarsis Calamoncosis minima Neochaplegis tarsata Elachiptera cornuta

Erdös, 1957a, 1957b, 1958; Fulmek, 1968 Tryapitsyn, 1978; Dyurich, 1980 Talitskiy & Kuslitskiy, 1990

Oscinella frit

Schander & Meyer, 1924; Simmonds, 1952; Fulmek, 1968

Lipara lucens

Blair, 1932; Séguy, 1934; Fulmek, 1968

Lipara lucens

Fulmek, 1968

Lipara lucens

Starke, 1940

Oscinella frit Lipara lucens

Chukanova & Kovalchuk, 1974 Giraud, 1863; Wagner, 1907

L. rufijitarsis L. pullitarsis L. similis Platycephala planifrons

Blair, 1932; Hedwig, 1950, 1955 Aerts, 1956; Fulmek, 1962, 1968 Waitzbauer, 1969; Chvála et al., 1974 Doskočil & Chvála, 1974; Imhof, 1979; Dyurich, 1980; Kasparjan, 1981; Grochowska, 1989; Talitskiy & Kuslitskiy, 1990 Wagner, 1907 Henriksen, 1918–1919; Blair, 1932, 1944b Starke, 1940; Šedivý, 1963; Fulmek, 1968 Waitzbauer, 1969; Raghi-Atri, 1980 Perkins, 1957; Mook, 1967; Waitzbauer, 1969 Waitzbauer et al., 1973; Chvála et al., 1974 Imhof, 1979; Fitton et al., 1988 Skuhravý & Skuhravá, 1978; Dyurich, 1980; Hudec et al., 1981; Skuhravý, 1981; Talitskiy & Kuslitskiy, 1990; Dely-Draskovits et al., 1993; Reader, 2003 De Bruyn, 1987

Lipara galls L. lucens L. similis L. rufijitarsis Lipara lucens L. similis L. rufijitarsis Platycephala planifrons

Lipara galls, primarily L. rufijitarsis Lipara lucens Platycephala planifrons

G. notabilis (Förster) Glypta sp.

Lipara lucens L. rufijitarsis Lipara lucens Oscinella frit

Phrygaedeon troglodytes Giraud ?Pimpla turionellae Linnaeus Promethes sulcator (Gravenhorst) Scambus planatus (Hartig)

Lipara rufijitarsis Chlorops pumilionis; C. strigulus Oscinella frit Lipara lucens

44

Henriksen, 1918–1919; Seyrig, 1927 Hellén, 1949; Valkeila, 1949; Fulmek, 1968; Waitzbauer, 1969; Waitzbauer et al., 1973; Kasparjan, 1981; Dely-Draskovits et al., 1994 Giraud, 1863 Henriksen, 1918–1919; Fulmek, 1968 Henriksen, 1918–1919; Blair, 1932 Fulmek, 1968; Waitzbauer, 1969; Chukanova & Kovalchuk, 1974 Aerts, 1940 Fulmek, 1968 Anonymous, 1963 Fulmek, 1968

Table 2 (Continued). Parasitoids S. sagax (Hartig) S. nigricans (Thomson) (= similis Bridgman) (var. habermehli Schmideknecht) S. nucum Ratzeburg Braconidae ?Aleiodes rufijicornis (Herrich-Schäffer) (= Rhogas dimidiatus Spinola) Baryproctus barypus (Marshall) ?Blacus (B.) interstitialis Ruthe Bracon discoideus Wesmael Bracon (B.) longicollis Wesmael B. (B.) longulus Thomson Bracon (B.) minutator (Fabricius) (= abscissor Nees) Bracon sp. Bracon (Glabrobracon) discoideus Westmael ?Cardiochiles saltator (Fabricius) var. brachialis Rondani Chasmodon apterus (Nees)

Chorebus lar Morley Chorebus sp. aff. ampliator (Nees) Chorebus sp. C. (Gyrocampa) pospelovi Kurdjumov Coelinidea nigra (Nees)

C. vidua (Curtis) ?Coeloides abdominalis (Zetterstedt) ?C. melanostigma Strand (= stigmaticus Hell) Dacnusa areolaris (Nees) Dacnusa sp. ?Gonatocerus sulphuripes Förster Microgaster sp.

Chloropidae hosts Lipara lucens L. similis Lipara lucens

Fulmek, 1968; Dely-Draskovits et al., 1974; Kasparyan, 1981

Lipara lucens

Starke, 1940

Lipara lucens

Starke, 1940; Hedwig, 1950; Fulmek, 1968

Lipara similis L. rufijitarsis Oscinella frit Chlorops pumilionis Chlorops pumilionis Oscinella frit Lipara lucens

Dyurich, 1980 Talitskiy & Kuslitskiy, 1990; Dely-Draskovits et al., 1994 Fischer, 1963; Haeselbarth, 1973; Achterberg, 1988; Koponen, 1991, 1993 Fulmek, 1968 Kearns, 1931; Fulmek, 1962, 1968 Ivatsik & Kovalchuk, 1975 Henriksen, 1918–1919

Lipara galls Chlorops pumilionis Chlorops pumilionis

Séguy, 1934; Fulmek, 1968 Fulmek, 1962, 1968 Fulmek, 1962, 1968

Chlorops pumilionis

Porchinsky, 1881; Fulmek, 1962, 1968

Oscinella frit Oscinella vastator; O. frit Meromyza saltatrix Lasiosina herpini

Chlorops pumilionis Lipara lucens

Cunliffe, 1921; Vojnovskaya-Kriger, 1929 Riggert, 1935; Simmonds, 1952 Jones, 1965, 1969b; Fulmek, 1968 Dyurich, 1980; Moore, 1983, 1984; Moore et al., 1986; Moore & Hunt, 1987; Moore & Ridout, 1987; Talitskiy & Kuslitskiy, 1990; Umoru, 1993a, 1993b Dyurich, 1980 Dyurich, 1980; Talitskiy & Kuslitskiy, 1990 Chukanova & Kovalchuk, 1974 Kurdjumov, 1912; Meyer, 1923; Schander & Meyer, 1925; Riggert, 1935 Goureau, 1861; Lampa, 1888 Marshall, 1891; Rörig, 1893 Wahlgren, 1918; Znamensky, 1923 Tzygankov, 1930 Kearns, 1931; Séguy, 1934 Balachowsky & Mesnil, 1935 Goodliffe, 1939, 1942; Fulmek, 1962, 1968; Anonymous, 1963; Dyurich, 1980; Talitskiy & Kuslitskiy, 1990 Fulmek, 1962, 1968 Fulmek, 1968

Lipara lucens

Starke, 1940; Hedwig, 1950; Fulmek, 1968

Chlorops pumilionis Chlorops pumilionis Oscinella frit Oscinella frit

Ruschka & Fulmek, 1915; Fulmek, 1962, 1968 Fulmek, 1962, 1968 Schander & Meyer, 1925 Chukanova & Kovalchuk, 1974

Oscinella frit Oscinella frit Oscinella frit Oscinella frit Oscinella frit Chlorops pumilionis C. novakii; C. strigulus C. laetus Lasiosina herpini Meromyza pratorum M. nigriventris

Authors Fulmek, 1968

45

Table 2 (Continued). Parasitoids Opius caesus Haliday O. irregularis Wesmael O. maculipes Wesmael

Chloropidae hosts Oscinella frit Oscinella frit Oscinella frit

O. piceus Thomson Opius sp. Phaenocarpa helophiliae Achterberg Polemochartus aboletus Papp

Oscinella frit Oscinella frit Calamoncosis glyceriae

P. breviventris (Telenga) P. liparae (Giraud)

Lipara sp. Lipara galls Preferable L. lucens L. pullitarsis L. similis L. rufijitarsis

P. melas (Giraud)

Lipara galls Preferable L. rufijitarsis

Oscinella frit

Papp, 1992; Dely-Draskovits et al., 1993, 1994; Umoru, 1993a, 1993b; Nielsen, 1994 Papp, 1992 Giraud, 1863; Marshall, 1891 Reynvaan & Docters van Leeuwen, 1906 Wagner, 1907; Blair, 1932, 1944b Seguy, 1934; Starke, 1940 Erdös, 1955, 1957a, 1957b; Ruppolt, 1956, 1957; Mook, 1961, 1967; Fulmek, 1968; Waitzbauer, 1969; Waitzbauer et al., 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 Giraud, 1863; Marshall, 1891 Reynvaan & Docters van Leeuwen, 1906; Fulmek, 1968; Waitzbauer et al., 1973, 1974 Chvála et al., 1974; Skuhravý, 1981; Maetô, 1983b; De Bruyn, 1987; Dely-Draskovits et al., 1993, 1994 Ruschka & Fulmek, 1915 Vojnovskaya-Kriger, 1929; Riggert, 1935; Simmonds, 1952; Fulmek, 1962, 1968 Maetô, 1983a Dyurich, 1980; Talitskiy & Kuslitskiy, 1990 Curtis, 1860 Rörig, 1893; Collin, 1918 Meyer, 1923; Riggert, 1935; Simmonds, 1952; Fulmek, 1962, 1968 Ivatsik & Kovalchuk, 1975

Oscinella frit

Cunliffe, 1921; Simmonds, 1952; Fulmek, 1968

Oscinella frit

Chukanova & Kovalchuk, 1962

Lipara similis

Platycephala planifrons Protodacnusa tristis (Nees)

Oscinella frit Chlorops pumilionis

Sarops rea Nixon

Cryptonevra sp. C. flavitarsis Oscinella frit Oscinella frit Chlorops pumilionis

Sigalphus caudatus (Curtis) ?Triaspis caudata (Nees)

Triaspis obscurella Nees Aphidiidae ?Aphidius avenae Haliday (= granarius Marshall) ?Aphidius sp.

Authors Anonymous, 1963 Anomymous, 1963 Simmonds, 1952; Fulmek, 1968; Dyurich, 1980; Talitskiy & Kuslitskiy, 1990 Anonymous, 1963 Bhaattacharyya, 1957; Umoru, 1993b Achterberg, 1998

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. The name “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 (Ichneumonidae), 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 (AlmaAta, Kazakhstan) and Dr M.D. Zerova (Schmalhausen Institut of Zoology, Kiev, Ukraine).

46

Ryzhkova (1962) found in the salivary glands and other organs of larvae of Oscinella frit and O. pusilla symbiotic bacteria which in her opinion are involved in the digestion of plant tissues. Later, Sasonov (1966) investigated in detail the relationship between micro-organisms and larvae of Oscinella species. He found no microorganisms in the salivary glands of the larvae using both light and electron microscope, but the bacteria Pseudomonas fluorescens, P. mycophaga and Chromobacter flavum were found in the larval gut and Bacterium ciliaerogenas on the larval body. He also found the following bacteria in plant tissues fed upon by the larvae: Mycobacterium globiforme, Pseudomonas fluorescens, P. herbicola, Bacterium aquatile acidifijicans, B. candicans lactium. These epiphytic and phytopatogenous bacteria are known as decomposers of complex plant proteins. Another group

of bacteria found in frit flies are considered as pathogenous. Of these, 26 species have been recorded from chloropid eggs, larvae, pupae and adults (Aleksandrov et al., 1969; Aleksandrov & Fedosov, 1971; Gritsenko & Fedosov, 1972). Only freshly laid eggs are sterile. The microflora of the larvae is formed by bacteria occurring in plant tissues. Some bacteria are transmitted from larvae to adults. Adults taken in the field contain up to 19 species of bacteria, adults hatching from puparia in the laboratory up to 5 species (Gritsenko & Fedosov, 1972). Two species of entomophagous fungi have been found in England on two species of Chloropidae: Entomophthora americana on Lipara lucens and E. culicis on Thaumatomyia glabra (Leatherdale, 1966, 1970).

Distribution and zoogeography The family Chloropidae is worldwide in distribution. Some Palearctic species extend northwards to the coast of the Arctic Ocean (Nartshuk, 2002b). Three species: Oscinella frit s.l., Cetema cereris and Chlorops hypostigma are recorded from Faroe Islands, one species: Oscinella frit from Iceland, and no species from Greenland (Mazon, 1890; Meijere, 1910; Becker, 1915; Nielsen & Tuxen, 1954; Lyneborg, 1968; Lindroth et al., 1973; Messersmith, 1982; Ólafsson, 1991). Mazon (1890) mentioned also Madiza palpora (sic!) Fln., now Lasiambia palposa (Fallén) from Iceland, but this has no been confirmed. Recently 484 specimens of Chloropidae were collected in Iceland (Prescher et al., 2005) that all proved to belong to O. frit (M. v. Tschirnhaus, pers. comm.). The diversity of chloropids in terms of genera and species increases southwards in Europe and remaining parts of the Palearctic Region. In Fennoscandia and Denmark are presently recorded 203 species in 48 genera. This is nearly 50% of the European species, but 80% of the genera. The subfamily Siphonellopsinae with two European genera is absent in North Europe. In Oscinellinae with 27 European genera only Aprometopis Becker is not found in North Europe. In the Chloropinae 8 genera (Assuania Becker, Capnoptera Loew, Elachiptereicus

Becker, Homalura Becker, Metopostigma Becker, Paraeurina Duda, Phyladelphus Becker and Platycephalisca Nartshuk) out of 28 do not reach North Europe. All missing genera except Homalura and Platycephalisca have a Mediterranean distribution. In comparison to the 203 species of chloropids found in Fennoscandia and Denmark, the following species numbers can be given for some European countries: Great Britain – 175 species (Ismay, 1998), Germany – 208 (Wendt, 1999; Schumann, 2002), Belgium – 112 (De Bruyn & Tschirnhaus, 1991), Netherlands – 140 (Beuk & Nartshuk, 2002), Poland – 170 (Nartshuk, 2002a), Latvia – 115 (Karpa, 2001), Lithuania – 97 (Nartshuk & Pakalniškis, 2004), Hungary – 217 (Dely-Draskovits, 2001), Schwitzerland – 134 (Dely-Draskovits, 1998), Czech and Slovak Republics – 151 (Roháček, 1997), Bulgaria – 138, all Balkan Peninsula – 180 (Beschovski, 1985), Italy – 190 (Nartshuk, 1995), Spain – 91 (De Bruyin & Báez, 2002; Nartshuk, 2004). The chloropid fauna of Fennoscandia and Denmark is composed of species with very different total distribution ranges: (1) widespread Old World species found in the Palearctic, Afrotropical and/or Oriental Regions – 5 (2.4%), Holarctic – 17 (8.3%), trans-Palearctic boreal – 67 (33.3%), southPalearctic eremial (steppes and semideserts) – 3 47

(1.4%), Eurasian – 43 (21.1%), Euro-Mediterranean 12 (5.9%), European (including Euro-Caucasian) – 55 (27.1%), and Mediterranean-Atlantic (coastal) – 1 species (0.5%). The actual distribution of the different chloropid species occurring in Fennoscandia and Denmark is insufficiently known, especially in Norway, Karelia and the Kola Peninsula. Still, some general remarks may be in place here. A preliminary analysis of species distributions within the subfamily Chloropinae was given earlier (Nartshuk, 2002b). The area of present Fennoscandia and Denmark was covered entirely with ice during the Riss glacial period, and only the western part of Jutland (Denmark) was ice free during the following Würm glacial period. Recolonization of this territory took place gradually during the retreat of the glacier, 12– 10 000 years ago. The latitudinal extend of Fennoscandia and Denmark is impressive (nearly 18 degrees from the southern border of Denmark to the Norwegion coast at the Arctic Ocean) and corresponds to the latitudinal extend of the remaining European territory. The main route of colonization of Fennoscandia came from the south through Denmark. As for many other insect groups it is evident that most chloropid species occurring in the northern parts of Fennoscandia have Holarctic or transPalearctic total distributions, whereas species confined to Denmark and the southern parts of Fennoscandia have a stronger representation of primarily European and Euro-Mediterranean species. As in most other groups of organisms the species diversity of Chloropidae decrease gradually to the north in Fennoscandia. Thus, for most species the northern border of their European distribution range goes across Fennoscandia at some latitude. Only 39 species, or 19.2% of the Scandinavian Chloropidae have been found north of the Arctic circle, and most of these have much wider distributions further southwards in Fennoscandia and Denmark. There are surprisingly few examples of chloropid species with exclusively high boreal to subarctic distributions in Fennoscandia: Conioscinella abiskoi, C. tornensis and C. messaurea and Aphanotrigonum norrbotticum. The Conioscinella species all belong to the C. frontella species complex, which suggests relatively recent speciation. Most chloropid species of Fennoscandia and Denmark can be referred to one of the following four groups determined by the position of their northernmost distribution range: 48

1. This group includes pronouncedly southern (nemoral) species extending no further north than Denmark and the southernmost provinces of Sweden (SK, HA, SM, Öl, GO). It consists of 1 species of Rhodesiellinae, 13 species of Oscinellinae and 15 species of Chloropinae. Some of the southern species have only been found on islands: Dicraeus raptus, Chlorops fijinitimus, C. pallidiventris, C. pannonicus, Chloropsina rohaceki, Neohaplegis glabra on Öland, D. ingratus on Bornholm and Gotland. This group mainly includes species with Euro-Mediterranean, some with coastal or eremial habitat preferences. Eutropha fulvifrons is a coastal, atlanticMediterranean species that properly belongs in this group despite a record from NB. 2. This group includes species distributed northwards to the 60°N latitude in Fennoscandia, which approximately coincides with the northern limit of the boreo-nemoral zone: in Sweden to VR and UP, in Finland to Al and Ka, in Russia to Vib and southern Kr. A diverse group consisting of 23 species of Oscinellinae and 20 species of Chloropinae. This group mainly includes species with European, Euro-Caucasian and Euro-Mediterranean distributions. 3. This group includes species extending northwards approximately to the 66°N latitude in Fennoscandia, which approximately coincides with the northern limit of the middle boreal zone: in Sweden to LU and NB, in Finland to Lk. This is the largest group consisting of 19 species of Oscinellinae and 38 species of Chloropinae. Some of the species, e.g. Trichieurina pubescens, Meromyza sibirica and Neohaplegis tarsata, may well extend further north, but there is no data yet. This group includes the diverse fauna of chloropids found associated with Common reed (Phragmites australis), a plant with a northern range coinciding well with that of the present group 3. 4. This group includes the most widely distributed species in Fennoscandia embracing the northern boreal zone, some of them (Oscinella frit, O. ventricosi, Chlorops planifrons, C. scutellaris, C. varsoviensis, Melanum laterale, Pseudopachychaeta rufijiceps, Thaumatomyia trifasciata) even subarctic tundra (Nartshuk, 2002b). The group is well represented by 19 species of Oscinellinae and 17 species of Chloropinae. This group consists mostly of widely distributed Holarctic or trans-Palearctic species

High latitude melanism. In several widely distributed species of Chloropidae it is very evident that specimens from the northern boreal zone of Fennoscandia tend to be significantly more darkcoloured as normal. Northern specimens of Gau-

rax, especially females, are often almost wholly black, and the usually discrete black scutal stripes seen in Chlorops speciosus and C. meigenii tend to widen and fuse together in northern specimens.

Taxonomy Monophyly and relationships The Chloropidae is a well defined family belonging to the Schizophora, a specious higher taxon traditionally divided in two major groups of families: Calyptratae and Acalyptratae. Chloropids belong to the acalyptrate grade. The monophyly of the family is widely accepted. Apomorphies for the family after Andersson (1977): 1. First and second pairs of orbital bristles lateroproclinate. 2. Propleuron with vertical carinate ridge 3. “Tibial organ” present on hind tibia. 4. Vein Sc reduced in apical half. 5. Cross-vein bm-cu absent. 6. Vein CuA1 slightly sinuate near middle of penultimate sector 7. Cell cup absent. 8. Male tergite 6 absent. Additional character states have been suggested as chloropid apomorphies by earlier authors, e.g. Malloch (1934), Hennig (1958) and Griffiths (1972), but they rather represent apomorphic states relative to the ground plan of all acalyptrate families. The relationship of the Chloropidae to the numerous other acalyptrate families has for long been controversial. Several earlier authors placed Chloropidae as an isolated group of “higher” Acalyptratae (Malloch, 1948; Oldroyd, 1954; Hennig, 1958; Harrison, 1959). Hennig (1958) was uncertain about the position of the Chloropidae. He suggested on one hand that the family might belong to either Drosophiloidea or Milichioidea, but ended up with erecting a superfamily Chloropoidea with a single family Chloropidae. Later, Hennig (1969) placed the family as a questionable member of the superfamily Milichioidea together with Milichiidae and Carnidae without specified evidence. Hennig (1971) united Milichiidae and Chloropidae

in the superfamily Chloropoidea based on the observation by Sturtevant (1926) that the females of both families possess a ventral pocket-like seminal receptacle and desclerotized spermathecae. A year later Hennig (1972) regretted this opinion, because sclerotized spermathecae were found in Carnidae, a family he considered most closely related to Milichiidae. Therefore, the absence of sclerotized spermathecae in Milichiidae and Chloropidae might be considered as a homoplasy. Speight (1969) made a comprehensive comparative study of the prosternum in Acalyptratae. He found that Chloropidae possess a well developed prosternal bridge and proposed the inclusion of the family in Drosophiloidea together with Drosophilidae (including Camillidae), Diastatidae, Canacidae, Ephydridae, Tethinidae and Braulidae. Hennig (1972) argued that antennal morphology spoke against uniting the Chloropidae and Milichiidae with Drosophilidae. However, Hackman & Väisänen (1985), based on characters of wing vein C, followed Speight (1969) and included Chloropidae in Drosophiloidea. Griffiths (1972), in a comparative study of male terminalia of cyclorrhaphous Diptera, considered the Chloropidae most closely related to the families Acartophthalmidae, Milichiidae and Carnidae. Wheeler (1994c) supported a sister group relationship between Chloropidae and Milichiidae and rejected a closer relationship between these families and the Risidae and Cryptochetidae. Brake (2000), in a cladistic analysis of Milichiidae, supported the idea of a sister group relationship between Chloropidae and Milichiidae. She also presented a detailed review of previous accounts on the relationships between Milichiidae, Carnidae and Chloropidae. It should be emphasized finally that the phylogenetic relationships of Chloropidae are far from being elucidated. The sister group of Chloropidae may not be found without a detailed analysis of species of the big genus Apotropina Hendel (sub49

Table 3. Assignments of the family Chloropidae in different classifications of Diptera. Taxon Suborder Infraorder Division Sectio Superfamily Prefamily Included families

Suborder Infraorder Division Sectio Superfamily Prefamily Family group Included families

Suborder Division Sectio Superfamily Included families

Stone et al., 1965 Griffiths, 1972* Cyclorrhapha

Delfinado & Hardy, 1977

Steyskal, 1974*

Brachycera

Cyclorrhapha

Schizophora Chloropoidea CHLOROPIDAE

Muscoidea Tephritoinea CHLOROPIDAE, Acartophthalmidae, Milichiidae, Carnidae

Crosskey, 1980 Cyclorrhapha

Evenhuis, 1989 Cyclorrhapha

Schizophora

Schizophora Acalyptratae

Drosophiloidea

Ephydroidea

CHLOROPIDAE, Camillidae, Ephydridae, Diastatidae, Curtonotidae, Drosophilidae, Milichiidae, Carnidae, Tethinidae, Canacidae

CHLOROPIDAE, Curtonotidae, Drosophilidae, Ephydridae, Cryptochetidae, Tethinidae, Canacidae, Neurochaetidae

Yeats & Wiegmann, 1999 Brachycera Muscomorpha Schizophora Acalyptratae Carnoidea CHLOROPIDAE, Canacidae, Carnidae, Cryptochetidae, Milichiidae, Tethinidae

Pape & Thompson, 2010 Cyclorrhapha Schizophora Acalyptratae

Schizophora Acalyptratae Chloropoidea CHLOROPIDAE, Milichiidae, Carnidae McAlpine, 1989 Brachycera Muscomorpha Schizophora Acalyptratae Ephydroidea

CHLOROPIDAE, Camillidae, Ephydridae, Diastatidae, Curtonotidae, Drosophilidae, Cryptochetidae, Tethinidae, Canacidae

Carnoidea CHLOROPIDAE, Acartophthalmidae, 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 characters of the family. The genus is mainly tropical and out of scope of our work. Therefore we only summarize how the family has been classified in catalogues and manuals of Diptera from different zoogeographical regions published the last decades to show diverse opinions (Table 3).

Classification and nomenclature The first species of the present family Chloropidae described with valid binominal names were classified in the genus Musca Linnaeus: Musca frit 50

and M. saltatrix by Linnaeus (1758, 1761), followed by M. umbelliferarum by Scopoli (1763), M. hordei, M. pumilionis and M. avenae by Bjerkander (1777, 1778, 1781), M. lineata, M. minuta, M. umbraculata and M. planifrons by Fabricius (1781, 1787, 1794, 1798), and M. nasuta by Schrank (1781). Five of these names are still valid and include some notorious pest species of cereals and cultured grasses that were first described from Sweden. The oldest names for chloropid genera are Chlorops proposed by Meigen (1803) and Oscinis by Latreille (1804). Fallén (1820) revised the Swedish fauna of these flies and referred them to their own family Oscinidae (as “Oscinides”) based on La-

treille’s name. It was established much later that Oscinis Latreille is a junior synonym of Chlorops Meigen. For that reason Becker (1909, 1910) proposed the genus Oscinella for Oscinis auct., nec Latreille, and replaced the family name Oscinidae with Chloropidae even if it is younger, as first proposed by Rondani (1856) in the family Agromyzidae as one of two chloropid “stirps” [groups] Chloropina and Oscinina. Schiner (1864) adopted the name Chloropina, but treated it as a subgroup of the family Muscidae. The name Chloropidae is now firmly established and valid for the family according to ICZN Art. 40.2. It is correctly cited with author and year as Chloropidae Rondani, 1856 (1820) according ICZN Recommandation 40A. Lioy (1864) divided his family “Heteromyziti” in seven subfamilies, four of them based on generic names of present Chloropidae: Elachipterini, Siphonellini, Meromyzini, and Chloropsini, but this classification was not followed by subsequent authors. Becker (1910) rather followed Rondani by dividing the family in two subfamilies: Oscinellinae and Chloropinae. Unfortunatly, the family-group names Elachipterini Lioy, 1864 and Siphonellini Lioy, 1864 have priority over Oscinellinae Becker, 1910. Therefore, an application to ICZN is required to preserve the well established name Oscinellinae for the subfamily. Duda (1930) divided Choropidae in three subfamilies: Oscinellinae, Chloropinae and Hippelatinae. Later, after inclusion of exotic forms, he split the family in two subfamilies: Siphonellopsinae and Chloropinae. The latter subfamily he split in two tribes Chloropini and Oscinellini; the latter tribe was then split in two superfamilies: Hippelatoidea and Oscinelloidea. Andersson (1977) proposed a new tribe Rhodesiellini in the subfamily Oscinellinae and divided all remaining genera of the Old World into several informal genus groups: 9 in the tribe Oscinellini and 12 in the subfamily Chloropinae. Kanmiya (1983) changed the contents of some of Andersson’s genus groups and erected 4 new genus groups. Nartshuk (1983, 1987) proposed to divide the family into 21 tribes. Andersson (1977, 1979) and Nartshuk (1987) presented some ideas on the phylogeny of Chloropidae, but stressed that much further work was needed. Andersson (1977, 1979) remarked that the family Chloropidae has evolved relatively recently

which, in combination with the size of the group, makes morphology based cladistic analysis of the family very difficult. He proposed a phylogeny for the family (Andersson, 1979), but stressed that it was weakly supported and implied many instances of homoplasy. In the present work we divide the family Chloropidae in three subfamilies: Rhodesiellinae, Oscinellinae and Chloropinae. A fourth subfamily Siphonellopsinae is extralimital and is not discussed further here. Rhodesiellinae was earlier proposed as a subfamily but without much argument (Nartshuk, 1983). It is plesiomorphic in terms of Chloropinae by having costal vein extended to vein M, and plesiomorphic in terms of Oscinellinae by having long, procurved and divergent ocellar setae. Rhodesiellinae and Oscinellinae share the following characters: 1. Costal vein goes to M (plesiomorphy). 2. Ocellar triangle large (not present in all Oscinellinae) (apomorphic). 3. Scutellum elongated with apical setae inserted on tubercles (only in a few species of Oscinellinae) (apomorphic). 4. Femoral organ present (apomorphic). 5. Hypandrium closed (only in a few species of Oscinellinae). Rhodesiellinae and Chloropinae share the following characters: 1. Ocellar setae long, procurved and divergent (plesiomorphic). 2. Tibial organ absent (except in some Chloropinae) (apomorphic). 3. Sclerites in postabdomen of male usually reduced (apomorphic). 4. Cerci reduced or small and fused into mesolobus (apomorphic). 5. Distiphallus membranous and long (apomorphic). As we see it, there are two possible cladistics scenarios (Figs 113, 114): Chloropinae is sister group to Rhodesiellinae + Oscinellinae. Oscinellinae is sister group to Rhodesiellinae + 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

52

Three rather long, outcurved or procurved orb. vti stronger than vte. if 1–3. Two rather strong h, one recurved and one incurved. One strong seta on katepisternum. Lateral setae on scutellum situated closer to anterior margin of

–

scutellum than to apical setae . . . . . . . . Siphonellopsinae (not in the Nordic countries) orb numerous (5–10) and short. vte stronger than vti. if more numerous or if few then up-

2

–

right or recurved. Usually only 1 recurved h or without distinct h. No seta on katepisternum, bare or with some setulae. Usually several lateral setae on scutellum, if only one then situated closer to apical setae than to anterior margin of scutellum . . . . . . . . . . . . . . . . . . . . 2 Costa reaches R4+5 (except some Dicraeus in which 2nd costal sector is 3–4 times longer than 3rd costal sector) (Figs 35–40). Male cerci apically free (Figs 71, 73) . . . . . . . . . . . . 3 Costa not reaching R4+5 (Figs 41–44). Male cerci completely fused, forming a subrectangular sclerite (Figs 76, 80, 81) . . . Chloropinae

3

–

oc procurved and divergent. Male cerci absent. Anepisternum covered with setulae. Scutellum usually more or less triangular and flattened. Apical scutellar setae usually arising from warts or long projections . . . . . Rhodesiellinae (only genus Aspistyla in Nordic countries) oc upright or recurved, parallel, convergent or cruciate. Male cerci usually present. Anepisternum bare or with setulae. Scutellum usually rounded and convex. Apical scutellar setae rarely arising from warts . . . . Oscinellinae

Key to Fennoscandian genera 1 –

2 (1)

–

3 (2)

–

4 (3)

Wings fully developed, usually longer than abdomen . . . . . . . . . . . . . . . . . . . . 2 Wings reduced in size (Figs 2, 40), shorter than abdomen, or practically absent . . . . . . . . . . . . . . . . . . . . . . . . . . 52 Costal vein reaching vein M (Figs 35– 39), if ending between apices of R4+5 and M then vein R2+3 long and 2nd costal sector 3 times as long as 3rd costal sector (Fig. 37) . . . . . . . . . . . . . . . . . . . . 3 Costal vein extending only to R4+5 or slightly beyond, never reaching vein M (Figs 41–44). (Subfamily Chloropinae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 Ocellar setae rather long, directed anteriorly and divergent (Fig. 115). Body black, shining. Arista distinctly pubescent. (Subfamily Rhodesiellinae) . . . . . . . . . . . . . . . Aspistyla Duda (p. 59) Ocellar setae small, upright, convergent or parallel. Body black or partly yellow, shining or matt by dusting. (Subfamily Oscinellinae) . . . . . . . . . . . . . . . . . . . . . 4 Arista thick, cylindrical or flattened, or appearing thick due to long and dense pubescence, coloured black or white (Figs 21–23) . . . . . . . . . . . . . . . . . . . . . . . 5

– 5 (4)

– 6 (5)

–

7 (4)

–

8 (7)

Arista thin, without strong pubescence (Figs 18–20) . . . . . . . . . . . . . . . . . . . . . . . 7 Two long orb on each side of frons (Figs 270, 273). Scutum punctuated and scutellum with tubercles at base of apical and subapical setae (Figs 276, 282, 284, 287) . . . . . . . . . Elachiptera Macquart (p. 90) More than two orb on each side of frons being short and equal in size . . . . . . . . 6 First flagellomere rounded and black; arista subapical, black or white (Fig. 21) . . . . . . . . . Oscinella Becker p. p. (p. 124) First flagellomere angular on upper corner, predominantly yellow; arista apical (Fig. 23). Male wing with black spots (Figs 3, 38) . . . . . . . . . Gampsocera Schiner (p. 100) Large flies, body 4–8 mm long, antennae separated by a wide dusted keel (Figs 4, 9) . . . . . . . . . . . . . Lipara Meigen (p. 118) Small flies, body 1–4 mm long, antennae usually not separated, keel absent or very narrow (Figs 13–15) . . . . . . . . . . . . 8 Scutum with 3 or 5 deeply impressed punctuate lines . . . . . . . . . . . . . . . . Tricimba Lioy (p. 149) 53

–

9 (8)

–

10 (9)

– 11 (10) – 12 (11)

–

13 (12) –

14 (11) –

15 (14)

–

54

Scutum without punctuate lines, but punctuate depressions may be present along dorsocentral line on posterior part of scutum . . . . . . . . . . . . . . . . . . . . . . . . 9 Vein R2+3 long, 2nd costal sector 3–4 times longer than 3rd costal sector (Fig. 37) . . . . . . . . . . . . . Dicraeus Loew (p. 85) Vein R2+3 not so long, 2nd costal sector no more than 2 times longer than 3rd costal sector (Figs 35, 36, 39) . . . . . . . 10 Vein R2+3 very short, 2nd costal sector distinctly shorter than 3rd costal sector (Fig. 39) . . . . . . . . Siphunculina Rondani (p. 146) Vein R2+3 long, 2nd costal sector longer than 3rd costal sector (Figs 35, 36) . . . 11 Thorax dorsoventrally flattened . . . . . 12 Thorax not flattened . . . . . . . . . . . . . . . 14 Scutum and scutellum with coarse microsculpture. Apical setae of scutellum on small tubercles (Figs 473, 479) . . . . . . . . . . . . . Oscinisoma Lioy (p. 135) Scutum and scutellum smooth, without microsculpture, at most slightly sculptured before scutellum . . . . . . . . . . . . . 13 Scutum densely dusted and dull . . . . . . . . . . . . . . . Eribolus Becker (p. 96) Scutum without or with very thin dust, shining, sometimes with a green metallic lustre . . . . . Rhopalopterum Duda p. p. (p. 141) Ocellar triangle densely dusted or subshining through very thin dust . . . . . . 15 Ocellar triangle without any dusting, shining, sometimes with punctures and setulae on its surface . . . . . . . . . . . . . . 22 Body yellow with black or reddish stripes (Fig. 34), if stripes fused and scutum black then postpronota and pleura partly yellow. Crossvein dm-cu distinctly oblique; distance between crossveins equals length of dm-cu (Fig. 543) . . . . Trachysiphonella Enderlein (p. 148) Body black or densely grey-dusted. Crossvein dm-cu more upright and dis-

16 (15)

–

17 (16)

–

18 (16)

–

19 (18)

–

20 (19) – 21 (20)

–

22 (14)

tance between crossveins longer than dm-cu . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 Pleura dull by dusting, if shining below then all body black including genae and face . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Pleura shining, without dust except on upper part of anepisternum; if pleura dusted then proboscis long and geniculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 Genae wider than first flagellomere (Fig. 215) Colliniella Nartshuk & Andersson (p. 77) Genae at most as wide as first flagellomere (Figs 121, 126, 134) . . . . . . . . Aphanotrigonum Duda (p. 61) Vibrissal angle distinctly projecting beyond eye margin. Proboscis may be long and geniculate (Figs 460, 463, 469) . . . . . . . . . . . Oscinimorpha Lioy (p. 133) Vibrissal angle not projecting beyond eye margin and proboscis short (Figs 347, 360, 424, 530) . . . . . . . . . . . . . . . . 19 Frons except ocellar triangle with small shining punctures. Eyes densely haired. Scutellum with many marginal setae (Fig. 532) . . . . . . . . . Speccafrons Sabrosky (p. 148) Frons evenly haired, without punctures. Eyes sparsely short haired. Scutellum with 1 apical and 2–3 subapical pairs of setae . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 Scutum with setulae dispersed . . . . . . 21 Scutum with setulae in rows . . . . . . . . . . . Incertella Sabrosky (p. 109) Scutum densely grey-dusted, convex, if shining then setae and setulae yellow. Distance between crossveins shorter than basal sector of vein M1 . . . . . . . . Conioscinella Duda p. p. (p. 78) Scutum black, very thinly dusted, flattened. Distance between crossveins no shorter than basal sector of vein M1 (Fig. 514) . . . . . Rhopalopterum Duda p. p. (p. 141) Genae distinctly produced and antennae separated by keel. Proboscis very

–

23 (22)

–

24 (23)

–

25 (23)

–

26 (25)

–

27 (25)

–

long and geniculate (Figs 13, 27) . . . . . . . . . Siphonella Macquart (p. 145) Genae barely produced. Proboscis shorter. Female in doubtful cases with compressed, sclerotized cerci (Fig. 184) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 Postocellar setae (poc) parallel, frons distinctly punctuate (Figs 371, 380, 484) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 poc convergent or cruciate, frons not punctuate, dull or covered with silky hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 Antennae fully separated by thin keel (Figs 14, 486), wing cell br sometimes widened (Fig. 487) . . . . . . . . . . . . Polyodaspis Duda (p. 138) Antennal keel only developed in upper part of face (Figs 15, 382) . . . . . . . . Lasiambia Anonymous (p. 114) Ocellar triangle with setulae on its surface (Figs 149, 159, 170, 177, 198, 212, 342) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 Ocellar triangle without setulae on its surface, only on frons outside its side margins . . . . . . . . . . . . . . . . . . . . . . . . . 27 Ocellar triangle with setulae arranged in a full length row along its side margins (Fig. 342) . . . . . . . Hapleginella Duda (p. 108) (one species H. laevifrons) Ocellar triangle with setulae in 2–3 irregular rows, arising from distinct small punctures (Figs 149, 159, 170, 177, 198, 212) . . . . Calamoncosis Enderlein (p. 68) Body yellow with black stripes on scutum, or black, sometimes with yellow marks. Scutum shining, with dispersed, rather long setulae. Eyes and arista with long pubescence. If body entirely black, then it is shining and arista is white. Wing cell R1 rather short and broad, broader than cell R2+3 (Fig. 324) . . . . 28 Body uniformly black, setulae on scutum short, sometimes arranged in rows. Eyes and dark arista with short pubescence . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

28 (27)

–

29 (27)

–

30 (29)

–

31 (30)

–

32 (2)

–

33 (32)

–

Scutellum rounded with remote apical setae (Figs 323, 325) . . . . . . . . . . . . . . . . Gaurax Loew (p. 101) Scutellum conical with apical setae approximated (Fig. 493) . . . . . . . . Pseudogaurax Malloch (p. 139) Ocellar triangle reaching front margin of frons (Fig. 431). Setulae on scutum few, arranged in single rows along acrostichal and dorsocentral lines. Male cerci nearly round or rather large (Figs 426, 429, 434, 441, 445) . . . . . . . . . Oscinella Becker p. p. (p. 124) Ocellar triangle shorter than frons (Figs 223, 346, 362). Setulae on scutum more numerous and evenly dispersed, or at least arranged in multiple rows . . . . . 30 Body elongated, scutum flattened. Setulae on scutum dark and evenly dispersed. Body sometimes with light metallic luster . . . . . Rhopalopterum Duda p. p. (p. 141) Body short, scutum convex. Setulae on scutum either yellow or arranged in double rows . . . . . . . . . . . . . . . . . . . . . . . . . 31 Setulae on scutum yellow, abundant and dispersed . . . . . . . . Conioscinella Duda p. p. (p. 78) Setulae on scutum black, arranged in double rows . . . . . . . . Microcercis Beschovski (p. 122) R2+3 closely following and fused with R1 and C (Figs 42, 566). Apical scutellar setae placed on disc of scutellum (Fig. 565) . . . . . . . . . . . . Camarota Meigen (p. 154) R2+3 well separated from R1 and C (Fig. 41). Apical scutellar setae, if developed, placed on margin of scutellum . . . . . . 33 One to three orbital bristles distinctly longer than remaining ones (Figs 732, 764, 918). Body always yellow with black stripes that may become fused on scutum . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 Orbital bristles all about same size (Figs 569, 598, 602). Body yellow with black scutal stripes or wholly black . . . . . . . 35

55

34 (33)

–

35 (33)

– 36 (35)

–

37 (36) – 38 (37)

–

39 (37)

–

40 (39)

–

56

Crossveins of wing approximated, their separation not greater than length of dm-cu (Fig. 44). Small species . . . . Pseudopachychaeta Strobl (p. 223) Crossveins more separated from each other than length of dm-cu (Fig. 766) Larger species . . . . . . . . . . . . . Lasiosina Becker (p. 196) Hind legs with femora greatly enlarged and tibiae usually curved (Figs 5, 904) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 Hind legs without enlarged femora . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 Veins R1 and R2+3 straight (Fig. 902). First flagellomere much longer than deep (Figs 900, 907). Frons strongly produced. Body large, reddish yellow (Fig. 7a) . . . . . . . . Platycephala Fallén (p. 219) Veins R1 and R2+3 curved towards C. First flagellomere barely longer than deep. Frons weakly produced. Body smaller, greenish . . . . . . . . . . . Meromyza Meigen (p. 200) Head triangular in side view (Figs 745, 953) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 Head rounded or rectangular in side view (Figs 570, 591, 599, 728) . . . . . . . 39 Body grey-dusted, densely covered with long setae (Figs 952, 953). Anepisternum and anepimeron naked . . . . . . . . . Trichieurina Meigen (p. 230) Body brownish-yellow with only short setae. Anepisternum and anepimeron setulose . . . . . . . Eurina Meigen (p. 193) Crossveins r-m and dm-cu approximated, their separation not greater than length of dm-cu (Figs 43, 44, 729) . . . 40 Crossveins r-m and dm-cu more separated, their distance from each other greater than length of dm-cu . . . . . . . . 41 A row of setulae on ocellar triangle inside its lateral margin (Fig. 728). Surface of scutum rugose and shining . . . . . . . . . . . . . . Diplotoxa Loew (p. 190) A row of setulae on frons outside lateral margin of ocellar triangle (Fig. 732). Sur-

41 (39)

–

42 (41)

– 43 (42) –

44 (41)

–

45 (44)

– 46 (45) –

47 (45) – 48 (47)

–

face of scutum smooth, evenly dusted . . . . . . Diplotoxoides Andersson (p. 190) Ocellar triangle with one or more rows of setulae within side margins (Figs 705, 930, 939, 944, 948) . . . . . . . . . . . . . . . . 42 No setulae within side margins of ocellar triangle. Setulae present on frons outside ocellar triangle (Figs 569, 752) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 Scutum entirely yellow with black or red longitudinal stripes (Fig. 7b). Scutellum flattened, with apical setulae approximated . . . . . . . Thaumatomyia Zenker (p. 226) Scutum entirely black . . . . . . . . . . . . . 43 First flagellomere rounded apically . . . . . . . Neohaplegis Beschovski (p. 218) First flagellomere acuminate apically (Figs 706, 707) . . . . . . . . . . . . . Cryptonevra Lioy (p. 186) Arista shorter than remaining antenna, nearly bare (Fig. 754). Body setulae white . . . . . . . . . . . . . . Eutropha Loew (p. 194) Arista as long as or longer than remaining antenna. Body setulae usually black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45 First flagellomere longer than deep (Figs 20, 24, 570, 893, 894). Arista white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46 First flagellomere only as long as deep. Arista black or white . . . . . . . . . . . . . . 47 Scutum black, rugose . . . . . . . . . . Centorisoma Becker (p. 155) Scutum yellow, smooth, with five dull black stripes, lateral stripes with a velvety black spot anteriorly . . . . . . . . Parectecephala Becker (p. 218) Scutum yellow with black or red longitudinal stripes . . . . . . . . . . . . . . . . . . . . . 48 Scutum black, at most postpronota and notopleura partly yellow . . . . . . . . . . . 49 Genae linear (Figs 697, 700). Male genitalia: pre- and postgonites in line (Figs 699, 702) . . . Chloropsina Becker (p. 185) Genae wider, at least half as wide as first flagellomere (Figs 591, 595, 599,

49 (47) – 50 (49) – 51 (49)

–

603, 608). Male genitalia: postgonites situated outside of pregonites (Fig. 77) . . . . . . . . . Chlorops Meigen p. p. (p. 159) Surface of scutum smooth . . . . . . . . . 50 Surface of scutum rugose . . . . . . . . . . . 51 Vibrissal angle acute (Figs 17, 779) . . . . . . . . . . . . Melanum Becker (p. 200) Vibrissal angle obtuse (Figs 616, 619, 623, 639) . . . . . Chlorops Meigen p. p. (p. 159) Ocellar triangle reaching anterior margin of frons (Fig. 737). Middle tibiae with small apical spur . . . . . . . Epichlorops Zetterstedt (p. 192) Ocellar triangle not reaching anterior margin of frons. Middle tibiae with strong apical spur . . . . . . . . . . . . . . Cetema Hendel (p. 155)

52 (1)

–

53 (52)

–

oc setae procurved, divergent. Body fuscous yellow with black scutal stripes and black marks on pleura . . . . . . . . . Lasiosina Becker p. p. (p. 196) oc setae recurved or upright, convergent (Fig. 270). Body black with grey dust or thorax red and abdomen black . . . . . 53 Body black, grey-dusted. Scutellum without warts at base of marginal setae (Figs 238, 239) . . . . . . . . Conioscinella Duda p. p. (p. 78) Thorax red, abdomen black with tergites 1–2 extended. Scutellum with warts at base of marginal setae (as Fig. 276) . . . . . . . . . . Elachiptera Lioy p. p. (p. 90)

Preliminary key to genera for 3rd instar larvae Larval descriptions exist for only a small number of genera and species. The present key to genera, intended for mature (3rd instar) larvae is therefore very preliminarily. It can be used for puparia as well, as many larval characters remain intact on puparia: form of posterior end, spiracular zone of abdominal segments, cephalopharyngeal skeleton and anterior spiracles. 1

–

2 (1)

Larvae large, up to 8–12 mm, rather thick and white. First or three first body segments and last segment dorsally sclerotized and blackish (Figs 398, 403). Posterior spiracles sessile. Larvae formings galls on Phragmites australis (Figs 106, 107) . . . . . . . . . . . Lipara Meigen (p. 118) Larvae smaller and narrower, of white, yellowish or greenish colour, without dorsal sclerotisations (Figs 92, 185). Posterior spiracles sessile or born on posterior processes (Figs 454, 909, 815, 915, 963, 970, 973) . . . . . . . . . . . . . . . . . . . . . 2 Hind spiracles trilobate, sometimes more or less sclerotized (Figs 956, 957, 962) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

–

3 (2)

– 4 (3)

–

Hind spiracles not trilobate, sessile or born on posterior processes (Figs 454, 909, 915, 970, 973) . . . . . . . . . . . . . . . . . 5 Facial mask with numerous nearly parallel ridges. Anterior spiracles of rosette type with 10–12 buds (Fig. 958). Branched hairs on hind spiracles absent. All three thoracic segments without spicules, abdominal segments with many spicules on creeping welts (Fig. 959). Larvae in shoots of grasses . . . . . . . . . . . . Camarota Meigen (p. 154) Facial mask with few ridges or ridges practically absent (Fig. 961) . . . . . . . . . 4 Facial mask smooth. Cephalic segment large. Distal segment of antennae larger than basal one and round (Fig. 961). Three thoracic segments narrow and long (Fig. 960). Larvae among roots of plants . . . . . . . Thaumatomyia Zenker (p. 226) Facial mask simple, with maximally four crescent ridges. Larvae in shoots of grasses . . . . . . . Conioscinella p. p. (p. 78) 57

5 (2)

–

6 (5)

–

7 (6) –

8 (7)

–

9 (7)

58

Hypopharyngial and tentoropharyngeal sclerites articulated. Mandibles more or less slender, without or with accessory teeth on ventral side (Figs 96, 456, 964, 968, 971). Facial mask usually with complex pattern of ridges. Hind spiracles born on posterior processes (Figs 717, 725, 973, 979) . . . . . . . . . . . . . . . . . . . . . 6 Hypopharyngial and tentoropharyngeal sclerites fused (Figs 97, 911, 916, 997, 1003, 1010). Mandibles very massive, always with accessory teeth on ventral side. Facial mask usually with more simple pattern. Hind spiracles sessile or born on posterior processes . . . . . . . . 14 Mandible slender, without accessory teeth on ventral side (Fig. 964). Facial mask consists of several rows of parallel, often dentate ridges (Fig. 93) . . . . . . . . 7 Mandible rather massive and with accessory, sometimes very small teeth on ventral surface. Facial mask contains ridges uniting into cells (Figs 94, 449, 455) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Anus surrounded by coarse spicules (Fig. 725) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Anal area with fewer spicules above and/or below anus, or without spicules .................................9 Dental sclerite arched (as Fig. 97). 2nd and 3rd thoracic segments without spicules. Creeping welts covered with coarse and thin spicules, but anterior row interrupted in the middle. Larvae in shoots of grasses . . . . . . . Lasiosina Becker (p. 196) Dental sclerite small and of different form, or absent (Figs 712, 719). 2nd and 3rd thoracic segments without spicules. Creeping welts covered with coarse and thin spicules, anterior row usually entire (Figs 715, 716, 721, 722). Larvae in shoots of Pragmites australis, often in Lipara galls . . . . . . . . . Cryptonevra Lioy (p. 186) Anal area and thoracic segments without spicules. Spicules on ventral side of

– 10 (9)

–

11 (10)

–

12 (6)

–

13 (12)

abdominal segments large and brown (Figs 93, 964, 965). Larvae in different media . . . . . . . Polyodaspis Duda (p. 138) Some spicules present above and/or below anus . . . . . . . . . . . . . . . . . . . . . . . . 10 Larvae in shoots of grasses and other plants, usually monocots . . . . . . . . Aphanotrigonum Duda (p. 61), Elachiptera Macquart (p. 90), Incertella Sabrosky (p. 109) Larvae in rotten wood, bracket fungi, coniferous cones or in egg sacks of spiders . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Spicules on ventral side of abdominal segments large, acute and wide basally, arranged in two rows on ventral side of 2nd to 8th abdominal segments (Fig. 966). Larvae found in egg sacks of spiders . . . . Pseudogaurax Malloch (p. 139) Spicules on ventral side of abdominal segments smaller and apically obtuse, arranged in several rows on ventral side of 2nd to 8th abdominal segments (Figs 969, 972). Larvae found in rotten wood, bracket fungi, and coniferous cones . . . . Gaurax Loew (p. 101), Hapleginella Duda (p. 108) First three abdominal spicular zones consist ventrally of confluent rows of finely serrate striae; on the following abdominal segments these transverse fields of striae are accompanied in front by rows of small spicules (Fig. 976). Abdomen deeply divided posteriorly into two large spiracular lobes (Fig. 977) . . . . . . . . . . . . . . Cetema Hendel (p. 155) All abdominal spicular zones consist ventrally of several transverse rows of spicules of same or different size . . . . 13 Facial mask: frontal ridges between maxillary papillae absent; genal ridges forming only 3–4 plain cells (Figs 449, 455). Spicular fields or individual rows on 2nd and 3rd thoracic segments usually present, if absent then creeping welts on abdominal segments all armed with

–

14 (5)

– 15 (14)

–

16 (14)

coarse spicules (Figs 450–453) . . . . . . . . . . . . . Oscinella Becker (p. 124) Facial mask more complex: about 4 rows of conspicuous, transverse densely dentate frontal ridges between maxillary papillae; genal ridges forming more than 4 finely dentate cells (Fig. 978). Spicular zones on 2nd and 3rd thoracic segments absent (Fig. 980) . . . . . . . . . . . . Conioscinella Duda (p. 78) Spicular zones absent ventrally and dorsally on abdominal segments, or only with a single row of spicules . . . . . . . . 15 Spicular zones present on both thoracic and abdominal segments . . . . . . . . . . 16 Larvae slightly depressed dorsoventrally, up to 4 mm long, pale yellow, in seeds of grasses . . . . . . . . . Dicraeus Loew (p. 85) Larvae not depressed, white, in stems of Phragmites australis, both intact and stems with Lipara galls or infected by Platychephala planifrons; also found in other grasses . . . . . . . Calamoncosis Enderlein (p. 68) Spicular zone of abdominal segments consists ventrally of numerous homogenous, often slightly confluent ridges without well defined spicules present (Figs 987, 1009, 1013) . . . . . . . . . . . . . . 17

–

17 (16)

–

18 (16)

–

Spicular zone of abdominal segments at least to some extent consisting of well defined spicules (Figs 909, 915) . . . . . 18 Antennae widely separated, situated anterolaterally on the head lateral to maxillary papillae. Maxillary papillae completely surrounded by a thick dark brown ring (Figs 989, 992, 996, 999, 1002). Dentate sclerite absent or very small. Caudal end of body variable. Larvae white or yellowish, in shoots of grasses . . . . . . . Chlorops Meigen (p. 159) Antennae situated in front of maxillary papillae. Maxillary papillae not surrounded by thickened ring (Fig. 1008). Dentate sclerite arched (Figs 1010, 1011, 1014). Caudal end of body dorsally with a furrow (Fig. 1012). Larvae pale green . . . . . . . . . . . Meromyza Meigen (p. 200) Posterior spiracles born on processes from caudal end of body (Fig. 981). Larvae yellowish, in inflorescences of Cyperaceae . . . . Pseudopachychaeta Strobl (p. 223) Posterior spiracles sessile (Figs 909, 915). Larvae white, stocky, living in shoots of Phragmites australis . . . . . . . . . . . Platycephala Fallén (p. 219)

Subfamily RHODESIELLINAE Genus Aspistyla Duda, 1933 Macrostyla Lioy, 1864: 1126 (preocc. Winnertz, 1846). Type species: Chlorops plumiger Meigen, 1830 by monotypy. Aspistyla Duda, 1933: 224, as replacement name for Macrostyla Lioy, 1864. Type species: Chlorops plumiger Meigen, 1830, automatic.

lomere reniform, wider than long. Scutum shining, scutellum with apical and lateral setae on small tubercles. Tibial organ indistinct. Femoral organ formed by a group of small warts with short thin setae, presents in both sexes and in male on hind femora as well. Male genitalia: epandrium with terminal surstyli; pregonites with strong setae; distiphallus long and slender, curved.

Black species with large naked eyes, haired arista and linear genae. Ocellar triangle large shining, reaching front margin of frons. Setae of head long, ocellar setae procurved and divergent. First flagel-

Distribution. – Worldwide, most diverse in the Old World tropics with many species in the Oriental 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 (Meigen, 1830) Figs 46, 47, 115–119 Chlorops plumiger Meigen, 1830: 153; Meigen’s plate 229: 12 (Morge, 1976b). Lectotype in NHMW (Nartshuk, 1997a). Body black, shining. Ocellar triangle large shining, reaching front margin of frons, with long setulae along sides. Setae of head long, ocellar setae procurved and divergent. Eyes large, naked. Genae linear. First flagellomere wider than long with black long-haired arista. Palpi brown. Scutum and pleura shining, scutellum triangular, apical and subapical

60

setae long, standing on small tubercles, apical setae longer than scutellum. Legs with femora black, tibiae and tarsi yellow. Distal segment of tarsi darkened. Halteres yellow. Body length nearly 2 mm. Distribution. – Rare. Denmark: EJ: Svejbæk, 19.vii.1964 (V. Koch), 1 female. Sweden: SK, ÖL. Finland: Al. Not found in Norway or Russian parts of Fennoscandia. Trans-Palearctic species, known from the British Islands to the Far East of Russia. Biology. – Adults from end of June to September, in broad-leaf woods and bushes. It has been reared from a mushroom (Xerocomus chrysenteron) in Tatarstan, Russia (Khalidov, 1984; Yakovlev, 1994).

Subfamily OSCINELLINAE Genus Aphanotrigonum Duda, 1932 Aphanotrigonum Duda, 1932: 35. Type species: Chlorops trilineatus Meigen, 1830 by original designation. Small black, densely dusted species with mainly dusted pleura. Ocellar triangle small and dusted. Genae with produced vibrissal angle. poc short, upright and convergent. Face with distinct but short carina. First flagellomere small and thick basally. npl 1 + 2. Tibial organ oval. Femoral organ consists of two rows of warts with short thin setulae. Male genitalia: hypandrium closed. Distribution. – Holarctic, New Zealand, Oriental Region.

Key to species of Aphanotrigonum Duda 1 – 2

–

3

– 4

–

5

Wings shorter than abdomen. Male genitalia Fig. 120 . . . . . . . A. brachypterum (Zetterstedt) Wings longer than abdomen . . . . . . . . . . . . . 2 Scutellum with many marginal setae, apical setae short and not much wider apart than poc setae (Figs 135, 143) . . . . . . . . . . . . . . . . . . . . . 3 Scutellum with no more than two pairs of marginal setae, apical setae long and wider apart than poc setae (Fig. 122) . . . . . . . . . . . . 5 Three or five brown stripes on scutum and scutellum. Male genitalia Figs 146–148 . . . . . . . . . . . . . . . . . . . A. trilineatum (Meigen) Scutum evenly grey without brown stripes ......................................4 Male genitalia: epandrium large; cerci not tapering, with several longish setae; surstyli broad (Fig. 131) . . . . . . . . . . . . A. hungaricum Dely-Draskovits Male genitalia: epandrium small; cerci slightly tapering, with one long seta; surstyli narrow (Figs 136–138) . . . . . . A. nigripes (Zetterstedt) All body black except first flagellomere below. Scutellum rounded, apical setae rather close together. Two posterior npl setae. Anepister-

–

6 –

7 –

num shining. Male genitalia with strongly tapering and straight surstyli (Figs 139–141) . . . . . . . . . . . . . . . . . . . . . A. norrbotticum sp. n. Head and legs partly yellow. Scutellum trapezoid with apical setae wider apart (Fig. 127). One posterior npl seta. Pleura mainly dusted. Male genitalia with surstyli of different shape and directed mesad (Figs 123–125, 128, 129, 132, 133) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Hind tibiae with a small but distinct black spur. Prothoracic episternum shining . . . . . . 7 Hind tibiae without a spur. Prothoracic episternum dull. Male genitalia Figs 132, 133 . . . . . . . . . . . . . . . . . . . . . . . . . A. inerme Collin All coxae and femora dark. Male genitalia Figs 128–130 . . . . . . . . . . . . . . A. femorellum Collin At least anterior and middle femora partly yellow. Male genitalia Figs 123–125 . . . . . . . . . . . . . . . . A. cinctellum (Zetterstedt)

2. Aphanotrigonum brachypterum (Zetterstedt, 1848) Fig. 120 Oscinis brachyptera Zetterstedt, 1848: 2660. Holotype in ZMLU (Andersson, 1966; DelyDraskovits, 1981). Aphanotrigonum trilineatum var. microptera Duda, 1932: 36. Lectotype male by present designation, and paralectotypes 4 males and 6 females all from Denmark: NEZ: Tibirke, 19.x.1914 (W. Lundbeck), in ZMUC. Duda (1932) misspelled Tibirke as “Fibicke”. Syn. n. Body black, densely grey-dusted. Wings shortened, though extend nearly to the end of abdomen. Ocellar triangle dull, reaching halfway to front margin of frons. Frons brownish yellow in front. Scutellum with 8–10 small marginal setae, apical setae approximated. Pleura dusted. Femora mainly black. Body length 1.7–1.8 mm. Distribution. – Rare species. Denmark: NWJ: Hansted Reservatet, 08.vi.1960 (ZMUC Exp.), 1 female; NEZ: Without locality (R.W. Schlick), 1 male; Tibirke, 19.x.1914 (Krüger), 5 males, 6 females; B: 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 (holotype) (examined by Dely-Draskovits, 1981); Kullaberg, 16.viii.1966 (H. Andersson), 1 female. European species also known from Germany and Hungary. Note. – Andersson (1966) and Ismay (1980b) considered A. brachypterum as a short-winged form of A. nigripes (Zetterstedt), but Dely-Draskovits (1981), after examination of the type specimens, treated it as a separate species based on differences in the male genitalia.

3. Aphanotrigonum cinctellum (Zetterstedt, 1848) Figs 121–125 Oscinis cinctella Zetterstedt, 1848: 2659. Holotype male in ZMLU (Andersson, 1966). Oscinis fasciella Zetterstedt, 1855: 4808. Holotype female in ZMLU (Andersson, 1966). Syn. n. 62

Note. – Examination of a long series of specimens from different localities revealed a sexual variation in colour of legs: femora dark in males, yellow in females. Therefore we consider Oscinis cinctella Zetterstedt and O. fasciella Zetterstedt as male and female of the same species. Body mainly black, lightly grey-dusted. Frons yellow anteriorly; ocellar triangle black with grey dust, reaching halfway to front margin of frons, behind reaching eye margin. Face and genae yellow, vibrissal angle produced. Palpi yellow. Antennae yellow, first flagellomere darkened. Scutum grey, scutellum short, rectangular. Apical setae longer than scutellum, wide apart. Pleura dusted. npl 1 + 1. Base of abdomen yellow, remaining tergites with black bands. Halteres yellow. Legs in female extensively yellow, in male femora dark; hind tibiae with small dark spur. Body length 1.0–1.3 mm.

Distribution. – Denmark: SJ, NEJ, F, NEZ. Sweden: SK, SM, Öl, GO, BO, SÖ. Norway: HEn: Isterfossen. Finland: Al, Ab, Om, Ob. Russia: Kr: Petrozavodsk. Widely distributed in Palearctic, eastwards to Yakutia, East Siberia, southwards to Israel. Biology. – Adults in July to early September in dry habitats, mainly sandy beaches. Tschirnhaus (1981) recorded as host plant Puccinella maritima and Bährmann & Weipert (1989) reared the species from Elytrigia and Puccinella.

4. Aphanotrigonum femorellum Collin, 1946 Figs 126–130 Aphanotrigonum femorella Collin, 1946: 127. Lectotype male (Dely-Draskovits, 1981), paralectotypes 3 males, 9 females in OXUM (Pont, 1995). Similar to A. cinctellum in having npl setae 1 + 1, apical scutellar setae wide apart, hind tibiae with small dark spur. Main differences: first flagellomere larger and more extensively brownish on outer side, abdomen darker, pale hind margins on tergites 3–5 narrower. Legs much darker, all femora dark, tibiae with dark band, tarsi brownish. Body length 1.5–1.75 mm. Distribution. – Rare species. Denmark: WJ: Skallingen, 17.vii.1982 (E. Bro Larsen); F: Langeland, Keldsnor, 6.viii.1975 (Lyneborg, Martin & Michelsen). Sweden: SK: Foteviken, 25.viii.1960, coastal meadow, 28 specimens (H. Andersson). Not found in Norway, Finland and adjacent territories of Russia. Widely distributed but rare Palearctic species, known from Europe to Mongolia, and North Africa. Biology. – Adults in July and August in dry meadows and on sandy beaches. Tschirnhaus (1981) recorded as host plants Puccinella maritima and Festuca rubra.

5. Aphanotrinonum hungaricum Dely-Draskovits, 1981 Fig. 131 Aphanotrinonum hungaricum Dely-Draskovits, 1981: 131. Holotype in HNHM. Similar to A. nigripes. Body black, densely greydusted. Frons black, fuscous yellow in front. Ocellar triangle dull grey, reaching halfway to front margin of frons. A few tiny setulae arising from minute

puncture on sides of ocellar triangle. orb numerous and very short. Genae as wide as first flagellomere, yellow in front, Vibrissal angle produced. First flagellomere dark brown, yellowish at base. Scutum and pleura dusted bluish grey. Scutum and scutellum punctate. Scutellum with approximated apical setae and 4–5 subapical setae on each side. npl 1 + 2; postalar seta rather long. Abdomen broad and flat with lateral parts of tergites on ventral side of abdomen, greyish brown. Male abdominal sternites 3 and 4 wider than long. Halteres yellow. Legs mainly black. Male genitalia: epandrium large, cerci broad, surstyli with thick setulae at base. Body length 2.0 mm. Distribution. – Denmark: WJ, F. Sweden: SK. Finland: Al: Sund (R. Frey); Ab: Nyström; N: Tvärminne. Not found in Norway or Russian parts of Fennoscandia. Recorded from Europe and Afghanistan.

6. Aphanotrigonum inerme Collin, 1946 Figs 132, 133 Aphanotrigonum inerme Collin, 1946: 128. Lectotype male (Dely-Draskovits, 1981) and paralectotypes 12 males and 9 females in OXUM (Pont, 1995). Similar to A. femorella, but without an anteroventral spur on hind tibiae, prothoracic episternum dusted, abdomen without pale bands on hind margins of tergites except on female tergite 5. npl 1 + 1. Apical scutellar setae wide apart. Body length 1.5– 1.8 mm. Distribution. – Rare species. Denmark: WJ: Fanø, 30.v.1919, 10.vii.1921 (T. Mortensen), 2 males, 2 females; Ringkøbing, 24.vii.1951 (K.O. Leth), 1 female; NEZ: no further data (R.C. Stæger), 1 female. Sweden: SK: Revinge, 26.vi.1985 (H. Andersson), 1 female. Not found in Norway, Finland or Russian parts of Fennoscandia. European species. Biology. – Adults in May to July. In Bulgaria the species occurs on sand dunes with Phragmites australis (Beschovski, 1982).

7. Aphanotrigonum nigripes (Zetterstedt, 1848) Figs 29, 134–138 63

Figs 126–133. Aphanotrigonum femorellum Collin: 126: head, lateral view. 127: scutellum. 128: hypopygium, dorsal view. 129: hypopygium, 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

Oscinis nigripes Zetterstedt, 1848: 2652. Holotype in ZMLU (Andersson, 1966). Body black, densely grey-dusted. Frons black, fuscous yellow in front. Ocellar triangle dull grey, reaching halfway to front margin of frons. A few tiny setulae arising from minute punctures on sides of ocellar triangle. orb numerous and very short. Genae as wide as first flagellomere, yellow in front, vibrissal angle produced. First flagellomere dark brown, yellowish at base. Scutum and pleura dusted bluish grey. Scutum and scutellum punctate. Scutellum with approximated apical setae and 4–5 subapical setae on each side. npl 1 + 2 and a rather long postalar seta. Abdomen broad and flat with lateral parts of tergites on ventral side of abdomen, greyish brown. Halteres yellow. Legs mainly black. Body length 1.5–2.0 mm. Distribution. – Denmark: EJ, NWJ, NEZ, B. Sweden: north to NB and LU. Norway: Ø, AK, HEn, Bø, AAy, Ry. Finland: north to Ob and Ks. Russia: Vib, Kr. Eurasian species, known from Europe to Yakutia, East Siberia and Mongolia. Biology. – Adults from May to October, hibernate. Common in wet meadows, bogs and marshes. Larvae live in shoots of different grasses including Calamagrostis epigeios, Leymus arenarius, Avena, Puccinella maritima (Uffen & Chandler, 1978; Tschirnhaus, 1981). Collected on agricultural grasses in Denmark (Nielsen & Nielsen, 1984). Bread from stems of wheat and oats in Bulgaria (Krusteva & Beschovski, 2000, 2001). Kibric et al. (1999) listed the species as tyrphophilous. Adults have been found in nest of Microtus sp. (Collin, 1939).

8. Aphanotrigonum norrbotticum Nartshuk & Andersson, sp. nov.

dusted. It is characterized by the wholly black frons; antennae separated by very thin facial carina; pleura partly shining (contrary to all other species of the genus); male genitalia with long surstyli. The dark body colour, partly shining pleura and structure of male genitalia provide the best diagnostic characters of the new species. Description. – Body black, darkly dusted, except for shining parts of pleura. Only face and ventral part of first flagellomere fuscous yellowish. Frons a little longer than wide, dull. Ocellar triangle very small, nearly reaching halfway to front margin of frons. Setae of head very short. Genae distinctly produced beyond eye, wider than first flagellomere, brownish with black lower margin. First flagellomere very small, rounded, reddish below. Antennae separated by very thin facial carina. Proboscis rather long. Palpi darkened. Scutum nearly square with distinct puncture line in the middle and a pair of small depressions before scutellum. Scutellum flat, rounded, apical setae not longer than scutellum, two small subapical setae on each side. Distance between apical setae not longer than 1.5 times distance between poc. Pleura partly shining, only dusted on anepimeron and upper part of katepisternum. npl 1 + 2. Halteres yellow. Abdomen and legs entirely black. No spur on hind tibiae. Male genitalia: cerci small and wide apart; surstyli rather long, strongly tapering and nearly parallel; hypandrium closed, rather massive; 2 trichoid sensillae distinct in front of hypandrium. Body length 1.2– 1.25 mm. Etymology. – The species is named after the type locality.

9. Aphanotrigonum trilineatum (Meigen, 1830)

Figs 139–141

Figs 10, 11, 48, 142–148

Holotype, male: Sweden: Norrbotten: Högsön, Råneå, 01.vii.1972 (H. Andersson). Paratypes: 20 males, 31 females, same locality, 01, 06 and 07.vii.1972 (H. Andersson); 2 males (in alcohol), Lule Lapmark: Messaure, 08–15 and 22–29.vii.1974 (K. Müller). Holotype and 49 paratypes in ZMLU, 4 paratypes in ZISP.

Chlorops trilineatus Meigen, 1830: 162; Meigen’s plate 210: 11 (Morge, 1976b). Lectotype in NHMW (Nartshuk, 1997a). Aphanotrigonum beschovskii Dely-Draskovits, 1978: 128.

Diffferential diagnosis. – In general appearance most similar to species of the Aphanotrigonum trilineatum species group: genae with produced vibrissal angle, ocellar triangle small, dusted, frons

Body black, densely bluish grey-dusted. Frons dark brown behind and yellowish in front, covered with short pale setulae on minute black tubercles. Ocellar triangle poorly demarcated at sides, nearly reaching middle of frons, black, grey-dusted with 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: hypopygium, 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 produced. Antennae yellow, first flagellomere black above. Head setae black, short and stout. Scutum densely covered with bluish grey dust, with 3–5 brown longitudinal stripes along acrostichal and dorsocentral lines and sometimes on sides of scutum and scutellum. Scutellum slightly wider than long with short stout apical setae and a pair of subapical setae. Pleura dusted. npl 1(2) + 2. Legs with 66

femora all black, tibiae black with both ends yellowish. Body length 1.5–2.0 mm. Distribution. – Very common. Denmark: EJ, WJ, NEZ. Sweden: north to LU and NB. Norway: Ø, AK, HEn, HEs, Bø, VE, AAy. Finland: north to Li. Russia: Kr, Lr. Probably Holarctic, as Beaulier & Wheeler (2001) recorded the species with some reservation from

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, widespread species ranging from the British Isles to Japan. Biology. – Adults from March to October on bogs and other wetlands, also in meadows and agricultural fields. Hibernates as adults. Kallio (1950) listed it as a pest of cereals in Finland. Collected on agricultural fields in Denmark (Nielsen & Nielsen, 1984). Hackman (1963) found adults in borrows of voles (Microtus, Clethrionomys) in Finland.

It has been reared from stems of cereals (wheat, ray, barley, oats) and foddergrass. Johanssen (1960) reared it from shoots of cereals in Sweden, Rygg (1966a) from seedlings of oats and wheat in Norway. The following wild grasses and sedges are also known as host plants: Festuca pratensis, Calamagrostis epigeios, Leymus arenarius, Poa pratensis and Eriophorum vaginatum (Kreiter, 1928; Nartshuk, 1972; Uffen & Chandler, 1978; Krusteva & Beschovski, 2000, 2001). 67

Some larval structures were illustrated by Kreiter (1928) and Nartshuk (1987).

3 –

Genus Calamoncosis Enderlein, 1911 Calamoncosis Enderlein, 1911a: 235. Type species: Lipara rufijitarsis auct. nec Loew, 1858 (= Lipara minima Strobl, 1893) by original designation. Rhaphiopyga Nartshuk, 1971: 292. Type species: Calamoncosis glyceriae Nartshuk, 1958 by original designation. Black species with hairy eyes, surface of ocellar triangle shining with many fine setulae situated in punctures. Face concave with facial carina being narrowly triangular above and very thin below. Genae usually narrower than first flagellomere, vibrissal angle more or less produced. poc upright, convergent. First flagellomere rounded rectangular, wider than long; arista short. Scutum entirely polished, convex, covered with small setulae. Scutellum sometimes with scutellar setae on small or long tubercles. npl 1 + 1(2). Tibial organ large, oval. Femoral organ with numerous warts on a rounded area. Male genitalia: cerci basally fused, surstyli simple; postgonite relatively large and broad. The genus is divided into two subgenera: Calamoncosis s. str. has female cerci simple and soft, covered with numerous hairs; Rhaphiopyga Nartshuk, 1971 has female cerci laterally compressed, faintly sclerotized and sometimes elongated. Distribution. – Probably worldwide.

Key to species of Calamoncosis Enderlein 1

–

2

–

68

Ocellar triangle reaching 0.75–1.0 of distance to front margin of frons (Figs 159, 170, 192). 2–3 posterior npl setae . . . . . . . . . . . . . . . . . . . . . . 2 Ocellar triangle shorter, at most reaching slightly more than halfway to front margin of frons (Figs 149, 177, 212), if longer then only 1 posterior npl seta . . . . . . . . . . . . . . . . . . . . . . . 5 Frons except ocellar triangle dull. Notopleura shining. Male genitalia Figs 195–197 . . . . . . . . . . . . . . . . . . . . . . . . C. halterata sp. n. Frons covered with silky hairs. Notopleura dusted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

4

–

5

– 6

–

Scutellum longer than wide (Figs 161, 201). Wings and wing veins not whitish . . . . . . . . . 4 Scutellum as long as wide or shorter (Fig. 172). Wings and veins M whitish . . . . . . . . . . . . . . . . . . . . . . C. duinensis (Strobl) Scutellum long, triangular, basally flattened, with apical and several subapical setae on distinct tubercles (Fig. 161). Male genitalia Figs 152, 163 . . . . . . . . . . . . . . . . C. aspistylina Duda Scutellum shorter, convex at base, with apical and subapical setae on very small tubercles (Fig. 201). Male genitalia Figs 203, 204 . . . . . . . . . . . . . . . . . . . . . . . C. minima (Strobl) Only one posterior npl seta. Halteres and palpi black. Male genitalia Figs 152, 153 . . . . . . . . . . . . . . . . . . . . . . . C. aprica (Meigen) Two posterior npl setae. Halteres and palpi yellow or palpi more or less darkened . . . . . 6 Vibrissal angle distinctly projecting beyond level of eyes (Fig. 178). Setae and setulae on thorax black. Male genitalia Figs 180–182. Ovipositor laterally compressed and cerci sclerotized (Figs 183, 184) . . C. glyceriae Nartshuk Vibrissal angle obtuse and not projected (Fig. 213). Setae and setulae on thorax whitish. Ovipositor not compressed and cerci not sclerotized . . . . . . . . . . . . C. oscinella (Becker)

Species key to known 3rd instar larvae of Calamoncosis 1

–

2

–

Creeping welts ventrally on abdominal segments without spicules. Larvae (Figs 207–211) in galls of Lipara spp. on Phragmites . . . . . . . . . . . . . . . . . . . . . . . C. minima (Strobl) Creeping welts ventrally on abdominal segments with one or two rows of small spicules (Figs 167, 190) . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Stigmal plate of hind spiracles with 3 short processes and without interstigmatic tufts (Figs 164–169). Larvae in stems of Phalaris arundinaceae . . . . . . . . . . . C. aspistylina Duda Stigmal plate of hind spiracles with 4 processes and interstigmatic tufts; these tufts of-

3

–

ten broken in puparia, but indicated by scars from their bases (Fig. 189) . . . . . . . . . . . . . . . 3 Creeping welts ventrally on abdominal segments with two rows of spicules (Fig. 190). Larvae living gregariously in unsheathed inflorescences of Glyceria spp. . . . . . . . . . . . . . . . . . . . . . C. glyceriae Nartshuk Creeping welts ventrally on abdominal segments only with one short row of spicules (Fig. 158). Larvae in galls formed in stems of Phragmites by Platycephala planifrons . . . . . . . . . . . . . . . . . . . . . . . C. aprica (Meigen)

stem of reed contained from 1 to 52 larvae. Pupation takes place inside the host plant. Adults from a new generation were taken in traps from July 7 to August 11. Grochowska supposed that the species has two generations per year, but the host plant of the summer generation is unknown. Egg and larvae of three instars described by Grochowska (2002).

11. Calamoncosis (Calamoncosis) aspistylina Duda, 1935 Figs 159–169

10. Calamoncosis (Calamoncosis) aprica (Meigen, 1830) Figs 149–158 Chlorops apricus Meigen, 1830: 158; Meigen’s plate 211: 4 (Morge, 1976b). Lectotype in NHMW (Nartshuk, 1997a). Black, shining except for dull frons. Ocellar triangle short, extending 1/2 to 2/3 of distance to front margin of frons, with many setulae. Genae a little wider than first flagellomere. First flagellomere shorter than high, black. Palpi black. Pleura shining. npl 1 + 1. Halteres dark. Female cerci simple. Body length up to 3 mm. Distribution. – Denmark: SJ, EJ, WJ, F, SZ. Sweden: SK, SM, VR. Norway: AK, VE, AAy. Finland: Al, Ab, N, Sb. Not found in Russian parts of Fennoscandia. European species. Biology. – Adults from May to July in wet places. Larvae phytophagous or phytosaprophagous, living as inquilines in stems of Phragmites australis damaged by larvae of Platycephala planifrons or rarely by caterpillars of Archanara geminipuncta (Noctuidae) (Tschirnhaus, 1981; Grochowska, 2002). Uffen & Chandler (1978) recorded as host plant foliage of Draba incana, but this seems to be doubtful. Grochowska (2002) studied the life cycle of the species in Poland. Adults appear in middle of May and lay eggs on stems of Phragmites australis attacked by Platycephala planifrons. Most larvae of C. aprica were found among frass in existing larval feeding tunnels and had their alimentary tract filled with some brown substance, but some larvae were also found feeding on fresh plant tissue. Each

Calamoncosis aspistylina Duda, 1935: 158. Holotype in ZMHU (Wendt, 1975). Body black, shining. Frons longer than wide, covered with silky hairs. Ocellar triangle narrow, extending nearly to front margin of frons, punctate along side margins, median part smooth, shining. Genae as wide as first flagellomere is long. First flagellomere black, shorter than wide, basal antennal segments yellow. Scutum convex and punctate, evenly covered with black setulae. Scutellum elongated. Thick marginal scutellar setae inserted on large tubercles, 3–4 pairs of subapical setae. Pleura shining, anepisternum wrinkled. npl 1 + 2. Halteres black. Female cerci simple. Body length 2.5– 3.0 mm. Distribution. – Known only from southern Sweden: SK and BL. European species. Biology. – Adults from late May to early of August in wet meadows. Larvae phytophagous, live in stem of Phalaris arundinacea (Nartshuk, 1981; Sharipov, 1981). Larvae described by Nartshuk (1981).

12. Calamoncosis (Calamoncosis) duinensis (Strobl, 1909) Figs 170–176 Siphonella duinensis Strobl, 1909: 288. Syntypes 2 females in NMBA (Chvála, 2008). Body black, shining. Ocellar triangle reaching front margin of frons, shining and coarsely punctured 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 darkened, projecting beyond facial margin. Basal segment of antennae yellow, first flagellomere dark brown, broader than long. Scutellum short semicircular widely rounded apically. Apical setae shorter than scutellum. npl 1 + 2. Wing slightly whitish. Halteres with back macula on knob. Female cerci simple. Body length 2.0–2.5 mm. Distribution. – Denmark: WJ, NWJ, F, NEZ. Sweden: SK, HA, SM, VS. Finland: Ab, N. Russia: Vib. Not found in Norway. 70

Widespread Palearctic species, known from Europe eastwards to Japan and in North Africa. Biology. – Adults from May to beginning of September on Common reed (Phragmites australis). Larvae phytophagous, live as inquilines in Lipara galls on the folded leaves.

13. Calamoncosis (Rhaphiopyga) glyceriae Nartshuk, 1958 Figs 1, 177–191

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. Holotype in ZISP. Body black, only antennae and parts of legs yellow. Frons longer than wide, dull. Ocellar triangle reaching halfway to front margin of frons. Frons covered with rather short and thin setulae. Genae brown, narrower than first flagellomere, slightly produced at vibrissal angle. Antennae yellow with small round first flagellomere, arista shortly pubescent and shorter than frons. Palpi rather long, brown. Proboscis slightly elongated. Scutum and scutellum coarsely punctate, scutum shining, notopleura dusted, pleura shining. Apical scutellar setae shorter than scutellum, with one or two pairs of rather thin subapical setae. npl 1 + 2. Femora black except basally and at tips, tibiae yellow with a black band, tarsi yellow. Halteres yellow. Ovipositor laterally compressed with sclerotized cerci. Body length 2.5 mm. 72

Distribution. – Denmark: EJ, NWJ, NEZ. Sweden: SK, VS. Norway: Ø: Gansröd, Fredrikstad. Finland: Om. Russia: Vib, Kr. Eurasian species. Biology. – Adults are very abundant in wet places, near lakes and rivers, from May to September. Larvae phytophagous, feed gregariously in unsheathed inflorescences of Glyceria maxima, G. fluitans, G. aquatica (Collin, 1946, as “C. laminiformis (Becker)”; Nartshuk, 1958). Larvae described by Ismay & Rotheray (1998).

14. Calamoncosis (Calamoncosis) halterata Nartshuk & Andersson, sp. nov. Figs 192–197 Holotype, male: Sweden: Närke: Hjälmarsberg, Örebro, 26.vii.1941 (T. Nyholm). Paratype, female: Swe-

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.

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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 (R. Danielsson). Holotype and paratype in ZMLU. Diffferential diagnosis. – Similar to C. oscinella Becker, but distinguished by longer and narrower frons, longer ocellar triangle, shorter dark setulae on frons, setae and setulae on thorax black (not whitish), genae narrower, and scutellum more coarsely punctated. Description. – Male, female. Body entirely black except for yellow antennae and tarsi of all legs. Frons longer than wide, dull. Ocellar triangle narrow, extending nearly to front margin of frons, on its surface along side margins with two rows of setulae. Frons covered with rather short and thin setulae, 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 as first flagellomere. Antennae yellow with small round first flagellomere, arista shortly pubescent and shorter than frons. Eyes large oval with vertical long axis, shortly pubescent. Palpi short, black. 74

Scutum longer than wide, finely punctated and covered with short dense setulae. Scutellum triangular, longer than wide at base, with very small tubercles at base of apical and subapical setae. Scutellum coarser punctated than scutum and dusted. Apical scutellar setae lost in male, in female only one seta present shorter than scutellum; two or three pairs of rather thin subapical setae. Pleura shining, anepisternum rugose. npl 1 + 2. Wings longer than abdomen, slightly brownish. Cell br not widened. Second sector of costal vein 1.7 times longer than third, third costal sector 1.25 longer than fourth. Halteres with stem and knob black. Legs black except both ends of tibiae and all tarsi yellow. Body length 2.5 mm.

15. Calamoncosis (Calamoncosis) minima (Strob1, 1893) Figs 49, 198–211

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

Lipara minima Strobl, 1893: 225. Holotype male in NMBA (Chvála, 2008). Body black, shining. Ocellar triangle extending nearly to front margin of frons, shining, punctured except on median glabrous area. Genae as wide as first flagellomere. Palpi yellow to dark brown, projecting beyond facial margin. Basal segment of antennae yellow, first flagellomere dark brown, broader than long. Scutum longer than wide. Scutellum elongate triangular, convex basally; apical setae on small tubercles, shorter than scutellum. npl 1 + 2. Wings hyaline. Halteres with back macula on knob. Female cerci simple. Body length 2.5–3.0 mm. Distribution. – Denmark: WJ, LFM, NEZ. Sweden: SK, SM, VG, NB, LU. Norway: AK, HEs, Bø, VE. Finland: north to Om. Russia: Vib. Common and widespread, trans-Palearctic species. Biology. – Adults from April to September on Common reed (Phragmites australis).

Larvae phytophagous, feed on rolled young lives of Phragmites or live as inquilines in galls of Lipara species. Usually several larvae in a gall, feeding on folded leaves above chamber of the Lipara larva (Séguy, 1934; d’Aguillar, 1953; Valkeila, 1956; Nartshuk, 1972a). Larvae described by d’Aguillar (1953) and Grochowska (2002).

16. Calamoncosis (Calamoncosis) oscinella (Becker, 1910) Figs 212–214 Lipara oscinella Becker, 1910: 145. Holotype probably in DEIM (Rohlfien & Edward, 1970). Body black, shining with all setae and setulae white. Frons black, finely punctate and covered with whitish setulae. Ocellar triangle shining, extending slightly more than halfway to front margin 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.)

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Figs 212–214. Calamoncosis oscinella (Becker): 212: head, dorsal view. 213: head, lateral view. 214: scutellum.

wide as first flagellomere, not produced beyond eye margin. Antennae yellow, first flagellomere darkened above. Palpi yellow. Scutum shining, covered with whitish setulae. Notopleura and anepisternum in upper part thinly grey-dusted, remaining parts of pleura shining. Scutellum convex, as wide as long. Apical scutellar setae approximated and short, two pairs shorter subapical ones. Abdomen whitish pubescent. Femora black, tibiae yellow, hind tibiae with black band. Halteres yellow. Body length 2.0–2.3 mm.

eye vs. not so produced; profile of head slightly concave vs. straight; arista pubescent vs. naked; genae no wider than first flagellomere vs. wider than first flagellomere; femoral organ present vs. absent; tergites of abdomen ventrally bent and separated by ridge from dorsal parts vs. normally shaped. Duda’s species meijerei further differs from the genus Conioscinella by the dusted pleura, wide genae, naked arista and eyes and absence of femoral organ. Therefore, we erect a new genus Colliniella for it.

Distribution. – Rare species. Sweden: SK: Skäralid, 09.vi.1965 (H. Andersson), 1 male. Norway: HEs: Abborhøgda, Kongsvinger. Finland: N: Grankulla, 29.v.1918 (R. Frey), 1 female; Espoo (R. Frey), 1 female. Not found in Denmark or Russian parts of Fennoscandia. Known also from Germany.

Etymology. – The genus is named in honour of the famous British dipterist J.E. Collin, who contributed significantly to the knowledge of British Chloropidae. Gender feminine.

Genus Colliniella Nartshuk & Andersson, gen. nov. Type species: Conioscinella meijerei Duda, 1933 by present designation. Conioscinella meijerei was transferred to the genus Aphanotrigonum together with some other species by Collin (1946) because of the dusted pleura. However, Duda’s species species differs significantly from Chlorops trilineatus Meigen, the type species of Aphanotrigonum, in other respects. The main differences between Aphanotrigonum trilineatum (Meigen) and Conioscinella meijerei Duda are: Setae of head and thorax strong and black vs. weak and whitish; vibrissal angle produced beyond

Description. – Body densely dusted. Setae of head 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 halfway to front margin of frons. Eyes oval with diagonal long axis, without hairs. Genae broad. Vibrissal angle not projected. Facial margin not visible in side view. Facial carina triangular above, narrow below, not reaching the facial margin. Arista naked. Proboscis not prolonged. Scutum dusted with few small dispersed setulae, ac and dc setulae not differentiated. Thoracic setae: 1 h, 1 + 2 npl, 1 pa, 1 dc, all weak. Scutellum short, distinctly convex. Two pairs of marginal scutellar present. Legs slender, with a short spur on middle tibia. Tibial organ present, femoral organ absent. Second and third costal sectors of same length. R2+3 , R4+5 and M straight. Abdomen oval, dusted, tergites not bent on ventral side and not divided into three parts. Male genitalia: cerci sep77

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, not divided.

yellow. Male genitalia: gonites narrow and long, projecting distinctly behind the base of phallus. Body length 1.5–2.0 mm.

17. Colliniella meijerei (Duda, 1933), comb. n.

Distribution. – Denmark: NWJ, NEZ, B. Sweden: SK, HA, ÖL, GO. Russia: Vib. Not so far found in Norway and Finland, but expected there. The species is known from Europe and Mongolia.

Figs 215–219 Conioscinella meijerei Duda, 1933: 61. Lectotype in ZMAN (Nartshuk, 2003). Body black, covered with light grey dust. Frons yellow in front, remaining parts of head black. Ocellar triangle grey-dusted, extending 3/4 of distance to front margin of frons. Genae much wider than first flagellomere, vibrissal angle not produced. First flagellomere rounded or slightly angulate, darkened to black with yellowish base. Genae more than 1/3 as wide as eye height. Palpi yellow. All setae of head white. Scutum grey-dusted, with white hardly visible setulae. Pleura mainly dusted. npl 1 + 2. Abdomen grey, with pale setulae. Femora mainly black, tibiae yellow, medially black; middle tibiae with spur, hind tibiae without spur. Halteres 78

Biology. – Adults in late of May to August on sandy beaches with Leymus arenarius. Ardö (1957) reared the species from leaves of Honckenya peploides, Nartshuk (1962a) from ears of Leymus arenarius on the Karelian Isthmus.

Genus Conioscinella Duda, 1929 Conioscinella Duda, 1929: 166, 169. Type species: Oscinella soluta Becker, 1912 by designation of Sabrosky, 1941: 751. Black grey-dusted species with evenly arranged setulae on scutum. Ocellar triangle short, dusted.

Genae with weakly produced vibrissal angle. poc upright, cruciate. First flagellomere rounded oval. Pleura mainly shining. npl 1 + 2. Tibial organ oval. Femoral organ consists of a row of warts. Male genitalia: cerci and surstyli simple; hypandrium usually open, but sometimes closed.

7

Distribution. – Worldwide.

7.

–

Key to species of Conioscinella Duda – 1

– 2 – 3

–

4

–

5

–

6

–

Wings much shorter than abdomen, sometimes absent; even halteres sometimes absent (Fig. 40). Male genitalia Fig. 240 . . . . . . . . . . . . . . . . . . C. zetterstedti Andersson Wings normally developed, longer than abdomen, halteres always present . . . . . . . . . . . 2 Setae and setulae on head and sometimes on scutum yellowish . . . . . . . . . . . . . . . . . . . . . . . 3 Setae and setulae on head and on scutum black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Ocellar triangle and scutum shining, scutum densely covered with yellow setulae. Male genitalia Fig. 227 . . . . . . . . C. gallarum (Duda) Ocellar triangle and scutum dusted and dull. Setae and setulae on scutum black, rarely yellow . . . . . . . . . . . C. sordidella (Zetterstedt) Genae not wider than front tibiae. Vibrissal angle not strongly projected beyond eye margin (Fig. 224) . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Genae wider than fore tibiae, nearly equal to width of first flagellomere. Vibrissal angle strongly projected beyond eye margin . . . . . 6 Femora dark in both sexes, middle tibiae broadly darkened in males, narrowly darkened in females. Frontal setae long. Male genitalia Figs 225, 226 . . . . . . C. frontella (Fallén) Femora dark in males only, middle tibiae yellow in both sexes. Frontal setae short. Male genitalia Figs 232, 233 . . . . . C. mimula Collin Genae as wide as first flagellomere, vibrissal angle distinctly produced and proboscis long. Male genitalia Figs 230, 231 . . . . . . . . . . . . . . . . . . . . . C. messaurea sp. nov. Genae narrower than first flagellomere, vibrissal angle less produced and proboscis shorter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

Setae on head long, orb about 8. Male genitalia: epandrium enlarged (Figs 235–237) . . . . . . . . . . . . . . . . . . . . . . C. tornensis sp. nov. Setae on head short, orb no more than 6. Male genitalia: epandrium normal sized (Figs 220– 222, 228, 229) . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Male genitalia: cerci triangular, wide apart and directed ventrally (Figs 220–222) . . . . . . . . . . . . . . . . . . . . . . . . C. abiskoi sp. nov. Male genitalia: cerci quadrate and directed mesad (Fig. 228) . . . . . . . . . C. livida Nartshuk

Note. – A species group closely related to C. frontella (Fallén) occurs in the northern part of distribution area of the genus. These species are darker coloured than C. frontella and have wider genae. They are rather difficult to separate and examitation of the male genitalia is essential for reliable identification. Three species are described here, but more species may show up in northern Fennoscandia.

18. Conioscinella abiskoi Nartshuk & Andersson, sp. nov. Figs 220–222 Holotype, male: Sweden: Norrbotten: Tärendö, 22.vi.1983 (S.O. Ulefors). Paratypes: 1 female, Sweden: Torne Lapmark: Vittangi, 16.vii.1955, on Carex (H. Andersson); 2 females, Torne Lapmark: Abisko, Betula-forest SW of marble quarry, 22.vii.1983, 2500 m (H. Andersson). Holotype and paratypes in ZMLU. Diffferential diagnosis. – Similar to C. frontella, but darker coloured with more produced vibrissal angle. Male genitalia: cerci smaller and wider apart, hypandrium closed. Description. – Male, female. Frons black, narrowly yellow anteriorly. Ocellar triangle black, dusted, extending nearly to middle of frons. Basal antennal segments brown, reniform first flagellomere yellow, darkened above. Genae narrower than first flagellomere, vibrissal angle produced. Genae and face yellow, except genae black along lower margin. Palpi yellow. Thorax black, scutum dusted and evenly covered with setulae. Scutellum short. Distance between apical setae nearly the same as between apical and 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. Postnotum shining. Abdomen black. Legs black with tibiae at both ends and tarsi yellowish. Halteres yellow. Wings hyaline, venation unremarkable for the genus. Male genitalia: cerci small and rather wide apart, surstyli narrow, hypandrium closed.

19. Conioscinella frontella (Fallén, 1820) Figs 223–226, 978–980 Oscinis frontella Fallén, 1820: 7. Lectotype in ZMLU (Andersson, 1963). Body black, grey-dusted. Frons narrow, yellowish in front. Ocellar triangle black grey-dusted, extending 3/4 of frons. First flagellomere rounded, yel80

low, darkened above. Arista black. Eyes sparsely haired. Palpi yellow. Scutum and scutellum greydusted and evenly covered with setulae. Scutellum rounded, apical setae longer than scutellum. Pleura only dusted above, shining on remaining surface. npl 1 + 2. Abdomen brown, covered with black setulae. Sexual dimorphism in colour of legs: male legs mainly dark with knees, tibiae distally and tarsi except last segment yellowish; female legs paler with front coxae, all trochanteres, front and middle tibiae and hind tibiae basally and distally yellowish. Body length nearly 2.0 mm. Distribution. – Common species. Denmark: EJ, WJ, NEJ, LFM, NWZ, NEZ, B. Sweden: north to LU and NB. Norway: AK, HEs, HEn, Bø, AAy, VAy, Ry, HOi, STi, Nsi. Finland: north to Li. Russia: Vib, Kr, Lr.

Widely distributed Palearctic species, known from Europe to Mongolia and Israel. Biology. – Adults from middle of June to middle of August in meadows. Hackman (1963) found adults in burrows of voles (Microtus, Clethrionomys) in Finland. Larvae phytophagous, living in shoots of many grasses: Elytrigia repens, Anthoxanthum odoratum, Phleum pratense, Dactylis glomerata, Deschampsia caespitosa, D. flexuosa, Festuca ovina, Holcus lanatus (Balachowsky & Mesnil, 1935; Nye, 1958, 1959; Nartshuk, 1962a, 1972a; Karpova, 1972; Tschirnhaus, 1981, 1992; Bährmann & Weipert, 1989) and in oats and barley stems (Krusteva & Beschovski, 1998, 2000). A record from Silene dioica appears unreliable. Larvae hibernate in the basal tillers of grasses and pupate the following spring. Larvae described by Balachowsky & Mesnil (1935) and Nye (1958).

20. Conioscinella gallarum (Duda, 1933) Fig. 227 Tropidoscinis gallarum Duda, 1933: 82. Syntypes in ZMHU (Wendt, 1975). Body black, shining. Frons yellow anteriorly and gradually darkened posteriad. Ocellar triangle black, shining, very sparsely dusted, nearly reaching front margin of frons. Antennae yellow, first flagellomere slightly wider than long, infuscate dorsally. Genae about half as wide as first flagellomere, pale yellow and shining black behind. Scutum shining and evenly covered with short yellowish setulae. Scutellum wider than long, dusted, apical setae longer than scutellum, a pair of subapical setae shorter. Pleura shining. npl 1 + 2. All head and thoracic setae yellow. Halteres yellow. Legs entirely yellow. Body length 1.5–2.0 mm. Distribution. – Denmark: NEZ. Sweden: SK, HA, NÄ, SÖ, VS. Norway: AK: Toien, Oslo, 23.vi.1851 (Siebke). Finland: Ab, N, St. Russia: Vib. Trans-Palearctic species, known from the British Isles to Japan. Biology. – Adults from May to end of July in forests and bushes. Adults have been reared from galls of Andricus and Biorrhiza pallida (Hymenoptera, Cynipidae) on oak, from cones of Larix decidua (Duda, 1933; Nordlander & Grijpma, 1991), from

bud of Populus tremula in Finland and from Cirsium head in Karelia (Russia) (specimens in collections).

21. Conioscinella livida Nartshuk, 1970 Figs 228, 229 Conioscinella livida Nartshuk in Nartshuk et al., 1970: 418. Holotype in ZISP. Body black, bluish grey-dusted. Frons narrow, yellowish in front. Ocellar triangle black, grey-dusted, extending halfway to front margin of frons, poorly delimited from remaining frons. First flagellomere rounded, yellow, darkened above. Arista black. Eyes haired. Palpi yellow. Scutum and scutellum bluish grey-dusted and evenly covered with setulae. Scutellum rounded, apical setae shorter than scutellum. Distance between apicals a little longer than distance between apical and subapical setae. Pleura dusted above and shining on remaining surface. npl 1 + 2. Abdomen brown, covered with black setulae. Legs black. Halteres yellow. Body length nearly 2.0 mm. Distribution. – Finland: N: Espoo, Sammatti; Sb: Varpaisjärvi. Total 3 males, 4 females, all taken in June. Russia: Lr: Petsamo, 1 male. Not found in other Nordic countries. European species, known from Estonia, St. Petersburg district and Germany (Bavaria) (Tschirnhaus & Schacht, 2000). Biology. – Adults in bogs.

22. Conioscinella messaurea Nartshuk & Andersson, sp. nov. Figs 230, 231 Holotype, male: Sweden: Lule Lapmark: Messaure, 15–27.vii.1974 (K. Müller). Paratypes: 31 males, 57 females, same data as holotype except 17.vi– 15.viii.1974. All specimens in alcohol. Holotype and most paratypes in ZMLU, some paratypes in ZISP. Diffferential diagnosis. – Similar to C. frontella, but differs in following characters: genae wider, nearly as wide as first flagellomere, vibrissal angle distinctly produced beyond eye margin and proboscis longer. General colour darker, lower margin of genae blackish, frons entirely dark without yellow band in front, all femora and tibiae black in both 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.

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first flagellomere reniform, shorter than high, darkened, but not black; arista black, short pubescent. Male genitalia: differences from C. frontella rather subtle, but cerci wider apart.

Oscinis sordidella Zetterstedt, 1848: 2659. Holotype in ZMLU (Andersson, 1966). Conioscinella flavifrons Duda, 1933: 52, as var. of C. frontella (Fallén) (Tschirnhaus, 1992).

Description. – Male, female. Ocellar triangle dull, extending halfway to front margin of frons. Frons covered with small black setulae. All setae of head small. orb numerous, no less than 7–8 on each side of frons. Face and genae fuscous yellow. Genae distinctly produced, as wide as first flagellomere, blackish along lower margin. Palpi fuscous yellow to blackish. Proboscis with elongated labella. Scutum and scutellum black, grey-dusted. Scutum evenly covered with setulae. Scutellum rounded behind, a little shorter than wide at base. Apical setae longer than scutellum, subapical setae small, distance between apical and subapical setae equals 1/2 distance between apical setae. Pleura shining except on upper part of anepisternum. npl 1 + 2. All setulae and setae of head and thorax black. Abdomen brown. Wing venation typical for the genus. Halteres yellow. Legs black, tarsi sometimes partly yellowish. Body length 1.75–2.0 mm.

Black, grey-dusted with yellow setae and setulae on head and thorax. Frons yellow. Ocellar triangle black, grey-dusted, extending halfway to front margin of frons. First flagellomere rounded, yellow. Arista black. Eyes sparsely haired. Palpi yellow. Scutum and scutellum grey-dusted and evenly covered with pale setulae. Scutellum rounded, apical setae longer than scutellum. Pleura dusted above and shining on remaining surface. npl 1 + 2. Abdomen brownish. Legs entirely yellow, including coxae. Body length nearly 2.0 mm.

23. Conioscinella mimula Collin, 1946 Figs 232, 233 Conioscinella mimula Collin, 1946: 129. Syntypes 2 males, 9 females in OXUM (Pont, 1995). Very similar to C. frontella, but smaller, male legs with yellow front and middle tibiae, female legs entirely yellow. Frons in upper part dark, frontal and thoracic setae and setulae dark, first flagellomere yellow, brownish above. Body length 1.0–1.5 mm. Distribution. – Denmark: EJ, NEJ, F, LFM, NEZ, B. Sweden: north to JÄ. Finland: north to Ta. Russia: Vib. Not so far found in Norway. European species. Biology. – Adults from June to middle of August in meadows. Larvae phytophagous, live in shoots of Anthoxanthum odoratum, Bromus sp., Lolium perenne, hibernate in shoots of their host plant and pupate following spring (Nye, 1958, 1959; Tschirnhaus, 1981). Larvae described by Nye (1958).

24. Conioscinella sordidella (Zetterstedt, 1848) Fig. 234

Distribution. – Denmark: LFM, NEZ, B. Sweden: north to VS. Norway: AK, HEs (Thunes et al., 2004). Finland: Al, Ab, N. Russia: Vib, Kr. European species. Biology. – Adults from end of June to end of August in dry meadows; found in canopies of Pinus silvestris in Norway (Thunes et al., 2004). Larvae phytophagous, live in shoots of Dactylis glomerata, Lolium perenne, Leymus sp., Bromus sp., Anthoxanthum odoratum, Elytrigia repens (Karpova, 1972; Tschirnhaus, 1981, 1992).

25. Conioscinella tornensis Nartshuk & Andersson, sp. nov. Figs 235–237 Holotype, male, Sweden: TO: Stordalen, 18.vii.1957 (P.I. Persson). There is another male from the same locality and with the same date, but without head. Holotype in ZMLU. Diffferential diagnosis. – Larger than C. frontella and differs by darker body, longer orb and other setae, especially on femora and abdomen, darkened wings and larger epandrium. Description. – Male. Frons black, only narrowly brown in front. Ocellar triangle extending a little more than halfway to front margin of frons, densely dusted. orb numerous (c. 8) and rather long, longer than vti and oc, but thinner. Eyes ovoid, covered with short hairs. Genae as wide as first flagellomere, fuscous yellow, black on lower margin, with distinctly produced vibrissal angle. Antennae yellow, first flagellomere reniform; arista black, shortly 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 covered with setulae. Scutellum semicircular. Apical setae longer than scutellum, two subapical setae on each side. Pleura shining except on upper part of anepisternum. npl 1 + 2. Postnotum black, shining. Abdomen black, covered with rather long setae. Legs entirely black, including tarsi. Wing brownish with venation usual for the genus. Halteres yellow. Male genitalia: epandrium large, with rounded cerci and slightly tapering surstyli; hypandrium open; postgonite with long hairs. Body length 2.2 mm.

26. Conioscinella zetterstedti Andersson, 1966 Figs 40, 238–240 84

Conioscinella zetterstedti Andersson, 1966: 72. Holotype in ZMLU. Head black, front margin of frons, face and genae yellow. Ocellar triangle black, dusted, extending 3/4 of frons. Basal antennal segments yellow, first flagellomere rounded, yellow and darkened above, especially on outer side. Scutum smooth and dusted. Scutellum very short and varies in size. It is proportionally larger in full-winged specimens. Apical scutellar setae wide apart. Wings vary markedly in size. Venation reduced in short-winged specimens. Halteres yellow, smaller in shortwinged specimens. Rare full-winged specimens occur: Finland: Ta: Hartola, 11.vi.1987 (Koponen), 1 specimens; Ok: Paltamo, 03.vii.1991 (Koponen), 1 specimen. Femora black, front and middle tibiae

yellow, especially in females, hind tibiae black except yellow at ends. Body length 1.25–1.5 mm. Distribution. – Denmark: EJ, WJ, NWJ, NEJ, NEZ. Sweden: SK, GO, UP, along coast of the Baltic Sea to NB. Finland: N, Ta, Sa, Om, Ok, ObS, Li. Russia: Vib, Kr. Not found in Norway. Holarctic species. Biology. – Adults xerobiontic, from middle of May to end of September, preferable in coastal dune areas, but sometimes inland. Biology and variability in size of the wings and halteres have been studied by Brauns (1937, 1938, 1959, as Conioscinella brachyptera). Ardö (1957) found adults in dunes in dense tufts of Leymus and Ammophila. Flies are excellent jumpers and run rapidly on plants and on the sand surface. It has also been found in mammals’ nests, peat-bogs, swamps, shore debris and leaf litter (Mohrig, 1967; Wheeler, 1994b; Beaulieu & Wheeler, 2001). Adults were found in Sphagnum in February in the Netherlands (Pelt, 1926). Larvae have been found in rotting shoots of Ammophila arenaria injured by Meromyza pratorum (Balachowsky & Mesnil, 1935). Tschirnhaus (1981) listed Elytrigia, Ammophila and Molinia coerulea as host plants. Karpova (1972) reared the species from Agropyron cristatum. Beaulier & Wheeler (2001) found the species on Carex in Canada.

Genus Dicraeus Loew, 1873 Dicraeus Loew, 1873: 51. Type species: Dicraeus obscurus Loew, 1873 (= Oscinis raptus Haliday, 1838) by monotypy. Oedesiella Becker, 1910: 146. Type species: Oedesiella discolor Becker, 1910 by monotypy. Paroedesiella Enderlein, 1936: 187. Unavailable. Post-1930 name proposed without designation of a type species. Paroedesiella Anonymous, 1937: 394. Type species: Oscinella styriaca Strobl, 1898 by monotypy. Paroedesiella Sabrosky, 1941: 761. Type species: Oscinella styriaca Strobl, 1898 by original designation. Black or yellow species with black stripes on scutum, with short semicircular scutellum, rather broad genae, long vein R2+3 . poc upright and cruciate. Ocellar triangle usually thinly dusted. First flagellomere rounded oval, arista shortly pubescent. Tibial organ distinct, narrow. Femoral organ formed by a group of warts with short stout spines.

The genus is divided into several subgenera mainly based on differences in the male genitalia. The subgenus Dicraeus has slender male cerci in a lateral position near base of simple surstyli. The subgenus Oedesiella has similar male cerci, but surstyli are long, of complex shape and with distinct setae. In the subgenus Paroedesiella male cerci are minute and in median position or absent, and surstyli are short and broad with spine-like setae. Distribution. – Probably worldwide, but not so far found in the Neotropical Region.

Key to species of Dicraeus Loew 1

– 2 – 3

– 4

–

5 –

6 – 7 – 8 –

Costal vein ending between apices of veins R4+5 and M. Male genitalia Figs 243, 244 . . . . . . . . . . . . . . . . . . . . . . . D. ingratus (Loew) Costal vein extending to end of vein M . . . . . 2 Crossvein dm-cu absent (Fig. 247). Male genitalia Figs 248, 249 . . . . . . . D. raptus (Haliday) Crossvein dm-cu present (Fig. 37) . . . . . . . . . 3 Postpronota and pleura partly yellow, in dark specimens yellow parts not distinct. Male genitalia Figs 266–268 . . . . . . D. vagans (Meigen) Thorax entirely black . . . . . . . . . . . . . . . . . . . 4 Pleura dusted; only one posterior npl setae. Male genitalia Fig. 258 . . . . . . . . . . . . . . . . . . . . . D. tibialis (Macquart) Pleura mainly shining, dusted only on upper part of anepimeron and anepisternum; two posterior npl setae . . . . . . . . . . . . . . . . . . . . . . 5 Genae and front margin of frons yellow . . . . 6 Genae fuscous yellow or blackish, frons entirely black. Genae distinctly wider than first flagellomere (Figs 245, 246) . . . . . . . . . . . . . . . . . . . . D. nitidus (Wahlgren) Ocellar triangle with thin layer of dust . . . . . 7 Ocellar triangle shining, without dusting . . . 8 Male genitalia Fig. 242 . . . . . D. fennicus Duda Male genitalia Figs 250–252 . . . . . . . . . . . . . . . . . . . . D. rossicus Stackelberg Male genitalia Figs 254–256 . . . . . . . . . . . . . . . . . . . . . . D. styriacus (Strobl) Male genitalia Figs 259–261 . . . . . . . . . . . . . . . . . . . . . . . . . D. tjederi sp. nov. 85

27. Dicraeus (Oedesiella) fennicus Duda, 1933 Figs 105, 106, 241, 242 Dicraeus fennicus Duda, 1933: 67, as var. of Dicraeus styriacus Strobl. Syntypes in NHMH. Body black, thinly dusted. Ocellar triangle black, slightly dusted, extending nearly to front margin of frons. Face, frons except for hind part and genae yellow; genae wider than first flagellomere. Basal antennal segments yellow, first flagellomere yellow, darkened above. Palpi yellow. Scutum thinly dusted, pleura shining except on upper part of anepisternum. npl 1 + 2. Abdomen brown. Femora and hind tibiae black, other tibiae and tarsi yellow. Body length 2.0–2.5 mm. Distribution. – Rather common. Denmark: WJ, NEJ, NWZ, NEZ, B. Sweden: north to UP including GO, also NB. Norway: Ø, AK. Finland: Al, Ab, N, Ka, St, Ta, Om. Russia: Vib, Kr. Holarctic species, recorded in the Palearctic from Europe to Yakutia, East Siberia and Mongolia. Biology. – Adults from end of June to end of July, rare in August. Larvae phytophagous, feeding on the unripe seeds of Elytrigia repens, Elymus pungens, E. pycnanthus and Elymus sp. (Parmenter, 1955; Nartshuk, 1960, 1967; Tschirnhaus, 1981). Nielsen & Nielsen (1984) recorded this species on agricultural grasses in Denmark, but this is probably because Elytrigia repens is a common weed everywhere.

28. Dicraeus (Dicraeus) ingratus (Loew, 1866) Figs 243, 244 Eutropha ingrata Loew, 1866: 26. Type material not found in ZMHU (Wendt, 1975, 1978). Body black, thinly dusted. Frons fuscous yellow. Ocellar triangle black, dusted, extending to 4/5 of frons. Face and anterior part of genae fuscous yellow, clypeus black. Genae as wide as or wider than first flagellomere. Basal antennal segments brown, first flagellomere nearly black. Palpi black. Scutum thinly dusted, pleura shining except on upper part of anepisternum. npl 1 + 2. Abdomen black. Legs mainly black. Wings clear, costal vein not reaching end of vein M. Body length 2.0–2.5 mm. 86

Distribution. – Only found in Denmark: B: Bastemose, 26.vi.1964 (O. Martin & B.V. Petersen), 1 male, 1 female. Holarctic species, in Palearctic known from Europe, southern Siberia, Israel. Biology. – Larvae phytophagous, feeding on developing seeds of Bromus inermis and B. erectus (Agafonova, 1962; Nartshuk, 1967; Gołebiowska & Nawrot, 1970).

29. Dicraeus (Paroedesiella) nitidus Wahlgren, 1913 Figs 245, 246 Dicraeus nitidus Wahlgren, 1913: 52. Holotype female in ZMLU. Dicraeus napaeus Collin, 1946: 133. Holotype male in OXUM (Pont, 1995). Syn. n. Note. – Dicraeus nitidus Wahlgren was described on a female from Öland. We had the opportunity to examine males collected on Öland that coincide in all essential characters with the holotype of D. nitidus. The male genitalia of these specimens agree with a drawing of D. napaeus by Collin (1946). Body black, thinly grey-dusted. Ocellar triangle black. Frons in front, face and genae blackish. Genae wider than first flagellomere, lower part of genae darker and punctate. Antennae nearly black. Palpi brownish. Pleura mainly shining. npl 1 + 2. Legs black with yellowish ends of tibiae and yellowish brown tarsi. Abdomen black. Body length 2.0–2.3 mm. Distribution. – Denmark: NWJ: Tved Plantage; B. Sweden: north to UP. Finland: Al. Not found in Norway or Russian parts of Fennoscandia. European species. Biology. – Adults in June to July in meadows. Larvae phytophagous, but host plant previously unknown. We have examined specimens from Öland reared from unripe seeds of Helictotrichon pratense. The distribution of the species in the area investigated coincides well with the distribution of the host grass.

30. Dicraeus (Dicraeus) raptus (Haliday, 1838) Figs 247–249

Figs 241–252. Dicraeus fennicus Duda: 241: head, lateral view. 242: hypopygium, postero-lateral view. D. ingratus (Loew): 243: hypopygium, 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.)

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Oscinis raptus Haliday, 1838: 187. Type in BMNH. Body black, grey-dusted. Frons at front margin, face and genae yellow. Ocellar triangle black, thinly dusted and shining, extending to 2/3 of frons. Antennae and palpi yellow, first flagellomere darkened above. Genae wider than first flagellomere. Scutum black, dusted, pleura mainly shining, dusted above. npl 1 + 2. Abdomen brown with yellow base. Wing without crossvein dm-cu. Halteres yellow. Legs with all femora and hind tibiae black. Body length 2.0 mm. Distribution. – Only from Sweden: Öland: Köpinge alvar, 1 female, also recorded by Ringdahl (1947). European species. Biology. – Larvae phytophagous, associated with Bromus ramosus (Ismay, 1981d).

31. Dicraeus (Oedesiella) rossicus Stackelberg, 1955 Figs 250–252 Dicraeus rossicus Stackelberg, 1955: 332. Holotype in ZISP. Similar to D. fennicus, but ocellar triangle and scutum more shining. Body black, frons narrowly yellow in front. Face and genae yellow, occiput black. Ocellar triangle subshining black, extending 4/5 length of frons. Genae wider than first flagellomere. Palpi yellow. Scutum and scutellum thinly greydusted, subshining. Scutellum semicircular, apical setae longer than scutellum. Pleura shining except on upper part of anepisternum. Sulcus between anepisternum and anepimerom yellow. npl 1 + 2. Legs mainly black, front and middle tibiae yellowish. Halteres yellow. Body length 2.0 mm. Distribution. – Finland: Al: Jomala, 1 female; N: Helsinki, 2 females. Russia: Kr: Sonostroff (Suonsaari), Nemetskiy Kuzov Is. Lr: Kandalaksha, 1 female. The Finnish records are in need of verification as based on females only. Trans-Palearctic species, known from Europe to Japan. Biology. – Larvae phytophagous, feeding on developing seeds of Elymus caninus (Nartshuk, 1960, 1967).

32. Dicraeus (Paroedesiella) styriacus (Strobl, 1898) Figs 52, 253–256 88

Oscinis styriaca Strobl, 1898: 263. Syntype male in NMBA (Chvála, 2008). Dicraeus vallaris Collin, 1946: 133. Syntypes 8 males, 7 females in OXUM (Pont, 1995). Synonymy established by J.W. Ismay and published by Nartshuk (2002a). Body black, grey-dusted. Frons in front, face and genae in front yellow. Genae as wide as first flagellomere. Antennae with basal segments yellow or brownish yellow, first flagellomere brownish. Palpi brownish yellow. Pleura mainly shining with yellow anepimeral suture. npl 1 + 2. Legs black, with yellowish knees and tarsi. Abdomen brownish. Body length 2.0–2.3 mm. Distribution. – Denmark: B. Sweden: SK, BL, ÖL, 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). European species. Biology. – Adults from June to beginning of July in meadows. Larvae phytophagous, feeding on developing seeds of Helictotrichon pubescens (Nartshuk, 1960, 1967) and probably Leymus arenarius.

33. Dicraeus (Dicraeus) tibialis (Macquart, 1835) Figs 257, 258 Oscinis tibialis Macquart, 1835: 263. Types probably in MHNL. Body mainly black with face, genae, palpi, anterior part of frons, halteres and abdomen yellow. First flagellomere yellow basally. Ocellar triangle extending 1/2 length of frons, covered with thin dust. Genae slightly wider than first flagellomere. Scutum and pleura mainly dusted. npl 1 + 1. Femora black, anterior tibiae yellow to brownish. Abdomen yellow with lateral dark spots at base. Body length 1.8–2.0 mm. Distribution. – Denmark: SJ, LFM, NEZ. Sweden: SK, SM. Finland: Al, N. Russia: Kr. Not found in Norway. Holarctic species, found more recently in New Zealand (Ismay, 1991). Biology. – Adults in June to early July in meadows. Larvae phytophagous, feeding on developing seeds of Bromus inermis, B. erectus and Helictotrichon pubescens (Nartshuk, 1960, 1967; Agafonova,

Figs 253–261. Dicraeus styriacus (Strobl): 253: head, lateral view. 254: hypopygium, dorsal view. 255: surstylus, ventral view. 256: hypandrium 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) reared the species from a mole hole.

34. Dicraeus (Paroedesiella) tjederi Nartshuk & Andersson, sp. nov. Figs 259–261 Holotype, male, Sweden: SK: Klammersbäck, Raulunda, 15.vi.1948 (B. Tjeder). Holotype in ZMLU. Diffferential diagnosis. – Belongs to the subgenus Paroedesiella, because of male genitalia: cerci absent, surstyli short. It is most similar to D. styriacus but differs in structure of the male genitalia: incision of surstyli deeper in the new species, almost as in D. vagans, and lower margin of surstylus without a projection on anterior part as in D. vagans and D. styriacus. Description. – Male. Head black except yellow on face, genae and narrowly on anterior part of frons. Ocellar triangle shining without any dust. Chaetotaxy of head typical for the genus. Frons and face in the holotype rather deformed and antennae thus partly retracted. Antennae black. Genae not wider than first flagellomere, with a row of black setulae along lower margin. Thorax entirely black, scutum thinly dusted and subshining except posterior part more densely dusted and dull like scutellum. Scutum evenly covered with black setulae. Scutellum triangular, apical setae longer than scutellum, subapical setae inserted a little nearer to base of scutellum than to apical setae. Pleura shining. npl 1 + 2. Wings clear with brown veins. Costal vein reaching end of M. Second costal sector 3.5 times longer than third one. Third costal sector 1.5 times longer than fourth. Veins R4+5 and M parallel. Cell br not widened. Halteres yellow. Legs with all femora black, except at tips, front and middle tibiae fuscous yellow, hind tibiae with black band. Abdomen dark brown. Body length 2 mm.

partly yellow. In dark specimens postpronota with a dark spot or all thorax black. Head black on occiput and ocellar triangle, yellow in front. Ocellar triangle extending 3/4 length of frons. Antennae and palpi yellow. Genae wider than first flagellomere. npl 1 + 2. Abdomen yellow or more or less darkened. Legs mainly yellow. Body length nearly 2.0 mm. Distribution. – Denmark: common and widespread. Sweden: north to UP, also GO. Finland: Al, N. Russia: Vib: Bolshoi Beresovyi Is. near Primorsk. Not found in Norway. European species. Biology. – Adults from end of May to middle of July in meadows and at forest edges. Larvae phytophagous, feeding on developing seeds of Arrhenatherum elatius (Nartshuk, 1960, 1967).

Genus Elachiptera Macquart, 1835 Elachiptera Macquart, 1835: 621. Type species: Chlorops brevipennis Meigen, 1830 by original designation. New generic synonyms published by Nartshuk & Tschirnhaus (2012). Black or rarely yellow species with slightly flattened scutum and scutellum and scutellar setae arising from small warts. Scutum with rather coarse stellate microsculpture, evenly dusted or dust arranged in lines along dorsocentral lines. First flagellomere oval or reniform, arista usually broadly thickened with dense, rather long hairs. Tibial organ short, oval. Femoral organ present as two rows or a group of warts. Wings sometimes reduced. In some species basal tergites fused. Male genitalia with separate cerci, simple surstyli and broad triangular postgonites. Distribution. – All regions except Australasia.

35. Dicraeus vagans (Meigen, 1830) Figs 51, 262–268 Chlorops vagans Meigen, 1830: 389. Type lost, in MNHN only a pin with Meigen’s name label as already noted by Becker (1902). Body yellow with black scutum except for yellow postpronota and stripes along notopleura. Pleura 90

Key to species of Elachiptera Macquart 1

Wings shortened, barely reaching end of abdomen or shorter (Fig. 2). Body partly reddish yellow, abdomen black. Syntergite 1 + 2 of abdomen long, occupying basal half of abdomen. Male genitalia Figs 271, 272 . . . . . . . . . . . . . . . . . . . E. brevipennis (Meigen)

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

– 2 –

3

–

4

–

5 – 6 –

Wings normally developed, usually longer than abdomen . . . . . . . . . . . . . . . . . . . . . . . . . 2 Scutum evenly thinly dusted or shining only on sides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Scutum mainly shining, with two dusted stripes and sometimes dusted before scutellum (Fig. 280) . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Scutellum longer than wide with long tubercles at base of apical setae (Fig. 287). Arista thick, flattened. Legs yellow . . . . . . . . . . . . . . . . . . . . E. tuberculifera (Corti) Scutellum not as long as wide with small tubercles at base of apical setae (Fig. 284). Arista thin, not flattened but densely pubescent. Legs mainly dark . . . . . E. scrobiculata (Strobl) Apical scutellar setae nearly twice as wide apart as their distance to lateral setae (Fig. 282). Scutum dusted before scutellum . . . . . . . . . . . . . . . . . . . . . . . E. diastema Collin Apical scutellar setae close together, barely wider apart than their distance from lateral setae (Fig. 276). Scutum not dusted before scutellum. Male genitalia Figs 277–279 (E. cornuta species complex) . . . . . . . . . . . . . . . . . . 5 Arista thin, legs darkened . . . . . . . . . . . . . . . . . . . . . . . . E. cornuta form A Arista varying in thickness, legs yellow . . . . . 6 Arista thickened, aristal pubescence shorter than diameter of arista . . . . E. cornuta form B Arista variable in thickness but aristal pubescence not shorter than aristal diameter. . . . . . . . . . . . . . . . . . . . . . . . E. cornuta (Fallén)

36. Elachiptera brevipennis (Meigen, 1830) Figs 2, 66, 270–272 Chlorops brevipennis Meigen, 1830: 159; Meigen’s plates 211: 12; 237: 4; 245: 13 (Morge, 1976b). Lectotype in NHMW (Nartshuk, 1978a). Ground colour varies from yellowish red to dark reddish brown, abdomen being darker than thorax. Ocellar triangle shining, reaching front margin of frons. Arista thickened and pubescent, black. Two pairs of orb longer than remaining ones. Genae as wide as first flagellomere. Antennae brown. Scutum densely punctate. Scutellum black with 4 small tubercles at base of scutellar setae. Abdomen black, shining. Syntergite 1 + 2 of abdomen long, 92

occupying 1/2 length of abdomen. Wing length either nearly as long as abdomen or only extending to hind margin of syntergite 1 + 2, in this case with reduced venation. Body length 1.5–2.0 mm. Distribution. – Found only in Denmark: SJ, EJ, F, NEZ. European species. Biology. – Adults in May to September in fens, costal marches and other wetlands with Carex, Scirpus and Typha. Adults crawl on plants near to the ground (Mohrig, 1967). Hibernates as adults. Tschirnhaus (1981) reared the species from shoots of Dactylis glomerata. Some early authors (Smith, 1856; Brauer, 1883) reported that eggs of this species were found under the short wings of the bug Nabis subapterus, but this record seems be doubtful. Vimmer (1906) discussed reduction of venation in brachypterous specimens.

37. Elachiptera cornuta (Fallén, 1820) Figs 22, 33, 58, 273–279 Oscinis cornuta Fallén, 1820: 6. Lectotype in ZMLU (Andersson, 1963). Body black, shining. Frons yellow, ocellar triangle black, smooth and shining, extending 2/3 of frons. Two pairs of orb longer than remaining ones. Genae yellow with silvery dust, nearly as wide as first flagellomere. Antennae yellow, first flagellomere black at aristal base. Arista variable, widened and thickly pubescent. Palpi yellow. Scutum and scutellum shining, coarsely punctate, scutum with two longitudinal dusted stripes along dorsocentral lines. Apical and subapical setae of scutellum on small tubercles. Distance between apical setae as distance between apical and subapical setae. Pleura smooth, shining except on upper part. npl 1 + 1. Setae of head and abdomen yellow. Abdomen black, shining. Legs yellow or more or less darkened. Body length 2.0–3.0 mm. Distribution. – Throughhout Fennoscandia and Denmark, reaching the coast of the Arctic Ocean. Pan-Palearctic species. Biology. – Common species in diverse types of habitats. Hibernates as adults in birds’ nests, Lipara galls, under scales of coniferous cones, and between the leaves of Typha (Tschirnhaus, 1992; Krivokhatsky & Nartshuk, 2001). Larvae saprophytophagous, secondary invaders, develop in rotting tissue of different grasses: Phrag-

Figs 269–279. Elachiptera brevipennis (Meigen): 269: head, dorsal view. 270: head, lateral view. 271: hypopygium, dorsal view. 272: hypandrium 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.

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mites, Agrostis, Dactylis, Calamagrostis, Deschampsia, Festuca, etc., and all species of cereals. Also found in other mono- and dicotyledon plants: decaying leaves of Typha latifolia and T. angustifolia, Bolboschoenus maritimus, Scirpus, Schoenoplectes taberhaemontani, Triglochin maritimus, Atriplex triangularis, A. prostrata, Gladiolus communis, Solanum fruticosum, Oenanthe crocata, Cannabis sativa (Kreiter, 1927, 1928; Goidanich, 1928; Jepson & Southwood, 1960a; Vilkova, 1962; Tschirnhaus, 1981, 1992). Rygg (1966) reared it from seedlings of wheat damaged by Phorbia fumigata (Anthomyiidae) in Norway, and Johansson (1960) from cereal shoots in Sweden. Mayer (1949) recorded the species as a pest of tomatoes, and Show (1955) as a pest of gladioli. It has been reared from a Schoenoplectes stem and from a stem of Bolboschoenus maritimus infested by Bactra robustana (Lepidoptera) together with Eribolus danicus. The species has no economic importance in cereal fields (Walton, 1937; Vilkova, 1962). Larvae were described by Vimmer (1926, 1931), Kreiter (1928), and Balachowsky & Mesnil (1935). Jepson & Southwood (1960a) gave distinguishing characters of eggs and first instar larvae from those of Oscinella frit. Note. – The species is very variable in size, colour of legs and thickness of arista. We found two deviating forms of the present species: (1) Form A is found in northern Fennoscandia and differs from normal E. cornuta in having a smaller body, black legs and rather thin arista. (2) Form B differs from normal E. cornuta in being larger and having a very broad arista and bright yellow legs; it occurs in southern Scandinavia especially in small ponds on Typha. The taxonomic status of these forms is unsettled. They either represent distinct taxa or two extremes within one variable species. There are some differences in the male genitalia between these forms, but some specimens are not always separable from the typical form of E. cornuta. Their status is unclarified. Examination of specimens from other parts of the vast range of E. cornuta is clearly needed to find out is this is a species or species complex. Besides, there exist many names proposed as species, subspecies and forms that are tentatively considered as synonyms of E. cornuta.

38. Elachiptera diastema Collin, 1946 Figs 280–282 94

Elachiptera diastema Collin, 1946: 147. Syntypes 9 males and 6 females in OXUM, 1 syntype in the Manchester Museum (Pont, 1995). Similar to E. cornuta, but differs from it the following characters: wings shorter, dusted stripes on scutum joined behind by a prescutellar dusted band, scutellum shorter and more truncate, apical setae much farther apart, distance between apical setae nearly twice the distance between apical and subapical setae. Setulae of head and abdomen dark. Body length 2.0–2.5 mm. Distribution. – Denmark: common and widespread. Sweden: north to NB along the Baltic Coast. Norway: Ø, HEs. Finland: N, Ka, Ta, Ok, Ob, Ks. Russia: Vib, Kr. Euro-Mediterranean species. Biology. – Adults in May and more common in August and September in wet habitats. Larvae phytosaprophagous, developing in shoots of grasses (Dactylis glomerata and Festuca arundinaceae (Sharipov, 1981; Tschirnhaus, 1981)). Collected on agricultural grasses in Denmark (Nielsen & Nielsen, 1984). Kibric et al. (1999) listed the species as tyrphophilous.

39. Elachiptera scrobiculata (Strobl, 1901) Figs 283, 284 Oscinis scrobiculata Strobl, 1901: 233. Syntypes male and female not found in NMBA or the Klagenfurt Museum (Chvála, 2008). Elachiptera tenuiseta Frey, 1947: 69. Holotype in NHMH, but Nartshuk in 1990 found only the pin and labels without fly. Body black, shining. Frons dark brown to black. Ocellar triangle black, shining, extending 3/4 of frons. Two pairs of orb longer than remaining pairs. Face and genae dark brown, genae as wide as first flagellomere. First flagellomere black. Arista rather thickly pubescent, black. Palpi dark. Scutum dusted, scutellum wider than long, trapezoid, coarsely punctate. Apical setae wide apart, on small tubercles. Pleura shining. npl 1 + 1. Abdomen black. Legs partly black. Halteres yellow. Body length 1.7–2.0 mm. Distribution. – Sweden: north to LU and NB. Norway: AAy: Jåmåsknutene, Froland. Finland: north to LkE. Russia: Vib. Not so far found in Denmark.

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 British Isles to Far East of Russia. Everywhere, but not common.

40. Elachiptera tuberculifera (Corti, 1909)

Biology. – Adults from May to October, probably hibernating like other species of the genus, in wet habitats with Carex. Apparently, the present species was found by Bleton & Fieuzet (1943) in stems of sorghum damaged by Atherigona (Muscidae) in Morocco.

Crassiseta tuberculifera Corti, 1909: 132. Types not seen, possibly in the Genova Museum.

Figs 285–287

Body black. Frons yellow, ocellar triangle black, shining, extending nearly to front margin of frons. Genae yellow, postgenae black. Antennae yellow, 95

first flagellomere darkened on apex. Arista black, broadly thickened with short marginal hairs. Thorax black, scutum dusted, postpronota shining. Scutellum a little longer than wide, flattened and weakly shagreened, dusted, with 3 pairs of setae on finger like tubercles. Legs yellow. Body length 2.0– 2.5 mm. Distribution. – Denmark: NEZ. Sweden: north to LU and NB. Norway: Ø, AK, HEs, On, Bv, VE, AAy, Ry. Finland: north to Om. Russia: Vib, Kr, Lr. Trans-Palearctic, polyzonal species. Biology. – Adults in wetlands, meadows and moist places in forests, very common, taken in traps from March to September. Hibernating adults found in birds’ nests (Tschirnhaus, 1992; Krivokhatsky & Nartshuk, 2001). Taken in a light trap in December in Norway (Hågvar & Greve, 2003). Larvae phytosaprophagous, developing in insect borrows with decaying plant tissue and frass. Found as secondary invader of grasses and all cereals (Vilkova, 1962), but also in stems of other monocots: Liliaceae, Iridaceae, Typhaceae. It has been reared from stems of several cereals (wheat, barley, rye), mostly infested by Phorbia fumigata (Anthomyiidae) in Norway and Sweden (Johansson, 1960; Rygg, 1966).

Genus Eribolus Becker, 1910 Eribolus Becker, 1910: 127. Type species: Eribolus sudeticus Becker, 1910 (= Oscinis nana Zetterstedt, 1838) by designation of Enderlein, 1911: 206. Black, dusted or shining species with flattened scutum. Ocellar triangle usually dusted. Vibrissal angle rounded. First flagellomere squarish or rounded. poc upright and cruciate. Setulae on scutum arranged in lines. Pleura mainly shining. npl 1 + 1. Legs sometimes with thickened femora, more distinct in males. Tibial organ oval. Femoral organ a group of slightly differentiated short setulae. Male genitalia: cerci and surstyli simple, hypandrium closed behind, postgonites enlarged. Distribution. – Holarctic and Neotropical Regions.

Key to species of Eribolus Becker 1

96

Frons narrow but distinctly reddish-yellow anteriorly. Genae wide, almost as wide in front

–

2 – 3

–

4

–

as first flagellomere (Fig. 296). Ocellar triangle extending almost to front margin of frons (Fig. 295). Male genitalia Figs 299, 300 . . . . . . . . . . . . . . . . . . . . . E. nana (Zetterstedt) Frons entirely black or slightly brownish in front. Genae in front distinctly narrower than first flagellomere (Figs 289, 291, 292, 302) ......................................2 Ocellar triangle shining except ocellar tubercle . . . . . . . . . . . . . . . . . . . E. gracilior (Meijere) Ocellar triangle entirely dull . . . . . . . . . . . . . 3 Genae in front nearly as wide as front tibiae (Fig. 292), first flagellomere slightly reddishyellow at base. All tibiae yellow at both ends. Femora not enlarged. Palpi dark. Male genitalia Figs 293, 294 . . . . . E. hungaricus Becker Genae wider, nearly half width of first flagellomere. Femora enlarged, more distinctly so in males (Figs 303, 304) . . . . . . . . . . . . . . . . . . . 4 Larger, 1.5–2 mm. In males first flagellomere yellow, darkened at aristal base, palpi yellow. In females first flagellomere dark, but yellow at base, palpi brown. Femora moderately enlarged in males. Male genitalia Figs 305, 306 . . . . . . . . . . . . . . . . . . . . . E. slesvicensis Becker Smaller, 1.0 mm. In males first flagellomere dark, reddish-yellow at base, palpi brown and all femora greatly enlarged. In females first flagellomere and palpi entirely black, femora slightly enlarged . . . . . . . . . E. danicus sp. nov.

41. Eribolus danicus Nartshuk & Andersson, sp. nov. Figs 288–290 Holotype male and paratype female: Denmark: NEZ: Amager, bread from stem of Bolboschoenus maritimus infested by Bactra robustana (Lepidoptera), collected 04.vi.1921, fly emerged 05.vii.1921 (Gusmann). Other paratypes: Sweden: SK: Alnarp, 20.vi.1960 (H. Andersson), 1 female; Lomma, 16.viii. 1959 (H. Andersson), 1 female; Silvåkra, 26.v.1982 (C. Hansson), 1 female. Holotype and 1 paratype in ZMUC, 3 paratypes in ZMLU. Diffferential diagnosis. – Most similar to E. hungaricus Becker, but distinguished by smaller size, wider genae, more distinctly enlarged femora, especially in males.

Description. – Male, female. Small, black. Frons shorter than wide, dull. Ocellar triangle extending 2/3 length of frons, dull, behind nearly as wide as frons. Three rather long and thin orb on each side of frons. poc and vte stronger and little longer than orb. First flagellomere large and round, arista shortly pubescent, nearly as long as frons. Genae in front a little narrower than a half width of first flagellomere, distinctly widened posteriorly. Eyes short oval. Scutum flat, dusted. Scutellum short, triangular. Apical setae thin, shorter than scutellum. Pleura shining. npl 1 + 1. Wings little brownish, longer than abdomen. Second sector of costal vein nearly 1.5 longer than third sector which is 1.25 times longer than fourth sector. Cell br not widened. Veins R4+5 and M divergent apically. Halteres yellow. Femora of all legs strongly enlarged in male, distinctly less so in female. Femora black, tibiae black or a little paler at each end, tarsi yellowish. Body length 1 mm.

42. Eribolus gracilior (Meijere, 1918) Fig. 291 Oscinella gracilior Meijere, 1918: 140. Lectotype in ZMAN (Nartshuk, 2003). Small, very flattened species, entirely black. Frons dull, ocellar triangle shining, with square dusted spot on ocellar tubercle, extending halfway to front margin of frons. Antennae black, first flagellomere rounded. Genae a little narrower than first flagellomere. Scutum flat, scutum and scutellum thinly dusted and subshining. Pleura shining except on upper part of anepisternum. npl 1 + 1. Femora black, front tibiae yellow, other tibiae yellow with black band, tarsi yellow. Wings narrow, longer than abdomen. Veins R4+5 and M divergent apically. Body length 1.0 mm. Distribution. – Sweden: SK, HA, ÖL, VG, and also further north in NB. Finland: Ta: Forssa, 26.vi.1964 (M. Loikaaanen), Ok: Paltamo, 10.vii.1981, 30.vi.1991 (Koponen). Not so far found in Denmark, Norway or Russian parts of Fennoscandia. Eurasian species. Biology. – Adults in May to August in bogs and other wetlands, often swept from edges of open water without Phragmites (Ismay, 1980a).

43. Eribolus hungaricus Becker, 1910 Figs 292–294 Eribolus hungaricus Becker, 1910: 128. Syntype female in ZMHU (Wendt, 1975). Small, very flattened species with black dull body. Frons black, brownish in front. Ocellar triangle dull extending 3/4 of frons. Genae narrower than first flagellomere. First flagellomere black, yellowish at base. Palpi black. Scutum flat, dull. Scutellum short with apical setae short and wide apart, inserted near to subapical setae. Femora black, tibiae yellow with black band. Body length 1.25 mm. Distribution. – Denmark: SJ, WJ, F, NEZ. Sweden: SK, SM, ÖG. Norway: HOi. Finland: Al, Ab, N, Ta. Russia: Vib. European species. 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 is common in reed beds in saline dunes in England.

44. Eribolus nana (Zetterstedt, 1838) Figs 295–300 Oscinis nana Zetterstedt, 1838: 782. Lectotype in ZMLU (Andersson, 1966). Body black, dull, narrow and flattened. Frons reddish in front. Ocellar triangle dull, extending 3/5 of frons. Genae wider than first flagellomere. First flagellomere black, rounded. Palpi black. Scutum flattened, grey-dusted. Scutellum shorter than long with short apical and subapical setae. Pleura shining below. Femora black except yellow at apices, tibiae yellow with black band. Body length 1.5– 2.0 mm. Distribution. – The most abundant and widespread species of Eribolus. Denmark: EJ, NWJ, F, NEZ. Sweden: north to LU and NB. Finland: north to Ks. Russia: Vib, Kr. Not recorded yet from Norway. Holarctic species, widely distributed in the Palearctic. Biology. – Adults hibernate and may be caught from February to October in different kinds of wetlands: bogs, fens, marshes, especially on reeds (Ismay, 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 of Carex, Scirpus, Sparganium and probably other hygrophilous monocots with borrows by other insects, often Cordilura spp. (Scathophagidae) (Valley et al., 1969; Nartshuk, 1972a; Uffen & Chandler, 1978; Valley & Foote, 1996). Johansson (1943) reported it from cereals in Sweden, but this needs to be confirmed. Valley et al. (1969) found 10 larvae in a stem of Carex injured by Cordilura varicornis Curran in North America. 98

45. Eribolus slesvicensis Becker, 1910 Figs 301–306 Eribolus slesvicensis Becker, 1910: 128. Syntypes (1 male, 1 female) in ZMHU (Wendt, 1975). Eribolus crassipes Nartshuk, 1972c: 910. Holotype in ZISP. Syn. n. Small flattened species with black dull body and enlarged femora, especially in male. Frons black,

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, dull, extending 3/4 of frons. Genae nearly as wide as first flagellomere. Antennae mainly yellow, but first flagellomere darkened above. Palpi yellow. Scutum dull, but pleura shining below. Scutellum small, apical setae short, rather wide apart. Femora enlarged. Femora and hind tibiae black with yellow ends, front and middle tibiae mainly yellow. Body length 1.5–2.0 mm. Distribution. – Rare species. Sweden: SK: Landskrona, 17.vii, 1 male; Skälderviken, 05.viii.1955 (Ringdahl), 2 males. Finland: Al: Sund (R. Frey); Föglö, vii.1939 (A. Nordman); N: Tvärminne; Parvoo. Not found in Denmark, Norway or Russian parts of Fennoscandia. Eurasian species, eastward to Mongolia. Biology. – Adults in June and August in wetlands, associated with reed beds (Phragmites australis), especially at lakes and sea shores. Tschirnhaus (1981) listed as host plants Carex and Bolboschoenus maritimus. Ismay (1980a) found the species in saline habitats in England.

Genus Gampsocera Schiner, 1862 Gampsocera Schiner, 1862: 431. Type species: Chlorops numerata Heeger, 1858 by monotypy. Black species with obliquely prolonged first flagellomere and apically thickened arista. Ocellar triangle shining. poc upright, cruciate. Male wings with black marks. Tibial organ distinct, oval. Femoral organ absent. Male genitalia: epandrium flattened, 100

with small cerci and simple surstyli. Phallic complex fused. Distribution. – Palearctic, Oriental and Neotropical Regions.

46. Gampsocera numerata (Heeger, 1858) Figs 3, 23, 38, 307–309 Chlorops numerata Heeger, 1858: 302. Lectotype in NHMW (Nartshuk, 1997b). Frons, face and genae yellow. Frons broader than long. Setae and setulae black. Ocellar triangle black, shining, extending half of frons, yellow behind and brown in anterior part. Antennae and palpi yellow. Scapus very short. First flagellomere prolonged dorsodistally; arista short and black, apically tickened and densely haired. Genae with silvery dust, narrower in front than behind, with rounded vibrissal angle and long pale vibrissa. Thorax yellow 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 shining, yellow with black marks or nearly entirely black. Abdomen brownish black. Legs mainly black or with indistinct black marks. Wings hyaline in female and with black marks in male. Body length 2.0–2.5 mm. Distribution. – Rare. Sweden: UP: Uppsala, 27.x.1975 (L. Hedström), 1 female. Norway: AK, AAy. Finland: Ta: Tavastia (Palmén), 1 male. Not found in Denmark or Russian parts of Fennoscandia.

Eurasian species, known from Europe to Yakutia, East Siberia, but scarce everywhere. Biology. – Adults occur in forests. Larvae probably myxo-mycetophagous as once found under plasmodium of a slime mold. Heeger (1858) found larvae in a stem of Alcea rosea damaged by Apion curvirostris (Brentidae). He observed a pair in copula and a female laying eggs in the decaying stem with excrements of Apion.

–

5

Genus Gaurax Loew, 1863 Gaurax Loew, 1863: 35. Type species: Gaurax festivus Loew, 1863 by monotypy. Species black or yellow with black markings. Ocellar triangle shining. Eyes usually distinctly haired. Genae narrow, usually with silvery dust. poc upright, cruciate. First flagellomere oval or reniform. Arista usually densely pubescent, but not thickened. Tibial organ distinct, oval. Femoral organ absent. Male genitalia: cerci usually long, surstyli of complex structure with long setae, phallic complex fused. Species of Gaurax are very variable in body and size, and only examination of the male genitalia ensures correct identification. Distribution. – Worldwide.

– 6

–

7

Key to species of Gaurax Loew 1

– 2 – 3

– 4

First flagellomere black with white pubescence; arista white, slightly thickened. Male genitalia Figs 331–333 . . . . . . . . . . . . . . . . . . . . G. leucarista Nartshuk Antennae different . . . . . . . . . . . . . . . . . . . . . 2 First flagellomere rounded, nearly as wide as long (Figs 313, 334) . . . . . . . . . . . . . . . . . . . . . 3 First flagellomere reniform, wider than long (Fig. 338) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 First flagellomere large, nearly as wide as height of face (Fig. 334). Male genitalia Figs 335–337 . . . . . . . . . G. macrocerus (Nartshuk) First flagellomere smaller, narrower than height of face (Fig. 313) . . . . . . . . . . . . . . . . . . 4 Male genitalia: cerci narrow, rather long; surstyli strongly elongated, widened apically

– 8

–

9

–

(Figs 310, 311). Body mainly, sometimes entirely black . . . . . . . . . . . . . . G. borealis (Duda) Male genitalia: cerci shorter and wider apart, surstyli shorter (Figs 315, 316). Colour of body varied, but thorax normally yellow with black stripes fused on anterior part of scutum, scutellum normally yellow, pleura with four black spots . . . . . . . . . . . . . . . . G. dubius (Macquart) Apical part of wings slightly darkened especially in male, best seen in fresh specimens. Male genitalia Figs 339, 340 . . . . . . . . . . . . . . G. maculipennis (Zetterstedt) Wings without darkening. Male genitalia different . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Ocellar triangle yellow, shining, only on ocellar tubercle black. Hind tibiae with dark narrow band at base. Usually one dark spot on pleura, on anepisternum, but some females have 3 or 4 dark spots. Male genitalia: surstyli with narrow process directed backwards (Figs 326–328) . . . . . . . . . . . . . . . G. fascipes Becker Ocellar triangle mainly or entirely black, shining or dusted. Usually 4 dark spots on pleura, or thorax entirely black. All legs entirely yellow, rarely tibial organ on hind tibiae surrounded by dark, or entirely dark . . . . . . . . . 7 Body mainly black except reddish yellow at front margin of frons and on genae, or also scutellum, notopleura and hind part of postpronota yellow . . . . . . . . . . . . . . . . . . . . . . . . . 8 Ventral side of thorax yellow with black spots. Legs yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Body black except reddish yellow at front margin of frons and on genae. All legs darkened. Male genitalia Fig. 341 . . . . G. suecicus sp. nov. Thorax black, with scutellum, a large spot covering notopleura, hind part of postpronota and upper part of anepisternum yellow. Legs yellow with black mark on all femora and mid and hind tibiae . . . . G. flavomaculatus (Duda) Male genitalia: cerci tapering and parallel, close to each other, surstyli with acute process directed medially . . . . . . G. fungivorus sp. nov. Male genitalia: broader and wider apart, surstyli with some processes on lower margin (Figs 317–320) . . . . G. ephippium (Zetterstedt) 101

Note. – Flies of the genus Gaurax are not numerous in the collections. Species identification is difficult because of considerable variation in colour within species and between sexes. Therefore examination of the male genitalia is essential for reliable identification. Additional species, including undescribed ones, may well occur in the investigated area.

47. Gaurax borealis (Duda, 1933) Figs 310, 311 Botanobia borealis Duda, 1933: 80, as var. of B. dubia (Macquart). Syntypes (1 male, 2 females) in NHMH. Usually scutum and scutellum black, covered with pale setulae, postpronota and notopleura yellow, but postpronota with black spot. Pleura yellow with 4 large black spots, but females from northern localities entirely black. Head yellow with black occiput. Frons as long as wide. Ocellar triangle short, extending half length of frons, black, shining. First flagellomere nearly as wide as long, more or less darkened. Genae narrow with silvery dusting. Palpi yellow. Setae of head and thorax black. Abdomen darkened dorsally. Legs yellow. Halteres yellow. Male genitalia: cerci narrow and long, surstyli elongated. Body length 1.3–1.5 mm. Distribution. – Rare. Sweden: SÖ: Tyresta Nature Reserve, 28.v–15.vii.1999 (B. Viklund), 26.v– 27.vii.2000 (Viklund, Wikars & Ahnlund), 1 male, 1 female, Malaise traps. Finland: Ab, Ok, Ks, LkE. Russia: Vib: Jukki, 12.vi.1932 (Stackelberg), 1 female; Kr: Paanajärvi, females? (specimens darkened), recorded by Frey (1933) as Conioscinella gallarum. Not found in Denmark and Norway. Note. – Described as a var. of dubia Macquart from Finland, but Duda (1933) did not specify exact localities and number of specimens. Frey (1933), who published details about the Finnish Chloropidae identified by O. Duda, listed the following localities: Ab: Lojo (Krogerus); OK: Kajana (Wuoretanus); Ks: Kuusamo, Paanajärvi (Frey); LKem: Muonio (Palmén). We found in NHMH: 1 female, Kajana (Wuoretanus), type No 8150; 1 female, Lapponia (Palmén), type No 8151; and 1 male, Kuusamo (Frey), type No 8149. The last specimen is presently designated as lectotype. Eurasian species, from Europe to Mongolia. Biology. – Adults in forests. 102

48. Gaurax dubius (Macquart, 1835) Figs 312–316, 971–973 Oscinis dubius Macquart, 1835: 604. Type probably in MHNL. Body yellow with three black stripes on scutum and four black spots on pleura. Frons as long as wide. Ocellar triangle short, extending half length of frons, yellow with black spot on ocellar tubercle, shining. First flagellomere nearly as wide as long, yellow or more or less darkened above. Genae narrow with silvery dusting. Male genitalia: cerci rather short and wide apart, surstyli short. Body length 1.3–1.5 mm. Distribution. – Sweden: SK, SM; SÖ: Tyresta Nature Reserve, 11.vi–17.vii, 28.vii–15.viii.1999, 04.viii– 24.ix.2000, in Malaise traps (Viklund, Wikars & Ahnlund), 1 male, 4 females; probably also LU: Messaure, but only females available. Norway: AK, HEs (Komonen et al., 2004). Finland: Al, Ab, N, Li. Russia: Vib, Kr. Not found in Denmark and Norway. Holarctic species. Occurs in forests. Biology. – Larvae primary fungivorous, develop in bracket fungi Piptoporus betulinus and Fomitopsis pinicola (Smith, 1964; Pielou, 1966; Pielou & Pielou, 1968; Pielou & Verma, 1968; Økland & Hågvar, 1994; Komonen et al., 2004). Smith (1964) noted that flies were reared from old dry bracket fungi. Adults also visit other fungi, Coriolus and Bjerkanderia (Uffen & Chandler, 1978). Larvae and puparia were also found under bark of coniferous trees infested by the bark beetle Pityogenes chalcographus (Bogdanova, 1975; Nartshuk, 1987) and in cones of Picea abies (Saksons, 1973; Skrzypczynska, 1982), but may be the latter records may concern G. ephippium. Bogdanova (1975) considered the species as entomophagous. Larvae described by Smith (1964), some larval structures illustrated by Nartshuk (1987).

49. Gaurax ephippium (Zetterstedt, 1848), sp. rev. Figs 317–320 Oscinis ephippium Zetterstedt, 1848: 2664. Holotype in ZMLU (Andersson, 1966). Restored from synonymy with Gaurax dubius (Macquart, 1835).

Figs 310–320. Gaurax borealis (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 paratype in ZISP, other paratypes in Institute of Biology in Salaspils, Latvia. Syn. n. Body yellow with three black stripes on scutum and four black spots on pleura. Scutal stripes fused in anterior part of scutum, shining, covered with pale setulae. Scutellum yellow, apical setae longer than scutellum. Frons as long as wide. Ocellar triangle short, extending half length of frons, yellow with black spot on ocellar tubercle, shining. First flagellomere reniform, wider than long, yellow or more or less darkened above. Genae narrow with silvery dusting. Palpi yellow. Male genitalia: cerci rather short and wide, surstyli short with patch of tick setae on inner surface. Body length 1.3–1.5 mm. Distribution. – Sweden: SK, ÖG, VG, BO, SÖ, NB. Finland: Al: Saltvik (R. Frey); N: Ruotsinkylä, 20– 22.vi.1942 (Tahvonen), from cones of Picea, 28 specimens. Not found in Denmark, Norway or Russian parts of Fennoscandia. European species. Biology. – Adults in June to July in woodland. It has been reared from cones of Larix decidua and Picea abies and from a twig of Pinus silvestris (Karps, 1981; Nordlander & Grijpma, 1991; and specimens in collections seen by us). Note. – Oscinis ephippium Zetterstedt has for long been considered as a synonym of Oscinis dubius Macquart, 1835. Our re-examination of the male holotype in Coll. Zetterstedt in ZMLU revealed that Zetterstedt’s species is not identical with the current interpretation of Gaurax dubius (Smith, 1964; Nartshuk, 1970). First flagellomere of G. ephippium is reniform, not round as in G. dubius, and the male genitalia also differ, but coincide with those of G. strobilum Karps.

50. Gaurax fascipes Becker, 1910 Figs 321–328 Gaurax fascipes Becker, 1910: 116. Syntypes in HNHM and probably in ZMHU. Body yellow with three black partly fused stripes on scutum and one black spot on pleura. Sometimes stripes fused only in anterior part of scutum. Central stripe not reaching scutellum, but a blackish spot may be present before scutellum. Scutum around hind dc usually yellow. Frons as long as 104

wide. Ocellar triangle short, extending half length of frons, yellow shining, black spot only on ocellar tubercle. First flagellomere reniform, wider than long. Genae narrow with silvery dusting. Posterior tibiae with small black ring in basal half. Male genitalia: surstyli with rather long narrow process and incision in low angle. Body length 1.3–1.5 mm. Distribution. – Sweden: SK, ÖG, NÄ, SÖ, VS, HS. Norway: AK: Lyndøya, Oslo. Finland: N: Tvärminne (R. Frey). Russia: Vib: Vyborg. Not found in Denmark. Euro-Caucasian species. Biology. – Adults in woodland. Larvae develop in bracket fungi (Piptoporus betulinus), rotting wood and cones of Larix and Picea (Smith, 1964, 1967; Milišauskas, 1976; Skrzypczyńska, 1978, 1982; Grebenshchikova & Naumov, 1979). It has also been reared from crown gall disease caused by Agrobacterium tumefacies (Tschirnhaus, 1992). Collin (1939) recorded that it has been bred from nests of blackbird and linnet in England and also found it in windows of his house. Larvae described by Smith (1967). Some larval structures illustrated by Nartshuk (1987).

51. Gaurax flavomaculatus (Duda, 1933) Figs 329, 330 Goniopsita flavomaculata Duda, 1933: 107. Holotype in ZMHU. Gaurax britannicus Deeming, 1980: 93. Holotype in BMNH. Body black, shining. Head yellow, black on occiput, ocellar triangle black. Eyes uniformly covered with short hairs. Antennae and clypeus black. Face and frons with exception of ocellar triangle and black parts with silvery sheen. Thorax black except yellow on scutellum, notopleura, hind part of postpronota and upper part of anepisternum. Legs yellow with apical part of all femora and basal part of mid and hind tibia infuscate. Wings hyaline. Haltere yellow. Body length about 2 mm. Distribution. – Only Norway: AAy: Lillesand, Sfacing slope, viii–ix.2004, Malaise trap, 5 females. European species. Biology. – Adults in broad-leaf forest. The larvae live under bark of trees, probably feeding on frass or dead insects. In East Europe some specimens were

Figs 321–333. Gaurax fascipes (Meigen): 321: head, dorsal view. 322: head, lateral view. 323: thorax, dorsal view. 324: wing. 325: scutellum. 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 (Nartshuk, 2008).

53. Gaurax leucarista Nartshuk, 1962 Figs 332, 333

52. Gaurax fungivorus Nartshuk & Andersson, sp. nov. Holotype, male: Norway: HOy: Raudli, Os, 20.vi– 16.vii.1991 ex Peptoporus betulinus (K.H. Thunes). Paratypes: 3 males and 1 female pinned, 18 males and 16 females in alcohol, same data as holotype except 20.vi–23.vii.1991, 1 male. Sweden: SÖ: 5 km WSW Sköldinge, Flen, 10.iv.1967, ex Ganoderma applanum (I. Nordin), 1 male, 2 females; UP: Vallentuna, 31.viii.1958 (K.J. Hedquist), 1 male. Holotype, all Swedish paratypes and 11 male and 8 female paratypes from Norway in ZMLU; 2 paratypes from Norway in ZISP; 16 paratypes in alcohol in ZMHU. Diagnosis. – Male genitalia with short surstyli similar to G. macrocerus and G. maculipennis, but G. fungivorus differs from the former by a smaller first flagellomere and longer and narrower male cerci, and from the latter by having clear wings and a small pointed process on surstyli. Description. – Male, female. Head yellow with black occiput. Frons little longer than wide. Ocellar triangle black, thinly dusted, short, extending half length of frons. Setae and setulae of head yellow or black. First flagellomere reniform, wider than long, entirely yellow or darkened on outer margin. Arista black with pale base. Eyes haired. Genae in front equal to width of first flagellomere, widened behind. Palpi yellow. Scutum shining black, only narrow lateral margins yellow, evenly covered with rather long pale setulae. Postpronota yellow with black spot. Scutellum black, or in females yellow or with black sides basally. In females scutal stripes not fused behind leaving yellow spots around hind dc. Pleura yellow with four black, shining spots. npl 1 + 1. Setae of thorax black. Wings not coloured. Halteres yellow. Legs yellow, posterior tibiae darkened around sensory spot. Abdomen black dorsally, epandrium and female cerci black. Male genitalia: cerci rather long, surstyli short with acute process directed to the middle and numerous setae on inner surface. Body length 1.7–2.0 mm. Note. – Larval development of the present new species takes place in bracket fungi. It is thus possible, that existing records of other Gaurax species from bracket fungi refer to the present species, as species of Gaurax are difficult to identify. 106

Gaurax leucarista Nartshuk, 1962a: 255. Holotype in ZISP. Body black, shining. Frons square, with brown silky hairs. orb 5, rather long. Ocellar triangle extending 2/3 length of frons, black, shining. Eyes bare. Genae brown, in front half as wide as first flagellomere. Antennae black, first flagellomere with white pubescence. Arista densely white pubescent and seems to be thickened. Scutum and pleura shining, scutellum dusted. npl 1 + 1. Legs yellow with slight darkening on femora and hind tibia. Halteres white. Body length 1.5 mm. Distribution. – Only known from Russia: Kr: Kondopozhsky District: Kivach Nature Reserve, 1 male, 18–20.vi.1989, Malaise trap in Populus tremula forest (Polevoi). Rare European species described from southern part of Leningrad oblast, where it was caught on young Populus tremula, and further found in Moscow Oblast and Karelia.

54. Gaurax macrocerus (Nartshuk, 1962) Figs 334–337 Botanobia macrocera Nartshuk, 1962a: 253. Holotype in ZISP. Head yellow. Frons as long as wide. Ocellar triangle black, shining, extending half length of frons. Genae narrow with silvery dust. Antennae yellow, first flagellomere in male large, nearly half vertical diameter of eye. Arista black. Scutum black covered with pale dense setulae. Postpronota, notopleura and scutellum yellow or scutellum black. Pleura yellow with four black spots. npl 1 + 1, a second smaller upper hind seta sometimes present. Legs yellow. Abdomen black. Epandrium black with yellow surstyli. Body length 1.3–1.5 mm. Distribution. – Rare species. Sweden: HA, VG, SÖ. Finland: Al: Saltvik, 1 male; Oa: Bergo, 17–28.vi.1946 (Hellén), 1 male; Ks: Kuusamo, 1 male. Not found in Denmark, Norway or Russian parts of Fennoscandia, but described from locality 130 km south from St. Petersburg. European species. Biology. – Adults in woodland.

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 (Zetterstedt, 1848) Figs 338–340 Oscinis maculipennis Zetterstedt, 1848: 2665. Holotype in ZMLU (Andersson, 1966). Head yellow with black occiput. Frons as long as wide. Ocellar triangle short, extending half length of frons, yellow with black spot on ocellar tubercle, dusted. First flagellomere reniform, wider than long, yellow. Genae narrow with silvery dusting. Scutum with three black fused stripes. Postpronota and spots around hind dc yellow, postpronota with black spot. Scutellum blackish. Wings of male with darkening in apical part, well seen in fresh specimens, but nearly disappeared in specimens in collection. Legs yellow. Body length 1.3–1.5 mm.

Distribution. – Sweden: SK (type area); SÖ: Svinmossen, Katrineholm, 12.iv.1966 (O. Elofson), 1 male, 3 females, reared from Polyporus fumetaria on Betula. Finland: Al, Ab, N, Ta, Kb. Not found in Denmark, Norway or Russian parts of Fennoscandia. European boreal species. Biology. – Adults in woodlands, was reared in Finland (Lemland) from tree fungus (swamp, Corylus) (specimens in collection) and from Fomitopsis pinicola (Komonen et al., 2004).

56. Gaurax suecicus Nartshuk & Andersson, sp. nov. Fig. 341 Holotype, male, Sweden: Lule Lappmark: Gällivare, 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: hypandrium and phallic complex.

Diagnosis. – On entirely black thorax similar to G. niger Czerny, but differs by partly yellow head and bare eyes.

Distribution. – Only known from the holotype from Lule Lappmark in Sweden.

Genus Hapleginella Duda, 1933 Description. – Male. Head black with frons in front and genae reddish yellow. Ocellar triangle black, shining. Eyes bare. First flagellomere small, wider than long, reddish yellow, darkened above. Arista black with short pubescence. Setulae on genae pale, head setae brown. Palpi reddish yellow with pale setulae. Thorax all black, scutum shining, covered with long pale setulae, distinctly parted along acrostichal lines. Scutellum dusted, black in posterior half. Apical scutellar setae longer than scutellum, two pairs of short subapical setae. Dorsal setae and setulae of thorax brown, ventral setulae pale. Pleura and postnotum black, shining. Legs slender, blackish. Wings not coloured, costal sectors as 55:60:40:20. R2+3 slightly procurved. R4+5 and M almost parallel. Halteres yellow. Abdomen black, very thinly dusted. Epandrium large, cerci wide apart, surstyli with process directed behind. Body length 1.5 mm. 108

Hapleginella Duda, 1933: 77. Type species: Oscinis laevifrons Loew, 1858 by original designation. Black, shining species. Ocellar triangle large shining with a row of setulae on its surface along side margins. Frons faintly shining. Vibrissal angle right. poc upright, cruciate. First flagellomere rounded. Scutum evenly covered with very short setulae. Tibial organ oval. Femoral organ consists of two rows of warts. Male genitalia: cerci and surstyli simple, hypandrium open, phallapodemic sclerite bowlshaped. Distribution. – Holarctic.

57. Hapleginella laevifrons (Loew, 1858) Figs 18, 36, 53, 342–345, 967–970 Oscinis laevifrons Loew, 1858: 70. Type material not found ZMHU (Wendt, 1975, 1978).

Oscinella inquilina Hendel, 1933: 47. Lectotype in NHMW (Nartshuk, 1997b). Body black, shining. Ocellar triangle smooth shining with a row of setulae on its surface along side margins, reaching margin of frons. Frons faintly shining. First flagellomere yellow, strongly darkened above. Genae half as wide as first flagellomere, yellowish with white dust. Palpi yellow. Scutum shining evenly covered with very short setulae. npl 1 + 2. Scutellum shorter than wide at base, dusted. Apical scutellar setae longer than scutellum. Legs yellow or femora and hind tibiae darkened. Body length 1.5–1.7 mm. Distribution. – Sweden: north to LU. Norway: AK, HEs. Finland: north to Li. Russia: Vib, Kr, Lr. Not found in Denmark. Trans-Palearctic boreal species known from the British Isles to Far East Russia. Biology. – Adults from March to July in woodlands. The larvae develop in cones of different coniferous trees (Pinus, Picea, Abies, Larix) infested by pyralid and tortricid moths or weevils: Dioryctria abietella, Retinia perangustana, Zeiraphera griseana, Clepsis retiferana, Pissodes validirostris (Hendel, 1933; Gaidene & Nartshuk, 1963; Stadnitsky, 1969; Saksons, 1973; Gaidene, 1976; Milišauskas, 1976; Skrzypczyńka, 1978, 1987; Nordlander & Grijpma, 1991). Larvae can develop in other parts of coniferous trees. Nartshuk examined specimens reared from rotting top buds of young seedlings of Pinus sibirica cultivated on the Karelian Isthmus. The larvae are saprophagous scavengers feeding on excrements, dead insects or rotting plant tissues in borrows of primary plant feeders and do not cause primary damage of coniferous cones and seeds. Hibernates as larvae or/and pupae (Gaidene & Nartshuk, 1963; Stadnizky, 1969; Kaidis & Georgevits, 1972; Skrzypczinska, 1974, 1978, 1986; Rognes, 1983; Skrzypczinska et al., 1987; Nordlander & Grijpma, 1991). A note by Duda (1932–1933) that it has been reared from galls of Andricus terminalis (Cynipidae) is questionable in the opinion of Tschirnhaus (1992). Larvae were described by Gaidene & Nartshuk (1963).

Black with densely or thinly dusted scutum. Frons dull, ocellar triangle smooth, dusted. Front margin of frons convex. Vibrissal angle right. poc upright, cruciate. First flagellomere rounded, wider than long. Scutum covered with setulae arranged in distinct rows. Pleura mainly shining. npl 1 + 1 or 1 + 2. Tibial organ oval. Femoral organ consists of two rows of warts with short, stout setulae. Male genitalia: cerci short, surstyli rather broad. Hypandrium closed, rarely open. The species Incertella scotica, I. zuercheri and I. nigrifrons are quite similar to Oscinella and their generic position is not clear. We leave them for the moment in Incertella. Further study is much required to clarify the status and limits of the genusgroup taxa Incertella, Microcercis, Cyclocercula, and Oscinella and probably Rhopalopterum. Distribution. – Holarctic.

Key to species of Incertella Sabrosky 1

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Genus Incertella Sabrosky, 1980 Incertella Sabrosky, 1980: 420. Type species: Oscinella incertella Becker, 1912 by original designation.

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Frons anteriorly and genae yellow or yellowish, yellow part on frons sometimes very narrow. Body with rather thick grey dusting . . . 2 Frons and genae entirely dark. Body with rather thin dusting, subshining . . . . . . . . . . . 4 First flagellomere yellow, darkened only at aristal base. Genae narrow, not wider than front tibiae (Fig. 352). Front and middle tibiae mainly yellow . . . . . . I. antennata (Collin) First flagellomere entirely black . . . . . . . . . . 3 Anterior half of frons and palpi yellow. Genae wider than front tibiae (Fig. 347). All tibiae yellow at both ends. Male genitalia Figs 350, 351 . . . . . . . . . . . . . . . . . . I. albipalpis (Meigen) Frons very narrow yellow anteriorly, sometimes nearly entirely black, palpi black. Genae not wider than front tibiae (Fig. 360). Male genitalia Fig. 351 . . . . . . . . . I. kerteszi (Becker) Wing cell br widened in the middle. Ocellar triangle reaching front margin of frons. Male genitalia Fig. 367 . . . . . . . . I. zuercheri (Duda) Wing cell br not widened in the middle. Ocellar triangle reaching front margin of frons or only half length of frons (Figs 353, 362) . . . . 5 Ocellar triangle reaching front margin of frons, sometimes with shining spots near ocelli (Fig. 362). Anterior notopleural seta weaker 109

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than posterior. Male genitalia Figs 365, 364 . . . . . . . . . . . . . . . . . . . . . . . . I. scotica (Collin) Ocellar triangle extending half length of frons (Fig. 353) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Wings darkened (Figs 356, 357). Male genitalia Fig. 358 . . . . . . . I. karteshensis Nartshuk & Przhiboro Wings clear or slightly greyish . . . . . . . . . . . . . . . . . . . . . . I. nigrifrons (Duda)

(1966) reared it from seedlings of oats and wheat in Norway. Kallio (1950) listed it as a pest of cereals in Finland. It has also been reared from a disposal tip and compost (Werner, 1997), no doubt accidentally with its host plants. Some larval structures illustrated by Kreiter (1928).

59. Incertella antennata (Collin, 1946) Fig. 352

58. Incertella albipalpis (Meigen, 1830) Figs 54, 346–351 Chlorops albipalpis Meigen, 1830: 163; Meigen’s plate 211: 10 (Morge, 1976b). Lectotype in NHMW (Nartshuk, 1997a). Body black, grey-dusted. Frons entirely yellow, occiput and postgenae black grey-dusted. Ocellar triangle black densely grey-dusted, extending half length of frons. Antennae entirely black, first flagellomere large and rounded. Genae pale yellow, slightly narrower than first flagellomere, postgenae black. Palpi yellow. Scutum grey-dusted, setulae arranged in line. Scutellum semicircular, apical setae a little longer than scutellum, wide apart. Subapical setae nearly 1/2 as long as apical. Pleura mainly shining, dusted on upper part of anepisternum. npl 1 + 1. Legs with all femora dark except both ends, hind tibiae with dark band, other tibiae yellow or with dark band. Body length 1.5–1.8 mm. Distribution. – Denmark: common and widespread. Sweden: throughout the country. Norway: Ø, AK, HEs, Bv, VE, VAy, TRy, TRi. Finland: north to Lk. Russia: Vib, Kr, Lr. Trans-Palearctic polyzonal species, known from the British Isles to Japan. Biology. – Common species, adults from the end of May to October in diverse types of meadows, edges of forest, crop fields. Larvae phytophagous or phytosaprophagous, develop in shoots of diverse grasses and cereals (wheat, ray, barley, oats) often damaged by other insects (Kreiter, 1928). As host plants are listed Lolium perenne, Poa spp., Phleum pratense, Festuca pratense, F. rubra, Bromus inermis, Anthoxanthum odoratum, Elytrigia repens, Calamagrostis epigeios and Dactylis glomerata (Karpova, 1965; Tschirnhaus, 1981; Nielsen, 1994). Found on agricultural grasses in Denmark (Nielsen & Nielsen, 1984). Rygg 110

Tropidoscinis antennata Collin, 1946: 136. Syntypes 6 males and 3 females in OXUM (Pont, 1995). Body black, grey-dusted. Frons in front, face and genae yellow. Ocellar triangle reaching middle of frons, dusted. Genae narrow, whitish yellow near eye and dark along ventral margin with a single row of dark setulae. Palpi yellow. oc, orb and vti equal in length, vte a little longer. Antennae yellow with first flagellomere darkened especially on outer side. Setulae on scutum arranged more or less in lines. Pleura shining, except on upper part of anepisternum. All setae and setulae black. Wings slightly brownish below costa and at base. Halteres yellow. Femora black, hind tibiae yellow with black band, other tibiae and tarsi except dark last segments yellow. Body length 1.0–1.25 mm. Distribution. – Denmark: SJ: Sottrupskov, 12.viii.1896 (W. Wüstnei), 1 female; NEZ: Sortemose, Ganløse Orned, 16.vi.1961 (Exp. Zool. Mus.), 1 female. Sweden: SK, SM, VR. Not found in Norway, Finland or Russian parts of Fennoscandia. European species. Biology. – Not common, adults in June to August in meadows.

60. Incertella karteshensis Nartshuk & Przhiboro, 2009 Figs 353–359 Incertella kartashensis Nartshuk & Przhiboro, 2009: 5. Types in ZISP. Body black. Frons entirely black, anterior margin straigth. Ocellar triangle 2/3 as long as frons, densely dusted and not well limited from frons surface. Eyes with sparse fine hairs. Genae narrow. Antennae entirely black, first flagellomere small, wider than long. Arista rather long, short pubescent. Palpi black. Scutum with dense irregular rows of tiny setae. Scutellum small semicircular.

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. Abdomen black, shining. Wings longer than abdomen, intensively darkened except for a lighter area between veins M and CuA1 . Cell br not widened. All femora, front and middle tibiae dark brown to black, hind tibiae yellow with black band in the middle. Body length 1.8–2.2 mm. Distribution. – Found only in Russia: Kr: in the upper, intertidal zone of the White Sea.

dusted, extending half length of frons. Antennae entirely black, first flagellomere rounded. Genae brownish, as wide as 1/2 of first flagellomere. Palpi black. Scutum grey-dusted, setulae arranged in line. Scutellum as long as 1/2 basal width, rounded. Apical scutellar setae little longer than scutellum. Subapical setae nearly 1/3 of apical. Pleura mainly shining, dusted on upper part of anepisternum. npl 1 + 1. Legs mainly black with tarsi yellowish. Body length 1.5–1.8 mm.

Oscinella kerteszi Becker, 1910: 163. Types in HNHM.

Distribution. – Denmark: WJ, NEJ, F, LFM, NEZ, B. Sweden: north to ÅS. Norway: HEs: Abborhøgda, Kongsvinger. Finland: north to Ok. Russia: Vib, Kr, Lr. Trans-Palearctic boreal species, known from the British Isles to Kamchatka.

Body black, grey-dusted. Frons black, narrowly reddish in front. Ocellar triangle black densely grey-

Biology. – Adults from end of May to early October in wet meadows, edges of forests.

61. Incertella kerteszi (Becker, 1910) Figs 360, 361

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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 caespitosa (Nartshuk, 1972a).

Conioscinella nigrifrons Duda, 1933: 161. Syntypes in NHMH.

as wide as first flagellomere. Antennae entirely black, first flagellomere small. Arista very short pubescent. Palpi black. Scutum thinly dusted and subshining. Scutellum semicircular. Apical setae wide apart. Pleura dusted above and shining below. npl 1 + 1. Abdomen black, shining. Wings slightly greyish. Cell br not widened. Legs black, tarsi yellowish-brown. Body length 1.0–1.2 mm.

Smaller and darker than I. kerteszi. Body black. Frons entirely black. Ocellar triangle 2/3 as long as frons, dusted and not well limited from frons surface. Eyes short pubescent. Genae narrow, half

Distribution. – Sweden: SK, HA, SM, ÖL, UP. Finland: Ab, N, Ob. Russia: Lr. Not so far found in Norway and Denmark. Trans-Palearctic species.

62. Incertella nigrifrons (Duda, 1933)

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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 (Collin, 1946)

Biology. – Adults in June to beginning of August, on bogs with Carex. Tyrphobiont species.

Figs 362–365 Tropidoscinis scotica Collin, 1946: 137. Lectotype female and paralectotype female in OXUM (Ismay, 1993; Pont, 1995).

64. Incertella zuercheri (Duda, 1933) Figs 366, 367 Tropidoscinis zuercheri Duda, 1933: 82. Lectotype in ZMAN (Nartshuk, 2003).

Body black, shining, similar to species of the genus Oscinella. Ocellar triangle extends to front margin of frons, covered with very thin dust and subshining, with a row of 6–7 inwardly directed setulae just inside lateral margin and shining spots near ocelli. Genae slightly broader than anterior tibia. npl 1 + 1, posterior seta slightly longer and much stouter than anterior. Femoral comb on male middle femora a single row of setae. Halteres yellow. Legs black with pale femora-tibial joints of anterior and middle legs and all tarsi. Male genitalia: cerci nearly circular, surstyli broad basally with setose inner patch, narrowed apically, hypandrium open. Body length 1.0–1.3 mm.

Similar to species of Oscinella. Body black, shining. Ocellar triangle covered with very thin dust and subshining, reaching front margin of frons. Eyes more longer and densely haired than in Oscinella species. Genae half as wide as first flagellomere. Antennae and palpi black. Scutum and scutellum covered with thin dust and subshining. Apical setae wide apart, longer than scutellum. Pleura mainly shining. npl 1 + 1. Wings longer than abdomen. Halteres yellow. Femora black, hind femora slightly enlarged. Front and middle tibiae black or yellowish at both ends. Body length 1.0–1.3 mm.

Distribution. – Denmark: B. Sweden: SK, HA, SM, UP. Finland: Ab, N, Ob, Ks. Russia: Lr: Lutto River. Not so far found in Norway. European species.

Distribution. – Denmark: EJ: Without locality, 20.viii.1956 (H. Andersson), 1 female. Sweden: SK, HA, SM, UP, NB. Norway: AK, VE. Finland: Ab, N, Ta, Tb, Sb. Russia: Vib, Kr. 113

Eurasian species, known eastwards to Yakutia, East Siberia. Biology. – Adults from middle of May to September on Common reed (Phragmites australis). Larvae phytophagous, live as inquilines in Lipara galls between folded leaves in upper part of galls. Tschirnhaus (1981) listed as host plant also Bolboschoenus maritimus and Glyceria maxima, but these records need to be confirmed. Egg, all larval stages and puparium are described by Grochowska (2008b).

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Genus Lasiambia Anonymous, 1937 Lasiambia Enderlein, 1936: 188. Unavailable. Post1930 name proposed without designation of a type species. Lasiambia Anonymous, 1937: 393 (see Evenhuis et al., 2008: 17). Type species: Oscinella fycoperda Becker, 1910 by original designation. Lasiambia Sabrosky, 1941: 754. Type species: Oscinella fycoperda Becker, 1910 by original designation. Black, shining species with coarsely punctate scutum. Vibrissal angle produced anterior to anterior margin of eye. poc upright and parallel. Genae below the eye is obliquely divided into a rugose and shining posterior and ventral part and a silvery dusted anterior and dorsal part. Most setae are slightly thickened. Facial carina separates entirely or partly the base of antennae. Tibial organ oval. Femoral organ absent. Male genitalia: cerci basally fused, surstyli broad ovate pointed. Hypandrium open. Distribution. – Recorded from all regions, except Australasian.

Key to species of Lasiambia Anonymous, 1937 1

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Genae wider than first flagellomere and vibrissal angle distinctly produced beyond eye margin (Figs 378, 381) . . . . . . . . . . . . . . . . . . . 2 Genae not wider than first flagellomere and vibrissal angle not so distinctly produced beyond eye margin (Figs 368, 372) . . . . . . . . . . 3 Pregenital part of abdomen before epandrium with two membranous vesicules, one on each side, in dry specimens situated within ab-

domen (Fig. 68). Male genitalia Fig. 379 . . . . . . . . . . . . . . . . . . . . . . . . L. coxalis (Roser) Pregenital part of abdomen before epandrium without membranous vesicules (Fig. 387). Male genitalia Figs 385, 386 . . . . . . . . . . . . . . . . . . . . . . . L. palposa (Fallén) Genae more smooth with single row of short setulae along darkened lower margin (Fig. 372). Male genitalia Figs 375–377 . . . . . . . . . . . . . . . . . . . . . L. brevibucca (Duda) Genae with several rows of short setulae along lower margin and genae not so smooth (Fig. 368). Male genitalia Figs 369, 370 . . . . . . . . . . . . . . . . . . . . . . . . . L. baliola Collin

65. Lasiambia baliola (Collin, 1946) Figs 368–370 Fiebrigella baliola Collin, 1946: 60. Lectotype male and 8 paralectotypes in OXUM (Pont, 1995; Ismay, 2000). Body black. Ocellar triangle shining, extending little more than half length of frons. Genae narrower than width of first flagellomere, widened backwards, dark, with setulae in more than a single row. Palpi stout and brownish. Scutum with thin pubescence and rather shining. npl 1 + 2, dorsal hind seta smaller than other ones. Scutellum punctate, undusted on disk and shining. Tergite 5 of abdomen twice length of tergite 4. The apex of tergite 5 with stronger setulae than in L. brevibucca. Femora black, tibiae almost entirely black. Wings brownish along costal half. Halteres yellow. Body length 1.5–1.8 mm. Distribution. – Sweden: SK, SÖ, UP, NB. Finland: Al. Not so far found in Denmark, Norway or Russian parts of Fennoscandia. European species. Biology. – Adults in June to July and again in September in woodland and even in parks in metropolitan area; in London once found on rather large excrescence on the trunk of a wych elm (Allen, 1981a). Larvae develop in rotten wood, especially broad leaved wood. It has been reared from an ulcerous elm tree, sappy chestnut bark, sappy excrescence on trunk of a wych elm (Ulmus glabra) (Collin, 1946; Allen, 1981a; Godfrey, 1998).

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: hypandrium and phallic complex. 377: tip of male abdomen, ventral view. (368–370 after Ismay.)

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66. Lasiambia brevibucca (Duda, 1933) Figs 372–377 Goniopsita brevibucca Duda, 1933: 107. Syntypes in ZMHU, NHMH and HNHM (Wendt, 1975). Body black, shining. Frons shining, ocellar triangle shining as frons, extending 3/5 length of frons. Genae narrower than first flagellomere, yellow with whitish dust and with single row of setulae on narrowly darkened ventral margin. Genae slightly broader in female and setulae in more than one row. Palpi large, yellow. First flagellomere yellow darkened above. Scutum shining densely covered with setulae. Scutellum dusted and slightly rugose. Pleura shining. npl 1 + 2, dorsal hind seta is much smaller. Tergite 5 of abdomen 1.5 times length of tergite 4. Coxae yellow, femora black, tibiae distinctly pale at both ends. Halteres yellow. Body length 1.5–2.0 mm. Distribution. – Denmark: LFM, NWZ, NEZ. Sweden: SK, SM, VG, UP. Norway: AK: Lindøya, Oslo. Finland: Al: Eckerö, 17.vii.1922 (R. Frey), 3 males, 2 females (syntypes); Ab. Not found in Norway or Fennoscandian districts of Russia. European species. Biology. – Adults in June to August in woodland associated with damaged trees and occurs even in parks in metropolitan area; in London once found on rather large excrescence on the trunk of a wych elm (Allen, 1981a). Larvae develop in rotten wood, sap flows of damaged trees and in rot-holes. It has been reared from sappy horse-chestnut (Aesculus hippocastanum) (Godfrey, 1988).

and produced from oral cavity. Scutum shining, densely punctate. Scutellum rounded behind with apical setae shorter than scutellum, subapical setae nearly 1/2 length of apical. Pleura shining. npl 1 + 1. Legs mainly black, but coxae from bright yellow to little paler than remaining parts of legs, ends of femora and tibiae brownish. Male has two pairs of vesiculae of intersegmental membrane, between tergite 5 and synsternite 7 + 8. Usually these vesiculae are inside of abdomen in dry specimens. Body length 2.5 mm. Distribution. – Finland: Al. Russia: Kr. Not found in Denmark, Sweden or Norway. European species. Biology. – Larvae are parasitic and develop in egg masses of Chrysochaon dispar Germar (Acrididae) (Hennig, 1941) and possible of other Acrididae. There are many records of larvae of P. palposa developing in egg pods of Acrididae, but some of these may as well refer to L. coxalis. Hennig (1941) gave drawing of tentoropharyngeal skeleton of larva.

68. Lasiambia palposa (Fallén, 1820) Figs 380–387 Oscinis palposa Fallén, 1820: 6. Lectotype in NRMS (Andersson, 1963). Very similar to L. coxalis, differs in structure of male postabdomen: no vesiculae of intersegmental membrane, between tergite 5 and synsternite 7 + 8. Body length 2.5 mm.

Chlorops coxalis Roser, 1840: 62. Lectotype in SMNS (Nartshuk, 1994). Goniopsita oophila Hennig, 1941: 198 (Nartshuk, 1994). Type specimens in DEIM.

Distribution. – Denmark: WJ, NWJ, NEJ, F, NWZ, B. Sweden: north to VB. Norway: AK, HEs, Bø, VE, TEy, AAy. Finland: north to ObS. Russia: Vib, Kr. Widely distributed Palearctic species, known from Europe eastwards to Yakutia, East Siberia and Mongolia and southwards to Israel. It has been recorded in Iceland by early authors (Mazon, 1890; Meijere, 1910), but was not mentioned by more recent authors (Nielsen & Tuxen, 1954; Lindroth et al., 1973; Messersmith, 1982; Ólafsson, 1991).

Black, shining species. Frons shining, reddish yellow in front and greatly punctate. Ocellar triangle 3/5 length of frons, black smooth shining. First flagellomere rounded, reddish, black above. Genae wider than first flagellomere, whitish yellow along eye margin, brown below. Vibrissal angle produced. Palpi brown to black, large, especially in male

Biology. – Adults from June to the end of September in meadows, wood edges. Larvae carnivorous, live in and may feed on egg pods of Acrididae. As host following species were recorded: Chortippus biguttulus, C. albomarginatus, C. apricarius, Stenobothrus eurasius, S. nigromaculatus, Omocestus viridulus (Frauenfeld, 1863;

67. Lasiambia coxalis (Roser, 1840) Figs 68, 378, 379

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; Nartshuk, 1972b; Tschirnhaus, 1992). It is possible that larvae may feed other food. Adults are often taken in autumn on fallen trunks of Betula in Leningrad oblast and Gusev (1928) found this species on trunks of Picea infested by Ips sexdentatus (Coleoptera). It has been reared from mushroom Leccinum scabrum in Tatarstan (Russia) (Khalidov, 1984). Beschovski (1982) and Beschovski & Georgiev (1993) recorded rearing this species from fruit of Ficus carica and from galls of Paranthrene tabaniformis (Lepidoptera, Aegeridae) in Greece, but maybe this record concerns some other species.

Genus Lipara Meigen, 1830 Lipara Meigen, 1830: 1. Type species: Lipara lucens Meigen, 1830 by monotypy. Large stout black species with hairy eyes, broad facial carina and convex densely haired scutum. Head and thorax setae indistinct hairlike. Pleura with coarsely wrinkled or granulate microsculpture. Tibial organ formed by an oval group of warts. Femoral organ present. Male genitalia: cerci small, sometimes fused, surstyli broad simple, postgonites haired. Adults have sound communication by vibration, sound produced by males being specific, sound produced by females of different species are more similar (Mook & Bruggemann, 1968; Kanmiya, 1981). Assumed mechanoreceptor is situated on the mid coxa (Kanmiya, 1981). The genus includes the largest species of the family. Larvae develop in stems of Common reed (Phragmites australis). The biology of the four species in Europe was studied in detail by Grochowska (2011). Distribution. – Holarctic, Australia.

Key to species of Lipara Meigen

2 – 3 –

Species key to 3rd instar larvae of Lipara Meigen Mature larvae of Lipara 6–10 mm long and 2–3 mm at widest diameter. Body milky white or yellowish, cylindrical or slightly curved. One or three thoracic segments sclerotized and black on dorsal side. Facial mask with ridges in cells. Antennae two segmented, apical segment is situated inside of basal segment. Maxillar palpus surrounded by sclerotized half-ring. Cephalopharyngeal skeleton black, Mandibles with acessory teeth in two rows, hypostomal and tentoropharyngial sclerites fused. Small papillae and spicules are situated on thoracic and abdominal segments. Abdominal segments devoid of sclerotized spicules. Anterior spiracles of fantype, posterior spiracles sessile on surface of 8th abdominal segment, with branched interspiracular processes. Lipara similis Schiner, 1854 (Figs 403–408) from central and western Europe is included in the key, as the species may show up in Denmark and south Sweden. 1

– 2

Lipara similis Schiner, 1854 (Fig. 396) from central and western Europe is included in the key, as the species may show up in Denmark and south Sweden. 1 –

118

Setulae on scutum orientated uniformly backward. Body smaller, up to 4–5 mm . . . . . . . . 3 Setulae on scutum serially directed toward each other producing longitudinal stripes (Fig. 390). Body larger, up to 8 mm. Male genitalia: cerci separated (Figs 391, 396) . . . . . . . . . . . . 2

Setulae on body yellowish. Male genitalia: Figs 391, 392 . . . . . . . . . . . . . . . . . . L. lucens Meigen Setulae on body whitish. Male genitalia: Fig. 396 . . . . . . . . . . . . . . . . . . . . . L. similis Schiner Male genitalia: cerci fused and far projecting (Fig. 73) . . . . . L. pullitarsis Doskočil & Chvála Male genitalia: cerci separated (Fig. 395) . . . . . . . . . . . . . . . . . . . . . . . . L. rufijitarsis Loew

– 3

Only one or three anterior segments of body black dorsally, posterior segment not black (Figs 398, 399, 409, 410, 415, 416) . . . . . . . . . 2 Anterior and posterior segments of body black (Figs 403, 404) . . . . . . . . . . . . L. similis Schiner Three anterior segments black dorsally (Fig. 399). Mandibles with accessory teeth in two rows, 3 in each row (Fig. 402). Larva 8–10 mm long milky white in cigar-like gall with hard shortened internodes . . . . . . L. lucens Meigen Only one anterior segment black dorsally or dorsally and ventrally (Figs 410, 416) . . . . . . . 3 First thoracic segment of body black dorsally and ventrally (Fig. 416). There are small spicules on the second segment of body (first thoracic) ventrally and around anal opening

–

(Fig. 416). Mandibles with accessory teeth in two rows, a tooth in each row (Figs 417, 418). Anterior spiracles 6–9 buds (Fig. 419). Larva milky white flattened ventrally 8–9 mm long (Fig. 415) in slightly enlarged stem of Pragmites . . . . . . . . . . . . . . . . . . . . L. rufijitarsis Loew First segment of body black only dorsally (Fig. 410). There are small spicules on the first segment ventrally, on the second segment of body (first thoracic) dorsally and ventrally and on the third segment (the second thoracic) ventrally (Fig. 410). Mandibles usually with two additional teeth in each row (Figs 411, 412). Anterior spiracles 4–5 buds (Fig. 413). Larva 6–9 mm long milky white (Fig. 409) in cigarlike gall . . . . . . L. pullitarsis Doskočil & Chvála

69. Lipara lucens Meigen, 1830 Figs 388–394, 397–402 Lipara lucens Meigen, 1830: 1; Meigen’s plate 210: 5 (Morge, 1976b). Holotype in NHMW (Nartshuk, 1997a). Large robust species, the largest species of the genus and of the family in the Palearctic region. Body black densely covered with long yellowish hairs, arranged on scutum in distinct longitudinal stripes with hairs directed right and left. Antennae with basal segments yellowish, first flagellomere mainly blackish. Antennae divided by rather broad facial carina. Genae wide. Pleura roughly wrinkled. Legs darkened. Body length 5.5–8.0 mm. Distribution. – Denmark: SJ, EJ, WJ, NEJ, LFM, SZ, NEZ, B. Sweden: SK, SM, VG, UP, VS. Norway: AK, Bø, some specimens in NHMW without exact locality and data. Finland: N, Ta, Sa, Kb. Russia: Vib. Euro-Mediterranean species. Sabrosky (1958) considered the species as introduced in North America. Biology. – Adults in May to June on Phragmites australis, occurs predominantly on suboptimally developed reed growing on dry soil (Mook, 1971; Grochowska, 2011). Kontkanen (1939) described habitats in Finland. There is only one generation per year. Overwinter as larvae inside the gall. Adults emerge from the end of May until early June. Adults have sound communication by vibration, sounds produced

by males being specific, sounds produced by females of different species are more similar (Mook & Bruggemann, 1968). Kanmiya (1990) described geographical variation in the vibratory courtship signals of the species. Newly emerged female carry mature eggs and can mate immediately. Eggs are laid externally on the surface of the reed stems or leaves. Larvae phytophagous, producing terminal cigar- or spike-like gall on shoots of Phragmites australis. After hatching the larvae crawl to the top, enter under the edge of a leaf sheath and gnaw their way down through the enwrapped leaves until they reach the growth point. Larvae feed on the enwrapped leaves and later enter the gall chamber. At the end of summer the full grown larvae enter diapause. The following spring larvae pupate in the chamber. Galls rather thick and contain a hardened gall chamber. Common reed growing on land with a diameter about 3–4 mm are preferred as host plants (Valkeila, 1956; Mook, 1961, 1971; Theowald, 1961; De Bruyn, 1987; Wolcsánszky et al., 1993). Females avoid to lay eggs on reed standing in water. The cause of this behaviour is explained by anatomical structure and biology of the reed showing xerophilous and hygrophilous habit (Ambrus, 1977). The meristem of infested plants stops functioning. Several histological changes are caused by modified metabolism of the tissues: sclerotisation, development of lignine deposits, accumulation of amylum in the ground tissue and clogging of the phloem or rhexigenous vessels (Reynvaan & Docters van Leeuwen, 1906; Wolcsánszky et al., 1993). Gromyssz-Kałkowska & Hubicka (1978) studied oxygen consumption during larval development of the species. Morphological and histochemical analysis of galls and effect of gall formation on the host plant were studied by De Bruyn et al. (1998), Vandevyvere & De Bruyn (1998). These authors describe the process of cecidogenesis in detail. Gall formation starts when the larvae feed outside the actual developing gall, on the enwrapped leaves. During gall growth internodes elongation is reduced. Internally the pith parenchyma is destined to become the nutritive tissue. The tissue cylinder around the gall chamber widens up to three times normal size. When gall is completed, the larva gnaws through the growing point and enters the gall chamber, where it consumes the nutritive tissues. The sclerenchymatization process starts now resulting in an extremely hardened gall. Many species of Chloropidae, species of other families of Diptera and other groups of arthropods 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, C. duinensis, Incertella zuercheri, Elachiptera cornuta, Eribolus hungaricus, Oscinella angustipennis, Cryptonevra diadema, C. flavitarsis, C. consimilis. Two more European species of Chloropidae Homalura tarsata Meigen and C. nigritarsis (Duda) are associated with Lipara galls. They are not known from Scandinavia, but may show up in southern part of Denmark. Records of Oscinella frit and O. pusilla from galls (Giraud, 1863; Blair, 1932; Theowald, 1961) probably concern Incertella zuercheri. Some authors (Seguy, 1934; Wend, 1968) recorded Oscinella nitidissima from Lipara galls. Cecidomyiidae: Giraudiella inclusa (Frauenfeldt), Microlasioptera flexuosa (Winnertz), Lasioptera hungarica Möhn, L. arundinis Schiner, Asynapta phragmitis (Giraud). Anthomyzidae: Anthomyza collini Andersson, A. gracilis (Fallén). Agromyzidae: Phytomyza succisae Hering (Theowald, 1961). The last species is usually known as a leaf-miner on Succisa pratensis. Therefore, the record from the Lipara galls is doubtful. Many bees and wasps make their nests in 2-years vacated galls (see chapter ‘Role in land ecosystems and species of economic importance’). Several species of mites (Acarina) were recorded within galls of Lipara lucens and L. pullitarsis: Siteroptes (S.) avenae (Müller), S. (S.) reniformis Krantz (Pygmephoridae), Tarsonemus lacustris Schaarschmidt, Stenotarsonemus phragmitidis (v. Schlechtendal) and S. (Mahunkacus) gibber Suski (Tarsonemidae), Urodiaspis tecta (Kramer) (Urodiaspidae), Tydeus sp. (Tydeidae) (Imhof, 1979; Rach & Carstensen, 1981; Abraham & Carstensen, 1982). The bushcricket Conocephalus dorsalis (Latreille) (Conocephalidae) is also associated with Lipara galls (Waitzbauer, 1969). Recently Reader (2003) described interaction between L. lucens and Cryptonevra flavitarsis (on my opinion erroneously named as L. rufijitarsis in the paper) in the term of intraguild kleptoparasitism. It is difficult to agree with this point of view, as larvae of these species feed different part of plant and feeding of larvae Cryptonevra flavitarsis is not influenced on gall formers. For parasitoids, predators and other inhabitants in Lipara galls see the chapters ‘Parasitoids, predators and parasites’ and ‘Role in land ecosystems and species of economic importance’. The most detailed descriptions of egg and larvae are done by Waitzbauer (1969), Chvála et al. (1974) and Beschovski (1995).

70. Lipara pullitarsis Doskočil & Chvála, 1971 Figs 73, 409–414 Lipara pullitarsis Doskočil & Chvála, 1971: 102. Holotype in DECU, paratypes in DECU, NMPC and BMNH. Black, shining robust species, but distinctly smaller than L. lucens. Scutum evenly and densely covered with yellowish setulae not arranged in stripes. Antennae mainly black, basal segments sometimes brown. Facial carina broader than in L. lucens, half width of first flagellomere. Legs black with yellow base of tarsi. Body length 3.5–5.0 mm. Distribution. – Denmark: NEZ. Sweden: SK, HA, SM, SÖ, UP, DR. Norway: AK, Bø, VE, AAy. Finland: Ab, N. Russia: Vib. Euro-Mediterranean species. Biology. – Adults in May to June on Phragmites australis. Larvae phytophagous, produce terminal gall on shoots of Phragmites australis. Galls are soft and do not contain a hardened gall chamber. Flies preferable infest thicker stems of host plant than L. lucens (De Bruyn, 1987). Some species of Chloropidae and other insects live as inquilines in folded leaves in upper part of the gall. All species of inquilines are listed under L. lucens. Larvae described by Chvála et al. (1974), egg, all larval stages, puparium and biology described by Grochowska (2006, 2011).

71. Lipara rufijitarsis Loew, 1858 Figs 395, 415–420 Lipara rufijitarsis Loew, 1858: 57. Lectotype in ZMHU, designated by Doskočil & Chvála (1971). Wend (1975) listed the species in the Index, but not in the main text of her paper. Black, shining robust species, but distinctly smaller than L. lucens. Scutum evenly and densely covered with whitish setulae, which not arranged in stripes. Antennae mainly yellow, basal segments sometimes brown. Facial carina narrower than in L. pullitarsis, in the middle scarcely one third as wide a first flagellomere, but distinctly widening above and below. Legs black with bright yellow tarsi. Body 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 collected 03.iii.1951, flies emerged 22.iv.1951 (S. Johansson), 2 females and a puparium. Even if only females are available, we are confident about the identification. European species, known from Central Europe. Biology. – Larvae are phytophagous as in other species of the genus and produce a top gall on shoots of Phragmites australis. Usually galls are narrower than galls of L. lucens, but sometimes they are indistinguishable from galls of L. lucens (Chvála et al., 1974). Egg, all larval stages, puparium and biology described by Grochwska (2007a, 2011). 122

Genus Microcercis Beschovski, 1978 Microcercis Beschovski, 1978: 28, as subgenus of Oscinella Becker. Type species: Oscinella trigonella Duda, 1933 by original designation. Body black. Ocellar triangle shining, extending half length of frons. Genae narrow. Antennae and palpi black. First flagellomere round. Scutum shining with setulae arranged more or less in rows. Scutellum and hind part of scutum before scutellum dusted. Male genitalia: cerci small. Surstyli short wide, not tapering. Hypandrium closed. Distribution. – Palearctic.

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 Incertella, Conioscinella incerta Becker, is a nonPalearctic 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 1

–

Scutum shining except part before scutellum. Acrostichal and dorsocentral setulae arranged in double rows. Male genitalia Figs 421, 422 . . . . . . . . . . . . . . . . . . . . . M. trigonella (Duda) Scutum dusted as in most species of Oscinella. Acrostichal setae arranged in double rows . . . . . . . . . . . . . . . . . . . . . . . M. kroeberi (Duda)

72. Microcercis kroeberi (Duda, 1935), comb. & stat. nov. Oscinella kroeberi Duda, 1935: 158, as var. of O. frit (Linnaeus). Syntypes in ZMHU. Body black. Frons dull, brownish in front. Ocellar triangle shining, extending half length of frons. Genae as narrow as front tibia. Antennae and palpi black. First flagellomere round. Scutum dusted, with setulae arranged in double rows along acrostichal lines. Scutellum dusted, apical setae longer than scutellum, wide apart, one very short subapical seta on each side. Pleura mainly shining, except on upper part. npl 1 + 1. Legs black, except brownish tarsi. Wings slightly brownish. Body length 1.0– 1.5 mm.

Figs 421–423. Microcercis trigonella (Duda): 421: hypopygium, dorsal view. 422: surstylus, lateral view. 423: hypandrium and phallic complex.

1984b), where it was listed as a valid species in Oscinella. Unfortunately only females are present in the available material deposited in the Zoological Museum, Hamburg. Without being able to examine the male genitalia we provisionally assign kroeberi to the genus Microcercis based on following characters: frons dull and straight in front (not concave as in Oscinella), ocellar triangle short and shining, acrostichal setulae arranged in double rows, npl setae 1 + 1 (in Oscinella usually 1 + 2). For a discussion about relationships between Microcercis and Incertella see under Microcercis.

73. Microcercis trigonella (Duda, 1933) Figs 421–423 Oscinella trigonella Duda, 1933: 196. Syntypes in HNHM.

Distribution. – Denmark: SJ: Sønderborg, vi.1897 (W. Wüstnei), 1 female; WJ: Tipperne, viii.1973 (E. Rald), 2 females; NEJ: Kringelrøn & Hornfiskrøn, Læsø, 14.vii.1969 (Zool. Mus. Copenh. Exp.), 1 female; F: Keldsnor, Langeland, 06.viii.1975 (Lyneborg, Martin & Michelsen), 2 females. Russia: Kr: Kartesh, 21–25.vii.1993 (Sugonyaev), 1 female.

Body black. Frons dull, brownish in front. Ocellar triangle shining, extending half length of frons. Genae as narrow as front tibia, brownish. Antennae and palpi black. First flagellomere round. Scutum mainly shining, with setulae arranged in double rows along acrostichal and dorsocentral lines. Scutellum and hind part of scutum before scutellum dusted. npl 1 + 1. Legs mainly black, but tibiae yellowish at ends. Body length 1.0–1.5 mm.

Note. – Duda (1935) described kroeberi from northern Germany as a variety of Oscinella frit. This taxon has remained unrecognized in the literature except in the Catalogue of Palaearctic Diptera (Nartshuk,

Distribution. – Denmark: SJ, EJ, WJ, LFM. Sweden: north to TO. Norway: Bv. Finland: north to Le. Russia: Vib, Kr, Lr. Eurasian species. 123

Biology. – Adults from the end of May to early August in wet meadows. Collin (1939) recorded numerous specimens bred by Mr. Basden from mole’s nest in June in England. Larvae live in stems of Juncus efffusum, J. gerardii, Glyceria fluitans, Phalaris arundinaceae, Festuca arundinaceae and Deschampsia caespitosa (Rogochaya, 1960; Sharipov, 1981; Tschirnhaus, 1981).

Key to species of Oscinella Becker 1 – 2 –

Genus Oscinella Becker, 1909 Oscinella Becker, 1909: 120. Type species: Musca frit Linnaeus, 1758 by designation of ICZN, 1978: 203 (Opinion 1100). Pachychoeta Bezzi, 1895: 72 (preocc. Bigot, 1857). Type species: Elachiptera aterrima Strobl, 1880 (= Oscinis capreolus Haliday, 1838) by original designation. Melanochaeta Bezzi, 1906: 50, as replacement name for Pachychoeta Bezzi, 1895. Type species: Elachiptera aterrima Strobl, 1880 (= Oscinis capreolus Haliday, 1838), automatic. Pachychaetina Hendel, 1907: 98, as replacement name for Pachychoeta Bezzi, 1895. Type species: Elachiptera aterrima Strobl, 1880 (= Oscinis capreolus Haliday, 1838), automatic. Note. – An application to the ICZN (Case 3576) to conserve Oscinella Becker, 1909 by reversal of precedence with Melanochaeta Bezzi, 1906 and Pachychaetina Hendel, 1907 has been made by Tschirnhaus & Nartshuk (2012). Black species mainly with dusted or rarely shining scutum. Frons dull, ocellar triangle large shining. Front margin of frons concave. Vibrissal angle right or slightly retreated. poc upright, cruciate. First flagellomere rounded, as long as wide. Arista mainly slender dark with short pubescence or rarely white with dense pubescence or enlarged with black pubescence. Scutum convex with setulae arranged in lines. Pleura mainly shining. Tibial organ oval. Femoral organ about 6–9 warts in a row. Male genitalia: cerci discrete and large, surstyli simple, hypandrium open. The genus is divided into two subgenera: Oscinella s. str. has large, more or less triangular male cerci; Cyclocercula Beschovski, 1978 has nearly circular male cerci. Distribution. – Worldwide. 124

3 – 4 – 5 – 6

–

7

– 8 – 9

–

10

Scutum shining . . . . . O. nitidissima (Meigen) Scutum dusted, not shining, only postpronota may be shining . . . . . . . . . . . . . . . . . . . . . . . . . 2 Arista densely pubescent, enlarged, white or black (Fig. 21). Scutum densely dusted . . . . . 3 Arista dark, rather thinly pubescent. Scutum less densely dusted . . . . . . . . . . . . . . . . . . . . . 4 Arista white (Fig. 439) . . . . O. maura (Fallén) Arista black. Male genitalia Figs 429, 430 . . . . . . . . . . . . . . . . . . . . O. capreolus (Haliday) Postpronota shining, scutum dusted . . . . . . . . . . . . . . . . . . . . O. ventricosi Nartshuk Postpronota and scutum both dusted . . . . . . 5 Last segment of all tarsi widened (Fig. 448) ......................................6 Last segment of all tarsi not widened . . . . . . 7 Legs more extensively dark, only extreme tips of femora yellow, trochanter of front legs black. Vibrissal angle rounded (Fig. 424). Ocellar triangle dusted along sides . . . . . . . . . . . . . . . . . . . . . . . O. angularis Collin Legs less extensively dark, all femora with broadly yellow tips, trochanter of front legs yellow. Vibrissal angle rather angular (Fig. 447). Ocellar triangle not dusted along sides . . . . . . . . . . . . . . . . . . . O. trochanterata Collin Wings long and narrow, c. 3 times longer than wide (Fig. 425). Male genitalia Figs 426, 427 . . . . . . . . . . . . . . . . . . . . O. angustipennis Duda Wings wider, only about 2.5 times longer than wide (Fig. 433) . . . . . . . . . . . . . . . . . . . . . . . . . 8 Genae very narrow, not as wide as front tibiae . . . . . . . . . . . . . . . . . . . . . . O. cariciphila Collin Genae not so narrow, wider than front tibiae ......................................9 Tibiae of front and middle legs entirely yellow, tibiae of hind legs black on middle part. Male genitalia Figs 445, 446 . . . O. pusilla (Meigen) Legs variably but more extensively dark, tibiae never entirely yellow. (O. frit (Linnaeus) species complex) . . . . . . . . . . . . . . . . . . . . . . 10 Legs entirely black . . . . . . . . . . . . . . . . . . O. nigerrima (Macquart)

– 11

– 12

–

Legs not entirely black, at least part of front and middle tibiae yellow or yellowish . . . . . 11 Genae distinctly wider than first flagellomere. Wings darkened and usually not reaching end of abdomen . . . . . . . . . . . . O. vastator (Curtis) Genae not wider than first flagellomere . . . 12 Hind tibiae entirely dark, front and middle tibiae never entirely yellow. Genae wider, nearly as wide as first flagellomere. Wings clear and reaching end of abdomen. Male genitalia Figs 434–436 . . . . . . . . . . . . . . . . . O. frit (Linnaeus) Hind tibiae dark with yellowish base and tip, front and middle tibiae extensively or entirely yellow. Genae distinctly narrower than first flagellomere. Wings a little darkened . . . . . . . . . . . . . . . . . . . . . . . O. hortensis Collin

Note. – “Oscinella frit” in the traditional, broad sense is a complex of several very similar and variable species, e.g. in respect to the width of genae and colour of legs, and thus very difficult to identify. From this complex only species found regularly in crop fields are included into the key. The Oscinella frit complex consists of the following taxa either considered as distinct species or forms of doubtful taxonomical status, part of them described exclusively or predominantly on larval stages: O. agropyri Mesnil, described from France on larval stage in Balachowsky & Mesnil, 1935 O. alopecuri Mesnil, described from France on larval stage in Balachowsky & Mesnil, 1935 O. festuceae Mesnil, described from France on larval stage in Balachowsky & Mesnil, 1935 O. frit, form Arrhanatherum, described on larval stage by Jepson & Nye (1957) and Nye (1958) O. frit, form Anthoxanthum, described on larval stage by Jepson & Nye (1957) and Nye (1958) O. exigua Collin, 1946, described from England as var. of O. frit (Linnaeus) O. hortensis Collin, 1946; England O. nigerrima (Macquart, 1835); France O. phlei Nartshuk, 1956; European part of Russia O. pusilla (Meigen, 1830); Germany O. vastator (Curtis, 1845); England Several other names are customarily listed as synonyms of O. frit (Linnaeus), even though the type specimens no longer exist, or have never been adequately examined:

Musca hordei Bjerkander 1777; Sweden Musca avenae Bjerkander 1781; Sweden Oscinis flavipes Olivier 1813; France Oscinis nigra Olivier 1813; France Chlorops fumipennis Meigen 1830; Germany Oscinis brunnitarsis Macquart 1835; France Hydrellia rufijitarsis Meigen 1838; Germany Chlorops nigrita Meigen 1838; Germany Chlorops aenea Roser 1840; Germany Chlorops distincta Roser 1840; Germany Chlorops nigrimana Roser 1840; Germany Oscinis granarius Curtis 1846; England Oscinis frit var. nigripes Strobl 1894; Austria Oscinella frit var. plumiseta Duda 1933; Europe Oscinella sziladyi Duda 1933; Bulgaria The applied literature dealing with frit flies as a pest in crops of cereals and fodder grasses in Europe, notably from England, Germany, France, Denmark, Sweden and Norway, usually distinguishes between 3 species: Oscinella frit, O. pusilla and O. vastator, sometimes even O. hortensis. Diagnostic characters used to distinguish the first three species are given in the chapter ‘Role in land ecosystems and species of economic importance.’ The relative length of the second and third segment of arista helps to separate species occurring in agricultural fields or reared from cereals and cultivated grasses, but this character may not work in relation to other species of the complex. An attempt to solve the taxonomy of the O. frit complex using caryological methods (Michailova & Beschovski, 1985) did not give unambiguous results. The chromosomes found were long and slender and difficult to study. Extended breeding experiments with pure cultures and molecular studies may help to settle the outstanding taxonomic problems still pending within the Oscinella frit species complex.

Species key to known 3rd instar larvae of Oscinella 1

– 2

3rd thoracic segment on ventral side covered with one or several rows of spicules (Figs 451, 452, 453, 457, 458) . . . . . . . . . . . . . . . . . . . . . . 2 3rd thoracic segment without ventral rows of spicules (Fig. 450) . . . . . . . O. angularis Collin Ventral creeping welts of abdominal segments with one or two rows of enlarged spicules (Figs 453, 457, 458) . . . . . . . . . . . . . . . . . . . . . . . . . . 3 125

–

3

–

4

–

5

–

Ventral creeping welts of abdominal segments without rows of enlarged spicules, if some large specules present, they do not form discrete rows (Figs 451, 452) . . . . . . . . . . . . . . . . 5 Ventral creeping welts of abdominal segments with two rows of enlarged spicules (Fig. 458) . . . . . . . . . . . . . . . . . . . . O. ventricosi Nartshuk Ventral creeping welts of abdominal segments only with one row of enlarged spicules (Figs 453, 457) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 On creeping welts of abdominal segments behind of a row from larger spicules only a row of smaller spicules presents (Fig. 457) . . . . . . . . . . . . . . . . . . . . . . . O. pusilla (Meigen) On creeping welts of abdominal segments two rows from smaller spicules presents (Fig. 453) . . . . . . . . . . . . . . . . . . . O. nitidissima (Meigen) On creeping welts of abdominal segments small spicules form 3–4 rows (Fig. 452). Stigmophores closed, distance between them shorter than a stigmophore. Larvae yellowish . . . . . . . . . . . . . . . . . . . . . . . . O. maura (Fallén) On creeping welts of abdominal segments small spicules form only a row (Fig. 451)Larvae white . . . . . . . . . . . . . . . . O. frit (Linnaeus)

74. Oscinella (Oscinella) angularis Collin, 1946 Figs 424, 449, 450 Oscinella angularis Collin, 1946: 141. Syntypes: 13 males, 4 females in OXUM, 1 female in BMNH (Pont, 1995). Body black. Ocellar triangle extending nearly to front margin of frons, shining, but covered with thin dust at tip and along sides. Genae with rounded vibrissal angle and head in side view rather retreating. Scutum dusted. Legs black with knees and tip of tibiae yellowish. Middle and hind tarsi yellow at base. The last segments of all tarsi enlarged. Knob of halteres yellow, stem dark. Body length 1.5–2.0 mm. Distribution. – Denmark: F: Humble, Langeland, 20.vii.1964 (O. Martin), 1 female. Sweden: SK, SM. Finland: N, Ta, Om. Russia: Vib. Not so far found in Norway. European species, but found in North America, probably introduced (Nartshuk, 2001). 126

Biology. – Adults in June to July and in September in wet meadows and marshy habitats. Larvae phytophagous, live gregariously in shoots of Phalaris arundinaceae (Collin, 1946; Nartshuk, 1956; Svirina, 1984). The tissues of the host plant above the feeding site of larvae become brown and the central leaf dries up. Larva described by Nartshuk (1956).

75. Oscinella (Cyclocercula) angustipennis (Duda, 1932) Figs 425–427 Oscinella angustipennis Duda, 1932: 87. Lectotype in ZMAN (Nartshuk, 2003). Body black, shining. Frons dull. Ocellar triangle smooth shining extending 3/4 length of frons. Genae half as wide as first flagellomere. First flagellomere rounded. Arista longer haired than in O. frit. Scutum and scutellum dusted. Legs black with brownish tarsi. Wings long and narrow, 3 times longer than wide, sometimes slightly darkened. Vein R2+3 distinctly concave. Halteres yellow. Body length 1.0–1.3 mm. Distribution. – Found only in Russian parts of Fennoscandia: Vib: Jukki; Peri. European species. Biology. – Adult in June to July in wet meadows and Phragmites beds. It has been reared from Lipara galls in Netherlands (De Bruyn, 1985). Larvae live between the enwrapping leaves in upper part of gall. Ismay (1981b) found the species on Glyceria sp. on marsh. Uffen & Chandler (1978) recorded Phalaris arundinaceae as food plant, but Drake (1988) recorded no correlation between the occurrence of this plant and the fly in Great Britain.

76. Oscinella (Oscinella) capreolus (Haliday, 1838), comb. n. Figs 21, 428–430 Oscinis capreolus Haliday, 1838: 187. Described from ‘England’. Whereabouts of type material unknown. Ismay et al. (2001) did not mention this species when revising Haliday’s collection. Differs from other Oscinella species by flattened and densely pubescent black arista. Body black.

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, reaching front margin of frons. 4 pairs of orb, longer than remaining setulae of frons. First flagellomere rounded. Genae narrower than first flagellomere. Scutum and scutellum dusted, scutellum as long as wide at base with broadly rounded apex. Pleura mainly shining. npl 1 + 2. Wings brownish. Halteres dark. Body length nearly 2 mm. Distribution. – Rare species, found only in Denmark: LFM: Resle, 28.v.1882 (H.J. Hansen), 1 female. Euro-Caucasian species. Note. – Collin (1946) stated about Melanochaeta capreola (Haliday) that it “except for its arista, might easily pass as a species of Oscinella”. Tschirnhaus (1992) even expressed the opinion that Haliday’s species rightly belongs to Oscinella. Examination of the male genitalia from specimens kindly sent by Tschirnhaus to Nartshuk supports this opinion. Oscinis capreolus has male genitalia of Oscinella type with rather big, nearly triangular cerci and simple gonites – not widened in apical part as in the Palearctic Melanochaeta pubescens (Thalhammer). We thus propose the new combination Oscinella capreola (Haliday), and accordingly treat the following genus-group names, that have Oscinis capreolus Haliday, 1838 as type species, in synonymy with Oscinella Becker, 1909: Melanochaeta

Bezzi, 1906, Pachychoeta Bezzi, 1895, and Pachychaetina Hendel, 1907. As the generic name Melanochaeta Bezzi, 1906 has priority over the widely used generic name Oscinella Becker, 1909, an appeal has been made to the ICZN to conserve of the generic name Oscinella Becker, 1909 and include it into Official List of Generic Names in Zoology (Tschirnhaus & Nartshuk, 1912). Other Palearctic species currently listed within the genus Melanochaeta are now transferred to the genus Lasiochaeta Corti, 1909 (type species: Elachiptera pubescens Thalhammer), now restored from synonymy with Melanochaeta. The male genitalia of Lasiochaeta pubescens are of the Elachiptera type with apically widened surstyli, see Andersson (1977: fig. 31).

77. Oscinella (Oscinella) cariciphila Collin, 1946 Oscinella cariciphila Collin, 1946: 141. Syntypes: 20 males and 8 females in OXUM (Pont, 1995). Body black, shining. Ocellar triangle reaching front margin of frons. Genae not wider then fore tibia. Scutum and scutellum dusted, pleura mainly shining. npl 1 + 2. Legs with front and middle femora very narrowly yellow at tip, front and middle tibiae 127

entirely yellow, hind tibiae black and only narrowly fuscous yellow about base and tip. Wings rather short and broad. Halteres yellow with brownish stem. Body length 1.5–2.0 mm. Distribution. – Sweden: SK, HA, GO, NÄ, UP, VR, DR, HS. Norway: Ø, AK, HEs, VE, STi. Finland: N. Russia: Vib; Kr: Kondopozhsky District: Kon, Bukolnikov Is., 24.vi.2003 (Polevoi), 1 male; Kartesh, 16– 29.vii.1966 (Tanasijtshuk), 2 females. Not so far found in Denmark. Eurasian species. Biology. – Adults in June in wet habitats with Carex, probably larvae live in shoots of Carex.

78. Oscinella (Oscinella) frit (Linnaeus, 1758) Figs 25, 55, 85, 95, 96, 103, 431–437, 451 Musca frit Linnaeus, 1758: 598. Body black. Frons dull with smooth shining ocellar triangle reaching front margin of frons. First flagellomere rounded. Second aristal segment 0.5 times as long as third aristal segment. Warts in femoral comb situated close each other and rarely fused. Genae nearly as wide as first flagellomere or wider. Scutum and scutellum dusted. Pleura mainly shining. npl 1 + 2. Legs mainly black or front and middle tibiae partly yellow. Body length 1.5–2.0 mm. Distribution. – One of the commonest species of the family. Widely distributed species (or group of closely related species), known from all of Fennoscandia to the coast of Arctic Ocean. The species has been found in all zoogeographical regions of the World. Biology. – Adults in May to September in meadows, wood edges, crop fields. Two or three generation per year in Nordic countries. Overwinter as larvae within tillers of grasses. Adults have been found on xerothermic locality in November and December in Germany (Thuringia) (Bährmann, 1996). Adults are very vagile, found up to 300 m above ground (Johnson et al., 1962). Thunes et al. (2004) found adult in canopies of Pinus silvestris in Norway. Larvae phytophagous, develop in shoots and seeds of cereals: wheat, barley, rye, oats and in shoots of many grasses. The species damages cereals and fodder grasses in all countries where it occurs, and there exist numerous publications on the biology and control of this species from all Nordic 128

countries (see chapter ‘Role in land ecosystem and species of economic importance’). More than 60 species of wild grasses are recorded as host plants, but notably the species does not develop in shoots of Dactylis glomerata (Vickerman, 1978a). However, the uncertain taxonomy within the O. frit species complex suggests that some records refer to other, closely related species. Usually polyphagous herbivorous species exhibit a distinct preference for one or a few host species. Nartshuk (1956) assumed that O. frit prefers wild grasses of the tribe Avenae. Vickerman (1978b) found out that the survival of O. frit larvae was lower on barley than on either oats or wheat. The females laid most eggs on oats, Lolium perenne and Festuca rubra, but relatively few eggs on barley in laboratory experiments (Vickerman, 1978a). Larvae feeding on different grasses may differ in the arrangement of spicules on ventral side of the body. Jepson & Nye (1957, 1959) and Nye (1958) described such differences in larvae from Arrhenatherum and Anthoxanthum, respectively. It is not clear if these larvae belong to different species or exhibit variation within O. frit under influence of the different host plants. It has been shown at least that no cross-breading takes place between O. frit and O. pusilla (Le Berre & Chevin, 1963; Agapova, 1966). Le Berre & Chevin (1961) demonstrated in experiments that colour of tibiae and arrangement of spicules on thoracic and abdominal segments of larvae are characters dependent on both temperature and host plant. O. frit has been reared from disposal dumps and compost (Werner, 1997), without doubt from larvae occurring in host plant remains. Larvae described by many authors, but in most detail by Steel (1931, all instars), Balachowsky & Mesnil (1935), Riggert (1935), Nartshuk (1956, all instars), Jepson & Nye (1957), Nye (1958), Jepson & Southwood (1960a, first instar).

79. Oscinella (Oscinella) hortensis Collin, 1946 Fig. 438 Oscinella hortensis Collin, 1946: 142. Syntypes: 44 males and 14 females in OXUM (Pont, 1995). Similar to O. frit, but colour of legs paler. Collin wrote that in colour of legs and width of genae this species is intermediate between O. cariciphila and O. pusilla. Shining ocellar triangle almost reaching front margin of frons. First flagellomere small as in O. pusilla but genae narrower though not so narrow

as in O. cariciphila. Arista pubescence shorter. Scutum more finely punctuated and less dusted than in O. pusilla. Femora black, front and middle tibiae yellow with black ring about middle. Hind tibiae darker. Male genitalia: surstyli narrower than in O. pusilla. Body length 1.0–1.5 mm.

down inside a folded leaf (Mashek, 1955). It has been found on agricultural grasses in Denmark (Nielsen & Nielsen, 1984). Larvae described by Balachowsky & Mesnil (1935), Nartshuk (1956), and Nye (1958).

Distribution. – Probably in all Nordic countries, but the species is difficult distinguish from other species of O. frit group.

81. Oscinella (Oscinella) nigerrima (Macquart, 1835)

Biology. – Host plants Lolium perenne, Deschampsia flexuosa, Poa sp. (Nielsen, 1994). Two generations per year in England (Vickerman, 1980).

Oscinis nigerrima Macquart, 1835: 603. Types probably in MHNL.

Note. – Nartshuk (1997a) studied a type specimen of Oscinis vindicata Meigen, 1830, which appeared identical to Oscinella hortensis, but made no formal synonymisation, as the taxonomy of this group is insufficiently studied.

80. Oscinella (Oscinella) maura (Fallén, 1820) Figs 6, 439, 452 Oscinis maura Fallén, 1820: 6. Lectotype in ZMLU (Andersson, 1966). Black species with densely white pubescent arista, more densely punctured and dull scutum than in other species of the genus. Wings darkened and halteres often darkened. Legs usually black, but tibiae sometimes partly yellow (Ismay, 1981c). Kariotype 2n = 6 + 0 X. Morphology of chromosomes different from other species of the O. frit complex (Michailova & Beschovski, 1985). Body length 1.7– 2.3 mm. Distribution. – Common species. Denmark: from all districts except WJ. Sweden: north to DR. Norway: AK, On, Bø, VE, Ry, HOi. Finland: north to Oa and Sa. Russia: Vib, Kr. Known from Europe, Siberia, the Caucasus and Iran. Biology. – Adults from end of May to early of September, in meadows, wood edges, fields of fodder grasses, 2–3 generations per year, but Vickerman (1980) observed only one generation in England. Larvae phytophagous, live in shoots of Dactylis glomerata (Séguy, 1934; Mashek, 1955; Nartshuk, 1956; Jepson & Nye, 1957; Nye, 1959), but Wetzel (1967) listed also Bromus inermis Leyss. Females lay eggs on the inside of a folded leaf of grass. Newly hatched larvae feed on their egg chorion and moves

Similar to O. frit, but legs entirely black. Systematic position of the species is not clear, possible it is only a dark coloured spring form of O. frit. Distribution. – Not fully clarified as difficult to separate from other closely related species, but evidently widespread in Denmark and Fennoscandia. Biology. – The species develops on Festuca ovina (Nielsen, 1994) and Arrhenatherum elatius (Vickerman, 1978a). Vickerman (1978b) established that larvae of the species did not survive on oats or barley, but a few larvae survived on wheat. The first species of the genus to appear in spring. Two generations per year in England (Vickerman, 1980).

82. Oscinella (Oscinella) nitidissima (Meigen, 1838) Figs 440, 453, 454 Chlorops nitidissimus Meigen, 1838: 388. Lectotype female in MNHN (Nartshuk, 2006). Body black. Similar to O. frit, but scutum not dusted, glabrous and shining. Frons dull. Ocellar triangle smooth shining reaching front margin of frons. Antennae black. First flagellomere round. Genae nearly as wide as first flagellomere. Scutellum dusted. Apical scutellar setae longer than scutellum. Pleura shining. npl 1 + 2. Legs mainly black, but ends of tibiae and tarsi yellowish. Body length 1.5–1.8 mm. Distribution. – Denmark: widespread. Sweden: north to LU. Norway: Bø: Akerøy, Hvaler, 6.vi.1992; HOy: Apeltunv., Bergen, 02.viii.1936, 1 male. Finland: north to Ob and Ks. Russia: Vib, Kr, Lr. Holarctic species, widely distributed in Palearctic. 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 on diverse habitats: meadows, wood edges, crop fields. Found on agricultural grasses in Denmark (Nielsen & Nielsen, 1984). Two generations per year in England (Vickerman, 1978). Larvae phytophagous, develop in shoots of Poacea, preferable in different species of Agrostis (Nartshuk, 1956; Vickerman, 1978a), but some authors listed also Anthoxanthum odoratum, Dactylis glomerata, Festuca arundinaceae, F. pratensis, F. rubra, Lolium multiflorum, L. perenne, Phleum pratense, Poa annua, P. pratensis, P. trivialis, Hol130

cus lanatus, Alopecurus geniculatus (Jepson & Nye, 1957; Nye, 1958, 1959; Rogochaya, 1960; Wetzel, 1967; Mowat, 1974, 1975). Some authors reared the species from galls of Lipara lucens (Kramer, 1917), but probably this represents misidentified Incertella zuercheri. In Canada was recorded unusual damage of oats caused by larvae of this species (Comeau & Pelletier, 1977). Vickerman (1978b) established that larvae of the species did not survive on either oats, wheat or barley. Larvae described by Nartshuk (1956, as O. agrostis), and Nye (1958).

83. Oscinella (Cyclocercula) norrbottica Nartshuk & Andersson, sp. nov. Figs 441–444 Holotype, male, Sweden: Norrbotten: Jupukka, Pajala, 28.vii.1955 (H. Andersson). Paratypes: Norrbotten: Jupukka, Pajala, 27–31.vii.1955 (H. Andersson), 2 males, 5 females; Kaunisjärvi, 08.vii.1951 (Ander, Ardö, Berden & Dahl), 1 female; Norrbotten, Kengis, 29–30.vii.1955 (H. Andersson), 1 male, 1 female. Holotype and 8 paratypes in ZMLU, 2 paratypes in ZISP. Diffferential diagnosis. – Beschovski (1978) described Cyclocercula as a subgenus of Oscinella Becker with two included species O. (C.) angustipennis and O. (C.) nartshukiana Beschovski, the latter is type species of the subgenus. The rounded shape of the male cerci is the main defining character that also refers the present species to Cyclocercula. It is distinguished from O. (C.) angustipennis by shorter wings and body and from O. (C.) nartshukiana by rounded first flagellomere without an angular upper corner. Description. – Male, female. All body black. Frons square, dull. Ocellar triangle shining, nearly reaching front margin of frons. Genae narrow, no more than half as wide as first flagellomere. Vibrissal angle rounded. First flagellomere round, arista with rather long pubescence. Scutum, postpronota and scutellum dusted. Setulae on scutum arranged in rows. Scutellum semicircular, apical setae longer than scutellum, thick at base and twice as far apart than their distance to small subapical setae. Pleura shining except on upper part. npl 1 + 2. Wings brownish. Cell br not widened. Second and third costal sectors subequal. Veins R4+5 and M divergent distally. Halteres yellow with dark stem. Legs black. Male genitalia: cerci rounded; surstyli broad; hypandrium open. Etymology. – The species is named after the type area, the Swedish province of Norrbotten.

84. Oscinella (Oscinella) pusilla (Meigen, 1830) Figs 445, 446, 455–457 Chlorops pusillus Meigen, 1830: 157; Meigen’s plate 216: 17 (Morge, 1976b). Type specimen in NHMW (Nartshuk, 1997a).

Very similar to O. frit, but differs in colour of legs: front and middle tibiae entirely yellow, hind tibiae yellow with a black band in the middle. Second aristal segment 0.3 times as long as third aristal segment. Genae as wide as first flagellomere. Rows of warts of femoral comb in male wider apart than in O. frit and distance between warts longer. Body length 1.5–2.0 mm. Distribution. – Denmark: widespread. Sweden: north to TO. Norway: Ø, AK, HEn, Bø, HO, SFi, NT, Fn. Finland: north to Li. Russia: Vib, Kr, Lr. Widely distributed in the Palearctic Region. Biology. – Adults occur in diverse habitats including crop fields. Larvae phytophagous, live in shoots and more rarely in seeds of cereals, but usually avoid oats, and in shoots of many wild grasses, preferable from the tribe Hordeae (Nartshuk, 1956). O. pusilla and O. frit are not cross-breeding (Le Berre & Chevin, 1963; Agapova, 1966). A well known serious pest of cereals in many European countries. Larvae described by Balachowsky & Mesnil (1935, as O. grossa), Nartshuk (1956), Jepson & Nye (1957), and Nye (1958). Note. – Becker (1902) who studied Meigen’s types in Vienna and Paris wrote “1 zerstörte Type in Paris. 1 Exemplar in Wien. . .” (p. 319). Nartshuk (1997a) examined the specimen in NHMW and found that it coincides with O. cariciphila Collin, has narrow genae equal of diameter of front tibiae, all femora black, the anterior and middle tibiae and all tarsi yellow, the posterior tibiae black with a yellow ends. According to Meigen’s description, all tibiae and tarsi yellow (no Oscinella species with such coloration of legs in European fauna), but in Meigen’s figure (Morge, 1976b) the base of hind tibiae are slightly darkened, as in the specimen in NHMW. Nartshuk examined the specimen in MNHN as well. It is a male without head, part of legs and a wing. Front legs absent, middle tibiae all yellow and hind leg black except tarsus. Scutum very densely dusted, more than in other species of the genus. The name Oscinella pusilla Meigen is widely used in the applied literature. In order to conserve this important name it is necessary to request the ICZN to reject the existing syntypes and designate a neotype of Chlorops pusillus Meigen in accordance 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) trochanterata Collin, 1946 Figs 447, 448 Oscinella trochanterata Collin, 1946: 140. Syntypes 16 males and 12 females in OXUM, 1 male and 1 female, in BMNH (Pont, 1995). Body black. Ocellar triangle shining, extending 2/3–3/4 length of frons. Genae with rounded vibrissal angle and head in profile rather retreating. Scutum densely dusted. The last segments of all tarsi enlarged and fourth segment short and deeply excised on upper apical margin. Trochanters of front legs conspicuously yellow. All femora broadly yellow at tip, hind tibiae yellow at both ends. Stem and knob of halteres yellow. Body length 2.5– 2.8 mm. Distribution. – Denmark: SJ, NWJ, NEJ, NWZ, NEZ, B. Sweden: SK, BL, HA, ÖL. Russia: Vib. Not found in Norway and Finland. European species. 132

Biology. – Adults in June and August, probably two generations per year. Occur in wet meadows, edges of ponds and marshy habitats, more rare than O. angularis, developing in the same plant-host. Larvae phytophagous, develop in unshelled inflorescence of Phalaris arundinaceae (Collin, 1946; Nartshuk, 1956). The tissues of the infested plant turns brown above the feeding site of the larvae. Wetzel (1967) listed as food plant also Bromus inermis. Larvae described by Balachowsky & Mesnil (1935, as O. gracilior), and Nye (1958).

86. Oscinella (Oscinella) vastator (Curtis, 1845) Oscinis vastator Curtis, 1845: 493. Body black. Similar to O. frit, but larger up to 3.0 mm and genae wider than first flagellomere, middle tibiae yellow on both ends, but hind tibiae never so distinctly yellow at base and tip as in O. pusilla. Nye (1958) supposed that O. vastator and O. festucae Mesnil might be conspecific on base of structures of larvae. Body length 2.3–3.0 mm.

Distribution. – Probably everywhere, but individual specimens very difficult to distinguish from other closely related species of the genus. Found in meadows and agricultural fields. It is possible that the species occurs in all Nordic countries. According to many authors the species damages different cereals. Vickerman (1978a) established, that O. vastator is the most polyphagous species and its preferred hosts are Lolium multiflorum, L. perenne and Festuca pratensis, but females lay no eggs on cereals in the laboratory. Nielsen (1994) found this species in Denmark on Lolium perenne, L. multiflorum, Poa spp., mainly P. trivialis, but not on wheat. Other authors listed following grasses as host plants: Agrostis tenuis, Bromus inermis, Festuca pratensis, F. rubra, Alopecurus pratensis, Phleum pratense, Arrhenatherum elatius, Lolium perenne, L. multiflorum (Jepson & Heard, 1959; Nye, 1959; Wetzel, 1967). Larvae described by Jepson & Nye (1957), Nye (1958), and as O. festucae by Nartshuk (1956).

87. Oscinella (Oscinella) ventricosi Nartshuk, 1956 Fig. 458 Oscinella ventricosi Nartshuk, 1956: 873. Holotype in ZISP. Body black. Similar to O. frit. Ocellar triangle shining and reaching front margin of frons. Genae as wide as first flagellomere. Scutum dusted, but postpronota shining without dusting. Body length 1.5– 2.3 mm. Distribution. – Sweden: north to TO. Finland: Ta. Probably more widespread in Fennoscandia, but confused with other species of the O. frit complex. Eurasian species, eastwards to Yakutia, East Siberia; occurs preferably in the north. Biology. – Adults in July to August in rich meadows. Larvae phytophagous, develop in shoots of Alopecurus spp. (Nartshuk, 1956). Larva described by Nartshuk (1956).

Genus Oscinimorpha Lioy, 1864 Oscinimorpha Lioy, 1864: 1126. Type species: Oscinis obliqua Macquart, 1835 by monotypy. Black grey-dusted species with produced vibrissal angle and prolonged proboscis. Ocellar triangle

short dusted. Face with facial carina. First flagellomere rounded. Pleura mainly shining. npl 1 + 2. Tibial organ narrow oval. Femoral organ forming one row of warts. Male genitalia: cerci discrete, and surstyli simple, hypandrium open. Distribution. – Palearctic.

Key to species of Oscinimorpha Lioy 1

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2

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All setae and setulae whitish. Pleura dull. Vibrissal angle strongly produced and proboscis rather long and geniculate (Figs 460, 461) . . . . . . . . . . . . . . . . . . . . . . . O. albisetosa Duda All setae and setulae black. Pleura partly shining. Vibrissal angle less produced and proboscis not so long and not geniculate . . . . . . 2 Genae nearly a half as wide as first flagellomere, vibrissal angle more produced (Fig. 469). Body entirely black, including head and legs . . . . . . . . . . . . . . . . . . . O. sordidissima (Strobl) Genae distinctly narrower, not wider than front tibia, vibrissal angle less produced (Fig. 463). Head and legs partly yellow. Male genitalia Figs 466, 467 . . . . O. minutissima (Strobl)

88. Oscinimorpha albisetosa (Duda, 1932) Figs 459–461 Conioscinella albisetosa Duda, 1932: 47. Syntypes in ZMHU (Wendt, 1975). Small grey-dusted species. Head yellow with black occiput. Ocellar triangle small black grey-dusted and extending halfway to front margin of frons. Antennae yellow, first flagellomere small rounded. Genae as wide as first flagellomere, with strongly produced vibrissal angle. Proboscis thin long, palpi yellow, long narrow. Scutum and scutellum light 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 yellowish at base. Legs yellow or femora and tibiae partly darkened. Wings whitish with yellow veins. Halteres yellow. Body length 1.5–1.7 mm. Distribution. – Scarce. Denmark: SJ: Rømø; NWJ: Struer; F: Langeland. Sweden: SK, ÖL. Not found in 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 Europe, but the species goes much further north in continental East Siberia, near Yakutsk. Biology. – Adults in June to August in dry meadows and sandy habitats. Larvae probably phytophagous, as it has been reared from Elytrigia and Pucclinella (Bährmann & Weipert, 1989).

89. Oscinimorpha minutissima (Strobl, 1900) Figs 56, 462–467

and setulae black. Legs nearly entirely black. Body length 1.5–2.0 mm. Distribution. – Sweden: SK, ÖL, GO, UP, DR. Finland: N. Russia: Vib. Not so far found in Denmark or Norway. Eurasian species. Biology. – Adults in June to August in meadows. In Great Britain the species has a restricted distribution (Ismay, 1994b) and the conservation status of notable (Falk, 1991). It has been found in Scotland on flat area behind the narrow fore dunes with patches of short turf (Cole, 1996).

Siphonella minutissima Strobl, 1900: 631. Syntypes 2 males in NMBA (Chvála, 2008). Black grey-dusted species. Frons dull, narrow yellowish in front. Ocellar triangle black grey-dusted, extending 2/3 length of frons. Antennae yellow with first flagellomere darkened. Genae narrower than first flagellomere, distinctly produced. Proboscis little elongated. Palpi yellow. Scutum and scutellum dusted, pleura mainly shining. Scutellum semicircular, apical setae as long as scutellum or longer. All setae and setulae black. Legs with femora and hind tibiae mainly black, remaining parts yellow. Body length 1.0–1.5 mm. Distribution. – Denmark: SJ, NWJ, NEJ, F, SZ. Sweden: north to TO. Norway: AK, HEn, VAy, Ry. Finland: north to Om. Russia: Vib; Lr: Khibiny Mts, 16.viii.1928 (Cheburova). Euro-Mediterranean species. Biology. – Adults from June to middle of August on diverse meadows, wood edges. As host plants Salvia sclarea, Secale, Oryza and Silene dioica were listed (Nartshuk, 1972a; Tschirnhaus, 1981).

90. Oscinimorpha sordidissima (Strobl, 1893)

Genus Oscinisoma Lioy, 1864 Oscinisoma Lioy, 1864: 1125. Type species: Chlorops vitripennis Meigen, 1830 (= Chlorops cognatus Meigen, 1830) by designation of Coquillett, 1910: 582. Black species with flattened granulate scutum. Head much wider than long and deep. Ocellar triangle shining or dusted. Vibrissal angle right, not produced. poc upright, cruciate. First flagellomere rounded, wider than long. Scutum markedly rugose, evenly covered with dense setulae arising from small prominences. Pleura mainly shining. npl 1 + 2. Apical scutellar setae arising from small tubercles in most species. Tibial organ distinct oval. Femoral organ composed of group of warts. Male genitalia with large cerci and slender surstyli. Hypandrium closed, postgonites with median directed projection. Distribution. – Holarctic and Oriental.

Key to species of Oscinisoma Lioy 1

Figs 468–470 Siphonella sordidissima Strobl, 1893: 252. Syntype female in NMBA (Chvála, 2008). Body black, dark grey-dusted. Frons black, ocellar triangle black, dusted, extending half length of frons. Antennae and palpi black. Genae narrower than first flagellomere, with produced vibrissal angle. Scutum and scutellum finely punctuate and dusted. Pleura shining below. npl 1 + 2. All setae

– 2

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Ocellar triangle dusted and subshining. Male genitalia Figs 480, 481 . . . . . . . . . . . . . . . . . . . . . O. germanica (Duda) Ocellar triangle not dusted and shining . . . . 2 Postpronota shining. Wings wider and shorter. Male genitalia Figs 475, 476 . . . . . . . . . . . . . . . . . . . . O. cognatum (Meigen) Postpronota dusted. Wings narrower and longer. Male genitalia Figs 482, 483 . . . . . . . . . . . . . . . . . . . . . . . . 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 (Meigen, 1830) Figs 57, 471–476 Chlorops cognatus Meigen, 1830: 154; Meigen’s plate 216: 16 (Morge, 1976b). Lectotype in NHMW (Nartshuk, 1997a). Body black, shining. Body and wings are shorter than in O. gilvipes. Frons in front and face yellow. Ocellar triangle black, shining, extending 3/4 length of frons. First flagellomere varies in the colour, from entirely yellow to darkened above. Genae as wide as first flagellomere, yellow with whitish dusting but black below. Palpi yellow. Scu136

tum and scutellum flattened, coarsely punctuate, thinly dusted. Postpronota shining. Apical scutellar setae on very small tubercles rather wide apart. Legs and halteres yellow. Body length 1.5–2.0 mm. Distribution. – Denmark: SJ, F, SZ, NEZ. Sweden: north to VR and UP, and further in NB. Norway: VAy: Straumen, Farsund. Finland: Al, Ab, N, Ta, Sa, Om. Russia: Vib, Kr. The commonest species of the genus. TransPalearctic boreal species. Biology. – Adults from May to October, common on bogs, fens, marshes and other wetlands. Flies live near the ground in deep litter, hibernate as adults.

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 aquatica, G. maxima, Phalaris arundinaceae, Festuca arundinaceae, Carex sp., Scirpus, Sparganium, Typha (Rogochaya, 1960; Nartshuk, 1972a; Sharipov, 1981; Tschirnhaus, 1992).

microsculpture and distinctly dusted. Postpronota dusted. Colour of legs is strongly variable from entirely yellow to darkened femora and tibiae. The male genitalia are larger than in the other species of the genus.

92. Oscinisoma germanicum (Duda, 1932)

Distribution. – Rare. Found only in Sweden: SK: Skäralid, 20.vi.1941 (T. Nyholm), 1 male; SM: Rödsle, Oskarshamn, 22.vii.1983 (R. Danielsson), 1 male. European species.

Figs 477–481 Discogastrella germanica Duda, 1932: 42. Holotype in ZMHU (Wendt, 1975).

Biology. – Collin (1939) reported that Mr. Basden found the species alive in one of his artificial decoy 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. First flagellomere entirely yellow. Scutum not so flat as in other species of the genus, with weaker

93. Oscinisoma gilvipes (Loew, 1858) Figs 482, 483 137

Oscinis gilvipes Loew, 1858: 71. Syntype in ZMHU (Wendt, 1975). Frons dusted, black behind and yellow in anterior third. Ocellar triangle black, shining, extending 2/3–3/4 down frons. orb about 10, small, but 2– 3 may be longer. Eyes with pale short hairs. First flagellomere wider than long, in male yellow with apically darkened part, in female darkened on the upper half. Arista black. Genae in front as wide as first flagellomere, wider behind, yellow in front and black behind. Palpi yellow. Scutum longer than wide, shining, rugose, flattened in hind 3/4. Postpronota and notopleura dusted. npl 1 + 1. Most part of pleura shining. Scutellum black, wider than long, rugose. Subapical setae on small tubercles in some specimens. Wings longer than in O. cognata, longer than abdomen in some specimens. Halteres yellow. Body length 1.5–1.7 mm. Distribution. – Rare. Found only in Sweden: SK, SM, UP. European species. Biology. – Adults from May to the end of September in wet habitats, probably hibernating as adults. It has been reared from Glyceria maxima (Tschirnhaus, 1992).

Genus Polyodaspis Duda, 1933 Polyodaspis Duda, 1933: 61. Replacement name for Macrothorax Lioy, 1864 (preocc. Jaennicke, 1867). Type species: Siphonella rufijicornis Macquart, 1835, automatic. Black species with well developed facial carina, distinctly produced vibrissal angle, shining frons and ocellar triangle. poc strong upright parallel. Genae divided by oblique carina into a lower posterior darker part with coarse sculpture and numerous setulae and a smooth upper anterior part. Tibial organ small oval. Femoral organ absent. Male genitalia: cerci simple, surstyli broad, hypandrium open. Distribution. – Palearctic, Oriental and Afrotropical Regions.

Key to species of Polyodaspis Duda 1

138

Wing cell br widened in the middle (Fig. 487). Scutellum with numerous marginal setae and

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apical setae approximated (Fig. 487b). npl 1 + 2(3). Vibrissal angle not strongly produced and proboscis shorter. Male genitalia Figs 488, 489 . . . . . . . . . . . . . . . P. rufijicornis (Macquart) Wing cell br not widened in the middle. Scutellum with apical setae wide apart and no more than two lateral (subapical) on each side. Vibrissal angle strongly produced and proboscis longer (Fig. 490). npl 1 + 1. Male genitalia Figs 491, 492 . . . . . . . . . . . . . . . . . . . . P. sulcicollis (Meigen)

94. Polyodaspis rufijicornis (Macquart, 1835) Figs 14, 93, 484–489, 964, 965 Siphonella rufijicornis Macquart, 1835: 585. Types probably in MHNL. Body black. Frons brown covered with brownish setulae on distinct punctures. Ocellar triangle indistinct, shining, minutely shagreened, extending beyond 2/3 length of frons. poc, vte and vti subequal 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 brown, distinctly rugose, anterior part covered with more or less silvery dust. Vibrissal angle strongly produced. Facial carina well developed, triangular above, very thin below, separating the deep antennal foveae. Proboscis elongated. Palpi yellow with black setulae and one strong apical seta. Scutum subshining, densely shagreened. Scutellum rounded apically, weakly flattened and shagreened with a pair of approximated apical and 6–8 pair of short stout subapical setae on small tubercles. Pleura shining, wrinkled and covered with small setulae. npl 1 + 2(3). Halteres with large black knob. Legs mainly black. Abdomen black with short black setulae. In wing cell br widened. Male genitalia: epandrium small with squarish strongly sclerotized cerci and broad surstyli pointed at tip. Body length 2.5–3.0 mm. Distribution. – Uncommon, except in Finland. Denmark: ?SJ: No locality [Crepis Litocha] (W. Wüstnei). Norway: AK: Rakeie, Oslo; Kirkeby, Maridalen (bread from flowerhead of Cirsium helenioides). Finland: N, Ta, Sa, Oa, Sb. Russia: Kr; Lr: Kandalaksha; Kola; Yläluostari. Not found in Sweden.

Widely distributed in the Palearctic Region, more abundant in the Asian part of the Palearctic Region, recorded from the Oriental Region as well. Biology. – Adults in dry habitats. Larvae feed on different substances, usually enriched proteins, and exhibit very wide ecological plasticity in relation to the choice of food substrates and environmental conditions. They may be saprophagous, phytophagous, necrophagous or parasitic (Kiauka & Nartshuk, 1972; Nartshuk, 1972b). Flies were reared from inflorescence of Asteraceae: Knautia arvensis together with larvae of Larinus sp., Cirsium spp. together with caterpillars of Microlepidoptera and larvae of Tephritidae, walnut fruits, both intact and infested by other insects, large heads of artichoke (Cynara scolymus) together with larvae of Apion carduorum, stems of Cannabis sativa and Onopordon acanthium together with Lixus pollinosus, stems of maize usually damaged by other insects, from dead and living caterpillars of Pyrausta nubilalis living in stem of maize, from tachinid fly living as parasite of Pyrausta caterpillar, from Laspeyresia caterpillars on apple and plum, from rolling leaves with larvae of the beetle Byctiscus rugosus on aspen, from Apoderus coryli on river birch, found as ectoparasite on larvae of Hoplocampa (Hymenoptera, Tenthredinidae) on pear and plum trees, from galls of Paranthrene tabaniformis (Lepidoptera, Sesiidae) (Perris, 1830, 1839, 1873, 1876; Frauenfeld, 1863; Schiner, 1864; Perris & Laboulbéne, 1871; Vukasovič, 1926; Goidanich, 1928; Falcoz, 1930; Sachtleben, 1930, 1939; Martelli, 1933a, 1933b; Rogochaya, 1960; Nartshuk & Kiauka, 1963; Beschovski & Georgiev, 1993). The species is often introduced with damaged walnuts fruits. Larvae described by Kiauka (1974).

Proboscis long with narrow labella. Palpi from yellowish to black, as long as front tibia. Scutum and scutellum shining, slightly flattened. Scutellum with apical setae long and not approximated, subapical setae thin and short. Pleura with distinct microsculpture. npl 1 + 1. Male abdomen with membranous vesicula between 5 tergite and synsternite 7 + 8, seen only on macerated abdomen. Legs mainly black. Body length 1.5–1.8 mm. Distribution. – Uncommon. Denmark: B: Bagå, 16.vi and 19.vii.1964, 2 males. Sweden: SK, SM, GO, SÖ, UP. Norway: AK: Hovedøya and Lindøya, Oslo Fjord. Finland: Al, Ab, N. Not found in Russian parts of Fennoscandia. Widespread in W Palearctic, eastwards to Yakutia, East Siberia and Mongolia. Biology. – Adults in May to July in dry habitats. Larvae as in P. rufijicornis exhibit wide ecological plasticity in relation to the choice of food substrates and environmental conditions. The adults were reared from stem of broomrape (Orobanche speciosa D.C.) damaged by other insects (Martelli, 1935). Meijere (1938) suggested that the larvae are predators, but Martelli (1935) regarded them as saprophytophagous. Some larval structures illustrated by Nartshuk (1987).

Genus Pseudogaurax Malloch, 1915 Pseudogaurax Malloch, 1915: 159. Type species: Gaurax anchora Loew, 1866 by original designation.

95. Polyodaspis sulcicollis (Meigen, 1838)

Yellow species with black markings, long triangular scutellum with approximate apical scutellars. Ocellar triangle shining. First flagellomere obliquely prolonged with thickened pubescent arista. Tibial organ distinct, narrow oval. Femoral organ absent. Male genitalia: cerci fused basally, surstyli simple.

Figs 67, 490–492

Distribution. – Worldwide.

Chlorops sulcicollis Meigen, 1838: 387. Lectotype female in MNHN (Nartshuk, 2006). Body black, shining. Frons shining brownish and punctuate. Ocellar triangle extending half length of frons with a row setulae on lateral margins. Antennae from yellow to black, first flagellomere a little angulated. Arista black with short pubescence. Genae shining dark brown, black on below margin, vibrissal angle acute and distinctly produced.

96. Pseudogaurax venustus (Czerny, 1906) Fig. 493 Gaurax venustus Czerny, 1906: 299. Lectotype in NHMW (Nartshuk, 1997b). Body yellow with black, shining longitudinal stripes on scutum. Frons narrow, longer than wide, yel139

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 triangle yellow with black ocellar tubercle, smooth shining, extending half length of frons. Occiput with small black spot. Eyes large and dense and long pubescent. Antennae yellow, first flagellomere wider than long. Arista short, basally thick and yellow, apically thin and darkened. Genae narrow, less than half width of first flagellomere with rounded vibrissal angle. Scutum with 3 black, shining stripes covered with yellow setulae. Scutellum elongate triangular, yellow, apical setae longer than scutellum and approximated. Pleura smooth shining with black marks. npl 1 + 2. Legs and halteres yellow. Body length 2.0–2.5 mm. Distribution. – Rare. Only Russia: Kr: Kivach Nature Reserve, Malaise trap, 1 female (Nartshuk, 1999). European species, known from Austria and Hungary. Biology. – Life habits of this species unknown, other species of the genus are carnivorous, larvae develop in egg sack of Aranea.

2

– 3 – 4

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5

Genus Rhopalopterum Duda, 1929

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Rhopalopterum Duda, 1929: 167. Type species: Oscinella limitata Becker, 1912 by monotypy. Black, shining species with elongated thorax and relatively long wings. Frons silky shining, ocellar triangle large shining or in thin dust. Genae usually linear, vibrissal angle straight. poc upright, cruciate. First flagellomere rounded, wider than long. Scutum rather flat, sometimes distinctly flattened behind, shining or thinly dusted and evenly covered with setulae. Thorax sometimes with metallic green lustre. Pleura smooth, mainly shining. npl 1 + 1. Tibial organ oval. Femoral organ composed of two indistinct rows of warts. Male genitalia: cerci and surstyli simple, hypandrium open, postgonites usually with sclerotized projection. Distribution. – Holarctic and Neotropical Regions.

Key to species of Rhopalopterum Duda 1

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Scutum and ocellar triangle shining, without dusting, scutellum covered with thin dust ......................................2 Scutum and ocellar triangle subshining, covered with rather thin dust, scutellum ticker

6

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dusted than scutum. Male genitalia Figs 515, 516 . . . . . . . . . . . . . . . . . R. tomentosum sp. nov. Genae wider than front tibia. Scutum with dusting at sides and behind, covered with pale setulae. Legs entirely darkened, only basitarsi of all legs yellowish . . . . . . . . . . . . . . . . . R. atricillum (Zetterstedt) Genae narrow, near linear (Fig. 498). Legs yellow or darkened . . . . . . . . . . . . . . . . . . . . . . . . 3 Legs entirely yellow . . . . . . . . . . . . . . . . . . . . . 4 Legs more or less dark, at most all femora black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Body with distinct greenish lustre, especially on abdomen. Scutum not flattened before scutellum, covered with brown setulae. Male genitalia Figs 494–496 . . . . . . . . . . . . . . . . . R. anthracinum (Meigen) Body black, without metallic luster. Scutum flattened before scutellum. Male genitalia Figs 500–502 . . . . . . . . . . R. atricorne (Zetterstedt) Wings brownish (Fig. 504). Legs including all femora and most tibiae black . . . . . . . . . . . . . 6 Wings transparent not brownish. Front tibiae yellow, hind tibiae and sometimes middle tibiae with black band. Postpronota and notopleural depression dusted. Male genitalia Figs 508–510 . . . . . . . . . . . . . . R. fasciola (Meigen) Ocellar triangle with small black metallic shining half-moon-shaped spot before front ocellus. Scutum dusted at sides. No hairy spot on inner surface of epandrium. Hypandrium open. Male genitalia Figs 505–507 . . . . R. brunneipenne Beschovski & Lansbury Ocellar triangle shining without metallic spot. Postpronota and greater part of notopleural depression shining without dust. Hairy spot present on inner surface of epandrium. Hypandrium closed. Male genitalia Figs 511–513 . . . . . . . . . . . . . . . . . . . . . . . . R. femorale Collin

97. Rhopalopterum anthracinum (Meigen, 1830) Figs 494–496 Chlorops anthracinus Meigen, 1830: 156; Meigen’s plate 216: 14 (Morge, 1976b). Lectotype in 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, especially on abdomen. Frons black, silky shining, ocellar triangle shining, extending 3/4 length of frons or longer, genae linear. Antennae and palpi black. Scutum shining, evenly covered with brown setulae. Notopleural depression dusted. Scutellum rounded, dusted. Legs entirely and halteres yellow. Body length 1.8–2.0 mm.

98. Rhopalopterum atricillum (Zetterstedt, 1838) Oscinis atricilla Zetterstedt, 1838: 781. Lectotype in ZMLU (Andersson, 1966).

Distribution. – Denmark: F, NWZ, NEZ, B. Sweden: north to HS. Finland: north to Ob. Russia: Vib, Kr. Not recorded yet in Norway. Eurasian boreal species.

Body black, elongated. Frons dull, ocellar triangle shining, reaching 3/4 length of frons. Antennae and palpi black. Genae wider than front tibia. Scutum shining, evenly covered with pale setulae, but extensively dusted behind and especially at sides. Legs entirely darkened, only basitarsi of all legs yellowish. Halteres yellow. Body length 1.5–1.8 mm.

Biology. – Adults in June to August in wetlands with Carex spp., which are known as host plants (Nartshuk, 1972a). Tyrphophilous species.

Distribution. – Sweden: north to TO. Norway: HOi, Nsy, TRi. Finland: north to Li. Russia: Kr, Lr. Not so far found in Denmark. Eurasian boreal species.

142

Biology. – Adults from June to early September in wetlands with Carex spp., which probably represent the host plants. Tyrphobiont species.

101. Rhopalopterum fasciola (Meigen, 1830)

99. Rhopalopterum atricorne (Zetterstedt, 1838)

Chlorops fasciolus Meigen, 1830: 154; Meigen’s plate 237: 9 (Morge, 1976b). Lectotype in NHMW (Nartshuk, 1997a).

Figs 497–502 Oscinis atricornis Zetterstedt, 1838: 780. Lectotype in ZMLU (Andersson, 1966). Lioscinella platythorax Nartshuk, 1958: 99. (Nartshuk & Andersson, 2003). Body elongated, black, shining. Frons dull black, velvety; ocellar triangle shining, extending 3/4 length of frons, genae linear, but a little wider than in R. anthracinum. Antennae and palpi black. Scutum shining, evenly covered with brown setulae, distinctly flattened in hind part. Postpronota and notopleural depression dusted. Scutellum rounded dusted. Legs entirely yellow. Halteres yellow. Body length 1.8–2.0 mm. Distribution. – Sweden: north to TO. Finland: Ab, N, Ta, Sb. Russia: Vib, Kr. Not so far found in Norway and Denmark. Eurasian boreal species. Biology. – Adults in June to August in wetlands with Carex spp. Probably species of Carex are host plants (Nartshuk, 1972a). The species is more abundant than R. anthracinum.

100. Rhopalopterum brunneipenne Beschovski & Lansbury, 1987 Figs 503–507 Rhopalopterum brunneipenne Beschovski & Lansbury, 1987: 95. Holotype male in OXUM (Pont, 1995). Frons a little longer than wide. Ocellar triangle extending 2/3 length of frons, shining with small black metallic shining half-moon-shaped spot. Genae linear. First flagellomere rounded, palpi black. Scutum before scutellum, notopleura, upper part of anepisternum and scutellum dusted. Postpronota shining. Legs mainly black. Wings distinctly brownish, especially in the anterior half. Body length 2.0 mm. Distribution. – Finland: Ab, Sb. Russia: Kr. Described from England. Elsewhere only known from NW Russia and Lithuania.

Figs 508–510

Body elongated, black, shining. Frons dull, ocellar triangle shining, extending 3/4 length of frons, genae almost linear. Antennae black. Postpronota and notopleural depression dusted. Legs with femora black, hind tibiae and sometimes middle with black band. Body length 1.8–2.0 mm. Distribution. – Denmark: EJ, NEZ, B. Sweden: north to HS, also NB. Finland: Ab, Ta, Kb, Om. Russia: Vib, Kr. Not so far found in Norway. Biology. – Adults in June to August in wetlands with Carex. Probably species of Carex are host plants.

102. Rhopalopterum femorale Collin, 1946 Figs 511–513 Lioscinella femoralis Collin, 1946: 145. Lectotype male and 6 males, 8 females paralectotypes in OXUM (Beschovski & Lansbury, 1987; Pont, 1995). Body elongated, black, shining. Frons dull, ocellar triangle shining, extending 3/4 length of frons, genae almost linear. Antennae black. Postpronota and greater part of notopleural depression shining, without dust. Wings faintly brownish. Legs with femora black, all tibiae with black band. Body length 1.8–2.0 mm. Distribution. – Sweden: north to TO. Finland: north to Lk. Russia: Vib, Kr. Not so far found in Norway and Denmark. Eurasian boreal species, known eastwards to Mongolia. Biology. – Adults in June to August in wetlands with Carex spp. Probably species of Carex are host plants (Nartshuk, 1972a). Tyrphophilous species.

103. Rhopalopterum tomentosum Nartshuk & Andersson, sp. nov. Figs 514–516 Holotype male, Sweden: SK: Åsljunga, 05.vi.1960 (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 ZMLU, 2 paratypes in ZISP. Diffferential diagnosis. – Differs from other species of the genus by thin dust on ocellar triangle and thorax and structure of epandrium. Description. – Male, female. Body black, subshining with rather thin dust. Head wider than thorax. Frons as long as wide, narrowed anteriorly, dull. Front margin of frons straight. Ocellar triangle extending 2/3 length of frons, subshining, covered with thin dust as in Incertella zuercheri. Head setae rather long, 5 thin orb, vte little longer than vti and poc. Genae a little wider than anterior tibiae and widened posteriad. Vibrissal angle rounded with short vi. Antennae black, first flagellomere small and rounded, arista with rather long pubescence. Palpi short black. Scutum longer than wide, covered with thin dust. Scutellum short, semicircular, thicker dusted than scutum. Apical scutellar setae longer than scutellum. Two pairs of subapical setae. Pleura shining. npl 1 + 1, but some female have second thin posterior seta. Wings brownish. Costal sectors as 30:25:18:10. Halteres yellow. Male genitalia: cerci more or less transversal. Hypandrium open. Body length 1 mm. Note. – The new species is provisionally placed in the genus Rhopalopterum, even though it shares some characters in dusting of ocellar triangle and scutum with black Incertella species. It differs from species of Incertella by the longer and less convex scutum. Also the shape of the hypandrium and epandrium is more like in other species assigned to the genus Rhopalopterum. The new species belongs to a complex of small, entirely black species which are rather difficult to distinguish. There exist some other, probably new species within this group in our material. We refrain however from describing more species at this moment, as more material is needed to assess character variation and and habitat preferences.

Genus Siphonella Macquart, 1835 Siphonella Macquart, 1835: 584. Type species: Madiza oscinina Fallén, 1820 by original designation. Black, shining species with strongly produced vibrissal angle, distinct high facial carina and geniculate proboscis with long labella. Ocellar triangle

shining. poc upright convergent. Scutellum rather long triangular, apical setae on small tubercles. npl 1 + 2. Tibial organ distinct oval. Femoral organ consists of a dense group of long setulae. Male genitalia: cerci close together, surstyli simple. Hypandrium closed. Distribution. – Probably worldwide, but not known in the Neotropical and Australasian Regions.

104. Siphonella oscinina (Fallén, 1820) Figs 12, 13, 27, 28, 59, 517–521 Madiza oscinina Fallén, 1820: 6. Lectotype in ZMLU (Andersson, 1963). Body black, shining. Ocellar triangle narrow, extending half length of frons. orb numerous. Face with narrowly triangular carina above, which after narrowing below and widening again reaches facial margin. Vibrissal angle strongly produced, proboscis long. Palpi long cylindrical yellowish. First flagellomere rounded, wider than long, rather thick below, yellow, darkened above. Scutum coarsely punctate, evenly covered with numerous short setulae arising from stellate granulae. Scutellum with the same microsculpture. Pleura smooth shining, only anepisternum wrinkled and dusted above. Halteres brown. Body length 2.0–3.0 mm. Distribution. – Denmark: SJ, EJ, WJ, NEJ, LFM, NEZ. Sweden: north to TO. Norway: Ø, AK, HEs, Bø, VE, VAy, Ry, SFi, STi, Fn. Finland: north to Li. Russia: Vib, Kr, Lr. Holarctic species. Biology. – Adults autumnal, from the end of July to September. Data on bionomics of the species controversial. Coquillett (1898) reported that from egg cocoon of Aranea (species not named) were reared 4 specimens of S. oscinina. Nartshuk (1962a) proposed that larvae feed on coccids Greenisca glyceriae (Green), living on Agrostis vulgaris. Note by Duda (1932–1933) that Schwangart (1906) reported parasitism of larvae in egg cocoon of Aranea virgata Hahn is not correct. Schwangart not exactly named the species, but considered it as species related to S. oscinina. Probably Schwangart’s note refers to Lasiambia coxalis or L. palposa. Adults were found on Common stinkhorn, Phallus impudicis in Finland (Luther, 1946; Smith, 1956). 145

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

Genus Siphunculina Rondani, 1856 Siphunculina Rondani, 1856: 128. Type species: Siphunculina brevinervis Rondani, 1856 (= Siphonella aenea Macquart, 1835) by original designation. Black, shining species or grey pollinose with short vein R2+3 . Ocellar triangle shining or dusted. Vibrissal angle produced. poc upright convergent. Face with carina completely dividing antennal foveae. Tibial organ small oval. Femoral organ absent. The genus has been associated with dung and birds nest. Distribution. – Worldwide.

Key to species of Siphunculina Rondani 1

146

Body shiny black, all setae and setulae of body black. Male genitalia Figs 525, 526 . . . . . . . . . . . . . . . . . . . . . S. aenea (Macquart)

–

Body grey pollinose, all setae and setulae of body whitish. Male genitalia Figs 527–529 . . . . . . . . . . . . . . . . . . . . . . S. nidicola Nartshuk

105. Siphunculina aenea (Macquart, 1835) Figs 26, 39, 522–526 Siphonella aenea Macquart, 1835: 585. Types probably in MHNL. Diastata albinervis Roser, 1840. Lectotype in SMNS (Nartshuk, 1994). Body black. Frons black and evenly dusted, head setae black and short. Ocellar triangle black, shining, reaching margin of frons, with a shallow median furrow. Antennae yellow, first flagellomere more or less darkened, wider than long, thick basally. Facial carina broadly triangular between antennae, then narrowing and again broadening before facial margin. Genae slightly narrower than first flagellomere. Vibrissal angle weakly produced. Scutum shining with fine setiferous punctuations, de-

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. Scutellum rather short, apical and two pairs of subapical setae very short and stout. Femora black, tibiae yellow with black band. Halteres black. Body length 1.5–2.0 mm. Distribution. – Sweden: HA, NÄ, DR. Norway: AK: Konglungen, Asker, 4.vi.1997 (G.E.E. Söli). Finland: north to Om. Russia: Vib. Not found in Denmark. Widely distributed species in the Palearctic Region eastwards to Japan. Biology. – Adults from May to August, mainly in woodland, often on flowers. Also found in a swallow’s nest (Krivokhatsky & Nartshuk, 2001). It has been reared from faeces of wild brown bear and cow dung in Japan (Kanmiya, 1982, 1989).

106. Siphunculina nidicola Nartshuk, 1971 Figs 527–529

Body black grey pollinose. Frons black, ocellar triangle shining, dusted on ocellar tubercle only, extending 2/3–3/4 length of frons, with a shallow median furrow. Head setae and setulae whitish. Antennae and palpi yellow. Arista short pubescent, nearly bare. Genae slightly narrower than first flagellomere. Vibrissal angle weakly produced. Scutum dull and with dispersed short whitish setulae, setae of thorax whitish. Pleura black, shining except on upper part of anepisternum. Notopleural setae 1 + 1. Scutellum trapezoid, apical setae little shorter than scutellum. subapical setae very close to apicals. Legs yellow, femora and tibiae of hind legs slightly darkened. Femora black, tibiae yellow with black band. Halteres brown. Body length 2.0 mm. Distribution. – Not yet found in Denmark and Fennoscandia, but expected in Finland. Eurasian species described from Russia: Leningrad Oblast and Mongolia. Biology. – In Mongolia bread from a kite’s (Milvus) nest.

Siphunculina nidicola Nartshuk, 1971: 290. 147

Genus Speccafrons Sabrosky, 1980 Speccafrons Sabrosky, 1980: 424. Type species: Oscinella mallochi Sabrosky, 1938 by original designation. Body black, shining or dusted. Eyes densely pubescent. Frons with numerous small bare spots scattered 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 brownish 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. Body black grey-dusted. Frons dull, brownish black behind and reddish yellow in front. Setulae of frons arise from shining punctures. Ocellar triangle small, extending half length of frons, black grey-dusted. Eyes densely haired. Antennae yellow brownish, first flagellomere wider than long. Genae as wide as first flagellomere or narrower. Palpi rather thick and long, yellowish. Scutum with setulae not arranged in rows. Scutellum semicircular, with numerous setae on hind margin. Pleura mainly shining. npl 1 + 2. Legs mainly black, but front and middle tibiae yellow at ends. Body length 1.8–2.0 mm. Distribution. – Denmark: SZ, NEZ, B. Sweden: north to LU. Finland: Ab, N, Sb. Russia: Vib, Kr. Not so far found in Norway. Trans-Palearctic species, known from the British Isles to Russian Far East. Biology. – Adults in June to September, in meadows and wetlands with Phragmites australis. Larvae carnivorous, developing in egg sacks of the spiders Larinioides sericatus and Singa nitidula (Schwangart, 1906; Pereleshina, 1928, as “Conioscinella frontella”; Séguy, 1934 and Vachon, 1952, as “Oscinisoma frontellum”; Nartshuk, 1972b). Schwangart (1906) determined the flies reared from eggs sacks as a species near Siphonella oscinina, but 148

Figs 530–534. Speccafrons halophila (Duda): 530: head, lateral view. 531: antenna. 532: scutellum. 533: hypopygium, dorsal view. 534: hypandrium and phallic complex.

Collin (1946) found it identical with the present species.

Genus Trachysiphonella Enderlein, 1936 Trachysiphonella Enderlein, 1936: 187. Type species: Oscinis pumilio auct. nec Zetterstedt, 1848 (= Chlorops scutellatus Roser, 1840) by monotypy. Yellow species with 3–5 black or red stripes on scutum and black marks on pleura similar in colour to those of Chlorops species. Frons dull with dusted small ocellar triangle. Vibrissal angle produced. Proboscis prolonged. Face with distinct narrow carina. npl 1 + 2. Tibial organ distinct. Femoral organ present as oval group of warts. Male genitalia: cerci and surstyli simple, hypandrium open, postgonites with sclerotized projections. Distribution. – The Palearctic Region.

Key to species of Trachysiphonella Enderlein

109. Trachysiphonella scutellata (Roser, 1840) Figs 60, 539–546

1

–

Body larger, reaching 2.0 mm. Stripes on scutum black sometimes fused. Pleura with four black marks. Male genitalia Figs 544–546 . . . . . . . . . . . . . . . . . . . . . . T. scutellata (Roser) Body smaller, reaching 1.5 mm. Stripes on scutum black or red. Pleura only with one black or red mark. Male genitalia Fig. 538 . . . . . . . . . . . . . . . . . . . . T. rufijiceps (Macquart)

108. Trachysiphonella rufijiceps (Macquart, 1835) Figs 535–538 Siphonella rufijiceps Macquart, 1835: 586. Types probably in MHNL. Chlorops pygmaeus Meigen, 1838: 385, syn. n. Lectotype, male, in MNHN (Nartshuk, 2006). Body yellow with black to red marks. Frons yellow and ocellar triangle very small, extending no more than 1/3 length of frons, blackish and dusted. Antennae and palpi yellow. Genae produced to different degree and proboscis elongated correspondingly. Scutum yellow with black to red longitudinal stripes. Scutellum yellow. Pleura with only one black spot. Legs and halteres yellow. Body length 1.5 mm.

Chlorops scutellatus Roser, 1840: 63. Syntypes in SMNS (Andersson, 1966; Nartshuk, 1994). Body yellow with black marks. Frons yellow and ocellar triangle very small, extending no more than 1/3 length of frons, black and dusted. Antennae and palpi yellow, first flagellomere darkened above. Genae distinctly produced and proboscis elongated. Scutum yellow with black longitudinal stripes, sometimes fused in dark coloured specimens. Scutellum yellow or darkened. Pleura with 4 black spots. Legs mainly yellow. Halteres yellow. Body length 1.8–2.0 mm. Distribution. – Denmark: LFM, NEZ. Sweden: north to LU. Norway: AK: Oslofjord: Lindøya; Hovedøya; Baerum; Oksenøya; Storøykilen. Finland: north to Lk. Russia: Vib; Kr: 3 km SW Konchozero, 15.vii.2002 (Polevoi); Kr: Kivach, 04.vii.2001 and 03.ix.2002 (Polevoi). Eurasian species known from Europe eastwards to Yakutia, East Siberia and Mongolia and southwards to Israel. Biology. – Adults in June to July in meadows.

Genus Tricimba Lioy, 1864

Biology. – Adults in June to August in meadows.

Tricimba Lioy, 1864: 1125. Type species: Oscinis lineella Fallén, 1820: 8 by designation Enderlein, 1911: 207. Nartshukiella Beschovski, 1981c: 119, as subgenus of Tricimba Lioy, 1864. Type species: Chlorops cincta Meigen, 1830 by original designation.

Note. – The present species is very variable in colour, projection of genae and correlated length of proboscis, but we were unable to find distinct differences in the structure of male genitalia supporting a subdivision into several species. Therefore, we only recognized two species in our large Palearctic material based on size, number of setulae on scutellum and distance between apical setae on scutellum (Figs 537, 542): the larger Trachysiphonella scutellata (Roser) and the smaller T. rufijiceps (Macquart, 1835). Oscinis flavella Zetterstedt, 1848, and Siphonella diplotoxoides Strobl, 1893 are both considered as likely synonyms of T. rufijiceps (Macquart, 1835).

Black densely dusted species with 3–5 longitudinal punctuate lines on scutum. Ocellar triangle dusted. Vibrissal angle produced. poc upright convergent. First flagellomere rounded. npl usually 1 + 2. Scutellum sometimes thick with convex side margins, and scutellar apicals stout. Tibial organ oval. Femoral organ consists of two rows of warts or absent. Male genitalia: cerci separated or more or less fused, hypandrium closed or open, pre- and postgonites indistinctly separated. Female with ovipositor membranous. European species of the genus Tricimba are referred to 2 subgenera: Tricimba s. str. having marginal scutellar setae short, located at the same

Distribution. – Denmark: SJ, NEZ. Sweden: north to VS. Finland: Al, Ab, N. Russia: Vib. Not so far found in Norway. Eurasian species.

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 tubercles 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 no tubercles at the base, cell br not widened, male cerci separated, hypandrium closed. A third subgenus, Schumaniella Beschovski, 1981, is confined to North Africa and the Near East. Distribution. – Worldwide. 150

Key to species of Tricimba Lioy 1

–

Ocellar triangle with shining spot before anterior ocellus. Postpronota yellow. Male genitalia Figs 552, 553 . . . T. (N.) humeralis (Loew) Ocellar triangle entirely dull without shining spot. Postpronota coloured as remaining parts of thorax . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2

–

Three faintly impressed punctuate lines on scutum. Scutellum flat and apical setae not stout, scutellum often yellow apically (Fig. 548). Male genitalia Figs 549, 550 . . . . . . . . . . . . . . . . . . . . T. (N.) cincta (Meigen) Five deeply impressed punctuate lines on scutum. Scutellum and apical setae stout, scutellum coloured as remaining parts of thorax (Figs 556, 557). Cell br widened in the middle (Fig. 558). Male genitalia Figs 560, 561 . . . . . . . . . . . . . . . . . . . . T. (T.) lineella (Fallén)

110. Tricimba (Nartshukiella) cincta (Meigen, 1830) Figs 61, 547–550 Chlorops cinctus Meigen, 1830: 162; Meigen’s plate 227: 12 (Morge, 1976b). Lectotype in NHMW (Nartshuk, 1997a). Tricimba fungicola Dely-Draskovits, 1983: 349 (Nartshuk, 1997a). Body black grey-dusted. Frons mainly yellow, infuscate behind. Ocellar triangle black grey-dusted, extending halfway to front margin of frons. Antennae yellow, first flagellomere orbicular, slightly darkened apically. Head setae from yellow to black. Genae yellow, narrower than first flagellomere. Vibrissal angle weakly produced. Palpi yellow. Scutum densely grey-dusted with 3 longitudinal sulci of punctuations along acrostichal and dorsocentral lines. Scutellum somewhat conical, slightly wider than long, distinctly flattened, usually with yellow tip, but specimens from N Fennoscandia have no yellow tip. Apical scutellar setae close-set, shorter than scutellum, three pairs of subapical setae. Scutellar setae from yellow to black. Pleura mainly shining. npl 1 + 2. Legs in males mainly yellow, in females femora darkened. Body length 1.5–2.0 mm. Distribution. – Denmark: from all districts except SJ, WJ and SZ. Sweden: north to TO. Norway: AK, HEs, Bø, AAy, STi. Finland: north to Li. Russia: Vib, Kr, Lr. Holarctic species. Biology. – Adults from June to September in meadows, in forests, from canopies of Pinus silvestris in Norway (Thunes et al., 2004), and in agricultural fields in Denmark (Nielsen & Nielsen, 1984). Hackman (1963) found adults in borrows of Microtus

and Clethrionomys in Finland. Hibernate as adults. It has been recorded many times indoors in the autumn together with Thaumatomyia notata. Adults were recorded also from birds’ nests (Hicks, 1953; Krivokhatsky & Nartshuk, 2001). The larvae are exceptionally diverse in feeding habits. They have been reared from stems of cereals infested by other insects by many authors, including seedlings of wheat in Norway (Rygg, 1966), from rotting spathes of skunk cabbage (Symplocarpus foetidus), from various mushrooms (Boletaceae, Agaricaceae, Russulaceae), from rotting bark of Alnus, berries of Sambucus racemosa, dead mollusks and meat baits (Brown, 1956; Myhályi, 1965; Dely-Draskovits, 1972; Krivosheina, 1974; Schatzman, 1977; Grimaldi & Jennike, 1983; Krivosheina et al., 1986; Tschirnhaus, 1992; Yakovlev, 1994). Anterior and posterior spiracles of larvae are figured by Krivosheina (2005). Note. – A lectotype designation for Chlorops cinctus Meigen by Dely-Draskovits (1983) is invalid, because she designated a specimen that cannot belong to the syntypes series, as labeled “Madrid 7. August”. Meigen stated “Im September auf der Blüthe der Möhren selten”. According to Pont (1986) this indicates specimens collected in Stolberg near Achen, Germany.

111. Tricimba (Nartshukiella) humeralis (Loew, 1858) Figs 551–553 Oscinis humeralis Loew, 1858: 59. Syntypes in ZMHU (Wendt, 1975). Body black grey-dusted. Frons mainly yellow with yellow setae and setulae. Ocellar triangle black grey-dusted, extending half length of frons with shining spot in apical part between front ocellus and tip of triangle. Antennae yellow, first flagellomere orbicular, slightly darkened apically. Arista yellow at base. Genae yellow, as wide as first flagellomere. Vibrissal angle weakly produced. Palpi yellow. Scutum black with postpronota yellow, densely grey-dusted with 3 longitudinal sulci of punctuations along acrostichal and dorsocentral lines. Scutellum slightly wider than long with yellow tip. Apical scutellar setae as long as scutellum. Pleura shining below. npl 1 + 2. Abdomen yellow at base. 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– 2.0 mm. Distribution. – Rare. Sweden: SK: Alnarp, 26.vi.1960 (H. Andersson), 1 male. Norway: 7 specimens in NHMW labelled “Norway”. Widely distributed species in the southern Palearctic and Afrotropical Regions. Biology. – Hibernate as adults. In Germany recorded many times inside buildings together with Thaumatomyia notata Meigen (Tschirnhaus, 1992). Also found in birds’ nests (Turdus philomelos and Hirundo rustica) (Zuska, 1966; Krivokhatsky & Nartshuk, 2001). Some specimens were reared from a disposal tip (Werner, 1997).

112. Tricimba (Tricimba) lineella (Fallén, 1820) Figs 554–561 Oscinis lineella Fallén, 1820: 8. Lectotype in NRMS (Andersson, 1963). 152

Body black grey-dusted, except shining lower part of pleura. Frons yellow in front, infuscate behind. Ocellar triangle thick grey-dusted, nearly extending half length of frons. Antennae yellow, first flagellomere orbicular, yellow with small dark spot at aristal base. Arista dark. Head setae short brown. Genae yellow, about half as wide as first flagellomere. Vibrissal angle weakly produced. Palpi yellow. Scutum densely grey-dusted with 5 longitudinal sulci of punctuations along acrostichal and dorsocentral lines and laterally from the latter ones. Scutellum somewhat conical, slightly wider than long, very thick, with scutellar setae arising on lower surface. Apical scutellar setae shorter than scutellum, two pairs of subapical setae, all stout. Pleura mainly shining. npl 1 + 2. Legs mainly yellow, middle and posterior femora and tibiae with dark strikes. Body length 1.1–1.5 mm. Distribution. – Denmark: SJ, NEZ. Sweden: north to UP. Norway: AK: Hovedøya, Oslofjord. Finland: Al. Holarctic species, known in the Palearctic Region from Europe to Japan and Israel.

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, mainly in forests, often on flowers, but also in parks in cities (Allen, 1981b). Adults hibernate in old birds’ nests in America (Malloch, 1934, as T. spinigera). The larvae develop in many different fungi: Cantharellus cibarius, Calocybe gambosa, Collybia dryophila, Amanita verna, Cortinarius uraceus, In-

ocybe bresadolae, Russula pectinata, Xerocomus chrysenteron, Leccinum calopus, Paxillus involutus, Lycoperdon gemmatum, and probably Langermannia gigantea (Falcoz, 1930; Dely-Draskowits, 1972; Allen, 1981b; Krivosheina et al., 1986; Yakovlev, 1994). Also from rotting spathes of skunk cabbage Symplocarpus foetidus (Araceae) in North America (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 Camarota Meigen, 1830: 7. Type species: Camarota flavitarsis Meigen, 1830 (= Oscinis curvipennis Latreille, 1805) by monotypy. Black species with triangular head, thick arista, large ocellar triangle and curved wings with thick veins, vein R2+3 fused with R1 and costa. Frons distinctly projected, face with narrow carina. In eyes anterior ommatidia larger than posterior. First flagellomere trapezoid. Tibial organ present. Male genitalia: mesolobus short wide, middle lobe of surstylus with hook-like apical end. Gonites divided into post- and pregonites. Distribution. – Palearctic and Afrotropical Regions. 154

113. Camarota curvipennis (Latreille, 1805) Figs 42, 562–568, 956–959 Oscinis curvipennis Latreille, 1805: 383. Syntype, 1 specimen, in MNHN (Ismay, 1999). Body black with partly yellow head and legs. Head longer than wide with very short setae. Eyes oval with horizontal long axis. Antennae mainly black, but first flagellomere with yellow spot around arista insertion. Arista broad lanceolate black. First flagellomere 1.5 times longer than wide. Genae broad with black spot. Ocellar triangle black reaching margin of frons, with a shallow depression in front of ocellar tubercle. oc short. vte and vti indistinct. Palpi yellow. Scutum black subshining with coarse microsculpture, covered with pale setulae. Scutel-

lum rounded wider than long. Apical setae short convergent, situated on the middle of the disk. Pleura mainly dusted and smooth. Anepisternum with shining wrinkled area below. Katepisternum shining at middle and in front below, remaining part silvery dusted. Wings brownish, convex, and are folded over the abdomen at rest and give the fly a beetle-like appearance. Halteres black. Legs with femora black, tibiae darkened in the middle. Tibial organ narrow oval. Body length 2.5–3.0 mm.

Tibial organ present. Intra-epandrial sclerite well developed. Mesolobus large. Surstyli with distinct upper lobe and hook-like middle lobe. Gonites divided into pre- and postgonites.

Distribution. – Nartshuk have seen two females in MNHN labeled “Denmark”. We have not seen other material from the territory investigated. Duda (1933) mentioned specimens from Finland and Sweden, but Nartshuk did not find specimens in NHMH and NRMS, and there are no specimens in ZMLU. There is only one species of the genus in the Palearctic Region. The species is known from all Europe except for the northern parts, and from the Caucasus, southern part of Palearctic Asia and North Africa.

Figs 24, 569–575

Biology. – Larvae phytophagous, develop in shoots and ears of different cereals: barley, oats, rye, wheat, but also on Elytrigria repens and possible other grasses. Four or five larvae may live in a single stem. Infestation of young shoots results in arrested growth and no development of ears. The species is considered as a pest of cereals in some countries of southern Europe (Italy, France, former Yugoslavia, Bulgaria) (Rondani, 1873; Marchal, 1894, 1897; Osten-Sacken, 1896; Balachowsky & Mesnil, 1935; Lyubenov, 1960; Grujičić, 1961; Zamfirov, 1962; Krusteva & Beschovski, 1998, 2000). Larvae of Elachiptera cornuta have been reported living together with the present species. Larvae described by Marchal (1884), Balachowsky & Mesnil (1935), and description repeated by Nye (1958).

Genus Centorisoma Becker, 1910 Centorisoma Becker, 1910: 106. Type species: Centorisoma elegantulum Becker, 1910 by monotypy. Body elongated, usually black with coarsely sculptured scutum and scutellum, more rarely thorax yellow and scutum smooth. First flagellomere more or less prolonged with thick usually white arista. Ocellar triangle large smooth shining. Face and genae with central dark shining sclerotized area.

Distribution. – Palearctic Region, most species found in eastern part.

114. Centorisoma elegantulum Becker, 1910 Centorisoma elegantulum Becker, 1910: 107. Syntypes in HNHM. Body black with partly yellow head and legs. Ocellar triangle large, behind broad as frons, rounded off at apex, reaching margin of frons, black, shining. Head setae small. First flagellomere 1.5 times as long as wide. Arista thick white haired. Genae as wide as first flagellomere, with black spot. Face with black spot. Scutum with coarse granulate sculpture, dusted behind. Scutellum dusted. Apical scutellar setae short and rather far apart. Legs yellow, middle tibiae with rather long black spur. Halteres yellow. Body length 2.0–2.5 mm. Distribution. – Only from Vib: Bolshoy Beresovy Is., Vyborg (Nartshuk, 1984a). Eurasian species, known from Europe and West Siberia eastwards to the Baikal Lake. Biology. – Rather scarce. Adults in dry meadows and on sandy beaches, often caught on Elytrigia repens. Larvae develop in shoots of Elytrigia repens (Karpova, 1972; Nartshuk, 1972a).

Genus Cetema Hendel, 1907 Cetema Hendel, 1907: 98, replacement name for Centor Loew, 1866 (preocc. Schoenherr, 1848). Type species: Oscinis cereris Fallén, 1820, automatic. Black species with partly yellow head, pleura, scutellum and legs. Body elongated. Ocellar triangle large, shining. First flagellomere rounded squarish, as long as wide. Arista white or black. Scutum moderately convex, coarsely sculptured. Middle tibiae with rather long spur. Front and middle tibiae in males sometimes with long hairs. Tibial organ 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 epandrium with posterior lateral corners prolonged and tapering. Middle lobe of surstyli serrate at apex; pregonites very long, postgonites with short hook at posterior end. In females sternum 5 very large.

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Distribution. – Holarctic. –

Key to species of Cetema Hendel 1

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156

Arista white. Anterior and middle tibiae of male with long hairs (Fig. 577). Male genitalia Figs 578, 579 . . . . . . . . . . . . . C. cereris (Fallén) Arista black. Anterior and middle tibiae of male with or without long hairs . . . . . . . . . . 2 Anterior and middle tibiae of male without long hairs. Epandrium of male genitalia with straight processes (Figs 581–584) . . . . . . . . . . . . . . . . . . . C. elongatum (Meigen)

Anterior and middle tibiae of male with long hairs (as in Fig. 577) . . . . . . . . . . . . . . . . . . . . . 3 Male genitalia: processes of epandrium straight, long and tapering (Fig. 587). Pregenital sclerite yellowish . . . . . . . . . . . . . . . . . . . . . C. neglectum Tonnoir Male genitalia: processes of epandrium curved mesad (Figs 585, 586). Pregenital sclerite black . . . . . . . . . . . . . . . . . . . . . C. myopinum (Loew)

115. Cetema cereris (Fallén, 1820) Figs 576–580 Oscinis cereris Fallén, 1820: 3. Lectotype in ZMLU (Andersson, 1963). Frons almost as long as wide, ocellar triangle black, shining. Genae half as wide as first flagellomere or

a little wider than that. First flagellomere slightly longer than wide, mainly yellow with infuscate dorsal margin. Arista white. Scutum black shagreened, postpronota yellow with black spot. Pleura yellow with black spots on all parts. Legs almost yellow except front tarsus and 2–3 last tarsal segments of other legs. Front and middle tibiae and their basitarsi with long white hairs in males and black spur on tip. Terminal segment of male abdomen weakly curved ventrad and clubbed in lateral view. Tergites 4 and 5 longer than tergite 3, synsternite 7 + 8 rather large and sclerotized in male. Sternite 5 of female abdomen enlarged. Body length 3.0–3.5 mm. Distribution. – Rather common species. Denmark: found in most districts. Sweden: everywhere northwards to LY. Norway: AK, HEs, HEn, Os, Bø, Bv, VE, AAy, HOy, HOi, Nnø. Finland: northwards to Li. Russia: Vib, Kr, Lr. Trans-Palearctic species in boreal and temperate zones, extending from Great Britain to Sakhalin (Russia) and Japan. Séguy (1934) recorded the species from 2000 m in Norway.

Biology. – Adults in June to August in meadows and edges of forests. Univoltine species, overwinters as larva. Larvae phytophagous, develop in shoots of Poaceae. Host plants: Dactylis glomerata, Holcus lanatus, Elytrigia repens, Lolium perenne, L. multiflorum, Hordeum murinum, Agrostis tenuis, A. alba, A. stolonifera, A. canina, Poa annua, P. pratensis, P. nemoralis, P. trivialis, Phleum pratense (Belachowsky & Mesnil, 1935; Nye, 1958; Karpova, 1972; Nartshuk, 1972a; Mowat, 1975; Tschirnhaus, 1981, 1992; Nielsen, 1994). Larvae described by Kreiter (1928) and Balachowsky & Mesnil (1935). A photo of the anterior end of the larva is given by Sol (1974).

117. Cetema myopinum (Loew, 1866) Figs 585, 586 Centor myopinus Loew, 1866: 9. Type material not found in ZMHU (Wendt, 1975, 1978).

Biology. – Adults in June to August, in meadows, forest edges and clearings. Johansson (1960) remarked briefly on a habitat in Sweden. Larvae phytophagous, develop in shoots of Poacea: Poa, Alopecurus, Festuca, Agrostis, Secale (Balachowsky & Mesnil, 1935; Nye, 1958; Nartshuk, 1972a; Tschirnhaus, 1981, 1992).

Body stout and in male hind end very large and curved ventrally. Frons yellow, ocellar triangle shining black, extending 2/3 length of frons. Arista black. In male front and middle tibiae with long white hairs. In female the 5th sternite of abdomen large. Setulae of abdomen pale. Body length 2.5– 3.0 mm.

116. Cetema elongatum (Meigen, 1830)

Distribution. – Denmark: SJ, WJ, ?NEZ. Sweden: northwards to JÄ. Norway: HEs, Bø, Bv. Finland: northwards to Om. Russia: Vib, Kr. Eurasian species, known from Great Britain to the Baikal Lake and northern Mongolia.

Figs 581–584 Chlorops elongatus Meigen, 1830: 151; Meigen’s plate 211: 2 (Morge, 1976b). Lectotype in NHMW (Nartshuk, 1997a). Cetema obliqua Beschovski, 1984: 213. Holotype in ZMBA. Cetema simile Ismay, 1985: 35. Holotype in OXUM. Smaller with narrower body than in preceding species. Front and middle male legs without long white hairs. Arista black. Setulae of abdomen dark. Male genitalia: epandrium small, slightly laterally compressed. Body length 2.0–2.5 mm. Distribution. – Denmark: found in all districts. Sweden: northwards to LY. Norway: northwards to Nsy. Finland: northwards to Om. Russia: Vib, Kr. European species.

Biology. – Adults in June to August in meadows. Larvae phytophagous, develop in shoots of many grasses, but prefer species of Agrostis (Nartshuk, 1962a). Hodson (1926) recorded the species from spring oats. He found several larvae living together in one shoot; they also pupated in the same shoot. Newly emerged adults copulated in their cage, but no eggs or larvae were found on oats shoots presented to them. Nye (1958) considered this attack due to larval migration from a newly ploughed grassfield.

118. Cetema neglectum Tonnoir, 1921 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 MHNB. Similar to C. cereris, but arista black and epandrium in male yellowish. Male front and middle tibiae with long white hairs. Setulae of abdomen pale. Pregenital sclerite yellowish. Body length 2.8– 3.0 mm. Distribution. – Denmark: from all districts except NWJ and B. Sweden: northwards to ÅN. Norway: AK: Landøya; Bø: Verksøya, Hurum. Finland: only in southern part: Al, Ab, N, Sa. Russia: Vib. European species. Biology. – Adults in June to August in meadows. Univoltine species. Larvae phytophagous, develop in shoots of Poaceae. Host plants: Lolium perenne, Festuca pratensis, Poa trivialis, P. pratensis, Agrostis stolonifera, Elymus pycnanthus. Larvae hibernate and pupate the following spring. Hodson (1926) found larvae in tillers of spring oats that he identified with C. myopinum, but Collin (1966) indicated that they belong to the present species. Probably this attack was due to larval migration from a newly ploughed grassfield nearby (Nye, 1958). Larvae described by Nye (1958).

has modified male front tarsi with shortened basitarsi and asymmetrical claws and a sclerotized aedeagus Distribution. – Worldwide.

Key to species of Chlorops Meigen 1

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2 (1)

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Genus Chlorops Meigen, 1830 Chlorops Meigen, 1830: 278. Type species: Musca pumilionis Bjerkander, 1778 by designation of ICZN, 1955: 423. Opinion 348. Name No 868 in Official List of Names in Zoology. Yellow species usually with black or rarely with yellow-red stripes on scutum and spots on pleura. Ocellar triangle differs in shape, usually shining, sometimes with ridges along lateral margins or sulcus in the middle. Antennae yellow to black. First flagellomere rounded, as long as or a little longer than wide. Arista slender, black or white. Palpi short or long and thick, usually yellow, rarely entirely black or black on tip. Scutum smooth, dusted or shining. Stripes black or red, sometimes partly or completely fused. Tibial organ absent. Male genitalia: middle lobe of surstylus with large granulate or hook like apex; lower lobe long hairy; pregonites and postgonites situated parallel to each another, and often equal in length. A species group within Chlorops, often recognized as subgenus Sclerophallus Beschovski, 1978,

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4 (1) –

Scutum entirely black, postpronota black or yellow with black spots. Scutellum sometimes black or darkened . . . . 2 Scutum yellow with 3–5 black or reddish stripes. Postpronota usually yellow with black spot. Scutellum usually yellow, very rarely with dark middle . . . . . 4 Basal segments of antennae yellow or fuscous yellow. Genae wider than first flagellomere, often with black setulae. Yellow parts of body usually darkened .................................3 Antennae entirely black. Genae at most as wide as first flagellomere. Yellow parts of body sharply contrasted with black parts. Ocellar triangle with acute apex and not reaching front margin of frons (Fig. 673). Scutum smooth and polished. Male genitalia Figs 675, 876 . . . . . . . . . . . . C. scutellaris (Zetterstedt) Ocellar triangle black with strongly demarcated outline, sometimes with windows of lighter colour outside ocelli, but these tend to disappear in dark specimens (Fig. 679). Male genitalia: outer gonites short (Figs 681, 682) . . . . . . . C. speciosus Meigen (dark form) Border of ocellar triangle not so strongly outlined, the tip often yellowish, but no light windows on the sides of ocelli. Male genitalia: outer gonites long (Figs 636, 637) . . . . . C. meigenii Loew (dark form) Palpi long and thick, sometimes black on tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Palpi not long, yellow, sometimes black on tip or entirely black, in that case arista white and markedly pubescent .................................9 159

5 (4)

Stripes on scutum dusted. Palpi greatly darkened in both sexes (Fig. 640). Male genitalia Figs 641, 642 . . . . . . . . . . . . . . . . C. nigripalpis (Duda)

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Stripes on scutum more or less shining. Palpi yellow, darkened at tip in one sex, only in male or only in female . . . . . . . 6

6 (5)

Genae not wider than first flagellomere. Ocellar triangle with more or less distinct groove in apical part. Palpi, especially in male, darkened at tip . . . . . . . . . . . . . . . . . C. palpatus Smirnov

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Genae wider than first flagellomere. Ocellar triangle without groove in apical part. Palpi yellow or darkened only in female . . . . . . . . . . . . . . . . . . . . . . . . . 7 Spot on katepisternum yellow . . . . . . . 8

7 (6) –

8 (7)

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9 (4)

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Spot on katepisternum black. Abdomen yellow with distinct narrow black bands on front margins of tergites 2–5 and sometimes with black median line. First flagellomere from yellow to almost black. Stripes on scutum shining or subshining, only thinly pollinose. Male genitalia Figs 671, 672 . . . C. scalaris Meigen Ocellar triangle mainly yellow with small dark spot only on ocellar tubercle and around it. Stripes on scutum shining. Abdomen predominantly yellow. Palpi yellow in both sexes, palpi of female longer and thicker, than in C. ringens . . . . . . . . . . . C. centromaculatus (Duda)

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10 (9)

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11 (10)

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12 (9)

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13 (12)

Ocellar triangle predominantly black, yellow only along side margins and sometimes with yellow window spots. Stripes on scutum subshining, thinly pollinose. Abdomen yellow with black bands, sometimes connected by median line. Palpi in female darkened at tip. Male genitalia Figs 663, 664 . . . . . . . . . . . . . . . . . . . . . C. ringens Loew

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Ocellar triangle smooth, shining, yellow with two small black spots, one on ocellar tubercle and another on apex of ocellar triangle, sometimes black spots are connected by dark line . . . . . . . . . . . . 10

14 (13)

Ocellar triangle yellow or black, often black spot occupies only central part of triangle leaving sides yellow. If small black spots are on ocellar tubercle and on triangle then surface of triangle not smooth but wrinkled along sides . . . . 12 Antennae and palpi black. Arista white, slightly thickened by dense pubescence. Black ocellar and apical spots on ocellar triangle connected by dark line. Male genitalia Figs 592, 593 . . . . . . . . . . C. anthracophagoides Strobl Basal antennal segments and palpi always yellow. Arista black, grey or whitish and slender with short pubescence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 First flagellomere yellow. Stripes on scutum usually red, but sometimes black, grey-dusted. Male genitalia Figs 666, 667 . . . . . . . . . . . . . . C. rufijinus (Zetterstedt) First flagellomere black. Stripes on scutum black, grey-dusted . . . . . . . . . . . . . C. planifrons Loew forma triangularis Becker Genae narrower, subequal to width of first flagellomere or narrower (Figs 619, 639, 693) . . . . . . . . . . . . . . . . . . . . . . . . 13 Genae broader, equal to width of first flagellomere or usually distinctly wider (Fig. 623) . . . . . . . . . . . . . . . . . . . . . . . . 17 Male front metatarsus long, 5 times longer than thick, front claws symmetrical. Stripes on scutum shining and abdomen predominantly yellow with black setulae or stripes on scutum dusted and abdomen dark dorsally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Male front metatarsus short, no more than 3 times longer than thick, front claws asymmetrical. Stripes on scutum dusted. Abdomen blackish or in male pale yellow with white setae or darkened with black setulae, in female reddish brown . . . . . . . . . . . . . . . . . . . . . . . 15 Stripes on scutum black, shining. abdomen predominantly yellow with black setulae. Basal segments of antennae yel-

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15 (13)

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16 (15)

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17 (12)

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18 (15) – 19 (18)

low. Ocellar triangle only with central black spot, leaving sides yellow. Male genitalia Figs 620, 621 . . . . . . . . . . . . . . . C. hypostigma Meigen Stripes on scutum black, dusted, abdomen dark dorsally. Antennae and ocellar triangle entirely black . . . . . . . . . . . . . . . . . C. bjerkanderi sp. n. Anepisternum with some small setulae, which are usually black in female and white in male. Ocellar triangle with yellow sides . . . . . . . . . . . . . . . . . . . . . . . . 16 Anepisternum bare without setulae. Ocellar triangle entirely black. Palpi yellow. Basal antennal segments yellow. First flagellomere rather large, black. Borders of ocellar triangle sharply outlined. Abdomen blackish (Figs 631, 632) . . . . . . . . . . . . . . . . . . C. limbatus Meigen Setulae on scutum blackish. Ocellar triangle mainly black, side margins and hinter corners brownish or sometimes yellow. Antennae black or brownish below. Palpi black or yellow. Abdomen reddish brown in female and darkened dorsally in male. Male genitalia Figs 694, 695 . . . . . . . . . . . . C. varsoviensis Becker Setulae on scutum whitish. Ocellar triangle widely yellow at sides. First flagellomere yellow below. Palpi yellow. Abdomen yellow with small black spots on the second tergite. Male genitalia Fig. 647 . . . . . . . . . . . C. pallidiventris (Duda) Stripes on scutum black, shining, without dusting or subshining and thinly dusted. Spot on katepisternum black, shining or reddish yellow without dusting . . . . . . . . . . . . . . . . . . . . . . . . . . 18 Stripes on scutum black densely greydusted. Spot on katepisternum greydusted, shining black or reddish yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 Wings weakly darkened, greyish or brownish . . . . . . . . . . . . . . . . . . . . . . . . 19 Wings uncoloured . . . . . . . . . . . . . . . . 20 First flagellomere and spot on katepisternum black. Ocellar triangle predom-

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20 (18)

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21 (20)

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inantly black. Outside margin of inner lateral stripes on scutum with triangular projection behind postpronota (Fig. 665) . . . . . . . . . . . . . . C. rossicus Smirnov First flagellomere yellow or darkened on upper and outer sides. Spot on katepisternum yellow or brownish on upper part. Black spot on ocellar triangle not occupying apical part, side margins and hind corners of triangle. Outside margin of inner lateral stripes on scutum more or less even, without projection (Fig. 635). Male genitalia Figs 636, 637 . . . . . . . . . . . . . . . . . . . . C. meigenii Loew Stripes on scutum shining. Ocellar triangle entirely black or with fuscous yellow window spots laterally from central line, but side margins and hind corners always black. Borders of ocellar triangle sharply outlined. Genae with black hairs. Body larger, 3.5–6.0 mm. Male genitalia Figs 681, 682 . . . . . . . . . . . . . . . . . C. speciosus Meigen Stripes on scutum subshining with thin dusting. Ocellar triangle black or yellow along side margins and on hind corners, usually without light window spots. Border of black parts not so sharply outlined. Genae without black hairs. Body smaller 3.0–3.5 mm . . . . . . . . . . . . . . . 21 Abdomen brown or black dorsally. Basal segments of antennae brownish. Ocellar triangle entirely black or narrowly yellow along side margins and on hind corners. Black stripes on scutum wider. Female cerci triangular if adjoined (dorsal view). Male genitalia Fig. 691 . . . . . . . . . . . C. troglodytes (Zetterstedt) Abdomen dorsally yellow with black bands. Ocellar triangle broadly yellow along side margins and on hinter corners. Basal segments of antennae yellow. Black stripes on scutum narrower. Female cerci truncate if adjoined (dorsal view). Male genitalia as Fig. 597 . . . . . . . . . . . . . . . . . C. calceatus Meigen

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23 (22)

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25 (22)

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26 (25)

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Pubescence on first flagellomere longer than aristal pubescence (Fig. 599). Ocellar triangle entirely black to mainly yellow. Anepisternum often with some black or white setulae . . . . . . . . . . . . . 23 First flagellomere with shorter pubescence. Anepisternum with some white or black setulae or without setulae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 Anepisternum without setulae. First flagellomere yellow to brownish, Palpi from yellow to black. Abdomen with wide brown bands on tergites. Male genitalia Fig. 600 . . . . . . C. fasciatus Meigen Anepisternum with setulae . . . . . . . . 24 Antennae and palpi black. Ocellar triangle mainly black. Scutellum often darkened. Male genitalia Figs 589, 590 . . . . . . . . . . . . . . . . . C. adjunctus Becker Antennae and palpi yellow, first flagellomere slightly darkened on upper side. Ocellar triangle yellow with darkening on corners and along sides to brownish, with small groove from anterior ocellus to apex. Scutellum yellow . . . . . . . . . . . . . . . . . . C. dasycerus Loew Arista white, slender or thickened with dense pubescence. Ocellar triangle wrinkled on apex and along sides. Scutellum often darkened. Palpus sometimes black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 Arista black, slender with short pubescence. Ocellar triangle black or partly yellow, smooth, not wrinkled. Palpi yellow . . . . . . . . . . . . . . . . . . . . . . . . . . 29 Antennae predominantly yellow, first flagellomere darkened on upper side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 Antennae entirely or only first flagellomere black . . . . . . . . . . . . . . . . . . . . . . . . 28 First flagellomere slightly longer than wide. Palpi and scutellum yellow. Spot on katepisternum yellow. On scutum stripes M2 and M3 connected in front (Fig. 650). Male genitalia Figs 651, 652 . . . . . . . . . . . . . . . . C. pannonicus Strobl

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30 (29)

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31 (29)

First flagellomere as long as wide. Palpi and middle of scutellum black or brownish. Spot on katepisternum black, greydusted. Male genitalia Figs 688, 690 . . . . . . . . . . . . . . . C. strigulus (Fabricius) Basal segments of antennae yellow. Central scutal stripe (M1 ) extending through scutellum. Scutal stripes M2 and M3 connected only in front (Fig. 604). Body smaller, 3.0–3.5 mm. Male genitalia Figs 605, 606 . . . . . . . . . . . C. fijinitimus Becker Basal segments of antennae black. Both stripes on each side of scutum fused. Body larger, 4.0–4.5 mm. Male genitalia Figs 610, 611 . . . . . . . C. frontosus Becker Spot on katepisternum black and densely grey, dusted . . . . . . . . . . . . . . . . . . . 30 Spot on katepisternum shining or subshining black without dusting, or reddish yellow . . . . . . . . . . . . . . . . . . . . . . . 31 Anepisternum with some black setulae. Basal segment of antennae yellow, first flagellomere yellow with darkened upper side. Front metatarsus and tip of front tibiae yellow. Ocellar triangle mainly yellow with black central line and without median groove before front ocellus. Male genitalia Fig. 628 . . . . . . . . . . . . . . . . . . . . C. laetus Meigen Anepisternum bare, without setulae. Basal segment of antennae black or strongly darkened, first flagellomere black, very rarely fuscous yellow at base on inner side. Front metatarsus and tip of front tibiae blackish. Ocellar triangle entirely black to almost yellow with black median line and black ocellar tubercle, in front of front ocellus with a groove that transforms into a low ridge in apical part of the triangle. Male genitalia Figs 661, 662 . . . . . . . . . . . C. pumilionis (Bjerkander) First flagellomere black or at most slightly brownish at base on inner side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32

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32 (31) – 33 (32) –

34 (33)

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35 (33)

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36 (32)

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37 (36)

First flagellomere yellow or at most darkened only on upper and outer sides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 Spot on katepisternum black . . . . . . . 33 Spot on katepisternum reddish yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 Ocellar triangle wrinkled on sides or also in apical part . . . . . . . . . . . . . . . . . . . . 34 Ocellar triangle smooth, at most with central groove before of front ocellus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 Ocellar triangle yellow, shining with small dark spot on ocellar tubercle. Black stripes on scutum narrow, divided by wide yellow ones. Scutellum yellow . . . . . . . . . . . . . . . . . C. pallifrons (Duda) Ocellar triangle black, shining. Black stripes on scutum wide, divided by narrow yellow ones. Scutellum often darkened . . . C. obscurellus (Zetterstedt) Ocellar triangle large with short linear part which is no more than 1/4–1/5 length of frons. Postnotum evenly dusted. Female cerci long and narrow, apart in dry specimens . . . . . . C. planifrons Loew (typical form) Ocellar triangle shorter, linear part longer, nearly 1/2 length of frons. Dusted parts on postnotum divided by shining part. Female cerci short . . . . . . . . . . . . . . . . . C. riparius Smirnov Ocellar triangle large with short linear part no more than 1/4–1/5 length of frons. Female cerci long and narrow and apart in dry specimens. Male genitalia Figs 655–657 C. planifrons Loew (intermediate form) Ocellar triangle with linear part nearly 1/2 length of frons. Female cerci short . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 Head higher than long (Fig. 677). Genae broad. Arista black. Black spot on ocellar triangle small, usually with two window spots near ocelli. Palpi yellow. Male genitalia Fig. 678 . . . . . . . . . C. serenus Loew

–

38 (31)

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39 (38) – 40 (39)

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41 (39)

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Head as high as long. Arista white. Palpus usually black at tip. Black spot on ocellar triangle variable, usually with yellow window spots, sometimes reduced to small spots near ocelli and median line . . . . . . . . . . . . C. zernyi (Duda) Ocellar triangle large, smooth, shining, reaching front margin of frons, without distinct linear part (Fig. 622). Stripes on scutum often partly reddish brown. Male genitalia Fig. 624 . . . . . . . . . . . . . . . C. interruptus Meigen Ocellar triangle with more or less expressed linear part, smooth or wrinkled or swelled along lateral sides, often with whitish spots in front of ocellar tubercle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 Ocellar triangle wrinkled at apex and along side margins (Figs 612, 625) . . . 40 Ocellar triangle smooth, not wrinkled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 Ocellar triangle dull, short, extending 0.5 length of frons and continues to anterior margin of front as a line (Fig. 625). Lateral sides of ocellar triangle swelled. Black spots situated on ocellar tubercle, on side margins and on apex. Palpi short and thick. Male genitalia as Figs 626, 627 . . . . . . . . . . . C. kirigaminensis Kanmiya Ocellar triangle shining, long, linear part not so distinct (Fig. 612). Median part of ocellar triangle whitish. Male genitalia as Figs 613, 614 . . . C. geminatus Meigen Ocellar triangle more or less evenly narrowing to apex, blackish. Anepisternum with some yellow setulae. Body smaller, 3–4 mm. Male genitalia Fig. 601 . . . . . . . . . . . . . C. fijiguratus (Zetterstedt) Ocellar triangle sharply narrowed in middle of frons with long linear part, often with whitish spots before ocelli (Fig. 615). No setulae on anepisternum. Body larger, 4–6 mm. Male genitalia Fig. 617 . . . . . . . . . . . . . . . . . . . C. gracilis Meigen

163

Species key to regionally known 3rd instar larvae of Chlorops Meigen 1

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2 (1) –

3 (2)

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4 (3)

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5 (2)

164

Posterior spiracles situated on distinct lobes of anal segment, sometimes lobes rather small or brownish coloured (Figs 990, 993–995, 998, 1000, 1004). Body of larvae cylindrical elongated (Figs 988, 991) ...................................2 Posterior spiracles situated on surface of anal segment (Fig. 987). Body of larvae nearly oval and slightly flattened (Fig. 986). Larvae 4–5 mm long, in galls on Brachypodium and Elytrigia . . . . . . . . . . . . . . . . . . C. strigulus Fabricius Lobes of anal segment longer than wide (Fig. 993) . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Lobes of anal segment not longer than wide, sometimes very small, brownish coloured (Figs 998, 1000) . . . . . . . . . . . . . 5 Posterior spiracles situated on the end of lobes (Figs 993, 994). Maxillary brown papillae close to each another (Fig. 992). Anus wide, occupies all width of segment ...................................4 Posterior spiracles situated on the inner surface of lobes (Fig. 990) and without interspiracular processes. Maxillary brown papillae wide apart (Fig. 989). Larvae up to 10 mm long, in shoots of large species of Carex . . . . . . . . . . . . . . . C. frontosus Meigen Maxillary palpi (Fig. 992) large, brown ring surrounding them narrow. Posterior spiracles with short interspiracular processes. Larvae up to 10 mm long, in shoots of Phalaris arundinaceae . . . . . . . . . . . . . . . . . . . . C. limbatus Meigen Brown ring surrounding maxillary palpi wide, especially on inner side. Posterior spiracles without interspiracular processes. Larvae 8–10 mm long, in shoots of Carex . . . . . . . . . . . . . . . . . . . . . . C. zernyi (Duda) Lobes of anal segment brown and tapering, posterior spiracles probably not functional (Fig. 995). Larvae 7–10 mm long, in shoots of large species of Carex . . . . . . . . . . . . . . . . . . . . C. planifrons Loew

–

6 (5)

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7 (6)

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Lobes of anal segment small, not brown or tapering (Fig. 998), but posterior spiracles may be brown (Fig. 1001) . . . . . . . . . . . . . . 6 Posterior spiracles not coloured, rather wide apart on edges of anal segment (Fig. 998). Spicular zones rather narrow. Larvae 5.5–7.5 mm long, in shoots of cereals and Elytrigia repens . . . . . . . . . . . . . . C. pumilionis (Bjerkander) Posterior spiracles coloured (Fig. 1001), situated on edges of anal segment or on inner sides of incision of anal segment (Figs 1000, 1004) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Posterior spiracles (Fig. 1001) somewhat cap-shaped with 3–4 projections and brown. Anus small, not more than 1/6 of width of anal segment (Fig. 1000). Larvae 5–6 mm long, in shoots of Elytrigia repens . . . . . . . . . . . . . . . . . . C. interruptus Meigen Posterior spiracles situated on inner side of incision of anal segment; anus wide, occupies 2/3–3/4 of width of anal segment (Fig. 1004). Larvae up to 10 mm long, in shoots of Deschampsia caespitosa . . . . . . . . . . . . . . . . . . . C. speciosus Meigen

119. Chlorops adjunctus Becker, 1910 Figs 589, 590 Chlorops adjunctus Becker, 1910: 90. Syntypes in HNHM. Ocellar triangle and antennae black. First flagellomere with white pubescence. Palpi blackish. Scutal stripes broad, nearly confluent, black densely dusted. Central stripe continues onto scutellum and scutellum darkened. Anepisternum with several black setulae. All pleural spots black. Abdomen dorsally black and with distinct setulae. Body length 3.0–4.0 mm. Distribution. – Rare species. Only from Sweden: SK: Vitemölla, 25.vii.1919 (E. Wahlgren), 1 male. European species.

120. Chlorops anthracophagoides Strobl, 1901 Figs 591–593

Chlorops anthracophagoides Strobl, 1901: 231. Holotype female in NMBA (Chvála, 2008). Ocellar triangle yellow with black side margins and two black spots, one on ocellar tubercle and another on tip. Both black spots connected by black line, as result forming two yellow windows on ocellar triangle. Antennae black, first flagellomere rather large. Arista thick white. Genae narrower than first flagellomere. Palpi black. Scutal stripes broad black, dusted, scutum before scutellum darkened. Katepisternum with black spot. Legs yellow. Body length 2.0–2.5 mm. Distribution. – Rare species. Denmark: Sandager (“Sandacker”), Rinkenæs, 24.vi.1896 (W. Wüstnei), 1 male. Sweden: NB: Eriksberg (in NHMH, Nartshuk, 1998). Russia: Vib, Kr. Not known yet from Norway and Finland. Euro-Caucasian species.

121. Chlorops bjerkanderi Nartshuk & Andersson, sp. nov. Figs 594–596

Etymology. – The species is named in honor of the Swedish scientist C. Bjerkander, who described the first species of the genus.

122. Chlorops calceatus Meigen, 1830 Fig. 597 Chlorops calceatus Meigen, 1830: 146; Meigen’s plate 160: 2 (Morge, 1976a). Lectotype, 1 female, in MNHN (Nartshuk, 2006). Ocellar triangle extending half length of frons, smooth shining black, but hind corners yellow. Antennae with basal segments yellow, first flagellomere black. Arista black. Genae wider than first flagellomere. Palpi small yellow. Scutal stripes black, thinly dusted and subshining. Spot on katepisternum black, shining. Abdomen yellow with narrow black bands on tergites and central black line. Legs yellow with black last tarsal segments. Cerci of female black. Body length 2.5 mm.

Holotype, female, Sweden: SK: Rövarekulan, Rolsberga, 01.ix.1974 (H. Andersson). Holotype in ZMLU.

Distribution. – Very common. Denmark: from all districts except NWJ and SZ. Sweden: northwards to LY. Norway: northwards to STi. Finland: northwards to ObS. Russia: Vib, Kr. Eurasian species.

Diagnosis. – A new species characterized by the following combination of characters not seen in any other Palearctic species: ocellar triangle and antennae entirely black, genae narrow, scutal stripes narrow, dusted, spot on katepisternum black, shining.

Biology. – Adults in June to early August, in meadows and forest edges. Larvae phytophagous, living on Festuca rubra (Tschirnhaus, 1981) and possibly other grasses.

Description. – Body yellow with black marks. Frons longer than wide, yellow, covered with short black setulae. All head setae black, rather short, vte, vti and poc equal in size. Ocellar triangle entirely black, shining, extending 2/3 length of frons. Occiput with wide black stripe leaving narrow yellow stripes along eyes margin. Antennae entirely black. Palpi yellow. Scutum longer than wide, ratio length:width as 3.0:2.3. Scutal stripes narrow, dusted, central one not extending to scutellum. Scutellum yellow. Pleura with small black, shining mark on anepisternum, black, shining spot on katepisternum and black, dusted spot on katepimeron. Postnotum black, dusted. Abdomen brown dorsally. Wings transparent, venation usual for the genus. Halteres yellow. Legs yellow with darkened tarsi. Cerci of ovipositor wide black. Body length 3.5 mm.

123. Chlorops centromaculatus (Duda, 1933) Oscinis centromaculatus Duda, 1933: 61. Syntypes in NHMW, HNHM and probably other museums. Ocellar triangle extending 3/4 length of frons and continues as line to front margin of frons, mainly yellow, black only on ocellar tubercle and here often with yellow windows. Genae wider than first flagellomere. Basal segments of antennae yellow, first flagellomere black, arista black. Palpi rather enlarged yellow. Scutal stripes black, shining, central stripe short. Spot on katepisternum reddishyellow. Abdomen mainly yellow with very narrow black bands on tergites. Legs yellow, last tarsal segment darkened. Cerci of female yellow. Body length 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: Kvistbro, Mullhyttan, 30.vii.1941 (T. Nyholt), 1 male; UP: (Boheman), 1 female; Forsmark, 18.vii.1975 (R. Lenneborg), 1 male. Finland: N: Helsinki, 02.viii.1976, 23.viii.1977, 21.ix.1979 (M. Koponen); Ta: Karkola, 15.viii.1974 (M. Koponen). Russia: Vib: Lake Pionerskoye, Polarny, 07.viii.1998 (Przhiboro), 1 male, 1 female. Not recorded from Denmark and Norway. European species. Biology. – Probably autumnal species, adults from late July to September.

124. Chlorops dasycerus Loew, 1866 Chlorops dasycerus Loew, 1866: 46. Type material not found ZMHU (Wendt, 1975, 1978). Ocellar triangle black with pale windows. First flagellomere yellow to brownish, with longish white pubescence. Arista black. Palpi yellow or blackish. Genae wider than first flagellomere. Scutal stripes black and densely dusted. Scutellum darkened at base. Anepisternum with several black setulae. All pleural spots black. Abdomen dorsally black or brown. Body length 3.5–4.0 mm. Distribution. – Rare species. Denmark: NEJ: Rødstensklit, Skagen (Bræstrup), 1 female. Sweden: SK: Vitemölla, 15.ix.1959 (H. Andersson), 1 female. Not found in Norway, Finland or Fennoscandian districts of Russia. Euro-Caucasian species.

125. Chlorops fasciatus Meigen, 1830 Figs 598–600 Chlorops fasciatus Meigen, 1830: 143; Meigen’s plate 276: 14 (Morge, 1976b). Types not found (Becker, 1902). Ocellar triangle black with yellow side margins and posterior corners. First flagellomere from yellow to brownish, in white long pubescence. Arista black. Palpi yellow or blackish. Genae wider than first flagellomere. Scutal stripes rather narrow and densely dusted, central stripe sometimes continued onto scutellum. Scutellum yellow or with black stripe. Anepisternum without black setulae. All

pleural spots black, shining. Abdomen dorsally with black bands on tergites. Body length 3.5– 4.0 mm. Distribution. – Not yet found in Denmark and Fennoscandia, but expected in SE Fennoscandia. Eurasian species.

126. Chlorops fijiguratus (Zetterstedt, 1848) Fig. 601 Oscinis fijiguratus Zetterstedt, 1848: 2631. Holotype in ZMLU (Andersson, 1966). Chlorops zonulatus Wahlgren, 1913: 52. Synonymy by Duda (1933), confirmed by Nartshuk & Andersson (2002). Lectotype in ZMLU (Nartshuk & Andersson, 2002). Ocellar triangle extending half length of frons and continues as line to front margin of frons, mainly yellow, black on ocellar tubercle and with more or less distinct black median line. Genae distinctly wider than small first flagellomere. Antennae yellow, first flagellomere sometimes darkened above, arista black. Palpi yellow. Scutal stripes black, dusted, rather short, not reaching scutellum. Spot on katepisternum yellow or reddish-yellow. Abdomen yellow, often with very narrow black bands on tergites and black central line. Legs yellow, last tarsal segment darkened. Cerci of female yellow. Body length 2.5–3.0 mm. Distribution. – Sweden: SK, Vittskövle, female holotype of Oscinis fijiguratus. ÖL: Borgholms Alvar, 22.vi.1910, type specimens of C. zonulatus Wahlgren. Russia: Kr: Paanajärvi (Frey), 1 male. Not found in Denmark or Norway. Eurasian species. Biology. – Unknown. A record by Kryshtal (1947) that larvae damaged stems and ears of Elytrigia repens probably refers to C. novakii Strobl.

127. Chlorops fijinitimus Becker, 1910 Figs 602–606 Chlorops fijinitimus Becker, 1910: 96. Syntypes in ZMHU and DEIM (Rohlfien & Ewald, 1970; 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 of frons, black, with yellow hind corners, wrinkled along side margins, sometimes with yellow marks lateral to ocelli. Genae distinctly wider than first flagellomere. Basal segments of antennae fuscous yellow, first flagellomere black, slightly longer than wide. Arista whitish. Palpi basally yellow, apically blackish. Scutal stripes black, dusted, sublateral stripes connected with lateral in front. Scutum before scutellum darkened and this diffuse spot occupies scutellum. Spot on katepisternum reddishyellow, often with black upper part. Abdomen yellow with black bands on tergites and black central line. Legs mainly yellow, femora partly black, last tarsal segment darkened. Cerci of female yellow. Body length 2.5–3.0 mm. Distribution. – Rare species, known only from Sweden: ÖL: Byxelkrok, 05.viii.1958 (B.H. Hansen), 2 males, 1 female; Halltorps Hage, 22.vii.1978 (H. Andersson), 1 male, 1 female. The species is known from Europe, the Caucasus and Central Asia. Biology. – Adults in July to August, in dry meadows.

128. Chlorops frontosus Meigen, 1830 Figs 607–611, 988–990 Chlorops frontosa Meigen, 1830: 148; Meigen’s plates 213: 10; 276: 15 (Morge, 1976b). Lectotype in NHMW (Nartshuk, 1997a). Ocellar triangle extending nearly to front margin of frons, black, wrinkled along side margins and with central sulcus. Genae distinctly wider than first flagellomere, yellow with black lower margin. Antennae black, first flagellomere longer than wide. Aristal pubescence thick and white. Palpi black. Scutal stripes wide black, dusted, outer lateral stripes fused with inner lateral ones. Scutellum blackish. Spot on katepisternum black, dusted. Abdomen dorsally black. Legs mainly black, coxae and tibiae partly yellow. Body length 3.0–3.5 mm. Distribution. – Scarce. Denmark: SJ: Gråsten (W. Wüstnei); NEZ: Høed Skov. Sweden: SK, SÖ, UP. Russia: Kr: Kondopozhsky Distr.: Malyi Lednikovski Is., 24.vi.2003 (Polevoi), 1 female. European species. Biology. – Adults in May to June, on fens and marshes with Carex.

Larvae phytophagous, develop in shoots of large species of Carex (Séguy, 1934; d’Aguillar, 1943). Larvae described by d’Aguillar (1943).

129. Chlorops geminatus Meigen, 1830 Figs 612–614 Chlorops geminatus Meigen, 1830: 141; Meigen’s plates 159: 10; 211: 19 (Morge, 1976b). Lectotype, female, in MNHN (Nartshuk, 2006). Specimen in NHMW does not belong to the type series (Nartshuk, 1997a). Ocellar triangle extending nearly to front margin of frons, yellow, shining, wrinkled along side margins. Ocellar tubercle and tip of triangle black and sometimes central black line connects these black spots. Genae 1.5 times wider than first flagellomere, yellow. Antennae entirely yellow, first flagellomere sometimes darkened above. Arista black or brown. Palpi rather thick yellow. Scutal stripes black, dusted, rather short. Spot on katepisternum yellow. Abdomen yellow with narrow black bands on tergites. Legs yellow. Body length 3.5–4.0 mm. Distribution. – Rare. Denmark: WJ: Tipperne, Ringkøbing Fjord (E. Rald). Sweden: ÖL: Mörbylånga, 09.vii.1917 (Tullgren), 1 female; GO: Hammars, Norrlanda, 12–18.viii.1979 (R. Danielsson), 1 male, 1 female; ÖG: Kastad, Örberga, 20.vii.1979 (H.W. Waldem), 1 female. Russia: Kr: Ruskeala, 28.vii.1949 (Nyland), 1 male. Not found in Norway or Finland. Eurasian species. Biology. – Adults in July to August in meadows. It has been reared from Hierochloe odorata (Karpova, 1972).

130. Chlorops gracilis Meigen, 1830 Figs 615–617 Chlorops gracilis Meigen, 1830: 140; Meigen’s plate 160: 1 (Morge, 1976a). Holotype, specimen without abdomen and hind legs, in MNHN (Nartshuk, 2006). Ocellar triangle extending nearly half length of frons, yellow, with median slightly darkened sulcus, black on ocellar tubercle and sometimes with yellow 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 yellow. Arista rather long haired, black, but basally yellow. Palpi yellow. Scutal stripes black, densely dusted, not extended to scutellum. Spot on katepisternum yellow. Abdomen yellow shining, sometimes with narrow black stripes on tergites. Legs yellow. Body length 4.5–5.0 mm. Distribution. – Rather common in dry habitats. Sweden: northwards to JÄ. Finland: only in southern regions: Ab, N, St, Ta, Sa, Tb, Kb. Russia: Vib; Kr: Kondopozhsky Distr.: Shoksha, 13.vii.2004 (Polevoi), 1 male. Not so far recorded from Denmark and Norway. Eurasian species. Biology. – Adults in June to July in dry meadows. One generation per year. Hibernates as larvae in shoots of its host plant. Larvae phytophagous, develop in shoots of Calamagrostis epigeios (Kröber, 1910; Nartshuk, 1962a; Uffen & Chandler, 1978; Dubbert et al., 1998). Larvae of other Diptera, mostly chloropids, live in shoots infested by this species: Aphanotrigonum trilineatum, A. nigripes, Lasiosina herpini, and Asteia concinna Meigen (Asteiidae).

131. Chlorops hypostigma Meigen, 1830 Figs 618–621 Chlorops hypostigma Meigen, 1830: 141; Meigen’s plates 159: 12; 211: 20; 213: 12 (Morge, 1976b). Lectotype male, and paralectotypes 2 males and 1 female in MNHN (Nartshuk, 2006). Ocellar triangle extending nearly half length of frons, black with rather wide yellow side margins, sometimes with small yellow window before front ocellus. Genae slightly narrower than first flagellomere. Basal segments of antennae fuscous yellow, first flagellomere black. Arista black. Palpi yellow. Scutal stripes black, shining, rather narrow and not extend scutellum. Spot on katepisternum black, shining. Abdomen yellow or sometimes with narrow black stripes on tergites. Legs yellow, last tarsal segments blackish. Body length 2.0–2.5 mm. Distribution. – Common. Denmark: widespread. Sweden: northwards to LY. Norway: AK, HEs, VAy, Ri, HOi, MRy. Finland: northwards to ObS. Russia: Vib, Kr. Euro-Caucasian species.

Biology. – Adults from late May to August, in meadows, edges of forests. Univoltine species, overwintering as third instar larva within shoots of grasses. Larvae phytophagous, develop in shoots of Dactylis glomerata, Poa sp., P. trivialis, Festuca sp. (Séguy, 1934; Mowat, 1975; Uffen & Chandler, 1978; Tschirnhaus, 1981).

132. Chlorops interruptus Meigen, 1830 Figs 622–624, 999–1001 Chlorops interruptus Meigen, 1830: 145; Meigen’s plate 213: 9 (Morge, 1976b). Lectotype in NHMW (Nartshuk, 1997a). Ocellar triangle large, reaching front margin of frons, yellow shining, medially with furrow and black line from ocellar tubercle to front end. Genae nearly 1.5 times wider than first flagellomere. Antennae entirely yellow, arista black. Palpi yellow. Scutal stripes black, dusted, rather narrow and not extended to scutellum, sometimes stripes partly brownish-red. Spot on katepisternum yellow. Abdomen yellow. Legs yellow, last tarsal segments darkened. Body length 2.5–3.5 mm. Distribution. – Only from Sweden: UP: Hässelby, Börje, 03.ix.1965 (Kurs). Euro-Mediterranean species. Biology. – Two generations per year in Central Europe. Larvae phytophagous, develop in shoots of Elytrigia and ?Phragmites (Séguy, 1934; Balachowsky & Mesnil, 1935; Uffen & Chandler, 1978). The record of Phragmites as host plant needs verification. Larvae described by Balachowsky & Mesnil (1935).

133. Chlorops kirigaminensis Kanmiya, 1978 Figs 625–627 Chlorops kirigaminensis Kanmiya, 1978: 55. Holotype and paratypes in Entomological Laboratory Faculty of Agriculture Kyushu University (Kanmiya, 1978). Paratype, female in USNM, Washington, D.C. 171

“Chlorops zonulatus Wahlgren”; Smirnov, 1958; Smirnov & Fedoseeva, 1976; Nartshuk, 1998. Misidentifications. Frons wider than long. Ocellar triangle short, extending to middle of frons and then continues to front margin as a line. Ocellar triangle shining, centrally yellow and with black spots on ocellar tubercle, in front of ocelli, on side margins and on apex. Surface of triangle between black spots whitishyellow and sulcate medially and laterally. Genae as wide as first flagellomere. First flagellomere yellow, darkened at base of arista, arista black. Palpi yellow. Scutum with 5 black densely grey-dusted stripes covered with yellowish setae anteriorly and black setulae posteriorly. Scutellum yellow with 6 setae. Spot on katepisternum yellow, spot on anepisternum black. Postnotum black, dusted. Body length 3.5–4.0 mm Distribution. – Sweden: LU: lake NW of upper course of river Vietasjokk, 14.vii.1961 (Dahm, Landin & Löfqvist), 1 male. More common in Finland: Al, Ab, N, St, Ta, Sa, Oa, Om. Russia: Vib; Kr: Paanajärvi. Not found in Denmark and Norway. Widely distributed in Asian part of the Palearctic eastwards to Japan and in East Europe, but not recorded in West Europe except for Sweden and Finland. An eastern Eurasian species. Biology. – Adults in July to August in meadows. Note. – The species was described from Japan (Kanmiya, 1978), then synonymized with C. geminatus by Kanmiya (1983) but subsequently restored from synonymy by Nartshuk (2002a).

134. Chlorops laetus Meigen, 1830 Fig. 628 Chlorops laetus Meigen, 1830: 143; Meigen’s plate 211: 15 (Morge, 1976b). Type specimens not found in Coll. Winthen in NHMW, probably destroyed. Frons wider than long. Ocellar triangle narrow, extending to middle of frons followed by linear extension to front margin of frons. Ocellar triangle with yellow windows and black spots in front of ocelli, on side margins and on apex. Genae 1.5 times as wide as first flagellomere. Basal segments of antennae fuscous yellow. First flagellomere basally yellow, black above. Arista black. Palpi yellow. Scutum 172

with 5 black densely grey-dusted stripes. Scutellum yellow. Spot on katepisternum yellow, anepisternum with some black setulae. Abdomen yellow with narrow black bands on tergites and black median line. Body length 3.5–4.0 mm. Distribution. – Not common. Sweden: SM, ÖL, ÖG, SÖ, UP. Norway: AK, HOy. Finland: Sa. Russia: Vib, Kr. Not found in Denmark. The species is known from Europe and Kazakhstan. Biology. – Autumnal species, adults in August to September.

135. Chlorops limbatus Meigen, 1830 Figs 629–632, 991–993 Chlorops limbatus Meigen, 1830: 150; Meigen’s plates 211: 13; 213: 11 (Morge, 1976b). Lectotype in NHMW (Nartshuk, 1997a). Chlorops discicornis Zetterstedt, 1848: 2625, non Loew, 1866: 66 (Nartshuk & Andersson, 2002). Chlorops fulviceps Roser, 1840: 62 (Nartshuk, 1994). Chlorops brevimanus Loew, 1866 (Nartshuk, 1997b). Frons wider than long. Ocellar triangle extending half length of frons, black smooth with little raised side margins. Genae narrow, about 2/3 width of first flagellomere. First flagellomere black, rather large, slightly broader than long. Arista black. Palpi rather stout yellow. Scutum with 5 black densely grey-dusted rather wide stripes. Scutellum yellow. Spot on katepisternum black. Abdomen yellow with black bands on tergites or sometimes blackish dorsally. Legs yellow with medially black hind tibiae and darkened tarsi. Front basitarsus of male short and claws of front legs asymmetrical. Male genitalia with sclerotized aedeagus. Body length 2.5–3.0 mm. Distribution. – Rather common. Denmark: from all districts except WJ and LFM. Sweden: northwards to UP and VR, and further north in NB. Norway: MRy, NTi. Finland: northwards to ObN. Russia: Vib, Kr. Trans-Palearctic species, known from the British Isles to Japan. Biology. – Adults in May to September, in wet meadows and on banks of rivers and lakes. Two generations per year.

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, dorsal view. 630: head, lateral view. 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.

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Larvae phytophagous, develop in shoots of Phalaris arundinacea, larvae of the summer generation on exposed panicles. The leaves above the stem-feeding larvae dry out and turn brown. Larvae hibernate in basal and aerial shoots and pupate the following spring (Nartshuk, 1962a; Sharipov, 1981). Records of other host grasses belonging to Alopecurus and Phleum pratense (Wetzel, 1967; Tschirnhaus, 1981) need verification. Larvae described by Balachowsky & Mesnil (1935), Oettingen (1935) and Nye (1958).

136. Chlorops meigenii Loew, 1860 Figs 633–638 Chlorops meigenii Loew, 1860: 43. The name is included in Official List of Names in Zoology, Opinion 2040 (Case 3190), 2003: 116. Ocellar triangle predominantly black, shining, reaching front margin of frons and yellowish at apex. Genae as wide as first flagellomere or a little wider. Basal antennal segments fuscous yellow. First flagellomere small rounded, mainly dark brown or yellow with darkening dorsally. Arista black. Palpi small yellow. Scutum with 5 black, shining stripes, central one with indistinct outline posteriorly. Inner lateral stripes without projection on outer margin. Stripes nearly fused in some dark specimens collected in northern Fennoscandia. Spot on katepisternum reddish brown or blackish dorsally. Abdomen yellow with black bands. Wings brownish. Legs mainly yellow, but femora and hind tibiae partly black, last tarsal segments darkened. Body length 3.0–3.5 mm. Distribution. – One of the most common species of the genus. Denmark: SJ, F, LFM, NWZ, NEZ. Sweden: northwards to TO. Norway: Ø, AK, HEs, AAy, HOy, HOi, SFy, SFi, Nsy. Finland: northwards to Li. Russia: Vib, Kr, Lr. Trans-Palearctic species ranging from Europe to Kamchatka. Biology. – Adults in May to August, in wet meadows and forest clearings. Adults were found in a swallow’s nest (Hicks, 1959). Probable host plants are some species of Calamagrostis growing in wet places, other than C. epigeios.

137. Chlorops nigripalpis (Duda, 1933) Figs 639–642 174

Oscinis nigripalpis Duda, 1933: 186. Holotype in NHMH, currently on loan in ZMLU. Chlorops crassipalpis Smirnov, 1958: 1166. Holotype in ZMMU (Nartshuk & Andersson, 2002). Frons longer than wide. Ocellar triangle extending 2/3 length of frons followed by linear extension to front margin of frons, predominantly yellow with median black spot, occupying ocellar tubercle and narrowing to apex. Genae 1.5 times as wide as first flagellomere. Basal antennal segments fuscous yellow. First flagellomere and arista black. Palpi yellow basally, black in apical half. Scutum with 5 black, dusted stripes not extended to scutellum. Scutellum yellow. Spot on katepisternum black, anepisternum with small black spot. Abdomen yellow with black bands on tergites. Legs mainly yellow, last tarsal segments darkened. Body length 3.0 mm. Distribution. – Rare species. Sweden: NB: Högsön, Råneå, 31.vii.1978 (H. Andersson), 1 female; Jupukka, Rajala, 28.vii.1955 (H. Andersson), 5 males. Finland: N: Helsinki (Ahnger), 1 female. Russia: Kr: female holotype of Oscinis nigripalpis. The species is known from Europe and Kazakhstan.

138. Chlorops obscurellus (Zetterstedt, 1838) Figs 644–646 Oscinis obscurellus Zetterstedt, 1838: 779. Holotype in ZMLU (Andersson, 1966). Frons as long as wide. Ocellar triangle large, behind touching eyes, extending nearly to front margin of frons, black, except hind corners, smooth and shining, wrinkled on apex. Genae a little wider than first flagellomere. Basal antennal segments brown, first flagellomere and arista black. Palpi yellow. Scutum with 5 black, dusted stripes, three central ones very broad and extended to scutellum. Scutellum yellow, basally black. Spot on katepisternum black, shining. Abdomen brown to black dorsally with narrow yellow side margins. Legs yellow, femora and hind tibiae partly black, last tarsal segments darkened. Body length 2.0–3.0 mm. Distribution. – Denmark: SJ, LFM, NEZ. Sweden: northwards to NB. Finland: Ta, Oa, Kb, Om. Russia: Vib. Not found in Norway. Eurasian species.

Figs 639–652. Chlorops nigripalpis (Duda): 639: head, dorsal view. 640: head, lateral view. 641: hypopygium, ventral view. 642: hypandrium 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.

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Biology. – Adults in June to August, in wet meadows and swamps. Larvae phytophagous, develop in shoots of Carex riparia without causing external changes (De Bruyn, 1989).

139. Chlorops pallidiventris (Duda, 1933) Fig. 647 Oscinis pallidiventris Duda, 1933: 164. Syntype, 1 female, in ZMHU (Wendt, 1975). Frons as long as wide or a little longer. Ocellar triangle short, extending nearly 1/3 length of frons followed by linear extension to front margin of frons. Black spot on ocellar triangle occupies central part; side margins narrowly and hind corners widely yellow. Genae narrower than first flagellomere. Antennae yellow, first flagellomere black above, arista black. Palpi yellow. Scutum with 5 black, dusted rather narrow stripes not extended to scutellum. Setulae of scutum yellow. Scutellum yellow, basally blackish. Spot on katepisternum black, shining. Anepisternum with some setulae. Abdomen whitish-yellow covered with yellow setulae. Legs yellow. Basitarsus of front legs in male short, only 2 times as long as broad and claws of front legs asymmetrical. Body length 2.0–2.5 mm.

Abdomen yellow with narrow black bands on tergites. Legs yellow, last tarsal segments darkened. Body length 3.0–3.5 mm. Distribution. – Rare. Only from Sweden: VS: Skäret, Valskog, 11.vii.1989 (H. Andersson), 1 female; VR: Storfors, 28.vi.1988 (H. Andersson), 2 females. European species.

141. Chlorops palpatus Smirnov, 1958 Chlorops palpatus Smirnov, 1958: 1169, 1174. Syntypes probably in ZMMU. Frons longer than wide. Ocellar triangle extending 2/3–3/4 length of frons followed by linear extension to front margin of frons, shallow in the middle. Black spot on ocellar triangle occupies central part, side margins and hind corners yellow. Basal segments of antennae yellow, first flagellomere black, arista black. Palpi rather long and broad, brownish to black. Scutum with 5 black, shining stripes, central stripes not extended to scutellum. Scutellum yellow. Spot on katepisternum shining, reddish yellow, with brown stripe above. Abdomen yellow, shining, covered with whitish setulae. Legs yellow. Body length 2.0–3.0 mm.

Distribution. – Rare. Only from Sweden: ÖL: Halltorps Hage, 19–23.v.1976 (H. Andersson & R. Danielsson), 1 male. European species, known elsewhere only from Hungary and Bulgaria.

Distribution. – Rare. Sweden: HA: Årnilt, Enslöv, 04.viii.1972 (H. Andersson), 2 males, 1 female; UP: (Boheman), 1 specimen. Norway: VE: Sandefjord (Siebke). Eurasian species, known from European part of Russia.

140. Chlorops pallifrons Strobl, 1909

142. Chlorops pannonicus Strobl, 1893 Figs 648–652

Chlorops pallifrons Strobl, 1909: 287. Syntypes 2 males, 4 females in NMBA (Chvála, 2008). Frons wider than long. Ocellar triangle a little longer than half length of frons, yellow, sulcate along side margins, continuing as a yellow line to front margin of frons. Ocellar tubercle black. Genae 1.5 times wider than first flagellomere. Basal antennal segments yellow, first flagellomere and arista black. Palpi yellow. Scutum with 5 black, dusted stripes, central one extended to scutellum. Scutellum yellow. Spot on katepisternum black. 176

Chlorops pannonica Strobl, 1893: 228. Syntypes 5 males, 5 females in NMBA (Chvála, 2008). Chlorops balticus Wahlgren, 1913: 51. Holotype in ZMLU (Duda, 1933; Nartshuk & Andersson, 2002). Frons longer than wide. Ocellar triangle extending nearly to front margin of frons with median sulcus and often wrinkled along side margins. Black spot on ocellar triangle occupies nearly all surface except hind corners. Genae a little wider than

first flagellomere. Antennae yellow, first flagellomere black above, a little longer than wide. Arista whitish. Palpi small yellow. Scutum with 5 black, dusted stripes, outer lateral stripes connected with inner lateral ones in front and fused with them behind. Scutellum yellow, basally at sides blackish. Spot on katepisternum from yellow to black, covered with thin dust and subshining. Abdomen brown or blackish. Legs yellow, femora partly black and last tarsal segments darkened. Body length 2.0–2.5 mm. Distribution. – Rare. Only from Sweden: ÖL: female holotype of Chlorops balticus and another female from July 1916 also collected by Wahlgren; GO: one female from collection of Boheman (NRMS). Holotype of Chlorops balticus was collected on the alvar (Wahlgren, 1913). Xerophilous species widely distributed in the Palearctic steppe zone from Hungary to the Far East, with local extensions northwards in dry, sandy habitats. Finding places nearest to Gotland and Öland in Estonia: Islands of Kõinastu and Saaremaa (Nartshuk, 1998). Note. – Duda (1933: 189) considered Chlorops balticus Wahlgren as a synonym of C. pannonicus. Nartshuk (1984b) listed the species as a separate species. Now, after examination of holotype of Chlorops balticus, we accept this synonymy.

143. Chlorops planifrons Loew, 1866

on katepisternum black or yellow, but intermediate colour also occurs: yellow with black above. Abdomen yellow with narrow black bands on tergites and often with median black line. Legs yellow, last tarsal segments darkened. Cerci of female narrow and long, in dry specimens divergent. Body length 2.5–3.0 mm. The light coloured form was described as C. triangularis Becker, 1910: 97, which Duda (1933) considered as variety of C. planifrons. Distribution. – Very common and widespread. Denmark: SJ, EJ, NEJ, NEZ. Sweden: northwards to TO. Norway: HEn. Finland: north to Li. Russia: Vib, Kr, Lr. Trans-Palearctic, widely distributed polyzonal species, known from the British Isles to the Far East of Russia, in Europe southwards to Spain. Biology. – Adults from middle June to middle August, on fens and marshes with large tussocks of Carex. One generation per year. Larvae phytophagous, develop in shoots of Carex, e.g. C. inflata, C. rostrata (Nartshuk, 1962a; Zhabinskaya, 1963). Larvae of the first instar hibernate in shoots of its host plant and continue feeding the following spring. Larvae of the summer generation make a furrow on stem that begins immediately below inflorescence and gradually widens downwards. At some point is an interruption of the furrow 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 feeding on another stem (Nartshuk, 1962a). Larvae described by Zhabinskaya (1963).

Figs 653–657, 995 Chlorops planifrons Loew, 1866: 55. Syntypes in ZMHU (Wendt, 1975). Chlorops triangularis Becker, 1910: 97, as var. of C. planifrons Loew. Syntypes in ZMHU (Wendt, 1975). Body yellow or sometimes in specimens collected in autumn reddish. Frons longer than wide. Ocellar triangle large, extending 4/5 length of frons, smooth shining. Black spot on ocellar triangle rather variable: occupies all triangle or divided into two spots, one on ocellar tubercle and another rhomboid in form at the apex of triangle. Genae a little wider than first flagellomere. Basal segments of antennae yellow, first flagellomere rather large black. Arista slightly whitish. Palpi yellow. Scutum with 5 black, dusted stripes. Scutellum yellow. Spot

144. Chlorops pumilionis (Bjerkander, 1778) Figs 104, 658–662, 996–998 Musca pumilionis Bjerkander, 1778: 240. Types not found; see under Meromyza saltatrix. Chlorops pumilionis gotlandica Holmgren, 1887: 75. Types not found (Nartshuk & Andersson, 2002). Body yellow or sometimes in specimens collected in autumn reddish. Frons as long as wide. Ocellar triangle extending slightly more than half length of frons and continues as line to front margin of frons. Its surface black or brown, smooth shining with median sulcus before front ocellus and median ridges in apical part. In light coloured specimens 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.

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black. Genae wider than first flagellomere. Basal segments of antennae brown or black, first flagellomere and arista black. Palpi yellow. Scutum with 5 wide black, dusted stripes. Scutellum yellow. Spot on katepisternum black, dusted. Abdomen yellow with black bands on tergites or blackish. Legs mainly yellow, apical part of tibiae and basitarsus or all tarsus of front legs black, last tarsal segments of other legs darkened. Body length 2.5– 3.5 mm.

yellow windows. Genae wider than small first flagellomere. Basal segments of antennae yellow or brownish, first flagellomere black or brownish below. Arista black. Palpi long yellow and black apically in females. Scutum with 5 black thinly dusted stripes. Scutellum yellow or rarely with black spot. Spot on katepisternum yellow. Abdomen yellow with narrow black bands on tergites. Legs yellow, hind tibiae with black band. Two last tarsal segments darkened. Body length 2.5–3.5 mm.

Distribution. – Denmark: from all districts. Sweden: north to ÅN. Norway: Ø, AK, HEs, Os, Bv, Bø, VAi, HOi. Finland: north to Om. Russia: Vib, Kr. Eurasian temperate species, widespread from Europe to Iran, Israel, West Siberia and Mongolia.

Distribution. – Denmark: NEZ, B. Sweden: north to DR. Norway: AK. Finland: Ab, N, Ka, Sa. Russia: Vib, Kr. Eurasian temperate species, known from southern Sweden and Finland southwards to Bulgaria and eastwards to Yakutia, East Siberia.

Biology. – Adults from end of May to early September, in two generations. Adults of the second generation were found in a swallow’s nest (Krivokhatsky & 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 of cereals causing thickening of the attacked shoots (“gout swelling”), while larvae of the summer generation feed on the exposed, developing ear and stem of cereals (Porchinsky, 1881). Plant reactions to the larval attacks were studied specially by Goidanich (1956). Many species of wild grasses have been recorded as possible host plants of the Goats fly by different authors, but most likely only Elytrigia repens and Aegylops sp. are genuine host plants. Injuries on cereals caused by Chlorops pumilionis have been reported from several Nordic countries (Lampa, 1888; Johansson, 1960; Rygg, 1966a), but the species appears to be of little economic importance. Larval descriptions are given by many authors, notably by Frew (1923a), Balachowsky & Mesnil (1935), Nye (1958) and Zhabinskaya (1963).

145. Chlorops ringens Loew, 1866 Figs 663, 664 Chlorops ringens Loew, 1866: 46, 60. Syntypes in ZMHU (Wendt, 1975). Frons as wide as long. Ocellar triangle extending 2/3 length of frons followed by linear extension to front margin of frons. Black spots on ocellar triangle rather small triangular and usually with

Biology. – Adults in meadows. Autumnal species occurring in August and September.

146. Chlorops riparius Smirnov, 1958 Fig. 107 Chlorops riparius Smirnov, 1958: 1167, 1174, as subspecies of C. nigripalpis (Duda). Syntypes probably in ZMMU. Frons wider as long. Ocellar triangle extending nearly half length of frons or little longer followed by linear extension to front margin of frons, with sulcus from front ocellus to apex. Black spots on ocellar triangle rather narrow, side margins of triangle widely yellow. Genae wider than small first flagellomere. Basal segments of antennae yellow or brownish, first flagellomere black or brownish below. Arista black, distinctly pubescent. Palpi yellow short and rather stout. Scutum with 5 black densely dusted stripes, central stripe not extending to scutellum. Scutellum yellow. Spot on katepisternum black and dusted. Abdomen yellow or with very narrow black bands on tergites. Legs yellow, two last tarsal segments darkened. Body length 3.0–4.0 mm. Distribution. – Sweden: VG, UP. Norway: SFi: Vassbygda, Aurland, 16.viii.1941 (N. Knaben); Loen, Stryn, 02.viii.1942 (N. Knaben), 1 male. Not found in Denmark, Finland or Russian parts of Fennoscandia. Eurasian species. 179

Biology. – Species occurring sporadically in August, near small streams in forest, probably one generation per year. Larvae phytophagous, live in shoots of Elymus caninus, hibernate as larvae of first or second instar. The mature larvae produce following spring a gall-like swelling near the ground on young shoots (Nartshuk, 1962a). Some inquilines live within the galls: Elachiptera cornuta and Asteia concinna Meigen (Asteiidae) (Nartshuk, 1962a).

147. Chlorops rossicus Smirnov, 1955 Fig. 665 Chlorops rossicus Smirnov, 1955: 334. Holotype probably in ZMMU. Frons usually wider than long with black upright setulae. Ocellar triangle black, extending half length of frons or little longer followed by linear extension to front margin of frons. Black spot on ocellar triangle occupies nearly all triangle except narrow yellow lines along side margins and hind angles, black middle line goes from front ocellus to apex of triangle. Genae with some black setulae, wider than first flagellomere. Basal segments of antennae yellow or brownish, first flagellomere and arista black. Palpi yellow. Scutum with 5 wide black, shining stripes and often blackish before scutellum. Inner lateral stripes widened in front and outer margin with triangular incision opposite notopleura. Scutellum yellow. Spot on katepisternum black, shining. Wings brownish, costal vein extending beyond tip of R4+5 . Abdomen blackish dorsally. Legs yellow, or with black stripes on hind tibiae. Two last tarsal segments darkened. Body length 2.5–3.5 mm.

148. Chlorops rufijinus (Zetterstedt, 1848) Figs 666–668 Oscinis rufijinus Zetterstedt, 1848: 2628. Holotype in ZMLU (Andersson, 1966). Oscinis citrinellus Zetterstedt, 1848: 2630 (Nartshuk & Andersson, 2002). Oscinis bipunctus Duda, 1933: 173. Oscinis bipunctus hungaricus Duda, 1933: 173. Oscinis bipunctus fennicus Duda, 1933: 173. Note. – Types of hungaricus not found in ZMHU (information of A. Soós in Zuska, 1960). Zuska (1960) designated a lectotype, 1 female, of bipunctus in NHMH (N 8159). Species very variable in colour of scutal stripes, spot on katepisternum and first flagellomere. Frons as wide as long or a little wider. Ocellar triangle extending 2/3–3/4 length of frons and continues as line to front margin of frons, yellow with two black spots: one on ocellar tubercle and another rhomboid spot on apex, both spots connected by black line, side margins of triangle narrowly black as well. Genae as wide as first flagellomere. Antennae yellow or first flagellomere darkened above. Arista whitish with yellow basal part. Palpi yellow. Scutum with 5 red, partly black or entirely black, thinly dusted stripes. Scutellum yellow. Spot on katepisternum from yellow to black, dusted. Abdomen yellow with narrow black bands on tergites. Legs yellow, last tarsal segments darkened. Body length 2.5–3.0 mm. Distribution. – Sweden: northwards to NB. Norway: AK, HOi. Finland: Al, Ta. Russia: Vib, Kr. Not recorded from Denmark. Widespread trans-Palearctic species, known southwards to Bulgaria and eastwards to Mongolia and Japan, but everywhere rare.

Distribution. – Denmark: NEJ: Danzigmand, Læsø, 12.vii.1969 (ZMUC-Exp.), 1 female; NEZ: Lyngby Mose, 03.vii.1964 (O. Martin), 1 male, 2 females, 21.vi.1972 (K. Schnack), 1 female. Sweden: 1 female. Sweden: SK: Revingehed, 06.vi.1993 (R. Danielsson), 1 female; SÖ: Trehörningen, Huddinge, 21.vi.1960 and 03.vii.1965 (L. Hedström), 2 females in ZMUU. Norway: SFy, STy, TRy. More common in Finland: N, Ka, St, Ta, Sb, Li. Russia: Vib, Lr. European species.

Biology. – Adults in July to August in meadows.

Biology. – Adults in June and July, in meadows and edges and clearings in forests.

Frons wider than long. Ocellar triangle extending 2/3 length of frons followed by linear exten-

180

149. Chlorops scalaris Meigen, 1830 Figs 669–672 Chlorops scalaris Meigen, 1830: 145; Meigen’s plate 159: 11 (Morge, 1976a). Lectotype and some paralectotypes in NHMW (Nartshuk, 1997a), other paralectotypes in MNHN.

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 median black spot; side margins, apex of triangle and windows near ocelli yellow. Genae wider than first flagellomere. Basal segments of antennae yellow, small first flagellomere yellow darkened above, rarely blackish. Arista black. Palpi long yellow. Scutum with 5 black thinly dusted stripes. Scutellum yellow. Spot on katepisternum black, shining. Abdomen yellow with narrow black bands on tergites. Legs yellow, last tarsal segments darkened. Body length 2.5–3.5 mm.

Distribution. – Denmark: common and widespread. Sweden: northwards to LU and NB. Norway: Ø, AK, HEs, HEn, Bø, VE, VAy, Ry, HOi, SFi, TRy, Fv, Fn. Finland: northwards to ObS. Russia: Vib, Kr. Eurasian polyzonal species, ranging southwards to Italy and Bulgaria and eastwards to East Siberia and Mongolia. Biology. – Adults in June and July, rather common everywhere in meadows. Larvae phytophagous, host plants Holcus lanatus, Poa pratensis (Tschirnhaus, 1981, 1992) and probably other grasses. 181

150. Chlorops scutellaris (Zetterstedt, 1838) Figs 673–676 Oscinis scutellaris Zetterstedt, 1838: 779. Lectotype in ZMLU (Andersson, 1966). Chlorops laevicollis Becker, 1910: 89. Holotype in ZISP (Nartshuk, 1998). Oscinis freyi Duda, 1933: 199. Lectotype in NHMH (Frey, 1944; Nartshuk, 1998). Rather dark species. Frons brown, wider than long. Ocellar triangle nearly reaching front margin of frons, entirely black, shining. Genae narrower than first flagellomere. Antennae entirely black, including arista. Palpi black. Scutum entirely black, shining. Scutellum and pleura darkened. Spot on katepisternum black, shining. Abdomen brown to black. Legs mainly black. Body length 2.0–2.5 mm. Distribution. – Sweden northwards to LY. Norway: Fn: Tana Bru. Finland: north to Li. Russia: Vib, Lr. Not found in Denmark. Northern Eurasian species, more likely of Siberian origin, known from North Europe eastwards to East Siberia. Biology. – Adults in July in wet meadows and fens.

151. Chlorops serenus Loew, 1866 Figs 677, 678 Chlorops serenus Loew, 1866: 62. Syntypes in ZMHU (Wendt, 1975). Frons wider than long. Ocellar triangle extending nearly to front margin of frons, yellow with small median black spot, side margins, apex of triangle and two windows near ocelli yellow. Genae distinctly wider than first flagellomere. Basal segments of antennae yellow, first flagellomere black. Arista black. Palpi yellow. Scutum with 5 black, dusted stripes, covered with yellow setulae. Scutellum yellow. Spot on katepisternum yellow, rarely blackish above, shining. Abdomen yellow with narrow black bands on tergites and median black line. Legs yellow, last tarsal segments darkened. Body length 3.0–3.5 mm. Distribution. – Scarce. Denmark: SJ, F, LFM, NEZ, B. Southern part of Sweden: SK, HA, and also UP. Norway: AK, AAy. Finland: Al: Kökar, 11.vi.1978 182

(M. Koponen), 1 female; Apalholm, Lemland, 12– 24.viii.1954 (H. Lindberg), 1 male, 1 female. Not found in Russian parts of Fennoscandia. Euro-Mediterranean species, known from middle and southern Europe, Israel and Algeria. Biology. – Adults in meadows in July and again in August to September, likely in two generations.

152. Chlorops speciosus Meigen, 1830 Figs 679–683, 1002–1004 Chlorops speciosus Meigen, 1830: 146; Meigen’s plate 159: 1 (Morge, 1976a). Lectotype in NHMW (Nartshuk, 1997a). Frons wider than long. Ocellar triangle extending 2/3–3/4 length of frons, black or brown smooth shining with very distinct borders. Genae distinctly wider than first flagellomere with many black setulae. Basal segments of antennae dark, first flagellomere black. Arista black. Palpi small yellow. Scutum with 3 broad black, shining stripes (in dark specimens from northern Fennoscandia stripes fused), covered with black setulae. Scutellum yellow or darkened. Spot on katepisternum black, shining. Abdomen yellow with black bands on tergites or blackish. Legs partly black. Body length 3.5–4.5 mm, some females up to 6.0 mm. Distribution. – The most common species of Chlorops occurring throughout Denmark and Fennoscandia. Eurasian species, found commonly in Europe southwards to Spain and Bulgaria, ranging eastwards to Altai, West Siberia. Biology. – Adults in June to August in wet meadows. Larvae phytophagous, live in stems of Deschampsia caespitosa and hibernate basally in the stem of their host plant. Larvae described by Balachowsky & Mesnil (1935), Nye, (1958) and Zhabinskaya (1963), descriptions of different authors differ in some details.

153. Chlorops strigulus (Fabricius, 1794) Figs 684–690, 986, 987

Figs 673–683. Chlorops scutellaris (Zetterstedt): 673: head, dorsal view. 674: head, lateral view. 675: hypopygium, dorsal view. 676: hypandrium 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 (Zimsen, 1964). Frons wider than long. Ocellar triangle reaching front margin of frons, black dull and sulcate along side margins and on apex. Genae distinctly wider than first flagellomere, yellow with black lower margin. Basal segments of antennae brown, first flagellomere longer than wide and black. Arista thick white pubescent with basal segment dark. Palpi black. Scutum with 5 black, dusted stripes, of which 3 central stripes are broad and sometimes fused before scutellum. Scutellum darkened. Anepisternum with many black setulae. Spot on katepisternum black, dusted. Abdomen dorsally brown to black with narrow yellow bands on tergites. Legs mainly black. Body length 2.5–3.5 mm.

Distribution. – Rare. Denmark: EJ, SZ, NEZ, B. Sweden: UP: Hässelby, Börje, 09.v.1959 (L. Hedström), 1 male. Not found in Norway, Finland or Russian parts of Fennoscandia. Eurasian species. Biology. – Adults from end of May to June, in dry meadows. Phytophagous larvae produce top galls on shoots of Brachypodium sylvaticum and B. retusum (Séguy, 1934; Balachowsky & Mesnil, 1935; Nye, 1958). Baudyš (1912) described galls on Elytrigia repens that in his opinion were produced by C. strigulus, but probably the gall former was C. novakii Strobl, a species not found in the Nordic countries. Larvae described by Rübsaamen (1895), Balachowsky & 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 (Zetterstedt, 1848) Fig. 691 Oscinis troglodytes Zetterstedt, 1848: 2626. Lectotype in ZMLU (Andersson, 1966). Frons wider than long. Ocellar triangle extending half length of frons, entirely black, shining. Genae wider than first flagellomere. Basal segments of antennae brown, first flagellomere and arista black. Palpi small yellow. Scutum with 5 black thinly dusted and slightly shining stripes. Scutellum yellow. Spot on katepisternum black, shining. Abdomen dorsally brown to black with narrow yellow bands on tergites. Cerci of ovipositor black. Legs yellow with darkened apical tarsomeres. Body length 2.0–2.5 mm. Distribution. – Rather common. Denmark: WJ, NWJ, LFM. Sweden: northwards to NB. Norway: 184

northwards to NTi. Finland: northwards to Ks. Russia: Vib, Kr, Lr. Eurasian polyzonal species, known from Great Britain eastwards to Yakutia, East Siberia and Mongolia. Biology. – Adults in June to August in meadows.

155. Chlorops varsoviensis Becker, 1910 Figs 692–695 Chlorops varsoviensis Becker, 1910: 92. Syntype, 1 male, in ZMHU (Wendt, 1975). Ocellar triangle medially with a large poplar leafshaped black spot. Genae narrower than width of first flagellomere. Antennae usually black or first flagellomere reddish ventrobasally. Palpi blackish. Scutum with 5 black stripes, sparsely grey-dusted.

Pleura with black spots. Anepisternum with setulae, white in males and black in females. Abdomen reddish-brown in females. Legs almost yellow with apical tarsomeres darkened. In male front basitarsus short, about 2.0–2.5 times as long as wide and front inner claw larger and more curved than outer one. Body length 3.0–3.5 mm. Distribution. – Sweden: northwards to TO. Norway: Fi: Gargia, Alta. Finland: northwards to Li. Russia: Vib, Kr. Lr. Not so far recorded from Denmark. Trans-Palearctic species, known from France to Japan, southwards to Spain, North Italy and Bulgaria, eastwards to Mongolia. Biology. – Adults in July and August in wet habitats with Carex. Larvae phytophagous develop in stems of Carex.

Yellow species with black stripes on scutum and black spots on pleura. Head broader than thorax. Eyes large, genae linear and postgenae narrow. Ocellar triangle smooth shining with convex sides and reaching front margin of frons, usually with a row of setulae on its surface along side margins. First flagellomere orbicular or slightly longer than wide. Scutum shining or rarely dusted. There are band-like sclerites along anus. Male genitalia: mesolobus small; surstyli closely alligned to epandrium; pre- and postgonites situated in line. Tibial organ present. Distribution. – Probably worldwide.

157. Chloropsina distinguenda (Frey, 1909) Figs 696–699

156. Chlorops zernyi (Duda, 1933) Oscinis zernyi Duda, 1933: 201. Holotype female in NHMW. Body yellow. Frons as long as wide, setae and setulae black. Ocellar triangle large, reaching front margin of frons, but narrow in distal third, smooth shining, mainly black, but yellow along sides, hind angles and on tip, wrinkled on hind angles. Genae a little wider than first flagellomere. Parafacialia as wide as gena. Basal segments of antennae brown, first flagellomere black. Arista white, thickly pubescent. Palpi large, blackish on tip. Scutum with 5 black, dusted stripes, covered with yellow setulae, not extended to scutellum. Scutellum yellow. Spot on katepisternum yellow-brownish, shining. Abdomen yellow with narrow black bands on tergites, covered with black setulae. Legs yellow. Cerci of female yellow. Body length 4.0–4.5 mm. Distribution. – Russia: Vib: Beloostrov, 15.viii.1956 (Nartshuk), 2 females. Eurasian species. Biology. – Usually in Carex beds.

Chlorops distinguendus Frey, 1909: 11. Holotype in NHMH. Head wider than thorax. Frons narrowed in front. Ocellar triangle large, smooth black, shining, reaching front margin of frons. vte setae on yellow spots. Genae linear, not wider than palpus. Antennae including arista and palpi black. Scutum nearly square, black, shining, scutal stripes nearly fused. Scutellum yellow. Pleura with several black, shining marks. Abdomen black dorsally and yellow ventrally. Legs mainly yellow with darkened femora and tarsi. Body length 2.3 mm. Distribution. – Very rare. Sweden SK: Hässleholm, 21.vii.1981 (H. Andersson). Finland: Al: Tinström, Sund (R. Frey), female holotype; Ab: Jurmo, Korpo, 04.viii.1960 (Meinander); N: Linna, Helsinki (R. Frey), 1 female; Oa: Berga, 17.vi.1946 (Hellén). Elsewhere only recorded from the Czech Republic.

158. Chloropsina rohaceki Nartshuk, 2000 Figs 700–702 Chloropsina rohaceki Nartshuk, 2000a: 256. Holotype in ZMOC, paratypes in ZISP and ZMLU.

Genus Chloropsina Becker, 1911 Chloropsina Becker, 1911: 51. Type species: Chloropsina oculata Becker, 1911 by designation of Malloch, 1931: 76.

Head wider than thorax. Frons longer than wide. Ocellar triangle large, smooth black, shining, reaching front margin of frons. vte setae on yellow spots. Genae narrower than palpus. First flagellomere yellow, 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, median stripe reaching scutellum as black spot with indistinct margins. Scutellum yellow with black side margins. Pleura with several black, shining marks. Abdomen black dorsally and yellow ventrally. Legs entirely yellow. Body length 2 mm. Distribution. – Only Sweden: BL: Kristianopel, 08.vii.1987 (R. Danielsson). European species, known elsewhere from Slovakia.

Genus Cryptonevra Lioy, 1864 Cryptonevra Lioy, 1864: 1125. Type species: Chlorops flavitarsis Meigen, 1830 by monotypy. 186

Black species, only frons, antennae and legs partly yellow. Head setae well developed, vte longer than vti. Ocellar triangle large, broad behind, almost reaching front margin of frons, with reticulate microsculpture, not dusted. Face with carina, triangular between antennae and narrow below. First flagellomere about as long as wide with pointed upper corner. Scutum black with faint granulate microsculpture, dusted and evenly covered with numerous setulae. Femoral and tibial organs present. Male genitalia: middle lobe of surstylus granulate. No sclerites lateral to anus. Postgonites short, connected with long pregonites. Distiphallus long complex with coronated sclerites. Contrary to Neohaplegis testes in male tubular as usual in Chloropinae, not round. Distribution. – Palearctic and Oriental Regions.

Key to species of Cryptonevra Lioy 1 – 2

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Frons yellow in front. Male genitalia Fig. 711 . . . . . . . . . . . . . . . . . . . . . C. diadema (Meigen) Frons entirely black . . . . . . . . . . . . . . . . . . . . . 2 Setulae on head, scutum, abdomen and legs black. Scutum shining. Front tibiae black. Male genitalia Figs 709, 710 . . . . . . . . . . . . . . . . . . . . C. flavitarsis (Meigen) Setulae on head, scutum, abdomen and legs pale. Scutum dull. Front tibiae pale. Male genitalia Figs 703, 704 . . . . . C. consimilis (Collin)

Species key to known 3rd instar larvae of Cryptonevra Mature larvae white cylindrical 3–5 mm long. Facial mask with linear branched ridges. Hypostomal and tentoropharyngeal sclerites separate. Mandibles with small accessory teeth in two rows. Body with numerous lines of spicules of different size on locomotory creeping welts on ventral side of body. Anal opening surrounded by spicules. Anterior spiracles of fan type with 4–9 buds. Posterior spiracles on stigmophores and with branched interspiracular processes. 1

–

Anterior lines of spicules on ventral locomotory creeping welts of abdominal segments continuous (Figs 721, 722). Anterior spiracles with 6–8 buds (Fig. 720) . . . . . . . . . . . . . . . . . . . . . C. diadema (Meigen) Anterior lines of spicules on ventral locomotory creeping welts of abdominal segments not continuous, separated laterally (Figs 715, 716). Anterior spiracles with 4–9 buds (Fig. 713) . . . . . . . . . . . . . . . . . C. flavitarsis (Meigen)

159. Cryptonevra consimilis (Collin, 1932) Figs 703, 704 Haplegis consimilis Collin, 1932: 113. Lectotype male and 1 male, 1 female paralectotypes in OXUM, 3 males, 2 females paralectotypes in BMNH, all bread from reed stem inhabited by Lipara similis (Ismay, 1994a; Pont, 1995).

Similar to C. flavitarsis, but differs in having paler front tibiae and all short pubescence of head, thorax, abdomen and legs pale. Only vte setae on head; thoracic and scutellar setae black. Frons slightly longer than wide, black (rarely with reddish-brown anterior margin). Ocellar triangle black, nearly reaching front margin of frons. Its surface minutely punctuate and subshining. Basal segments of antennae black or rarely reddish brown. First flagellomere slightly longer than wide, distinctly angulated dorsodistally, black. Arista black and minutely pubescent basally, distally glabrous. Scutum black, minutely shagreened, dusted on notopleura and before scutellum, evenly clothed with pale setulae. Scutellum grey-dusted. Pleura mainly shining. Wings slightly whitish. Veins R4+5 and M divergent distally. Abdomen basally brownish yellow, remaining part black grey-dusted and covered with white setulae. Femora black except for yellow knees, tibiae and tarsi yellow. Male with tarsal segment 4 of middle legs with black setae on ventral surface. Body length 2.5–2.8 mm. Distribution. – Known only from southern Sweden: SK: Krankesjön, 30.iv.1986, 11.v.1986 (B. Stille); SM: Kalmar, 07.v and 10.v.1986 (B. Stille). European species. Biology. – The larvae live as inquilines in galls of Lipara similis on Phragmites australis. They feed inside the hollow of the reed stem, and before pupation most of them pass down below the L. similis larvae (Collin, 1932).

160. Cryptonevra diadema (Meigen, 1830) Figs 711, 719–726 Chlorops diadema Meigen, 1830: 158; Meigen’s plate 211: 3 (Morge, 1976b). Syntype, a female in NHMW (Ismay, 1994a). Frons slightly longer than wide, broadly reddish yellow on anterior part and black posterad. Ocellar triangle predominantly black, reddish brown on apex and nearly reaching front margin of frons. Its surface minutely punctuate and subshining. Basal segments of antennae reddish brown. First flagellomere slightly longer than wide, distinctly angulated dorsodistally, basally and ventrally reddish brown to yellow, black dorsally. Arista black and minutely pubescent basally, distally glabrous. Scutum black, minutely shagreened, dusted on notopleura 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 mainly shining. Wings slightly whitish. Veins R4+5 and M divergent distally. Femora black except for yellow knees, anterior tibiae yellow, all tarsi yellow. Male with tarsal segments 2–4 of middle legs with black setae on ventral surface. Abdomen basally brownish yellow, remaining part black grey-dusted. Body length 2.5–2.8 mm. Distribution. – Denmark: SJ, WJ, NWJ, NEJ, F, NEZ. Sweden: SK, BL, HA, SM, VG, UP. Finland: north to Om along the Baltic coast. Russia: only Vib. Not recorded from Norway. Trans-Palearctic species, known from Europe to Japan, in southern part of Eurasia and in North Africa. Biology. – Adults in May to July. 188

The larvae live in shoots of Phragmites australis, usually as inquilines in Lipara galls. Egg, all larval stages and pupurium described by Grichowska (2008c).

161. Cryptonevra flavitarsis (Meigen, 1830) Figs 705–710, 712–718 Chlorops flavitarsis Meigen, 1830: 161; Meigen’s plate 211: 6 (Morge, 1976b). Lectotype in NHMW (Nartshuk, 1997a). Frons slightly longer than wide, black. Ocellar triangle black dull, nearly reaching front margin of frons. Genae nearly as wide as first flagellomere. First flagellomere slightly longer than wide, dis-

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 and minutely pubescent. Scutum black, minutely shagreened and shining, thinly dusted on notopleura and before scutellum, evenly clothed with blackish setulae. Scutellum grey-dusted. Pleura mainly shining. Wings slightly greyish. Veins R4+5 and M divergent distally. Femora black except for yellow knees, anterior tibiae black, tarsi yellow or brown. Male with tarsal segment 4 (rarely also 3) of middle legs with black setae on ventral surface. Abdomen basally brownish yellow, remaining part black grey-dusted, abdominal setae dark. Body length 1.5–2.5 mm. Distribution. – Common. Denmark: from all districts except SZ and NWZ. Sweden: north to JÄ. Norway: Ø, AK, VE. Finland: north to Om. Russia: Vib, Kr. Trans-Palearctic species, known from the British Isles eastwards to China and southwards to North Africa. Biology. – Adults from the end of April to beginning of August on Common reed (Phragmites australis). The larvae develop as inquilines in Lipara galls (Giraud, 1863), repeated by many authors. Recently Reader (2003) characterized the activity of the Cryptonevra flavitarsis larvae in the galls of Lipara lucens (wrongly given as L. rufijitarsis?) as “intraguild kleptoparasitism”. This seems questionable, however, as the larvae of these two species feed on different parts of the plant, and it does not seem that the presence of Cryptonevra larvae affects the well-being of the gall-inducing L. lucens larvae. Egg, all larval stages and pupurium described by Grichowska (2008a).

Genus Diplotoxa Loew, 1863 Diplotoxa Loew, 1863: 54. Type species: Diplotoxa versicolor Loew, 1863 by monotypy. Species of Chlorops-like general shape. Body mainly yellow with black nearly confluent scutal stripes, 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 R2+3 and R4+5 strongly procurved, R4+5 and M divergent and crossveins approximated. Middle tibiae with slightly curved spur. Tibial organ absent. Male genitalia: phallus, pre- and postgonites 190

fused into a rigid aedegal complex which articulates with hypandrium and probably functions as “pseudophallus”. Distribution. – Holarctic, Australia.

162. Diplotoxa messoria (Fallén, 1820) Figs 727–731 Oscinis messoria Fallén, 1820: 5. Lectotype in ZMLU (Andersson, 1963). Frons as long as wide, yellow. Ocellar triangle black, shining, extending nearly to front margin of frons, behind nearly touching eyes. Genae and face yellow, genae a little narrower than first flagellomere. Antennae black, first flagellomere round, arista white. Palpi yellow. Scutum with 3 wide, sometimes nearly fused black, dusted stripes, which extend to scutellum. Pleura yellow with black spots on anepisternum, katepisternum and katepimeron. Scutellum brown to black, apical setae shorter than scutellum. Abdomen dark brown to black, shining. Femora and hind tibiae black, front and middle tibiae fuscous yellow, tarsi darkened. Body length 2.5–3.0 mm. Distribution. – Denmark: from most districts. Sweden: north to NB along coast of Baltic Sea. Norway: AK: Oslo (Siebke). Finland: north to ObS. Russia: Vib, Kr. Holarctic species, in the Palearctic from the British Isles to the Far East of Russia. Biology. – Adults from middle of June to the beginning of September, on marshy habitats with Eleocharis. Some specimens were reared from this plant moved into laboratory (Nartshuk, 1962a). The species develops in rotting stems of Eleocharis macrostachya in North America (Valley et al., 1969). Tschirnhaus (1981) listed as host plants Eleocharis palustris and E. uniglumis. Other Nearctic species of Diplotoxa are also associated with species of Eleocharis (Wearsch & Foote, 1994).

Genus Diplotoxoides Andersson, 1977 Diplotoxoides Andersson, 1977: 150, as subgenus of Diplotoxa Loew. Type species: Diplotoxa dalmatina Strobl, 1900 by original designation.

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 as high as long. Anterior ommatidia larger than posterior ones. Ocellar triangle large smooth shining. Head setae well developed. First flagellomere rounded. Scutum smooth, dusted. The wings have veins R2+3 and R4+5 procurved, R4+5 and M divergent and crossveins approximated. Middle tibiae with slightly curved spur. Tibial organ absent. Male genitalia: phallus, pre- and postgonites fused into aedegal complex which articulates with hypandrium. Distribution. – Not clear, Palearctic?

163. Diplotoxoides dalmatina (Strobl, 1900) Figs 732–736 Diplotoxa dalmatina Strobl, 1900: 628. Syntypes not found in NMBA or the Sarajevo Museum (Chvála, 2008). Frons yellow, as long as wide in the middle and narrowed in front. orb 6–7 short recurved. poc shorter than oc. Ocellar triangle black, shining, extending nearly to front margin of frons, behind nearly 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. Antennae black, first flagellomere round, arista black. Palpi yellow. Scutum with wide, usually fused black thinly dusted subshining stripes, which extend to scutellum. Scutellum often darkened, apical setae longer than scutellum, wide apart. Pleura yellow with black spots on anepisternum, katepisternum and katepimeron. Abdomen dark brown. Coxae yellow, femora and tibiae mainly black, tarsi darkened. Body length 1.5 mm.

Pleura smooth yellow. Wings of Chlorops-type. Tibial organ absent. Male genitalia: pre- and postgonites situated parallel to each other.

Distribution. – Rare species. Denmark: NWJ: Thy National Park. Sweden: SK. Finland: AB: Lohja; N: Hanko, Helsinki; Om: Nykarleby, Pietarsaari. Russia: Kr: Bolshie Gory. Not recorded yet in Norway. Trans-Palearctic species, found from British Isles to Far East of Russia.

Oscinis puncticollis Zetterstedt, 1848: 2636. Lectotype in ZMLU (Andersson, 1966).

Biology. – Adults from June to August, in wetlands.

Genus Epichlorops Becker, 1910 Epichlorops Becker, 1910: 77. Type species: Oscinis puncticollis Zetterstedt, 1848 by original designation. Body black with partly yellow head, pleura and legs. Head as long as high with short setae and setulae. Ocellar triangle very large, behind almost as broad as frons, reaching front margin of frons in a rather broad tip. First flagellomere rounded. Scutellum rounded, black with stellate microsculpture. 192

Distribution. – Holarctic Region.

164. Epichlorops puncticollis (Zetterstedt, 1848) Figs 19, 737–743

Frons as long as wide, yellow. Ocellar triangle reaching front margin of frons, black smooth shining, anterior apex obtuse. Genae slightly wider than first flagellomere. Antennae and arista black. Palpi yellow. Scutum longer than wide, black except for lateral margins, undusted with strong stellate microsculpture and evenly clothed with black setulae. Postpronota yellow with black spot. Scutellum yellow, apical setae rather approximated. Pleura yellow with several black spots. Abdomen completely black dorsally except for extreme lateral margins. Legs mainly yellow, hind tibiae medially with black band and all tarsi black, sometimes legs extensively darkened. Body length 2.6–3.5 mm. Distribution. – Common species. Denmark: widespread, but absent from northern districts of Jutland. Sweden: north to NB. Norway: VE: Tønsberg (Siebke). Finland: north to Om. Russia: Vib, Kr. Holarctic species.

Biology. – Adults from June to the beginning of September in wet meadows and marshes with Carex. In North America E. exilis (Coquillett) was reared from Carex crinita (Rogers et al., 1991).

Genus Eurina Meigen, 1830 Eurina Meigen, 1830: 3. Type species: Eurina lurida Meigen, 1830 by designation of Westwood, 1840: 147. Body brown dull elongated. Head longer than high with frons strongly projecting, about as long as eye. Face retreating. Eyes small rounded. Ocellar triangle large, reaching front margin of frons, with longitudinal groove, surface covered with small setulae. Head setae very small. Antennae prolonged, first flagellomere trapezoid, longer than wide. Scutum flattened, evenly covered with short setulae. Scutellum rounded flat. Tibial organ absent. Male genitalia: pre-, postgonites, phallapodeme, and phallapodemic sclerites fused into a very broad disc-like structure. Distribution. – Probably worldwide.

Key to species of Eurina Meigen 1

–

Body dull grey, scutum with numerous lack shining spots at base of setulae. Ocellar triangle wide with central sulcus and numerous punctures at base of setulae, shining along central sulcus and dusted on sides (Fig. 744). Male genitalia Figs 747, 748 . . . . E. calva Egger Body reddish with distinct black stripes on scutum and spots at base of setulae. Ocellar triangle shorter with central sulcus and numerous punctures at base of setulae, shining on whole surface (Fig. 749). Male genitalia Figs 750, 751 . . . . . . . . . . . . . . . . . . E. lurida Meigen

165. Eurina calva Egger, 1862 Figs 744–748 Eurina calva Egger, 1862: 778. Lectotype in NHMW (Nartshuk, 1997b). Body brown covered with thick greyish dust. Head nearly triangular in side view. Frons strongly pro-

jected. Ocellar triangle reaching front margin of frons, with median black sulcus and coarse black punctuations. Genae broad and punctuate. Antennae from brown to black, scapus long triangular, first flagellomere 1.5 times longer than wide. Scutum longer than wide, with four black punctuate stripes. Scutellum rounded, apical setae very short. Pleura densely dusted. All thoracic setae short. Wings grey brownish. Halteres fuscous yellow. Body length 6.0 mm. Distribution. – Only from Denmark: SJ: Sønderborg, vi.1900 (W. Wüstnei), 1 male, 1 female; Sottrupskov, 12 and 15.vi.1893, 08.vi.1895, 04.vi.1898 (W. Wüstnei), 2 males, 4 females. Euro-Mediterranean species.

166. Eurina lurida Meigen, 1830 Figs 749–751 Eurina lurida Meigen, 1830: 5; Meigen’s plate 146: 11 (Morge, 1976a). Lectotype male in MNHN (Nartshuk, 2006). Body brown dull. Head longer than high with frons strongly projecting, longer than wide. Face retreating. Eyes small rounded, bare. Ocellar triangle large, straight-sided, reaching front margin of frons, with a conspicuous central groove and sulci along side margins. Ocellar tubercle black. Genae as wide as eye, covered with small setulae. Antennae prolonged, first flagellomere trapezoid, longer than wide, brown to black, arista black. Scutum flattened with 3 black, dusted longitudinal stripes, evenly covered with short setulae. Scutellum rounded flat with short apical setae. Anepisternum covered with many short black setulae. Abdomen reddish-brown dorsally, yellow ventrally. Wings slightly brownish. Legs yellow. Body length 6.0–6.5 mm. Distribution. – Denmark: LFM: Ulfshale, 27– 29.v.1983 (Andersen & Michelsen), 2 females. Sweden: SK: Klagshamn, 25.v.1981 (H. Andersson), 2 males; BO: Bro, 17.vii.1945 (E. Wieslander), 1 male. Norway: AK: Konglungen, Asker, 04.vi.1997 (G.E.E. Söli). Not found in Finland or Russian parts of Fennoscandia. Euro-Mediterranean species. Biology. – Adults on Common reed (Phragmites australis) and other plants in coastal, saline habitats (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: hypopygium, 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 controversial: Séguy (1934) mentioned Phragmites australis as host plant. Tschirnhaus (1981) records Bolboschoenus maritimus. In Israel larvae of this species develop on Eleocharis dulcis without producing top galls (Kaplan et al., 1986), but Ismay (1999), based on pers. comm. from A. Freidberg, listed Bolboschoenus maritimus as host plant but failed to find galls on that plant in England.

shorter than antennae. Scutum moderately convex, smooth. Scutellum rounded. All setae and setulae white. Wings of Chlorops-type. Middle tibiae with spur. Tibial organ present. Male genitalia: surstyli with pointed hook-like middle lobe; pregonites straight, postgonites short curved.

Genus Eutropha Loew, 1866

Key to species of Eutropha Loew

Distribution. – Palearctic, Afrotropical and Australasian Regions.

Eutropha Loew, 1866: 26. Type species: Chlorops fulvifrons Haliday, 1833 by designation of Becker, 1910: 59.

1

Head as long as high with short setae. Ocellar triangle smooth shining. Genae very wide. First flagellomere large rounded squarish. Arista very short,

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194

Pleura and scutellum yellow. Scutum with grey-dusted stripes fused in some specimens . . . . . . . . . . . . . . . . . . . . . . . . E. variegata Loew Thorax and abdomen black, shining. Only sides of scutum a little dusted. Male genitalia Fig. 756 . . . . . . . . . . . . . . E. fulvifrons (Haliday)

Not so far recorded in Russian parts of Fennoscandia. 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 secondary 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 view. 753: head, lateral view. 754: antenna. 755: wing. 756: male genitalia, ventral view.

167. Eutropha fulvifrons (Haliday, 1833) Figs 752–756 Chlorops fulvifrons Haliday, 1833: 172. Lectotype in NMID (Ismay et al., 2001). Body almost entirely shining brownish-black with white setae and setulae. Frons longer than wide, yellow in front and brown behind, or entirely brown, shining with white setae and setulae. Ocellar triangle extending 2/3 length of frons followed by linear extension to front margin of frons, black, shining. Face and genae yellow, the latter with black below margin. Genae 1.2–2 times as wide as first flagellomere. Antennae black or first flagellomere yellowish below. First flagellomere round or slightly angulated. Arista shorter than antennae, black, nearly bare. Palpi yellow. Scutum entirely black, shining, scutellum from yellow to black. Setae and setulae of thorax white. Abdomen black, shining, white pubescent. Legs mainly black. Halteres yellow. Body length 2.0–2.5 mm. Distribution. – Denmark: WJ, NWJ, NEJ, NWZ, NEZ. Sweden: SK, HA, ÖL, GO, NB. Norway: Ø, AK, VE, Ry. Finland: N: Hanko, 25.vii.1927 (Krogerus), 1 male.

Frons as long as wide, yellow with white setae and setulae. Ocellar triangle extending nearly to front margin of frons, yellow shining with median sulcus. Ocellar tubercle black. Face and genae yellow. Genae in male as wide as first flagellomere, in female 1.2 times wider than first flagellomere. Antennae yellow in male and black in female. First flagellomere large, especially in male, with upper corner. Arista shorter than antennae, black or whitish, nearly bare. Palpi yellow. Scutum yellow with 5 blackish densely grey-dusted and white pubescent stripes. Scutellum yellow. Pleura with black marks. Setae of thorax white. Abdomen brown, white pubescent. Halteres yellow. Body length 2.0–2.5 mm. Distribution. – Rare species. Norway: Ø: Larkollen, 06.vii.1955 (Ardö), 1 female. Finland: Ab: Vihti; N: Tvärminne, Hanko. Russia: Vib. Not so far found in Denmark and Sweden. Palearctic species, known from Europe to Central Asia. Biology. – Adults in July and August on sand dunes along sea coasts. Flies were found under leaves of Honckenya peploides. It is known that some tropical species of the genus are also associated with sea coast and develop in mammalian carrion (Kirk-Spriggs et al., 2001).

195

Genus Lasiosina Becker, 1910

5

Lasiosina Becker, 1910: 73. Type species: Chlorops cinctipes Meigen, 1830 by original designation.

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Body yellow with black stripes and spots. Head as 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 margin of frons, often with black rhomboid spot at tip, dusted but with shining areas at sides behind and on apical black spot. There is sexual dimorphism in colour in some species: antennae and palpi in males yellow, in females black. First flagellomere rounded. Scutum yellow with black stripes, dusted. Wings of Chlorops-type. Tibial organ absent. Male genitalia: surstyli fused with epandrium, granulate at apex. Postgonites apically curved upwards. Distribution. – Worldwide.

Key to species of Lasiosina Becker 1

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2

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196

Ocellar triangle without shining rhomboid spot apically. Wings usually shorter than abdomen. Male genitalia Fig. 774 . . . . . . . . . . . . . . . . . . . . . . L. parvipennis Duda Ocellar triangle with shining rhomboid spot apically (Fig. 764). Wings normal, longer than abdomen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Only one posterior npl seta. First flagellomere and palpi in male yellow, in female black. Male genitalia Fig. 760 . . . . . . . . . . L. brevisurstylata Dely-Draskovits Two posterior npl setae . . . . . . . . . . . . . . . . . . 3 Antennae black and palpi yellow in both sexes or only in males. Abdomen with white or black setulae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Antennae and palpi yellow in male and black in female. Abdomen with black setulae . . . . 5 Abdomen with white setulae, only some setulae black. Male genitalia Figs 757–759 . . . . . . . . . . . . . . . . . . . . . . . . L. albipila (Loew) Abdomen with black setulae. Male genitalia Figs 772, 773, 775–777 . . . . . . . . . . . . . L. intermedia Dely-Draskovits, L. subnigripes Dely-Draskovits

Male genitalia: surstyli shorter and their tip dentate and directed to the middle (Figs 767– 771) . . . . . . . . . . L. herpini (Guérin-Méneville) Male genitalia: surstyli longer and directed straight (Figs 761, 762) . . . . . . . L. danielssoni Nartshuk & Andersson

Species key to known 3rd instar larvae of Lasiosina Becker 1

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Body length 3 mm. Mandible with 1 triangular middle tooth; dental sclerite more or less arched (Fig. 984). On ventral side of abdomen creeping welts consist of 4–5 incomplete rows of spicules. In front of perianal opening a row of backward directed spicules (Fig. 985) . . . . . . . . . . L. brevisurstylata Dely-Draskovits Body length 4 mm. Mandible with 3–4 trapezoid middle teeth; dental sclerite more or less conical or semicircular (Fig. 982). On ventral side of abdomen creeping welts consist of 5–8 complete rows of spicules, usually uniformly situated. In front of perianal opening no spicules (Fig. 983) . . . . . . . . . . . . . . L. herpini (Guérin-Méneville)

169. Lasiosina albipila (Loew, 1866) Figs 757–759 Diplotoxa albipila Loew, 1866: 37. Type material not found in ZMHU (Wendt, 1975, 1978). Head broader than deep or long. Frons wider than long, yellow with black setae and setulae. Ocellar triangle extending half length of frons followed by linear extension to front margin of frons, yellow with two black spots, one on ocellar tubercle and another rhomboid spot on apex. Genae wider than first flagellomere. Antennae black, first flagellomere rounded. Palpi yellow. Scutum with 3 black densely grey-dusted stripes, scutellum yellow. Pleura with several black marks. Spots on anepisternum and katepisternum shining. Abdomen brown, covered with white setulae, but some black setulae present. Legs partly black. Body length 2.5–3.0 mm. Distribution. – Rare species. Sweden: SK: Ivö klack, Ivön, 19.v.1998 (R. Danielsson), 1 female; SM; UP.

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 found in Denmark and Norway. European species.

shining on upper part and dusted below. npl 1 + 1. Abdomen blackish dorsally, covered with black setulae. Legs yellow. Body length 2.0–2.6 mm.

Biology. – Adults in meadows. It was found in Switzerland that adults feed on secreta from chrysomelid larvae of Agelastica alni (Pschorn-Walcher, 1956).

Distribution. – Found only in Finland: St: Eura, 16.vi.1982 (Koponen), 1 female. Eurasian species known eastwards to Yakutia, East Siberia.

170. Lasiosina brevisurstylata Dely-Draskovits, 1977

Biology. – In Bulgaria reared from shoots of winter wheat, oats and spring barley together with L. herpini (Beschovski & Krusteva, 1995, 1997; Krusteva & Beschovski, 1998, 2000, 2001). Pupa described by Beschovski & Krusteva (1997).

Figs 760, 984, 985 Lasiosina brevisurstylata Dely-Draskovits, 1977: 267. Holotype and paratypes in HNHM. Frons as wide as long, yellow with black setae and setulae. Ocellar triangle nearly 1/3 as wide as frons, yellow shining, extending little more than half length of frons with shining black rhomboid spot on apex. Three pairs of long orb. vte longer than vti. Genae wider than first flagellomere. In male antennae and palpi yellow, first flagellomere little darkened above. In female first flagellomere and palpi black. Scutum with 3 black grey-dusted stripes, narrowed behind, scutellum yellow. Pleura with several black marks. Spot on katepisternum

171. Lasiosina danielssoni Nartshuk & Andersson, sp. nov. Figs 761–763 Holotype male, Sweden: GO: Lummelunda bruk, 30.viii.1992 (R. Danielsson). Holotype in ZMLU. Diagnosis. – New species belongs to L. herpini group (= cinctipes auct.) and is similar to L. cinctipes in colour of frons, antennae, palpi and legs. It differs by structure of male genitalia with longer surstyli with straight ends not curved towards middle and elongated hypandrium. 197

Description. – Male. Body yellow with black greydusted stripes on scutum and four spots on pleura. Frons as long as wide, little narrowed anteriorly. Base of ocellar triangle occupies 1/3 of frontal width. Ocellar triangle extending little more than half length of frons followed by linear extension to front margin of frons. Ocellar tubercle black, dull, another black, shining rhomboid spot situated on apex of triangle. Lateral sides narrow and line between anterior ocellus and tip of ocellar triangle brownish. Two long orb. vte, vti and oc nearly equal, poc little shorter. Frons covered with short black setulae. Eyes bare, strongly oblique. Genae wider than first flagellomere. Antennae yellow, first flagellomere round, with small dark spot at aristal base. Arista black. Scutum little longer than wide, yellow, with three wide black, dusted stripes. Central stripe narrowed behind and not extended to scutellum. Postpronota with small brownish spot. Scutellum yellow, apical setae longer than scutellum. Pleura with black, shining spot on anepisternum and entirely dusted spot on katepisternum. Remaining two spots black, dusted. npl 1 + 2. Abdomen brownish dorsally. Legs yellow with little darkened tarsi, especially on anterior legs. Wings transparent with venation typical for the genus. Halteres yellow. All setae and setulae of head and thorax black. Male genitalia: surstyli rather long and little curved in lateral view; distally dentate on inner side but not curved towards middle. Hypandrium elongated, pregonites long, postgonites with two setae. Body length 2.5 mm. Female unknown.

Genae wider than first flagellomere. First flagellomere rounded, yellow, darkened around insertion of arista in male and entirely black in female. Palpi yellow in male and black in female. Scutum with 3 black densely grey-dusted stripes, scutellum yellow. Pleura with several black marks. Spot on katepisternum black, completely dusted or shining at anterior and posterior ends in some specimens. Abdomen covered with black setulae. Legs yellow. Body length 2.5–3.0 mm. Distribution. – The most common species of the genus. Sweden: SK, SM, ÖL, GO, ÖG, UP. Norway: AK: (Rygg, 1966a, as L. cinctipes). Finland: Ab, N, St. Russia: Vib, Kr. Not so far from Denmark. Trans-Palearctic species.

Etymology. – The species is named in honor of R. Danielsson who found the only specimen known so far.

Biology. – Adults in July to September in dry meadows and cereal fields. The larvae develop in shoots of different grasses, including cereals: rye, wheat, oats, barley as secondary invaders, often in the same shoots as Chlorops pumilionis (Tzygankov, 1929, 1930; Balachowsky & Mesnil, 1935; Goodliffe, 1939; Nye, 1958; Krusteva & Beschovski, 2000, 2001). It has been recorded as a pest of cereals in Finland (Kallio, 1950) and in Sweden (Johansson, 1960). Rygg (1966a) reared it from barley shoots in Norway. As host plants were listed grasses from the genera Calamagrostis, Agropyron, Bromus, Festuca, Glyceria, Hordeum (Nartshuk, 1972a; Tschirnhaus, 1981). Larvae described by Kreiter (1928), Tzygankov (1929, 1930), Balachowsky & Mesnil (1935), Oettingen (1935), Nye (1958), Beschovski & Krusteva (1997). The species has been misidentified as L. cinctipes in nearly all applied publications and descriptions of larvae.

172. Lasiosina herpini (Guérin-Méneville, 1843)

173. Lasiosina intermedia Dely-Draskovits, 1977

Figs 764–771, 982, 983

Figs 772, 773

Chlorops herpini Guérin-Méneville, 1843: 30. Syntypes in MNHN. Lasiosina cinctipes auct., nec Meigen, 1830 (Nartshuk, 1997a).

Lasiosina intermedia Dely-Draskovits, 1977: 269. Holotype and paratypes in HNHM.

Head broader than deep or long. Frons wider than long, yellow with black setae and setulae. Ocellar triangle extending half length of frons followed by linear extension to front margin of frons, yellow with two black spots, one on ocellar tubercle and another rhomboid shining spot on apex. 198

Head broader than deep or long. Frons wider than long, yellow with black setae and setulae. Ocellar triangle extending half length of frons followed by linear extension to front margin of frons, yellow with two black spots, one on ocellar tubercle and another rhomboid shining spot on apex. Genae nearly as wide as first flagellomere. First flagellomere rounded, extensively darkened in both sexes,

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 distiphallus. 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 from yellow to brown, in female black. Scutum with 3 black densely grey-dusted stripes, scutellum yellow. Pleura with several black marks. Spot on katepisternum black below and yellow above, mainly shining, dusted only on lower angle. Abdomen covered with black setulae. Legs extensively darkened. Body length 2.5–3.0 mm. Distribution. – Found only in southern Sweden: SK, HA, ÖL, GO. European species.

Biology. – Adults in May, and again in July and August in meadows.

174. Lasiosina parvipennis Duda, 1933 Fig. 774 Lasiosina parvipennis Duda, 1933: 140. Lectotype in NHMH (Nartshuk, 1999). Frons longer than wide, yellow, with black setae and setulae. Ocellar triangle brown dull, main 199

part extending half length of frons, apical part linear. Face and genae yellow, genae as wide as first flagellomere. First flagellomere, arista and palpi black. Scutum yellow with black nearly fused densely dusted stripes. Scutellum yellow or darkened. Wings variable in length: shorter or equal of abdomen. Legs with femora black, tibiae and tarsi yellow. Halteres yellow. Body length 1.5 mm. Distribution. – Rare species. Found only in Finland: Ab, St, Ta, Sa, Kb, Om and Russia: Vib, Kr. Eurasian species, known from North Europe to East Siberia and Mongolia. Biology. – Adults in May, and again in September on bogs and marshes. Probably two generations per year.

175. Lasiosina subnigripes Dely-Draskovits, 1977 Figs 775–777 Lasiosina subnigripes Dely-Draskovits, 1977: 274. Holotype and paratype in HNHM. Head broader than deep or long. Frons as wide as long, yellow with black setae and setulae. Ocellar triangle extending half length of frons followed by linear extension to front margin of frons, yellow with two black spots, one on ocellar tubercle and another rhomboid shining spot on apex. Genae narrower than first flagellomere. In male first flagellomere rounded, extensively darkened, palpi yellow. (Female unknown). Scutum with 3 black densely grey-dusted stripes, central one narrowed behind. Scutellum yellow. Pleura with common black marks. Spot on katepisternum black, shining. npl 1 + 2. Abdomen covered with black setulae. Legs yellow. Body length 2.5 mm.

produced acutely. Proboscis somewhat prolonged. First flagellomere rounded. Scutum rather convex, smooth, dusted. Scutellum rounded. Wings of Chlorops-type. Tibial organ absent. Female cerci very short. Male genitalia: epandrium flattened posteriorly with a strong marginal carina. Surstyli apically black and densely sclerotized. Middle lobe of surstyli rectangular. Postgonites very long, pregonites indistinct. Distribution. – Palearctic Region and New Zealand.

176. Melanum laterale (Haliday, 1833) Figs 17, 778–782 Chlorops lateralis Haliday, 1833: 172. Lectotype in NMID (Ismay et al., 2001). Frons yellow, black pubescent. Ocellar triangle black, shining, dusted on ocellar tubercle. Ocellar triangle extending nearly to front margin of frons with acute apex. Face and genae yellow, genae as wide as first flagellomere, with strongly projected and acute vibrissal angle. Antennae black, first flagellomere as long as wide. Palpi yellow. Scutum black, dusted covered with short black setulae. Scutellum yellow. Pleura yellow with usual black marks. Spot on katepisternum dusted. Abdomen dorsally black with yellow lateral margins. Legs mainly yellow. Body length 2.5–3.0 mm. Distribution. – Denmark: from all districts. Sweden: along Baltic coast north to NB. Norway: AK, On, VA, Ry, HOi, STy, STi. Finland: north to ObS. Russia: Vib; Kr; Lr: Ponoj, 1 female. Trans-Palearctic species, known from the British Isles to Japan.

Distribution. – Rare species. Found only in Sweden: UP: Hässelby, Böje, 20.v.1969, 20.v.1970, 08.v.1972 (Kur), 5 specimens. European species.

Biology. – Adults from June to September in wetlands, especially on coastal salt marshes, but also on shores of lakes and banks of rivers inland. The larvae probably phytophagous. Tschirnhaus (1972) recorded Juncus gerardi as host plant.

Genus Melanum Becker, 1910

Genus Meromyza Meigen, 1830

Melanum Becker, 1910: 50. Type species: Chlorops lateralis Haliday, 1833 by original designation.

Meromyza Meigen, 1830: 163. Type species: Musca saltatrix Linnaeus, 1761 by designation of Macquart, 1835: 589.

Head as long as high, yellow with black. Ocellar triangle large, behind almost as broad as frons, with acute tip, smooth shining, black. Vibrissal angle 200

Body light green or yellowish green with dusted black or red stripes and maculae. Head as long

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 anterior part of eye larger than in posterior. Ocellar 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 concave with indistinct carina. Scutum moderately convex, usually smooth and dusted, with black or red stripes. Scutellum rounded. Wings with veins R2+3 and R4+5 strongly procurved. Veins R4+5 and M distinctly divergent. Legs stout, hind femora strongly thickened, ventrally with two rows of black spines. Hind tibiae correspondingly curved. Tibial organ absent. Male genitalia: postgonites usually large, strongly sclerotized and black, rarely small and nearly transparent. Structure of male genitalia, especially shape and colour of postgonites and surstyli are the only reliable characters to distinguish species of the genus. Larvae phytophagous, develop in shoots of Poaceae, including cereals.

some species on the borders of the Oriental and Neotropical Regions.

Key to species of Meromyza Meigen (males only) 1 – 2 (1)

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3 (2) Distribution. – Predominantly Holarctic, but one species is known in the Afrotropcal Region and

Palpi pale, at most only the tip a little darkened . . . . . . . . . . . . . . . . . . . . . . . . . 2 Palpi mainly black, at least on apical half . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .8 Abdomen with a central dark stripe and two dark lateral spots at the base, no lateral dark marking on tergites 3–5. Surstyli rather long and expanded medially with long hairs (Figs 808, 829, 846) . . . . . . . . 3 Abdomen with three dark stripes or nearly black dorsally. Surstyli different . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5 Aedeagus swollen at base, basally nearly twice as broad as at middle. Lower margin of anterior process of postgonite 201

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4 (3)

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5 (2)

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6 (5) –

7 (6)

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8 (1)

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202

strongly convex medially, the tip of anterior process turned a little laterally. Frons distinctly produced, hypopleural marks yellow to reddish. Genitalia Figs 846–850 . . . . . . . . . . . . . . . . . M. pratorum Meigen Aedeagus not swollen at base, basally not broader than at middle. Lower margin of anterior process of postgonite more straight and not turned laterally. Frons less produced and hypopleural mark black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Lower margin of anterior process of gonite a little convex and the tip less pointed. Genitalia Figs 808–811 . . . . . . . . . . . . . . . . . . M. ingrica Nartshuk Lower margin of anterior process of postgonite more straight and the tip pointed. Genitalia Figs 829–833 . . . . . . . . . . . . . . M. ornata (Wiedemann) Hind femora more swollen, 4 times as wide as hind tibia. Genitalia Figs 852– 855 . . . . . . . . . M. rohdendorfiji Fedoseeva Hind femora less swollen and only 2.5– 3 times wider than hind tibia. Genitalia different . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Central scutal stripe does not extend to the scutellum . . . . . . . . . . . . . . . . . . . . . . 7 Central scutal stripe extending over the scutellum. Surstyli short and straight. Genitalia Figs 812–815 . . . . . . . . . . . . . M. mosquensis Fedoseeva Surstyli long and expanded at tip laterally. Postgonites more massive with small tooth at tip. Upper margin of anterior process of postgonites slightly convex. Genitalia Figs 882–885 . . . . . . . . . . . . . . . . M. zimzerla Nartshuk Surstyli short, not expanded laterally. Postgonites with acute tip. Upper margin of anterior process of postgonites strongly convex. Genitalia Figs 789–792 . . . . . . . . . . . . . . M. bohemica Fedoseeva Frontal triangle with its boundary linearly darkened. Genae with black hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .9 Frontal triangle not darkened laterally. Genae without black hairs . . . . . . . . . . 10

9 (8)

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10 (8) –

11 (10)

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12 (11)

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13 (12)

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14 (13)

Frontal triangle 1.5 times longer than broad at base with concave side margins, and wrinkled surface. Postgonite more massive, posterior process long. Genitalia Figs 798–801. Body larger, 3.5–4.0 mm . . . . . . . . . . . . . . . . . M. elbergi Fedoseeva Frontal triangle nearly equilateral, with straight side margins. Postgonite small. Genitalia Figs 871–875. Body smaller 2.5– 3.0 mm . . . . . . M. triangulina Fedoseeva Anterior process of postgonite elongated and pointed at the tip . . . . . . . . . . . . . . 11 Anterior process of postgonite broad at the tip, sometimes depressed and turned laterally, postgonite more massive . . . . 16 Posterior process of postgonite long and directed posteriorly, basal part of aedeagus swollen and nearly square. Central mesonotal stripe usually ending before scutellum. Genitalia Figs 793–797 . . . . . . . . . . . M. curvinervis (Zetterstedt) Posterior process of postgonite shorter and directed downwards or a little anteriorly, sometimes hardly separated from anterior process. Aedeagus not swollen basally . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Postgonite scarcely darkened, nearly transparent or a little brown, posterior process small, sometimes hardly separated from anterior process. Hind femora more swollen, nearly 3 times wider than hind tibia . . . . . . . . . . . . . . . . . . . . . . . . 13 Postgonite more strongly sclerotized dark brown or almost black, posterior process well developed. Hind femora usually less swollen, not more than 2–2.5 times wider than hind tibia . . . . . . . . . . . . . . . . . . . . 15 Surstyli rather narrow and has a group of black thickened bristles at the base. Genitalia Figs 816–821 . . . . . . . . . . . . . . . M. nigriseta Fedoseeva Surstyli moderately broad, sometimes long but without black thickened bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Surstyli longer than epandrium. Anterior process of postgonite more elongated and directed downwards. Central scutal

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15 (12)

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16 (10)

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17 (16)

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18 (17)

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stripe ends before scutellum. Genitalia Figs 866–870 . . . . . M. sibirica Fedoseeva Surstylus shorter than epandrium. Anterior process of postgonite shorter and directed anteriorly. Central scutal stripe extending to scutellum. Black spot on ocellar triangle elongated anteriorly. Genitalia Figs 840–844 . . . M. pluriseta Péterfi Surstyli rather short and broad; postgonite small. Genitalia Figs 835–839 . . . . . . . . . . . . . . . . M. palposa Fedoseeva Surstyli narrower, expanded medially; postgonite longer. Genitalia Figs 822–826 . . . . . . . . . . . . . M. nigriventris Macquart Postgonite massive, posterior process of postgonite divided into two parts, anterior process at tip straight, not pointed or depressed. Ocellar triangle short (Fig. 783). Genitalia Figs 785–788 . . . . . . . . . . . . . . . . . . M. bifurcata sp. nov. Posterior process of postgonite not divided, tip of anterior process more or less pointed or depressed. Ocellar triangle longer . . . . . . . . . . . . . . . . . . . . . . . . . 17 Scutal stripes mainly or partly red. Tip of anterior process of postgonites not depressed . . . . . . . . . . . . . . . . . . . . . . . . 18 Scutal stripes black. Surstyli expanded medially. Postgonite massive with tip of anterior process depressed. Aedeagus broadened basally. Genitalia Figs 859– 865 . . . . . . . . . . . . M. saltatrix (Linnaeus) Hind femora less swollen, 2.5–3.0 times thicker than tibia. Surstyli long, expanded a little laterally. Postgonite narrower. Aedeagus with bifid process basally. Genitalia Figs 876–881 . . . . . . . . . . . . . . . . . M. variegata Meigen Hind femora strongly swollen, 4 times thicker than tibia. Surstyli not expanded laterally. Postgonite massive with concave tip and small tooth-like projection. Base of aedeagus without bifid process. Genitalia Figs 802–807 . . . . . . . . . . . . . . . M. femorata Macquart

Species key to known 3rd instar larvae of Meromyza Meigen 1 – 2

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3 – 4 –

5

Distal segment of antennae brownish and retracted within basal segment . . . . . . . . . . . . . 2 Antennae not coloured . . . . . . . . . . . . . . . . . . 3 Anterior spiracles of transversal rosette type with 10–12 buds on two lobes (Fig. 1005). Spicular zones on abdominal segments consist of numerous lines on anterior half of segment and a few lines on hind margin segments (Fig. 1006). In addition wings of dentate sclerite point laterally instead of anterolaterally. Larvae light green, in shoots of Calamagrostis epigeios and Ammophila arenaria . . . . . . . . . . . . . . . . . . . . . M. pratorum Meigen Anterior spiracles of small rosette type with 7–9 buds (Fig. 1007). Spicular zones on abdominal segments consist of 20–30 lines on anterior half of segment in a broadly triangular field plus a few lines on hind margin (Fig. 1009). Wings of dentate sclerite point anterolaterally. Posterior spiracles not borne on lobes and close together. Larvae blue-green, in shoots of Dactylis glomerata . . . . . . . . . . . . . . . . . . . . . M. variegata Meigen Two pairs of additional teeth on inner side of mandible . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Three pairs of additional teeth on inner side of mandible (Fig. 1011) . . . . . . . . . . . . . . . . . . . . . 6 Anterior spiracles with 7–8 buds . . . . . . . . . . 5 Anterior spiracles with 9–10 buds. Spicular zones of abdominal segments triangular, occupy only anterior part of segment, on posterior part only 2–3 short lines present. Interspiracular processes branched 2 or 3 times. Larvae greenish yellow, 6–7 mm long, in shoots of Poa pratensis, Festuca rubra, Agrostis vulgaris . . . . . . . . . . M. mosquensis Fedoseeva Interspiracular processes on hind spiracles not branched. Spicular zones semicircular, occupy only anterior part of abdominal segments, on posterior part of segment only 2–3 short lines present. Larvae greenish yellow, 5 mm long, in shoots of Dactylis glomerata and Festuca rubra . . . . . . M. triangulina Fedoseeva 203

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6

– 7

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8

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Interspiracular processes on hind spiracles forked at tips. Larvae greenish 6–7 mm long, in shoots of Lolium perenne . . . . . . . . . . . . . . . . . . . M. bohemica Fedoseeva Anterior spiracles with 11–12 buds. Parastomal bar with lanceolar widening. Larvae bright green, 7–10 mm long, in shoots of Deschampsia caespitosa . . . . . . M. ornata (Wiedemann) Anterior spiracles with 7–10 (11) buds . . . . . . 7 Spicular zones on ventral side of all abdominal segments occupy only anterior part of segments, no additional spicular lines at posterior margin, but some striation present on dorsal side. Larvae green, 7–8 mm long, in shoots wheat, rye, barley, Elytrigia repens, Phleum pratense . . . . . . . . . . M. nigriventris Macquart Short additional spicular lines present on posterior part of segments besides main area on anterior part of all segments. Spicular zones short, not occupying half of segments . . . . . 8 Larvae greenish-yellow, anterior spiracles with 8–9 buds. Dentate sclerite long, broadly flattened. Larvae 6–7 mm long, in shoots of Poa pratensis, Festuca rubra . . . . . . . . . . . . . . . . . . . M. saltatrix (Linnaeus) Larvae bright green, anterior spiracles with 9– 10 (11) buds. Dentate sclerite short, sharply curved. Larvae 7–8 mm long, in shoots of Elytrigia repens, Roegneria ramosum, Hierochloe odorata . . . . . . . . . . . . . M. nigriseta Fedoseeva

177. Meromyza bifurcata Nartshuk & Andersson, sp. nov. Figs 783–788 Holotype male, Sweden: SK: Kullaberg, 02.vii.1974 (H. Andersson). Holotype in ZMLU. Diagnosis. – New species belongs to the saltatrix group of species with black palpi, black stripes on scutum and scutellum, three black stripes on abdomen, moderately thickened hind femora and strongly sclerotized massive postgonites. It differs from all known species by bifidly divided front process of postgonites and also rather short ocellar triangle. Description. – Ground colour of body light yellow. Head longer than deep with almost horizontal 204

long axis of eye. Frons produced beyond eye and parafacial plates equal half width of genae. Genae as wide as first flagellomere is long, with yellow setulae. Frons longer than wide, covered with black setulae. Only vte distinct among head setae. Ocellar triangle short, extending little more than half length of frons, yellow shining, with black, dusted ocellar tubercle and small brown points along side margins. Antennae yellow, first flagellomere little longer than deep, slightly darkened above. Arista brown, nearly bare. Occiput with brown stripe and two brownish spots laterally. Palpi black in apical part. Scutum longer than wide with three wide black, dusted stripes, central one extending over scutellum. Pleura with black, shining spot on anepimeron, black, dusted spot on katepimeron and reddish with black upper angles on katepisternum. Legs mainly yellow, front femora with black stripe. Hind femora 2.5 times thicker than tibiae. Wings transparent with venation typical for the genus. Abdomen with three black spots on tergites. Male genitalia: surstyli distinctly angulated on inner side. Postgonites black, hind process large triangular, front process bifidly divided, one twig longer than the other. Aedeagus with enlarged base. Body length 3.0 mm.

178. Meromyza bohemica Fedoseeva, 1962 Figs 789–792 Meromyza bohemica Fedoseeva, 1962: 472. Holotype in ZISP. Ground colour yellow. Ocellar triangle light, shining, with slightly darkened side margins. First flagellomere longer than wide, yellow darkened above. Palpi narrow, yellow, rarely slightly darkened at tip. Genae with some black setulae in a row. Stripes of scutum reddish centrally, dusted, outer margin of inner lateral stripes black. Central stripe not extending over scutellum. Pleura with reddish marks, except oval black mark on anepisternum. Abdomen with central brown to black stripe and spots on 2–5 tergites. Hind femora 2.5–3.0 times wider than tibiae. Body length 2.5–3.0 mm. Distribution. – Found only in Denmark: EJ, LFM, NWZ and southern Sweden: SK, HA, SM, ÖL, GO. European species. Biology. – Adults in July to September in meadows. Larvae develop in shoots of Lolium perenne (Fedoseeva, 1974).

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: hypopygium, 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 M. lolii Hubicka).

179. Meromyza curvinervis (Zetterstedt, 1848) Figs 793–797 Oscinis curvinervis Zetterstedt, 1848: 2620. Holotype in ZMLU (Andersson, 1966). Ground colour yellow. Frons longer than wide. Ocellar triangle extending 2/3 length of frons. Palpi

widely black on apical half. Stripes of scutum black, densely grey-dusted. Central stripe ends before scutellum, but sometimes before scutellum a black mark with indistinct borders. Sometimes central stripe partly reddish and scutellum with black mark. Spot on katepisternum black and mainly dull. Abdomen with only central reddish brown to black stripe. Hind femora 2.0–2.5 times wider than tibiae. Phallus swollen at base. Body length 2.0– 3.0 mm. 205

Distribution. – Scarce, only in southern Fennoscandia. Sweden: SK, ÖL, GO. Finland: Ab, Ka. Russia: Vib. Not so far found in Norway and Denmark. Eurasian species known from the British Isles to Mongolia. Biology. – Adults in July and August in meadows. Tschirnhaus (1981) proposed as host plants Ammophila arenaria and Corynephorus canescens.

180. Meromyza elbergi Fedoseeva, 1979 Figs 798–801 Meromyza elbergi Fedoseeva, 1979: 118. Holotype in ZISP. Ground colour fuscous yellow. Ocellar triangle wrinkled, shining, with distinctly concave, slightly darkened side margins. First flagellomere longer than wide, yellow with darkening above. Palpi narrow, black in distal half. Genae 1.5 times wider than wide of first flagellomere, with some black setulae in irregular rows. Stripes of scutum black, or central stripe reddish, dusted. Central stripe extending over scutellum. Pleura with reddish spots, except black marks on anepisternum and katepimeron. Abdomen with three brown to black stripes. Hind femora about 3 times as thick as tibiae. Body length 3.5–4.0 mm. Distribution. – Sweden: GO, NÄ, UP. Norway: AK. Finland: Al. Not found in Denmark or Fennoscandian districts of Russia. Central and eastern European species. Biology. – Adults in July and August in dry meadows.

181. Meromyza femorata Macquart, 1835 Figs 802–807 Meromyza femorata Macquart, 1835: 589. Types probably in MHNL. Ground colour yellowish. Frons produced in front no more than 1/2 breadth of first flagellomere. Ocellar triangle shining, rugose in apical part and black on ocellar tubercle only. Palpi black on apical half. Stripes of scutum red, grey-dusted, central stripe reaching scutellum and scutellum with red mark. Pleura with red marks, except small black mark on the anepisternum. Abdomen with central black stripe and small spots on 3–5 tergites. Hind 206

femora strongly swollen, nearly 4 times as thick as tibiae. Body length 3.5–4.0 mm. Distribution. – Southern species. Denmark: from all districts except NWJ and SZ. Sweden: SK, BL, SM, ÖL. Norway: AK, Ø. European species. Biology. – Adults from the end of June to September in meadows. Larvae phytophagous, develop in shoots of Dactylis glomerata.

182. Meromyza ingrica Nartshuk, 1992 Figs 808–811 Meromyza ingrica Nartshuk, 1992: 125. Holotype and 1 paratype in ZISP, other paratypes in NHMH. Ground colour greenish. Body elongated, tip of abdomen reaching well behind tip of hind femora. Head longer than deep. Frons produced in front as long as breadth of first flagellomere. Ocellar triangle longer than wide at base, with wrinkled surface and black on ocellar tubercle only. Palpi greenish. Stripes of scutum black, densely grey-dusted. Central stripe reaching scutellum and scutellum with black mark. Katepimeron with black or dark brown mark. Abdomen with only a central black stripe. Hind femora moderately swollen, about 2.5 times as thick as tibiae. Hypandrium covered with long hairs. Body length to 4.0 mm. Distribution. – Sweden: HS, JÄ, NB, LY. Norway: HOy, HOi. Finland: north to Ob. Russia: Vib. Not found in Denmark. Known till now only from the Fennoscandian districts listed above. Possibly more widely distributed, but not distinguished from other species of the M. pratorum species group. Biology. – Adults in July to August in wet meadows.

183. Meromyza mosquensis Fedoseeva, 1960 Figs 812–815 Meromyza mosquensis Fedoseeva, 1960: 457. Syntypes in ZISP. Ground colour greenish-yellow. Frons produced in front no more than 1/2 breadth of first flagellomere. Face more upright than in most Meromyza species. Ocellar triangle with wrinkled rhomboid

Figs 798–811. Meromyza elbergi Fedoseeva: 798: hypopygium, dorsal view. 799: hypopygium, lateral view. 800: postgonite. 801: basiphallus. 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.

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dark coloured apical part and black ocellar tubercle. Palpi yellow. Stripes on scutum rather wide and sometimes touching in front, black densely grey-dusted or central one reddish brown. Central stripe reaching scutellum and scutellum with dark brown mark. Pleura with brown to black marks. Abdomen short, dorsally shining, with 3 black stripes, in dark specimens entirely black dorsally. Hind femora moderately swollen, 2.5 times as thick as tibiae. Body length 3.0–3.5 mm.

Host plants: Elytrigia repens, E. tenerum, Leymus ramosus, Hierochloe odorata, Agrostis vulgaris, rarely larvae found in shoots of summer wheat and winter rye (Karpova, 1965, 1972; Nartshuk, 1972a; Fedoseeva, 2003). Larvae of third instar described by Fedoseeva (1966).

Distribution. – Denmark: SJ, WJ, F, LFM, NEZ, B. Sweden: SK, BL, HA, SM ÖL, GO. Norway: Ø, AK. Finland: Al, Ab, N, Ta, Sa. Russia: Vib, Kr, Lr. European species, but not in the southernmost parts.

Figs 822–826, 1010–1012

Biology. – Adults in June to August in meadows. Host plants: Poa pratensis, Agrostis vulgaris, Festuca rubra, F. pratensis, Elytrigia repens (Nartshuk, 1962a; Fedoseeva, 1966, 2003; Karpova, 1972). Hibernates as larvae of third instar. Larvae of all three instars are described by Fedoseeva (1966).

184. Meromyza nigriseta Fedoseeva, 1960 Figs 30, 816–821 Meromyza nigriseta Fedoseeva, 1960: 456. Syntype in ZISP. Ground colour greenish-yellow. Frons produced in front no more than 1/2 breadth of first flagellomere. Ocellar triangle slightly wrinkled in apical part and with rather large black spot on ocellar tubercle. Palpi black slightly thick in apical part. Stripes of scutum rather narrow, black densely grey-dusted or central one reddish brown. Central stripe reaches scutellum and scutellum with black mark. Abdomen with 3 black stripes or only one central stripe and with spots on tergites 3–5. Hind femora moderately swollen, no more than 3 times as thick as tibiae. Epandrium covered with black setae and surstyli with thickened black setae in several rows on basal part. Body length 3.5–4.0 mm. Distribution. – Denmark: NEJ, F, NEZ, B. Sweden: north to DR. Norway: AK. Finland: Ab, N, Sa. Russia: Vib. Eurasian species, known from the British Isles to Mongolia. Biology. – Adults from the end of June to beginning of August, in meadows. 208

185. Meromyza nigriventris Macquart, 1835 Meromyza nigriventris Macquart, 1835: 590. Type probably in MHNL. Meromyza basalis Roser, 1840: 63, Type 1 male in SMNS (Ismay, 1981a; Nartshuk, 1994). Meromyza cerealium Reuter, 1902a: 84. Type not found in NHMH (Nartshuk, 1992). Mesnil (1931) considered M. cerealium as a summer form of M. nigriventris (as M. saltatrix). Meromyza rostrata Hubicka, 1966: 16. Holotype in ZIWP. The species is very variable in colour, showing two distinct colour forms: Summer form. Ground colour yellow. Frons produced in front no more than 1/2 breadth of first flagellomere. Ocellar triangle no longer than wide at base, yellow, wrinkled, black spot on ocellar tubercle small. Palpi black in apical part and a little swollen. Stripes of scutum entirely reddish or central stripe behind and outer marginal stripes brown to black. Central stripe reaches scutellum and scutellum with red or brown mark. Pleura with brownish marks, except small black spot on anepisternum. Abdomen with 3 reddish to brown stripes. Hind femora moderately swollen, no more than 3 times as thick as tibiae. Spring form dark coloured. Ocellar triangle entirely black, or with large square black spot at base and occiput black. Scutal stripes wide and black and sometimes fused, leaving scutum black with small marginal yellow marks. Abdomen black dorsally, only hind margins of tergites narrowly yellow. Legs partly black. Body length 3.0–3.5 mm. Distribution. – Denmark: SJ, WJ, NWJ, NEJ, LFM, NWZ, NEZ. Sweden: along Baltic coast north to NB. Norway: AK, VE. Finland: north to Tb. Russia: Vib, Kr. Holarctic species, in the Palearctic widely distributed from the British Isles to Japan, in the Nearctic confined to western North America.

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.

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Biology. – Adults from May to June and August to September, two generations per year. Larvae develop in different cereals: wheat, barley, rye and many wild grasses: Aegelops sp., Elytrigia repens, E. tenerum, Agropyron cristatum, Lolium perenne, Arrhanatherum elatius, Phleum pratense, Hierochloe odorata, Calamagrostis epigeios (Karpova, 1972; Fedoseeva, 2003). Hibernates as larvae of the third instar within shoots of grasses. Larvae damage shoots and ears of cereals. This species (not M. saltatrix) is a pest of cereals (Fedoseeva, 1969; Fedoseeva & Perfiljeva, 1988; Krusteva, 1995). Reared from cereals (wheat, barley) in Sweden (Johanssen, 1960), Finland (Reuter, 1902a, as M. cerealium) and Norway (Rygg, 1966a, as M. saltatrix var. variegata and M. pratorum var. decora). Some specimens were reared from compost (Werner, 1997). Without doubt larvae occurred there with grass remains. Larvae were described by Kreiter (1929), Tzygankov (1930), Balachowsky & Mesnil (1935), Nishijima (1960), usually under name M. saltatrix. Full descriptions of all three larval instars are given by Fedoseeva (1966). The species is named as M. saltatrix in most applied publications (misidentification).

186. Meromyza ornata (Wiedemann, 1817) Figs 827–834 Oscinis ornata Wiedemann, 1817: 86. Holotype in NHMW (Nartshuk, 1997b). Meromyza sororcula Fedoseeva, 1962: 470. Holotype in ZISP. Ground colour greenish. Body elongated, head longer than deep. Frons produced in front nearly as long as breadth of first flagellomere. Ocellar triangle longer than wide at base, with wrinkled surface and black on ocellar tubercle only. Occiput black. Head setae and setulae predominantly black. Palpi greenish. Stripes of scutum wide, black, densely grey-dusted. Central stripe continuing over scutellum. Katepisternum with pale spot, katepimeron with blackish mark. Abdomen with only central black stripe. Hind femora moderately swollen, 2.5 times as thick as tibiae. Epandrium covered with long pale setulae. Body length to 4.0–4.5 mm. Distribution. – Denmark: widespread. Sweden: north to LU. Norway: AK: Enebakk, Oslo; VE: Tønsberg. Finland: north to ObN. Russia: Vib, Kr. 210

Eurasian species, ranging from the British Isles to Yakutia, East Siberia and Mongolia. Biology. – Adults from end of June to August in wet meadows. Larvae live in shoots of Deschampsia caespitosa. Larvae of the first instar hibernate and in spring move into another plant to continue development. Larvae of all three instars are described by Fedoseeva (1966, as M. sororcula).

187. Meromyza palposa Fedoseeva, 1960 Figs 835–839 Meromyza palposa Fedoseeva, 1960: 452. Syntype in ZISP. Ground colour yellow. Frons slightly produced in front. Ocellar triangle as long as wide at base or little longer, with indistinct apical part, with large black spot on ocellar tubercle. First flagellomere as wide as long. Palpi black, rather broad. Genae with pale setulae. Stripes of scutum black, densely grey-dusted or central stripe brownish in the middle. Central stripe reaches scutellum and scutellum with black or brown mark. Abdomen with 3 wide black stripes. Hind femora slightly swollen, no more than in 2 times as thick as tibiae. Surstyli broad, densely covered with short setulae, gonites rather small. Body length to 2.5–3.5 mm. Distribution. – Denmark: WJ, NWJ, NEJ, B. Sweden: north to NB along Baltic coast. Norway: present, but district uncertain. Finland: north to Obs along Baltic coast. Russia: Vib. European species. Biology. – Adults in June to August in dry meadows. Larvae develop in shoots of Nardus stricta (Fedoseeva, 2003).

188. Meromyza pluriseta Péterfi, 1961 Figs 840–844 Meromyza pluriseta Péterfi, 1961: 167. Type depository unknown. Ground colour fuscous yellow. Frons produced in front as long as 1/2 breadth of first flagellomere. Ocellar triangle as long as wide at base or a little longer, with dark wrinkled rhomboid spot in apical part, which connects with extended large black spot of ocellar tubercle in dark specimens. In light

Figs 827–839. Meromyza ornata (Wiedemann): 827: head, dorsal view. 828: head, lateral view. 829: hypopygium, dorsal view. 830: hypopygium, 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: basiphallus.

211

Figs 840–851. Meromyza pluriseta Fedoseeva: 840: hypopygium, dorsal view. 841: hypopygium, lateral view. 842: postgonite. 843: postgonite, 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 small. Palpi black except yellow at base. Stripes of scutum wide black, densely grey-dusted or lateral stripes brownish in front. Central stripe reaching scutellum and scutellum with black or brown mark. Pleura with large black marks. Abdomen with 3 wide black stripes, sometimes partly fused. Hind femora 3 times as thick as tibiae. Front femora darkened. Male genitalia: surstyli broad, densely covered with short setulae. Gonites small and light. Body length 3.5–4.0 mm. Distribution. – Denmark: NWJ, NWZ, NEZ, B. Sweden: SK, ÖL. Norway: On: Dovre. Finland: north to ObS. Russia: Vib, Kr, Lr. Eurasian species ranging from the British Isles to Yakutia, East Siberia and Mongolia. Biology. – Adults in June to August in meadows. Larvae develop in shoots of Hierochloe odorata, Elytrigia repens (Karpova, 1972; Fedoseeva, 2003) and probably other grasses.

189. Meromyza pratorum Meigen, 1830 Figs 5, 845–851, 1005, 1006 Meromyza pratorum Meigen, 1830: 165; Meigen’s plate 162: 11 (Morge, 1976a). Lectotype female and paralectotype female in MNHN (Nartshuk, 2006). Ground colour greenish. Body elongated, tip of abdomen reaching well behind tip of hind femora. Head longer than deep. Frons produced in front as long as breadth of first flagellomere. Ocellar triangle longer than wide at base, with wrinkled surface and black on ocellar tubercle only. Palpi greenish. Stripes of scutum black, densely grey-dusted. Central stripe reaching scutellum and scutellum with black mark. Katepimeron with yellow mark. Abdomen with only central black stripe. Hind femora moderately swollen, no more than 2.5 times as thick as tibiae. Hypandrium covered with long hairs. Phallus swollen at base. Body length 4.0– 6.0 mm. Distribution. – Denmark: widespread. Sweden: north to NB. Norway: Ø, AK, VE, Fn. Finland: north to ObN. Russia: Vib, Kr. Holarctic species: widely distributed from the British Isles to Japan, and also in western North America.

Biology. – Adults in June to August in dry meadows and sandy, coastal habitats. Larvae develop in shoots of Calamagrostis epigeios, C. neglecta, Ammophila arenaria and Hierochloe odorata (Nartshuk, 1962a; Karpova, 1972; Huiskes, 1979). One generation per year. Females lay eggs on leaves. Larvae damage upper meristem and central leaves. Larvae described by Balachowsky & Mesnil (1935).

190. Meromyza rohdendorfiji Fedoseeva, 1974 Figs 852–856 Meromyza rohdendorfiji Fedoseeva, 1974: 916. Holotype in ZISP. Ground colour brownish yellow. Frons produced in front no more than 1/2 breadth of first flagellomere. Ocellar triangle longer than wide at base, shining, wrinkled in apical part and black on ocellar tubercle only. Palpi yellow, slightly widened at tip. Stripes of scutum densely grey-dusted, central one reddish brown, inner and outer lateral stripes black. Central stripe reaching scutellum and scutellum with black or brown mark. Pleura with red marks, except black mark on anepisternum. Abdomen with central black stripe and small spots on tergites 2–5. Hind femora strongly swollen, nearly 4 times as thick as tibiae. Body length 3.5–4.0 mm. Distribution. – Denmark: widespread. Sweden: north to UP. Norway: Ø: Larkollen (Siebke); VAy: Lindesnes, Spangereid. Finland: Ab: Dragsfjärd, Nagu. European species. Biology. – Adults in July to August, in meadows.

191. Meromyza saltatrix (Linnaeus, 1761) Figs 79, 857–865, 1013, 1014 Musca saltatrix Linnaeus, 1761: 555. The only specimen in Coll. Linnaeus, Linnean Society, London, under Musca saltatrix is a female of Chlorops pumilionis (Bjerkander), but Ismay (1981a) found it questionable that this is an original specimen, as it disagrees with Linnaeus’ 1761description saying “Femora postica crassa”, a character omitted in the 1767-edition of his Systema Naturae. An application should be made to the ICZN for permission to designate of a neotype 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, ventral 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.

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usage of the genus Meromyza that has Musca saltatrix Linnaeus as type species.

genitalia: postgonites light-coloured, elongated and curved. Body length 3.0–3.5 mm.

Ground colour greenish. Frons produced in front by no more than half width of first flagellomere. Ocellar triangle longer than wide at base, with wrinkled surface and black on ocellar tubercle only. Occiput darkened. Head setae and setulae black. Palpi blackened in apical half. Stripes of scutum black, densely grey-dusted. Central stripe reaching scutellum and scutellum with black mark. Pleural marks reddish to black, spot on katepisternum dusted. Abdomen with three black stripes. Hind femora 3 times as thick as tibiae. Male genitalia: postgonites black, large triangular with tip of posterior process depressed. Phallus swollen at base. Body length 3.5–4.0 mm.

Distribution. – Only Sweden: 1 male without exact locality. Finland: Sb: Maaninka (Tuovilankas), 1 male. The species is rather common in Palearctic Asia, but rare in Europe. A few specimens are known from Austria, Germany, Poland, Estonia, Lithuania and European part of Russia.

Distribution. – Very common species occurring throughout Denmark and Fennoscandia. Holarctic: widely distributed trans-Palearctic and western Nearctic species.

Ground colour yellowish or light greenish. Frons produced in front less than 1/2 breadth of first flagellomere. First flagellomere little longer than deep, darkened above. Ocellar triangle as long as wide at base or little longer, with large black spot on ocellar tubercle and darkened side margins. Palpi black an apical half, narrow. Genae with black setulae on lower margin. Stripes of scutum black, densely grey-dusted, central stripe narrowed behind and reaching scutellum, and scutellum with black mark. Abdomen usually with 3 black stripes, rarely only one stripe present. Hind femora rather thin, nearly 2 times as thick as tibiae. Male genitalia: postgonites rather small, black. Body length 2.5–3.0 mm.

Biology. – Adults from June to August, in meadows. Larvae phytophagous. As host plants were recorded Poa pratensis, Elytrigia repens and Festuca rubra (Fedoseeva, 1966, 2003). Hibernates as larvae of third instar. Larvae of all three instars described by Fedoseeva (1966). Earlier authors frequently recorded the present species as a pest of cereals, but these records are all based on misidentifications of M. nigriventris (Fedoseeva, 1969; Fedoseeva & Perfiljeva, 1988).

192. Meromyza sibirica Fedoseeva, 1961 Figs 866–870 Meromyza sibirica Fedoseeva, 1961: 706. Paratypes in ZISP. Ground colour yellow. Frons slightly produced in front. Ocellar triangle little longer than wide at base, yellow, wrinkled at tip. Black spot on ocellar tubercle small. First flagellomere a little longer than deep, darkened above. Palpi black and slightly widened on apical half. Stripes of scutum wide, black or dark brown laterally, densely grey-dusted. Central stripe narrowed behind and does not reach scutellum, and scutellum yellow, sometimes with narrow fuscous yellow stripe. Abdomen usually with one black stripe, but sometimes with some lateral spots on tergites. Hind femora rather thick, more than 3 times as thick as tibiae. Male

193. Meromyza triangulina Fedoseeva, 1960 Figs 871–875 Meromyza triangulina Fedoseeva, 1960: 457. Syntype in ZISP.

Distribution. – Denmark: all districts except SJ and EJ. Sweden: north to NB along Baltic coast. Norway: AK: Ryenberg, 22.vi.1850 (Siebke). Finland: Al, Ab, N, St. Russia: Vib, Kr. Widespread in Europe. Biology. – Adults in June to July, and again in September in meadows, probably two generations. Larvae live in shoots of Dactylis glomerata, Festuca ovina, F. rubra (Fedoseeva, 1966, 2003). Hibernates as larvae of the third instar. Larvae of the third instar described by Fedoseeva (1966) based on puparium.

194. Meromyza variegata Meigen, 1830 Figs 876–881, 1007–1009 Meromyza variegata Meigen, 1830: 165. Syntype in 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: postgonite. 874: postgonite, another specimen. 875: base of basiphallus.

Meromyza lidiae Nartshuk, 1992: 126. Holotype and some paratypes in ZISP, other paratypes in NHMH (Nartshuk, 2006). Ground colour yellowish. Frons produced in front no more than 1/2 breadth of first flagellomere. Ocellar triangle shining, rugose in apical part and black on ocellar tubercle only. Palpi black on apical part. Stripes of scutum black, densely greydusted or central one reddish brown. Central stripe ends before scutellum and scutellum yellow. Pleura with brown to black marks. Abdomen with central black stripe and small spots on tergites 3–5. Hind femora moderately swollen, 2.5 times as thick as tibiae. Aedeagus with bifurcate process at base. Body length 3.5–4.0 mm. 216

Distribution. – Denmark: SJ, NWJ, F, LFM, NEZ. Sweden: SK. Finland: Ab, N. Russia: Vib. Not found in Norway. European species distributed southwards to Bulgaria. Biology. – Adults in July to August in dry meadows. Proposed host plant Phleum phleoides.

195. Meromyza zimzerla Nartshuk, 1992 Figs 882–885 Meromyza zimzerla Nartshuk, 1992: 135. Holotype and some paratypes in ZISP, remaining paratypes in NHMH.

Figs 876–885. Meromyza variegata Meigen: 876: hypopygium, dorsal view. 877: hypopygium, lateral view. 878: postgonite. 879: basiphallus, 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 more than half width of first flagellomere. Ocellar triangle shining wrinkled in apical part, ocellar tubercle with black spot. Palpi yellow. Stripes of scutum black to reddish brown, densely grey-dusted. Central stripe not reaching scutellum, and scutellum yellow. Pleura with red mark on katepisternum and brown mark on katepimeron. Abdomen usually with 3 black or brown stripes. Hind femora 3 times as thick as tibiae. Male genitalia: postgonites

rather large and black. Surstyli long. Body length 3.5–4.0 mm. Distribution. – Denmark: WJ, NEJ. Sweden: SK, SM, ÖL, NÄ. Norway: AK: Drøbak (Siebke). Finland: Al, N, Ka. Not found in Russian parts of Fennoscandia. European species. Biology. – Adults in July to August, in dry meadows with Phleum phleoides, probably its host plant.

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Genus Neohaplegis Beschovski, 1981 Neohaplegis Beschovski, 1981a: 51. Type species: Oscinis tarsata Fallén, 1820 by original designation. Species with entirely black body. Ocellar triangle large and shining. vte and vti setae subequal in size. First flagellomere of antennae rounded. Tibial organ present. Male genitalia: surstyli short, granulated; anus flanked by a pair setulose sclerites; distiphallus simple; testes globular, not tubular as in Cryptonevra and other genera of Chloropinae. Distribution. – Palearctic Region.

a broadly rounded apex. Genae narrower than first flagellomere. Scutum dull, covered posteriorly with grey dust not or just reaching level of anterior npl setae. Legs black with yellow to black trochanters. Body length 2.0–2.5 mm. Distribution. – Denmark: NEZ: without further data (R.C. Stæger), 1 male, 1 female; Utterslev Mose, 17.vi.1906 (W. Lundbeck), 1 female. Sweden: north to NB. Finland: north to Om: Pietarsaari. Russia: Vib, Kr. Not so far found in Norway. Trans-Palearctic species. Biology. – Adults in June to July in wetlands with Carex. Larvae live in shoots of Carex (Nartshuk, 1972a).

Key to species of Neohaplegis 1

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Ocellar triangle smooth and shining, with straight side margins and acute apex. Legs black with brown to black trochanters. Male genitalia Figs 886–889 . . . . . N. glabra (Duda) Ocellar triangle subshining, with convex side margins and a rounded apex. Legs black with yellow to black trochanters. Male genitalia Figs 890, 891 . . . . . . . . . . . . N. tarsata (Fallén)

196. Neohaplegis glabra (Duda, 1933) Figs 886–889 Haplegis glabra Duda, 1933: 131, as var. of H. tarsata (Fallén). Lectotype and paratype in ZMHU (Ismay, 1994a). Body black. Frons dull. Ocellar triangle smooth and shining, with straight side margins and acute apex. Genae narrower than first flagellomere. Scutum subshining, rather densely setulose, covered with brown dust posteriorly extending beyond anterior npl setae. Legs black with brown to black trochanters. Body length 2.0–2.5 mm. Distribution. – Found only in Sweden: ÖL. Distributed in Europe and Kasakhstan.

197. Neohaplegis tarsata (Fallén, 1820) Figs 890, 891 Oscinis tarsata Fallén, 1820: 5. Lectotype in ZMLU (Andersson, 1963). Body black. Frons dull. Ocellar triangle subshining, with convex side margins and anteriorly ending in 218

Genus Parectecephala Becker, 1910 Parectecephala Becker, 1910: 105. Type species: Oscinis longicornis Fallén, 1820 by designation of Duda, 1933: 208. Body of Chlorops-type with elongated first flagellomere and velvety black spot in anterior part of inner lateral stripes. Head higher than long. Ocellar triangle large, reaching front margin of frons. First flagellomere squarish, longer than wide. Arista white pubescent. Scutum yellow with black stripes, smoothly dusted. Scutellum rounded, convex. Wings of Chlorops-type. Tibial organ absent. Male genitalia: middle lobe of surstyli with small, flat apical part; pre- and postgonites parallel to each other. Distribution. – Worldwide.

198. Parectecephala longicornis (Fallén, 1820) Figs 20, 892–898 Oscinis longicornis Fallén, 1820: 4, as var. of O. lineata (Fabricius). Lectotype in ZMLU (Andersson, 1963, 1966). Andersson referred the name longicornis to Zetterstedt and accordingly designated a lectotype from Zetterstedt’s collection. All specimens from Fallén’s collection belong to other species (Andersson, 1963: 104). However, current usage refers the name to Fallén. Frons shorter than wide, yellow along eye margins and brown near ocellar triangle. Ocellar triangle reaching front margin of frons, narrowed anteriorly, yellow or brown with large black shiny spot.

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 flagellomere. First flagellomere 1.5 times longer than wide, yellow, darkened above. Arista white, thickened. Palpi black. Thorax yellow or slightly reddish. Scutum with 5 black, grey-dusted stripes. Lateral stripes anteriorly with round velvety spot. Scutellum yellow with median black stripe. Pleura with black marks, spot on katepisternum sometimes reddish. Anepisternum with setulae below. Abdomen brown. Legs mainly yellow, femora with black stripes. Body length 2.5–3.0 mm. Distribution. – Denmark: widespread. Sweden: SK, HA, ÖL. Not found in Norway, Finland or Russian parts of Fennoscandia. Central and South Europe, Near East and Asia. Biology. – Adults from the end of July to September in dry, sandy meadows. Tschirnhaus (1981) suggested Corynephorus canescens, Carex arenaria or Ammophila arenaria as possible host plants.

Genus Platycephala Fallén, 1820

ifrons Fabricius, 1798) by designation of Curtis, 1839: 725. Body brown, elongated, hind femora thickened. Head longer than high. Frons strongly produced for at least half length of eyes. Eyes with horizontal long axis. Ocellar triangle large, occupying almost entire frons, broadly reaching and semicircular at front margin of frons. Surface of frons covered with numerous setulae. Face sloping backwards, more or less keel-shaped. Antennae prolonged, first flagellomere longer than wide, sometimes narrowed in apical part. Scutum flat, shining with brown or red stripes and covered with numerous setulae. Scutellum oval, flat, apical setae present or absent. Pleura extensively haired. Middle tibiae with 1–3 apical spurs. Hind femora thickened, ventrally in apical part with a ridge covered with granules. Hind tibiae correspondingly curved. Male genitalia: middle lobe of surstyli flat, squarish or triangular; pregonites slender, postgonites short. Distribution. – Palearctic and Oriental Regions.

Platycephala Fallén, 1820: 2. Type species: Platycephala culmorum Fallén, 1820 (= Musca plan219

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

Key to species of Platycephala Fallén 1

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First flagellomere longer and much narrower at tip than at base (Fig. 907). Frons without punctures. Male genitalia Fig. 908 . . . . . . . . . . . . . . . . P. umbraculata (Fabricius) First flagellomere shorter and slightly narrower at tip than at base (Fig. 900). Frons with black punctures. Male genitalia Fig. 905 . . . . . . . . . . . . . . . . . . . P. planifrons (Fabricius)

Species key to known 3rd instar larvae of Platycephala Larvae 6–8 mm long, body cylindrical, milky white, with or without a small sclerotization on the first thoracic segment. Cephalopharyngeal skeleton black, hypostomal and tentoropharyngeal segments fused. Mandibles with accessory teeth. Anterior spiracles transverse, of rosette type. Posterior spiracles sessile on surface of 8th abdominal segment, with interspiracular branched processes.

Body with numerous spicules dorsally or dorsally and ventrally. 1

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Abdominal segments dorsally and ventrally with several rows of sclerotized spicules (Fig. 909). Spicules different in size, some of them large and hook like (Fig. 914). Mandibles with 3 large accessory teeth in each row (Fig. 911) . . . . . . . . . . . . . . . . . . . P. planifrons (Fabricius) Abdominal segments only dorsally with several rows of sclerotized spicules (Fig. 915). Spicules uniformly small (Fig. 917). Mandibles with small accessory teeth only (Fig. 916) . . . . . . . . . . . . . . . . P. umbraculata (Fabricius)

199. Platycephala planifrons (Fabricius, 1798) Figs 7a, 899–905, 909–914 Musca planifrons Fabricius, 1798: 565. Type lost, in Coll. Fabricius, ZMUC, only a name label (Zimsen, 1964). Body yellowish brown. Frons longer than wide, produced anterior to eyes for 1/2 length of eye. Face with a median carina. Ocellar triangle large, occupying entire frons, semicircular anteriorly, with median sulcus and covered with black punctures each inserted by a small setula. First flagellomere about 2 times longer than deep, rounded apically. Apical segment of arista distinctly pubescent, white. Genae roughly wrinkled, behind as wide as eye. vi setae absent. Scutum with three dark brown stripes, roughly punctured and separated by pale unpunctured stripes along dorsocentral lines. Scutellum nearly squarish, apical setae absent. Pleura with coarse black punctures and pale setulae. Abdomen flattened, long and narrow, dusted and with pale setulae. Body length 6–8 mm.

Grochowska, 1986, 1994, 1995). Only one generation per year. Grochowska (1986, 1994) described the life-cycle of the species in Poland, Skuhravý & Skuhravá (1978) reported than the species Hibernates as adults in galls of Lipara in former Czechoslovakia. Larvae of Calamoncosis aprica live as inquilines in stems of reed infested by P. planifrons (Grochowska, 2002). Some larval parts described by Wandolleck (1899) and Nartshuk (1987), egg, all larval instars and pupa by Grochowska (1994).

200. Platycephala umbraculata (Fabricius, 1794) Figs 906–908, 915–917 Musca umbraculata Fabricius, 1794: 348. Type lost, in Coll. Fabricius, ZMUC, only a name label (Zimsen, 1964). Platycephala nigra Meigen, 1830: 26 (Nartshuk, 1997a). Body yellowish brown. Frons as long as wide, slightly produced anteriorly. Face with a median carina. Ocellar triangle large, occupying nearly entire frons, but side margins slightly incurved, with shallow median sulcus, otherwise smooth with fine punctures each inserted by a small setula. First flagellomere 3 times longer than deep, narrowed at apex. Apical segment of arista distinctly pubescent, white. Genae roughly wrinkled, behind as wide as eye. vi absent. Scutum shining, with brownish stripes, roughly punctured. Thoracic setae longer than in P. planifrons. Scutellum rounded behind with apical setae nearly a half as long as scutellum, subapical setae absent. Pleura with pale punctures. Abdomen flattened, long and narrow. Body length 4–6 mm.

Distribution. – Denmark: all districts except NWJ and B. Sweden: north to HS. Norway: Ø, AK, VE, AAy. Finland: north to ObS. Russia: Vib, Kr. Eurasian species.

Distribution. – Denmark: SJ, EJ, WJ, NEJ, F, LFM, NEZ. Sweden: SK, BL, HA, UP. Norway: Ø, VE. Finland: Al, Ab, N, Ta, Om. Not found in the Russian parts of Fennoscandia. Trans-Palearctic species, known from the British Isles to Far East of Russia. More abundant in Asian part of range.

Biology. – Adults from June to September in beds of Common reed. One generation per year. Larvae phytophagous, living in stems of Common reed (Phragmites australis). They induce little visible stem galls that make the central leaf wither and shorten two internodes of the infested stem (Wandolleck, 1899; Skuhravý & Skuhravá, 1978;

Biology. – Adults in July to August. Less common and more demanding than P. planifrons, in beds of Common reed growing in relatively dry habitats. Larvae phytophagous, living in young shoots of Phragmites australis at or slightly above ground level (Nartshuk, 1987; Grochowska, 1998). 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.

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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 Strobl, 1902 Pseudopachychaeta Strobl, 1902: 500, as subgenus of Diplotoxa Loew, 1863. Type species: Diplotoxa (Pseudopachychaeta) pachycera Strobl, 1902 by monotypy. Small species with procurved veins R2+3 and R4+5 and crossveins close together. Head higher behind than in front. Ocellar triangle shining or thinly dusted. oc very strong, procurved. vte twice as long as vti. poc short, cruciate. 2–3 very strong, recurved orb. Genae widest behind, at middle not as broad as first flagellomere. First flagellomere rounded. Arista thickened or slender. Scutum dusted, yellow with black, sometimes very wide and fused stripes. Tibial organ absent. Male genitalia: surstyli fused with epandrium, coarsely granulate; post- and pregonites fused. Distribution. – Worldwide.

Key to species of Pseudopachychaeta Strobl 1

Scutum with dusted black stripes fused. Ocellar triangle entirely darkened Male genitalia

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2

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Fig. 927. Ovipositor Figs 928, 929 . . . . . . . . . . . . . . . . . . . P. rufijiceps (Zetterstedt) Scutum yellow with 3 black stripes distinctly separated. Ocellar triangle yellow or darkened ......................................2 Female cerci narrower, soft and densely pubescent (Figs 921, 922). Male genitalia Fig. 920 . . . . . . P. approximatonervis (Zetterstedt) Female cerci deeper, compressed, strongly sclerotized and without pubescence (Figs 925, 926). Male genitalia Figs 923, 924 . . . . . . . . . . . . . . . . . . . . . . . P. oscinina (Fallén)

201. Pseudopachychaeta approximatonervis (Zetterstedt, 1848) Figs 918–922 Oscinis approximatonervis Zetterstedt, 1848: 2622. Lectotype in ZMLU (Andersson, 1966). Frons as long as wide, yellow, black haired with two long orb. Ocellar triangle reaching half length of frons followed by linear extension to front margin of frons, yellow or brownish, thinly dusted. Face 223

and genae yellow, genae narrowed in front. All setae on genae of females usually pale. Antennae brown, first flagellomere darkened, arista black. Scutum yellow with 3 black, dusted stripes, median stripe not reaching scutellum. Scutellum yellow. Abdomen yellow to brown, dusted. Legs mainly yellow. Female cerci cylindrical, covered in rather long, dense pubescence.

Known from Europe southwards to Spain (Nartshuk, 2004) and eastwards to Kazakhstan and Yakutia, East Siberia.

Distribution. – Denmark: NWJ, LFM, NEZ. Sweden: north to TO. Finland: north to Le. Russia: Vib, Kr. Not so far found in Norway. Holarctic species.

203. Pseudopachychaeta rufijiceps (Zetterstedt, 1838)

Biology. – Adults in May to June and again in September, probably overwintering. In freshwater marches and other wet habitats. Larvae phytophagous, living in inflorescences of various Cyperacea. Different species of Eriophorum other than E. vaginatum according to Nartshuk (1962a). Tschirnhaus (1981) listed as host plant Schoenoplectus tabernaemontani. The larvae are in North America (Valley et al., 1969) reported from inflorescences of Eleocharis obtusa and E. smalli feeding on unripe seeds.

Oscinis rufijiceps Zetterstedt, 1838: 780. Lectotype in ZMLU (Andersson, 1966).

202. Pseudopachychaeta oscinina (Fallén, 1823) Figs 923–926 Phytomyza oscinina Fallén, 1823: 4. Holotype in ZMLU (Nartshuk & Andersson, 2002). Lasiosina heleocharis Nartshuk, 1964: 321 (Nartshuk & Andersson, 2002). Frons as long as wide, yellow, black haired with two long orb. Ocellar triangle reaching half length of frons followed by linear extension to front margin of frons, yellow or slightly brownish, thinly dusted. Face and genae yellow, genae narrowed in front. Part of setae on genae dark. Antennae brown, first flagellomere strongly darkened, arista black. Scutum yellow with 3 wide black, dusted stripes, median stripe not reaching scutellum. Scutellum yellow. Pleura darkened, spot on katepisternum and postnotum black, dusted. Abdomen brownish. Legs mainly yellow. Female cerci shining, bladelike, laterally compressed, almost without pubescence. Distribution. – Sweden: SK, VG, UP, NB. Finland: Ta. Russia: Vib. Not so far found in Denmark and Norway. 224

Biology. – Adults from the end of May to July, found in wet habitats. Larvae phytophagous, living in inflorescences of Eleocharis spp. (Nartshuk, 1964).

Figs 927–929, 981

Frons as long as wide, yellow, black haired with two long orb. Ocellar triangle reaching half length of frons followed by linear extension to front margin of frons, brownish to black, dusted. Face and genae yellow to brownish. Genae narrowed in front. Antennae brown, first flagellomere darkened, arista black. Scutum yellow with 3 black, dusted fused stripes, median stripe reaching scutellum. In dark coloured specimens scutal stripes more or less confluent with yellowish patches only on notopleura and anterior part of postpronota. Scutellum yellow to brownish. Abdomen and legs darkened. Female cerci cylindrical, covered with setulae. Distribution. – The most abundant species of the genus, especially to the north. Sweden: north to TO. Norway: Bv, HOy, HOi, Fn. Finland: north to Li. Russia: Vib, Kr. Lr. Not from Denmark. Trans-Palearctic arcto-boreal species, ranging from northern parts of the British Islands to the Chukotsky Peninsula, Far East Russia. Biology. – Overwintered adults appear in spring, new generation active from July to September (Nartshuk, 1962a). Tyrphobiont, mainly in raised bogs. Females lay eggs in early spring on inflorescences of Eriophorum vaginatum and other species of Eriophorum (e.g. E. polystachyum, E. latifolium, but not on the delicate E. gracilis). Larvae phytophagous, living 4–6 larvae together in one inflorescence, feeding on the developing seeds. They pupate within the inflorescence. Tentoropharyngeal sclerites and pupa are illustrated by Nartshuk (1987).

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.

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Genus Thaumatomyia Zenker, 1833 Thaumatomyia Zenker, 1833: 344. Type species: Thaumatomyia prodigiosa Zenker, 1833 (= Chlorops notatus Meigen, 1830) by monotypy. Chlorops-like yellow species with black or red stripes on scutum, flattened scutellum with approximated apical scutellar setae. Frons somewhat projecting, as broad as long. Ocellar triangle large with straight or convex sides, reaching or almost reaching front margin of frons, with one or more rows of setulae. First flagellomere rounded, usually longer than wide. Scutum rather convex with 5 black or red stripes, smooth and shining, almost bare or covered with short setulae. Scutellum rounded or somewhat triangular, flat, bare or with numerous setulae. Apical scutellar setae close together. Tibial organ present. Middle lobe of surstyli with nail- or hook-shaped apex; pre- and postgonites situated in line. Males of some species have eversible vesicles in the intersegmental membrane of the postabdomen that become inflated during the premating period. These vesicles probably emit short-range sex pheromones. Distribution. – Worldwide.

Key to species of Thaumatomyia Zenker 1

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Genae narrow, no more than half width of first flagellomere; ocellar triangle with two rows of setulae inside lateral margins (Figs 939, 940). Male abdomen with two lateral membranous vesicules seen usually in macerated abdomen (Fig. 70) . . . . . . . . . . . . . . . . T. notata (Meigen) Genae as wide as or wider than first flagellomere; ocellar triangle with one row of setulae inside lateral margins (Figs 930, 931, 944, 945, 948, 949) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Ocellar triangle carinate in apical part; genae very wide, wider than first flagellomere (Figs 948, 949). Male genitalia Figs 950, 951 . . . . . . . . . . . . . . . . . T. trifasciata (Zetterstedt) Ocellar triangle without carina in apical part, genae not wider than first flagellomere (Figs 930, 931, 944, 945) . . . . . . . . . . . . . . . . . . . . . . 3 Scutum and scutellum with sparse setulae. Scutum with black, shining stripes, middle stripe reaching scutellum. Male abdomen

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with arched membranous vesicula seen best in macerated abdomen (Fig. 69) . . . . . . . . . . . . . . . . . . . . . . . T. glabra (Meigen) Scutum and scutellum distinctly haired. Scutum with black or red stripes, middle stripe usually ending before scutellum. Male abdomen without membranous vesicula . . . . . 4 Body yellow with black or red stripes on scutum. Male genitalia Figs 946, 947 . . . . . . . . . . . . . . . . . . . . . . . T. rufa (Macquart) Body darkened with black stripes. Male genitalia Figs 937, 938 . . . T. hallandica Andersson

204. Thaumatomyia glabra (Meigen, 1830) Figs 69, 930–936, 960–962 Chlorops glabra Meigen, 1830: 149; Meigen’s plates 162: 4, 211: 18 (Morge, 1976b). Lectotype in NHMW (Nartshuk, 1997a). Frons slightly longer than wide. Ocellar triangle large, mainly black, linearly yellow at side margins and posterolateral corners. Genae very wide. First flagellomere about 1.2 times as long as wide, distally brown, basally yellow. Only vte setae distinct on head. Scutum glabrous with three broad black stripes, all reaching scutellum. Setulae covering scutum minute. Scutellum flat, nearly bare, apical setae black, approximated. Abdomen black dorsally. Male postabdomen with large eversible vesicles. Body length 2.3–2.6 mm. Distribution. – Common species. Denmark: found in all districts except SZ. Sweden: north to TO. Norway: Ø, AK, HEs, Bv, TEy, VAy, Ry, HOy, HOi, SFy, STi, NTi. Finland: north to Le. Russia: Vib, Kr, Lr. Holarctic species. Biology. – Adults from June to September, abundant everywhere, found in a variety of habitats including freshwater marshes, very often on flowering Medicago (Fabaceae). Aggregating males have been observed on the inflorescence of Wisteria sinensis in California, with single males occupying an individual blossom (Sorensen & Sorensen, 1997). Steyskal (1945) described the behavior of a North American subspecies of this species (T. glabra var. bistriata Walker). He observed that males with exposed terminal vesicles would walk about on sugar maple leaves fluttering their wings and swelling their vesicles. On approaching another (male?) individual they would go infight us-

Figs 930–938. Thaumatomyia glabra (Meigen): 930: head, dorsal view. 931: head, lateral view. 932: antenna. 933: wing. 934: male genitalia, 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 eventually one of them would be pushed over the edge of the leaf and forced to escape. It was found that the males were attracted by methyl anthranilate (Landolt et al., 2000). Sabrosky (1935) mentioned that this species was taken in a light trap, although chloropids are seldom taken at light. Larvae carnivorous, living among roots of plants feeding on root aphids, especially Pemphigidae, as reported by Parker (1918) and in many subsequent publications, see Nartshuk (1972b). There is one record of the species from a spider egg sac (Sabrosky, 1943). It has been reared from compost (Werner, 1997) that no doubt contained larvae imported with plant material. Larvae described by Balachowsky & Mesnil (1935), puparium figured by Nartshuk (1987).

205. Thaumatomyia hallandica Andersson, 1966 Figs 937, 938 Thaumatomyia hallandica Andersson, 1966: 63. Holotype in ZMLU. Body generally infuscate. Frons slightly longer than wide, dull. Ocellar triangle large, almost reaching front margin of frons, shining, brownish black, with straight side margins. Genae a little wider than first flagellomere. First flagellomere about 1.2 times as long as wide, black or basally yellow. Arista brown, shortly pubescent. vte strong. Palpi yellow to brown. Scutum glabrous with 5 broad black stripes, middle one ending at middle of scutum. Scutellum flat, apical setae black, approximated. Abdomen brown dorsally. Male without postabdominal 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 except SZ and NWZ. Sweden: north to LU. Norway: north to Nnø. Finland: north to Le. Russia: Vib, Kr. Eurasian species, ranging from the British Isles to Mongolia and Central Asia. Biology. – Adults from June to August in dry meadows and sand dunes. Larvae carnivorous, living among roots of plants feeding on root aphids.

206. Thaumatomyia notata (Meigen, 1830) Figs 7b, 70, 110–114, 939–943 Chlorops notata Meigen, 1830: 144; Meigen’s plate 211: 17 (Morge, 1976). Lectotype in NHMW (Nartshuk, 1997a). Frons nearly as long as wide. Ocellar triangle from black to yellow with median black stripes of dif228

ferent width, with 2–3 irregular rows of setulae along side margins. Genae narrow, nearly 1/3 of first flagellomere. First flagellomere nearly orbicular or sometimes very slightly wider than long, almost entirely black or brownish basally. vte and vti setae both developed and rather strong. Scutum with 5 black, rarely partly reddish, stripes not reaching scutellum. Scutellum flat, apical setae black, approximated. Pleura with black or red spot on katepisternum. Male with pair of eversible postabdominal vesicles. Body length 2.5–3.0 mm Distribution. – Very common. Denmark: SJ, LFM, NEZ. Sweden: north to NB along the Baltic coast. Norway: AK, Os, HOi. Finland: north to Om. Russia: Vib, Kr. Widespread species, recorded from the Palearctic, Afrotropical and Oriental Regions. Palearctic distribution from the British Isles to Japan.

Biology. – Adults from May to October, hibernate, found in a variety of habitats, often visit flowers. In Norway found by canopy fogging of Pinus silvestris (Thunes et al., 2004). In Switzerland 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 between scales of Picea cones for overwintering (Krivokhatsky & Nartshuk, 2001). Occationally, several millions of specimens enter houses in the autumn for overwintering. This has repeatedly been recorded from various parts of Europe including the Nordic countries: Finland (Haartman, 1950) and Sweden (T. Pape, pers. comm.). Lampa (1893) described 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), undoubtedly from larvae imported with roots of plants. Larvae described by Balachowsky & Mesnil (1935), some larval structures illustrated by Nartshuk (1987).

207. Thaumatomyia rufa (Macquart, 1835) Figs 944–947 Chlorops rufa Macquart, 1835: 593. Type probably in MHNL. Frons slightly longer than wide. Ocellar triangle entirely brownish yellow to blackish, reaching front margin of frons, with one row of setulae along slightly concave side margins. Genae nearly as wide as first flagellomere. First flagellomere distinctly longer than wide, mainly yellow, darkened dorsally. Head only with distinct vte setae. Scutum with 5 stripes not reaching scutellum. Stripes black or sometimes reddish brown. Spot on anepisternum black, remaining pleural spots usually reddish brown. Male without postabdominal vesicles. Body length 2.0–2.5 mm. Distribution. – Denmark: SJ, WJ, F, LFM, NEZ, B. Sweden: north to TO. Norway: O, VE, HOi. Finland: north to Om. Russia: Vib, Kr. Trans-Palearctic species, known from the British Isles to Japan and southwards to North Africa.

Figs 948–951. Thaumatomyia trifasciata (Zetterstedt): 948: head, dorsal view. 949: head, lateral view. 950: surstylus. 951: hypandrium and phallic complex.

Biology. – Adults from middle of June to August, in meadows. Larvae carnivorous, living in the ground among roots of plants where feeding on root aphids. It has been reared from compost (Werner, 1997). Anterior spiracles of larvae illustrated by Nartshuk (1987).

208. Thaumatomyia trifasciata (Zetterstedt, 1848) Figs 948–951 Oscinis trifasciata Zetterstedt 1848: 2609. Lectotype in ZMLU (Andersson, 1966). Frons slightly longer than wide. Ocellar triangle usually black, extending nearly to front margin of frons, with one row of setulae along side margins and on anterior half of central ridge. Genae 1.5–2.0 times wider than first flagellomere is deep. First flagellomere slightly longer than deep, mainly yellow, darkened dorsally. vte setae absent. Scutum with 5 black stripes, central one not reaching scutellum. Pleural spots black. Male without postabdominal vesicles. Body length 3.5–4.0 mm. Distribution. – Widespread in Denmark and Fennoscandia. 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 places such as bogs, marshes, swamps, fens and littoral zone of lakes. In Norway found by canopy fogging of Pinus silvestris (Thunes et al., 2004). Larvae as in other species of the genus carnivorous, feeding on root aphids among roots of plants. Meijere (1950) found larvae among roots of Cichorium intybus (Asteraceae) in the Netherlands. A. Przhiboro (pers. comm.) reared the species from plant roots growing at the littoral zone of the White Sea.

Genus Trichieurina Duda, 1933 Trichieurina Duda, 1933: 126. Type species: Eurina pubescens Meigen, 1830 by original designation. Greyish species with strongly produced frons. Body covered with very thick dust and extremely long slender hairs. Head triangular in side view. Eyes small. Genae very wide. Ocellar triangle reaching front margin of frons, covered with long hairs. First flagellomere longer than wide, arista nearly bare. Tibial organ absent. Male genitalia: middle lobe of surstyli serrate, postgonites and pregonites distinct, the latter transversally directed. 230

209. Trichieurina pubescens (Meigen, 1830) Figs 952–955 Eurina pubescens Meigen, 1830: 4; Meigen’s plate 209: 4 (Morge, 1976b). Lectotype in NHMW (Nartshuk, 1997a). Densely grey-dusted species, body covered with long hairs. Frons projecting beyond eyes as long as length of an eye. Ocellar triangle reaching front margin of frons, dusted, with a longitudinal median groove and long hairs along sides. Second antennal segment as long as wide, first flagellomere longer than wide; arista dark, basally whitish. Scutum dark grey-dusted with double central line brown, covered with long hairs. Pleura, abdomen and legs completely dusted. Distribution. – Found only in Finland: Kb: Hammaslahti, Pyhäselkä, 15.vi.1935, 04.vi.1938, 07.vii.1939 (Kontkanen), 2 males, 5 females. The species is known from Central and East Europe and Kazakhstan. Biology. – Kontkanen (1935) described in detail the habitat in Finland. The species occurs in swamps with sedges. Carex spp. are likely host plants.

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 Balachowsky & Mésnil, 1935; 964, 965 after Kiauka, 1974; 966 after Kanmiya, 1972.)

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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.)

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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: posterior 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.)

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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: cephalopharyngeal 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.)

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Abbreviations aed: aedeagus ane: anepisternum ant1 , ant2 , ant3 : first–third antennal segment apog: anterior process of postgonite ag: accessory gland bph: basiphallus c: cercus cph: cephalopharyngeal sclerite cx1 , cx2 , cx3 : first–third coxa dph: distiphallus ds: dental sclerite egt: egg tube ej ap: ejaculatory apodeme ep: epandrium ep s: ventral epandrial sclerite f1 , f2 , f3 : first–third femur fc: facial carina fl1: first flagellomere, postpedicel fp: frontal palpi hyp: hypandrium hys: hypostomal sclerite kepml: ketepimerom m: mesonotum md: mandilula (mouth hook) of larva mes: mesolobus, fused cerci mg1 , mg2 , mg3 , mg4 : first–fourth costal segment mn: metanotum mp: maxillary palpi mt1 , mt2 , mt3 : first–third metatarsus npl: notopleuron ph: phallapodeme ph s: phallapodemic sclerite pl: paralobes pog: postgonite pph: phallic paraphyses ppog: posterior process of postgonite prg: pregonite prpl: propleuron psc: pregenital sclerite pst: parastomal sclerite sp: spiracle spt: spermatheca sur: surstylus t1 , t2 , t3 : first–third tibia ts: testes v: vesicula

C: costal vein CuA1 : fifth longitudinal vein M: medial, fourth longitudinal vein R1 : first radial, first longitudinal vein R2+3 : second + third radial, second longitudinal vein R4+5 : fourth + fifth radial, third longitudinal vein Sc: subcostal vein ta: r-m crossvein tp: dm-cu crossvein acr: acrostical setae a npl: anterior notopleural seta a pa: anterior postalar seta ap sc: apical scutellar setae dc: dorsocentral setae fr: frontal setulae h: humeral seta if : interfrontal setulae la sc: lateral scutellar setae oc: ocellar setae orb: orbital setae p npl: posterior notopleural setae p pa: posterior postalar setae pra: prealar seta prpl: propleural seta prs: presutural setae psc: prescutellar setae pst: peristomal setulae pooc: postocellar, postvertical setae sa: supraalar seta sc: scutellar setulae scp: scapular seta s tr: sensillum trichodeum vi: vibrissa vte: outer vertical setae vti: inner vertical setae ANPL: anepimeral spot H: humeral spot KEMSP: katepimeral spot KESSP: katepisternal spot M1 : central scutal (mesonotal) stripe M2 : inner lateral scutal (mesonotal) stripe 235

M3 : outer lateral scutal (mesonotal) stripe MN: metanotal spot PRPL: propleural spot

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PTPL: pteropleural spot S, s: sternite T, t: tergite

Catalogue

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

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Germany Great Britain SJ EJ WJ NWJ NEJ F LFM SZ NWZ NEZ B SK BL HA SM ÖL GO GS ÖG VG BO DS NÄ SÖ UP VS VR DR GA HS ME HR JÄ Conioscinella abiskoi Nartshuk & Andersson Conioscinella frontella (Fallén) Conioscinella gallarum (Duda) Conioscinella livida Nartshuk Conioscinella messaurea Nartshuk & Andersson Conioscinella mimula Collin Conioscinella sordidella (Zetterstedt) Conioscinella tornensis Nartshuk & Andersson Conioscinella zetterstedti Andersson Cryptonevra consimilis Collin Cryptonevra diadema (Meigen) Cryptonevra flavitarsis (Meigen) Dicraeus fennicus Duda Dicraeus ingratus Loew Dicraeus nitidus Wahlgren Dicraeus raptus Haliday Dicraeus rossicus Stackelberg Dicraeus styriacus (Strobl) Dicraeus tibialis Macquart Dicraeus tjederi Nartshuk & Andersson Dicraeus vagans (Meigen) Diplotoxa messoria (Zetterstedt) Diplotoxoides dalmatina (Strobl) Elachiptera brevipennis (Meigen) Elachiptera cornuta (Fallén) Elachiptera diastema Collin Elachiptera scrobiculata (Strobl) Elachiptera tuberculifera (Corti) Epichlorops puncticollis (Zetterstedt) Eribolus danicus Nartshuk & Andersson Eribolus gracilior (Mejiere) Eribolus hungaricus Becker Eribolus nana (Zetterstedt) Eribolus slesvicensis Becker Eurina calva Egger Eurina lurida Meigen Eutropha fulvifrons (Haliday) Eutropha variegata Loew Gampsocera numerata (Heeger) Gaurax borealis Duda Gaurax dubius (Macquart) Gaurax ephippium (Zetterstedt) Gaurax fascipes Becker Gaurax flavomaculatus (Duda) Gaurax fungivorus Nartshuk & Andersson Gaurax leucarista Nartshuk Gaurax macrocerus Nartshuk Gaurax maculipennis (Zetterstedt) Gaurax suecicus Nartshuk & Andersson Hapleginella laevifrons (Loew) Incertella albipalpis (Meigen) Incertella antennata (Collin) Incertella karteshensis Nartshuk & Przhiboro Incertella kerteszi (Becker) Incertella nigrifrons (Duda) Incertella scotica (Collin) Incertella zuercheri (Duda) Lasiambia baliola (Collin) Lasiambia brevibucca (Duda) Lasiambia coxalis (Roser) Lasiambia palposa (Fallén) Lasiosina albipila (Loew) Lasiosina brevisurstylata Dely-Draskovits

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Al Ab N Ka St Ta Sa Oa Tb Sb Kb Om Ok ObS ObN Ks LkW LkE Le Li Vib Kr Lr

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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)

Norway

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Germany Great Britain SJ EJ WJ NWJ NEJ F LFM SZ NWZ NEZ B SK BL HA SM ÖL GO GS ÖG VG BO DS NÄ SÖ UP VS VR DR GA HS ME HR JÄ Lasiosina danielssoni Nartshuk & Andersson Lasiosina herpini (Guérin-Méneville) Lasiosina intermedia Dely-Draskovits Lasiosina parvipennis Duda Lasiosina subnigripes Dely-Draskovits Lipara lucens Meigen Lipara pullitarsis Doskočil & Chvála Lipara rufijitarsis Loew Melanum laterale (Haliday) Meromyza bifurcata Nartshuk & Andersson Meromyza bohemica Fedoseeva Meromyza curvinervis (Zetterstedt) Meromyza elbergi Fedoseeva Meromyza femorata Macquart Meromyza ingrica Nartshuk Meromyza mosquensis Fedoseeva Meromyza nigriseta Fedoseeva Meromyza nigriventris Macquart Meromyza ornata (Wiedemann) Meromyza palposa Fedoseeva Meromyza pluriseta Péterfi Meromyza pratorum Meigen Meromyza rohdendorfiji Fedoseeva Meromyza saltatrix (Linnaeus) Meromyza sibirica Fedoseeva Meromyza triangulina Fedoseeva Meromyza variegata Meigen Meromyza zimzerla Nartshuk Microcercis kroeberi (Duda) Microcercis trigonella (Duda) Neohaplegis glabra (Duda) Neohaplegis tarsata (Fallén) Oscinella angularis Collin Oscinella angustipennis Duda Oscinella capreolus (Haliday) Oscinella cariciphila Collin Oscinella frit s. lat. Oscinella maura (Fallén) Oscinella nitidissima (Meigen) Oscinella norrbottica Nartshuk & Andersson Oscinella pusilla (Meigen) Oscinella trochanterata Collin Oscinella ventricosi Nartshuk Oscinimorpha albisetosa (Duda) Oscinimorpha minutissima (Strobl) Oscinimorpha sordidissima (Strobl) Oscinisoma cognatum (Meigen) Oscinisoma germanicum (Duda) Oscinisoma gilvipes (Loew) Parectecephala longicornis (Zetterstedt) Platycephala planifrons (Fabricius) Platycephala umbraculata (Fabricius) Polyodaspis rufijicornis (Macquart) Polyodaspis sulcicollis (Meigen) Pseudogaurax venustus (Czerny) Pseudopachychaeta approximatonervis (Zetterstedt) Pseudopachychaeta oscinina (Fallén) Pseudopachychaeta rufijiceps (Zetterstedt) Rhopalopterum anthracinum (Meigen) Rhopalopterum atricillum (Zetterstedt) Rhopalopterum atricorne (Zetterstedt) Rhopalopterum brunneipenne Beschovski & Lansbury

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Al Ab N Ka St Ta Sa Oa Tb Sb Kb Om Ok ObS ObN Ks LkW LkE Le Li Vib Kr Lr

Finland

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)

Norway

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Germany Great Britain SJ EJ WJ NWJ NEJ F LFM SZ NWZ NEZ B SK BL HA SM ÖL GO GS ÖG VG BO DS NÄ SÖ UP VS VR DR GA HS ME HR JÄ Rhopalopterum fasciola (Meigen) Rhopalopterum femorale (Collin) Rhopalopterum tomentosum Nartshuk & Andersson Siphonella oscinina (Fallén) Siphunculina aenea (Meigen) Speccafrons halophila (Duda) Thaumatomyia glabra (Meigen) Thaumatomyia hallandica Andersson Thaumatomyia notata (Meigen) Thaumatomyia rufa (Macquart) Thaumatomyia trifasciata (Zetterstedt) Trachysiphonella rufijiceps (Macquart) Trachysiphonella scutellata (Roser) Trichieurina pubescens (Meigen) Tricimba cincta (Meigen) Tricimba humeralis (Loew) Tricimba lineella (Fallén)

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NB: Oscinella frit s. lat. represents a complex of species with an unsettled taxonomy: Oscinella frit (Linnaeus), C. hortensis Collin, C. nigerrima (Macquart) and C. vastator (Curtis).

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Finland

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Norway

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Literature Abraham, R., 1983. Racosina deplanata (Hymenoptera, Pteromalidae) in der Haseldorfer Marsch bei Hamburg. – Entomologische Mitteilungen aus dem Zoologischen Museum Hamburg, 7 (112): 397–398. Abraham, R. & B. Carstensen, 1982. Die Schilfgallen von Lipara-Arten (Diptera: Chloropidae) und ihre Bewohner im Schilf der Haseldorfer Marsch bei Hamburg. – Entomologische Mitteilungen aus dem Zoologischen Museum Hamburg, 7 (116): 269–277. Achterberg, C. van, 1988. Revision of the subfamily Blacinae Förster (Hymenoptera, Braconidae). – Zoologische Verhandlungen, Leiden, 249: 1–324. Achterberg, C. van, 1998. Phaenocarpa helophilae spec. nov. (Hymenoptera: Braconidae), a gregarious parasitoid of Helophilus larvae (Diptera: Syrphidae), with notes on related species. – Zoologische Mededelingen, Leiden, 72: 15–27. Agafonova, Z.J., 1962. [New pests of brome grass, flies of the genus Dicraeus (Diptera, Chloropidae).] – Entomologicheskoe Obozrenie, 41: 402–504. (In Russian.) Agapova, E.G., 1966. [Role of host plants in divergence of Oscinella flies.] – Trudy Vsesojusnogo Instituta Zhashchity Rasteniy, Leningrad, 26: 185–192. (In Russian.) d’Aguilar, J., 1943. Contribution a l’étude des larves de Chlorops Meig. (Dipt. Chloropidae). – Bulletin de la Société Entomologique de France, 48: 153–155. d’Aguilar, J., 1953. Contribution a l’étude des larves des Diptères (9e Note). La larve de Calamoncosis minima Strobl (Dipt., Chloropidae). – Bulletin de la Société Entomologique de France, 58: 72–76. Ahlberg, O., 1937. En erinran om fritflugeundersökningen. – Meddelanden – Statens Växtskyddsanstalt, 2: 27–28. Åkerman, Å., 1951. Några erfarenheter rörande angrepp av fritflugelarver påhavre. – Sveriges Utsädesförenings Tidskrift, 61: 108–118. Aldrich, J.R. & T.M. Barros, 1995. Chemical attraction of male crab spiders (Aranea, Thiomizidae) and kleptoparasitic flies (Diptera: Milichiidae and Chloropidae). – Journal of Arachnology, 23: 212–214. Alexander, K.N.A., 1983. Eurina lurida Meigen (Dipt., Chloropidae) in Pembrokeshire. – Entomologist’s Monthly Magazine, 119: 90.

Aleksandrov, Yu.P. & N.A. Fedosov, 1971. [Patogenes of frit fly.] – Zhashchita rasteniy, Moskva, 3: 49–50. (In Russian.) Aleksandrov, Yu.P., N.A. Fedosov, N.V. Ushakova & E.V. Yudina, 1969. [Bacterioses of frit fly.] – Zhashchita Rasteniy, Moskva, 8: 52–53. (In Russian.) Allen, A.A., 1981a. The two British species of Lasiambia End. (Dipt., Chloropidae) in SE London. – Entomologist’s Monthly Magazine, 117: 144. Allen, A.A., 1981b. Tricimba lineella (Fall.) (Dipt., Chloropidae) on giant puffball fungus in S.E. London. – Entomologist’s Monthly Magazine, 117: 127. Allen, A.A., 1982. Eurina lurida Meig. (Dipt., Chloropidae) recaptured in N.W. Kent after nearly 80 years. – Entomologist’s Monthly Magazine, 118: 238. Ambrus, B., 1977. A nad-gubacslegy gazdanoveny válogatása (Diptera, Chloropidae). – Folia Entomologica Hungarica, 30 (2): 5–7. Andersen, J., 1988. Postglacial immigration of plants and animals in Scandinavia. – Fauna, Oslo, 41: 1–11. Andersson, H., 1956. Undersökningar rörande vanliga fritflugan Oscinella frit L. – Sveriges Utsädesförenings Tidskrift, 66 (5): 249–280. Andersson, H., 1963. A revision of the species of Oscinides described by C.F. Fallén in Diptera Sueciae (Dipt., Chloropidae). – Opuscula Entomologica, 28: 103–110. Andersson, H., 1966. A revision of the species of Oscinides described by J.W. Zetterstedt (Dipt., Chloropidae). – Opuscula Entomologica, 31 (1–2): 61–76. Andersson, H., 1967. Inventering av förekomst av fritflugor (Diptera: Chloropidae) i stråsäd 1961–1963. [The occurrence of frit flies (Diptera, Chloropidae) in cereal crops in Sweden 1961–1963.] – Meddelande – Statens Växtskyddsanstalt, 14 (115): 1–27. Andersson, H., 1972. En snabb och billig metod att montera genitalpreparat i glycerin. – Entomologen, Lund, 1: 1–2. Andersson, H., 1977. Taxonomic and phylogenetic studies on Chloropidae (Diptera) with special reference to Old World genera. – Entomologica Scandinavica, Supplement, 8: 1–200. Andersson, H., 1979. Problem vid kladistisk analys av flugfamilijen Chloropidae. – Entomologisk Tidskrift, 100: 180–187.

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

abiskoi Nartshuk & Andersson 79 adjunctus Becker 164 aenea Meigen 146 albinervis Roser 146 albipalpis Meigen 110 albipila Loew 196 albisetosa Duda 133 angularis Collin 126 angustipennis Duda 126 antennata Collin 110 anthracinum Meigen 141 anthracophagoides Strobl 164 Aphanotrigonum Duda 61 aprica Meigen 69 approximatonervis Zetterstedt 223 Aspistyla Duda 59 aspistylina Duda 69 atricillum Zetterstedt 142 atricorne Zetterstedt 143 baliola Collin 114 balticus Wahlgren 176 basalis Roser 208 beschovskii Dely-Draskovits 66 bifurcata Nartshuk & Andersson 204 bipunctus Duda 180 bjerkanderi Nartshuk & Andersson 165 bohemica Fedoseeva 204 borealis Duda 102 brachypterum Zetterstedt 61 brevibucca Duda 116 brevimanus Loew 172 brevipennis Meigen 92 brevisurstylata Dely-Draskovits 197 britannicus Deeming 104 brunneipenne Beshovski & Lansbury 143 Calamoncosis Enderlein 68 calceatus Meigen 165

calva Egger 193 Camarota Meigen 154 capreolus Haliday 126 cariciphila Collin 127 Centorisoma Becker 155 centromaculatus Duda 165 cerealium Reuter 208 cereris Fallén 156 Cetema Hendel 155 Chloropinae 154 Chlorops Meigen 159 Chloropsina Becker 185 cincta Meigen 151 cinctellum Zetterstedt 62 cinctipes auct. 198 citrinellus Zetterstedt 180 cognatum Meigen 136 Colliniella Nartshuk & Andersson 77 Conioscinella Duda 78 consimilis Collin 187 cornuta Fallén 92 coxalis Roser 116 crassipalpis Smirnov 174 crassipes Nartshuk 98 Cryptonevra Lioy 186 curvinervis Zetterstedt 205 curvipennis Latreille 154 Cyclocercula Beschovski 124 dalmatina Strobl 191 danicus Nartshuk & Andersson 96 danielssoni Nartshuk & Andersson 197 dasycerus Loew 167 diadema Meigen 187 diastema Collin 94 Dicraeus Loew 85 Diplotoxa Loew 190 Diplotoxoides Andersson 190 discicornis Zetterstedt 172 279

distinguenda Frey 185 dubius Macquart 102 duinensis Strobl 69 Elachiptera Macquart 90 elbergi Fedoseeva 206 elegantulum Becker 155 elongatum Meigen 157 ephippium Zetterstedt 102 Epichlorops Becker 192 Eribolus Becker 96 Eurina Meigen 193 Eutropha Loew 194 fasciatus Meigen 167 fasciella Zetterstedt 61 fasciola Meigen 143 fascipes Becker 104 femorale Collin 143 femorata Macquart 206 femorellum Collin 63 fennicus Duda 86 fennicus Duda 180 figuratus Zetterstedt 167 finitimus Becker 167 flavifrons Duda 83 flavitarsis Meigen 188 flavomaculatus Duda 104 freyi Duda 182 frit Linnaeus 128 frontella Fallén 80 frontosus Meigen 169 fulviceps Roser 172 fulvifrons Haliday 195 fungicola Dely-Draskovits 151 fungivorus Nartshuk & Andersson 106 gallarum Duda 81 Gampsocera Schiner 100 Gaurax Loew 101 geminatus Meigen 169 germanicum Duda 137 gilvipes Loew 137 glabra Duda 218 glabra Meigen 226 glyceriae Nartshuk 70 gotlandica Holmgren 177 gracilior Mejiere 97 gracilis Meigen 169 hallandica Andersson 227 halophila Duda 148 halterata Nartshuk & Andersson 72 280

Hapleginella Duda 108 heleocharis Nartshuk 224 herpini Guérin-Méneville 198 hortensis Collin 128 humeralis Loew 151 hungaricum Dely-Draskovits 63 hungaricus Becker 97 hungaricus Duda 180 hypostigma Meigen 171 Incertella Sabrosky 109 inerme Collin 63 ingratus Loew 86 ingrica Nartshuk 206 inquilina Hendel 109 intermedia Dely-Draskovits 198 interruptus Meigen 171 karteshensis Nartshuk & Przhiboro 110 kerteszi Becker 111 kirigaminensis Kanmiya 171 kroeberi Duda 123 laetus Meigen 172 laevicollis Becker 182 laevifrons Loew 108 Lasiambia Anonymous 114 Lasiambia Enderlein 114 Lasiambia Sabrosky 114 Lasiosina Becker 196 laterale Haliday 200 leucarista Nartshuk 106 limbatus Meigen 172 lineella Fallén 152 Lipara Meigen 118 livida Nartshuk 81 longicornis Zetterstedt 218 lucens Meigen 119 lurida Meigen 193 macrocerus Nartshuk 106 Macrostyla Lioy 59 maculipennis Zetterstedt 107 maura Fallén 129 meigenii Loew 174 meijerei Duda 78 Melanochaeta Bezzi 124 Melanum Becker 200 Meromyza Meigen 200 messaurea Nartshuk & Andersson 81 messoria Zetterstedt 190 Microcercis Beschovski 122 microptera Duda 61

mimula Collin 83 minima Strobl 74 minutissima Strobl 135 mosquensis Fedoseeva 206 myopinum Loew 157 nana Zetterstedt 97 napaeus Collin 86 Nartshukiella Beschovski 149 neglectum Tonnoir 157 Neohaplegis Beschovski 218 nidicola Nartshuk 147 nigerrima Macquart 129 nigra Meigen 221 nigrifrons Duda 112 nigripalpis Duda 174 nigripes Zetterstedt 63 nigriseta Fedoseeva 208 nigriventris Macquart 208 nitidissima Meigen 129 nitidus Wahlgren 86 norrbottica Nartshuk & Andersson 131 norrbotticum Nartshuk & Andersson 65 notata Meigen 228 numerata Heeger 100 obliqua Beschovski 157 obscurellus Zetterstedt 174 Oedesiella Becker 85 oophila Hennig 116 ornata Wiedemann 210 oscinella Becker 75 Oscinella Becker 124 Oscinellinae 61 Oscinimorpha Lioy 133 oscinina Fallén 145 oscinina Fallén 224 Oscinisoma Lioy 135 Pachychaetina Hendel 124 Pachychoeta Bezzi 124 pallidiventris Duda 176 pallifrons Strobl 176 palpatus Smirnov 176 palposa Fallén 116 palposa Fedoseeva 210 pannonicus Strobl 176 Parectecephala Becker 218 Paroedesiella Anonymous 85 Paroedesiella Enderlein 85 Paroedesiella Sabrosky 85 parvipennis Duda 199 planifrons Fabricius 221

planifrons Loew 177 Platycephala Fallén 219 platythorax Nartshuk 143 plumiger Meigen 60 pluriseta Péterfi 210 Polyodaspis Duda 138 pratorum Meigen 213 Pseudogaurax Malloch 139 Pseudopachychaeta Strobl 223 pubescens Meigen 230 pullitarsis Doskočil & Chvála 121 pumilionis Bjerkander 177 puncticollis Zetterstedt 192 pusilla Meigen 131 pygmaeus Meigen 149 raptus Haliday 86 Rhaphiopyga Nartshuk 68 Rhodesiellinae 59 Rhopalopterum Duda 141 ringens Loew 179 riparius Smirnov 179 rohaceki Nartshuk 185 rohdendorfi Fedoseeva 213 rossicus Smirnov 180 rossicus Stackelberg 88 rostrata Hubicka 208 rufa Macquart 229 ruficeps Macquart 149 ruficeps Zetterstedt 224 ruficornis Macquart 138 rufinus Zetterstedt 180 rufitarsis Loew 121 saltatrix Linnaeus 213 scalaris Meigen 180 scotica Collin 113 scrobiculata Strobl 94 scutellaris Zetterstedt 182 scutellata Roser 149 serenus Loew 182 sibirica Fedoseeva 215 simile Ismay 157 similis Schiner 118 Siphonella Macquart 145 Siphunculina Rondani 146 slesvicensis Becker 98 sordidella Zetterstedt 83 sordidissima Strobl 135 sororcula Fedoseeva 210 Speccafrons Sabrosky 148 speciosus Meigen 182 strigulus Fabricius 182 281

strobilum Karps 104 styriacus Strobl 88 subnigripes Dely-Draskovits 200 suecicus Nartshuk & Andersson 107 sulcicollis Meigen 139 tarsata Fallén 218 tenuiseta Frey 94 Thaumatomyia Zenker 226 tibialis Macquart 88 tjederi Nartshuk & Andersson 90 tomentosum Nartshuk & Andersson 143 tornensis Nartshuk & Andersson 83 Trachysiphonella Enderlein 148 triangularis Becker 177 triangulina Fedoseeva 215 Trichieurina Duda 230 Tricimba Lioy 149 trifasciata Zetterstedt 229 trigonella Duda 123 trilineatum Meigen 65 trochanterata Collin 132

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troglodytes Zetterstedt 184 tuberculifera Corti 95 umbraculata Fabricius 221 vagans Meigen 90 vallaris Collin 88 variegata Loew 195 variegata Meigen 215 varsoviensis Becker 184 vastator Curtis 132 ventricosi Nartshuk 133 venustus Czerny 139 zernyi Duda 185 zetterstedti Andersson 84 zimzerla Nartshuk 216 zonulatus auct. 172 zonulatus Wahlgren 167 zuercheri Duda 113