The empidoidea (Diptera) of Fennoscandia and Denmark. III. Genus Empis [1/1, 1 ed.] 9004096639

Citation preview

noscandia ark. Ill ius Empis

by

M. Chvala

List of abbreviations for the provinces used throughout the text, on the map and in the following tables. DENMARK SJ South Jutland EJ East Jutland WJ West Jutland NWJ North West Jutland NEJ North East Jutland F Funen

LFM SZ NWZ NEZ B

Lolland, Falster, Mgn South Zealand North West Zealand North East Zealand Bornholm

SWEDEN Sk. Bl. Hall. Sm.

Skane Blekinge Halland Smaland

Vrm Dir. Gstr Hls.

Varmland Dalarna Gastrikland Halsingland

Ol.

Oland

Med

Medelpad

Gtl. G. Sand. Og. Vg. Boh. DIsl. Nrk. Sdm. Upl. Vstm.

Gotland Gotska Sandén Ostergétland Vastergétland Bohuslan Dalsland Narke Sédermanland Uppland Vastmanland

Hrj. Harjedalen Jmt. Jamtland Ang. Angermanland Vb. Vasterbotten Nb. Norrbotten As. Lpm. Asele Lappmark Ly. Lpm. Lycksele Lappmark P. Lpm. © Pite Lappmark Lu. Lpm. Lule Lappmark T. Lpm. Torne Lappmark

NORWAY

4)

Ostfold

HO

Hordaland

AK HE O B VE TE AA VA R n northern

Akershus Hedmark Oppland Buskerud Vestfold Telemark Aust-Agder Vest-Agder Rogaland s southern

SF MR sii NT Ns Nn TR F

Sogn og Fjordane M@re og Romsdal Sor-Tréndelag Nord-Tr@ndelag southern Nordland northern Nordland Troms Finnmark

FINLAND Al Ab N Ka

Alandia Regio aboensis Nylandia Karelia australis

Kb Om Ok ObS

Karelia borealis Ostrobottnia media Ostrobottnia kajanensis Ostrobottnia borealis, S part

Satakunta

ObN

Ostrobottnia borealis, N part

Tavastia australis Savonia australis Ostrobottnia australis Tavastia borealis Savonia borealis

Ks LkW LkE Li Le

Kuusamo Lapponia Lapponia Lapponia Lapponia

St

Ta Sa Oa Tb Sb

@ eastern

Vv western

y outer

kemensis, W part kemensis, E part inarensis enontekiensis

USSR

Vib

Regio Viburgensis

Kr

Karelia rossica

Lr

1 inner

_ Lapponia rossica

FAUNA

ENTOMOLOGICA

SCANDINAVICA

Volume 29

1994

The Empidoidea (Diptera) of Fennoscandia and Denmark. III Genus Empis by

M. Chvala

E.J. Brill Leiden

- New York

- Koln

The paper in this book meets the guidelines for permanence and durability of the Committee on Production Guidelines for Book Longevity of the Council on Library Resources. © Copyright 1994 by E. J. Brill, Leiden, The Netherlands

All rights reserved. No part of this book may be reproduced or translated in any form, by print, photoprint, microfilm, microfiche or any others means without written permission from the publisher. Authorization to photocopy items for internal or personal use is granted by E.J. Brill provided that the appropriate fees are paid directly to Copyright Clearance Center, 27 Congress Street, SALEM MA 01970, USA. Fees are subject to change. PRINTED IN THE NETHERLANDS Editor-in-chief: N.P. Kristensen Desk editor: V. Michelsen

ISBN 90 04 09663 9 ISSN 0106-8377 Library of Congress Cataloging-in-Publication Data (Revised for vol. 3)

Chvala, Milan. The Empidoidea (Diptera) of Fennoscandia and Denmark. (Fauna entomologica Scandinavica, 0106-8377 ; V.

)

Sequel to: The Tachydromiinae (Dipt. Empididae) of Fennoscandia and Denmark. Vol. 3 has imprint: Leiden : New York : E.J. Brill. Includes bibliographical references and indexes. Contents: -- 2. General part, the families Hybotidae, Atelestidae, and Microphoridae -- c. Genus Empis. 1. Diptera--Scandinavia--Classification. 2. Insects--Scandinavia--Classification. I. Chvala,

Milan. Tachydromiinae (Dipt. Empididae) of Fennoscandia and Denmark. II. Title. II. Title: Empidoidea of Fennoscandia and Denmark.

QL535.4.S34C47 1983 ISBN 90-04-09663-9

595.771

85-126754

Die Deutsche Bibliothek - CIP-Einheitsaufnahme

The Empididae (Diptera) of Leiden ; New York ; K6ln:

Fennoscandia and Denmark. -

Brill

3. Genus Empis / by M. Chyala - 1994 (Fauna entomologica Scandinavica ; Vol. 29)

ISBN 90-04-09663-9 NE: Chvdla, Milan; GT

Cover illustration: Empis (Platyptera) borealis L. Author’s address: Milan Chvala Department of Zoology Charles University Viniéna 7, 128 44 Praha 2 Czech Republic

Contents MROIC TCU OLE Me eet re nodes eens ices Meee Shee os ace ee ie ete bee we oe Ea ee aNd Mla ENORMONVICCECIC TSH mee eat REM Aleta ees oy vs HO eee or ou eee eee ee eAGY Eaea es Ati alkanTiCaic LhOUS Magee terete eta ene ess os PRES se Mee eae EE WMS ae cae eA EMME OUDMOLOPY maeat eat ee elele eee Pe kas. 5.)sea SE AE OEM a aes eee a UoO COV FAlICUIC ASSILCATLOM Renee nem ete tea hes a o's Seo tee ete el lead Meare Sad os Sie [LMG TS KOT es co Rh Racca i aa oe Ac mo PMA ALUTCISLA CCS EM Meares rere eevee ae as SHU i ohn EEN ome PRES tag POOLING, 6 5 cupulgei Carediet Canc a oh Ait elau ea? RP A a ee ae RACAL AD LESNAR CHS ee Lem, SPEC Sg as cule No ene LE WR, SPIO, SY ea “SWEPTDYES GCC SEN cc Aa TSIEN OPER ts ra a OR cea CEC ce Cat ig ECCAOOOLADLY MARTA TNS ee eR ee Saco SN enna ge Eee eee eee ees ests emrencObramiulesOn Em pididacm memati nee... wi-.o See eee eee oe Sos eb ee eae ees tate CMEORS CHCA OMEN PIGiINacen mete RAT. ac one MOM ae meee eee I aL | ee erm UEC A Ol W111 PISh ea ta Mae ener s eee aneyc Ste cigs ate ag aaao we cle ew MeN bee kek aoe t LR" “o! SBGGHES) SA Nr BiG Sis cin hid eetulcuins Ole, cana eee ke ad ee a a rer ILO COMUSECHIP iSMERCVaremty, ee Re ATS sore aha eto Rites seed alee ke ASE, GE SOP EMS LACEY TCTICIStCPICISmene sith Mae tac: ayocM eee ies ett ale eee ee PEED SENUSHL OLY DICDIIAHISNCZZIRM etree As,«iso. echo oe ees Ran ye rele rea ee A ESET CHIUISCAIT ILC1717150COL gM Meee Ne ae ova. eo 2A OR Doe el oe FE LEE ES PEST SCIUSEAMACKOSLICHILS DCZZIMA Maye Ct eee no. c Tee ee ele Seles SRR tales Ba eo PEED OCIUS IAL DIETa MCIZCNeeme eS. on 5 = De Hie melons 6 a oe TUE aS ek PmSUNCenUSHE EN teripis:‘ Collins mires & owas Site neers ae ea be be anda alantd ake eins PRS CMUSHLI/IDIS SSL Nee cats Are eS co ninicre Se ee re tee ee a, Eee Se BSL SCMUSKCODLOPNICDIGIDEZZIN Me htt retiat: «30a Moe amen ees ea tee cae eas oe bale PERE OCIILISEMQINMEHIPISADEZZN nts ee aiatie sis 5 cma a one Caner eee Gh gees SS EY STIISHET SS C//LPISI CZZIM Me MEME Ie sic a ssiohe eee Mee UM 2 eth Dodie a aah adlaenll JLSLEE SRE 2,o.dsbucublonn Creu deri oe ge ee ee ae eee (CACLUOHMS 6 o,ye lhatoet oae Mi aes ite AR ie ona ee ee a IGE . oo 6 2056.0 dyitieeia Biren Oe ae eae, ar crear eee oer Arr eran aaa,

6 7 8 8 11 12 12, 12 12 12 15 15 19 19 19 20 21 30 36 43 46 56 61 69 131 146 174 178 181 186

Introduction This volume covers the Scandinavian species of the genus Empis Linné of the family Empididae, and is the third ‘empidid’ volume ofthe ‘Fauna entomologica scandinavica’. The first empidid volume (volume 3 of the series) appeared in 1975 and dealt with the subfamily Tachydromiinae, now classified in the family Hybotidae. The second empidid volume (volume 12 of the series) appeared in 1983: in that volume, the former paraphyletic family ‘Empididae’ was split into four families, and the reasons for this were set out in a comprehensive ‘General Part’. Consequently, the 1983 volume had the superfamily name ‘Empidoidea’ in the title, and dealt with the rest of the newly-established family Hybotidae (subfamilies Hybotinae and Ocydromiinae) and with the two smaller families Atelestidae and Microphoridae. The present volume covers only the large genus Empis of the family Empididae (as newly restricted), and it is therefore the first volume actually dealing with the restricted family Empididae. It is, however, a direct continuation of volume 12, published in 1983, where the family Empididae was discussed in full: its systematic position, classification and phylogeny were covered, based on its detailed morphology as exemplified by Empis pennipes Linné, the type-species of the genus and family. For this reason the introductory sections here are brief, and the reader is referred for more detailed information to volume 12 of this series. The major part of this volume is the systematic treatment, which follows the format of the preceding two volumes. Each species begins with its valid name, followed by the original combination and a chronological list of synonyms with all the relevant bibliographic references. The descriptive part begins with a short diagnosis for rapid orientation, followed by a more detailed description of each sex to ensure their correct differentiation from related species. If necessary, further diagnostic notes are given. Information on types is given, including the identification of holotypes or lectotype designations or references to published revisionary notes. There is a paragraph for each species covering its distribution in Fennoscandia and Denmark, its flight period, general distribution, and data on biology, feeding habits and epigamic behaviour if known. The main diagnostic features for each species are illustrated, the male genitalia in

6

particular, and each subgenus is prefaced by a habitus drawing of the type-species. The subgenera are briefly diagnosed, with notes on phylogeny and classification, distribution and biology, and keys for the identification of species are given. The genus Empis consists of very attractive flies, varying in size from nearly 2 mm to 12 mm, and with unusual features in their habits and epigamic behaviour. The majority of Empis species occur in spring and their conspicuous swarming habits, combined with prey transfer by the male as a ‘wedding present’ to the female during mating, attract many collectors. Furthermore, females with their strikingly pennate legs spread out in flight, which enlarges the silhouette of an animal that is already large, are unique among flies and indeed among insects. Many species are very abundant, occurring in large numbers in a small area and often just fora short period in spring. Some species, however, such as the large Empis borealis Linné or Empis lucida Zetterstedt, are among the most common and abundant species in the north, far beyond the Arctic Circle, and are some of the most abundant arctic insects. In all, 65 species of Empis in eleven subgenera are treated in the present volume, though in taxonomically very difficult groups such as Empis s.str., Coptophlebia and Xanthempis, a few additional closely-allied species are discussed. The species are known from the cooler northern parts of Europe, along the coasts of the North Sea and Baltic Sea, but only 38 have been found in Fennoscandia

and Denmark. However, more intensive investigations in the Scandinavian countries and the use of modern mass-trapping methods will certainly lead to a better knowledge of the Scandinavian fauna. Such a possibility is clearly demonstrated by the recent collecting activity of Dutch Dipterists, especially of Mr V.S. van der Goot and Mr Van Aartsen. In 1989, for example, seven Malaise traps were run

for the whole season in the Netherlands, and some ten thousand specimens of Empis s.str. were examined from the catches (van der Goot, letter of 29.2.1990). This is about the number of specimens now available from all the Scandinavian countries! The results obtained from this material are fully commensurate with the effort and the collecting methods: nine species of Empis s.str. (E. lepidopus Meigen, E. acinerea Chvala, E. melanotricha

Loew, E. tanysphyra Loew, E. woodi Collin, E. gootisp.n., E. laminata Collin, E. dasyprocta Loew

and E. levis Loew) have been found for the first time in the Netherlands during the last five years. It is hoped that the present revision will lead to the discovery of further species in the Scandinavian countries, and will also stimulate and increase

the interest of Scandinavian Dipterists in this family of flies. It is also hoped it may assist further studies of Empis in Great Britain. The British fauna is rather well known, thanks

the excellent monograph of the Empididae published by Collin (1961) in the series ‘British Flies’, but in Scandinavia the last revisional works date back to the early years of this century. Lundbeck (1910) recorded 28 species of Empis from Denmark, and Frey (1913) 15 species from Finland and the adjacent Russian territory. Apart from scattered faunistic records by Scandinavian Dipterists and the more recent Finnish check-list by Hackman (1980), there are several new faunistic records from Norway (Jonassen 1987, 1988, 1992) and the

present author (Chvala 1991) has published a preliminary paper of the Finnish species of Empis s.str. and Coptophlebia, together with more detailed data on the distribution within Finland and NW Russia and forming a revision of Frey’s (1913) paper. The scope of the area being studied will be clear from the distribution ‘Catalogue’ at the end of the book. In addition to Great Britain, the Catalogue also includes the Netherlands — instead of N. Germany of the preceding two volumes, as the Dutch fauna is much better known (see Goot 1989), and an eastern territory covering Russian Karelia and Lapland. In the following text, the area of Fennoscandia and Denmark, which comprises Norway, Sweden, Finland and Denmark, is called Scandinavia for brevity as it was in volume 12, also matching the title of the series — Fauna entomologica scandinavica. Fennoscandia is used as a term of distribution to cover the Scandinavian Peninsula and Finland, including the adjacent Russian territory northwards to the Kola Peninsula.

Acknowledgements For the loan or donation of material, for help with literature, and for valuable information on collec-

tions I am very grateful to the following colleagues (in alphabetical order): M. Baez (La Laguna — Tenerife), M. Bartak (Prague), V. Beschovski (Sofia), P. Beuk (Leiden), I. Ceianu (Bucarest), R. Contreras-Lichtenberg (Vienna), A. Freidberg (Tel-Aviv), T. Jonassen (Sjernargy), L. Papp (Budapest), A.C. Pont (Goring-on-Thames), R. Rozkosny (Brno), W. Schacht (Munich), S.P. Schembri (Msida, Malta), Scott C. Sherman (New York), K.G.V. Smith (London), B.R. Stuckenberg (Pietermaritzburg), Bo G. Svensson (Uppsala), O. Syrovatka (Ceské Budéjovice), H. Troger (Innsbruck), F. Vaillant (Montbonnot), J.R. Vockeroth (Ottawa) and R. Wagner (Schlitz). I also wish to thank G.C.D. Griffiths (Edmonton) and H. Ulrich (Bonn) for their valuable advice and for many important suggestions during my morphological and phylogenetic studies. I am very indebted to L. Lyneborg (Copenhagen), L. Greve-Jensen (Bergen), H. Andersson and R. Danielsson (Lund), P.I. Persson (Stockholm), and W. Hackman and A. Albrecht (Helsinki), for their kind permission to study the Scandinavian collections under their care, as well as for _ their kind hospitality during my stays in Denmark, Sweden and Finland. These studies have been sup-

ported by grants from the Swedish National Sci-

ence Research Council, the Zoo-Tax in Stockholm, and by a cultural agreement between Charles University, Prague, and Helsinki University. I am also indebted to H. Schumann (Berlin) for his assistance during my studies of the collections of H. Loew and T. Becker in the Zoologisches Museum of the Humboldt University, Berlin. P. Grootaert (Bruxelles) kindly arranged for a visit to Belgium in 1987, supported by a grant from the Institut Royal des Sciences Naturelles de Belgique. I am especially grateful to V.S. van der Goot (Amsterdam) who arranged for two visits to the Instituut voor Taxonomische Zodlogie of Amsterdam in 1987 and 1990, supported by the Dutch U-E Foundation (Uyttenboogaart-Eliasen Stichting). I am particularly indebted to Mr van der Goot, whose enthusiastic help and collecting activity in the Netherlands has enabled a more thorough and complete study of the NW European Empididae fauna to be made. Finally, I also wish to thank Mr A. Veltman (Zevenaar) who has prepared the habitus drawings of the Empis species, especially for the patience and willingness with which he has accepted all my suggestions and alterations during his work on the illustrations. Mr Adrian C. Pont has kindly checked the English of the manuscript and offered many valuable suggestions.

Material and methods This monograph is based on an examination of nearly 10,000 specimens of Empis from the standard Scandinavian collections, which have been studied by the author over the last fifteen years or more, and a much larger number of specimens from other parts of Europe. The Scandinavian collections that I have studied are those in Copenhagen (with the historic collections of Fabricius, Staeger and Lundbeck), Lund

(with the large type collection of Zetterstedt, and also Wallengren, Roth and Ringdahl), Stockholm (with the old collection of Fallén, and additional specimens from Wahlberg and Boheman) and, finally, Helsinki (with the Frey collection, divided into separate ‘Finnish’ and ‘Palaearctic’ Collections). Thanks to Dr L. Greve-Jensen, I have been able to

study most of the collection in Bergen, including material of the taxonomically difficult groups of Empis, the alcohol material of the Hardangervidda IBP project, and some specimens from the Bidenkap Collection. I have also re-examined some species collected in Norway by Terje Jonassen. As will be clear from the above list, I have not

seen any private collection except for that of Terje Jonassen. However, I am sure that there are many Scandinavian dipterists, such as the Swedish Eskilstuna group, who are active in the field and must have many empidids in their possession. Apart from these Scandinavian collections I

have, of course, studied all the main European collections including the types of Meigen (Paris), Collin (Oxford, London), de Meijere (Amsterdam) and Loew and Becker (Berlin). I have not seen the types of Bezzi in Milan or those of Strobl at Admont, but I have studied specimens in other collections collected and/or identified by these two dipterists and I believe that their species are correctly recognised. The length of specimens was measured with an ocular scale grid to one-tenth of a millimetre, and was taken from the front of the head (without antennae) to the tip of the genital lamellae in males or tip of the cerci in females; if the male aedeagus

was long, extending beyond the tip of the genitalia, it was not included in the total length. Immature specimens, with a weakly sclerotised abdomen that shrivelled when dried, were not included in the range of body length. However, it should also be borne in mind that specimens preserved in alcohol, with the body telescopically expanded, will often be unnaturally large in size. All figures were drawn using a binocular microscope with an ocular grid. Male genitalia were dissected and boiled in a 10% solution of potassium hydroxide and studied in glycerol. Very exceptionally (in rare species, with single specimens available) the genitalia were drawn ‘in situ’.

Adult morphology Species variable in size from very small, little more

than 2 mm in length, to very large and, with a body length of almost 12 mm, the largest forms in the family. Generally black, more or less dusted grey or shining species, rarely yellow (Xanthempis and some Polyblepharis and Leptempis), with body almost bare, or more or less pubescent or bristled. Legs armed with hairs or bristles in male, certain parts (basitarsi, femora, tibiae) sometimes swollen or thickened; in females legs often curiously pennate. Head with a long slender proboscis, wings with vein R,,, apically forked, and metapleura with a cluster of bristles. For detailed comparative morphology, see volume 12 of the Fauna (Chvala 1983).

Head

Head rounded or rather higher than either long or wide, occiput flat or slightly concave, especially be-

8

low neck, and large compound eyes. The species of Xanthempis have small eyes, always dichoptic in both sexes, and occiput convex, broadly produced posteriorly. Males are holoptic in the ground-plan, with upper ommatidia more or less enlarged, very strongly so in most Empis s.str. and Coptophlebia, though male eyes may be narrowly approximated (some Euempis and Polyblepharis) with a tendency towards a decrease in the upper ommatidia, or eyes broadly separated on frons (Pachymeria, Xanthempis and some Polyblepharis) as is obligatory in the female sex, and with all ommatidia

small, of the apposition type. Antennae situated at about middle of head in profile, with basal two segments bristled or at least covered with fine bristly-hairs, 2nd segment always globular and as long as deep; basal segment as long (Empis s.str., Coptophlebia), but usually distinctly longer, rarely very elongate (Xanthempis). 3rd segment moderately long in ground-plan

and conical, with a terminal 2-segmented style (with a small basal segment) shorter than the length of 3rd segment or at most as long (except for the non-Scandinavian Rhadinempis, with a very short 3rd segment and long style), less often 3rd segment very elongate and style short (Xanthempis, Lissempis and some Anacrostichus). Mouthparts very conspicuous, with a long labrum which is at least slightly longer than head is high, although usually much longer, up to three times as long as head is high in some Empis s.str. species and, if a long labium is porrect, then the proboscis is much longer still. The stylet-like hypopharynx and the paired maxillae (laciniae) as long as labrum, maxillary palps often small, slender, rarely longer or distinctly bristled.

Thorax

Primitive forms of Empis have the mesonotum densely covered with non-differentiated hairs or bristly-hairs, as in some Euempis, Polyblepharis and even Empis s.str. species, with a gradual differentiation and specialisation of sensory bristles and their consequent reduction. The most specialised forms have the mesonotum almost bare with strongly differentiated marginal bristles and reduced acrostichal and dorsocentral bristles, right up to the complete loss of acrostichal bristles (Xanthempis and a few species in almost all the more advanced subgenera), and to the almost complete reduction of all bristles as in E. (Xanthempis). digramma. Metapleural bristles always present, even if only a few and/or small and inconspicuous. Prothorax always with a large precoxal bridge, the prosternum fused with prothoracic episterna, completely covered with hairs in primitive forms. Prothoracic episterna in the form of small discrete sclerites between upper side-tips of prosternum and pronotum (prothoracic collar), and with specific bristles or pubescence.

rower wing with longish discal cell produced apically, all veins complete and reaching wing-margin, radial fork (apical fork of vein R,,,) rather acute with upper branch (vein R,) slightly curved, at least not straight, and axillary lobe large, consequently with an acute axillary excision. The most specialised wing venation (the apomorphous state) is present in some Empis s.str. species, where the wings are rather broad, clear or even whitish with fine inconspicuous veins except for strong (darkened) veins along costal margin (C, R,,;, Ry,;), radial fork broadly ‘open’, with upper branch straight and almost at right-angles to vein R., and a short, broad,

apically truncate discal cell. On the other hand, the very weak development of the axillary lobe, or even its absence, with a very obtuse axillary excision as in Lissempis or Xanthempis, is also considered to be an apomorphous state. Full information on the wing-venation in the Empididae, and the characteristics of plesiomorphous and apomorphous states, are given in the preceding empidid volume of the Fauna (Chvala 1983). However, there are frequent venational aberra-

tions in Empis species that sometimes cause serious problems with identification, even at the generic level, as with the partial or complete loss of the radial fork. An incomplete radial fork on one wing, or even its complete absence on one wing, is a fairly common situation in practically all Empis subgenera, but specimens without any indication of a radial fork on either wing will be identified as species of Rhamphomyia. It has been found, for example, that Rhamphomyia gentilis Loew is actually Empis (Polyblepharis) fallax Egger without a radial fork on either wing, and, conversely, the type of Empis s.str. pulicaria Loew has proved to be Rhamphomyia (Aclonempis) longipes Meigen with a complete ‘Empis’-like radial fork on one wing. Venational aberrations may also involve the abbreviation of veins M, and/or M,, which in some specimens do not reach the wing-margin (as is customary in Coptophlebia and most Kritempis species). Sometimes an extra crossvein r-m is present,

Wings Wings in Empis, so far as known to me, always well-developed, clear, very whitish to milky-white, or uniformly clouded to deep brown, or distinctly spotted. Wings rarely conspicuously broadened in “females and with parallel infuscation, within the

European fauna only in the subgenus Platyptera _ anda few Empis s.str. species. It is suggested that the plesiomorphous state of the wing-venation is represented by a rather nar-

or a crossvein m-m connecting veins M, and M, behind the discal cell, forming an additional small cell.

Legs The legs are generally long and slender, rather densely covered with short or more or less long hairs, or with very specific bristles. Female legs are often shorter, sometimes rather flattened laterally,

especially on posterior femora and tibiae, and strikingly pennate, covered with flattened pennate

y

bristles or fringed with flattened hairs. In males of Empiss.str.and Coptophlebia, the basitarsi on fore and/or hind legs are often more or less swollen, and in some species-groups of Anacrostichus and Pachymeria the hind femora are usually conspicuously thickened and ventrally spinose in both sexes. The mid legs are generally very slender and much smaller, especially when compared with the enlarged and elongated hind pair. Almost all species of Empis have at the distal end of hind tibia a posteroventral comb-like row of bristle-like

setae

or spines

called

a ctenidium,

which is sometimes very strong and distinct, and Collin (1961) used this character (or that of a spine set in the ‘comb’) for distinguishing some Empis subgenera. The bristling of the legs in the male, especially on the mid pair, where unusual long sensory bristles are often present, undoubtedly offers a good and phylogenetically useful feature for separating monophyletic lineages at the specific level. On the other hand, the bristling of the female legs in the form of a ciliation of short stout bristles, flattened hairs or bristles (lanceolate bristles), or the very characteristic strong pennation (pennate, apically rounded bristles), appears to be a rather recent adaptation. It is believed that pennation of the legs in females has evolved only as an adaptation for specifically distinct epigamic behaviour, as a very useful signal for the process of ethological isolation among closely related species to prevent interbreeding. It is, therefore, an adaptive character with no phylogenetic value, bringing together quite unrelated species, such as E. acinerea and E. laminata within Empis s.str., as is clear in the Key to females.

Abdomen

The female abdomen is always telescopic, with more or less elongate slender cerci at tip. The last two visible segments (7th and 8th) are sometimes elongated and narrowed, ovipositor-like. The male abdomen consists of 8 pregenital segments; in general, the 8th tergum is a small discrete sclerite and the corresponding sternum is enlarged, but insome Xanthempis species both sclerites of the 8th segment have been fused into a single ring-like sclerite. One or two pregenital segments are often more or less modified, armed with specifically distinct protuberances, tubercles, and excisions.

10

Male genitalia The male genitalia of the Empididae were fully discussed in the ‘General Part’ of the preceding empidid volume of the Fauna (Chvala 1983) and will not be discussed here again in detail. The epandrium (9th tergum) is primitively a discrete dorsal sclerite, more or less deeply cleft into two lamellae (dorsal lamellae) which are usually small and more or less concealed between the lateral lamellae, less often enlarged, very strongly so in Kritempis, or of complicated structure (Leptempis, some Empis s.str.). The hypandrium (9th sternum), the so-called ventral lamella, is often very reduced, shielding the base of aedeagus from below.

The main visible elements of the male genitalia are the /ateral lamellae, the gonopods, or more precisely their basal segment, the gonocoxites. The gonopods are paired, originally clasping organs, shielding the aedeagal complex from the side in the form of large lateral sclerites that vary species-specifically in size, shape and structure. The homology of the individual genital sclerites in the Empididae is still, however, a subject of some controversy, and the terminology has not yet been unified and cleared up satisfactorily in the literature. Some authors interpret the dorsal lamellae (epandrium) in Empis as cerci, and the lateral lamellae (gonopods) either as the epandrium or as fused gonopods with epandrium. The ventral lamella (hypandrium) has very recently been interpreted by some authors as a fused hypandrium with gonopods. To avoid possible misunderstandings with the terminology of these structures, and as the primary purpose of this book is to facilitate the simple and straightforward identification of the north European fauna of Empis, the common and universal term ‘lamellae’ (dorsal, lateral, ventral) is used in the text that follows. Aedeagus rather short and moderately stout in the ground-plan, almost wholly concealed within the lateral lamellae, as for instance in Euempis, in the E. tessellata-complex. There is a tendency towards elongation of the aedeagus, and the long, free, and often strikingly bent or looped aedeagus represents an apomorphous state. An unusually long and thin, thread-like or wire-like aedeagus is present in some Empis s.str. species (acinerea, albopilosa, decora), but also in Pachymeria, for in-

stance in the E. femorata-complex.

Phylogeny and classification The phylogeny of the Empidoidea, including the family Empididae, has been fully analysed by the present author (Chvala 1983). Since that time a paper has appeared by Waters (1989), containing the description of the fossil Empis orapaensis, the oldest known representative of the subfamily Empidinae, tribe Empidini, described from the Afro-

tropical region, from the Mesozoic middle Cretaceous fossiliferous sediments of the Orapa diamond mine, Botswana. This finding supports previous conclusions, except that the diversification of the empidid subfamilies, and that of the subfamilies Empidinae and Hemerodromiinae in particular, must have taken place earlier and during the lower Cretaceous period. There are few diagnostic features preserved on the mudstone compression fossil of the holotype male of Empis orapaensis, as the head has been completely lost, and the legs and tip of abdomen are very indistinct. However, the wings are wellpreserved and all the characters of the wing venation are clearly plesiomorphous, as given for the supposedly plesiomorphous condition of Euempis species in the section on adult morphology. Waters (1989) classified Empis orapaensis in the subgenus Empis s.str., but the only basis for this was that ’all branches of vein M issuing from cell dm (i.e. discal cell) are complete and reaching the wing margin’. Her assignment of this species to Empis ‘sensu stricto’ should therefore be regarded as the classification of a form with an unmodified wing venation, unlike that of Coptophlebia for instance. The Scandinavian Empis species are here classified into eleven larger systematic units, traditionally treated as ‘subgenera’ in the Palaearctic fauna. This subgeneric classification was founded by Bezzi (1909), and has been followed by all subsequent dipterists including R. Frey in Finland and J.E. Collin in England. This classification was also adopted in the Palaearctic Catalogue (Chvala and Wagner 1989) where the eleven northern European ‘subgenera’ were recognised together with the central European Rhadinempis Collin and Argyrandrus Bezzi, and the East Asian Planempis Frey. However, this ‘Palaearctic’ subgeneric classification is not applicable to other geographical regions, and not even to the North American fauna. There are even difficulties in classifying some of the Mediterranean species, as is clearly illustrated by the group of ‘Unplaced species of Empis’ in the _ Palaearctic Catalogue.

I am well aware that the Palaearctic ‘subgenera’ are very artificial and unnatural units, but I did not

want to change the classification familiar in Europe without proposing a new natural classification. There are perhaps two or three natural lineages forming monophyletic groups among the recent Empis ‘subgenera’, such as Xanthempis Pachymeria,

but Lissempis

for instance

or

is more

closely related to Rhamphomyia subgenus Lundstroemiella than to any other subgenus of Empis, and these two together clearly form another monophyletic group. The same also applies to the complex of Empis s.str., Coptophlebia, and Rhamphomyia subgenus Aclonempis. The crux of the problem is with the correct generic differentiation of Empis and Rhamphomyia, as the only distinguishing (though practical) character lies in the presence or absence of the radial fork on the wing, a character not infrequently subject to aberrant development. The generic classification of the Empidini is not just a problem of the Palaearctic fauna, but also involves other ‘genera’

with extralimital species. The example of Empis s.str. and Aclonempis in Europe, with a practically identical structure of the proboscis, is quite obvious. However, any attempt at a new classification must be based on a thorough knowledge of the world fauna, and this also applies to the subgeneric classification of Empis. The primitive forms of the genus Empis should be found among the species classified in the subgenus Euempis. The supposed monophyly of Pachymeria is based on the structure of male genitalia, and on further characters in the structure and bristling of the legs and abdomen. The subgenus Xanthempis can be characterised by several autapomorphies in the structure of the head, including the mouthparts, wings and legs. It is suggested that the rest of the European species have evolved along several phylogenetic lineages, one (of boreal origin) leading to Anacrostichus and Platyptera, another to Polyblepharis and Kritempis (evidently of subtropical and/or Asian origin), and the last one through Leptempis to Empis s.str. and Coptophlebia. The informal groups (or complexes) of species within the ‘subgenera’, if such divisions are natural or necessary, are mentioned under each higher taxon.

11

Life history Immature stages Practically nothing is known about the immature stages of Empis. The few records available indicate that in cool and temperate Europe larvae are probably exclusively predaceous, living in soil, and that the larva is the overwintering stage. Collin (1961) refers to the larva of E. (Coptophlebia) vitripennis bred from a ‘Puff-ball’ fungus in England, and quotes Beling who bred E. (Xanthempis) trigramma larvae from a cow pak from the previous year. I myself have identified many Empis species from various biotopes in central Europe from soil emergence traps, including specimens of E. trigramma, the larvae of which evidently develop in soil. At the beginning of April 1965, I collected numerous empidid larvae in soil beneath a willow tree in southern Bohemia, and from these two Empis species, E. (Euempis) tessellata and Empis s.str. beckeriana Engel, emerged from pupae in the first half of May the same year.

Adults

The biology of adult empidids is much better known, and better than in most other dipterous families, because of their very exceptional ethology. For a long time the Empididae, the danceflies, have been the subject of general interest because of their swarming activity and unique habits of parallel prey presentation, a transfer of the socalled ‘wedding present’ or ‘nuptial gift’ during mating in aerial aggregations. Empis species, as well as Hilara and Rhamphomyia, exhibit a wide range of habits that have recently been known as important ethological isolating mechanisms to prevent interbreeding and to secure a quick meeting of different sexes of the same species. The epigamic behaviour of Empis species, as well as that of other empidines, is closely connected with their feeding habits, ecology and phenology, as isolating ethological mechanisms are important only if other non-biological barriers are not effective.

(Chvala 1976). The nectar-feeding habit evolved secondarily from the original predatory activity, and this new feeding habit prevailed in some groups, and especially in Empis. Nectar-feeding may be regarded as the main feeding habit in recent Empis species, although in some groups the original predatory activity has been partly (some Euempis) or wholly (Xanthempis) retained during the whole adult life in both sexes or just for a short period of mating activity, when females accept insect food from males as a ‘wedding present’. In many Empis species, this undoubtedly serves as a supplementary source of protein-rich food required by females for ovarian maturation, unless the whole process has been ritualised and the ‘prey’ is only a stimulus or signal for mating. Very recently Svensson, Petersson and Frisk (1990) have concluded that the time spent in copula in E. (Platyptera) borealis was positively correlated with the volume of the nuptial gift transferred by male to female. This means that the prey presentation during copulation still serves in some species as a food for females. A rather short and stout proboscis indicates strong predatory activity, but even the characteristic long and slender ‘Empis-like’ proboscis, as for instance in E. s.str. pennipes, readily serves to imbibe sweet nectar from flowers, as well as for very

active predation on other insects, piercing and sucking liquified animal tissue. As regards the role of Empis species in nature, their predatory activity is obviously far less important than that of some predaceous hybotids such as Platypalpus, Drapetis and others. The economic importance of Empis species clearly rests on their nectar-feeding activity, as many species are undoubtedly very important spring-time pollinators, especially of fruit-trees. Waters (1989) was undoubtedly right in suggesting that the diversification of the Empidoidea was correlated with the appearance of the angiosperms and that empidids served as early pollinators of the flowering plants. This must have been true throughout the long evolution of Empis.

Swarming and mating

Feeding habits It is generally accepted that the original forms of Mesozoic Empis (and of the family Empididae) were predaceous because of the absence of angiosperms with nectaria that secreted sweet substances

12

Swarming in the air is a primitive but indispensable habit in Empis to bring together different sexes of conspecific populations and to enable emerging females to mate immediately. There is an extensive literature on this phenomenon, and more detailed

Table 1.

Type of distri-

Great Britain

Europe

bution

Eng-

Scot-

north- north- central south- south

south-

(cf. Figs

land

land

west

east

east

Asia

west

Denmark

and Fenno-

1,2)

Euempis 1. tessellata 2. sericans 3. picipes 4. ciliata Pachymeria 5. femorata 6. scotica 7. tumida

Polyblepharis 8. Opaca 9. strigata Kritempis 10. livida Anacrostichus 11. lucida

12. verralli 13. nitida 14. bistortae Platyptera 15. borealis Leptempis 16. rustica 17. nigricans 18. variegata 19. grisea Empis 20. lepidopus 21. albopilosa 22. serotina 23. decora

scandia

Cc (A) B (A)

+

(A) (B) (A)

+ + +

+

A (G)

+

+

C

aE

4

1

aE

+

(A) D (A)

+

+

D

4

A (A) (A) A (A) (A) A (A)

Flight period in

+

4

a

+ + + +

+

+

+

+ + + +

+ +

+ + +

+

i

+

+ + + +f

4 +

+ + +

+

V-VIII (V-VI) V-VII (V-VI)

+

(V-VI) (VII-VIII) (VII-IX)

+

V-VI (VI)

+

+ +

+

+

+

VI-IX

+

1

+p

V-VIII

+ +

+

a +

ste

i:

+

i

+ 4p +

+

(V-VI) V-VIII (VI-VI)

+

ta

+

+ + + +

=f: + + +

+

+

tf

IV-VII

VI-VIII (V) (V-VI) VI-VIII +

+

(IV-V) (V-VI) VI-IX (V-VI)

+ +

24, acinerea

B

+

+

+

IV-VI

25. 26. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37. 38. 39. 40. 41. 42.

Cc (A) A A B (A) (A) (A) (A) D A € A D A B @

+ + + +

+

+ + ‘ts 4 +

VII-IX (VI-IX) V-VI V-VI V-VI (VI-VIII) (V-VI) (V-VI) (V-VI) V-VII VI V-VII V (VI) V-VII vV-VI V-VI VI-VII

syrovatkai genualis rufiventris planetica staegeri limata melanotricha tanysphyra woodi nigripes nitidula laminata gooti bicuspidata nuntia chioptera prodromus

+ te

+

+

+

+ +

3

4°

+

+ 4b + +E +

+ + + +

43. dasyprocta

(Ay.

44. caudatula

A

ie

45. aestiva 46. praevia 47. levis

A (A) (A)

ap +

+

+ + se

+

4

+ + + + 4 +

+ + + +

+

+ + 4p +

+ +

oF

+ + + + + +

+

+

(VII-VHI)

te

3

+

+

+

ie

+

V-VII

oF +

+

VI-VII (V) (V-VI)

13

Table 1.

Type of

Europe

Great Britain

Asia

distri-

north-

north- central south-

bution

Eng-

Scot-

(cf. Figs

land

land

west

48. hyalipennis

D

+

oe

“

+

a

49. impennis

(A)

+

EF

=f.

50. albinervis

A

~

+

51. vitripennis 52. volucris

A (A)

+ ~

2:

53. pilimana

(A)

ee

a fe Ee ae a

+

4

i

east

south

west

Flight period in

south-

Denmark

east

and Fennoscandia

+

VII-IX

Coptophlebia

Xanthempis 54. scutellata 55. laetabilis 56. lutea

(A) B A

a +

albifrons rohaceki

ze +

+ -

(V-VI1)

+

VI

+

+

+ ae

=

(A)

+

+

58. stercorea 59. aemula

D A

+ +

+ -

us +

-

60. univittata 61. trigramma

B A

-

~

+

+

+

+

62. punctata

B

-

+

+

(A)

fe

+ =

+

63. testacea

4:

64. digramma

B

=f

4

n

Lissempis 65. nigritarsis

A

~

information is given under individual subgenera and species. However, in Empis, as well as in Hilara and Rhamphomyia, mating in the air is connected with feeding, as the secondarily nectar-feeding females receive protein-rich food from males during copulation as a ‘wedding present’ or ‘nuptial gift’. As discussed in the chapter on feeding habits, the prey presented to the female originally served as a food, but its original meaning has gradually been lost and the prey either became a stimulus for mating, as the female is not receptive to a male without prey, or its function was a visual signal to the female of the characteristically enlarged male carrying a prey. The primitive forms assigned to the subgenus Euempis are very primitive in their behaviour, as species that are still both predaceous and nectarfeeding mate without prey transfer, whereas in the purely nectarivorous species such as E. tessellata prey transfer during copulation is already obligatory. On the other hand, in the otherwise very specialised Xanthempis species that have also retained the primitive predatory habit during adult life but are also parallel nectar feeders, mating activity is not connected with prey transfer. More-

14

+

+ +

=e

~

VIL-VIUI

+ = ~

57. concolor

ae

(VI-VII) VLVII VITI-IX (VI-VII) (VI-VIII)

ES

fe

as

a8

+

(VI-VII) +

~ -

+

45

V-VII

VI VLVII

+

V-VI

+

V-VII (V-VI)

+

Es

V-VI

+

+

V-VII

over, mating in this group has been transferred to the ground, a unique habit within the Empidinae. All other European Empis species, except perhaps Lissempis, produce aerial swarms, usually a very species-specific synorchesia (sensu Gruhl 1955) where, with few exceptions, males are the ac-

tive elements, flying in stable swarms and waiting (usually with prey) for emerging females. Modified swarms are known only in the species of Polyblepharis, Kritempis and Anacrostichus, usually the typical polyorchesia (sensu Gruhl 1955), an occasional rhythmic dancing-perching-dancing by several individuals, that may be also dispersed over a larger mating area. The different sexes meet in the air and start immediately to copulate, but pairs settle on nearby vegetation and do not fly in copula in the air (except when disturbed) as is common in many species of Hilara and even Rhamphomyia. In some species of Polyblepharis, males offer females inanimate objects such as seeds or flower petals, which are readily accepted by females during copulation. The so-called ‘balloon-flies’, where males produce curious objects made of silk as ‘wedding presents’, remain unknown in the Palaearctic (or at least western Palaearctic) Empis

species, but are known among certain Nearctic Empis species and Palaearctic Hilara. The whole process of swarming and mating in Empis has become ritualised in some way over a long period of evolution and has become an effective ethological isolating mechanism and a very good way ofisolating related sympatric populations (Chvala 1990).

Phenology In principle most Empis species are typical spring species, occurring mainly in the second half of May and in June. This is also the peak period when most species are on the wing and when Empis, especially the common species, are very abundant almost everywhere, being commonly found during the day on flowers and performing their regular characteristic aerial swarms. There is no doubt that the earliest spring species of Empis (and of the whole family) is E. (Platyptera) borealis, recorded even in Scandinavia as early

as 28 March. It is rather common in April in lowlands in the south, although in the mountains and in the north it can still be found in July. Other early spring species, on the wing in Scandinavia and/or England by the end of April, are E. s.str. chioptera, acinerea and /epidopus, and E. (Lissempis) nigritarsis. Three other species of Empis s.str., E. albopilosa, woodi and tanysphyra, are also early spring species, occurring at the very beginning of May.

As stated above, the majority of species fly for a rather limited spring period in May and June, although there are two very characteristic species, Empis s.str. albopilosa and E. (Leptempis) nigricans, which are found only for two or three weeks in May. On the other hand, there are several species that have an unusually long flight period lasting from May to August, such as E. (Euempis) tessellata, E. (Anacrostichus) nitida and lucida, or Empis s.str. nigripes. A rather long flight period is also characteristic of a few species that occur in high summer, that are on the wing mainly in July and August, such as Empis s.str. aestiva, dasyprocta and limata. There is also a group of typical summer species that continue flying until September, such as E. (Kritempis) livida, Empis s.str. syrovatkai and genualis, or E. (Coptophlebia) hyalipennis. However, Empis s.str. serotina and E. (Coptophlebia) vitripennis are the only two Scandinavian species that can be recorded as typical late summer or even autumn species, being on the wing only in August and September. _ Table 1 shows the flight period of all Empis species within Scandinavia. For species not yet found in Scandinavia, a presumed flight period is given in brackets based on the seasonal occurrence in Great Britain or the Netherlands. In temperate central Europe, flight periods in most species are often two or three weeks earlier, and in southern

Europe at least a month earlier.

Zoogeography The genus Empis is worldwide in distribution though it evidently prefers warmer southern and subtropical areas as can be clearly seen from the distribution pattern of Empis within Europe. According to the recent Palaearctic Catalogue (Chvala and Wagner 1989), 210 species of Empis have been described and named from Europe and only 65 of these include the cooler northern areas in their distribution. However, in view of the fact that the fauna of northern and central parts of Europe is much better known and that there are still very many species in the southern warmer areas and the Mediterranean awaiting description, the prevalence of species inhabiting southern areas will be much greater. Unlike Rhamphomyia, the typical empidids of boreal and even circumpolar distribution, the ge-

~ nus Empis most probably originated in the tropics and subtropics. This can also be clearly seen from the distribution-pattern of the 65 northern Eu-

ropean species summarised in Table 1. As in the previous two empidid volumes, the Scandinavian species are again classified into six distributionpatterns (A-G), as shown in Figs 1 and 2. A- Atlantic and central European elements extending into Denmark and the extreme south of Sweden. The largest group of species, almost half of the Scandinavian Empis species (45%), belongs here. There is a small group of Empis s.str. species (planetica, aestiva) and Coptophlebia (albinervis, vitripennis) that are also included in this group although they extend further northwards in the west and into the south of Norway, thus not exactly matching line A in Fig. 1. B-— Species with a southern distribution within a larger area covering Denmark, the southern parts of Norway and Sweden, and also the

Baltic coast of Finland, reaching approxi-

15

the northern and central parts of Scandinavia, but absent in the south. This is an unusual type of distribution within Empis and is represented only by E. (Anacrostichus) lucida, a very common species in the north, a boreomontane element, known outside Scandinavia from Great Britain and isolated localities in the Alps. This type of distribution, including the extreme northern species distributed beyond the Arctic Circle, is unknown in Empis. Continental eastern and central European species, extending through the NW parts of European Russia to the eastern parts of Scandinavia, especially to Finland. This group is represented by only one non-Scandinavian

mately 62° N. Only 8 species, representing 21% of the Scandinavian fauna. Species with a similar type of distribution as group B, but extending further northwards to the central parts of Scandinavia, to approximately 65-66° N. E.°(Euempis)

tessellata, E.

(Kritempis) livida, and four species of Empis s.str. (pennipes, syrovatkai, laminata and prodromus) belong here. Species widely distributed throughout Scandinavia, even occurring in the very cold parts of the extreme north, in Lapland. A small group, represented by E. (Platyptera) borealis, E. (Anacrostichus) nitida, Empis s.str. ni-

E-

gripes and bicuspidata, E. (Coptophlebia) hyalipennis and E. (Xanthempis) stercorea. Exclusively northern species, distributed in 102ee

2

is

ES “4

N

ibe

18°)

202

22°

\ Saas

\ \

(&

Ne

7

\

Ang

Fig. 1. Distribution pattern of Fennoscandian species. — Groups A-C.

16

524%

926%

28°

302)

1322

34°

36°

38°

species, E. (Polyblepharis) strigata, which may well occur in eastern Finland. Quite logically, the two commonest Scandinavian Empis species, E. (Platyptera) borealis and E. (Anacrostichus) lucida, are widely distributed spe-

(Anacrostichus)

verralli and Empis

s.str. limata,

which still appear to be endemic in the British Isles. 40 species of Empis are now known from Great Britain and 48 from the Netherlands, which is the best-investigated country at present. Fourteen species found in the Netherlands have not yet been

cies even in the north, and are classified in distribu-

tion-patterns D and E respectively. As regards the extra 27 Empis species treated here as possible Scandinavian species (their distribution symbols are given in Table 1 in brackets), they all belong to distribution-pattern A, to the typical rather southern Atlantic elements, except for E. (Polyblepharis) strigata in group G and E. (Pachymeria) scotica which should be provisionally classified in group B. Of the 65 species treated here, eight have so far been found only in Great Britain, including two, E.

recorded from Scandinavia, but they are included

in the present study as expected Scandinavian species of distribution-pattern (A) in Table 1. Although the distribution data for Empis species in the Palaearctic Catalogue (Chvala and Wagner 1989) is very up-to-date, thanks to the late Arpad Soo6s and to Laszl6 Papp (the editors of the Catalogue) who readily accepted all the new data sent by the present author even at the final proof stage, there are several species found very recently in the Netherlands that were not included in the Cata-

6 Sand

ot

ee

(f (

Fig. 2. Distribution pattern of Fennoscandian species. — Groups E-G.

17

logue. They are E. (Anacrostichus) bistortae, known also from Belgium, Empis s.str. levis and lepidopus, and E. (Xanthempis) testacea. On the other hand, there are four species, E. (Anacrostichus) lucida, Empis s.str. caudatula, E. (Xanthempis) laetabilis and E. (Lissernpis) nigritarsis, known from Scandinavia and also recorded by Collin

(1961) from Great Britain, but not yet found in the Netherlands. Empis s.str. nitidula is the only exclusively Scandinavian species, known so far only from Sweden, and there are three further species, E. (Leptempis) rustica, Empis s.str. staegeri and E. (Xanthempis) univittata, that are known from central Europe and

Figs 3-10. Radial fork (R, and R,) and vein M, in Empis. — 3: E. (Euempis) tessellata; 4: E. (Xanthempis) stercorea; 5: E. (Lissempis) nigritarsis. Axillary excision of wing in Empis. — 6: E. tessellata; 7: E. stercorea; 8: E. nigritarsis. Antenna of Empis. — 9: E. stercorea; 10: E. nigritarsis. Scale: 1 mm, 0.2 mm in Figs 5, 8-10.

18

extend north to Scandinavia but are unknown from both the Netherlands and the British Isles.

—

Key to subfamilies of Empididae Prosternum small, in the form of an isolated sclerite between fore coxae. Empidine-like species with costal vein running round the entire wing as an ambient vein. Male genitalia with well developed unmodified soft cerci o 0 } Sveum cope aeakea decpend a ae Oreogetoninae Prosternum large, fused with episterna... 2 2(1) Costa ending at wing-tip, running to vein R,,. or M,, vein Sc incomplete (except Hilara). Axillary lobe of wing usually well developed, the vein closing anal vein (vein Cu,,) strongly recurrent. Male genitalia with invisible small vestigial cerci, but epandrium (dorsal lamella) cleft and usually remarkably modified .. 0 2 00 Gcatolnci tiecapeee wena ae kar Empidinae Costa running round the entire wing as an ambient vein, vein Sc complete, reaching costa. Axillary lobe and axillary angle not developed, the wings very narrow at base (except Brachystoma) and the vein closing anal cell not very recurrent, usually joining anal vein at GIN CUMGUIS: «oS sh,aeicre nee at eae treat, baron enes 3 3(2) Fore legs raptorial, with coxae very elongated oct} LOCA ae Hemerodromiinae POReMCOSMOULAPtOLal 2... asses sen es 4 4(3) Posterior four legs remarkably shifted posteriorly (as in Hemerodromiinae). Anal cell small, usually rounded apically, or completely ADS CHUM Teh a ene Soe Clinocerinae Posterior four legs not shifted posteriorly. Anal cell very long (in European species) and the vein closing it (vein Cu,,) convex, of the MV DOSAVPC™ ease... Brachystomatinae

Key to genera of Empidinae _Metapleural bristles present (Empidini) .. 2 - Metapleural bristles absent (Hilarini). Vein R,,; forked, radial fork rather long and acute; proboscis short and rather soft, labrum only in exceptional cases as long as head is high oo hod TES R a Sa ee Hilara Meigen 2(1) Vein R,,; simple, not forked. Proboscis rather short and stout (except Aclonempis), usually

shorter than head is high, labella generally soft and stout ..... Rhamphomyia Meigen Vein R,,, forked. Proboscis long and very slender, heavily sclerotised, labrum much longer than head is high, labella slender .... RCA Arar c detnedhe tsaehinietes Empis Linné

Key to subgenera of Empis 1

Axillary lobe of wing well developed, axillary excision acute (Fig. 6); in doubtful cases, then antennal style long, at least half as long as 3rd CLEVER AVSSI Ge eae SeROU aA

CREAR NL

ote oP

2

—

Axillary lobe less developed, axillary excision obtuse, at least 90° or more (Fig. 7). 3rd antennal segment elongate, style very short. Large yellow species (Xanthempis) with long slender legs and no acrostichal bristles, or small black species (Lissempis) with hind tibiae apically and hind basitarsi very dilated ..... 10 2(1) Veins R,,; and M, very diverging at wing-tip, radial fork acute, and vein R, often S-shaped (Fig. 3). Usually strongly bristled species, at least hind tibiae spinose and (except Pachymeria) abdomen with distinct hind marginal DEISULCS ena Ae cot ea se egcis x eckn ohic 3 — Veins R,,; and M, almost parallel towards wing-tip, or at least not very diverging, radial fork not acute, rather ‘open’ (Fig. 4). Not very bristly species, legs covered rather with bristly-hairs, and no hind-marginal bristles on abdominaliterGagner sent taser 6 3(2) Prosternum completely covered with hairs. Dark mesonotal stripes, if present, usually along the lines of bristles! 235-45-..-000- 4 — Only sides of prosternum with hairs, central part between fore coxae bare (several species of the Empis s.str pilosa-group with prosternum completely covered with hairs do not occur in NW Europe and Scandinavia) ..... > 4(3) Eyes meeting on frons in males, females dichoptic. Large to medium-sized strongly bristled species. Legs strong, but hind femora not swollen, and hind tibiae spinose. Abdomen with distinct hind-marginal bristles ........ BE eS RTS BS SEE 1. Euempis — Eyes broadly dichoptic in both sexes. Medium-sized, rather short but densely pubescent species. Legs not distinctly bristled, hind femora often strongly dilated and hind tibiae with a ‘bent knee’ (Fig. 26). No hind-marginal bristles Gn abdomen = eae eerre 2. Pachymeria

19

5(3) All veins from discal cell reaching wing-margin. Dark mesonotal stripes between the lines of bristles (in Scandinavian species), or yellow species. Male: 8th sternum enlarged, genitalia upturned and dorsal lamellae small ............ ibaa od Bee eee ...... 3. Polyblepharis — Upper two veins from discal cell (veins M, and M,) abbreviated, not reaching wing-margin. Dark mesonotal stripes along the lines of bristles. Male: 8th sternum small, dorsal lamellae Very enlaroed ime tee ret te 4. Kritempis 6(2) Prothoracic episterna bare; if fine haris present, then with dark (or, in black species, shining) mesonotal stripes between the lines of bristles. Male genitalia in ventral position, footshaped, aedeagus simple. Generally large speCIES. Sok Gialete dsevags ued a) = eaten oe tate eee 7 — Prothoracic episterna covered with hairs. Mesonotum, if striped, with darker or more shining stripes along the lines of bristles .... 8 7(6) Face bare. Legs uniformly black; if bicoloured, then hind femora thickened and acrostichal bristles absent. Male: ventral genital lamella simple, without appendages. Female: wings not broadened ....... 5. Anacrostichus — Face covered with hairs. Legs bicoloured, contrasting black and reddish-brown, hind femora slender. Male: ventral lamella with two long slender appendages. Female: wings broadened, deep browne oe. eee or 6. Platyptera 8(6) Large species, about 7-10 mm (except E. grisea), often light grey dusted, with labella of proboscis rather soft and covered with hairs about as long as labellum is wide. Male genitalia: lateral lamellae elongate, dorsal lamella deeply bifurcated ancdiwithiLons, OLIStLeS ae ETP ES AOS 2 re Sipe RES ARO 7. Leptempis — Generally small (3-5 mm), usually black to blackish-grey coloured species, labella narrowly pointed, heavily sclerotised, covered with sparse minute hairs. Male genitalia differently shaped 9 9(8) Vein M, (the upper vein from discal cell) complete, reaching wing-margin ... 8. Empis s.str. — Vein M, abbreviated, not reaching wing-marOUD ete soca loe ORME eee nts 9. Coptophlebia 10(1) Generally large (4-8mm) species with ground-colour yellow. Legs mostly yellow, long and slender, covered with sparse, minute hairs

and bristles. Acrostichal bristles usually absent BI oh. SRRRY Spennlrra Shae 10. Xanthempis — Smaller (about 4 mm) black species. Legs yellow with a black pattern, covered with long dense pubescence, and hind tibiae and tarsi dilated. Acrostichal bristles present (in Scandinavial Species), «cue dah ere 11. Lissempis

20

List of species subgenus Euempis Frey, 1953 1. E. tessellata Fabricius, 1794 2. E. sericans Brullé, 1832 3. E. picipes Meigen, 1804 4. E. ciliata Fabricius, 1787

subgenus Pachymeria Stephens, 1829 5. E. femorata Fabricius, 1798 6. E. scotica Curtis, 1835 7. E. tumida Meigen, 1822

subgenus Polyblepharis Bezzi, 1909 8. E. opaca Meigen, 1804 9. E. strigata Loew, 1867

subgenus Kritempis Collin, 1926 10. E. livida Linné, 1758 subgenus Anacrostichus Bezzi, 1909 11. E. lucida Zetterstedt, 1838

12. E. verralli Collin, 1927

13. E. nitida Meigen, 1804 14. E. bistortae Meigen, 1822

subgenus Platyptera Meigen, 1803 15. E. borealis Linné, 1758

subgenus Leptempis Collin, 1926 16. E. rustica Fallén, 1816

17. E. nigricans Meigen, 1804 18. E. variegata Meigen, 1804 19. E. grisea Fallén, 1816

subgenus Empis s.str. 20. E. lepidopus Meigen, 1822 21. E. albopilosa de Meijere, 1935 22. E. serotina Loew, 1867

23. E. decora Meigen, 1822 24. E. acinerea Chvala, 1985

25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35.

E. E. E. E. E. E. E. E. E. E. E.

syrovatkai Chvala, 1985 genualis Strobl, 1893 pennipes Linné, 1758 rufiventris Meigen, 1838 planetica Collin, 1927 staegeri Collin, 1963 limata Collin, 1927 melanotricha Loew, 1873 tanysphyra Loew, 1873 woodi Collin, 1927 nigripes Fabricius, 1794

36. E. nitidula Zetterstedt, 1859

37. E. laminata Collin, 1927 38. E. gooti sp.n. 39. E. bicuspidata Collin, 1927 40. E. nuntia Meigen, 1838

41. E. chioptera Meigen, 1804 42. E. prodromus Loew, 1867 43. E. dasyprocta Loew, 1867

44. E. caudatula Loew, 1867 45. E. aestiva Loew, 1867

9.

10.

46. E. 47. E. subgenus 48. E. 49. E. 50. E.

praevia Collin, 1927 levis Loew, 1873 Coptophlebia Bezzi, 1909 hyalipennis Fallén, 1816 impennis Strobl, 1902 albinervis Meigen, 1822

number of notopleural bristles (often merging with supra-alar ones) and scutellar, though in some species all bristles including humeral, posthumeral and postalar are well differentiated. Metapleural fan always composed of numerous long bristles.

51. E. vitripennis Meigen, 1822 52. E. volucris Wiedemann, 1822

hairs

always

numerous,

acrostichal

53. E. pilimana Loew, 1869 subgenus Xanthempis Bezzi, 1909

bristly-hairs at sides), or all mesonotal hairs uni-

54. E. scutellata Curtis, 1835

formly spread over mesonotum and not differentiated. Usually three darker stripes on mesonotum, along the lines of bristles. Wings always large, with thickened veins, radial fork very acute, ending well before the wing-tip, veins R,,; and M, broadly diverging. Costal bristle strong. Anal vein (vein Cu) always complete. Axillary lobe well developed, axillary excision acute. Legs strong, generally densely bristled, especially on tibiae, shorter bristles usually present on femora beneath. The bristles are always more spine-

55. E. laetabilis Collin, 1926 56. E. lutea Meigen, 1804 E. albifrons Bezzi, 1909

11.

Mesonotal

bristles at least irregularly 2- to 3-serial, often pluriserial, generally well separated from multiserial dorsocentrals (which usually merge into similar

E. 57. E. 58. E. 59. E. 60. E. 61. E. 62. E. 63. E. 64. E. subgenus 65. E.

rohaceki sp.n. concolor Verrall, 1872 stercorea Linné, 1761 aemula Loew, 1873 univittata Loew, 1867 trigramma Wiedemann, 1822 punctata Meigen, 1804 testacea Fabricius, 1805 digramma Meigen, 1835 Lissempis Bezzi, 1909 nigritarsis Meigen, 1804

Subgenus Euempis Frey, 1953 Euempis Frey, 1953:32 (as subgenus of Empis). Type-species: Empis tessellata Fabricius, 1794 (orig. des.). Generally large (well over 10mm, the smallersized species about 5 mm long), robust, densely pubescent and long-bristled species. Head distinctly higher than wide. Eyes in male either touching for a long distance and then upper ommatidia often slightly enlarged (though never as much as in Empis s.str.), or touching at one point, or ‘nearly’ touching. In most cases all ommatidia equally small as in broadly dichoptic females. Antennae distinctly bristled on basal two segments, 3rd segment never very elongated, style rather long, sometimes not very much shorter than 3rd segment. Palpi long, slender, more or less bristled, sometimes very densely so, especially beneath. Proboscis strong, labrum distinctly longer than head is high, often twice as long. Thorax robust, humeri and postalar calli dis-

tinct. Pronotum in the form of a discrete curved sclerite, not bilobed, with long bristles, prosternum

completely covered with hairs. Large strong bristles sometimes not well differentiated from numerous densely-set long bristly-hairs, except for a

like than usual, strong and rather short, hind tibiae

with a posteroventral comb-like row of spines. Abdomen more or less densely pubescent, hindmarginal bristles well differentiated and strong in males, in females often present on Ist tergum only. There are two types of male genitalia in the ground-plan: (1) the E. tessellata-group, with large compact genitalia, where the small simple dorsal lamellae and a rather short stout aedeagus are completely hidden within the enlarged triangularshaped lateral lamellae. E. picipes belongs to this group. (2) the E. ciliata-group, with small lateral lamellae, but dorsal lamella broadly cleft and often bifurcate, clearly visible and of complicated structure, aedeagus thin and very long. There are some intermediate forms (for instance E. sericans) that show tendencies towards the ground-plan condition of Pachymeria. On the other hand, the type of male genitalia of EF. ciliata, E. kerteszi and others

closely resembles that of Leptempis. Collin (1961) has characterised Euempis (together with Pachymeria) as having dark mesonotal stripes along the lines of bristles (i.e. acrostichal and dorsocentral bristles), unlike the subgenera Polyblepharis, Platyptera and Anacrostichus where the dark stripes should be between the lines of bristles. However, this only applies to a few British species and seems to be a somewhat misleading character. For instance, E. ciliata has polished stripes between the lines of bristles (even if only theoretically, as the hairs are evenly spread over the mesonotum), and in Polyblepharis the position

of the dark mesonotal stripes alters from species to

21

vTmit

. poe OB TTT ray TAT ae

Ras Ws

f

eo eer

4

ASRS MSE

eed

Fig. 11. Empis (Euempis) tessellata F., male. Total length: 9-11 mm (A. Veltman del.).

species, sometimes even changing according to the point-of-view. Phylogeny and classification. In view of their robust size, the almost uniform and still largely nondifferentiated pubescence and bristling on body and legs, and their biology (prey presentation during mating has not yet developed), Ewempis species are regarded as the most primitive members of the genus Empis. The species classified in Euempis are rather uniform, and the two groups of species,

oe

the tessellata- and ciliata-groups, are hardly separated on the basis of the structure of the male genitalia. However, it seems very probable that Euempis includes rather relict primitive forms from which at least two evolutionary lineages have developed: one, represented by E. tessellata, led to Polyblepharis and its allies, and the second, the E. ciliata-type, to Leptempis. Collin (1961), pointing out that the thoracic prosternum is completely pubescent in Euempis and Pachymeria (though this character is also pre-

Figs 12-14. Antennae of Empis subgenus Euempis. — 12: E. tessellata; 13: E. sericans; 14: E. picipes. Scale: 0.2 mm.

sent in some species of Empis s.str), classified these two groups in the single subgenus Pachymeria (the oldest available name). However, I do not think that this synonymy was well-founded because Pachymeria (as well as Xanthempis) seems to be-an exceptionally well-defined monophyletic group with Empis (and Rhamphomyia) and certainly deserves subgeneric status.

Distribution. The centre of distribution of Euempis seems to be in the southern warm regions of the Mediterranean, including north Africa and the Near East. Of the twenty two Palaearctic species, four are known from Jordan and Palestine, and

such as hawthorn blossom, and also blossoming fruit-trees; they are undoubtedly effective pollinators, thanks to the long proboscis which gains substantially in length when the labium is extruded. Euempis species clearly belong to the group of Empis where mating in species-specific swarms has not yet been fully developed and, therefore, all intermediate habits can be observed. In the primitive forms where both sexes are both predators and nectar feeders, copulation is achieved without prey transfer, but in species that visit flowers alone the males capture prey that is presented to the female during copulation.

seven from the Far East, two being endemic to Ja-

pan. Most of the European species are known from SW Europe (including several new and still undescribed species), from where they have expanded into the warmer parts of central Europe and along

Key to Euempis species 1

Halteres pale, metapleural bristles black. Thoracic spiracles pale yellow. Body dulled by greyish or brownish dust, abdominal pubescence black. Female legs not pennate ...

—

Halteres black, metapleural bristles partly whitish. Thoracic spiracles blackish. Body shining black, base of abdomen with whitish

the Atlantic coast of NW Europe, to the Netherlands, and even to Denmark and southern Sweden.

Two species only, E. picipes and the widely distributed E. tessellata, are native to northern Europe.

Biology. The adults are very good fliers. They fly very rapidly and for long distances, and when carrying large prey the males look very much like asilids. They are predators, preying on whatever insects are available, and the prey is sometimes almost as large as the fly itself. However, both sexes are also frequent visitors to flowers, and their den-

_sely pubescent body can become coated with pollen grains. The adults of the large-sized species often visit small flowers such as dandelion, bushes

hairs. Legs black, pennate in female. Acros-

tichal and dorsocentral bristles uniformly distributed over mesonotum.

Large, 9-12 mm

lonlgeegreh) te okey eee 4. E. ciliata 2(1) Large species, 8-12 mm. Palpi black, 4th tergum simple in male. Antennae moderately long, lst segment at least twice as long as deep —

Small species, about

5 mm.

Palpi brown or

23

yellowish apically, 4th tergum in male with conspicuous lateral projections. Antennae short, lst segment as long as deep and as long as 2nd segment. Legs mostly brownish-yellow on femora and tibiae ......... 3. E. picipes 3(2) Larger, robust species about 9-12 mm long. Legs more or less yellowish-brown, at least tibiae brownish. Wings deep brownish in both sexes. Male genitalia large and compact, aedeagus not visible, short and stout.........

—

se ata atebrs aloPRES cela: ordasloet TaNSFen 1. E. tessellata Generally smaller, about 8-10 mm long. Legs uniformly black, subshining. Wings brownish in male, almost clear in female. Male genitalia small, aedeagus long and very thin, black, clearly visible ete-. ee 2. E. sericans

1. Empis (Euempis) tessellata Fabricius, 1794 Figs 3, 6, 11-12, 15-16.

strong black bristles merging posteriorly with 3 to 4 supra-alar bristles, usually 2 strong postalar bristles; scutellum with 6 to 10 strong marginal bristles. Metapleural fan composed of at least 15 very long black bristles and further numerous finer bristlyhairs. Prosternum densely covered with long black bristly-hairs, sides with longer and coarser bristles. Wings light brownish with a tendency to be yellowish along costal margin, veins dark brown. Squamae and halteres yellowish. Legs usually black on coxae and femora, yellowish on tibiae and base of tarsi, tips of tarsi blackish.

The leg colour varies from very dark specimens having only tibiae dark brown, to pale-legged forms with legs completely yellowish except for dusky coxae. Fore femora with long hairs beneath, posterior four femora with short stubby spines in two ventral rows, and all tibiae with longer spinose bristles in two rows dorsally, on mid tibiae also

Empis tessellata Fabricius, 1794:405. Empis fraterna Loew, 1865:239. Empis nigripes Strobl, 1880:9 (as var. of tessellata). Empis tipuloides: Strobl, 1893:63 (as var. of tessellata). Empis genuina Strobl, 1893:63 (as var. of tessellata).

ventrally, all spines slightly longer than tibiae are deep. Abdomen black, terga covered with dense greyish-bronze to brassy tessellate tomentum; pubescence rather long, black, hind-marginal bristles strong. Genitalia subshining black, compact; large triangular lateral lamellae covered with microscopic reddish pubescence and long black hairs, dorsal lamellae small and narrow, closely attached

Empis atripes Strobl, 1893:63 (as var. of tessellata).

to upper margin of side lamellae; aedeagus yellowish, short and rather stout, concealed within lamel-

Very large, robust species, generally well over 10 mm, with brown wings, pale halteres, and black metapleural fan. Body strongly black bristled, dull, abdomen marbled bronze-grey. Legs varying in colour, at least tibiae brownish. 3. Eyes meeting for a short distance, upper ommatidia very indistinctly enlarged. Face broad, dull grey. Occiput dull blackish-grey, densely covered all over with long bristly-hairs, those below neck finer. Antennae black, lst segment about 3 times as long as deep, with long black bristles; 3rd segment slender, as long (without style) as basal two segments combined, style nearly half as long. Palpi black, covered with fine black bristly-hairs, much longer ventrally. Labrum almost black, much longer than head is high. Thorax

black

in ground-colour,

mesonotum

dark brownish-grey dusted with three broad, velvety-black stripes along the lines of bristles, pleura lighter grey; spiracles yellowish. Acrostichal bristles irregularly 5-serial, separated by bare stripes from multiserial dorsocentrals, all long (about as

long as antennal style) and rather hair-like. Humeri densely covered with numerous long black bristly-hairs, notopleural depression with more than 10

24

lae. Length: body 9-11 mm, wing 9.3—-11 mm. ?. Eyes widely separated by greyish dusted frons, sides along orbital margins with a row of black adpressed bristly-hairs. Tibiae with obviously longer and more numerous black spinose bris-

tles, and with additional ventral rows on fore and hind tibiae. Wings as in male. Abdomen with a similar pattern but covered with shorter pubescence, tapering apically. Length: body 9-13 mm, wing 8.7-11 mm. A very variable species as regards the leg colour, as clearly reflected in the list of colour varieties described by Strobl. There is a large literature discussing the pale- and dark-legged forms of E. tessellata in temperate and cool Europe, which has recently been summarised by Laurence (1992). The pale-legged form is dominant in southern areas and is frequently misidentified as E. morio Fabricius; however, the latter is exclusively a Mediterranean species known from Spain east to Lebanon, and can be separated immediately by its yellow palpi. Note. The types in the Fabricius Collection in Co-

Figs 15-16. Male genitalia of E. (Euempis) tessellata F. (CZ, Bohemia, Davle). — 15: postabdomen; 16: aedeagus with

hypandrium. Scale: 0.5 mm.

penhagen are lost, but FE. tessellata has probably been correctly recognised, which was also Collin’s (1961:507) opinion. Empis fraterna Loew was described from Georgia (Kutaisi), but in the Loew Collection in Berlin there is a single female labelled ‘Amasia/10476’, originating very probably from Turkey (Amasya). Distribution. Common

in southern and central

parts of Scandinavia; more than 600 specimens have been examined. In the north rather rare, dis-

tributed approximately to 66° N, in Norway to Ns, in Sweden to As. Lpm., and to ObN in Finland. With an extensive flight-period, with dates ranging from 9 May (Norway, HOi) to 19 August (both in the north and south), but mainly in June and in the first half of July. — Throughout the Palaearctic region from Portugal (absent from Canary Islands) through North Africa east to Japan. In temperate Europe very common mainly in May and June, in Portugal even at the end of March. Biology. The adults are nectar feeders and are commonly found on flowers, but apparently only the males are predators, catching the prey (usually dipterous) which is transferred to female during copulation, though the male itself sucks the prey when at rest. Theré are many published records on - the feeding and mating habits of E. tessellata (Poulton 1913, Gruhl 1924, Laurence 1950, Parmenter _ 1951, Smith 1952, Hobby and Smith 1961) and it can

be concluded, in agreement with my own observations, that males hunt larger prey that is often as

large as they are themselves or even larger; that they await females in unorganised aerial aggregations of several individuals above bushes or small trees; that pairs meet in the air and then settle on nearby vegetation; and that females suck the prey during copulation. Copulation has been observed to last from 19 minutes to more than half-an-hour (Hamm 1909, Gruhl 1924). It is thought that the female herself does not hunt prey (Hobby and Smith 1961).

2. Empis (Euempis) sericans Brullé, 1832 Figs 13, 17-18. Empis sericans Brullé, 1832:298. Empis nepticula Loew, 1869:259.

Resembling a small E. tessellata but legs completely black, mid tibiae with longer spine-like bristles dorsally and female wings almost clear. Aedeagus in male long and thin, black, free and clearly visible. 3d. Head black, occiput blackish-grey dusted and with dense long black bristles, eyes practically touching on frons, with upper ommatidia only slightly enlarged. Face very broad, dark grey dusted. Antennae black, Ist segment about twice as long as deep and with long bristles above, 3rd segment

rather shorter and broadened,

style two-

thirds as long as 3rd segment. Palpi very long (as long as antennae without style) and very slender, with long black bristles beneath, the longest bristles as long as palpus. Labrum polished black, al-

25

Figs 17-18. Male genitalia of E. (Euempis) sericans Brullé (NL, Oosterbeek). — 17: postabdomen; 18: aedeagus. Scale: 0.5 mm.

most twice as long as head is high, though proboscis much longer if labium is porrect. Thorax more uniformly darker grey dusted than in E. tessellata, even on pleura. Mesonotum with three dull black stripes when viewed from above and from behind, which disappear in anterior view. Black mesonotal hairs and bristles as in E. tessellata but acrostichal bristles less numerous, irregularly 3- to 4-serial, usually only 4 strong notopleural and only 4 or 6 scutellar bristles. Prosternum covered with long black hairs, metapleural fan composed of numerous very long black bristles. Spiracles pale yellow. Wings faintly brownish-grey clouded, costal cell darker brown, and a distinct brown stigma at tip of

vein R,. Squamae light brown, margins not fringed. Halteres yellowish-brown, knobs clear yellow. Legs uniformly black, subshining. Femora with rather numerous dense hairs ventrally, except for an anteroventral row of short black spines on hind femora. Fore tibiae with a row of long postero-dorsal (or rather posterior) black bristles, about twice as long as tibia is deep; hind tibiae with similar spinose bristles antero- and posterodorsally, but mid tibiae more bristled also antero- and posteroventrally, the 4 to S spine-like bristles in posterodorsal row the longest, at least 3 times as long as tibia is deep. Posterior four basitarsi spinose beneath, and all tibiae and basitarsi with circlets of preapical spines. Abdomen dull black with greyish-bronze tessellations, distinct black hairs at sides of anterior two

marginal bristles. Genitalia small, black; dorsal lamellae simple and very small, closely attached to upper margin of small rectangular lateral lamellae, that are level with only upper half of 8th segment; lateral lamellae with a ventral row of long black bristly-hairs, otherwise covered with minute hairs. Aedeagus free, black and very thin, slightly undulating below and ending in a very slender tip above lamellae. Length: body 8.2—9.6 mm, wing 8-8.5 mm. 2. Frons very broad, dull dark grey, both sides with a row of rather long downcurved black hairs. Palpi with shorter bristles beneath. Wings almost clear, costal cell yellowish. Abdomen similarly coloured and with the same pattern as in male, but entirely bare, only Ist tergum with rather short black hind-marginal bristles. Length: body 8-10 mm, wing 8-8.8 mm. With its pale halteres, black metapleural fan and rather large size, together with black palpi and 3 dark mesonotal stripes, the species needs comparison only with E. tessellata and this was done in the above redescription. I have not seen Brullé’s types but believe that the species has been correctly recognised. Loew (1869) described Empis nepticula from Sicily and Corfu; there are 19 specimens under E. nepticula in the Loew Collection, including

a pair from Corfu (leg. Erber) which are undoubtedly types, and further specimens from Dalmatia, Calabria and Orsova. Distribution. A species of southern distribution,

terga, dorsum almost bare, but all terga at sides

over a wide area from Spain to Greece and the

and all sterna with conspicuous, long black hind-

Black Sea, also known from temperate central Eu-

26

rope and extending northwards perhaps along the Atlantic coast to the Netherlands; not found in Great Britain. It could easily reach Danish Jutland along the North Sea coast, as it has recently been collected in large numbers in Malaise traps, in May 1990, at Wageningen and Oosterbeek in the Netherlands. In May and June, locally common. Nothing is known of the biology.

3. Empis (Euempis) picipes Meigen, 1804 Figs 14, 19-20. Empis Empis Empis Empis Empis Empis

picipes Meigen, 1804:226. sulcipes Meigen, 1822:19. maculipes Zetterstedt, 1842:374. incompta Zetterstedt, 1842:374 (MS name). brevicornis Loew, 1869:263. lasionota Loew, 1869:264.

Rather small (about 5 mm), dull grey species with pale halteres and black metapleural bristles, pubescence and bristles black. Antennae unusually short, and palpi yellowish at tip. Male with a curious, spur-like, long, bristly process at side of 4th

tergum. 3d. Head black, densely dark grey dusted. Eyes meeting on frons for a long distance, upper half of eyes with distinctly though not very enlarged ommatidia (the largest enlargement of ommatidia within Euempis). Face broad, dull grey. Occiput densely covered with rather long black bristlyhairs. Antennae black, Ist segment very small, scarcely longer than deep, 2nd segment globular, both with fine, short, black hairs; 3rd segment broadened and rather short, style about two-thirds

length of 3rd segment. Palpi small, apically yellowish or at least brown, with short bristly-hairs beneath.

Labrum

blackish-brown,

not very much

longer than head is high. Thorax grey dusted on pleura, more brownishgrey on mesonotum, which is rather densely covered with rather long black bristly-hairs, representing irregularly 2- to 4-serial acrostichal and irregularly 3-serial dorsocentral bristles, the latter extending out to humeri and to notopleural depression, and ending in 2 pairs of strong bristles in prescutellar depression. Similar strong bristles are a humeral, 1-2 posthumerals, 3-4 notopleurals, a

supra-alar, a postalar, and 2 pairs of scutellar bristles. Prosternum completely covered with long black bristly-hairs, sides with longer coarser bristles. Metapleural fan black, spiracles pale yellow. Wings brownish clouded, veins dark, and a dis-

tinct deeper brown stigma. Squamae dirty greyish-

brown with short black fringes, halteres with yellowish knobs, stalks brown.

Legs varying in colour to some extent, generally brownish to brownish-yellow with femora more or less darkened or with blackish streaks, tarsi blackish towards tip, and coxae always black, densely grey dusted. Pubescence and bristles black. All femora rather long pubescent but only posterior two pairs with short ventral spine-like bristles. All tibiae and tarsi distinctly spinose, and all tarsal segments with circlets of long preapical spine-like bristles. Fore tibiae with not very long or stout bristles dorsally, but posterior four tibiae spinose both dorsally and ventrally, the spines in posterodorsal row the longest, about twice as long as tibiae are deep, on mid tibiae longer still. Abdomen dull velvety-black when viewed from above, greyish pollinose in side view. Pubescence (particularly long at sides of anterior four terga) and bristles black, hind-marginal bristles conspicuous at sides of all terga and on sterna. 4th tergum broadly excised laterally below and produced into a long, subshining black, pointed process with very long and strong black spines at tip. Genitalia compact, very much as in E. tessellata, but ventral lamella enlarged, polished black, partly shielding base of stout yellowish aedeagus, which (unlike tessellata) is apically almost black, thin and clearly visible. Length: body 4-5.5 mm, wing 4.6-5.6 mm. 2. Palpi yellowish, covered with only fine minute hairs, and all hairs and bristles on occiput, thorax and abdomen distinctly shorter than in male. Thorax more greyish and the dark mesonotal stripes more distinct. Wings paler, rather light greyish, costal cell and stigma yellowish. Legs generally paler than in male, often uniformly yellowish except for darkened tarsi, with similar bristling but posterior femora without the ventral spines. Abdomen more uniformly greyish or subshining black in some views, pubescence much shorter but

hind-marginal bristles distinct. Length: body 5.5—6.5 mm, wing 5.1-5.3 mm. Rather a variable species in leg colour. However, the large number of synonyms has been caused by its small size and its general resemblance to the appearance of Empis s.str. species, with which it is often misidentified. In keys to Empis s.str., E. picipes runs readily to E. pilosa and its allies. The wing venation with an acute radial fork and diverging veins R,,; and M,, and spinose tibiae, provide decisive support for its assignment to Euempis. Even the structure of the male genitalia clearly shows its close relationship to E. tessellata. Type revisions and lectotype designation. Mei-

PE

Figs 19-20. Male genitalia of E. (Euempis) picipes Meigen (S, Vstm: Sala, Vasterfarnebo).—19: postabdomen; 20: aedeagus with hypandrium. Scale: 0.2 mm.

gen’s types in Paris have been revised by Collin (1961). Zetterstedt described this species as E. maculipes from several localities in Sweden. There are 2 males and 3 females from the original syntypic series in the Zetterstedt Collection in Lund, 2 pairs from ‘Ostrogothia’ and a male labelled ‘Gestr.’ (Gestricia), all conspecific and identical with E. picipes Meigen. A male labelled ‘Fredensborg’ and ‘E. maculipes do. Ostr.g.’ is herewith designated as lectotype of Empis maculipes Zetterstedt, 1842, and was labelled accordingly in 1992. I have seen also the types of Loew’s two species in Berlin, those of FE. brevicornis a pair from Bavaria (Augsberg, Kittel) and of E. lasionota from Munich (Bavaria) and Corfu (leg. Erber), and they all are also identical with E. picipes; lectotypes have not been designated.

species and often misidentified in northern collections; specimens in the Helsinki Finnish Collection were even identified as Empis livida, or Hilara gallica. On dates ranging from 15 May to 10 July, but mainly in June. — In Great Britain only from England (Collin 1961), widely distributed in temperate central parts of Europe but not at all common,

mainly in May. The only available records from southern Europe are from Italy and Greece. Biology. The adults are frequently found on flowers sucking nectar. Mating has not been observed,

but Smith (1952) observed a pair in copula with a Liriomyza sp. (Agromyzidae) as prey.

4. Empis (Euempis) ciliata Fabricius, 1787 Figs 21-22.

Distribution. A species of southern distribution in Scandinavia, rarely extending to approximately 60° N, in Sweden to Dir. and Gstr., in Finland to St and Ta. Rather uncommon, with only 124 specimens examined, though apparently commoner in eastern areas as 65 specimens were collected in

Empis ciliata Fabricius, 1787:365. Empis laurata Villers, 1789:571.

Empis boja Schrank, 1803:170. Empis elongata Meigen, 1838:84.

southern Finland. There is a male from the Bidenkap Collection in Bergen, with an identification la-

bel ‘Empis grisea’. According to Lita Greve Jensen (letter of 10.2.1989) this is very probably a specimen from Vestfold, recorded by Bidenkap (1892) under ‘Empis grisea Fall.’. Jonassen (in litt.) recorded it in Norway from @stfold. An overlooked

28

Large (9-12 mm), uniformly black species with black halteres and legs, female legs pennate. Thoracic pubescence long, black, but metapleura and base of abdomen with mostly whitish bristly-hairs. 3d. Eyes meeting at one point, all ommatidia equally small. Face very broad, subshining black,

as well as the densely black pubescent occiput. Antennae rather small, Ist segment twice as long as

deep, 3rd segment slender, style two-thirds as long as 3rd segment. Palpi black, with long but fine black bristly-hairs beneath. Labrum_blackishbrown, nearly twice as long as head is high. Thorax uniformly subshining black even on pleura, mesonotum densely covered with uniform black hairs about as long as antennal style, the same pubescence also on humeri; acrostichal and dorsocentral bristles not differentiated. Mesono-

brown with a tuft of dark hairs in a comb at tip above, halteres black. Legs completely black and almost shining, densely short black pubescent and spinose. Femora beneath with rather spine-like bristles, the charac-

teristic spines in rows only on tibiae dorsally, all scarcely longer than tibiae are deep. Tibiae and all tarsal segments with circlets of preapical spines, and basitarsi spinose beneath, though the spines are not as strong and conspicuous as in E. tessellata.

tum unstriped, though viewed from in front there

Abdomen shining black, dorsum short black pu-

are two narrow, more shining black stripes lying (theoretically) between the hypothetical acrostichal and dorsocentral bristles. Large bristles confined to about 12 bristles in notopleural depres-

bescent, but sides covered with long hairs, those on

sion, a postalar and about 6 to 10 scutellar bristles

intermixed with long black hairs. Prosternum completely covered with dense black pubescence, metapleural fan composed mainly of whitish bristles, or black bristles becoming white at tips. Spiracles black. Wings very faintly greyish-brown clouded, rather light, with a contrasting deep brown costal cell and almost black costal stigma. Squamae blackish-

anterior two segments and on venter whitish; hindmarginal bristles black, merging with the long pubescence. Genitalia elongate, of the Leptempistype, with a thin dark aedeagus clearly visible below, and even above the curiously bifurcate dorsal lamellae. Lateral lamellae pointed posteriorly, polished black, ventrally with dense long black bristles. Length: body 9.5-12.2 mm, wing 9-10.5 mm. 2. Everywhere shorter pubescent, and consequently more shining black, mesonotum with two polished black stripes clearly visible and extending from pronotum to scutellum, and with two addi-

pe} Z, YA

; Figs 21-22. Male genitalia of E. (Euempis) ciliata F. (NL, Tongeren). — 21: postabdomen; 22: aedeagus with hypandrium,

lamellae dotted. Scale: 0.5 mm. :

PAS)

tional shorter and much broader polished stripes behind humeri. Frons broad, subshining black, gradually widening below and continuous with face, margins with short black hairs. Wings decidedly deeper brown than in male, but the darkened costal margin also distinct. Posterior four femora and hind tibiae broadly black pennate on both sides, the tibial spines being retained as they were in male, mid tibiae with several flattened hairs at base beneath. Abdomen practically bare and polished black on dorsum, sides of basal two terga with rather long whitish pubescence, sterna greyish dusted and finely pale pubescent. Length: body 9-13 mm, wing 9.3-11.5 mm. A very characteristic species with a quite different, Leptempis-like male genitalia and pennate legs in female, the most closely related and very similar species being the central European E. kerteszi Bezzi. Distribution. A similar type of distribution to E. sericans, rather common in SW Europe, widely distributed in temperate central Europe, where it is locally common, especially in southern Germany (Bavaria), and extending probably along the Atlantic coast to the Netherlands. Its occurrence in the southern parts of Scandinavia, in Denmark in particular, seems very probable. A spring species, in May and June.

Type-species: Empis ruralis Meigen, 1822 (automatic). Medium-sized (generally about 5-6 mm long), rather uniformly black species with predominantly short dense pubescence, not at all strongly bristled. Eyes with all facets equally small and broadly separated on frons in both sexes. Hind femora often swollen, and male genitalia often unusually large, with an unusually long thin aedeagus. Head small, eyes dichoptic in both sexes. Antennae with rather short 3rd segment, style half as long as this. Proboscis about twice as long as head is high, but very much longer if labium is porrect. Thorax rather densely short pubescent, marginal bristles not very prominent; acrostichal bristles usually 2- to 3-serial and distinctly separated from the more numerous dorsocentral bristles, all rath-

er small and hair-like. Prosternum between fore coxae completely covered with hairs, metapleural bristles always long and numerous. Wings very much as in Euempis, mostly clear with distinct dark veins, radial fork very acute,

Subgenus Pachymeria Stephens, 1829

veins R,,; and M, diverging at wing-tip; anal vein complete, reaching wing-margin, and axillary lobe large, consequently axillary excision very acute. Anterior four femora slightly thickened but hind femora very distinctly so and usually spinose beneath. Tibiae always slender, only finely pubescent, without spines or spine-like bristles. Hind tibiae usually characteristically connected by a ‘bent knee’ with the swollen hind femora. Abdomen clothed with short hairs or almost bare, no hind-marginal bristles, and always differently shaped in males and females. Female abdomen practically devoid of pubescence, either densely silvery-grey pollinose, or with bare polished patches. Male genitalia very conspicuous and of almost uniform structure in all species: dorsal lamellae small, simple, attached above at base of the characteristically-shaped large lateral lamellae, which may be very elongate, flattened, somewhat bean- or boat-shaped, covering the entire thin and very long aedeagus, or lamellae shortened beneath with base of aedeagus clearly visible.

Pachymeria Stephens, 1829:262. Type-species: Empis ruralis Meigen, 1822 (by monotypy) = Empis femorata Fabricius, 1798. Pachymerina Macquart, 1834:333. Type-species: Empis femorata Fabricius, 1798 (orig. des.). Zapolia Gistl, 1848: x (new name for Pachymeria Stephens).

Phylogeny. Pachymeria species clearly form a monophyletic group within Empis of specialised forms with a series of autapomorphies, but also sharing a number of very primitive plesiomorphous characters and having much in common with the subgenus Euempis. Unlike the latter, the species of Pachymeria have attained a high level of specialisation in courtship, forming aerial swarms

Biology. The adults frequently suck the sweet nectar of flowers, commonly hawthorn blossoms, but they undoubtedly prey on other insects as well. The mating behaviour and swarming have not been observed, but the epigamic behaviour of the very closely related central European E. kerteszi has been observed by the present author in May 1964 in southern Slovakia: males swarm during the day on hill-tops between tree canopies, in large dispersed swarms without prey, and readily mate with females, entering the swarm.

30

Fig. 23. Empis (Pachymeria) femorata F., male. Total length: 4.5-7.2 mm (A. Veltman del.).

with astable organisation and making with parallel prey presentation by male to female.

Denmark and the south of Scandinavia is extreme-

ly likely. The rather boreomontane E. scotica may well also occur in central Scandinavia.

Distribution. Thirteen species of Pachymeria are known

from

the

west

Palaearctic,

distributed

mainly in southern Europe, North Africa, and extending eastwards to steppe and semidesert biotopes in Russia and the Near East. Five species are known in temperate central Europe, but two are exclusively mountain species in the Alps (E. picena Bezzi and E. ringdahli Collin). The remaining three species are distributed further northwards, especially along the Atlantic coast to Great Britain and NW Europe. The occurrence of all three in

Biology. So far as is known, Pachymeria species are nectar feeders during their adult life, and are commonly found on flowers. Their short-pubescent body is often completely covered with pollen grains, even in pinned specimens in collections. Predation is apparently restricted to the mating period when males offer females large items of edible insect prey, often as large as the male himself. Some species, such as the common European E. femorata, inhabit shaded biotopes, particularly the

31

margins of deciduous forests, but some are characteristic inhabitants of open biotopes such as steppes, grasslands and even mountain meadows.

—

Paipi bright yellow to orange-yellow, small, clothed with sparse dark bristly-hairs .... 2 2(1) Halteres yellow. Abdominal pubescence pale, acrostichal

Note. The three Pachymeria species treated in this monograph have not yet been found in Scandinavia or Denmark, although their occurrence there is to be expected. As all three have been found in Great Britain and have been fully redescribed by Collin (1961), only brief characterisations are given here.

—

and dorsocentral

bristles small,

dark and hair-like. Hind femora very thickened, black, brownish at extreme tip. Metapleural fan composed of black and pale hairs, with pale bristly-hairs often dominant. Male genitalia very large, boat-shaped, aedeagus practically invisible ........ 5. E. femorata Knobofhalteres blackish. Abdominal pubescence

black,

acrostichal

and

dorsocentral

bristles black and bristle-like. Hind femora

: 2 Key to Pachymeria species 1‘

Palpi black to densely black dominal terga rather small, basal half sca

blackish-brown, very long and bristled inamale>Basalitwo'aba with black hairs. Male genitalia aedeagus clearly visible in its \.Aeitetere eee 6. E. scotica

rather slender, curved, brownish at least on apical third. Metapleural fan composed of entirely black bristly-hairs. Male genitalia small, aedeagus clearly visible in its basal half .... ee oe eee eet 7. E. tumida

Figs 24-30. E. (Pachymeria) femorata F. — 24: wing-tip; 25: antenna; 26: hind leg in anterior view. E. (Pachymeria) scotica Curtis. — 27: antenna; 28: hind leg in anterior view; 29: palpus. E. (Pachymeria) tumida Meigen. — 30: antenna. Scale: 0.5 mm, 0.2 mm for antennae and palpus.

32

5. Empis (Pachymeria) femorata Fabricius, 1798 Figs 23-26, 31-32. Empis femorata Fabricius, 1798:568. Empis ruralis Meigen, 1822:40. Empis quinquevittata Macquart, 1827:129. Dark grey to silvery-grey species with short, hairlike bristling on mesonotum, very thickened hind femora, and unusually large male genitalia. 3. Palpi yellowish-brown, covered with sparse, minute black hairs. All hairs and bristles on head and thorax black, rather fine and short, except for about 3 notopleurals, a supra-alar, a postalar, a prescutellar pair of dorsocentrals, and 6-10 scutellar bristles. Broad side-margins of occiput covered with dense and distinctly flattened bristles. Mesonotum with three blackish stripes (more distinct when viewed from behind) along the lines of the irregularly 3-serial acrostichal and 4-serial dorsocentral bristles, all very diverging. Metapleural fan varying in colour from entirely black (central Europe) to almost wholly pale (Great Britain and NW Europe). Wings clear, squamae and halteres very pale yellow. Legs black on coxae and femora, but tips of femora and all tibiae and tarsi brownishyellow. Hind femora very thickened, ventrally with black spine-like bristles, hind tibiae bent at knee

and very slender. Abdomen almost velvety-black or subshining when viewed from above, glittering light grey when viewed from the side due to the microscopic greyish pubescence on terga 3-7, basal two segments with pale hairs on sides. Genitalia conspicuously large, lateral lamellae greatly produced posteriorly, somewhat laterally compressed, shielding the extremely long and thin wire-like aedeagus. Length: body 4.5—7.2 mm, wing 4.3-6 mm. 2. Everywhere

with shorter hairs and bristles,

hind femora less swollen and legs often darker, uniformly more brownish, even the black parts of femora often blackish-brown. Abdomen silverygrey, terga narrowly polished black on anterior margins. Length: body 4.5-6.5 mm, wing 4.5-6.5 mm. Varying greatly in size in both sexes, and even in a single population the smallest specimens are only one-half the size of the large individuals.

Distribution. Great Britain including Ireland, Belgium, the Netherlands, and through temperate parts of central Europe south to France and Italy. It is still unknown from the northern and eastern parts of Europe, but its occurrence in Denmark

and southern Scandinavia may be expected. Very common in temperate Europe, a typical spring species, mainly in May and first half of June. Biology. Inhabiting forested areas mainly in the lowlands, common in deciduous forests, along tracks and in parks. Males produce small stable swarms usually combining several individuals, steadily hovering with their prey under overhanging branches of bushes and trees at a height of 0.5 to 2 metres. The prey is often very large, usually large muscids, anthomyiids, bibionids or therevids. Females entering the swarm copulate with males in the air, though copulation continues on the nearby vegetation. Swarming behaviour has been described, for example, by Gruhl (1924), Laurence (1950, mistakenly as E. grisea Fallén) and Parmenter (1951). The so-called ‘offering’ swarming of individual females may also be observed when males are not available.

6. Empis (Pachymeria) scotica Curtis, 1835 Figs 27-29, 33-34.

Empis scotica Curtis, 1824:plate 18 (nomen dum). Empis scotica Curtis, 1835:plate 18. Pachymeria palparis Egger, 1860:345.

nu-

Differing from E. femorata by generally blacker colour, denser and longer bristling on thorax, and thinner legs, even hind femora. Basal abdominal segments with black hairs and male genitalia very distinctive, aedeagus clearly visible. 3d. Palpi long, black, projecting forwards and with very dense short black bristles. Bristling on thorax longer, acrostichal and dorsocentral bristles longer than antennal style, anterior part of mesonotum, humeri and posthumeral region, covered with very dense, coarse long bristling resembling the flattened bristles on occipital margins. Mesonotal stripes as in E. femorata. Wings almost clear, squamae and halteres pale yellow. Legs similar in colour to E. femorata but more slender, tibiae especially very thin. Hind femora not as swollen and very narrowed on apical third, viewed from above distinctly curved, and with very dense long black bristles anterodorsally. Abdomen rather subshining black when viewed from above, terga 3-7 almost bare and in side view velvety silverygrey as in E. femorata; basal two terga covered with black hairs. Genitalia subshining black, small, lateral lamellae shortened and not extending far backwards, consequently base of the thin blackish

38

Figs 31-32. Male genitalia of E. (Pachymeria) femorata F. (SLK, Muran). — 31: postabdomen; 32: aedeagus with hypandrium. Scale: 0.5 mm.

aedeagus clearly visible. Dorsal lamellae blunter at tip and broader than in E. femorata. Length: body 5—7.5 mm, wing 5.2-6.8 mm. 2. Palpi smaller with a tendency to be brownishblack, covered with only a few black bristly-hairs. Wings very clear. Legs generally darker, more

brownish,

hind

femora

less

thickened,

conse-

quently less curved and without the constriction in apical third, ventrally with several black bristly spines. Similar but shorter bristles also on posterior four tibiae dorsally, the unusual anterodorsal

bristling on hind femora in male replaced in female

Figs 33-34. Male genitalia of E. (Pachymeria) scotica Curtis (SLK, Mala Fatra Mts, Bystfitka). — 33: postabdomen; 34: aedeagus with hypandrium. Scale: 0.5 mm.

34

Figs 35-36. Male genitalia of E. (Pachymeria) tumida Meigen (E, Area di Cadi). — 35: postabdomen; 36: aedeagus with hypandrium. Scale: 0.5 mm.

by a row of about 5 spine-like bristles. Abdomen completely and very densely light silvery-grey dusted and almost bare, except for the black pubescence at sides of basal two terga. Last (8th) segment and cerci black. Length: body 4.8-7.3 mm, wing 4.8-6.3 mm.

grey, the three black mesonotal stripes more distinct when viewed from behind. Acrostichal and dorsocentral bristles rather long and hair-like, very much as in E. scotica, but anterior part of meso-

notum including humeri less pubescent. Metapleural bristles long and numerous, uniformly black. Wings clear, halteres blackish, stalk dark brown.

Distribution. In Great Britain mainly in Scotland, on the continent at rather higher altitudes, and a

typical mountain species in warmer parts of central Europe and in the south (Italy, Albania). It is a common

species in Scotland, and its occurrence

even in the cooler areas of Scandinavia (Norway, Sweden) is very likely. A typical summer species, known to me only from July and August. Biology. In mountain meadows in central Europe, often common at altitudes above 1000 m. Males are often captured with prey and both sexes frequently visit flowers. Epigamic behaviour, however, still unknown.

7. Empis (Pachymeria) tumida Meigen, 1822

Figs 30, 35-36.

Empis tumida Meigen, 1822:39. Pachymeria erberi Nowicki, in Loew, 1873:217.

Resembling E. scotica but palpi orange-yellow, halteres black and abdomen in both sexes covered with black hairs on all segments. 3. Palpi very small, yellowish and almost completely hidden in the mouth-cavity, covered with only a few minute dark hairs. Mesonotum dark

Legs similar in colour to both E. scotica and E. femorata, rather slender, hind femora not as stout as in

E. femorata and not apically constricted and curved as in E. scotica. Legs covered instead with thin bristles or hairs, no spine-like bristles on hind femora beneath, and posterior four tibiae with only short fine bristles dorsally. Abdomen velvetyblack to somewhat subshining black, depending on the angle of view, viewed from above with grey tessellations and paler hind margins to terga; pubescence black, short, long at sides of anterior two ter-

ga. Genitalia superficially resembling those of E. scotica, lateral lamellae shortened at base and ae-

deagus clearly visible, very thin, blackish; dorsal lamellae small and pointed posteriorly, very much as in E. scotica. Length: body 4.7—7.2 mm, wing 5.2-7.3 mm. 2. Closely resembling male, pubescence and bristling scarcely shorter. Legs as in male, even hind femora with a similar slight thickening, though generally darker, the pale parts of legs rather brown. Wings clearly shorter than in male. Abdomen uniformly light grey dusted, with darker tessellations from some angles, all segments covered with short black pubescence (only basal two terga in E. scotica), basal segments with longer black hairs at sides. Length: body 6—7.6 mm, wing 5.8-6.3 mm.

35

Distribution. Widely distributed in Europe, in the south from Spain east to Albania and the Ukraine, but rather rare in central Europe. Collin (1961) recorded it from several counties in England from July to the end of September, and its occurrence in the south of Scandinavia seems likely. A typical summer and late summer species. Biology. According to Collin (1961), males are often captured with prey as large as themselves, but swarming and mating have not been recorded.

Subgenus Polyblepharis Bezzi, 1909 Polyblepharis Bezzi, 1909:95 (as subgenus of Empis). Type-species: Empis albicans Meigen, 1822 (orig. des.). Medium-sized to very large, robust species, antennae and wing venation as in Euempis, but male genitalia including structure of pregenital segments very distinctive. Eyes meeting on frons in male, or closely approximated, or males broadly dichoptic like the females (E. albicans-complex). Upper ommatidia

ss

Fig. 37. Empis (Polyblepharis) opaca Meigen, male. Total length: 6-9.5 mm (A. Veltman del.).

36

never considerably enlarged. Antennae with basal segment longer than deep, and 3rd segment usually rather long, pointed apically, style long, sometimes almost as long as 3rd segment. Palpi small but usually densely pubescent or bristled. The ground-plan condition is the densely pubescent thorax with long, hair-like and unmodified acrostichal and dorsocentral bristles evenly distributed over mesonotum and with stronger bristles hardly differentiated (E. albicans-complex); the specialised forms have prominent and well differentiated marginal bristles, and small acrostichal

and dorsocentral bristles less numerous, even uniserial. Only sides of prosternum covered with bristly-hairs, prothoracic episterna usually bare or with a few minute hairs, pronotum with spinose bristles in a row. Metapleural bristles long, bristlelike, rarely thin (E. strigata-complex). The position of mesonotal stripes varies in Polyblepharis species, although practically all European species have darker stripes between the rows of bristles. Wings with venation as in Euempis, radial fork very acute, veins R,,, and M, distinctly diverging, and the vein R,,, ending far before wing-tip. Costal bristle small or absent. Axillary lobe well devel-

'

oped, axillary excision very acute, wider in the E.

strigata-complex. Anal vein complete, though sometimes only fine. Legs strong, sometimes densely covered with long hairs in addition to the usual spine-like bristles, hind femora rarely enlarged (E. albicanscomplex), but usually all femora long and simple, distinctly spinose beneath, at least on hind pair. Tibiae almost always distinctly spinose, hind tibiae with an apical comb of spines posteriorly at tip. Femora in females often short pennate or fringed ventrally. Abdomen usually with strong hind-marginal bristles, often covered with long dense pubescence, or abdominal pubescence and bristling very reduced. Male 6th tergum always more or less modified, large, usually devoid of pubescence and velvety-grey or microscopically brownish pubescent, very different in colour from other segments. The next two terga reduced, but 8th sternum on the contrary very enlarged and upturned, pushing the genitalia from below so that they project upwards and partly backwards above the modified 6th tergum. Dorsal lamellae not very large but placed dorsally of the large simple lateral lamellae (not in an inner position), ventral lamella very reduced. Aedeagus short and stout in ground-plan, often bulbous on basal half, the apical narrower

: Figs 38-41. Wing venation of Empis subgenus Polyblepharis.— E. opaca — 38: wing-tip; 39: axillary excision. E. strigata—40: wing-tip; 41: axillary excision. Scale: 0.5 mm.

at

Figs 42-46. E. (Polyblepharis) opaca Meigen. — 42: antenna. E. (Polyblepharis) strigata Loew. — 43: antenna; 44: male

genitalia in dorsal view (tip of aedeagus and dorsal lamella). E. (Kritempis) livida L. — 45: antenna; 46: wing-tip. Scale: 0.2 mm, 0.5 mm in Fig. 46.

part bent back and pointing to the rear, though aedeagus usually completely hidden within lateral lamellae. Female abdomen conical, apical two segments narrowing, usually polished and consequently differently coloured, cerci generally unusually short.

Phylogeny and classification. The subgenus Polyblepharis includes primitive forms of Empis having much in common with Euempis and Kritempis, as is indicated by the series of plesiomorphous characters given in the diagnosis above. However, itis not a natural group at all and it is impossible to give areal differential diagnosis based on constitutive characters that would characterise all the species placed in Polyblepharis. Collin (1961:477) recognised three groups of species, represented by E. strigata, E. albicans (the type-species) and E. opaca, but there are many species from the central parts of Asia recently assigned to Polyblepharis that should be separated into further natural groups of species or species-complexes. This clearly indicates that Polyblepharis is a paraphyletic group as even the structure of the male genitalia,

38

and of the aedeagus in particular, is far from being constant (as was supposed by Collin), and that the structure of the male genitalia is not an autapomorphous character of Polyblepharis as it is now understood. The male genitalia and aedeagus are of very different structure in some Asiatic ‘Polyblepharis’ species, which include forms with a slender curved ‘Empis-like’ aedeagus though the structure of the 6th tergum and 8th sternum suggests typical Polyblepharis species. Collin (1961:478) suggested that Empimorpha Coquillett, 1896 (described as a genus) should be accepted as another closely related subgeneric taxon for those ‘Polyblepharis’ species with the prosternum completely covered with hairs; in addition to Nearctic species, this would also include several

eastern Palaearctic species recently placed in Polyblepharis and also the Mediterranean E. hoffmannseggi, discussed under Kritempis. Furthermore, the classification of Polyblepharis

species is also complicated by the (admittedly rare) partial or complete loss of the radial fork on the wings. Superficially, Polyblepharis species resemble species of Rhamphomyia s.str. (including

Dasyrhamphomyia and Eorhamphomyia) in many respects and, when the radial fork is absent, their identification is very difficult for non-specialists. Quite by chance I found recently during a brief review of the types in the Loew Collection in Berlin that Rhamphomyias.str. gentilis Loew, 1871, described from the south of European Russia (Sarepta), is actually a specimen of Empis (Polyblepharis) fallax Egger, 1860, in which the upper vein of the radial fork has been completely lost on both wings — a synonymy discussed here for the first time although it was recorded without explanation in the Palaearctic Catalogue (Chvala and Wagner

1989). Distribution. A Holarctic group; most of the European species of Polyblepharis are of Mediterranean distribution (nearly 30 species), some of them extending to the Near East or, through the lowland steppes of southern Russia, as far east as Middle Asia; 2 species are known from Siberia and 6 from the Far East and/or Japan. Two species in Europe have a truly northern distribution (E. opaca and E. strigata) and are treated here. Five further species are known from temperate Europe: three are typical southern steppe species, extending northwards to the Ukraine, Hungary, Slovakia, Moravia and Austria (E. albicans Meigen, E. eversmanni Loew and E. fallax Egger), whilst two are ecologically very atypical species that occur high in the mountains in alpine meadows (E. crassa Nowicki and E. gravipes Loew). All these species (except E. strigata) have four dark stripes on the mesonotum, with the central two between the lines of bristles. Altogether forty species of Polyblepharis are known to occur in the Palaearctic region, but the group is particularly rich in species in the steppes and semideserts of Middle Asia, as more than 15 new species are included ina

revision of Polyblepharis now being completed by the author. Biology. In the adult stage all species are active nectar feeders and commonly visit flowers. The mating behaviour has been fully described in two species, in E. opaca by Hobby (1932), Laurence (1950) and Hobby and Smith (1962), and in E. albicans by Chvdla (1978). The males perform modified swarms: they do not fly in the usual stable aggregations of the synorchesic type, but they fly individually with their preyin a horizontal (E. opaca) or vertical (E£. albicans) flight above the vegetation, attracting receptive virgin females resting in the grasses. This is undoubtedly an adaptation by species inhabiting biotopes of the open

countryside, without trees or bushes that could be used as swarm-markers. The prey is usually very large, often of the same size as the predator, and cannibalism has been recorded in both species. Although the female sucks the prey during copulation, it is apparently not an essential source of protein for her, as Hobby (1932) observed successful mating in E. opaca even with males carrying inanimate objects, such as flower seeds. It seems that

the large male, carrying an almost equally large prey or some other object, forms a more effective signal for the female who is attracted by the large silhouette of the patrolling male who is actually a moving swarm-marker.

Key to Polyblepharis species 1

—

Larger (6-10 mm) black species, mesonotum blackish-grey with 4 dark brown stripes, the inner two between the lines of bristles. Legs reddish-yellow, tarsi and tips of femora and tiblaciblack#aeneae eres eee 8. E. opaca Smaller (5-6 mm) yellow species, mesonotum with a single narrow dark median stripe. Legs yellow, tarsi brownish ... 9. E. strigata

8. Empis (Polyblepharis) opaca Meigen, 1804 Figs 37-39, 42, 47-48. Empis opaca Meigen, 1804:226. Empis opaca Fabricius, 1805:138 (not Meigen, 1804). Empis affinis Stephens, 1829:263. Empis apteropus Bezzi, 1899:130 (as subsp. of E. opaca).

Large species (6-10 mm), dorsum of abdomen mostly polished black, mesonotum dark grey with 2 dark brown stripes between the rows of bristles. Legs extensively reddish. 3d. Eyes meeting on frons or very narrowly approximated; face dull grey, mouth-cavity polished black. Occiput dusted blackish-grey, with rather dense irregular black bristles, with longer whitish bristly-hairs below. Antennae black. 3rd segment moderately long, about three times as long as deep at base, narrowed apically; style long, not very much shorter than 3rd segment. Palpi reddish-yellow, covered with sparse but rather long, coarse, black bristles. Labrum strong, mostly black, nearly twice as long as head is high. Thorax dark grey dusted even on pleura, mesonotum with two rather broad dark chestnut-brown

Be}

stripes between lines of bristles, and another two broader and shorter ones outside dorsocentral bristles. Humeri polished on the outside, strongly black spinose and with one large humeral bristle posteriorly; 4 equally strong notopleurals, a supraalar, a postalar and usually 6 strong scutellar bristles; no posthumeral bristle. Acrostichal bristles irregularly biserial, dorsocentrals irregularly 4-serial at middle, more numerous anteriorly and almost uniserial behind, all rather short, the former hairlike, the latter bristle-like; last 2 or 3 pairs of prescutellar dorsocentrals long and strong. Sides of prosternum with a tuft of black and white bristlyhairs, pronotum black spinose, sides and adjacent edge of prothoracic episterna with longer whitish hairs, and much smaller fine pale hairs anteriorly in notopleural depression. Metapleural fan composed of black and pale bristles. Prothoracic spiracles yellow. Wings faintly brownish-yellow clouded, more intensely so in costal cell; costal bristle very small. Axillary excision acute, anal vein complete but fine. Squamae and halteres yellowish. Legs reddish-brown to brownish-yellow, only coxae, tips of femora and tibiae, and all tarsi, black. Fore femora with short black antero- and posteroventral bristles, with equally short but more spinose bristles in two ventral rows on posterior four femora; all tibiae dorsally with two irregular rows of black spinose bristles, shorter than tibiae are deep, with distinct circlets of preapical spine-like bristles on tibiae, shorter circlets on all tarsal seg-

ments, and all basitarsi with short, strong spinose bristles ventrally. Abdomen mostly polished black on dorsum, but 1st tergum, basal third of 2nd tergum, narrow sides of the following terga and all sterna densely very light grey dusted; 6th tergum velvety-brownish to greyish, and the enlarged 8th sternum subshining black. Pubescence short, black, hind marginal bris-

tles rather fine, not at all prominent, and sides of anterior segments with longer pale hairs. Genitalia small, upturned, both lamellae subshining and with fine short black pubescence; aedeagus brown, partly visible at base, but the unusually bent tip hidden within lamellae. Length: body 6-9.5 mm, wing 7.2—9 mm. ?. Frons dark grey. All hairs on thorax, legs and abdomen

shorter and less numerous,

but tibiae

spinose almost as in male. All femora with posteroventral bristly-hairs more flattened, mid femora almost distinctly short pennate posteroventrally, but the bristles in anteroventral row unmodified; specimens without pennation or flattened hairs were described as a ‘subspecies’ apteropus. Abdomen with all terga narrowly grey dusted anteriorly, more broadly so at sides, otherwise polished black, and apical three terga and two sterna completely polished black. Length: body 6.8-10 mm, wing 7-9 mm. Empis opaca is a very characteristic Polyblepharis species in Europe, especially because of the contrasting light grey dusting and polished black colour, and the generally short pubescence. The

Figs 47-48. Male genitalia of E. (Polyblepharis) opaca Meigen (CZ, Moravia, Palava). — 47: postabdomen; 48: aedeagus. Scale: 0.2 mm.

40

only very similar and closely related species is the southern European E. fallax Egger, which is a distinct species, as correctly stated by Collin (1961), and not a subspecies of E. opaca as suggested by Engel (1941). For full synonymy, see Collin (1961:482). Distribution. A rare species with a southern distribution in Scandinavia, known to me only from3 3 and5 9 takenin Denmark, and1 2 from the south

of Sweden (Sk., Lomma) on dates ranging from 12 May to 2 June; this roughly corresponds with the short flight period given by Collin (1961) for Great Britain. — A species of cool and temperate Europe, from the Baltic coast south only to central France,

for the polished oral margin) and occiput greyish pollinose, occiput with an irregular circlet of sparse, rather long black bristly-hairs, lower part with a few whitish hairs. Eyes meeting on frons for a long distance, ommatidia on the upper half slightly enlarged, hardly twice as large as those below. Antennae yellowish on basal two segments, 3rd segment black or at most narrowly yellowish at extreme base; Ist segment about twice as long as deep, covered with fine dark bristly-hairs, 2nd segment globular, slightly broader; 3rd segment pointed, almost 3 times as long as wide at base, style as long as 3rd segment. Palpi small, very dark brown, contrasting with the yellow labrum, clothed with several minute dark hairs. Labrum

northern Italy, Austria, and also the northern and

twice as long as head is high, rather slender, and

central parts of European Russia. A typical spring species, in temperate Europe on dates ranging from 21 April to 27 May.

completely shining yellow. Thorax yellow, finely greyish dusted even on pleura, mesonotum slightly more brownish to reddish-yellow, with a narrow, rather indistinct blackish-grey median stripe. Mesonotum almost bare, acrostichal bristles (situated on the dark median stripe) minute, very closely biserial, dorsocentrals only slightly longer, hair-like, ending in a long prescutellar pair, all black. A rather fine humeral bristle with additional small bristly-hairs anteriorly, an equally fine posthumeral, 3 strong notopleurals, one strong and another finer supra-alar, a strong postalar, and only 2 scutellar bristles. Prosternum bare, but lateral lobes with a tuft of fine, pale, rather long hairs, episterna bare, and pronotum with about 10 rather short black bristly-hairs, no spines. Metapleural fan usually composed of 3 long dark brown to blackish bristles, a larger number of simi-

Biology. A species of grassy lowland biotopes, preferring drier open habitats and steppes. The adults are nectar feeders, frequently visiting the flowers of Taraxacum,

Euphorbia

and Ranunculus.

The

mating behaviour was fully described by Hobby (1932) and later by Hobby and Smith (1962). Males fly rapidly with a large prey or even an inanimate object over and about meadow grasses, flying horizontally backwards and forwards low in the herbage or up to about 3 metres high according to Laurence (1950). Receptive females resting in the grasses are attracted by the male display and fly up to join the male in the air, and then the pairs settle. Copulation has been observed to last from 20 seconds to 8 minutes. The prey is mainly bibionids (Bibio, Dilophus) but even includes specimens of its own species (as has also been observed in E. albicans). After copulation, the males swarm again after a short rest and carry the same prey if it was not dropped when the couple separated.

lar pale bristles, and some smaller whitish hairs.

Spiracles whitish. Wings practically clear with brownish, not very dark veins, radial fork not very acute (Fig. 40), costal bristle absent. Axillary lobe well developed but axillary excision not as acute as usual, though less

than 90°. Anal vein fine but complete. Squamae

9. Empis (Polyblepharis) strigata

white with concolorous fringes, halteres whitish-

Loew, 1867 Figs 40-41, 43-44, 49-50.

yellow. Legs completely yellow, only tarsi darker brownish, rather long and slender, but hind tibiae somewhat broadened apically (as deep at tip as femur is wide) and hind basitarsi slightly swollen, as deep as tibia is deep. Anterior four legs almost devoid of bristles, clothed with short hairs only, but tibiae with a few rather fine dorsal bristles and a preap-

Empis strigata Loew, 1867: 159 (@ ). Empis pittoprocta Loew, 1873:229 (¢). Medium-sized (5-6 mm) completely yellow species, but occiput blackish grey. Mesonotum with a narrow dark median line on the closely biserial very minute acrostichal bristles. The spinose legs and the characteristically shaped male genitalia ~ are as in other Polyblepharis species. 36. Head black in ground-colour, face (except

ical circlet of still more distinct black bristles. Hind femora ventrally on apical two-thirds with a double row of black bristles, shorter than femur is deep; hind tibiae with two rows of dorsal bristles,

those in anterodorsal row stronger but hardly as

41

49

Figs 49-50. Male genitalia of E. (Polyblepharis) strigata Loew (CZ, Bohemia). —49: postabdomen; 50: aedeagus, hypandrium dotted. Scale: 0.2 mm.

long as tibia is deep. Fore basitarsi microscopically pale pubescent beneath, but mid and hind basitarsi spinose ventrally, hind basitarsus very distinctly so, and with additional strong adpressed bristles dorsally. Abdomen subshining yellow, 6th tergum densely silvery-grey dusted in some lights. Pubescence pale, rather long at sides, especially on basal segments, no distinct hind-marginal bristles but 1st tergum with a row of blackish bristly-hairs on hind margin, and some dark bristly-hairs also on hind margin of 7th tergum and on the whole surface of the very enlarged 8th sternum. Genitalia Polyblepharis-like, shining yellow, with broadened, dorsally situated and somewhat laterally open dorsal lamellae, and with the characteristically shaped, apically bent and curved aedeagus with an unusual paler yellow shining bulbous widening at middle. Length: body 5.2-6 mm, wing 7-7.3 mm. ?. Frons blackish-grey, clypeus polished black. Resembling male but hind legs distinctly narrower, hind femora with short black bristles ventrally towards tip only, and both hind femur and basitarsus much more slender, the latter more distinctly spinose beneath. Hind tibiae with a very distinct comb-like row of spines at tip beneath as in male. Abdomen almost shining yellow, pubescence very sparse and fine, apical two segments polished dark brown. Length: body 5.5-6 mm, wing 5.6-6.2 mm. Superficially resembling a small Xanthempis

42

species, especially because of its yellow colour and the dark median stripe on mesonotum. However, the morphological characters unambiguously exclude any relationship with Xanthempis. The most reliable distinguishing characters at first sight are the structure of the head, closely set on the thorax, the short 3rd antennal segment with equally long style, the presence of biserial acrostichal bristles, the wing venation, the spinose legs, and the structure of the male genitalia. Holotype identification. Described twice by Loew, first from a female in 1867 from Sarepta (southern Russia) as Empis strigata, and 6 years later from a male as Empis pittoprocta from the ‘Gegend von Kultuk’ near Lake Baikal. In the Loew Collection in Berlin there are 2 d and 3 @ without locality label standing under ‘E. strigata m.’, none of them perhaps the type, and one male labelled ‘Kultuk/10514/Typus’ under ‘E. pittoprocta Lw.’, undoubtedly the holotype. All these specimens are conspecific and belong to the species described here as Empis strigata Loew.

Distribution. Distributed over a wide area, but obviously an overlooked species and often misidentified as Xanthempis. Known from central Europe (Bohemia), northern and southern parts of European Russia, eastwards through Siberia as far as

the Far East. In June 1990 I found a single female of E. strigata in the Palaearctic Collection in Helsinki Museum taken by Hellén at Archangelsk, NW

Russia, included among unidentified Xanthempis

species. Although it has not yet been recorded from Scandinavia,

its occurrence

in the central

parts of Finland and Sweden seems very likely. Obviously rare everywhere as I have seen only single specimens in collections, all captured in June.

Subgenus Kritempis Collin, 1926 Kritempis Collin, 1926:236 (as subgenus of Empis). Type-species: Empis algira Macquart, 1838 (orig. des.).

Generally large, rather less pubescent species with well differentiated bristles, tibiae spinose and both

Fig. 51. Empis (Kritempis) livida L., male. Total length: 7.5—9.3 mm (A. Veltman del.).

43

upper veins from the discal cell (veins M, and M,) often abbreviated. Eyes meeting on frons in male, upper ommatidia usually slightly enlarged (at most twice), females dichoptic. Antennae Euempis-like, with 3rd segment long conical and with a long, rather stout style at least half as long as 3rd segment. Ist segment moderately long, two to three times as long as deep. Palpi small, covered with only a few minute hairs, not. bristled. Proboscis strong, labrum at most twice as long as head is high. Thorax distinctly bristled, mesonotum with well differentiated strong marginal bristles, acrostichal and dorsocentral bristles rather long and hair-like, usually pluriserial; if less numerous, as in E. livida, then dorsocentrals bristle-like. Prosternum bare between coxae, but sides (epimera) with a tuft of

bristly-hairs, and prothoracic episterna usually with a few hairs; pronotum simple, with a single row of short spine-like bristles. Metapleural fan composed of numerous very long bristles. Mesonotum, if striped, with dark stripes along the lines of bristles, alternating with lighter dull grey stripes between the rows. Wing venation in many respects as in Euempis, Polyblepharis or Pachymeria, with radial fork acute and veins R,,, and M, diverging, but at least one of the two upper veins from discal cell not reaching wing-margin, and anal vein (vein Cu) fine and incomplete. Costal bristle long, squamae fringed, and axillary excision very acute. Legs rather long and slender, not as strong as in Euempis or with the incrassate hind femora of Pachymeria. Short pubescence rather sparse and coarse, femora in male without distinct spine-like

bristles beneath, but all tibiae spinose, the spines not at all numerous, but rather long, often much longer than tibiae are deep, particularly on mid legs; femora in female usually short pennate. Abdomen more or less long pubescent in male, at least anteriorly at sides, terga with distinct and usually strong (except EF. livida) hind-marginal bristles. Male genitalia compact, globose and not very large, aedeagus short and often unusually bent or curved, though mostly concealed within very enlarged dorsal lamellae.

phasising the enormous development of the dorsal lamellae of the male genitalia, the abbreviated veins from the discal cell, and the somewhat unusual anterior and posterior bristling (or pennation in females) or fore femora. Collin believed Kritempis, as defined above, to be a subgenus that appears to form a natural group, with the single true European representative E. livida as ‘somewhat aberrant’. He also included in Kritempis the species E. macropalpa Egger, E. sibillina Bezzi, E. macquarti Becker (geniculata Macquart not Zetterstedt) and E. nigrimana Becker. However, the last three species are arranged in the Palaearctic Catalogue (Chvala and Wagner 1989) as ‘unplaced species of Empis’. They all have the characteristically bristled legs of E. algira, but the male genitalia are very different, having a long, slender undulating aedeagus that curves backwards over the abdomen. Moreover, E. macquarti has a small dorsal lamella, and Collin clearly misinterpreted the long bristled hypandrium (ventral lamella) as the lateral lamella. All these species show a closer affinity to Polyblepharis than to Kritempis — if the latter is accepted as a ‘natural group’ represented by E. algira and E. macropalpa. It should be noted that Engel (1941) in ‘Lindner’ placed E. livida in Polyblepharis, an assignment criticised by Collin (1961:473) who diagnosed Polyblepharis mainly by the peculiar structure of the male aedeagus. E. hoffmannseggi Loew (E. grisea Wiedemann not Fallén) is included in the Palaearctic Catalogue as a fourth species of Kritempis, but Iam now almost convinced that, in spite of the abbreviated vein M,, the entirely pubescent prosternum and the structure of the male genitalia indicate Euempis rather than Kritempis. There can be not doubt that all these species are primitive and phylogenetically old forms of Empis, with the wing-venation in a plesiomorphous state like other characters such as the structure of the head and antennae, bristling on body and legs, etc. It is also clear that except for Pachymeria, which is a natural group, the other undoubtedly phylogenetically primitive Empis subgenera Euempis, Polyblepharis and Kritempis are not at all clearly separated and can scarcely be defined as monophyletic groups. Moreover, the abbreviation

Phylogeny and classification. Kritempis is undoubtedly the most problematic ‘subgenus’ within Empis and, in my opinion, includes very heterogeneous forms whose relationships are unclear. The main differential feature used in many classifications seems to be the abbreviated veins M, and/ or M,. Collin (1926) designated the Mediterranean Empis algira Macquart as type of Kritempis, em-

44

of the M

veins occurs widely among unrelated speciesgroups and cannot be regarded as a good taxonomic character. Apart from the species of Coptophlebia and E. livida, itis found as acommon reduction of the wing-venation in mainly tropical and subtropical species (see Smith 1971), and is present for example in sub-Saharan Rhamphomyia species; it

is also believed that many Afrotropical ‘Coptophlebia’ are also wrongly assigned subgenerically.

Distribution. Empis livida is the only species widely distributed in temperate and northern Europe, as all the other species provisionally retained in Kritempis are typical Mediterranean elements, distributed especially along the African coast. Biology. The adults are nectar feeders, often visiting flowers, and pinned specimens in collections are often densely covered with pollen grains. The epigamic behaviour has been observed many times in E. livida, and in general very closely resembles that of Polyblepharis species.

10. Empis (Kritempis) livida Linné, 1758 Figs 4546, 51-53. Empis livida Linné, 1758:604.

before prescutellar depression; dorsocentrals uniserial, longer and bristle-like, more broadly spaced in the row and ending in 2 or 3 strong pairs. Wings faintly brownish, veins M, and M, very thin, disappearing before wing-margin, as well as anal vein. Squamae and halteres light yellowish, the squamae very finely pale fringed. Legs yellowish including coxae, basitarsi very darkened and rest of tarsi black. All femora with extremely short black bristles, fore tibiae practically without bristles, mid tibiae with a few anterodorsal and ventral bristles about twice as long as tibia is deep, and hind tibiae with much shorter

black bristles in two close rows, similar but longer bristles also on hind basitarsus dorsally. All basitarsi spinose ventrally, very distinctly so on hind legs. Abdomen brown, finely pollinose or almost subshining, covered with sparse small black hairs and distinct, even though not very long, hair-like hind-

Asilus tipuloides Linné, 1758:606.

marginal bristles. Genitalia small, globose, formed

Empis Empis Asilus Empis Empis and

mainly by the enlarged brownish dorsal lamella, lateral lamellae small, shining brown, with a tuft of black bristly-hairs, and almost hidden between dorsal lamella and the enlarged 8th sternum. Aedeagus concealed within dorsal lamellae, short and not very stout, tip bent back upon itself very much as in Polyblepharis, pointing to the rear. Length: body 7.5—-9.3 mm, wing 7-8.7 mm. 2. Frons dark grey dusted. Thorax as in male but abdomen dark blackish-grey, greyish tessellate from some angles, pubescence extremely short and fine, hind-marginal bristles rather indistinct. Wings very conspicuously clear, somewhat iridescent and unusually pellucid, veins distinct except for whitish anal vein. Legs coloured as in male but all bristles much shorter, those on mid tibia scarcely as long as tibia is deep, and hind tibiae with an anterior row of very small stubby spine-like bristles. Fore femora anteriorly with a row of short pennate hairs, denser but equally short pennation also on hind femora posteroventrally. Length: body 7.5-10.2 mm, wing 6.8-8.6 mm. This large Empis species can easily be recognised by the abbreviated upper two veins from the discal cell, a character present in the Scandinavian fauna only in small Coptophlebia species. For synonymy, see Collin (1961:476).

constans Harris, 1776:150. fugeo Harris, 1776:150. reticulata Fourcroy, 1785:465. lineata Villers, 1789:571. lineata Fabricius, 1805:141 (not Villers 1789 Meigen 1804).

Empis trilineata Wiedemann, 1818:25 (as trilineata

Pallas). Large (about 8-10 mm) with yellowish legs, wings with the upper two veins from discal cell abbreviated. Male abdomen brownish and wings faintly brown clouded, female abdomen grey and wings clear. 6d. Eyes meeting for a long distance, ommatidia on upper third of eyes almost twice as large as those below. Face dull grey, clypeus polished black. Occiput densely grey dusted, with two circlets of black bristles above neck, becoming irregular, longer and white below. Antennae black, as in Fig.

45, 2nd segment brownish. Palpi small, brownishyellow, clothed with several minute dark hairs.

Thorax grey on pleura, rather brownish dusted on mesonotum,

with three almost black stripes

along the lines of bristles which disappear when viewed from in front; postalar calli and scutellum more or less yellowish. All thoracic bristles black (a humeral with several small bristles, a small posthumeral, 4-5 strong notopleurals, a supra-alar, a postalar, 4-6 scutellars), but small bristly-hairs at sides of prosternum mostly pale, and several long whitish bristles present in the otherwise black -metapleural fan. Acrostichal bristles very small, hair-like, arranged in two close rows and ending

Distribution. Very common in southern and even central parts of Scandinavia; more than 750 specimens have been examined. The northern limit of its distribution lies at approximately 64° N, in Norway in NT and Nsy, in Sweden in Jmt., in Finland in Oa, Tb, Sb (Kiuruvesi) and Kb; common

also in

45

Figs 52-53. Male genitalia of E. (Kritempis) livida L. (CZ, Bohemia, Davle). — 52: postabdomen; 53: aedeagus. Scale: 0.5 mm.

Russian Karelia. A typical summer species, from June to the end of August, though mainly in July and rarely even in September. The earliest captures are from 5 June (Smaland and Russian Karelia), and the latest dates were recorded in Finland,

on 20 September (Ka, Vehkalahti) and on 26 September (Ta, Forssa). — Distribution in Europe very similar to Empis (Platyptera) borealis: common and widespread throughout Great Britain, very common in central parts of Europe, but extending southwards only to central France, Switzerland, northern Italy, Austria and Hungary; also north and central parts of European Russia and the Ukraine. In temperate Europe a common summer species, from June to mid September; in warm steppe biotopes as early as the second half of May.

Biology. A species of meadows, field margins and even drier steppe biotopes, and its epigamic behaviour is an adaptation to flat open areas, thus closely resembling that of Polyblepharis species. Adults are active during the whole day, sucking nectar at flowers, commonly at Scabious, and performing a special, very modified aerial courtship. The mating activity of E. livida has been described by many authors, such as Hamm (1908), Poulton (1913), Gruhl (1924) and Smith (1949), with a general concensus that females are active and fly in swarms, waiting for males with prey. However, Tuomikoski (1939) was right when he described males with prey as swarming for short periods of 10 to 12 seconds. The males are usually the active elements in the epigamic behaviour of E.

46

livida, making short flights with prey to above 1 metre over the vegetation. Usually several more males immediately take flight as in a chain reaction, and if no receptive female flies up to join a male then the males settle again after several seconds. Females very often make similar short circling flights above ground, which are evidently ‘offering’ flights when males are unavailable or are few in number and the female has to advertise her presence. This type of aerial mating in very modified swarms, or rather in ‘offering’ flights, is undoubtedly an adaptation for biotopes without trees or bushes and without distinct swarm markers. This is very similar to the habits of the steppeinhabiting Polyblepharis species. Copulating males always carry a large insect prey (usually large syrphids) that is often as large as the male himself. Females suck the prey during a long copulation on vegetation, which in my experience may last for more than 45 minutes.

Subgenus Anacrostichus Bezzi, 1909 Anacrostichus Bezzi, 1909:93 (as subgenus of Em-

pis). Type-species: Empis nitida Meigen, 1804 (orig. des.). Medium-sized

(4-5 mm) or large (up to 8 mm)

species with elongate 3rd antennal segment and short style. Femora often thickened and male genitalia shifted to a ventral position, foot-shaped. Eyes broadly separated on frons in male, or closely approximated to almost touching, upper ommatidia at most slightly enlarged, in broadly dichoptic males equally small; females dichoptic, frons generally very broad. Head almost globular, not very much higher than broad, occiput convex and either densely covered with very long hairs (E.

lucida-group) or with shorter bristles (E. nitidagroup). Face always very broad. Antennae with basal segment longer than deep (at most 4 times), covered with bristly-hairs like the globular 2nd segment; 3rd segment very long and rather evenly stout right up to tip, style short, at most one-third as long as 3rd segment, generally much shorter. Palpi very small, practically concealed in the mouth cavity, covered with minute hairs. Labrum at most twice as long as head is high, often shorter,

Fig. 54. Empis (Anacrostichus) nitida Meigen, male. Total length: 3.8-4.8 mm (A. Veltman del.).

47

Figs 55-60. E. (Anacrostichus) lucida Zetterstedt. — 55: male head in frontal view; 56: antenna. E. (Anacrostichus) nitida

Meigen. — 57: male head in frontal view; 58: antenna; 59: hind leg in anterior view. E. (Anacrostichus) bistortae Meigen. — 60: male head in frontal view. Scale: 0.5 mm, 0.2 mm for antennae.

rather slender and heavily sclerotised, extreme base almost bulbous. Thorax with well developed, large, and distinctly convex humeri and postalar calli. Mesonotum either densely long pubescent with multiserial long hair-like dorsocentral and acrostichal bristles that are hardly differentiated from the thoracic pubescence (E. lucida-group), or acrostichal and dorsocentral bristles reduced, former absent, latter at

most biserial, and large thoracic bristles well differentiated (E. nitida-group). Humeri always with long bristly-hairs anterior to humeral bristle. Prosternum bare between coxae, but sides with long bristly-hairs, episterna sometimes bare, and pronotum with bristly (not spinose) hairs becoming longer and fine at sides. Metapleural fan always dense, composed of numerous long bristles. Wings with radial fork more or less acute, veins R,,; and M, almost parallel to wing-tip, or at most slightly diverging, anal vein usually complete. However, there are frequent venation aberrations. Axillary angle acute, rarely somewhat obtuse, costal bristle usually small or if long then weak, rarely distinct. Squamae long-fringed. Legs rather long and slender, at most with hind

48

tibiae towards tip and hind basitarsi slightly swollen (E. lucida-group), or hind femora thickened and hind tibiae with a bent ‘knee’ (EF. nitidagroup). Femora ventrally and tibiae dorsally more or less bristled, basitarsi often spinose beneath. Hind tibia posteroventrally with an apical comb of small setulae, absent in E. verralli. Abdomen generally polished, clothed with more or less dense hairs, hind-marginal bristles not developed or at least not well differentiated from the other pubescence. Male postabdomen always modified: 5th segment with special excavations and projections on both tergum and sternum posteriorly, 6th and 7th segments reduced and downcurved, 8th segment of normal size again, apically broadened and funnel-like, supporting the genitalia which, together with the 8th segment, form a curious hoof-shaped hypopygium. Genitalia formed by small simple dorsal lamellae placed above elongated and upturned lateral lamellae; aedeagus rather short and stout, concealed within

lateral lamellae, base often visible, only partly shielded by usually well developed ventral lamella. Female abdomen pointed apically, at least apical

Figs 61-64. Wing venation of Empis subgenus Anacrostichus. — E. lucida — 61: wing-tip; 62: axillary excision. E. nitida — 63: wing-tip; 64: axillary excision. Scale: 0.5 mm.

two segments narrowed, usually three, 6th to 8th segments ovipositor-like, cerci very long, slender.

Phylogeny and classification. The species of Anacrostichus are in many respects a phylogenetically primitive group of Empis, but on the other hand share a series of characters in an apomorphous state (3rd antennal segment elongate, short style, modified pregenital segments, structure of male genitalia). They are closely related to Platyptera, which is undoubtedly the sister-group of Anacrostichus and which differs in several phylogenetically insignificant characters such as the pubescent face, modified broadened wings of the female, modified

hypandrium in male: these are all unsatisfactory as a basis for subgeneric separation. As stated by Collin (1961), Anacrostichus species can easily be divided into two well-defined species-groups. The first is the E. /ucida-group, which is undoubtedly older phylogenetically, with closely approximated eyes in the male and upper ommatidia slightly enlarged, non-differentiated hair-like pubescence on head and thorax, more acute radial fork on wing, and long slender unmodified legs. This group is represented in Europe by two species, E. lucida and E. verralli. The second

group is represented by the type-species E. nitida, and is undoubtedly a much more specialised group that includes species with dichoptic males, more specialised bristling on head and thorax, wing venation rather Empis s.str.-like with more open radial fork, and hind legs being more or less modified into special clasping organs in both sexes. This group is represented in Europe by three species, E. nitida, E. bistortae and E. monticola.

Distribution. Anacrostichus is essentially a boreal group, absent in the south, and represented by only twelve species in the Palaearctic region. Five species occur in Europe, and the other seven are east Palaearctic, known from Siberia, the Far East and

Japan. The European species are typical inhabitants of cool and temperate Europe: four species are ‘northern’ and are treated here, whilst the fifth,

E. monticola Loew, is a central European species that is found only in the high mountains.

Biology. Adults are nectar feeders, frequently in large numbers in flowering mountain meadows and especially on Polygonum. The lowland E. nitida is common

in orchards, visiting blossoming

49

fruit-trees, and is undoubtedly an important pollinator of apple-trees. Although the European species are often very abundant, frequently occurring in large numbers, little is known about their epigamic behaviour. Typical compact aggregations of swarming individuals, the synorchesia, were not observed and it is thought that most of the species mate in modified swarms, in rhythmic solitary flights (polyorchesia), as described below under the individual

species.

Key to Anacrostichus species 1

~Acrostichal and dorsocentral bristles numerous, long and hair-like, the former at least 2serial, the latter 4-serial. Hind femora slender and hind tibiae without a bent knee. Thoracic stripes polished. Eyes almost touching in MALES sas 55 doc cose 5 nu OP ee 2 — Acrostichal bristles absent, dorsocentrals 1to 2-serial. Hind femora more or less strongly swollen and hind tibiae with a bent knee. Thoracic stripes dull. Eyes distinctly separated on fronsimamales agers eee 3 2(1) Larger, 6-8 mm long. Hind femora with short black spines beneath. 3rd antennal segment very long, nearly 4 times as long as deep, style very short. Frons in female very shining black < SETENTS MEE SO EE 11. E. lucida — Smaller, 4-5 mm long. Hind femora not spinose beneath, clothed with fine dark hairs. 3rd antennal segment shorter, less than 3 times as long as deep, and style longer, onethird length of 3rd segment. Frons in female rather subshining, margins pollinose ....... is (Rin ee ee 12. E. verralli 3(1) Smaller, 4-6mm long. Prothoracic hairs, metapleural bristles and abdominal pubescence black. Hind femora very thickened, legs rather brownish, femora brown at tip. d:

—

50

eyes separated by a broad frons about as deep as the distance between posterior ocelli. @: 4th abdominal tergum simple . 13. E. nitida Larger, 6-8 mm long. Metapleural bristles pale, prothoracic bristly-hairs and abdominal pubescence yellowish. Hind femora only slightly thickened, legs more blackish, femora entirely black, d: Eyes narrowly separated, frons about as wide as anterior ocellus. 2: 4th abdominal tergum with a large projection on each side ema eee 14. E. bistortae

11. Empis (Anacrostichus) lucida Zetterstedt, 1838 Figs 55-56, 61-62, 65-66.

Empis lucida Zetterstedt, 1838:561.

Large, about 6-8 mm, shining black species with blackish legs, pluriserial, long, hair-like acrostichal and dorsocentral bristles separated by bare polished stripes, hind femora slender and spinose ventrally. Female frons brilliantly shining black. 3d. Eyes closely approximated, often touching for a short distance, ommatidia on upper half of eyes slightly enlarged, about twice as large. Occiput greyish-black, finely dusted, covered with numerous, rather long, fine, black bristly-hairs. Antennae black, 3rd segment very long, style short, stout, pointed apically. Palpi small, blackish, covered with sparse but very long black bristly-hairs. Labrum not very long, barely one-and-a-half times length of head. Thorax black, subshining, pleura more greyish. Mesonotum covered with dense, long bristly-hairs, large bristles as long and scarcely differentiated. Acrostichal and dorsocentral bristles long, hairlike, almost as long as basal two antennal segments combined, former irregularly 4-serial, latter multiserial anteriorly, almost 2-serial posteriorly, and ending in a row of stouter prescutellar bristles almost as long as other marginal bristles, several notopleurals (not differentiated from supra-alar), a postalar, and usually more than 10 scutellars. Sides of prosternum with very long black bristly-hairs, pronotum with coarser shorter bristles (but not spines), metapleural fan black, long and dense. Spiracles black. Wings faintly brownish-yellow, veins brown, radial fork rather acute; no costal bristle. Squamae dirty dark grey, narrow margins and fringes black. Halteres yellow, stalk often very dark. Legs shining black to very dark brownish-black, coxae grey dusted; all hairs and bristles black. All femora slender, especially hind femora long, hind tibiae slightly dilated towards tip and basitarsus slightly swollen. Fore femora with hairs beneath except for a few bristly-hairs before tip, mid femora with spinose bristles antero- and posteroventrally, hind femur with only an anteroventral row of bristles, posteroventrally somewhat reddish pubescent; femoral spines much shorter than femora

are deep. Tibiae with dorsal bristles arranged in two rows on all pairs, the bristles at least as long as tibiae are deep, usually longer, very distinctly so on mid tibia; short spine-like ventral bristles present on mid tibia and posterior four basitarsi.

Abdomen almost shining black, covered with long black hairs on anterior segments at sides, 2nd to 4th tergum with stronger hind-marginal bristles, corresponding sterna with long straggling hairs, Sth sternum with an apical comb of strong black spines between two finger-like lateral projections. Genitalia as in Fig. 65, lamellae finely grey pollinose, lateral lamellae with long black hairs cover-

ing almost the whole surface. Aedeagus concealed, stout and yellowish at base. Length: body 6.5-8 mm, wing 6.3—7.6 mm. 2. Very much like male though generally smaller, eyes broadly separated by very shining black frons, dusted grey above antennae only, and all hairs and bristles shorter and less numerous, usually only 6 scutellar bristles. Abdomen polished black, pubescence sparse and shorter, apical three segments narrowed. Length: body 5.6—7.8 mm, wing 6-7 mm. Not a very variable species; the legs may be wholly black or even very dark brownish. I have seen several specimens with the radial fork of the wings completely missing, though always on one side only. Lectotype designation. Described by Zetterstedt (1838) from eleven localities from northern

Sweden (Lapland, Jamtland) and Norway (Nordland, Finnmark). There are 9 d and5 @ from the syntypic series in the Zetterstedt Collection at Lund, all from Jukkasjarvi (T.Lpm.) except for 2 3 from Lycksele (Ly.Lpm.). A male in perfect condition labelled ‘E. lucida 3. Juckasj.’ from the Géteborg Collection was labelled in 1992 and is herewith designated as lectotype of Empis lucida Zetterstedt. Two males and a female from Jukkasjarvi are labelled as ‘var.b’, and one of these males has a symmetrically incomplete vein M, as in the subgenus Coptophlebia.

Distribution. One of the commonest species in Scandinavia; more than 1,200 specimens have been examined. Very common in northern and central parts, extending southwards to approximately 60° N, unknown from Denmark and with one exceptional capture in southern Sweden (Hall., Enslov, 29 May 1955, a female taken by H. Andersson). According to Lita Greve Jensen (by letter), E. lucida is evidently an alpine species in Norway, though found also in the lowlands and in coastal biotopes; it has even been observed high in the mountains, on 5 August 1965, on snow at 1900 m at

Vaga, Rondvassh¢ggda (On). With a long flight-pe-

as —

Figs 65-66. Male genitalia of E. (Anacrostichus) lucida Zetterstedt (S, Lapland, Vassijaure).—65: postabdomen; 66: aedeagus, hypandrium dotted. Scale: 0.5 mm.

i

riod, on dates from 18 May to 16 August, mostly in July, and only 15 specimens have been collected in the second half of May, in the southern warmer areas of Norway (HOi, HOy), Sweden (Hall., Dir.) and Finland (N).— A species of northern distribution in Europe, apart from Fennoscandia known also from Great Britain, Iceland and the Faroes, from NW Russia, through Siberia as far as the Far East, and at isolated localities in the Austrian and Swiss Alps. A species of boreo-alpine distribution in Europe. Biology. According to Tuomikoski (1939), swarming males fly rapidly in the air, perhaps with prey, but the flight is always very high and fast. Copulation takes place on the branches of coniferous trees. Tuomikoski recorded Rhamphomyia sulcata as the prey of E. lucida, and I have seen Diptera (Muscidae) and Plecoptera (Nemuridae) in collec-

tions as prey.

12. Empis (Anacrostichus) verralli Collin, 1927

Figs 67-68.

Empis (Anacrostichus) verralli Collin, 1927:21.

Resembling E. lucida, but a smaller (4-5 mm) and more brownish species with longer and denser pubescence everywhere, and hind femora finely pubescent beneath, not spinose. 3. Eyes very closely approximated on frons, all

ommatidia almost equally small. Antennae shorter than in other Anacrostichus species, basal segment not very much longer than the broader globular 2nd segment, 3rd segment broad at base, tapering apically and barely 3 times as long as deep, style nearly one-third length of 3rd segment. Labrum brownish, as long as in EF. lucida but much narrower at base. Thorax more brownish-black on pleura, thinly grey dusted. Acrostichal bristles irregularly 2-serial, dorsocentrals 4-serial, anteriorly more numerous, all very long and hair-like, the longest hairs as long as 3rd antennal segment with style. Humeri densely long-pubescent as in E. /ucida but marginal bristles even thinner and less distinct, scutellar bristles less numerous. Wings somewhat deeper brown clouded, often with a long hair replacing costal bristle, and axillary excision less acute. Squamae almost blackish and halteres yellowish-brown, stalk very dark at base. Legs similar in shape to those of E. lucida but more shining dark brown, all femora finely longpubescent beneath, without spines or bristles. Fore tibiae with several long posterodorsal bristles that are more than twice as long as tibia is deep, posterior four tibiae with similar though finer bristling than in E. lucida. Abdomen

almost shining dark brown, covered

with very dense but not very long black pubescence, only sides of basal two segments with longer hairs, and hind-marginal bristles distinctly longer

Figs 67-68. Male genitalia of E. (Anacrostichus) verralli Collin (GB, Scotland, Inverness: Loch Einich). — 67: postabdomen; 68. aedeagus with hypandrium. Scale: 0.2 mm

bs

although fine. Anterior five segments unmodified, 6th and 7th smaller and turned down, bearing the foot-shaped hypopygium. Lateral lamella elongate, subshining, with dense long black pubescence, the very slender tip of aedeagus often visible above dorsal lamellae; aedeagus simple, as in Fig. 68. Length: body 4-5 mm, wing about 4.5 mm. ?. Resembling male but all hairs and bristles much shorter and less numerous, bristling on legs finer and shorter. Frons shining black at middle, margins broadly pollinose. Abdomen blackishbrown, subshining, duller than in male, and all

hairs short. Length: body 3.7-4.5 mm, wing 3.8-4.2 mm. I have seen only a few specimens, but Collin (1961) referred to the very unstable wing venation: he recorded both veins M, and Cu abbreviated,

and even the rudiments or complete loss of the radial fork.

short, almost invisible, with 2 or 3 long black hairs

apically. Labrum brownish-black, about twice as long as head is high, its base very swollen. Thorax dark blackish-grey dusted, mesonotum rather finely greyish-brown pollinose, with a black triangular patch (with margins narrowly silverygrey) in prescutellar depression clearly visible in anterior view. All marginal bristles (a humeral with several not much shorter bristles anteriorly, a

posthumeral, usually 3 notopleurals, 2 supra-alars, a postalar with another bristle anteriorly, and 1 or 2 pairs of scutellars with several hairs each side) not very long and rather weak; no acrostichal bristles, dorsocentrals uniserial, small and hair-like, ending

in one bristle-like pair. Sides of prosternum with a tuft of rather long black bristles, pronotum with shorter and coarser bristles; metapleural fan black, dense but no conspicuously long. Spiracles black. Wings almost clear, with very dark and distinctly contrasting veins. Axillary excision unusually obtuse, sometimes

Distribution. Known so far from Great Britain and Ireland only, but it could well be a more widely distributed but overlooked species on the continent, including Denmark and southern Scandinavia. My records of its occurrence are from the end of May and the first half of June. The record of Meyer and Heydemann (1990) from Schleswig-Holstein needs confirmation.

13. Empis (Anacrostichus) nitida Meigen, 1804 Figs 54, 57-59, 63-64, 69-70. Empis nitida Meigen, 1804:223.

Small species, 4 to 5 mm long, females up to 6 mm, all hairs and bristles black, no acrostichal bristles, dorsocentrals

uniserial,

and

hind

femora

very

swollen, yellowish-brown at tip. 3. Eyes broadly separated on frons, all ommatidia equally small. Frons and face dark grey pollinose, frons with minute dark hairs on each side;

clypeus polished black. Occiput black, rather densely lighter grey dusted, clothed with irregular black spine-like bristles that are hardly longer than anterior ocellar bristles. Antennae black, Ist segment slender, more than twice as long as deep and like the globular 2nd segment covered with only very small bristly-hairs; 3rd segment very long, ~ evenly narrowed towards tip, and together with the very short style about twice as long as basal two segments together. Palpi blackish, extremely

almost

90°. Squamae

blackish-

grey with rather long black fringes, halteres clearly yellow, stalk much darker. Legs rather yellowish-brown on all tibiae and also fore femora, mid and hind femora shining black, yellowish at tip, tarsi darkened towards tip. Anterior four femora slender apically but on basal half slightly swollen, hind femora very stout, and hind tibiae with a bent ‘knee’. Fore femora almost bare beneath except for 2 or 3 small black spines at tip; posterior four femora with antero- and posteroventral rows of spinose bristles, between them with numerous minute black spines, interlocking with the small ventral bristly-hairs of the corresponding tibiae and forming very functional clasping organs. Anterior four tibiae with only short hairs dorsally, hind tibiae with longer dorsal bristly-hairs towards tip, and a posteroventral apical comb of setulae. Tarsal segments with only fine bristles in preapical circlets, basitarsi not spinose beneath. Abdomen very polished black, covered with fine short black pubescence, without hind-margi-

nal bristles, only sides of basal terga and posterior margin of 8th sternum with longer hairs. Genitalia small, lateral lamellae finely black pubescent, rather short, shining black, dorsal lamellae brownish, pointed, covered with minute dense hairs; ae-

deagus concealed within lamellae. Length: body 3.8-4.8 mm, wing 4.3-5.3 mm. 2. Very much like male but somewhat larger, with broader frons, and wings distinctly brownish clouded especially on basal half, apex mere hya-

line. Abdomen with shorter and finer black pubes-

53

Figs 69-70. Male genitalia of E. (Anacrostichus) nitida Meigen (CZ, Bohemia, Alber). — 69: postabdomen; 70: aedeagus. Scale: 0.2 mm.

cence, shining black, apical two segments very narrowed and cerci slender, very long. Length: body 4-6 mm, wing 4.5-5.8 mm. Abdomen rather sickle-shaped and downcurved in both sexes so that it is difficult to give the real body-length. As in the preceding species, the wing-venation is unstable and I have seen incomplete radial forks in several specimens, or a radial fork completely absent on one side. Distribution. Uncommon (25 6 and 30 2 examined) but widely distributed in Scandinavia. Commoner in the south, in Denmark, and at isolated localities in central and northern areas, the most

northern locality being in Norway (Fi, Kautokeino, 3 July 1979, 1 3, leg.L.Greve). On dates ranging from 8 May to 7 August, in southern areas in May and June, in central parts of Sweden (Jmt.) and Norway mainly in July, the only two August captures from Norway (TEi, Nsi). Not yet recorded from Finland but I have seen specimens from Russian Karelia. - The Netherlands, but not known from Great Britain, rather common in central Eu-

rope including the north of France and Italy. In temperate Europe in May and the first half of June, especially common on blossoming fruit-trees, in particular on apple-trees.

54

14. Empis (Anacrostichus) bistortae Meigen, 1822 Figs 60, 71-73.

Empis bistortae Meigen, 1822:29. Empis truncata Wiedemann in Meigen, 1822:31. Rather large, generally about 6-8 mm, almost uniformly black species, legs black, metapleural fan and abdominal pubescence pale. No acrostichal bristles, 4th tergum in female with large protuberances at sides. d. Frons about as wide above as anterior ocellus, widening out below, almost subshining black but somewhat wrinkled; face much broader and

grey dusted, blackish-grey

clypeus polished black. Occiput pollinose, densely black spinose

above neck, with longer bristles at about middle

becoming still longer and whitish below. Antennae black, basal segments with sparse, short black bristles, 3rd segment about 3 times as long as deep at base, style one-third of its length. Palpi not very small, blackish with brown tip, ventrally with several long pale hairs. Labrum about twice as long as head is high, brown, base very swollen, bulbous and polished black.

Thorax black, finely greyish pollinose, mesono-

tum with more brownish dusted stripes on the (hypothetical) lines of bristles, making the two dull darker stripes between the lines of bristles more conspicuous. Acrostichal bristles absent, dorsocentrals rather long, bristle-like, irregularly biserial anteriorly, uniserial posteriorly and ending in 2 or 3 pairs of strong prescutellar bristles, all black. Large bristles strong, a humeral with numerous shorter bristles anteriorly, a posthumeral, usually 6 notopleurals and 2 supra-alars arranged in a row, a postalar, and 4 to 6scutellars. Pronotum with a row of stout black bristles, but sides of prosternum and prothoracic episterna with long pale bristly-hairs. Metapleural fan composed of dense, long, pale bristly-hairs, usually a few dark brown or even black bristles intermixed. Spiracles blackish. Wings large, clear, with contrasting blackish veins, a distinct black costal bristle, and radial fork

not very acute, more as in FE. nitida. However, axillary excision very acute, unlike E. nitida. Squamae light brownish with pale fringes, halteres yellow, stalk dark at base. Legs shining black but coxae dull grey, at least hind tibiae at base, or often on the whole dorsal

surface, reddish to chestnut-brown. All hairs and bristles black, pubescence short and rather coarse, all femora (fore femora towards tip only) with a double row of short black ventral spines, and all tibiae bristled dorsally, the bristles barely longer than tibiae are deep. Mid and hind basitarsi spinose beneath. Hind femora slightly swollen (also mid femora more thickened than usual), though not at all as incrassate as in E. nitida and hind tibiae without a bent ‘knee’. Abdomen polished black, rather long pale pubescent except for dorsum, the modified (downcurved) 6th and 7th segments finely grey dusted, at least on margins, the funnel-like 8th segment more shining on sternal part and clothed with long pale hair-like bristles, 5th sternum with two finger-like processes posteriorly. Genitalia very much as in E. nitida, lateral lamellae with longer black bristles on posterior margin, and two similar upcurved bristly-hairs at tip of hypandrium (ventral lamella).

Length: body 5.8—7.5 mm, wing 6.6-8 mm. 2. Resembling male but frons broader and legs more brownish, hind tibiae yellowish-brown on at

~

Figs 71-73. Male genitalia of E. (Anacrostichus) bistortae Meigen (CZ, Bohemia, KrkonoSe Mts). — 71: postabdomen; 72: hypopygium in lateral view; 73: aedeagus, hypandrium dotted. Scale: 0.2 mm.

BS

least basal half, and mid tibiae at least at base, usually more. Wings also clear but veins lighter brownish. Legs as in male, abdomen densely but shorter pale pubescent. 4th tergum with very conspicuous, prominent, rounded protuberances at sides, the next two segments much smaller, and apical two (7th and 8th) segments very slender, ovipositorlike, cerci very long and slender. Length: body 6-8.2 mm, wing 6—7.8 mm. Distribution. Known from the Iberian Peninsula (Portugal, Spain) and distributed along the Atlantic coast to Belgium. Not yet known in England, Denmark or southern Scandinavia, but its occurrence there is probable. In temperate central parts of Europe, it is common in mountains from north-

ern France east to the Ukraine; very common in mountain meadows in June and July, on dates ranging from 9 June to 2 August.

choptic and all ommatidia equally small. Occiput and palpi covered with long, fine black hairs. Antennae long, 3rd segment elongate, style very short; basal segment about 3 times as long as deep, covered with long hairs. Proboscis about twice as long as head is high. Thorax covered with long black pubescence, large marginal bristles not very well differentiated, long and thin; female with shorter mesonotal pubescence, and marginal bristles consequently more distinct; usually 3 notopleurals, a supra-alar, a postalar, 6 scutellars, and 3 strong pairs of prescutellar dorsocentral bristles. Acrostichal and dorsocentral bristles pluriserial, between them with two dull darker stripes. Prosternum bare between coxae, but with long hairs on lateral lobes, episterna bare, or at most with a few minute hairs. Metapleu-

ral fan composed of long, thin, numerous bristlyhairs. Wings very conspicuous, in female very broadened, rounded at tip, but the shape not absolutely

Biology. The adults are nectar feeders, and are especially abundant on flowering Polygonum. The swarming was described by Gruhl (1955), who considered E. bistortae to be a typical species with the so-called polyorchesic swarming, a modified mating flight, a rhythmic alternation of dancing — perching — dancing, the males flying rhythmically against the wind above meadows. Gruhl also observed females flying about 2 to3 metres above the

constant. Radial fork not at all acute, and more ob-

ground,

femora ventrally and tibiae dorsally with rows of bristle-like hairs, not spinose or strongly bristled. Hind tibiae with a brush of short bristly-hairs ventrally at tip. Female legs with ordinary unmodified fine bristly-hairs or pennate. Abdomen finely pubescent, no hind-marginal bristles, sterna with long straggling bristly-hairs. 7th segment reduced and curved downwards in male, so that the somewhat foot-shaped hypopygium lies ventrally. Lamellae simple and moderately

with

the

lateral

abdominal

‘air sacs’

broadened in a manner be described in detail.

Subgenus Platyptera Meigen, 1803 Platyptera Meigen, 1803:269. Type-species: Empis borealis Linné, 1758 (subsequent des. by Meigen, 1804:219). Platypterygia Stephens, 1829:263 (new name for Platyptera Meigen, emend.). Type species: Empis borealis Linné, 1758 (automatic). Not very large species, about 6 to 7 mm long, with fine but long black pubescence, simple slender legs, male hypopygium foot-shaped and ventrally placed, and very broadened dark wings in female. Face below antennae broad, clothed with long black hairs on each side. Eyes very narrowly separated on frons in male, almost touching, upper ommatidia slightly enlarged. Females broadly di-

56

tuse in females, veins R,,,and M, rather distant but ending almost parallel to wing-tip in both sexes. Axillary lobe well developed, axillary excision acute, anal vein complete, and a distinct costal bris-

tle. Legs long and slender, simple, hind pair the longest, very elongate but both femora and tibiae not at all swollen. Pubescence longer in male, fine,

large, dorsal lamellae smaller than lateral ones,

ventral lamella with long projections; aedeagus concealed within lamellae, apically thin and rugose at sides before tip, base broadened and armed with lateral projections. The male hypopygium of Platyptera has been studied in detail by Ulrich (1972).

Phylogeny. Several characters, such as the structure of head (eyes in male very narrowly separated, not actually touching), the elongate 3rd antennal segment with very short style, absence of spinose bristles, the wing-venation (radial fork not very acute, veins R,,, and M, nearly parallel), and especially the ventrally shifted foot-like male hypopygium (very similar even in structural details)

Fig. 74. Empis (Platyptera) borealis L., male. Total length: 5.5—7.3 mm (A. Veltman del.).

suggest that Platyptera together with Anacrostichus forms a monophyletic group within Empis. The constitutive autapomorphous characters of Platyptera are the presence of hairs on the face, the

longs to Platyptera. Two species are known from Europe, both with a rather boreal distribution, oc-

cupying cool and temperate regions, and absent in the south.

unusually broadened wings in female, and the two

long slender projections ofthe inner wall of hypandrium in male. Distribution. A Holarctic group, as the Nearctic Empis barbata Loew, previously assigned to the genus Empimorpha Coquillett, undoubtedly be-

Biology. The adults are nectar feeders, frequently visiting flowers or flowering catkins in spring. Mating in aerial synorchesic swarms performed by females (an unusual habit in Empis); males capture an insect prey that is transferred to the female during copulation.

BS

76

/

Figs 75-77. E. (Platyptera) borealis L.—75: male wing-tip; 76: female wing-tip; 77: antenna. Scale: 0.5 mm for wing, 0.2 mm for antenna.

15. Empis (Platyptera) borealis

half of fore femora. Mid and hind femora with antero- and posteroventral rows of very short bristles, much longer bristly-hairs in two dorsal rows

Linné, 1758

Figs 74-81.

on hind tibiae, and on hind basitarsi ventrally.

Empis borealis Linné, 1758:603. Rather large species, about 6-7 mm long, finely long black pubescent all over, face covered with long hairs. Wings long, brown, very broadened and apically rounded in female. 3d. Head dark grey dusted and clothed with very long, fine black hairs on occiput; ocellar tubercle, facial margins, and even the linear frons with minute hairs. Antennae

black, basal two segments

long black pubescent, similar hairs on black palps. Thorax

black

in ground-colour,

mesonotum

dulled by greyish-brown dust, pleura greyer. All thoracic hairs and bristles long, black; acrostichal bristles irregularly 2- to 3-serial, dorsocentrals 4serial, extending anteriorly to humeri and at sides to notopleural depression. Mesonotum with 4 dull black stripes, the two middle ones between the lines of bristles longer and narrower. Wings brown, long, but not at all broadened. Squamae brownish, margins with long black fringes. Halteres bright yellow, base of stalk darkened. Legs long and slender, rather densely long black pubescent, especially anterior four tibiae dorsally. Legs mostly black, greyish dusted on coxae; posterior four femora except apical third or less, and basal third of hind tibiae, yellowish-brown to reddish-brown; hind tibiae often only translucent brownish ventrally at base, as is also basal third to

58

Abdomen somewhat laterally compressed, subshining black from some angles, distinctly brownish pollinose when viewed from above. Pubescence fine, uniformly black, especially long at sides of basal terga, and with long, very thin, straggling bristly-hairs on sternal part. Genitalia placed ventrally, foot-shaped, ventral lamella with two long slender rod-like projections, hanging beneath hypopygium. Length: body 5.5—7.3 mm, wing 7.6-8.3 mm long, 2.5-2.8 mm broad. 2. Differing chiefly by the broad dark grey frons and very large, broadened, and apically broadly rounded deep brown wings. All hairs on head and thorax shorter, legs shorter pubescent, with a tend-

ency (on anterior four tibiae in particular) to have rather short bristles. Legs distinctly paler than in male, practically all femora almost completely yellowish-brown, as well as tibiae except tips. Abdomen more subshining black, dusting finer and more greyish, the black abdominal pubescence not so dense, but hardly shorter. Length: body 6—7.8 mm, wing 8.2—9.6 mm long, 3.7-4.3 mm broad.

In addition to the variable colour of the legs, the wing-venation is also not at all constant. The radial fork may be incomplete on one side, and I have examined 2 females from Sweden (T.Lpm., Kilpis-

Figs 78-81. Male genitalia of E. (Platyptera) borealis L. (CZ, Bohemia, Veseli n.Luznici). —78: postabdomen; 79: aedeagal complex with hpandrium; 80: aedeagus in lateral view; 81: the same in dorsal view. Scale: 0.2 mm.

jarvi) with the upper branch of radial fork (vein R,) again forked apically before reaching costa. The second European species of Platyptera, Empis latiptera Chvala from Austria (female only), has the main diagnostic characters of E. borealis but differs chiefly by the pale metapleural fan, pale abdominal pubescence, white-fringed squamae, and sides of prosternum with pale hairs, but it may be recognised at first sight by the pennate legs in female. Distribution. Undoubtediy the commonest empid_ idin Scandinavia (nearly 1,500 specimens examined), though much more common in the colder central and northern parts. There are only a few records from Denmark and southern Sweden. It is a typical early spring species and the first Empis

species to emerge after winter. In the south mainly in May, in the north in June, though recorded over a much longer period on dates ranging from 28 March (Nrk., Markarret 1 d, and on1 April 8 d in

Og., Nykib. Stensj6fall) to 31 July (Finland, Ka, Vehkalahti); however, this last record is very problematic and may represent a very exceptional second generation as the other sparse records from July are all from Lapland. Collin (1961) recorded the latest date of capture in Great Britain as 19 June. — Very common in temperate central Europe on dates ranging from 2 April (lowlands) to 30 June (mountains). The southern limit of its distribution is in northern France, Switzerland, Austria,

Hungary and the Ukraine; known also from West Siberia.

39

o is &

Fig. 82. Empis (Leptempis) rustica Fallén, male. Total length: 7.6-10.5 mm (A. Veltman del.).

60

YS

86

Figs 83-86. Antennae of Empis subgenus Leptempis. — 83: E. rustica; 84: E. variegata; 85: E. grisea. Male head in frontal view. — 86: E. grisea. Scale: 0.2 mm.

Biology. The adults may be found in early spring, both in the lowlands and high in the mountains, sucking nectar from flowers and often in large numbers on willow catkins. Mating activity has been thoroughly studied and there are many reports in the literature, the most recent and most

complete being by Svensson and Petersson (1987, 1988, 1990). E. borealis is a rare exception within

the genus Empis, as the adults perform female synorchesia. Female swarms both large and small take place close to the overhanging branches of trees or bushes (in the mountains, closely associated with

dwarf pines) and males entering the swarm carry whatever insects available as prey: I have even caught males carrying spiders as the nuptial gift.

long wings and very distinctive male genitalia; the species with maculated wings very closely resemble rhagionids. Eyes broadly separated on frons with all ommatidia equally small (E. grisea-group), or touching on frons with upper ommatidia considerably enlarged (E. rustica-group). Females always broadly dichoptic. Antennae with 3rd segment conical, pointed apically and moderately long, style long and, unlike the preceding subgenera, slender and at least half as long as 3rd segment. Palpi rather long and slender, finely pubescent, not strongly bristled. Labrum at least as long as head is high, usually much longer. Thorax dull, if striped then the darker stripes are along the lines of bristles, often disappearing in anterior view. Thorax never densely long pubescent, large marginal bristles strong and well differentiat-

Subgenus Leptempis

ed, usually in full number, acrostichal and dorso-

Collin, 1926

central bristles generally. biserial, never multise-

Leptempis Collin, 1926:235 (as subgenus of Em-

longer bristles, sides of prosternum and prothoracic episterna with numerous long bristly-hairs;

rial. Pronotum

Pai

2h

fee

Fanon: (orig. des.).

Empis rustica Fallén, 1816 (orig. Engel, 1943:28 (as subgenus of Em-

Empis eh

Fabricius, 1781

Medium-sized (about 5 mm) to very large (much over 10 mm) species with long, mostly slender legs,

not spinose, covered with rather

metapleural bristles rather fine, numerous, but not

very long. Wings generally long, tapering apically and not at all broad, sometimes (E. maculata-complex) more or less brown maculated. At most with a small costal bristle. Radial fork not very acute but upper branch (vein R,) S-shaped, veins R,,;and M, almost parallel to wing-tip or only slightly diverging. All veins complete, not abbreviated, axillary excision acute or at least much less than 90°.

61

ee

Figs 87-90. Wing venation of Empis subgenus Leptempis. — E. rustica — 87: wing-tip; 88: axillary excision. E. grisea — 89: wing-tip; 90: axillary excision. Scale: 0.5 mm.

Legs long and slender, rarely (E. grisea-group) hind femora incrassate. Neither femora nor tibiae spinose, at most covered with rather fine bristles, or densely pubescent, hind tibiae postero-ventrally at tip with only a small, indistinct comb of fine setulae. Femora in females often fringed, covered with ventral flattened hairs, or more distinctly pennate. Abdomen rather long and slender, cylindrical, covered with hairs, no strong hind-marginal bristles. Male genitalia very distinctive, long, projecting distally along the longitudinal axis, formed by elongate slender lateral lamellae, often densely long bristled beneath and at tip; aedeagus placed ventrally and turned up at tip of hypopygium, concealed within lamellae or clearly visible, always long and slender, and often unusually undulating. Upper lamellae of complicated structure, bilobed and often bifurcate on each side, more or less con-

cealed within lateral lamellae. Female abdomen evenly narrowed apically, cerci long and slender.

Phylogeny and classification. The subgenus Lep-

62

tempis is undoubtedly a well differentiated group containing the most specialised large Empis species, sharing a series of apomorphous characters and having much in common with the phylogenetically progressive Empis s.str. species. Apart from their large size, which is not a significant differential character at all, Leptempis species only differ by the broadened, fleshy labial labella, the somewhat acute radial fork with vein R, S-shaped (also present in some phylogenetically primitive Empis s.str. species), the generally more ample wings, and especially the unique structure of the male genitalia (the structure of the dorsal lamellae in particular). However, similarly shaped male genitalia are also found in the E. ciliata-group of Euempis and the E. lepidopus-complex of Empis s.str., and synapomorphy cannot be ruled out. Leptempis has evolved along two phylogenetic lines: one retains holoptic males and unmodified slender legs, the EF. rustica-group, and the second contains dichoptic males (obviously a secondary condition) and swollen hind femora, the E. griseagroup. The latter is represented in the Palaearctic

region by three exclusively European species (E. grisea Fallén, E. flavitarsis von Roser and E. pteropoda Egger) that have often been incorrectly placed in the subgenus Pachymeria, for example by Engel (1943) and Frey (1954). In general terms, Leptempis appears to be a natural group of species within Empis because of the structure of the male genitalia and other rather consistent characters, even though the large E. rustica-group includes species that are very different in appearance, with yellow species superficially resembling Xanthempis species (E. dimidiata-complex) or the large, light grey dusted, rather southern European species with maculated wings (E. maculata-complex). Distribution. The species of Leptempis are widely distributed in the Palaearctic region: of the 28 described species (and several others are awaiting description), four are known from the Far East and Japan, and the remaining 24 are European in distribution. Apart from the four species treated here, which are found in cool and temperate Europe, 9 species are exclusively central European and another 11 are Mediterranean in distribution. Only one species, Empis grisea, has been found in Great Britain. This and E. rustica are known to me from Scandinavia, but another two (E. nigricans and E. variegata) have recently been found in the Netherlands and their occurrence in Denmark or southern Sweden seems very likely. Biology. Both males and females frequently visit flowers, imbibing sweet nectar and, presumably, also feeding on the protein-rich pollen grains. Swarming and mating have only been fully described in E. confusa Loew (Chvala 1980): males swarm in organised aerial aggregations and carry a small insect prey. This is clearly not a protein-rich meal for mating females, adequate for ovarian maturation, especially as the observed copulations were very short and lasted no longer than 30 to 90 seconds. No further data on epigamic behaviour in Leptempis are available.

Key to Leptempis species 1

—

larged, legs not pennate in female. Hind femOraislende nm iHeis oat ech Stee Nek 2 2(1) Wings uniformly faintly clouded......... 3 — Wings brown maculated. Coxae black, densely grey dusted, femora and tibiae brownishyellowsteien aches tae 18. E. variegata 3(2) Labrum long, more than twice as long as head is high. Coxae brownish, basal antennal segments rather yellowish. d: Dorsum of abdomen shining black; genitalia elongate, aedeagus clearly visible beneath ... 16. E. rustica — Labrum shorter, not very much longer than head is high, at most 1.5 times. Coxae black, dusted grey, antennae black on basal segments. 6: Dorsum of abdomen grey dusted, dull; genitalia small, aedeagus concealed within lamella yaar erase te 17. E. nigricans

16. Empis (Leptempis) rustica Fallén, 1816 Figs 82-83, 87-88, 91-92. Empis rufipes Fabricius, 1805:138 1763 and Gmelin, 1788).

(not Scopoli,

Empis rustica Fallén, 1816:18.

Large, 8-11 mm long, long-legged species with basal antennal segments brown and wings uniformly brownish clouded. Eyes meeting on frons in male and abdomen polished black on dorsum. Labrum long, about twice as long as head is high. 3d. Eyes contiguous on frons with upper ommatidia considerably enlarged, at least twice as large above as below. Face broad, upper mouth edge polished black, otherwise densely grey dusted like the occiput, the latter with two rows of black bristles becoming much longer and pale below. Antennae brown on basal two segments, or at least 2nd segment brown in ground-colour, 3rd segment black, not very long, pointed apically, and arista more than half as long as 3rd segment. Palpi rather long, slender, widened towards tip, yellowishbrown in colour and covered with sparse dark bristly-hairs. Proboscis long, labrum 1.9—2.2 times

as long as head is high. Mesonotum black in ground-colour, pleura densely lighter grey dusted, mesonotum more brownish tinged and with 3 darker stripes along

Small, about S—6 mm long. Eyes broadly sep-

the lines of bristles, contrasting black when viewed

arated on frons in male with all ommatidia

from behind but almost disappearing in anterior

equally small, posterior four femora shortpennate in female. Hind femora thickened . 2 Pole 5 SOE rot Cree Cee ere 19. E. grisea Large species, 8-11 mm long. Eyes meeting _on frons in male with upper ommatidia en-

view. Acrostichal and dorsocentral bristles black, hair-like, as long as basal antennal segment, former

2-serial, latter irregularly 3-serial at about middle and ending in 3 strong prescutellar pairs. Large bristles strong, black:

a humeral, with numerous

63

shorter bristles anteriorly, a posthumeral, usually 4 notopleurals, 2 smaller and one strong supra-alars, a postalar, and usually 4-8 scutellar bristles. Pronotum with about 6 rather long bristles on each side becoming pale and hair-like at sides, prothoracic episterna and sides of prosternum with long pale bristly-hairs. Metapleural fan almost orangeyellow, long and dense. Spiracles pale yellow. Wings uniformly light brown clouded, veins black towards tip. Radial fork rather acute, a small black costal bristle, and axillary excision acute. Squamae and halteres pale yellow, squamal fringes concolorous. Legs brownish-yellow in ground-colour including coxae, dorsum and tips of femora more or less darkened, coxae often darkened, and tips of tibiae and all tarsi black. All femora and tibiae slender. Fore femora almost bare, with longish dark hairs anteriorly, posterior four femora with a double row of outstanding long black bristles beneath longer than femora are deep, those on hind femora becoming shorter towards tip, and between them with numerous shorter black bristly-hairs; mid femora anteriorly, and hind femora both anteriorly and posteriorly, with long fine dark pubescence. All tibiae with irregular black dorsal bristles at least as long as tibiae are deep, but only mid tibia long pubescent beneath. Tarsal segments with preapical circlets of short black bristles, and hind basitarsus spinose ventrally. Abdomen shining black on terga, only basal tergum greyish dusted. Pubescence short on dorsum, sides with dense, long, pale bristly-hairs, sterna

grey dusted. Genitalia large, polished black; lateral lamellae long, slender, concave above and blunttipped, ventrally at base with long pale bristles becoming black towards tip; dorsal lamellae large, deeply excised, with lower lobe broad, upper one slender. Aedeagus polished brown, straight on basal half and often visible below lamellae, then curved up and very thin apically. Length: body 7.6-10.5 mm, wing 8.6-10 mm. 2. Eyes separated by a broad, dark grey dusted frons. Labrum still longer, 1.9-2.5 times as long as head is high. Wings almost clear. Legs clearly more extensively yellowish than in male and femora without the ventral bristles and lateral hairs, only tibiae with dorsal bristles, though much smaller. Abdomen thinly greyish dusted on terga, not at all shining black, and all pubescence short and sparse. Cerci very long, slender. Length: body 8-11 mm, wing 8-9.6 mm. Very much like the next species, E. nigricans, which is compared with it in the description that follows. Six European species of the E. rusticagroup with non-maculated wings were revised by Chvala (1977). Lundbeck (1910) misidentified this species in Denmark as E. nigricans. Type revision. Described by Fallén (1816) as ‘In Carduis Scaniae mense Augusto rarius visa’, from both sexes. In 1985 I examined 2 specimens under this name in the Fallén Collection in Stockholm: a male, which has lost its head but is otherwise well preserved and is the species described above as E. rustica, and a conspecific female. A lectotype has

Figs 91-92. Male genitalia of E. (Leptempis) rustica Fallén (SLK, Mala Fatra Mts, Bystii¢ka). — 91: postabdomen; 92: aedeagus. Scale: 0.5 mm.

64

not been designated, but required the male should be selected. Distribution. Apart from the two type specimens from Swedish Skane I have seen only 11 d and6 @, from Denmark, collected on dates ranging from 27 June to 7 August. — Not yet found in Great Britain or the Netherlands, but it has the same continental distribution as E. nigricans, occurring southwards to Austria, Hungary, Slovenia and Romania. A typical summer species; in temperate Europe with a long flight-period, as in Denmark, from the second half of June to the middle of August, but mainly in July, preferring higher altitudes.

17. Empis (Leptempis) nigricans Meigen, 1804 Figs 93-94. Empis nigricans Meigen, 1804:234. Resembling E. rustica but labrum shorter, not very much longer than head is high, antennae black even on basal segments, legs with fewer and shorter bristles on femora beneath, coxae grey, and male genitalia smaller with aedeagus hardly visible. 3. Eyes meeting on frons but upper ommatidia much smaller than in E. rustica, only slightly enlarged. Antennae black, only very exceptionally basal segment translucent brownish at tip. The deciding distinguishing feature on the head is the much shorter proboscis, labrum only 1.1-1.4 times as long as head is high. Thorax with the same pubescence and bristling as in E. rustica but mesonotum obviously more greyish, without the brownish tinge, and mesonotal stripes more distinct, contrasting velvetybrown even when viewed from in front. Metapleural fan apparently paler, more whitish-yellow. Legs light brownish to yellowish-brown on femora and tibiae, only knees and tips of tibiae blackish, but coxae black in ground-colour, densely grey concolorous

with

thoracic

shape to E. rustica but much smaller, lateral lamel-

lae with long black bristles along ventral margin only, and aedeagus completely concealed within lamellae, only a short, thread-like upcurved tip visible between dorsal lamellae. Length: body 7.5-9.7 mm, wing 8—9.2 mm. ?. Labrum slightly longer, about 1.3-1.7 times as long as head is high. Pubescence and bristling shorter everywhere, posterior femora almost without bristles, but corresponding tibiae also with short irregular bristles ventrally. Abdomen more shining black, the terga in particular less dusted, and all hairs very short even on venter. Length: body 7.3-9.6 mm, wing 7.2-9.1 mm. The most closely allied species is E. rustica, with which it is compared in the description above. The central European E. cognata Egger is very similar, having an equally short labrum, but the coxae are brownish as in E. rustica, and the abdomen is shining black above and pale yellow on venter.

Distribution. A species of the temperate and cooler parts of Europe, with the southern limit of its distribution

in Austria,

northern

Italy, Slovenia

and Romania, extending north through Germany to the Netherlands. Its occurrence in Denmark or southern Sweden is very likely. A typical spring lowland species, flying for a short period from the end of April to the end of May, only rarely in June. Common on flowers, very often on blossoming fruit-trees, and very possibly an important pollinator when present in large numbers. A common species in suitable warm lowland biotopes.

18. Empis (Leptempis) variegata Meigen, 1804 Figs 84, 95-96.

Wings less clouded, almost clear.

dusted,

dull grey, not at all shining, dorsum with short black, rather coarser pubescence, sides of terga and whole of sterna covered with pale bristlyhairs, much longer on sterna. Genitalia similar in

pleura; tarsi

black. Fore and mid femora covered only with short hairs except for a double row of rather short black bristles on mid femur, the bristles in posterior row distinctly longer, but at most as long as femur is deep. Hind femora with similar long hairs anteriorly and posteriorly to those in E. rustica but the ventral black bristles in a double row very short _and no additional bristling between them. Armature of tibiae and tarsi as in E. rustica. Abdomen distinctly greyish dusted on terga,

Empis variegata Meigen, 1804:225.

Large, generally about 10 mm, with brown maculated wings, blackish grey coxae and short labrum. Abdomen shining black on dorsum in male, genitalia large with a clearly visible slender aedeagus. 3. Eyes meeting on frons, with upper ommatidia slightly enlarged, face very broad, polished black on lower half and on clypeus. Occiput dull grey, with short but strong black bristles above, below neck with long, fine pale hairs. Antennae black, basal segments sometimes translucent brownish, 3rd segment rather long and slender,

65

Figs 93-94. Male genitalia of E. (Leptempis) nigricans Meigen (CZ, Bohemia, Hostomice p.Brdy).—93: postabdomen; 94: aedeagus. Scale: 0.2 mm.

style not very much shorter than 3rd segment. Palpi reddish-brown, ventrally with long dark hairs. Labrum rather strong and short, at most 1.5 times as long as head is high. Thorax rather darker grey and densely dusted, mesonotum with 3 darker stripes along lines of bristles, the central one very narrow, lateral stripes very broad; all disappearing or very light brownish in anterior view, almost velvety-black and coalescent posteriorly when viewed from behind. Acrostichal bristles very small (about as long as 2nd antennal segment), very closely biserial or even in one alternating row, dorsocentral bristles more bristle-like, at least twice as long and distinctly biserial. Large marginal bristles rather short but very strong, in full number including a group of usually about 8 notopleural bristles merging with up to 4 supra-alars. Pronotum with 5 strong black bristles

on each side, laterally with long pale bristly-hairs, similar to those on lateral lobes of prosternum and on episterna. Metapleural fan long, pale, spiracles pale yellow. Wings large, almost clear, with distinct brown maculations on crossveins and bifurcations, and darkenings along veins, costal stigma blackish. Costal bristle very small, but base of costa with

dense short bristles. Radial fork rather obtuse, but vein R, distinctly curved on apical half, axillary excision acute. Squamae (with concolorous fringes) and halteres pale yellow. Legs similar in colour to E. nigricans, trochanters, knees, tips of tibiae and all tarsi contrasting black, otherwise legs pale yellowish-brown, coxae

66

black in ground-colour. Bristling as in E. rustica but ventral bristles on hind femora shorter, posteroventral bristles on mid femora very long and strong, similar strong bristles also in a double row on mid tibia ventrally. Hind tibiae with longer adpressed pubescence beneath. Abdomen shining black on dorsum, whole of Ist tergum and narrow posterior margins on following terga greyish pollinose, sterna dull grey. Dorsum with short black hairs, sides and venter clothed with long whitish-yellow hairs. Genitalia of very complicated structure, lateral lamellae slender, upper margin straight, tip broadened and curved backwards (in dried specimens), lower margin and at least apical third of lamella with dense strong black bristles. Dorsal lamellae bifurcate on apical half, sides deeply excised, covered with rather finer and shorter black bristles; aedeagus very long and thin, free above lamellae and curved backwards. Length: body 9-11 mm, wing 9.6-10.6 mm. ?. Resembling male but eyes separated by a broad dull grey frons, and all ommatidia equally small. Marginal bristles on mesonotum longer and thinner, but all bristles on legs much shorter, and long bristly-hairs at sides of posterior femora absent. Abdomen finely grey pollinose on dorsum, not shining, and pubescence short even at sides and ventrally. Length: body 8-11 mm, wing 8.8-10.3 mm. In many respects (shorter labrum, blackish-grey coxae, black antennae, slightly enlarged upper ommatidia in male), this species more closely resem-

bles E. nigricans, but the wings are distinctly

Figs 95-96. Male genitalia of E. (Leptempis) variegata Meigen (D, Schwarzwald, Wehrtal). — 95: postabdomen; 96: aedeagus. Scale: 0.5 mm.

brown maculated, thoracic bristling shorter, male abdomen shining black on dorsum, and genitalia very distinctive with the aedeagus very much like that of E. cognata Egger.

Distribution. In NW Europe known from Belgium and the Netherlands, extending south to Austria, Slovakia and the central parts of European Russia. Its occurrence in the extreme south of Scandinavia and in Denmark is to be expected. In the second half of May and in June in central temperate Europe, mostly in lowlands.

3. Eyes broadly separated by a dark grey dusted frons that is about as wide above as ocellar triangle, broadened below, above antennae as wide as the similarly dark grey dusted face, clypeus polished black. Occiput dull blackish-grey, with dense long black bristles, below neck with longer whitish bristly-hairs. Antennae brownish-yellow on basal segments, lst segment not very much longer than deep and covered with a few short black bristlyhairs; 3rd segment black, very narrowed apically and rather short, style about half as long as 3rd segment. Palpi very small, slender, yellowish to almost whitish, with several fine hairs beneath. Labrum

19. Empis (Leptempis) grisea

brownish, slender, barely twice as long as head is

Fallén, 1816

high.

Figs 85-86, 89-90, 97-98.

Empis grisea Fallén, 1816:22. Empis pilipes Meigen, 1822:31. Empis nodosa Beling, 1882:208. Empis rufiventris Strobl, 1893:85 (as var. of E. grisea). Empis nigriventris Strobl, 1893:85 (as var. of E. gri-

sea).

oo

Thorax almost uniformly dulled by greyish dust, darker mesonotal stripes along the lines of bristles very inconspicuous, completely disappearing in anterior view. Acrostichal bristles 2-serial, diverging posteriorly, dorsocentrals irregularly 2- to 3-serial, ending in 3 strong prescutellar pairs, all black and bristle-like, about as long as antennal style. Humeri with dense black bristles, posteriorly with 2 strong bristles about as long as the numerous marginal bristles represented by merging post-

Rather small, about 5-6 mm, with large swollen

humeral, notopleural, supra-alar and postalar bris-

hind femora densely bristled beneath in male, short pennate posteroventrally in female. Eyes broadly separated on frons in male, and aedeagus long, undulating.

tles, with additional small bristly-hairs, and usually 4 scutellar bristles. Pronotum with a row of black bristles, but episterna and sides of prosternum with longer pale bristly-hairs, rarely with addition-

67

al black bristles. Metapleural fan dense, varying from completely pale, through intermixed pale and black bristles, to completely black. Spiracles yellowish. Wings large and almost clear, veins dark, apex rather pointed. A small black costal bristle, axillary excision varying in shape but never very acute. Squamae pale with concolorous fringes, halteres yellowish. Legs often uniformly yellowish including coxae and tarsi, or with darker brown markings on coxae, or even femora and tarsi. Anterior four femora rather small and slender, but hind femora much longer and distinctly thickened, somewhat spindle-shaped. Fore femora with hairs only, some longer hair-like bristles posteriorly, mid and hind femora with a double row of black ventral bristles, those on hind femora longer, between the spinose bristles with numerous shorter hairs and bristles. Tibiae not bristled, covered with short hairs, ven-

trally with very dense, soft outstanding fine pubescence, more distinct on posterior four tibiae. Tarsi with only short hairs. Abdomen either completely brownish-yellow or entirely black, subshining, covered with longer hairs on basal segments at sides, mostly pale pubescent in light coloured specimens, or with black pubescence in dark forms. Genitalia with the characteristic structure of Leptempis, yellow or extensively darkened; lateral lamellae very long and slender, ventrally and at tip with dense black bristles, dorsal lamellae deeply excised. Aedeagus long and slender, undulating, hypandrium (ventral lamella) projecting as a tongue-like, microscopically pilose process.

Length: body 4.5-6.3 mm, wing 4.8-6.6 mm. 2. Frons broader, as wide above as face, and all hairs and bristles slightly shorter and less numerous. Posterior four femora anterodorsally with numerous irregular flattened hairs, posteroventrally distinctly pennate towards tip. Abdomen less shining, finely grey pollinose, pubescence shorter and variable in colour, depending on the colour of the abdomen. Length: body 4.2-6.5 mm, wing 5.2—6.6 mm. A very variable species in the colour of the abdomen. The forms with a yellowish abdomen are generally named as var. nodosa or var. rufiventris, and those with a black abdomen as var. nigriventris. The abdominal pubescence also varies from very pale to entirely black, and the legs may be completely yellowish, with distinct dark markings, to very extensively darkened. Metapleural bristles vary from entirely black to very pale, with all intermediate transitions. Females are often confused with females of E. (Euempis) picipes; however, E. picipes has the acrostichal and dorsocentral bristles much shorter, the latter very numerous anteriorly, and the posterior four femora not so stout and without pennation or flattened hairs. Type revision. Described by Fallén (1816) from ‘Ostrogothia’, Sweden. There are 2 specimens in the Fallén Collection in Stockholm under EF. grisea,

as was stated by Collin (1961), which I studied in 1985: a male, which is the species described above (which should be selected as lectotype if so required), anda female of E. (Euempis) picipes Meigen. For full synonymy, see Collin (1961:496).

Figs 97-98. Male genitalia of E. (Leptempis) grisea Fallén (CH, Palagnedra, Lago). — 97: postabdomen; 98: aedeagus. Scale: 0.5 mm.

68

Distribution. A species of southern distribution in Scandinavia, so far known only from a small area in the extreme south though it is fairly common there; more than 150 specimens have been examined, 40 ¢ and 57 2 from Denmark, and 30 6 and 23 2 from Swedish Skane. Two further males were collected by R. Danielsson at Vg., G6teborg, Moludal. Bidenkap (1892) recorded this species from Norway (probably VE), but study of his material showed that it was E. (Euempis) picipes (for details, see under that species). On dates from 10

June to 15 August, but mainly end of June and first half of July. There is one very early record from May in the Copenhagen collection (SJ, Sgnderborg 14.5.1889, 2 2, Wiistnei) which is rather problematic or very exceptional. — Great Britain including Ireland, the Netherlands, and south into central Europe, to Switzerland, northern Italy and Romania. In warm lowland biotopes from the beginning of May to the first third of August, but mainly in June.

Subgenus Empis s.str. Empis Linné, 1758:603 Type-species: Empis pennipes Linné, 1758 (des. by Latreille, 1810). Pterempis Bezzi, 1909:87 (as subgenus of Empis). Type-species: Empis genualis Strobl, 1893 (des. by Collin, 1961 by subsequent monotypy). Generally small, about 2-4 mm;

the largest Pal-

aearctic species measure about 5-6 mm, but sucha

size is exceptional. Mostly black species, shining or dulled by greyish dust, rarely dark brown in colour, never yellow. Legs uniformly black to brownish, often pennate in females. Head small, rounded, males holoptic with upper ommatidia usually very enlarged, much larger above than below, females broadly dichoptic with all ommatidia equally small. Antennae with small basal segments, Ist segment not much longer than deep, slender; 2nd segment usually broad, globular, both covered with fine minute bristles. 3rd segment moderately long, narrowed apically, style slender and at least one-quarter length of 3rd segment, to almost as long. Palpi small, covered with fine bristles. Proboscis long and slender, all parts heavily sclerotised; labrum always much longer than head is high, even more than three times, labi-

um always slender with terminal labella scarcely distinguishable, very sclerotised and only micro-

scopically pubescent. Thorax never distinctly striped as is usual in

other subgenera, and the stripes, if present, alter their position or disappear depending on the point-of-view and the angle of light. Large bristles usually in full number, acrostichal and dorsocentral bristles always present and varying from unito multiserial. The number, length and thickness of the thoracic hairs and bristles are species-specific and are always of great diagnostic importance. Prosternum bare on middle between coxae, except for a small group of species close to Empis pilosa Loew, sides of prosternum (epimera), as well as prothoracic episterna, covered with minute fine bristly-hairs, pronotum usually with stouter bristles. Metapleural fan always long and rather dense. Wings very characteristic because of a widely open radial fork and a rather short and wide discal cell that is very truncate at tip. The upper vein of the radial fork (vein R,) always straight, perpendicular, almost at right-angles to vein R,, or slightly oblique. All veins from the discal cell reaching wing-margin in the ground-plan condition, anal vein complete or abbreviated. Aberrant spécimens with an incomplete or entirely absent radial fork, or with an abbreviated vein M, as in Copto-

phlebia, appear infrequently in almost all species. Such venation aberrations are usually on one side, and only rarely on both. Axillary lobe always large, but axillary excision not always acute. Legs either long and slender with tips of tarsi often characteristically curved (E. pennipes-group), or legs short and rather stout, femora never thickened, but hind tibiae towards tip and hind and/or fore basitarsi sometimes distinctly swollen (E. chioptera-group). Bristling of legs very distinctive for each species, as is the frequent pennation of legs in females. Abdomen more or less pubescent, the hairs may be longer on posterior margins, but abdomen never distinctly bristled and hind-marginal bristles not developed. Male genitalia varying in structure but never very large, all lamellae rather small, dorsal lamella, even if complex, never enlarged. Aedeagus long and thin, evenly bowed or undulating and clearly visible (the so-called ‘open’ genitalia of the E. pennipes-group), or short and usually stouter, more or less concealed within lateral lamellae (the so-called ‘closed’ genitalia for instance of the E. chioptera-group). Female abdomen pointed with moderately long slender cerci. Phylogeny and classification. When compared with the undoubtedly primitive forms of Empis in the subgenera Euempis and Pachymeria, Empis s.str. possesses a series of apomorphous characters and is in many respects much more specialised.

69

Fig. 99. Empis (s.str.) pennipes L., male. Total length: 3.6-5.2 mm (A. Veltman del.).

This applies not only to the morphological characters but also to the biology, feeding habits and epigamic behaviour. The apomorphous characters of Empis s.str. may be summarised as follows: (1) Differentiation of sensory bristles on thorax and legs; primitive forms with more uniform hairing are represented by the E. pilosa-complex of species. (2) Structure of antennae, especially the shortening of the basal segment and lengthening of antennal style. (3) The long, heavily sclerotised proboscis, especially the labium with its slender, bare, sclerotised labella. (4) Bare prosternum, with only the sides (epimera)

70

covered with fine hairs (except for the E. pilosacomplex). (5) Specialised wing venation, especially the strengthening of veins along costal margin (Sc, R) and consequent ‘opening’ of radial fork. (6) The almost right-angled radial fork with straight vein R, which reflects the obvious shortening and broadening of the wings, including the shortening of the apically truncate discal cell and the enlargement of axillary lobe. (7) Structure of legs, development of sensory bristles and frequent thickening of basal tarsal segments covered with sensory setae. (8) Structural simplification of the male genitalia (originally with a long thin aedeagus), an ad-

Fig. 100. Empis (s.str.) pennipes L., female. Total length: 3.8-5.2 mm (A. Veltman del.).

aptation for obligatory mating in the very specialised aerial aggregations of the synorchesic type. The very characteristic pennation of legs in females is an apomorphous state but it cannot be regarded as an autapomorphy of this group, as flattened pennate bristling isa. common feature in females of phylogenetically older groups of Empis. Even in Empis s.str., the development of this bristling in an uninterrupted series from fringes and flattened hairs up to very broad pennate bristles has no phylogenetic value as it is only a specifically

useful adaptation — a very functional species-specific visual signal from a flying female ready for copulation. As was recently been shown (Chvala 1990), it is the most closely related species that differ significantly in the development of pennate bristles in females. Although the subgenus Empis s.str. as currently conceived includes rather uniform small Empis species that are well characterised by the series of apomorphous characters ennumerated above, it is not a natural monophyletic group at all. Its status

aL

as a ‘subgenus’ merely underlines the very unnatural classification of empidine genera, as Empis subgenus Coptophlebia and Rhamphomyia subgenus Aclonempis only differ from it in slight venational differences, the abbreviated vein M, and the

unforked vein R,,, respectively. Moreover, venation aberrations are not'uncommon in species of Empis s.str., Coptophlebia and Aclonempis, and the weak taxonomic differentiation may be entirely obliterated when the aberration appears on both wings. Such specimens cause real problems in identification for non-specialists, as the useful and practical differential characters of the ‘subgenus’ key are lost. Several attempts have been made to group the large number of Empis s.str. species into smaller units, the species-groups. Most of these attempts used artificial but practical characters such as the colour of the halteres and abdominal pubescence, usually in combination with the colour of the metapleural fan. Permutations of these two main characters were used by Engel (1946) in Lindner’s ‘Die

Fliegen’ to arrange all the Palaearctic species into four species-groups which had considerable practiced value. Such a practical classification is used here too in the key to species. The first serious attempt at a classification into natural groups, based mainly on the structure of the male genitalia, was published by Syrovatka (1980). At that time he recognised four groups, the E. pennipes-, E. pilosa-, E. chioptera- and E. alpicola-groups, based entirely on structural and/or proportional differences (sometimes a little significance) in the genital lamellae. Later (Syrovatka 1991), he himself abandoned this classification and proposed three groups of species (named as complexes), the E. pennipes-, E. nigripes- and E. chioptera-complex, which in my opinion are more natural. This classification is in general agreement with my own ideas. However, his concept of the E. nigripes-complex, based on the presence of distinct projections and tubercles on the 8th abdominal tergum in male, seems unnatural as it covers phylogenetically unrelated species, some of which

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105

Figs 101-107. Wing venation of Empis s.str.—101: E. lepidopus; 102: E. albopilosa, radial fork; 103: E. pennipes, radial fork; 104: E. serotina, radial fork; 105: the same, discal cell; 106: E. chioptera, radial fork; 107: the same, discal cell. Scale: 0.5 mm.

Te

C

Figs 108-112. Male mid leg in anterior view of Empis s.str. — 108: E. lepidopus; 109: E. albopilosa; 110: E. acinerea; 111: E. syrovatkai; 112: E. genualis. Scale: 0.5 mm.

should be provisionally placed in the E. pennipesgroup. There are also some objections to his concept of E. chioptera-complex. A truly natural classification into subordinate groups of species or species-complexes must include the species currently included in the ‘subgenera’ Coptophlebia and Aclonempis. ‘The Empis s.str. species described by Loew and Becker have been fully revised by Syrovatka (1991), based on a study. of the types.

Distribution. The species currently included in the subgenus Empis s.str. are worldwide in distribution and are found throughout the Palaearctic region. 106 species of Empis s.str. are recognised as

valid in the Palaearctic Catalogue (Chvala and Wagner 1989) but this number is probably scarcely half the number of species actually in existence. I have seen, for instance, more than twenty new species of Empis s.str. awaiting description from Middle Asia, collected during two short expeditions to the mountains of Tadzhikistan, Uzbekistan, Kirgizia and Kazakhstan, a huge area of Asia from

which not a single species has previously been recorded. The same is undoubtedly true of Japan and the Far East, as only three species of Empis s.str. have been recorded as eastern Palaearctic. Species of Empis s.str. are distributed mainly in the southern parts of Europe and in the Mediterranean, including North Africa and the Near East;

73

Figs 113-116. Male mid leg in anterior view of Empis s.str. — 113: E. pennipes; 114: E. rufiventris; 115: E. planetica; 116: E. staegeri. Scale: 0.2 mm.

two species are known from the Canary Islands. However, there are still many undescribed species known to me even from southern Europe. Seventeen species have been found in Scandinavia (if Empis nitidula Zetterstedt is regarded as valid species), and another eleven with a northern distribu-

tion, known along the coasts of the North Sea, are also included here. The Finnish species of Empis s.str. have recently been dealt with (Chvala 1991), with complete listings of distribution data and with distribution maps.

Biology. Most of the species of Empis s.str. are typical spring species, occurring mainly in May and

74

June, and E. chioptera is actually the earliest spring species of the genus, together with E. (Platyptera) borealis. Only a few species are of summer or even of late summer occurrence. The adults are frequently found in large numbers on flowers where they suck the sweet nectar, as the structure of proboscis clearly enables them to imbibe fluids of both plant and animal origins. The adults are not predators, except for a short period during mating activity, when males transfer the prey without consuming it to the female during copulation. So far as I know, the prey is always edible and the female sucks it during copulation, though obligatory mating without prey transfer has also been recognised in several species.

The species of Empis s.str. exhibit a wide range of very characteristic species-specific aerial mating aggregations, the typical synorchesia that is also known in other Empis subgenera such as Pachymeria, Leptempis or Platyptera. There is an extensive literature on the epigamic behaviour of Palaearctic Empis s.str. (Tuomikoski 1939, Laurence 1950, Parmenter

1951, Smith 1952, Chvala

1980,

and others), and many observations are also available from the Nearctic region. The very different epigamic behaviour of two closely related species, E. nigripes and E. nuntia, has been discussed by Chvala (1991) as an example of an effective ethological isolating mechanism in insects.

Key to Empis s.str. species

Males 1

pubescence black and very long proboscis . . —

Halteres black; if somewhat dark brown (dasyprocta), then small species, 2.5-3.5 mm long, with abdominal pubescence pale ..... Pg TPN. eis meet Med Petales BR Ra a ee HL 23 2(1) Labrum very long, about 3 times as long as head is high. Metapleural bristles and abdominal pubescence black. Halteres rather yellowish-brown or very darkened. Wings brownish, aedeagus very long and slender, evenly bowed. Large species, about 4-5 mm long .. Se ai ten oa ies ee Oe 27. E. pennipes — _Labrum shorter, at most slightly more than twice as long as head is high (care must be taken with a long labium if it is porrect). Halteres extensively pale. If there is doubt as to the length of labrum (rufiventris), then metapleural bristles and abdominal pubescence pale .

Halteres pale; if darkened (pennipes), then large species, 4-5 mm long, with abdominal

; Figs 117-119. Male legs in anterior view of Empis s.str. — 117: E. melanotricha, mid leg; 118: E. tanysphyra, mid leg; 119: E. woodi, hind leg. Scale: 0.2 mm.

ah)

Figs 120-125. Legs of Empis s.str. in anterior view. — 120: E. chioptera, male mid leg; 121: E. prodromus, male mid leg; 122: the same, male hind leg; 123: the same, female hind leg; 124: E. dasyprocta, male mid leg; 125: the same, female hind leg. Scale: 0.2 mm.

of abdomen, also hairs at sides of prosternum

and on prothoracic episterna pale (except rufiventris, acinerea and gooti) ............ 4 Abdominal pubescence black or blackishbrown, including base of abdomen and prothorax. Metapleural bristles black ...... 21 4(3) Front half of mesonotum densely clothed with long, fine, pale hairs; acrostichal and dorsocentral bristles multiserial, very narrowly separated. Aedeagus very long and thin, evenly bowed. Large species, 5-6 mm long er TE en a ee i OTe 23. E. decora Front half of mesonotum with the usual dark

76

bristly-hairs; if whitish (albopilosa), then anterior quarter of mesonotum bare, and acrostichal and dorsocentral bristles biserial .. 5

5(4) Dorsum of abdomen dulled by greyish dust, not shining black; if in doubt (genualis, syro-

vatkai), the species have been included in both sections sr. eee ene 6 Dorsum of abdomen shining black or at least subshining, basal segments sometimes finely pollinose mae Wet ee. eee eee 15 6(5) 8th tergum with prominent lateral tubercles, 7th tergum with corresponding excisions on hind margin at sides. Aedeagus short, without

loops or bends, 4 scutellar bristles. Mediumsized species, 3-4mm long (nigripes-comDIS oe eer Ae Ae oats... 7 8th tergum simple, without tubercles, and excisions on posterior sides of 7th tergum indistinct; if a small tubercle is present (planetica),

then aedeagus with a loop and 2 scutellar bris(HSS "yD oe ghode ok ead tac Pee 10 7(6) Aedeagus very stout, apically broadened (Fig. Se OAT EU RA SRS ea ete a 8 Aedeagus rather slender and apically disftmetLyenarrowed(Fig.195)) Saeedackent. 9 8(7) Lateral tubercles on 8th tergum rounded, side excisions of 7th tergum rather shallow. Dorsum of abdomen mainly with black hairs. Mid

tibiae with 2-3 long dorsal bristles on basal half, posterior four femora with long anteroGorsallbnstlespee eae ee 35. E. nigripes Lateral tubercles on 8th tergum flattened and spoon-shaped, pointing forwards into very deep excisions of 7th tergum. Dorsum of abdomen with pale hairs. Mid tibiae with 4—5 long dorsal bristles, posterior four femora with short bristly-hairs dorsally ...........

et: ee ee

ies Ticlaminata

9(7) 9th tergum with bi-pointed tubercle pointing forwards and backwards. Metapleural fan mostly pale, dorsum of abdomen with pale hairs except for dark hind-marginal hairs .. . nfSER RNS A ee, AG 39. E. bicuspidata

Figs 126-130. Male legs of Empis s.str. —126: E. caudatula, mid leg in anterior view; 127: E. aestiva, the same; 128: E. praevia, the same; 129: E. praevia, fore tarsus in dorsal view; 130: E. levis, mid leg in anterior view. Scale: 0.2 mm.

77

—

8th tergum with one-pointed tubercle pointing forwards. Metapleural fan black, dorsum of abdomen black pubescent, only sides and venter with pale hairs......... 38. E. gooti 10(6) Generally smaller species, at most 3.5—4 mm long. Mid tibiae with l.or 2 conspicuously long anterodorsal bristles on basal half, and another equally long preapical bristle, ventrally with short bristles not much longer than tibia isdeepads sadace te sae gene ee ees aoe 11 — Generally larger species, 3.5—5 mm long. Mid tibiae anterodorsally with 4-5 very strong bristles and equally long bristles also beneath 11(10) Aedeagus short, slightly overlapping genital lamellae, stouter at base, slender apically, and with a loop at middle. Mesonotum with 2 indefinite darker stripes between the rows of bristles when viewed from in front. Legs with a tendency to be light brownish. 4 scutellar bristles, outer pair very small. Larger, 3.5-4 mm lone... - se eeeeee 40. E. nuntia — Aedeagus long and slender. Mesonotum unstriped, almost uniformly grey dusted. Legs blackish. 2 scutellar bristles. Smaller, less than, 3:5.mm“( 2 ee we ce eee 12 12(11) Aedeagus with a distinct loop at middle, 8th tergum with a small, simple tubercle on each side. Wings clear, veins M and Cu distinct, DLOWIShEe see eee 29. E. planetica — Aedeagus evenly bowed above body, without loop, 8th tergum simple, without tubercles. Wings almost whitish, veins pale and very inconspicuous, especially veins Mand Cu .... Sirs He he ADA per pore 30. E. staegeri 13(10) Metapleural fan black, mid tibiae beneath with 4—5 strong black bristles. Thorax and abdomen uniformly dark grey dusted, dorsocentral

bristles

biserial,

hair-like.

Long-

legged species with conspicuously large genitalia, aedeagus very long and thin, threadlike, with 3 loops at base ... 24. E. acinerea — Metapleural fan whitish-yellow, mid tibiae beneath with about 10 strong black bristles. Thorax dull grey, abdomen contrasting subshining black, dorsocentral bristles irregularly triserial, bristle-like. Rather robust stronglegged species with small genitalia, aedeagus short and stout, not overlapping lamellae .. AS See Be PR es RS 14 14(13) Knees broadly yellowish, usually posterior four basitarsi also yellowish-brown. Wings brownish. Fore basitarsus with long black pu-

78

bescence posteriorly, the hairs longer than basitarsus is'deepie.. --..22- 26. E. genualis Legs entirely black. Wings almost clear or faintly greyish. Fore basitarsus with very short pubescence posteriorly, the hairs scarcely half as long as basitarsus is deep ... Dd See ae ee 25. E. syrovatkai 15(5) Metapleural bristles pale (with additional black bristles in nitidula and lepidopus). Aedeagus short, not extending much above genital lamellac:. Hawt Hes SOR a eee eee 16 Metapleural bristles black. Aedeagus long and slender, evenly bowed; if short (/imata), then withailoopmeeereent: Seer 20 16(15) 8th tergum at sides with large rounded tubercles, aedeagus rather stout, broadened apically. Acrostichal and dorsocentral bristles black, biserial. Smaller, shining black species about3-5 mim lone eee eer 36. E. nitidula 8th tergum simple, without tubercles. Large species, 4-5 mm long, if smaller (albopilosa) then aedeagus very slender towards tip, wings milky-white, and acrostichal and dorsocentral bristles whitish anteriorly, latter quadriSETI ys s.00< spd each Sees eee ily 17(16) Aedeagus very slender towards tip. Acrostichal bristles biserial, dorsocentral quadriserial, all numerous and very long, hair-like . eee ea ree, |) wee, BAT ac 18 Aedeagus very stout, broadened at tip. Acrostichal bristles biserial, dorsocentrals irregularly triserial, black, less numerous and bristle-like. Mid tibiae usually with 5 strong black anterodorsal bristles, and numerous similar long bristles in a double row ventrally. 4-6 scutellar bristles. Large species, 4-5 mm LONE» 5 net aeieeeele': = oh ese eee 19 18(17) Smaller, about 3-3.5 mm. Wings almost milky-white, veins whitish, and acrostichal and dorsocentral bristles mostly pale anteriorly. Mid tibiae with a dense row of at least 10 fine posterodorsal bristly-hairs more than twice as long as tibia is deep, ventral hairs adpressed and much shorter. 24 scutellar bristes) 4. Saree 21. E. albopilosa Larger, about 4.5mm. Wings almost clear with dark veins, and acrostichal and dorsocentral bristles black. Mid tibiae with 4 rather short bristles antero- and posterodorsally, those in posterior row longer, at least as long as tibia is deep, otherwise tibiae almost bare. 8-10 scutellar bristles ..... 20. E. lepidopus

19(17) Knees and basitarsi broadly yellowish (see SCCLIONMA) at Cee eine oesIe 26. E. genualis — Legs entirely black (see section 14) ........ See ee Se 25. E. syrovatkai 20(15) Smaller, about 3.5mm long. Aedeagus short, mostly concealed within genital lamellae, and with a loop before tip. Upper lobe of dorsal lamella with a brush of black hairs. Wings clear, abdomen completely black .... ON See Lane aE 31. E. limata — Larger, 4.5-5.5 mm long. Aedeagus long and slender, pennipes-like, evenly bowed. Dorsal genital lamella simple, long and slender. Wings faintly brownish, abdomen yellowish A DASH SA. foreets SH Nee 28. E. rufiventris 21(3) Smaller, 3-3.5 mm long. Genitalia ‘open’, pennipes-like, aedeagus long and slender, evenly bowed ........ 32. E. melanotricha — Larger species, 44.5 mm long. Genitalia ‘closed’, aedeagus short and stout or, if longer and more slender, then with a loop at about Ea: MINES OE he PO BRE. 22 22(21) Aedeagus long and slender, with a loop at middle, very thin at tip, and often concealed within lamellae. Hind femora with short bristly-hairs dorsally, much shorter than femur is SGD > ole Se iaee 33. E. tanysphyra — Aedeagus short and stout, widening at tip. Hind femora with a row of long anterodorsal bristles as long as, or longer than, femur is EMEC ee EEE 34. E. woodi 23(1) Abdominal pubescence whitish to yellowish, PRAVCINICOMplets AMM Aware n0t4 24 — Abdominal pubescence black. Small species, ern lOnge: eww. 28g 27 24(23) Large, about 5 mm long, with golden-yellow (rarely almost whitish) pubescence on abdomen, coxae, metapleura and humeri. All basitarsi slender. Aedeagus short, broad at base, and concealed within lamellae .......

0 'y.htho esQE a 22. E. serotina S$mall species, 2.5-3.5 mm long, abdominal pubescence always whitish. Genitalia small and closed, base of short aedeagus usually invisible (chioptera-complex) ........... 25 25(24) Humeri and notopleural depression with -minute white hairs in addition to the black bristles. Metapleural fan completely pale. Body rather densely grey to bluish-grey dusted. Wings milky-white, fore and hind basitarsi swollen. Antennae with a short style, onethird length of 3rd segment. 2 scutellar bris: LES bod A a oe 41. E. chioptera — Humeri and notopleural depression with

—

black hairs. Metapleural fan black or brownish, sometimes with additional pale bristles (sometimes mostly pale in northern populations). Rather subshining black species, fore and hind basitarsi not very stout, slightly deeper than corresponding tibiae at tip. Antennal style long, almost as long as 3rd segIDOLOS Ac iis or can in PA 26 26(25) Mesonotum shining black when viewed from above; 2 scutellar bristles. Wings faintly milky-white, halteres blackish. Mid tibiae with 2 dorsal bristles (near base and at middle), about 3 times as long as tibia is deep, a long preapical bristle, and 3 anteroventral bristles about middle twice as long as tibia is deep. Hypandrium bare . 42. E. prodromus — Mesonotum finely grey dusted from all points of view; 2—4 scutellar bristles. Wings almost clear, halteres rather brownish, at least on stalk. Mid tibiae with 4—5 long dorsal bristles on basal half in addition to preapical bristle, ventrally with shorter bristles arranged in pairs. Hypandrium with long bristly-hairs .. NERA RL Oe RIA AME 43. E. dasyprocta 27(23) Anal vein complete, reaching wing-margin. Fore and hind basitarsi cylindrical, scarcely stouter than tips of tibiae. Subshining black species, lateral genital lamellae with a long, curved, apical seta. Wings rather milky-white, veins fine and pale except darkened C, R, and ated eiaah lara een oO 44. E. caudatula — Anal vein abbreviated, disappearing before wing-margin. Wings clear ............. 28 28(27) Fore tibiae dorsally with fine short hairs not longer than tibia is deep. Mid tibiae with 1 or 2 long dorsal bristles at base and in basal third (very rarely 3 bristles), otherwise with short hairs, preapical bristles scarcely differentiated. Aedeagus rather long and very thin apically, free at base. Usually 2 scutellar bristles REPRE Se ST ae 45. E. aestiva (The closely similar species E. (Coptophlebia) vitripennis Meigen has, in addition to an abbreviated vein M,, aedeagus yellowish and very stout at base, and mid tibiae with 1-4 dorsal bristles on basal half and an equally long preapical bristle.) — Fore tibiae dorsally with bristly-hairs much longer than tibia is deep. Mid tibiae usually with 4-5 long dorsal bristles arranged evenly alongithe whole length Way seas eee 29 29(28) Fore tarsi and mid basitarsus without preapical bristles. Hind basitarsus with short bristly-hairs above, not longer than basitarsus

79

—

is deep. Aedeagus very slender at tip, hypandrium small. Dorsocentral bristles uni- to biserial, rather short and fine; usually 4 scutellar bristles xetiaind ase 47. E. levis Fore basitarsus with a circlet of strong preapical bristles, 2nd and 3rd tarsal segments with long preapical bristles as long as following tarsal segment; mid basitarsus with a strong preapical bristle. Hind basitarsus with strong dorsal bristles, longer than basitarsus is deep. Aedeagus not narrowed at tip, its basal half enclosed in the large hypandrium. Dorsocentral bristles bi- to triserial, long and bristlelikesusually 2iscutellar bristles eer

Females 1 Halteres pale; if brownish (pennipes), the species is included in both sections ...... 2 — Halteres black, at most dark brown..... 21 2(1) Legs not pennate, covered with ordinary hairs and bristles, without flattened hairs. Abdominal pubescence pale, at least on basal segments, .Pckea scious eae eee 3 — Legs distinctly pennate, or at least hind legs withitlattened| hairs: ere err 5 3(2) Metapleural fan pale, dorsocentral bristles quadriserial. Abdomen polished _ black.

—

Wings indistinctly brownish, almost clear. Small species, 3-3.5 mm) long -..7........-. kuin cae etic 21. E. albopilosa Metapleural fan black, or at least partly black, dorsocentral bristles biserial. Abdomen dull blackish-grey, at most subshining

(4(3) Smaller, 3-4 mm long. Wings clear. Hind femora compressed, twice as deep as hind tibiae, dorsal hairs with a tendency to be flattened; netted atts 37. E. laminata — Larger, generally over 4mm long. Wings deep brown. Hind femora slender, narrowly conical, as deep as hind tibiae Stel cs eee aera ee 24. E. acinerea 5(2) Labrum very long, about 3 times as long as head is high. Abdominal pubescence and metapleural bristles black. Halteres often darkened, wings dark brown. Legs strongly pennate on posterior four femora, hind tibiae, and on mid tibia dorsally; shorter flattened

hairs on fore tibiae and anterior four basitarsi dorsally, and on mid tibia beneath at base. Large species, 4-S mm long ..............

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—

6(5)

—

7(6) —

8(7)

—

9(7)

—

Labrum shorter, at most slightly more than twice as long as head is high (care must be taken with a long labium if it is porrect) .... 6 Abdominal pubescence and metapleural bristles black. Medium-sized species, generally about 3.5—4.5 mm long Abdominal pubescence pale, at least at base of abdomen, metapleural bristles pale or black @ iniccaeise ee eee 10 Posterior four femora broadly pennate on bothisidesien®:, aie. cee eee 8 Posterior four femora at most with short adpressed pennation, mostly with slender flattened hairs, the longest on hind femur beneath towards tip at most as long as femur is deep. Hind tibiae with ordinary (scarcely flattened) bristles above, and with additional long bristly-hairs intermixed Hind tibiae with distinct pennation of equal length dorsally and on basal half beneath. Fore tibiae and basitarsi with ordinary bristlyhairs dorsally, mid tibiae fringed above. Wings light brownish ........ 34. E. woodi Hind tibiae short pennate dorsally, with very short flattened hairs beneath on basal half. Anterior four tibiae and fore basitarsi with small flattened hairs dorsally. Wings darker brownaeecnea eee 33. E. tanysphyra Mid basitarsus long, much more than half length of mid tibia; posterior four basitarsi with 4—5 pairs of stubby ventral spines as long as basitarsi are deep. Hind femora with flattened hairs, without distinct pennation. Prescutellar depression and interspaces dark brownish-grey dusted when viewed from behind. Wings uniformly brownish .......... aan Ane eet: Be 32. E. melanotricha Mid basitarsus half as long as mid tibia; posterior four basitarsi with short adpressed bristly-hairs. Hind femora pennate, with a short

adpressed pennation dorsally along the whole length, on apical two-thirds beneath with pennate fringes almost as long as femur is deep. Prescutellar depression and interspaces light grey in posterior view. Wings subhyaline, almost clear except faintly brownish costal.marginikisnaeven meh Somat 38. E. gooti 10(6) Legs strongly pennate, also on fore femora and.mid coxaéase.qietaunies sedeen ee 11 — Fore femora and mid coxae with only ordinary hairs and bristles, without pennation ..

wticcince Tate eient bat. he ee ae 12 11(10) Front half of mesonotum densely clothed

with fine, long, pale hairs. Acrostichal and dorsocentral bristles multiserial, very narrowly separated. Legs blackish. Large species, 5-6 mmlong

..........

23. E. decora

—

Front margin of mesonotum bare, without whitish hairs. Acrostichal and dorsocentral bristles biserial, widely separated. Legs rather brownish, with knees and posterior basitarsi often yellowish-brown. Somewhat smaller, 4-5 mm long ...... 26. E. genualis 12(10) Hind femora pennate; legs more or less pennate also on mid pair, or mid femora clothed miinelatteneqdubristlesemt stesis. 13

—

Femora without pennation or flattened hairs,

but hind tibiae with dorsal pennation about as long as tibia is deep, and with shorter fringes on basal half beneath. Mesonotum light grey dusted, dorsum of abdomen shining black. arcer,about 4.5 mnt lone j.sG6 oes « Pes iSie fdas ese alls 20. E. lepidopus 13(12) Fore legs with only ordinary hairs or bristles, no pennation or flattened bristles. Medium-sized species, about 3-4 mm long ... 14 — Fore tibiae and/or fore basitarsi dorsally with distinct flattened bristles; if in doubt, a spe-

cies with somewhat broadened brown wings and 2 scutellar bristles (planetica) belongs [ETE So oll pads eeety ea ee ee er ee 18 14(13) Shining black species, mesonotum with greyish dusted notopleural depression and prescutellar area. Abdominal pubescence pale, 4 scutellar bristles, outer pair long, only slightly shorter than the inner pair ........ Berar chance eee & slelafassbans «is 36. E. nitidula — Dull grey, densely dusted species on thorax READ COMIC Mse-4.feleys ale2s,..ceaetseh ait aes» 15 15(14) Mesonotum unstriped, or with 2 indefinite darker stripes between the rows of bristles. Abdomen covered with pale hairs. 2 scutellar bristles, if 4, then outer pair small, less than

—

amaeneth Of inner pairs.) 20.56.6265 16 Mesonotum (viewed from above) with 3 indefinite darker stripes along the lines of bristles. Dorsum of abdomen with some additional black hairs. 4 scutellar bristles, outer pair long, not very much shorter than inner Rolie Cee relent Sa wuists lyk PySete Se 1)

16(15) 2 scutellar bristles. Mesonotum uniformly dark grey, without stripes. Metapleural bristles often blackish: Legs black, hind femora

and tibiae with long dorsal pennation about as long as corresponding femora and tibiae

are deep. Wings almost clear (distinctly brown in planetica). Generally smaller, about SFSou sogoancagopEe 30. E. staegeri — 4 scutellar bristles, outer pair very small. Mesonotum with 2 indefinite darker stripes between the rows of bristles when viewed from in front. Metapleural bristles entirely pale. Legs rather lighter brown, hind femora with much shorter pennation and hind tibiae only fringed above (as in nigripes and bicuspidata). Wings faintly brownish. Generally larger, about4mm long ..... 40. E. nuntia 17(15) Metapleural bristles mostly black, often with additional pale bristles. Hind tibiae dorsally with rather long flattened bristles, and some additional longer simple bristly-hairs. Dorsum of abdomen with numerous black harstesdey tse actaist eta 35. E. nigripes — Metapleural fan mostly pale, rarely with additional black bristles. Hind tibiae with more uniform short pennation (or short flattened bristles) dorsally, often mixed with several longer, flattened bristly-hairs. Dorsum of abdomen mostly with pale hairs............. 5 oct Seat 3,ROM ERC ee TOS 39. E. bicuspidata 18(13) Large species, about 4.5 mm long. Posterior four femora and tibiae strongly pennate.... a5 es asRee OR ee 19 — Smaller species, about 3-3.5 mm long. Legs lessidistinctly pennatey.. eee te 20 19(18) Larger, 4.5-5.5 mm, long-legged species, metapleural fan black. Fore tibiae above with long flattened bristles, fore basitarsus without pennation or flattened hairs. Wings light brownish. Mesonotum dull grey, abdomen brownish, shining........ 28. E. rufiventris — Smaller, about 44.5 mm, robust species with stronger legs, metapleural fan pale, rarely with a few black additional bristles. Fore tibiae with flattened bristles dorsally on apical half only, fore basitarsus distinctly fringed along its whole length. Wings darker brown. Abdomen subshining black .............. i SER, oer 25. E. syrovatkai 20(18) 2 scutellar bristles, thorax and abdomen dull dark grey. Metapleural fan pale, with additional

—

black

bristles.

Wings

brownish,

somewhat broadened ..... 29. E. planetica 4scutellar bristles (outer pair small), thorax and abdomen subshining black. Metapleural fan black. Wings very dark brown, not broadEMCI Preoecaatasie lenteiaensenvotu: 31. E. limata

81

21(1) Abdominal pubescence pale. Anal vein com-

plete, reaching the wing-margin........ 22 — Abdominal pubescence blackish ....... Ds 22(21) Large species, about 5 mm long. Abdomen, coxae and metapleura covered with goldenyellow hairs (rarely almost whitish). Mid and hind femora short:pennate, hind tibiae on basal half beneath and mid tibiae at base beneath with finer pennation; fore tibiae dorsally fringed. Humeri with pale hairs and one contrasting black bristle .... 22. E. serotina — Smaller species, 2.5-3.5 mm long. Abdominal pubescence always distinctly whitish ... sani cchen Oe eal eee CR eae 23 23(22) Humeri and notopleural depression anteriorly covered with minute whitish hairs in addition to black bristles. Metapleural fan whitish. Mesonotum greyish dusted, wings brownish, broadened. Hind femora fringed above, pennate on apical half posteroventrally, with several long black bristly-hairs anteroventrally before tip. Hind tibiae finely fringed above, mid femora with similar fringes above and posteroventrally on apical half ASSL eae ie 41. E. chioptera — Humeri and notopleural depression with entirely black hairs and bristles. Metapleural fan black or brownish, sometimes with addi-

tional pale bristles, rarely mostly pale. Wings notbroadenedieeaneeneenee eee 24 24(23) Mesonotum rather shining black. Hind femora convex above and distinctly fringed there, ventrally almost straight and clothed with minute dark hairs except for a few longer anteroventral bristly-hairs towards tip. Hind tibiae stout, distinctly fringed above. Wings faintly brownish, almost clear apically. 2 scutellambiriStlesiaeenteeie 42. E. prodromus — Mesonotum greyish dusted. Hind femora and tibiae rather slender, femora pennate beneath, finely fringed above, tibiae without dorsal fringes. Wings uniformly faintly brownish clouded. Halteres with a tendency to be paler, brownish. 2-4 scutellar bristles ...... St EIR 43. E. dasyprocta 25(21) Large, about 4-S mm long. Labrum very long, about 3 times as long as head is high. Legs strongly pennate on posterior four femora, hind tibiae, and also on anterior four tib-

—

82

iae above. Colour of halteres varying from yellowish to dark brown (see section 5) .... sitters Enh Lotivs Sr a ee es 27. E. pennipes Small species, about 2.5-3.5 mm long. Labrum much shorter. Anterior four tibiae

without pennation or flattened hairs, halteres blackishvraiiera Oe. eee 26 26(25) Anal vein complete, reaching the wingmargin. Mesonotum thinly grey dusted. Posterior four femora pennate on both sides; hind femora distinctly convex above, anteriorly before tip with 3-4 long flattened hairs, hind tibiae without pennation or flattened bristles. Wings faintly yellowish-brown at base, clear towards tip .... 44. E. caudatula — Anal vein abbreviated, not reaching wingmargin. Mesonotum almost shining black. Legs with a different combination of characters os. edd OR 2 27(26) Posterior four femora and hind tibiae strongly pennate. 2 scutellar bristles, wings father brownishete 45. E. aestiva — Legs not pennate, covered with only ordinary hairs and bristles, without flattened hairs . . 12g hae ed. Be ee 28 28(27) Usually 4 scutellar bristles. Wings rather dark brown. Hind femora rather long and slender, and practically bare ... 47. E. levis — Usually 2 scutellar bristles. Wings almost clear. Hind femora shorter and stouter, covered with several anterodorsal bristly-hairs, and similar anteroventral ones on apical third or more, the hairs not as long as femur is deep Fee eae Cs is A 46. E. praevia

20. Empis s.str. lepidopus Meigen, 1822 Figs 101, 108, 131-134. Empis lepidopus Meigen, 1822:23.

Generally large (4.2—5.8 mm) with dull grey thorax and shining black abdomen, pale halteres, thoracic hairs and bristles black, but metapleural fan and abdominal pubescence whitish. Legs shining black, tibiae in male with a few short spine-like bristles, female with only hind tibiae above broadly pennate. 3d. Eyes meeting on frons but upper ommatidia only slightly enlarged, very closely resembling EF. albopilosa in this respect. Occiput dull grey, with rather dense black bristles above, on lower third with longer and finer whitish-yellow hairs. Antennae black, arista slender, about two-thirds length of 3rd segment. Palpi black, covered with several fine yellowish hairs. Proboscis rather long, labrum black and nearly twice as long as head is high. Thorax densely grey dusted, more brownishgrey on mesonotum, with darker (rather velvety-

black) not very distinct stripes along the lines of bristles, clearly visible from all points of view. Mesonotum with dense bristles, acrostichal and dorsocentral bristles black, very long and hair-like (as long as 3rd antennal segment without style), very numerous, former 2-serial, latter broadly 4serial, becoming irregularly 2-serial and very long and bristle-like on posterior half of mesonotum. Large bristles conspicuous, a humeral with numerous additional small black hairs, a posthumeral, 3 to 4 strong notopleurals with additional black and yellowish hairs, 1 or 2 supra-alars, a postalar, and about 10 scutellar bristles, usually 3 pairs very long and strong. Pronotum with rather short black bristles in a row, but sides of prosternum and episterna with long yellowish hairs. Metapleural fan dense, long, whitish-yellow, often with a few additional black bristles. Wings clear with dark veins becoming yellowish

axillary excision acute. Squamae pale with very white fringes, halteres yellowish. Legs long and slender, including all basitarsi, very shining black except for dark grey dusted coxae. All femora conspicuously bare, ventrally with fine yellowish hairs, the longest on mid femur in basal third (about as long as femur is deep), mid femora with a few, about 3 to 5, long black anterodorsal bristles and with silver pile beneath. Fore tibiae with a posterodorsal row of black bristly-hairs about as long as tibia is deep, anterodorsally with 2 shorter but stouter black bristles in apical half. Mid tibiae antero- and posterodorsally with about 4 evenly spaced short black spine-like bristles, with similar spines ventrally, with the same bristling on hind tibiae but ventral spines absent, replaced by minute dark pubescence beneath. All basitarsi black spinose beneath, basal three tarsal segments with equally long black spine-like bristles in terminal circlets. Abdomen polished black on dorsum, sterna finely grey dusted, pubescence whitish-yellow, longer at sides of anterior segments. Genitalia

at base, a distinct black costal bristle. Radial fork

unusually acute for Empis s.str. and vein R, slightly bowed (not quite straight), anal vein complete and

133 Figs 131-134. Male genitalia of Empis (s.str.) lepidopus Meigen (NL, Enspijk, Mariénwaard). — 131: postabdomen; 132: lamellae in lateral view; 133: dorsal lamella in anterodorsal view; 134: aedeagus with hypandrium. Scale: 0.2 mm.

83

small, in general structure resembling those of E. albopilosa, but lateral lamellae smaller, rather triangular, with long black bristles on hind margin, rather like 8th sternum; dorsal lamellae with short,

mark or southern Sweden seems very likely. A typical early spring species; my records are from 27 April to 11 May.

fine, dense, black pubescence above. Aedeagus

21. Empis s.str. albopilosa

slender, yellowish-brown

de Meijere, 1935

on the free basal half,

then turned up and concéaled within lamellae. Length: body 4.2-5.2 mm, wing 5.3-5.6 mm. 2. The main distinguishing features as in male, but mesonotal pubescence (acrostichal and dorsocentral bristles) very short, wings dark brownish, and legs differently bristled. Legs shining black, especially on femora that are almost bare, ventral pale hairs practically absent, but mid femur with posterodorsal black bristles as in male, and hind femur beneath at tip with 2 or 3 flattened bristles. Fore tibiae with similar black posterodorsal bristles, perhaps more densely set than in male, and anterodorsal spine-like bristles on mid tibiae replaced by a row of flattened bristly-hairs. Hind tibiae somewhat compressed laterally, posterodorsal spine-like bristles of male present, but anterodorsally tibia long pennate, though pennation is slightly shorter than the laterally-compressed tibia is deep, and ventrally with similar flattened bristlyhairs as on mid tibia anterodorsally. Basitarsi spinose as in male, terminal circlets smaller. Tarsi of-

ten distinctly curved at tip. Abdomen shining black except for grey dusted apical two segments and cerci, pubescence not very much shorter than in male. Length: body 4.6-5.8 mm, wing 5-6 mm. E. lepidopus has much in common with E. albopilosa and both species are among the most primitive forms of the subgenus, with several characters in their plesiomorphous states. The small upper ommatidia of the eyes, and the rather acute radial fork on the wings with curved vein R,, make E. lepidopus somewhat exceptional and tend to set it aside from Empis s.str. and rather closer to Leptempis. E. albopilosa may easily be distinguished by the white acrostichal and dorsocentral bristles in the male, the milky-white wings and long bristles on tibiae, whilst the hind tibiae in the female are not pennate. The holotype female of E. lepidopus was identified and redescribed by Syrovatka and Chvala (1986).

Distribution. A widely distributed but only locally common species known from Spain and Italy in the south, from the central parts of Europe (Germany, Switzerland, Hungary), and from the Netherlands where it has been recently collected on several occasions in large numbers by B.V. Aartsen at Mariénwaard (Enspijk Gld.). Its occurrence in Den-

84

Figs 102, 109, 135-137.

Empis albopilosa de Meijere, 1935:126. Rather smaller (3.2-4.3 mm), mostly pale pubescent and bristled species with yellowish halteres, thorax dull grey, abdomen contrasting polished black. Male with milky-white wings and densely long-bristled legs, female wings faintly clouded and legs not pennate. 3. Eyes touching for a long distance on frons but upper ommatidia not very enlarged, only about twice as large as those below. Occiput rather dark grey dusted, covered with dense rather fine bristlyhairs, black above, whitish (or at least light brownish) below. Antennae black, style long and slender, not very much shorter than 3rd segment. Palpi small, black, with a few long dark hairs at tip. Labrum rather short, not very much longer than head is high, barely 1.5 times as long. Thorax rather uniformly densely grey dusted, viewed from above and behind with 3 blackish stripes along the lines of bristles, mesonotal bristling long and fine. Large marginal bristles blackish (a humeral, a posthumeral, an intrahumeral, 3

notopleurals, a supra-alar, a post-alar), as well as last 3 pairs of prescutellar dorsocentral bristles, and two scutellar bristles, often with a minute dark

or pale hair each side. Acrostichal and dorsocentral bristles whitish or at least light brownish, long, hair-like, nearly as long as 3rd antennal segment without style; former 2-serial, latter irregularly 3-

to 4-serial for a short distance; anterior part of mesonotum between humeri bare (unlike E. decora). Pronotum with a row of small blackish bristles,

but sides of prosternum and episterna with longer whitish hairs. Metapleural fan whitish-yellow. Wings milky-white with whitish veins, only costa blackish and vein R, yellowish, no costal bristle.

Radial fork not at all right-angled. Anal vein very fine, incomplete, disappearing before reaching wing-margin. Squamae pale with concolorous fringes, halteres whitish yellow, stalk brownish.

Legs long and slender, polished black or blackish-brown (except for greyish coxae), only hind tibiae slightly dilated on apical half and hind basitarsus also stouter, as deep as tibia is broad at tip. All femora almost bare, covered with fine pale hairs, ventrally shining, without silver pubescence;

mid femora with anteroventral yellowish hairs becoming longer towards tip, and both posterior femora with outstanding pale bristly-hairs anterodorsally on basal half, those on mid femora about as

long as femur is deep. Fore tibiae with darkened posterodorsal bristly-hairs slightly longer than tibia is deep, and similar but shorter hairs also present on basitarsus. Mid tibiae posterodorsally with a row of at least 10 very long, fine black bristly-hairs, longest at middle (at least 3 times as long as tibia is deep), anterodorsally with more numerous but shorter reclinate bristles, ventral pubescence shorter than tibia is deep. Mid basitarsus dorsally with dark bristly-hairs slightly longer than basitarsus is deep. Hind tibiae with fine, long, dark bristles above, the bristly-hairs becoming longer towards tip, those in posterodorsal row longer and blacker, slightly longer than tibia is deep, and with similar bristles also on basitarsus; hind tibiae ventrally

with short adpressed hairs. No terminal circlets on tarsal segments, at most longer fine hairs at tip, and basitarsi not distinctly spinose beneath. Abdomen shining black, rather densely whitish pubescent, longer at sides of basal segments. Genitalia small, lateral lamellae long-ovate, smooth

and polished, hind margin with a row of pale bristles, dorsal lamellae practically concealed. Aedeagus clearly visible on a long basal part, polished black and slender, tip still narrowing, concealed within lamellae. Length: body 3.3-4.2 mm, wing 3.3-4.2 mm (lectotype body 3.5 mm, wing 4.0 mm). ?. The 2-serial acrostichal and 4-serial dorsocentral bristles very short and extensively darkened, almost blackish, large marginal bristles smaller than in male. Wings faintly greyish-brown clouded, not at all clear, veins brownish. Abdomen shining black, pubescence whitish but short and sparse, apical two segments and cerci dull grey. Legs shaped very much as in male but everywhere with short hairs and bristles, only posterior four tibiae dorsally with a few short (not as long as tibiae are deep), rather stout, black bristles, without

pennation or flattened hairs. Length: body 3.2-4.3 mm, wing 3.4-3.8 mm. E. albopilosa is undoubtedly most closely allied to E. lepidopus, but the latter has all the mesonotal hairs and bristles black in male, clear wings with

distinct dark veins, metapleural fan partly black, tibiae with only a few short black rather stubby

Figs 135-137. Male genitalia of Empis (s.str.) albopilosa de Meijere (CZ, Moravia, Trinec). — 135: postabdomen; 136: dorsal lamella in lateral view; 137: aedeagus with hypandrium. Scale: 0.2 mm.

85

spines, and genitalia with black bristles posteriorly; the female of E. /epidopus can be easily distinguished by the long pennate hind tibiae, and legs otherwise with only ordinary hairs and bristles. Lectotype designation. Described by de Meijere (1935) from Linschoten, the Netherlands. There are 4 d and1 2 syntypes in the de Meijere Collection in Amsterdam from Linschoten, one male from 5 June 1920, the other specimens from 14 May 1921. A male labelled ‘Linschoten 14. V.21 de Meijere’ was labelled by me in 1986 and is herewith designated as lectotype. Distribution. Overlooked, but undoubtedly a widely distributed species in cool and temperate Europe, rather common throughout the Netherlands (V.S. van der Goot, letter), and should also occur in the southern parts of Scandinavia; unknown from Great Britain. Apart from the Netherlands, known to me only from central Bohemia, where it is common in lowland biotopes in the vicinity of Prague, especially at the beginning of May. A typical early spring species; my records are from 2 May to 5 June.

22. Empis s.str. serotina Loew, 1867 Figs 104-105, 138-140.

Empis lineata Meigen, 1804:224 (not Scopoli, 1763 and Villers, 1789).

Empis serotina Loew, 1867:55, 49. Empis obscuripennis Strobl, 1893:66.

Rather large, about 5 mm, with blackish halteres and dense golden-yellow pubescence on coxae, metapleura and abdomen, thoracic bristles black. Mid and hind femora short pennate in female. 3. Eyes with ommatidia on upper half considerably enlarged, occiput with long black bristlyhairs above becoming longer and whitish-yellow below. Antennae black, style half as long as 3rd segment, palpi black. Proboscis rather short, labrum scarcely 1.5 times as long as head is high. Thorax densely and rather dark grey dusted, mesonotum uniformly grey in anterior view, or with somewhat brownish stripes along the lines of bristles, in dorsal and posterior views the stripes becoming almost velvety-black, and the interspaces and prescutellar depression very light grey. Large bristles black: a humeral and 3 posthumeral bristles small, other bristles stronger and longer, 3-4 notopleurals, a supra-alar, a postalar, and 4

scutellars. Acrostichal and dorsocentral bristles small and hair-like, black to dark brown, former

regularly 2-serial and slightly diverging, latter ir-

86

regularly 3-serial, ending in 4 long prescutellar bristles. Humeri, posthumeral region and notopleural depression covered with small yellowish hairs. Sides of prosternum and episterna with yellowish to golden-yellow hairs, pronotum with a row of erect short bristly-hairs varying from golden-yellow to wholly black. Metapleural fan long, golden-yellow to yellowish. Wings faintly brownish-grey clouded, with distinct blackish veins and large, dark brown costal stigma. The wings are considerably different from the typical shape in Empis s.str., being rather long, narrowed apically; radial fork consequently not as wide as usual, rather acute, and discal cell elongate,

not truncate. Anal vein complete, axillary excision acute. Squamae dirty grey with narrow blackish margin, fringes long, golden-yellow; halteres blackish, base of stalk brownish. Legs rather shining black except for greyish coxae, rather long and slender, basitarsi not swollen, though fore basitarsus obviously broader than tibia at tip. Anterior four legs including tarsi covered with soft whitish pile beneath, becoming reddish on anterior tibiae in front. Bristles and hairs black,

all rather short, but fine ventral hairs on fore femora pale, ending in 3 black bristly-hairs at tip. Posterior four femora with a double row of ventral bristles about half as long as femora are deep, tibiae dorsally with longer bristles arranged irregularly in two diverging rows, longest on hind tibiae anterodorsally, ventrally with a row of bristles on basal half nearly as long as tibia is deep, merging towards tip with shorter ventral pubescence. All tarsi with short black spine-like bristles in apical circlets, and hind basitarsi finely spinose beneath. Abdomen subshining black, finely brownish dusted, more densely so on posterior terga, pubescence golden-yellow to silvery-yellow, especially long and dense on basal segments at sides and on venter. Genitalia small, closed, aedeagus brownish and short, enclosed within the lamellae, very thin at tip. Lateral lamella ovate, with long hairs posteriorly, dorsal lamellae hidden, very black and rugose on inner surface; ventral lamella (hypandrium) shining, cornet-like, posteriorly with a striking pointed tuft of long black spinose bristles. Length: body 4.5-5.8 mm, wing S—6 mm. 2. Abdominal pubescence shorter and legs differently bristled, otherwise as in male, including

brownish clouded wings and bristling on thorax. Legs pennate, all femora laterally flattened and consequently broader than in male. Posterior four femora short pennate on both sides (half as long as femora are deep), mid femora ventrally on apical half only, additional flattened bristles also on basal

Figs 138-140. Male genitalia of Empis (s.str.) serotina Loew (NL, T. Harde Gld.).—138: postabdomen; 139: dorsal lamella in anterodorsal view; 140: aedeagus with hypandrium. Scale: 0.2 mm. :

third of hind tibiae beneath and on extreme base of mid tibiae. Fore femora with short black fringes dorsally and ventrally, fore tibiae with similar fringes posterodorsally. Length: body 4.5-5.8 mm, wing 4.6-5.8 mm. E. serotina is very distinct from all other species in the northern fauna because of the golden-yellow pubescence on thorax and abdomen together with black halteres. I have seen several specimens from the Netherlands in which the characteristic golden-yellow hairs on prothorax, coxae, metapleura and abdomen were rather whitish, not yellowish. A lectotype for E. serotina Loew was designated by Syrovatka (1991). Distribution. An overlooked species because of its very late summer occurrence, known to me from Denmark and the south of Sweden (Skane); only 7 3 and 8 @ have been examined. Mainly in the second half of August and first half of September, but there is one female in Copenhagen Museum collected by Lundbeck exceptionally early, on 31 July 1920 at Tinbaek Molle in the eastern Jutland. — Unknown from Great Britain but not uncommon in the Netherlands and Schleswig Holstein, south to northern France and Austria. Locally common in central Europe, but only during a short period in late summer; my dates are from 7 August to 9 September.

23. Empis s.str. decora Meigen, 1822 Fig. 141.

Empis decora Meigen, 1822:22. Empis villosa Macquart, 1823:161.

Empis maura Macquart, 1838:160 (not Fabricius, 1776). Empis dasythrix de Meijere, 1918:132. Empis (Pterempis) andalusiaca Engel, 1946:355. Empis (Pterempis) pallipes Engel, 1946:367 (as var. of E. decora).

Large (5-6 mm), with pale halteres, whitish metapleural fan and abdominal pubescence, anterior half of mesonotum densely clothed with long, soft, whitish hairs. Legs including coxae strongly pennate in female. 3d. Occiput greyish, rather densely clothed with long whitish hairs, only postocular bristles blackish. Antennae black, style not very much shorter than 3rd segment, labrum about twice as long as head is high. Thorax greyish dusted, mesonotum somewhat subshining from some angles, viewed from behind with darker stripes along the lines of bristles. Anterior half of mesonotum densely clothed with long, soft, whitish hairs leaving only narrow stripes between the multiserial acrostichal and dorsocentral bristles, all darkened posteriorly, the latter usually ending in 3 stouter black prescutellar pairs. Humeral bristle pale and similar to other pubescence, but 3 notopleurals, 1-2 supra-alars, a postalar and 2 pairs of scutellar bristles black and distinct. Prothorax with white hairs, metapleural fan whitish. Wings very faintly brownish clouded, costal margin and base rather yellowish, all veins dark, distinct. A small blackish costal bristle. Squamae and halteres yellowish, former with pale fringes. Legs subshining black, rather long and slender,

87

=~

Fig. 141. Male genitalia of Empis (s.str.) decora Meigen (from Collin, 1961).

hind tibiae dilated towards tip and fore and hind basitarsi slightly swollen. Fore femora with short hairs; mid femora with two rows of long yellowish bristles beneath, those in posterior row longer and more numerous, some additional long yellowish

shorter and darker at tip and above, dorsal lamel-

hairs anteriorly, and anterodorsally with a row of

very long darkened bristly-hairs; hind femora with short pale hairs beneath, similar but longer bristles

humeral bristle darkened, and wings hardly darker than in male. Legs rather blackish-brown, shining, and strongly pennate. Mid and hind coxae pennate

posteriorly at base, and black anterodorsal bris-

anteriorly, fore femora pennate at base both dor-

tles. Fore tibiae stouter than is usual, posterodorsally with bristly-hairs about as long as tibia is deep, posteriorly and ventrally densely short pubescent; mid tibiae scarcely narrower than fore tibiae, with about 10 very long bristles dorsally, with somewhat shorter bristles ventrally, the bristles in anteroventral row towards tip the longest, and anteriorly towards tip with dense long pubescence; hind tibiae with long dense bristles above, ventrally with short adpressed bristly-hairs. Mid and hind basitarsi with long bristles above, spinose beneath, mid basitarsus with several long anterior bristles

sally and ventrally, posterior four femora strongly pennate on both sides except for basal third of mid femora beneath; mid tibiae pennate dorsally and

on basal half, and all tarsi with circlets of rather

the dense, long, soft, whitish pubescence on ante-

long black preapical bristles. Abdomen subshining black when viewed from above, rather brownish-grey dusted in anterior view, pubescence long and whitish, especially long at sides of anterior segments; sterna with long pale

rior half of mesonotum. A variable species as regards structure and colour of bristling both on legs

hairs, 8th sternum with long bristles on hind mar-

gin. Genitalia large and very distinctive because of the very long thin aedeagus that is evenly bowed above lamellae. Lateral lamellae long ovate, hind

margin

88

with

long pale bristly-hairs

becoming

lae rather concealed, somewhat curved, and short brownish pubescent. Length: body 5S—6 mm, wing 5.2-5.7 mm.

2°. With shorter hairs on thorax and abdomen,

towards base ventrally, hind tibiae above on apical

half and ventrally at about middle; short pennation also on fore tibiae dorsally and ventrally towards tip, on basal three segments of fore tarsi above, and on hind basitarsus dorsally at base. All basitarsi spinose beneath. Length: body 5—6.2 mm, wing 5.2-5.5 mm. E. decora can be easily distinguished from all other larger-sized northern European species by

and thorax, as is also reflected in the large number

of synonyms. Meigen’s types were revised by Collin (1961), and a lectotype was designated by Syrovatka and Chvala (1986). I have re-examined the types of E. dasythrix de Meijere in Amsterdam and can confirm the long-accepted synonymy with E. decora.

Distribution. This does not seem to be rare in the south of England. It is also recorded from Belgium and the Netherlands (although recent captures since de Meijere are unknown to me), and its occurrence in Denmark and southern Sweden is very possible. The record of E. decora from Denmark given by Lundbeck (1910) refers to E. syrovatkai. There are nine specimens of E. decora in the Palaearctic Collection in Helsinki collected by H. Lindberg at Algeciras, Spain, identified as E. andalusiaca. In NW Europe in May and June, in the Mediterranean (Spain, France, North Africa) as early as April. Unknown from the central parts of temperate Europe, but clearly distributed northwards along the warmer Atlantic coast.

24. Empis s.str. acinerea Chvala, 1985 Figs 110, 142-144.

Empis cinerea Zetterstedt, 1855:4609 (not Fabricius, 1775 and Miiller, 1776). Empis (s.str.) acinerea Chvala, 1985:389 (new name).

Rather large (4-5 mm), densely grey dusted and long-legged species with pale halteres, whitish abdominal pubescence and black metapleural bristles. Male with clear wings and a remarkably long thread-like aedeagus; females with slightly broadened deep brown wings and simple legs, no pennation or flattened hairs. d. Occiput dull grey, covered with rather fine bristles that are black above, brownish below. An-

tennae black, style not very much more than half length of 3rd segment, 2nd segment globular and often brownish, basal segment longer and narrower. Palpi black, very small, with a fine black termi-

nal hair as long as palpus. Proboscis not very long, labrum scarcely 1.5 times as long as head is high. Thorax densely rather light grey dusted even on mesonotum, latter with three brownish stripes along the lines of bristles, the central one becom-

ing blackish in dorsal and posterior views. All thoracic hairs and bristles black. Acrostichal and dorsocentral bristles rather long and hair-like, at least

as long as antennal style, both narrowly biserial, the latter uniserial, longer and bristle-like posteri-

orly. One strong humeral bristle with a tuft of about

6 small

hairs

anteriorly,

a posthumeral,

about 5 notopleurals with additional smaller hairs, asupra-alar, a postalar, and 4 to 6scutellar bristles,

outer pair (or two pairs) more than half length of the inner pair. Metapleural fan composed of long black bristles with a tendency to be brownish at tips. Pronotum with black bristles, sides of pro-

sternum and episterna with fine blackish or dark brown hairs. Humeri anteriorly and pronotum with a tendency to be brown. Wings clear and somewhat iridescent, veins brownish, a long brown costal stigma and a small black costal bristle. Radial fork more acute than is usual in related species. Anal vein complete. Squamae pale with blackish-brown fringes, halteres whitish-yellow. Legs long and slender, blackish to blackishbrown, finely grey dusted, subshining from some angles; tips of tarsi often curved. All femora slender, ventrally with silver microscopic pile. Fore femora

almost

bare, clothed

with minute

dark

hairs, a pair of ventral preapical hairs the longest. Mid femora with anterodorsal row of short adpressed bristles, beneath with about 6 pairs of strong blackish bristles on basal half, apical half practically bare. Hind femora with several thin bristly-hairs on basal half anteriorly and posteriorly, and with 2 to 4 similar anteroventral bristlyhairs at about middle, all about as long as femur is

deep or longer. Fore tibiae dorsally and posterodorsally with fine bristles, the hairs slightly longer than tibia is deep, densely short dark pubescent ventrally as is also mid tibia, the latter with about 4 strong black dorsal and ventral bristles about 3 times as long as tibia is deep, and another equally long anterodorsal preapical one. Hind tibiae slightly dilated towards tip (about as deep at tip as femur), ventrally with short adpressed bristles, dorsally with long bristles in pairs becoming longer towards tip, those on apical third longer than tibia is deep. Hind basitarsi distinctly narrower than tibia, dorsally with several bristles about as long as the segment is deep, ventrally strongly spinose, more distinctly so than on middle pair, and tarsal segments on all legs with distinct long black bristles in preapical circlets. Abdomen light greyish dusted, concolorous with thorax, covered with whitish hairs that are longer at sides of anterior segments, sterna with long yellowish straggling bristles. Genitalia very large and conspicuous on account of the peculiar, very long and wire-like aedeagus wound round the posterior half of abdomen. Lateral lamellae elongate, obliquely positioned or almost perpendicular, forming together with the prolonged 8th sternum a long ‘keel’, armed posteriorly with a row of long black bristles becoming brownish on sternum. Dorsal lamella in the form of a broad collar with short fine pubescence above. Lateral lamella posteriorly above with a leaf-shaped process bearing a brush of apically-curved black bristles at tip.

89

Figs 142-144. Male genitalia of Empis (s.str.) acinerea Chvala (SF, AB: Vichtis). — 142: postabdomen; 143: lamellae in lateral view; 144: dorsal lamella from the inner side. Scale: 0.2 mm.

Length: body 4.2-5 mm (without aedeagus), wing 4.5—5 mm. 2. Resembling male but thoracic bristles and abdominal pubescence shorter, and wings very brown and slightly broadened, posterior margin often paler but costal half very deep brown, veins blackish. Legs unusually long and slender, shaped very much as in male, even hind tibiae dilated towards tip, but legs practically bare. Femora without silver pile beneath and no ventral bristles, anterior four tibiae dorsally with short reclinate black bristly-hairs, but hind tibiae with almost the

same bristling as in male, though shorter. All basitarsi spinose beneath; basal three segments and also tips of tibiae with terminal circlets of small black bristles. Length: body 3.6—-5 mm, wing 4.2-5 mm. The male of E. acinereais very characteristic and cannot be mistaken for any other species on account of its very remarkable genitalia. The female has not been correctly recognised, and needs to be compared especially with the female of E. laminata; the latter is a smaller species (about 3-4 mm) with more slender and almost clear wings, legs shorter and more obviously with short bristles, and

90

hind femora somewhat flattened and almost twice as broad as hind tibiae. The female of E. albopilosa has almost clear wings, contrasting polished black abdomen, and yellowish metapleural fan. There are occasional aberrations in venation: I have seen 2 males with an incomplete radial fork on one side, and one male in which the radial fork was completely missing. Holotype identification. Empis cinerea was described by Zetterstedt (1855) from a single female, taken by Holmgren in June 1852 on the island of Oland ‘in floribus Potentillae and Glémminge’ (Sweden). There is one female, undoubtedly the holotype, in the Wallengren Collection in Lund labelled ‘Emp. cinerea Zett. n.sp.?. Oland.52. Holmgren’. This female is in poor condition, without antennae, wings or abdomen; fore leg complete on the left side, mid leg on the right, and both hind legs are present. It belongs to the species redescribed above. Distribution. Denmark and southern Sweden, in Finland approximately to 63° N, to Tb, Sb and Kb,

on dates ranging from 29 April to 24 June. Not rare; 39 d and 29 2 have been examined, but most

of them (31 ¢ and 20 @ ) are from Finland. — Clearly an overlooked species, previously known to me only from the Netherlands (first capture on 22 April) and central Europe.

tibiae. Legs subshining on anterior two pairs, finely grey pollinose, coxae dull grey and covered with yellowish bristly-hairs, but hind legs very shining blackish. Fore legs covered with short hairs, also

25. Empis s.str. syrovatkai Chvala, 1985

posterodorsal row of small black bristly-hairs much shorter than tibia is deep. Mid femora with a double row of long black bristles beneath, those in anterior row longest about middle (at least twice as long as femur is deep), those in hind row long even on apical half, dorsally with a row of erect bristles, at base about as long as femur is deep, gradually shortening towards tip. Mid tibiae with strong black bristles above and beneath at least twice as long as tibia is deep, those below (arranged in two rows) more numerous, anterodorsally with a slightly longer terminal bristle. Hind femora with fine bristles beneath, those on basal half posteroventrally the longest, distinctly brownish and longer than femur is deep, anterodorsally on apical half with about 6 very long black bristles. Hind tibiae with numerous long black bristles dorsally in two rows, intermixed with shorter hairs, the bristles becoming longer on apical half (longer than tibia is deep) and continuing also onto basitarsus; venter of tibia with short adpressed bristly-hairs. Only hind basitarsus with long bristles dorsally, anterior four basitarsi with short hairs above, though all distinctly spinose beneath, like the following three tarsal segments; terminal circlets formed by short stubby spines, mid basitarsus with a long anterodorsal terminal bristle as long as next tarsal segment. Abdomen subshining black, contrasting shining when compared with the dull thorax, dorsum finely brownish-grey pollinose and covered with sparse short pale hairs, sides with dense long whitish hairs, and less numerous shorter hairs on sterna. 8th sternum enlarged, with very long black hind-marginal bristles, and with a distinct ventral brush of spine-like black bristles anteriorly. Genitalia robust, lateral lamellae rectangular, covered with minute hairs, ventral and hind margins with longer black bristles; dorsal lamellae slender, mostly concealed, densely short black pubescent above. Aedeagus conspicuously stout, short, shining brown and clearly visible except for the broad-

the rather slender fore basitarsi, only tibiae with a

Figs 111, 145-148. Empis plumipes Zetterstedt, 1842:382 (not Meigen, 1804). Empis decora: Lundbeck, 1910:112.

Empis syrovatkai Chvala, 1985:389 (new name). Robust species (about 3.5—5 mm), halteres pale, metapleural fan and abdominal pubescence whitish, legs black, strong, with long bristles. Male genitalia with a short stout aedeagus, female legs strongly pennate, also fore tibiae and basitarsi above with short pennation. d. Eyes with upper ommatidia very enlarged, occiput dull grey, with two circlets of black forwardly-curved bristles above, otherwise covered with long white bristly-hairs. Antennae black, style about two-thirds length of 3rd segment; palpi black, with a few black bristly-hairs apically. Labrum 1.5 times as long as head is high. Thorax dull grey, lighter on pleura, mesonotum more brownish-grey with indefinite brownish stripes along the lines of bristles in anterior view, much greyer and with stripes becoming broader and blackish in posterior view. Large bristles black and in full number, usually 2 pairs (rarely 3) of scutellar bristles, outer pair at least half as long as inner pair, often longer. Acrostichal bristles 2-serial,

dorsocentrals irregularly 3-serial, numerous and extending towards sides at middle, all black and

not strong, about as long as antennal style, dorsocentrals becoming uniserial, long and bristle-like posteriorly. Humeri, posthumeral region and notopleural depression anteriorly with numerous fine whitish hairs. Sides of prosternum and episterna with white hairs, and even pronotum with a row of small, pale erect bristles. Metapleural fan whitish-yellow, long and dense. Wings clear with distinct dark veins, all reaching margin, costal stigma light brownish, and a very small black costal bristle. Axillary excision acute, discal cell truncate, but radial fork more acute than

is usual in Empis s.str., resembling that of E. lepidopus or E. acinerea..Squamae pale with long _ whitish fringes, halteres yellowish.

Legs rather long but stout, hind legs especially large (twice as long as mid pair) with tibiae evenly dilated and somewhat laterally compressed, broader than femora, basitarsi at most as deep as

ened tip concealed within lamellae; ventral lamella

polished black, small, upturned posteriorly. Length: body 3.5—5 mm, wing 3.64.2 mm. 2. Resembling male except for the shorter whitish abdominal pubescence, much shorter and obvi-

ously more bristle-like black acrostichal and dorsocentral bristles, slightly brownish wings, and the very characteristic strongly pennate legs. All coxae

of

and fore femora simple, latter at most with short ciliation above and minute hairs below, but fore

tibiae and basitarsi distinctly pennate above, and short pennate fringes also on 2nd tarsal segment. Mid femora broadly pennate on both sides, especially long at base above and ventrally before tip (though barely longer than femur is deep), mid tibiae and basitarsi short pennate above, and tibiae also at base beneath. Hind femora broadly pennate on both sides, the longest pennation on apical two-thirds beneath; hind tibiae with broad pennation (as long as tibia is deep) dorsally and on basal half beneath, with several (about four) long simple posterodorsal bristles, much longer than pennate bristles, and another anterodorsal bristle at tip. Hind basitarsi with a very long bristle about middle above and a shorter preapical pair, and all basitarsi long spinose beneath. Length: body 3.3-4.8 mm, wing 3.3-4.3 mm. A very characteristic, robust, whitish pubescent species with a very stout aedeagus in male and strongly pennate legs in female, which needs comparison in the Scandinavian fauna only with E. genualis, as has been done in the description of the latter. However, there is another closely related species in the warmer parts of Europe, E. gymno-

poda Bezzi, that was erroneously reported from Norway, a mistake that was repeated in the Palaearctic Catalogue (Chvala and Wagner 1989:265). The male of E. gymnopoda has grey dusted abdomen and legs, less strongly bristled mid tibiae, hind basitarsus with only short bristles above, aedeagus narrowed at tip, whilst the female also has mid coxae and fore femora strongly pennate beneath. Lectotype designation. Empis plumipes was described by Zetterstedt (1842) from Sweden and Norway, in Sweden from Skane (Esperéd), Ostergotland (Larketorp) and Jamtland (Areskutan, Hofverberget, Asarne at Sather prope Krokumselfven), in Norway from Akershus (Naes). There are 11 specimens under E. plumipes in the Zetterstedt Collection in Lund, 4 6 and3 2 from Larketorp, a male from Areskutan, a female from Hofverberget, and a pair from Naes, Norway; all are conspecific and represent the species described

above, except for the female from Naes which is E. pennipes. One male in perfect condition from Larketorp, labelled ‘E. plumipes. 6. ciliata z. Fall. LarKet.’, was labelled by me in 1977 and is herewith designated as lectotype of E. plumipes Zetterstedt.

Figs 145-148. Male genitalia of Empis (s.str.) syrovatkai Chvala (CZ, Bohemia, Maxov near Nachod). — 145: postabdomen; 146: dorsal lamella in anterodorsal view; lamellae in lateral view; 148: aedeagus. Scale: 0.2 mm.

BPs

Distribution. Widely distributed in Scandinavia but uncommon; only a little over 100 specimens (60 3d and 43 2) have been examined. Southern and central parts, approximately to 63-64° N, in Norway to SFi and On, in Sweden to Jmt., in Finland north to Tb, Sb, Kb and Om, also Russian Karelia. A typical summer species, on dates from 2 July to 22 September, but mainly in the second half of July and in August. — In the Netherlands from the beginning of July, unknown from Great Britain, but in the warmer areas of central Europe very exceptionally in spring; the earliest capture known to me is from a very warm region in southern Moravia (Lednice) from 26 May. A species of cool and temperate Europe, distributed south only to the southern parts of Germany (Bavaria), Austria and Slovakia.

26. Empis s.str. genualis Strobl, 1893 Figs 112, 149-152. Empis genualis Strobl, 1893:39.

long, slightly curved, femora with an anterior curve and a thickening in apical third, tibiae slightly curved posteriorly near base. Generally legs more shining and with longer bristles than in E. syrovatkai. Fore femora with a few longer black bristly-hairs posteroventrally towards tip, tibiae covered with coarser black posterodorsal bristles and longer posterior pubescence, which is still longer posteriorly on basitarsus, the numerous rather thin black bristly-hairs distinctly longer than basitarsus is deep. Mid femora with more numerous and stronger anterodorsal black bristles, becoming shorter on apical third only, the ventral long black bristles very prominent, strong, densely set at base, those in anterior row not becoming much shorter on apical half, but the posterior bristles on the contrary long on basal third of femur only, very short and fine towards tip. Mid tibiae with about 5 strong black anterodorsal bristles (including a terminal one) and more numerous, about 10, identical and equally long black bristles beneath. Hind femur with several strong black anterior (hardly anterodorsal) bristles, longest in basal third, posteriorly on basal half of femur with long, dense pale pubescence at least as long as femur is deep. Hind tibiae and basitarsi with numerous long black bristles above, distinctly longer than femur and basitarsus are deep, femora with short adpressed hairs ventrally, basitarsi long spinose, tips of tarsi characteristically curved. Abdomen extensively shining black, but dorsum finely coppery pollinose, pubescence pale, especially long at base of anterior three segments, sterna with shorter, sparse, entirely pale hairs. 8th sternum with black hind-marginal bristles, elongate, and forming a ‘keel’ with genitalia, anteriorly above with a distinct projection. Genitalia small, lateral lamellae long triangular, the long lower edge with strong black bristles, tip and upper mar-

Resembling E. syrovatkai. A robust, mostly pale pubescent species (3.5-5 mm), with pale halteres and rather shining black abdomen, legs in male stout and almost yellowish-brown on all ‘knees’ and posterior basitarsi, female legs strongly pennate also on posterior four coxae. 3. Head as in E. syrovatkai with similar enlarged upper ommatidia and mostly whitish bristles on occiput, but antennal style longer, not very much shorter than 3rd segment. Proboscis long, labrum much more than one-and-a-half times length of head. Thorax rather dull grey dusted, lighter grey on pleura, more brownish-grey on mesonotum, though when viewed from behind mesonotum greyer and with more outstanding blackish stripes, as in E. syrovatkai. The black marginal bristles including 2-serial acrostichal and irregularly 3-serial dorsocentral bristles almost exactly as in E. syro-

ing black and almost entirely concealed, densely short pubescent. Aedeagus light brown, short and

vatkai, with additional whitish hairs anteriorly on

rather stout, though much narrower than in E. sy-

mesonotum, pale hairs on prothorax, including a row of pale bristly-hairs on pronotum. Metapleural fan long, whitish-yellow. Wings distinctly darkened, light brownish

rovatkai, base protected by a small polished ventral lamella, tip concealed within lamellae or projecting above, slightly widening. Length: body 44.6 mm, wing 4.5—5 mm. 2. Very similar to that of E. syrovatkai but wings deeper dark brown and legs more shining, ‘knees’ and posterior basitarsi lighter brownish, and legs differently pennate; posterior four femora strongly pennate on both sides, also tibiae dorsally, hind

clouded, not at all hyaline, but venation including a

somewhat acute radial-fork, and all other characters, as in E. syrovatkai. Legs perhaps even stouter, at least tibiae, and more shining black to blackish-brown even on anterior two pairs, with ‘knees’ (the extreme tip of

femora and base of tibiae) and posterior four basitarsi more or less yellowish-brown. Hind legs very

gin with finer hairs; dorsal lamellae slender, shin-

tibiae on more than basal half beneath, and mid

tibia at extreme base only, but mid basitarsus with short adpressed pennation above. However, there

u2

is a distinct outer pennation also on posterior four coxae, fore femora have distinct flattened bristles above, as well as fore tibiae beneath towards tip, and fore tibiae and basal two tarsal segments are distinctly pennate above. Length: body 3.5—5 mm, wing 3.8-4.8 mm. Distribution. Known from the Netherlands, and its occurrence in the south of Scandinavia is therefore possible. Not rare in central Europe, but especially common in the Mediterranean, from Spain in the west through Corsica and Italy to Macedonia, Greece and as far as Transcaucasia (Azerbaijan). Described by Strobl (1893) from the Adriatic island Hvar (as Lesina) and, although I have not seen the types, I have seen several specimens identified by Strobl and believe that the species 1s correctly recognised. There is a series of 21 specimens in the Palaearctic Collection in Helsinki taken by R. Frey at Torino, Italy. In the colder parts of Europe from July to the beginning of September; a typical steppe species in temperate and warm parts of Europe, and a late species even in the Mediterranean, mainly in June. My records are from 31 May to 10 September.

27. Empis s.str. pennipes Linné, 1758 Figs 99-100, 103, 113, 153-156. Empis pennipes Linné, 1758:604. Empis longirostris Meigen, 1804:223.

Large (about 4-S mm), shining black, long-legged species with brown wings in both sexes and conspicuously long proboscis. Metapleural bristles and abdominal pubescence black, male genitalia large with long, evenly bowed, slender aedeagus, female legs strongly pennate. d. Occiput black with greyish dusting and densely set, forwardly curved black bristles becoming brownish below, face polished black. Antennae black, style two-thirds length of 3rd segment.

Palpi small, black, directed

forwards

and

stouter apically, clothed with several fine black bristly-hairs. Proboscis conspicuously long, labrum about 3 times as long as head is high. Thorax subshining black when viewed from above, rather densely brownish-grey dusted in anterior and posterior views, pleura grey dusted. All hairs and bristles black, large marginal bristles very long but not strong, in full number, including2 or 4scutellar bristles, outer pair (if present) always much smaller. Acrostichal and dorsocentral bristles long and hair-like, about as long as antennal

Figs 149-152. Male genitalia of Empis (s.str.) genualis Strobl (SLK, Zadiel, Hrhov).—149: postabdomen; 150: dorsal lamella in anterodorsal view; 151: lamellae in lateral view; 152: aedeagus. Scale: 0.2 mm.

94

style, both biserial, dorsocentral becoming longer and uniserial posteriorly, ending with a long prescutellar pair. Prothoracic pubescence black, rath-

er long but very fine on episterna and sides of prosternum; metapleural fan very long, black. Wings distinctly brownish with dark veins, a deeper brown costal stigma, and a small costal bristle. Anal vein complete. Squamae dirty grey with fine black fringes; halteres never clear yellow, usually dirty to brownish-yellow, or even extensively darkened. Legs subshining black to brownish-black, long and slender, only hind tibiae slightly dilated (as deep as hind femur) and hind basitarsus, although obviously dilated, not as stout as tibia at tip. Fore femora with only a row of small fine hairs beneath, fore tibiae with posterodorsal bristles shorter than tibia is deep, ventrally and posteriorly with dense, rather brownish, soft, short pubescence. Mid femora ventrally on basal half with two rows of very long bristles (the longest more than twice as long as femur is deep), becoming shorter and finer towards tip, anterodorsally near base with several

longer adpressed bristles. Mid tibiae with about 4 to 5 spine-like bristles above, slightly longer than tibia is deep, another similar bristle at tip, ventrally with more numerous but much shorter bristlyhairs. Hind femora with a row of distinct anterodorsal adpressed bristles, ventrally with only short fine bristles; hind tibiae with short bristly-hairs beneath, dorsal bristling hardly longer than tibia is deep, similar dorsal bristles also on basitarsus. Fore basitarsus with soft dense brownish pubescence, but posterior four basitarsi distinctly spinose beneath, and only very short bristles in terminal circlets on each tarsal segment. Tarsi distinctly curved, very characteristically rolled at tip. Abdomen subshining brownish-black, thinly brassy-brown dusted from some angles, pubescence black, short, longer at sides of terga and below, sterna

with

long, fine hind-marginal

hairs.

Genitalia large, characteristically ‘open’, with a long, free, evenly bowed, slender aedeagus. Ventral lamella very small, invisible, lateral lamellae triangular, narrowly pointed apically and armed with several rather long, strong black bristles; dor-

Figs 153-156. Male genitalia of Empis (s.str.) pennipes L. (CZ, Moravia, Lednice). — 153: postabdomen; 154: lamellae in lateral view; 155: dorsal lamella in anterodorsal view, lateral lamella dotted; 156: aedeagus with hypandrium. Scale: 0.2 mm.

95

sal lamellae simple, strip-like and with short hairs only. Length: body 3.6-5S.2 mm, wing 3.64.8 mm. 2. Bristles on thorax and abdominal pubescence shorter than in male, wings hardly broader but deeper brown, hind margin somewhat paler. Legs very conspicuously and strongly dark brown pennate; pennation on posterior four femora and hind tibiae on both sides at least as long as legs are broad, mid tibiae similarly broadly pennate above along whole length, ventrally with short pennation at base only, and fore tibiae and anterior four basitarsi above clothed with distinct flattened bristles. Length: body 3.8-5.2 mm, wing 4-S.2 mm. There are two closely related species in cooler temperate Europe with a similar, very long proboscis, E. procera Loew and E. nitidiventris Loew, but both have metapleural bristles and abdominal pubescence pale. In Scandinavia E. pennipes needs to be compared only with E. rufiventris: the main distinguishing features are given in the key and in the description of the latter. This is rather a variable species in the colour of halteres, and venation aberrations are not uncom-

mon: the radial fork is sometimes incomplete or missing on one wing, but I have seen a female in which the radial fork has been entirely lost on both wings, as in Rhamphomyia, without any trace of vein R,. There is a female which is undoubtedly conspecific in my collection (Bohemia, Budislav, 18.6.1977, leg.O. Syrovatka) with mid tibiae beneath, and fore tibiae and anterior four basitarsi above, with only simple

unmodified

setae, and

without any pennation or flattened bristles present.

Distribution. Rather a southern species in Scandinavia; common in the south, but individually approximately to 65° N, in Norway to Nsy, in Sweden only to Dlr., and in Finland to Tb, Sb and Kb, also in Russian Karelia. About 400 specimens have been examined (162 d and 235 2) and, as is usual in collections, females are more commonly represented (ratio d : 2 =2:3) because of the obvious attractiveness of their strikingly pennate legs. My records are from 11 May to 8 July, but mainly in June, though there are two unusual dates on Wiistnei material in the Copenhagen Museum from Sgnderborg (SJ, Denmark), namely 9 April 1904 and 27 July 1903; the April 1904 date in particular seems to be an error. — Collin (1961) recorded E. pennipes in Great Britain from mid May to mid July, and a very ‘remarkable’ capture in Perthshire on 21 August, apparently a very exceptional second generation. Very common in central Europe

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on dates ranging from 8 May to 14 July, mainly from mid May to mid June. A species of cool and temperate Europe, south only to northern France, Italy and Romania (Mtii Rodnei); absent in the Mediterranean. Biology. Very common in spring, especially on the pink flowers of Geranium where both males and females imbibe nectar. The epigamic behaviour was fully described by Chvala (1980), with details on swarms, swarming sites, mating and prey presentation. Males swarm in compact aggregations of 10 to 30 individuals in the shelter of trees, and the prey is caught fresh and intact and is much smaller than the predator (1.6-3.2 mm). Contact is initiated in the swarm, but the copulating pair immediately leaves the swarm and settles on nearby vegetation. The longest period of mating observed was 10 minutes. From many points-of-view, this is the basic pattern for mating behaviour in Empis s.str.

28. Empis s.str. rufiventris Meigen, 1838 Figs 114, 157-160. Empis rufiventris Meigen, 1838:86. Empis ventralis Zetterstedt, 1842:381. Large (4.5-6 mm), long-legged species with pale halteres and black metapleural bristles. Resembling E. pennipes but labrum shorter, abdominal pubescence pale, and legs in female less strongly pennate. 3. Occiput dull, rather densely grey dusted, covered with not very dense fine bristling, black above, light brownish below. Antennae often brownish on basal segments, style not very much shorter than 3rd segment. Only upper margin of clypeus polished black to brown; palpi black, covered with a few dark hair-like bristles, labrum light

brownish towards tip, more than twice as long as head is high. Thorax rather uniformly grey dusted, viewed from above mesonotum with three broad, subshin-

ing black stripes along the lines of bristles. Large marginal bristles black, long, in full number (a humeral, a posthumeral, 3 notopleurals, 1-2 supraalars, a postalar, 4 scutellars), acrostichal and dor-

socentral bristles mostly blackish, both irregularly biserial, rather short and hair-like anteriorly, but

gradually becoming longer towards prescutellar depression (including acrostichal bristles), last two pairs of prescutellar dorsocentrals as long as mar-

ginal bristles. Prothorax with blackish to dark brown bristly-hairs, the bristles of metapleural fan black, with tips distinctly brownish. Wings almost clear, very slightly brownish clouded, costal stigma long and deep brown, veins dark. A small black costal bristle. Squamae whitish with blackish fringes, halteres clear whitish-yellow. Legs subshining blackish-brown to almost brownish, very finely silvery-grey pollinose, long and slender, hind legs very elongate and hind tibiae and basitarsi slightly swollen. Femora with whitish pile beneath, especially on mid pair. Fore femora practically bare, fore tibiae dorsally with fine black bristly-hairs, no longer than tibia is deep, posteriorly with shorter, dense brownish pubescence. Mid femora ventrally on basal half with a double row of conspicuously long bristly-hairs, about 3 bristles in posterior row remarkably long, about 4 times as long as femur is deep, those in anterior row more numerous and shorter, and apical half of femur with minute hairs. Mid tibiae dorsally usually with 3, rarely 4, rather stout black bristles, but no apical bristle, ventrally with two rows of more numerous but shorter black bristles longer than tibia is deep. Hind femora ventrally with a

double row of very fine dark hair-like bristles, longer at about middle (slightly longer than femur is deep), with similar bristly-hairs posteriorly, and about 4 to 5 anterodorsal bristly-hairs on basal half the longest. Hind tibiae with long black bristling dorsally, becoming longer apically, all bristles longer than tibia is deep, ventrally with short adpressed bristles; long dorsal bristles also on hind basitarsus. Basal two tarsal segments on all pairs black spinose beneath, more strongly so on hind legs, bristles in terminal circlets not very long. Abdomen contrasting subshining in comparison with the dull thorax, more or less broadly brownish to brownish-yellow at base and on venter, blackish

towards tip. Pubescence light brownish to almost pale, longer at sides of basal segments, the longer hind-marginal hairs on terga blackish, but hindmarginal bristles on sterna very pale and more prominent. Genitalia black, widely open, in general structure very much as in EF. pennipes, but aedeagus evenly bowed, without the slight inward curve of E. pennipes, and lateral lamellae broader and blunt-tipped. Ventral lamella large, shielding base of aedeagus, lateral lamellae with long black bristles on posterior margin, but dorsal lamellae above

; Figs 157-160. Male genitalia of Empis (s.str.) rufiventris Meigen (NL, Elsboo Lbg., Hoge Bos). — 157: postabdomen; 158: lamellae in lateral view; 159: dorsal lamella in anterodorsal view, lateral lamella dotted; 160: aedeagus with hypandrium. Scale: 0.2 mm.

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with very short, dense, rather reddish-brown pubescence. Length: body 4.5—6 mm, wing 5.3—-5.8 mm. 2. The black bristling on thorax as in male, abdominal pubescence obviously shorter and paler, mostly yellowish. Wings more yellowish-brown clouded. Legs broader, especially posterior four femora and tibiae somewhat laterally flattened and the greyish dusting more distinct. Mid and hind femora on both sides, and hind tibiae above, broadly and rather brownish pennate, but the pennation scarcely longer than legs are deep, hind tibiae pennate on basal half only, unlike E. pennipes, fore and mid tibiae dorsally with flattened hairs, mid tibia ventrally at extreme base only, and anterior four basitarsi with simple bristles dorsally. All basitarsi (and next tarsal segment) distinctly black spinose beneath. Length: body 5—6 mm, wing 5.5-6.5 mm. E. rufiventris needs comparison mainly with E. pennipes, as has been done in the above description. However, it can be immediately recognised by its shorter proboscis, pale abdominal pubescence, and distinctly near-yellowish base of abdomen. Lectotype designation. Zetterstedt (1842) described Empis ventralis from Hogestad in Skane (Sweden) and from ‘Dania’. There are 3 ¢ and1 @ in the Zetterstedt Collection in Lund. All are conspecific and belong to the species described above, as correctly stated by Lundbeck (1910) and Collin (1961); 2 ¢ and 1 @ are labelled ‘E. ventralis a Staeg.’ and are undoubtedly the Danish specimens sent to Zetterstedt by Staeger, whilst the third male is labelled H6gestad’ and is herewith designated as lectotype of E. ventralis; it was labelled accordingly in 1992. The lectotype male is in good condition, only 3rd segment on both antennae and tarsi on both hind legs missing. Meigen’s type of E. rufiventris was studied by Collin (1961).

Distribution. A species of extreme southern distribution in Scandinavia (the classic distribution-pattern A, as shown in Fig. 1), known from Denmark and Swedish Skane. Altogether 120 specimens (50 3 and 70 @ ) have been examined, occurring in the

second half of May and in June, on dates ranging from 17 May to 4 July. However, there are two very problematic dates of capture from Denmark in Copenhagen University Museum: 2 ¢ labelled by Schlick as from 26 April 1870 (NEZ, Ermelund),

and 3 d and 4 @ allegedly collected by Wiistnei at Sgnderborg (EJ), on the magic date 9 April 1904 (as E. pennipes); this is apparently an error, as in temperate central Europe my records are from 14

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May to 24 June. — A species of northern and central Europe known, from England including Ireland (Chandler 1986), the Netherlands, and south only to Bavaria (Germany), Austria, Croatia and Macedonia. Collin (1961) recorded it from England as ‘locally common’ and ‘probably a woodland species’. In central Europe rather rare, though commoner, as stated by Collin, in woodland areas and preferably at higher altitudes in submontane regions.

29. Empis s.str. planetica Collin, 1927 Figs 115, 161. Empis lamellicornis: Lundbeck, 1910:116.

Empis planetica Collin, 1927:23 (as subsp. of E. lamellicornis Becker = E. simulium Nowicki). Rather small (about 3 mm), dull, dark greyish species with pale halteres, pale abdominal pubescence, and 2 scutellar bristles. Male with a small pointed tubercle at sides of 8th tergum and aedeagus with a loop at middle; female legs broadly pennate, also fore tibiae with flattened hairs. 3. Eyes with upper ommatidia very enlarged, occiput dark grey, subshining from some angles, covered with black bristles above that become finer and paler below. A pair of very long ocellar bristles. Antennae black, 3rd segment distinctly constricted on more than apical half, style not very much shorter. Labrum only slightly longer than head is high. Thorax dull dark grey, or very slightly subshining in some lights, practically unstriped. Marginal bristles black, rather long but fine, 2 scutellar bristles. Acrostichal bristles very narrowly biserial, black, fine and hair-like, dorsocentrals slightly longer and diverging, biserial, posteriorly uniserial and ending in 2 pairs of long prescutellar bristles, prescutellar depression often lighter grey and more densely dusted. Prothorax (sides of prosternum and episterna) with minute pale hairs, pronotum with short black bristly-hairs. Metapleural fan composed of a few black bristly-hairs above and some finer pale hairs below. Wings quite clear, somewhat iridescent, veins blackish but those beyond R,,, finer and less distinct. Radial fork not very ‘open’, rather acute, but vein R, straight. A small black costal bristle, alular fringes whitish, anal vein complete. Squamae pale with concolorous fringes; halteres clear yellow, base of stalk dark brown. Legs rather slender, subshining blackish-brown with a fine covering of grey pollen; mid legs very

short, hind pair elongate, with tibiae dilated to-

Fig. 161. Male genitalia of Empis (s.str.) planetica Collin (N, AK; Baerum, Ost@ya). Scale: 0.2 mm.

wards tips, as deep at tip as femur, basitarsus equally stout, and even the next two tarsal segments slightly dilated. Fore legs with short fine hairs, only tibiae posterodorsally with scattered black bristly-hairs about as long about middle as tibia is deep, ventrally with dense, soft brownish pubescence; tarsal segments with very long black bristles in preapical circlets, very much as in E. praevia. Mid femora with an anteroventral row of about 6 long black bristles, more than twice as long as femur is deep, posteroventrally before middle with two even longer black bristles and much shorter hairs towards tip. Mid tibiae with a rather smaller anterodorsal bristle in basal quarter, another very long bristle in basal third (at least 3 times as long as tibia is deep) and a further perhaps even longer preapical seta (as long as basitarsus); ventrally with short adpressed bristling. Mid basitarsus with a long dorsal bristle in the apical circlet, the next tarsal segments with only short terminal circlets of bristly-hairs. Hind femora on apical half with 3-5 long adpressed anterodorsal bristles, longer than femur is deep, and a row of outstanding thinner anteroventral bristles becoming longer towards tip. Hind tibiae dorsally with bristly-hairs slightly longer than tibia is deep, similar long bristly-hairs also on the stout basitarsus, ventrally with short adpressed bristles. Hind tarsal segments with short terminal circlets. Abdomen rather densely grey pollinose, though somewhat subshining when viewed from the side, pubescence pale, dense, longer at sides of basal segments, venter with long whitish straggling bristles. 8th tergum with a small pointed tubercle at each side and 8th sternum with 5—6 hind-marginal bristles about as long as sternum is wide. Genitalia open, of the E. pennipes-type, with a long, slender,

brown, undulating aedeagus, with two bends anda recurved tip. Lateral lamellae triangular, pointing upwards, hind margin with a few long black bristles; dorsal lamellae very small, strip-shaped, projecting beyond or above apex of lateral lamellae. Length: body 2.8-3.2 mm, wing 3.3-3.5 mm. 2. Very much like male but thoracic bristling and abdominal pubescence shorter, and wings distinctly brownish, deeper brown towards costal margin, all veins distinct. Legs shorter than in male and somewhat laterally flattened, especially on hind pair, rather strongly pennate. Fore femora bare, but fore tibiae and basitarsi posterodorsally with flattened bristles (less distinct on basitarsus), mid femora with short pennation (not longer than femur is deep) dorsally and on apical half ventrally, mid tibiae very short pennate above and ciliated below, hind femora broadly pennate above along whole length and ventrally on apical two-thirds, hind tibiae with similar but slightly shorter pennation dorsally and on basal half ventrally. — Length: body 3-3.5 mm, wing 3.2-3.8 mm. E. planetica belongs to a very difficult group of several very closely related species; in addition to E. staegeri, with which it is compared below, and other undescribed species, there are two species in temperate Europe, EF. protarsalis Collin and E. simulium Nowicki (= E. lamellicornis Becker), which differ above all by the structure of male genitalia and the bristling of legs. The female of E. planeatica is very characteristic, with a row of distinct flattened bristles on fore tibiae above.

Distribution. A rare species in the southern parts of Scandinavia; I have examined only 14 3 and 13 ? from Denmark, Sweden (Skane) and Norway (AK, Baerum: Ostgya). Lundbeck (1910) recorded

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Fig. 162. Male genitalia of Empis (s.str.) staegeri Collin (N, AK: Baerum, Ostgya). Scale: 0.2 mm.

it from Denmark as E. lamellicornis Becker, and Jonassen (letter of21Dec. 1992) reported it in Norway from the counties Ostfold, Buskerud (¢) and far to the north in Nord-Trgndelag (1): the latter is an extraordinarily northern record. A typical spring species, on dates from 15 May to 23 June. — Rather a northern species in Europe, known from England, Ireland, the Netherlands and, in addition to Scandinavia, only from Germany.

30. Empis s.str. staegeri Collin, 1963 Figs 116, 162.

arated supra-alars, a postalar, and 2 long scutellar bristles. Acrostichal bristles scarcely shorter than dorsocentrals, about as long as antennal style and hair-like; former narrowly biserial, latter biserial about middle, uniserial anteriorly and posteriorly, last 3 pairs very long. Prothorax with whitish hairs, but pronotum with a row of minute black bristles. Metapleural fan pale, whitish-yellow, only rarely with additional blackish bristles. Wing venation as in E. planetica but wings rather whitish, only anterior veins (costa and radial veins) brownish, veins beyond R,,, whitish and very inconspicuous. Squamae pale with concolorous fringes,

Empis staegeri Collin, 1963:171.

Small (2.6-3.5 mm) dull grey species resembling E. planetica, with pale halteres, pale metapleural bristles and abdominal pubescence, and with 2 scu-

tellar bristles. Wings more whitish in male with indistinct medial veins, evenly bowed aedeagus without loops, and abdomen without tubercles on 8th tergum. Female legs shorter pennate and anterior tibiae with ordinary bristles above. 3d. Occiput dull grey, pubescence below neck wholly whitish. Antennae black, 3rd segment distinctly constricted at least on apical half or more, style slender, only slightly shorter than 3rd segment. Labrum shining black, not very much longer than head is high. Thorax with similar dusting and bristling asin E. planetica, mesonotum uniformly dark grey, with all bristles black. Marginal bristles long and rather fine, a humeral, 2 smaller intrahumerals, a long posthumeral, usually 2 notopleurals, 2 widely sep-

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halteres

whitish-yellow,

base

of stalk

brown. Legs very much as in E. planetica, differing only in details. Fore legs with similar posterodorsal bristly-hairs on tibiae and brownish soft pubescence beneath, terminal circlets on tarsal segments also long. Mid femora with short adpressed hairs dorsally and anteroventral bristles obviously shorter, but those posteroventrally almost as long as in E. planetica. Mid tibiae with the upper anterodorsal bristle slightly longer, not very much shorter than the other two bristles in basal third and at tip,

and with several (4-5) much longer anteroventral bristles at about middle. Hind femora with shorter bristles, dorsally with short adpressed hairs and anteroventrally with shorter hair-like bristles, shorter than femur is deep. Tibiae and hind basitarsus as in E. planetica. Abdomen rather densely light grey dusted, covered with dense whitish pubescence, longer at sides of anterior segments, venter with long, whit-

ish straggling bristles. 8th tergum simple, without

tubercles

with a few,

Slovakia (the former Czechoslovakia), where it is

about 3 to 4, very long pale marginal bristles, sternum produced and bilobed anteriorly above. Genitalia very different in detail from those of E. planetica, dorsal lamellae broadened at base and rather narrowly triangular (not strip-like), and hypandrium (ventral lamella) very long, slender, V-shaped in lateral view, distinctly shielding the base of the evenly bowed slender aedeagus with only extreme tip recurved (in E. planetica, aedeagus thickened basally and with a loop at about middle, near the base of its slender apical part). Length: body 2.6-3.5 mm, wing 3.2-3.8 mm. 2. Resembling EF. planetica but wings only slightly clouded, almost clear, medial veins fine, and metapleural bristles mostly pale. Legs less pennate, fore and mid tibiae without flattened hairs, fore tibiae above at most finely ciliated, mid femora with short anterodorsal pennation and only a few pennate bristles ventrally towards tip,

at sides, and 8th sternum

rather rare in the second half of May and at the beginning of June. One male was captured by chance in a male swarm of E. nigripes.

hind femora with similar but shorter pennation,

and hind tibiae above and on basal third ventrally with uniformly short pennation, shorter than tibia is deep. Length: body 2.8-3.5 mm, wing 3-3.5 mm. So far as the Scandinavian fauna is concerned,

E. staegeri needs to be compared only with E. planetica, as has been done in the description above. Both species are very characteristic, having a twobristled scutellum. E. staegeri is generally misidentified in Scandinavian collections, most often as E. lamellicornis (a synonym of E. simulium) or as E. planetica. Holotype identification. Described by Collin (1963) from a single male from the Staeger Collection in Copenhagen. I have seen the holotype male in Copenhagen, labelled ‘coll. Staeger’, standing over the name Empis staegeri Collin in the Lundbeck Collection, undoubtedly the holotype and so labelled by me in 1989. There is a female taken by Lundbeck on 18 May 1910 at Ermelund (NEZ), labelled ‘Empis sp. not lamellicornis and not planetica, det. Collin, 1963’ in Collin’s handwriting, un-

doubtedly a female of E. staegeri. Distribution. Southern parts of Scandinavia, approximately to 60° N, in Norway in a small area in

the south-east (VE, @, AK, Bg, HEs), in Sweden north to Upl., Vstm. and Dlr., and as an isolated

occurrence in Finnish Alandia (27 d and 11 2). A typical spring species; my records in Scandinavia _

arefrom15 May tol July; not uncommon, and alto-

gether 86 d and 53 9 have been examined. — A poorly known species, and apart from the above specimens known to me only from Bohemia and

31. Empis s.str. limata Collin, 1927

Fig. 163.

Empis limata, Collin, 1927:22. Rather smaller (about 3.5 mm), with pale halteres, pale abdominal pubescence, but metapleural bristles black, thorax subshining and abdomen strikingly polished black. Wings clear in male, brownish in female. 3d. Occiput dark grey, covered with black bristlyhairs becoming pale below. Antennae black, style shorter than 3rd segment, about two-thirds of its length. Labrum about 1.5 times as long as head is high. Thorax subshining black, only slightly greyish pollinose, and indistinctly striped from some angles. All thoracic bristles black, including not very long biserial acrostichal bristles, dorsocentrals slightly longer, biserial about middle, uniserial anteriorly and posteriorly and becoming longer behind, ending in a long prescutellar pair. Marginal bristles long, a humeral, a smaller intrahumeral, a

posthumeral, 3 notopleurals, a supra-alar, a postalar, and one pair of long black scutellar bristles with a small fine hair on each side. Pronotum with small black bristly-hairs, sides of prosternum and episterna with several pale to brownish hairs, metapleural fan black. Wings almost clear with all veins distinct, a small black costal bristle, and anal vein complete. Squamae greyish with pale fringes, halteres with knobs yellowish. Legs rather shining black, slender, but hind tibiae towards tip and hind basitarsus dilated. Fore femora with short hairs, fore tibiae with distinct posterodorsal bristly-hairs almost twice as long as tibia is deep, posteriorly and ventrally with dense rather short pubescence, the hairs also distinct on the slender fore basitarsus, tarsal circlets small. Mid femora ventrally with two rows of long black bristly-hairs about 3 times as long as femur is deep, those in posterior row about middle the longest, anterodorsally at base with much shorter, but distinct bristly-hairs. Mid tibiae with 2—3 very long anterodorsal bristles on basal half and another even longer one at tip, ventrally with a double row of densely-set bristles longer than tibia is deep, basitarsus with a long anterodorsal bristle at tip. Hind femora ventrally with rather short fine bristly-

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The above short description was based on my notes made in 1992 from the collection of the London Natural History Museum. E. limata is very characteristic on account of its polished black abdomen and subshining thorax, combined with the very distinctive structure of genitalia in male and the pennate legs in female. E. planetica and E. staegeri have the body dulled by greyish dust, a pale metapleural fan, two scutellar bristles, very different male genitalia and differently pennate legs in females, whereas E. melanotricha and its al-

lies (E. tanysphyra, E. woodi) have the abdomen black pubescent. Fig. 163. Male genitalia of Empis (s.str.) limata Collin (from Collin, 1961).

hairs, longer hairs anteriorly and posteriorly towards base, and a row of rather shorter anterodorsal adpressed bristles. Hind tibiae densely long bristled above, beneath with short adpressed brist-

ly-hairs. Hind basitarsus with bristles above similar to those on tibia, black spinose ventrally. Abdomen distinctly shining black both on dorsum and on venter, pubescence pale at sides of basal segments, dorsum rather short black pubescent, sterna with outstanding pale bristly-hairs on hind margins, 8th sternum posteriorly with black bristles. Genitalia small but very distinctive, lateral lamellae triangular in shape, covered with short hairs only, dorsal lamellae with a small upper process bearing at tip a comb of small black spinose bristles, lower part produced beyond lateral lamellae. Aedeagus clearly visible beneath, and with a distinct outward loop before being covered by lateral lamellae, very slender apically. ?. Wings brownish, distinctly darker than in male, mesonotum subshining black, very finely grey pollinose, more densely so when viewed from the side. Abdomen polished black as in male, but apical three narrowed segments and cerci dull dark grey. Posterior two pairs of legs distinctly pennate, fore femora with short hairs but fore tibiae distinctly fringed dorsally. Mid femora rather broadly pennate above (as long as femur is deep) and on apical half beneath, mid tibiae above with shorter pennation, and pennate bristles present also on basal third beneath. Hind femora with dorsal pennation shorter than femur is deep, ventrally longer pennate (at least as long as femur is deep) on the whole length. Hind tibiae broadly pennate along whole length above and on basal half beneath, those bristles above and at base beneath as long as tibia is deep. Length: body about 3.5 mm according to Collin (1961).

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Distribution. E. limata is still known only from a few specimens taken in England over a long period from June to mid-August. It remains unknown outside Great Britain, where it could well be an endemic species, but, as its occurrence on the continent cannot be excluded and it may well even occur in southern Scandinavia, it is included here.

32. Empis s.str. melanotricha Loew, 1873

Figs 117, 164-167.

Empis melanotricha Loew, 1873:39. Rather smaller (about 3.5 mm) dark grey dusted species with pale halteres, and all hairs and bristles including abdominal pubescence blackish. Male genitalia small, aedeagus clearly visible, slender and evenly bowed; female legs covered with flattened bristles on mid and hind pair, without pennation. 3. Eyes with upper ommatidia very conspicuously enlarged and sharply delimited above. Occiput rather subshining, finely dark grey dusted, occipital bristly-hairs rather dense and long, black and forwardly-curved above, finer and whitish below. Antennae black, style short, not very much more than half length of 3rd segment; palpi small, black, with a few minute dark hairs. Proboscis long, almost twice as long as head is high. Thorax rather densely grey dusted, mesonotum more brownish-grey in anterior view and with indistinct stripes, lighter grey when viewed from behind and with the stripes blacker. Large marginal bristles rather thin; a long humeral, a smaller intrahumeral, one strong posthumeral, notopleural depression bare, only posteriorly with two long notopleural bristles, the lower one coarser, a supraalar, a postalar, and 2 pairs of scutellar bristles, outer pair very small. Acrostichal and dorsocentral bristles fine and hair-like, shorter than antennal

style, former narrowly biserial, latter somewhat

longer and biserial at about middle, anteriorly and posteriorly uniserial, longer posteriorly and ending in 2 strong prescutellar pairs. Prothoracic pubescence and metapleural fan black. Wings clear with distinct dark veins, though medial veins fine, radial fork not at all right-an-

gled, and vein R, somewhat oblique; a short black hair-like costal bristle, anal vein complete. Squamae whitish with black fringes, halteres yellowish.

Legs blackish, not very subshining, rather densely greyish pollinose, conspicuously slender and especially mid pair short, hind legs distinctly elongated, twice as long as mid pair (both femora and tibiae), hind tibiae and basitarsi not swollen, as deep as hind femora. Tips of all tarsi usually very curved. Fore femora with short adpressed hairs above and almost bare beneath, fore tibiae with posterodorsal bristly-hairs nearly as long as tibia is deep, otherwise with short hairs, also on basitarsi,

terminal circlets inconspicuous, only short terminal bristles. Bristling on posterior two pairs very much as in E. planetica or E. staegeri, mid femur

with long bristles beneath, those posteroventrally before middle the longest, mid tibia with usually two (rarely three) very long anterodorsal bristles in basal third and at middle, and another still long-

er bristle at tip, the similar terminal bristle on basitarsus much shorter, and tibia ventrally with dense bristling, at about middle slightly longer than tibia is deep and shortening towards both tips. Hind tibiae andbasitarsi with sparse bristles above, those on tibiae longer than tibia is deep, basitarsi spinose beneath. Abdomen rather brownish-grey dusted when viewed from above, somewhat subshining in lateral view, pubescence entirely black, longer and denser at sides of basal terga, sterna with long black straggling hairs, and 8th sternum with a few long black hind-marginal bristles. Genitalia of the E. pennipes-type but small, lateral lamellae triangular in shape, lower posterior margin with longer black bristles, otherwise with short hairs; dorsal lamellae elongate, pointed apically beyond lateral lamellae, dorsum at base densely short black pu-

Figs 164-167. Male genitalia of Empis (s.str.) melanotricha Loew (CZ, Bohemia, Davle).—164: postabdomen; 165: lamellae in lateral view; 166: dorsal lamella in anterodorsal view, lateral lamella dotted; 167: aedeagus with hypandrium. Scale: 0.2 mm.

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bescent. Aedeagus slender and evenly bowed, more or less shielded on basal third by a large, shining blackish brown V-shaped hypandrium, very slender towards the free, slightly recurved tip. Length: body 2.6-3.3 mm, wing 3.5-3.8 mm. Syrovatka (1991) gives considerably larger measurements; see discussion below. 2. Acrostichal and dorsocentral bristles shorter than in male, abdomen more subshining even in dorsal view, and apical two segments (including cerci) elongate and densely grey dusted; pubescence short, rather dense, black. Wings much darker than in male, very brownish. Fore legs as in male, but posterodorsal bristles on tibiae slightly stouter. Mid femora anterodorsally with a row of not very long reclinate flattened bristles, beneath before tip with a few longer flattened bristles, mid tibiae with short ciliation above. Hind femora above with two rows (dorsal and anterodorsal) of reclinate flattened bristles, with similarly flattened but longer erect bristles beneath on apical half. Hind tibiae longer ciliated above (no flattened bristles), with 2 or 3 slightly longer bristles arranged dorsally in diverging pairs. Females longer than males because of the elongated two apical abdominal segments. Length: body 3.2-4.0 mm, wing 3.5-4.0 mm. In view of the black abdominal pubescence, EF. melanotricha needs comparison particularly with E. tanysphyra and E. woodi, as it has been done in the key. Females of the latter two species differ by the distinctly pennate legs. E. melanotricha was described from the southern slopes of the Romanian Carpathians (Baile Herculane), and a lectotype was designated by Syrovatka (1991). I have also studied the syntypic series in the Loew Collection in Berlin, as well as numerous additional specimens from central and north-eastern Europe, but all the specimens that I have seen were smaller than the measurements given by Syrovatka (1991): he has given body length for male 3.7-4.1 mm, and 3.8-4.3 mm for female, length of wing 4.54.6 mm and 4.3-4.6 mm respectively, almost 1 mm more than my measurements. Moreover, he described the female abdomen as ‘more densely grey dusted than in male’, just the opposite of my own description. The female of E. melanotricha also needs to be compared with females of E. gooti, a characteristic species of the E. nigripes-complex with abdominal pubescence practically all black, and thus very different from other pale pubescent females of that group. E. gooti is a greyer species, with longer antennal segments, legs shorter and more flattened, especially the hind pair, hind femora distinctly pennate above and on apical half beneath, and hind tibiae

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with several much longer single anterodorsal bristles well differentiated from the row of much shorter flattened bristles. In E. tanysphyra and E. woodi the hind tibiae are distinctly pennate above. However, there is another species very closely related to E. melanotricha in temperate Europe, E. ciliatopennata Strobl, which may well occur in the colder parts of Europe and in Scandinavia; it is a typical summer species occurring mainly in July and August, replacing E. melanotricha in the same biotopes later in the summer. There are slight differences in the male genitalia in E. ciliatopennata, longer hairing on femora, and the notopleural depression is distinctly pubescent anteriorly, whilst the female has longer and broader flattened bristles on hind femora. Distribution. North-western, central and southeastern Europe, from the Netherlands in the north southwards to northern Serbia and Romania; its occurrence in southern Scandinavia is very possible. A typical spring species; my records are from 15 May to 22 June.

33. Empis s.str. tanysphyra Loew, 1873 Figs 118, 168-171.

Empis tanysphyra Loew, 1873:38. Medium-sized (about 44.5 mm), grey dusted species with pale halteres and all hairs and bristles on body and legs black. Resembling a larger E. melanotricha, but aedeagus in male with a bend at about middle, and female legs pennate on posterior two pairs. 3d. Occiput dark grey dusted, occipital bristles blackish, at most dark brownish just above mouthopening. Antennae black, style about two-thirds length of 3rd segment; palpi with one or two long terminal bristly-hairs as long as palpus. Labrum black, light brown at tip, twice as long as head is high. Thorax densely rather light grey dusted, even on mesonotum, with three narrow brownish stripes along the lines of the bristles when viewed from in front, becoming broader and blacker in posterior view. All bristles black, a long humeral, a scarcely shorter intrahumeral, a posthumeral, 3-4 notopleurals (with 1 or 2 small hairs anteriorly), a supra-alar, a postalar, and 2 long scutellar bristles, rarely with a small hair each side. Acrostichal and dorsocentral bristles biserial, rather short and hairlike anteriorly, as long as antennal style, becoming

Figs 168-171. Male genitalia of Empis (s.str.) tanysphyra Loew (NL, Apeldoorn, Echoput). — 168: postabdomen; 169: lamellae in lateral view; 170: dorsal lamella in anterodorsal view, lateral lamella dotted; 171: aedeagus with hypandrium. Scale: 0.2 mm.

longer posteriorly, dorsocentrals ending in two long presutellar pairs. Prothoracic hairs and metapleural fan black. Wings very faintly brownish, not clear, veins brown, distinct, the broad apex of cell R, deep brown (a large costal stigma), radial fork with a tendency to be acute, and a small black costal bristle. Anal vein complete. Squamae pale with long black fringes, halteres yellowish. Legs rather subshining black, but similar in shape and bristling to E. melanotricha, including the very long hind legs and curved tips of tarsi. Fore tibiae with similar posterodorsal bristles, néarly as long as tibia is deep about middle, but bristles of tarsal terminal circlets longer, those on

2nd and 3rd segments as long as next tarsal segment. Mid femora with conspicuously long bristles beneath on basal half as in other related species,

those posteriorly before middle the longest, dorsum with short adpressed hairs (longer anterodorsal bristles in E. melanotricha). Mid tibiae with two, but often with three, very long anterodorsal bristles, the third in apical two-thirds, the fourth,

the terminal, the longest. Mid tibia with rather long bristles ventrally, 3 to 4 anteroventral bristles about middle twice as long as tibia is deep. Hind femora with only short reclinate bristles above (no longer anterodorsal bristles!), ventrally with fine hairs, those about middle the longest but still much shorter than femur is deep. Hind tibiae with irregular and unequal bristly-hairs above, becoming longer towards tip and only slightly longer there than tibia is deep, ventrally with short hairs, bas-

itarsus also with shorter rather reclinate bristles above and not very long spine-like bristles beneath. Abdomen more densely rather light grey dusted on dorsum, sides less pollinose, pubescence uniformly black, longer and denser at sides of basal segments, sterna with long straggling hair-like bristles and 8th sternum with long black hind-marginal bristles. Genitalia rather small, both lamellae cov-

ered with short fine black hairs; lateral lamellae broadly triangular, dorsal lamellae slender, mostly concealed and not overlapping side lamellae posteriorly. Aedeagus slender, brown, its base shield-

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ed by enlarged ventral lamella, otherwise aedeagus free, at middle with an outward loop, tip pointing backwards above dorsum of abdomen, not recurved. Length: body 3.5—4.8 mm, wing 3.9-4.6 mm; the wing length of 5.0-5.4mm given by Syrovatka (1991) seems to be a lapsus. 2. Mesonotum obviously more brownish-grey and all bristles shorter, black abdominal pubescence short, and wings conspicuously darkened, very brown, with well-differentiated deeper brown costal stigma. Legs very characteristically bristled: dorsum of fore and mid tibiae, and fore basitarsus, with short reclinate flattened bristles; mid femora rather broadly pennate dorsally along whole length and on apical half beneath (pennation scarcely as long as femur is deep), similar but stronger pennation on hind femora, the pennate bristles about middle above and before tip below slightly longer than femur is deep; hind tibiae short pennate above (shorter than tibia is deep) and with flattened bristles beneath at least on basal third. Length: body 3.6-4.8 mm, wing 4.0-4.6 mm. E. tanysphyra needs to be compared particularly with E. woodi, as has been done in the key and in the description of the latter. E. melanotricha, the third northern European species with pale halteres, dull grey body and black abdominal pubescence, is generally smaller and the differences have been discussed in its description above. EF. tanysphyra was described from the Romanian Carpathians, and a lectotype was designated by Syrovatka (1991). Distribution. Unknown from Scandinavia and Great Britain, but not uncommon in the Netherlands and possibly more widely distributed in the cooler parts of Europe. Almost throughout Europe, including the whole Mediterranean, from Spain in the west through Corsica eastwards to Greece (Corfu). Rather uncommon in the temperate central parts of Europe, on dates from 8 May to 4 June; an early spring species.

34. Empis s.str. woodi Collin, 1927

Figs 119, 172-175.

Empis woodi Collin, 1927:21.

3d. Antennae with style shorter, not very much more than half length of 3rd segment, and palpi with coarser black bristles at tip. Proboscis shorter than in E. tanysphyra, less than 1.5 times as long as head is high. Thorax obviously greyer in anterior view and practically unstriped, mesonotal bristling very similar but acrostichal and dorsocentral bristles slightly longer though thin and hair-like. Metapleural fan long, black. Wings slightly obscured, somewhat light brownish, not quite clear, veins dark brown and a deep brown costal stigma. Squamae dirty grey with black fringes, halteres orange-yellow. Legs similar in shape, long and rather slender, but hind tibiae more evenly dilated, at least as deep throughout as hind femur, basitarsus slightly narrower than tibia at tip. Fore tibiae more densely short bristled above, posterodorsal bristles more numerous (about 10 in a row) and stouter, longer than tibia is deep, continuing (though shorter) also on basitarsus; tibia with very dense, short, greyish pubescence posteriorly and beneath. Mid femora with longer anterodorsal (or nearly anterior) adpressed bristles on basal half (as in E. melanotricha), and the unusual long bristles beneath still long beyond middle of femur. Mid tibiae with 2 or 3 long anterodorsal bristles similar to those in E. tanysphyra, but the ventral bristling decidedly longer and stouter, 2 or 3 bristles beneath about middle more than twice as long as tibia is deep. Hind femora (unlike EF. tanysphyra) with long black bristles anterodorsally or nearly anteriorly, the bristles almost as long as femur is deep, and the fine ventral hairs also distinctly longer, again at least as long as femur is deep at about middle. Hind tibiae with long black bristles above (longer than tibia is deep), with similar but more adpressed dorsal bristles also on basitarsus, ventrally with black spines present also on the next three tarsal segments. Abdomen brownish-grey dusted, slightly subshining from some angles, but almost velvetyblack in posterior view, rather densely long black pubescent at sides of anterior segments, venter with fine, black straggling bristles. Genitalia small, lateral lamellae broadly triangular in shape, with longer black bristly-hairs on lower margin towards base; dorsal lamellae

Resembling FE. tanysphyra, but legs with longer and denser bristling in male, and aedeagus shorter and stouter, tip broadened. Female wings lighter brown and anterior four tibiae and basitarsi with only ordinary hairs above, without flattened bristles, posterior femora and hind tibiae pennate.

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covered

with short hairs,

slender and curved in dorsal view, not overlapping lateral lamellae posteriorly. Aedeagus shorter and stouter than in allied species, yellowish-brown, with a slight bend and an inward and outward thickening at about middle, tip broadened. Length: body 4.2-4.6 mm, wing 4.3-4.8 mm.

Figs 172-175. Male genitalia of Empis (s.str.) woodi Collin (NL, St. Pietersberg Lbg.).-—172: postabdomen; 173: lamellae in lateral view; 174: dorsal lamella in anterodorsal view, lateral lamella dotted; 175: aedeagus with hypandrium. Scale: 0.2 mm.

2. Very closely resembling female of E. tanysphyra, but wings lighter, hind margin in particular broadly paler, and legs with different bristles: anterior four tibiae and basitarsi above with only ordinary bristly-hairs, at most mid tibiae somewhat ciliated, and beneath near base sometimes with a few flattened hairs. Mid femora and hind legs more strongly pennate, mid femora with at least flat-

Pietersberg Lbg., 4 May 1988) with only vestiges of a radial fork on both wings. A full description was published by Collin (1961). Ihave studied syntypes from Herefordshire (a lectotype was not designated) in the Natural History Museum, London, and found them to be conspecific with NW and central European specimens. ji

tened bristles almost to base, and hind tibiae more

Distribution. A species with a similar pattern of distribution to that of E. tanysphyra; in the northwest known from England and the Netherlands, and through the central parts of Europe south to the Mediterranean; Collin (1961) recorded it from Sardinia. Recently it has been found locally common in the Netherlands (St. Pietersberg Lbg., Berg en Dal Gld.) and its occurrence in southern Scandinavia seems very likely. Collin (1961) gives the flight period in England as May and early June, and I have records of it in central Europe from 4 to 27 May; a typical early spring species.

uniformly short pennate also on basal half beneath. Length: body 4-5 mm, wing 44.5 mm. ' The leg pennation in females seems to be somewhat variable, and I have seen specimens with rather broadened bristles dorsally on fore basitarsus and on mid tibiae towards base, but fore tibiae

(unlike E. tanysphyra) always without flattened bristles. Some variation can also be observed on hind tibiae, where the dorsum is sometimes covered with long flattened bristles rather than a true pennation. I have also seen a female taken by B. Van Aartsen in a Malaise trap in the Netherlands (St.

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35. Empis s.str. nigripes Fabricius, 1794 Figs 176-180. Empis nigripes Fabricius, 1794:405. Empis pennaria Fallén, 1816:20. Empis vernalis Meigen, 1822:27. Empis flaviventris Strobl, 1898:410 (as var. of E. pennaria Fallén).

Medium-sized (about 3-4 mm) greyish dusted species with pale halteres, mostly pale pubescent abdomen and 4 scutellar bristles. 8th tergum in male with rounded tubercles at sides, aedeagus short and stout, tip widened. Female with short pennation on posterior four femora and tibiae. ¢o. Head dull grey on vertex and occiput, with rather long, black, forwardly-curved postocular bristles above, behind them two rows ofshorter occipital hairs, below with longer pale hairs. A pair of long black ocellar bristles as long as postocular ciliation. Antennae black, style shorter than 3rd segment, about two-thirds of its length. Palpi black, covered with minute dark hairs and 1 or 2 longer apical ones, labrum only slightly longer than head is high. Thorax rather densely dark grey dusted even on mesonotum, with three distinct dark stripes along the lines of bristles, clearly visible from all angles, more brownish in anterior view, and the interspaces always grey. All mesonotal hairs and bristles black, acrostichal and dorsocentral bristles irregularly biserial, latter uniserial anteriorly and behind, all barely as long as 3rd antennal segment without style, becoming longer posteriorly. A strong humeral bristle, a posthumeral, 3 notopleurals, a supra-alar (with 1 or 2 additional bristles anteriorly), a postalar, and 4 scutellar bristles: outer pair rather strong, more than half length of the inner pair, though very exceptionally 5 to 6 strong scutellar bristles, and I have examined one male from Sweden (Ly.lpm., Oltokjaure) with two scutellar bristles only. Metapleural fan composed of pale and black bristles, either pale or black may dominate in some specimens. Pronotum with a row of short black bristly-hairs, but sides of prosternum and episterna with several pale hairs. Wings clear, all veins complete including anal vein, all rather light brownish except for darkened costa and radial veins. Squamae pale with concolorous fringes, halteres yellowish. Legs black and somewhat subshining except for grey coxae, hairs and bristles mostly black, but fine hairs on inner sides of fore coxae before tip and microscopic pile beneath fore tibiae pale. Fore femora with short hairs except for 2 or 3 long pos-

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teroventral bristly-hairs at tip. Mid femora anterodorsally with 2 or 3 long bristles at base (longer than femur is deep), apical two-thirds with more numerous fine short anterodorsal hairs, ventrally at base with a cluster of long, irregularly-set spinose bristles continuing in two rows of antero- and posteroventral bristles, those in anterior row very long about middle (twice as long as femur is deep), followed by much shorter hairs placed closer together on apical third of femur, posteroventral bristles evenly spaced and barely as long as femur is deep. Hind femora with fine short reclinate hairs dorsally and posteriorly, those posteriorly at base becoming paler, anterodorsally with a row of outstanding bristles about as long as femur is deep, those near base finer, shorter and stouter apically; anteroventral bristles present on apical two-thirds of femur, those about middle the longest, longer than femur is deep, becoming shorter and more numerous towards tip. Fore tibiae with a row of posterodorsal bristly-hairs at least as long as tibia is deep, and several shorter bristles above becoming longer apically. Mid tibia with 2 to 3 (very exceptionally 4) very long anterodorsal bristles on basal half, and another conspicuously long preapical bristle, posterodorsal bristly-hairs short, ventrally with a double row of bristly-hairs, those in anterior row longer and coarser, 2 or 3 median bristles twice as long as tibia is deep. Hind tibiae gradually and evenly dilated towards tip, about as wide at tip as hind femur about middle, with a row of outstanding anterodorsal bristles at least as long as tibia is deep, posterodorsally with short bristlyhairs except for 2 or 3 longer apical ones, ventrally with short reclinate bristles and a terminal circlet of short bristles. Tarsi long and slender, basal segment on fore tarsi slightly dilated but barely as wide as tibia at tip, and about as long as next segment, and all segments with short terminal circlets of coarse bristles. Mid tarsus very slender, 2nd segment only slightly shorter than basitarsus, and all segments with even shorter terminal circlets of spines except for a very long anterodorsal terminal bristle on basitarsus. Hind basitarsi stout but not deeper than tibiae at tip, about as long as next two segments together, clothed with short hairs, ven-

trally spinose, terminal circlets very short, absent on apical three segments. Abdomen dull grey, rather long pale pubescent at sides and at base, including long straggling pale bristles on venter, but dorsum covered with mainly short black hairs, also longish hind-marginal bristly-hairs black. 7th tergum grey, posteriorly at sides with shallow excisions, but sides below them pro-

duced into slightly pointed lateral processes. 8th

tergum polished black, laterally with conspicuous double-headed microscopically pubescent tubercles. Genitalia as in Figs 176-180, aedeagus short and stout, widening at tip, lateral lamellae covered with sparse black hairs and a few longer bristles on outer margin. 8th tergum also almost bare except for a few, mostly fine, marginal bristles. Length: body 34.2 mm, wing 3.14.1 mm. ?. Labrum somewhat longer, usually one-anda-half times length of head. Thorax including mesonotal stripes as in male, but all hairs and bristles shorter and less prominent. Wings brownish, usually darker on costal half, with a faint brown stigma. Fore legs with entirely short and fine ordinary hairs, those in posterodorsal row on tibiae the longest, but much shorter than tibia is deep. Mid legs short pennate on femora, tibiae covered with short ordinary hairs and a terminal circlet of rather short, coarse bristles. Mid femur anterodorsally with a row of short pennate fringes barely half as long as femur is deep, becoming shorter towards

tip, apical fifth of femur with adpressed hairs; anteroventral ciliation of short flattened hairs present on apical two-thirds of femur. Hind femora dorsally with a similar pennation to that on mid pair, but with two rows of dorsal pennate fringes, those in anterior row longer and broader, disappearing on apical quarter. Hind tibiae with a row of dorsal flattened hairs almost throughout, the pennate fringes (often rather narrowly flattened hairs) rather shorter and intermixed among them with additional longer outstanding bristles nearly as long as tibia is deep, sometimes arranged in distinct diverging pairs, at least apically. Tarsi simple, all basitarsi long,second tarsal segment on all pairs about half as long as basitarsus. Abdomen with short hairs even on venter, black hairs predominant, pale pubescence confined to at least sides of anterior segments. Terga less grey dusted than in male, more subshining and distinctly blacker when viewed from behind. Last two abdominal segments and long slender cerci grey dusted.

Figs 176-180. Male genitalia of Empis (s.str.) nigripes F. (CZ, Bohemia, Jistebnice). — 176: postabdomen; 177: 7th and 8th tergum in dorsal view; 178: lateral lamella in lateral view; 179: dorsal lamella in lateral view; 180: aedeagus. Scale: 0.2 mm.

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Length: body 34.1 mm, wing 3.2-4.2 mm. This is the main (nominal) and most common representative of the taxonomically very difficult group of species differentiated here as the E. nigripes-complex. The male of E. nigripes, with a stout and apically broadened aedeagus, needs comparison in particular with E. laminata. All the other closely related species have the aedeagus either narrowed apically (E. bicuspidata, E. goott), or longer and slender throughout and with distinct loops (E. nuntia, E. limata, E. planetica). Further good distinguishing characters are the structure of the two pregenital segments in male, the bristling of the legs, and the colour of the abdominal pubescence and metapleural fan. E. nitidula, another

can be found at any hour of the day swarming in large, fairly dense aggregations. Male swarms were first described by Tuomikoski (1939), and the complete epigamic behaviour in comparison with the related E. nuntia was discussed by Chvala (1990) as an example of ethological isolation. Males of E. nigripes fly very rapidly in large aggregations, without apparent orientation to any swarm marker, and hunt small insects as prey. This is transferred to females in special loose female swarms in the vicinity; females fly rapidly in a horizontal flight very close to the ground, waiting for males.

very Closely related species (if specifically distinct from E. nigripes) has an extensively polished black thorax and abdomen. The identification of females in the E. nigripescomplex is much more difficult (except for E. laminata, with simple legs without pennation), and the very slight differences combined with the obvious variation in both the pennation of legs and the colour of body pubescence make it almost impossible to identify single females. The species of the E. nigripes-complex discussed in the following pages are compared with E. nigripes in their descriptions. There is also a frequent venation aberration in E. nigripes, with the radial fork often incomplete or entirely missing (often on one side), and even the number of large scutellar bristles varies to some extent. For many years E. nigripes was known as Empis pennaria Fallén, as Fabricius’ name nigripes was mistakenly used in the genus Rhamphomyia. For the full synonymy, see the detailed explanation given by Collin (1961:523-524).

Empis nitidula Zetterstedt, 1859:5008.

Distribution. Widely distributed throughout Scandinavia, common both in the south and in the north. One of the very common Empis species in Scandinavia, and known for a long time as E. pennaria or E. vernalis; | have studied more than 850 specimens (506 d and 347 @ ). A typical spring species, occurring mainly in June; the earliest data of capture of Scandinavia is 11 May (Skane), the latest 11 August (one male collected at Are in Jamtland by O. Ringdahl). — Throughout Europe including the Mediterranean (though rare in the south), east to West Siberia. Very common, undoubtedly the commonest Empis species in temperate central Europe, mainly in May and June.

36. Empis s.str. nitidula Zetterstedt, 1859

Very much like E. nigripes, with the same male genitalia and rounded tubercles on each side of 8th tergum, but thorax and abdomen extensively shining black, abdominal pubescence completely yellowish including hairs on genital lamellae; legs in female with similar fringes, but body shining as in male. Lectotype designation. A curious species known to me only from two syntypic males and a female in the Zetterstedt Collection in Lund; the males are in the Goteborg Collection, and the female in the Wallengren Collection. All have a small label ‘Ld 2/6’; one male (with its abdomen glued on a card) is labelled ‘Emp. nitidula Zett. n.sp. ¢. Lund’ and is herewith designated as lectotype; the female has an identical label but with a different sex sign. Zetterstedt (1859) described this species from ‘Scania ad Lund’, and these three specimens are undoubtedly syntypes. For a long time I thought that they might be aberrant or somewhat damaged (rubbed) specimens of E. nigripes, but the presence of wellpreserved bristles on a smooth and polished mesonotum indicates the specimens are undamaged. Nor were they kept in alcohol or some other liquid and later dried out, as the wings are also undam-

aged. Although I have examined hundreds of specimens of E. nigripes, including southern Scandinavia and Skane, I have never seen such shining black, non-pollinose individuals. All three syntypes were collected on one day (2 June) at Lund before 1859, and they could be very rare or even an extinct species, but I believe that E. nitidula should be redescribed here as a valid species of the E. nigripes-complex. 3 (lectotype). Head as in E. nigripes, with similar occipital bristling and structure of antennae,

Biology. Adults are common throughout the day on various flowers, sucking sweet nectar, but they

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but occiput mostly shining black or at least subshining above neck, narrow upper postocular mar-

gins greyish pollinose and also the lower part as far as the mouth opening. Labrum shining brown, only slightly longer than head is high. Thorax shining black on mesonotum, but notopleural depression right up to humeri, prescutellar depression (partly damaged by pin) and scutellum densely grey dusted, as are pleura. All thoracic bristles as in E. nigripes, black and rather long, sides of prosternum and prothoracic episterna with a few pale hairs, pronotum with blackish bristly-hairs. Metapleural fan completely whitish-yellow, with no additional dark bristles. Wings completely clear, with the same venation as in E. nigripes, costa and radial veins dark brown, other veins rather fine and inconspicuous, a long black costal bristle. Squamae translucent brownish, fringes pale, halteres whitish-yellow with base of stalks darker brown. Legs subshining black, with the same structure

and bristling as FE. nigripes, including rather long black bristly-hairs posterodorsally on fore tibiae, the irregularly densely-set and somewhat brushlike black bristles on mid femur ventrally at base becoming widely spaced and shorter towards tip; no differences in bristling of mid and hind tibiae. Abdomen (glued on card) shining black, only very indistinctly finely greyish pollinose, pubescence uniformly pale, whitish-yellow, denser and

longer at sides of anterior segments, only a few hind-marginal bristles which are fine, slightly longer and darker. 8th tergum polished like the preceding segments, with the characteristic rounded tubercles of E. nigripes at sides, including the corresponding excisions on 7th tergum. Genitalia of the same structure, aedeagus rather yellowish-brown,

with apex concealed within lateral lamellae, latter unlike E. nigripes, covered with uniformly pale hairs, including longer ones on lower margin before tip. Length: body 3.6 mm, wing 4.0 mm. 2. The single paralectotype female is not in very good condition, somewhat mouldy, with partly damaged wings, and of the hind legs only left femur is preserved. Although all legs on the right side are lost, the main characters are clearly visible and the structure of pennation and flattened bristles (partly damaged on mid femur beneath) seem to be exactly the same as in E. nigripes — only the hind tibiae and tarsi remain unknown. Thorax and abdomen shining black as in male, including the contrasting greyish pollinosity of the notopleural depression, base of scutellum and prescutellar depression, but margins

of scutellum

(unlike the

male) also shining. Metapleural fan (visible on left side) partly blackish.

Length: body 3.0 mm, wing 3.8 mm. The second male is in very poor condition: wings, abdomen and legs on the right side are lost (left wing glued on card), but otherwise all visible characters (including only the partial dusting of mesonotum in notopleural depression) are as in the lectotype male, except that the metapleural fan is not entirely pale as there are 3 blackish bristles intermixed on each side. I think that E. nitidulaisa good species related to E. nigripes, and hope that further specimens will be found in the future that will enable dissections of male genitalia to be made.

Distribution. Known only from two males and a female taken on 2 June at Lund in Skane, Sweden.

37. Empis s.str. laminata Collin, 1927 Figs 181-185. Empis laminata Collin, 1927: 22 (as subsp. of E. pennaria Fallén). Very much like E. nigripes, with a similar short aedeagus, but lateral tubercles on 8th tergum flattened and spoon-shaped, directed forward into deep excisions in the 7th tergum. A more bristly species, with metapleural fan mostly black and abdomen with entirely pale hairs on dorsum. Female legs without pennation or flattened hairs, covered with ordinary hairs and bristles. 6. Head as in E. nigripes but occipital bristles more numerous, longer and coarser, postocular bristles in particular conspicuously long and curved forwards, in side-view almost reaching front edge of eyes. Thorax rather uniformly dark grey, the dark longitudinal stripes distinct only when viewed from behind, in anterior view with traces of two faint

darker stripes between the rows of bristles. Acrostichal and dorsocentral bristles decidedly longer than in E. nigripes, former reaching further forward anteriorly, present even on front part of mesonotum, even if smaller. Outer pair of the four scutellar bristles prominent, not very much shorter than inner pair. Metapleural fan composed of pale and black bristles in northern populations, mostly black in southern specimens. The same applies to the colour of bristly-hairs at sides of prosternum; in southern populations the hairs are mostly black. Wings and halteres as in E. nigripes, squamae more dirty greyish. Fore legs with tibiae with longer and denser bristles above, especially posterodorsal bristles longer, about one-and-a-half times as long as tibia is deep.

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Figs 181-185. Male genitalia of Empis (s.str.) laminata Collin (NL, Reeuwijk).—181: postabdomen; 182: 7th and 8th tergum in dorsal view; 183: lateral laemella; 184: dorsal lamella in lateral view; 185: aedeagus. Scale: 0.2 mm.

Mid femora with ventral bristles arranged in two regular rows as far as base (those in anterior row longer), and even though the bristles are placed closer together at extreme base of femur they do not form a cluster of bristles as in E. nigripes. Unlike E. nigripes, dorsum of mid femora covered with short hairs, the distinct anterodorsal bristles of nigripes replaced by 2 or 3 fine anterodorsal hairs much shorter than femur is deep. Similarly,

hind femora with shorter bristly-hairs dorsally, the anterodorsal bristles much shorter, but with a few

longish fine bristly-hairs anteriorly and posteriorly in basal third to half, apical half of femur with a row

of anteroventral bristly-hairs as in E. nigripes. On the other hand, posterior four tibiae more distinct-

ly bristled: mid tibiae with 4 to 5 very long anterodorsal bristles and a still longer preapical one, and both posterodorsal and ventral bristly-hairs longer and coarser. Hind tibiae similar in shape to E. nigripes, but clothed dorsally with regular pairs of antero- and posterodorsal bristles, distinctly longer than tibia is deep. Anterior four tarsi with longer

112

basitarsi, 2nd tarsal segment much shorter, hardly more than half length of corresponding basitarsus. Hind tarsi as in E. nigripes. Abdomen lighter grey dusted when viewed from above and extensively whitish pubescent even on dorsum, leaving hind marginal bristles on Ist tergum and 8th segment black. 7th tergum deeper excised laterally, the central part thus appearing to be more produced posteriorly above the polished black 8th tergum; moreover, sides of 7th tergum not produced so far laterally as in E. nigripes, lining-up more with abdominal margin. The lateral tubercles on 8th tergum flattened and spoon-shaped, pointing anteriorly only, and fitting well into the deep excisions of 7th tergum. Genitalia with a stout and apically broadened aedeagus, similar to E. nigripes, dorsal lamellae almost identical, but lateral lamellae obviously taller, and

densely covered with long black bristles. More numerous black bristles also on ventral lateral part of 8th tergum (though absent as in E. nigripes in spec-

imens from Swedish Norrbotten) and on 8th sternum in particular. Length: body 3.0-3.7 mm, wing 3.2-4.2 mm. 2. All bristles on head distinctly shorter including postocular bristles. Labrum decidedly longer than in male, nearly twice as long as head is high. Thorax and mesonotal stripes as in male, but acrostichal and dorsocentral bristles less prominent, though the small anterior acrostichal bristles

reach right up to the front edge of mesonotum as in male, in the form of fine small hairs. Wings almost clear with distinct dark brown veins, but basal section of vein M (closing discal cell) very fine. Squamae extensively darkened, fringes pale. Legs simple, covered with only short ordinary hairs, without pennation or flattened hairs. All femora almost bare, mid femora with several small anteroventral bristles before tip, similar bristles also on apical half of hind femora. Anterior four tib-

and 25 @ ), collected during June, but two dates are exceptional: a male taken by R. Danielsson in Swe-

den (Sk., Dalby, O. Molle) on 4 May 1989, and two females captured by H. Lindberg in Finland (AI, Sottunga, Hus6) very late on 22 July 1952. — An overlooked species, known to me outside Scandinavia only from a few specimens from the Netherlands and central Europe (Bohemia). Collin (1927) described it from the continent, from Slovakia.

38. Empis s.str. gooti sp.n.

Figs 186-190.

in Sweden

A species of the E. nigripes-complex with a slender and apically narrowed aedeagus as in E. bicuspidata, but the tubercle on 8th tergum with a single, forwardly-directed point. Metapleural fan black, prothorax and dorsum of abdomen with entirely black hairs. Hind tibiae in female practically not pennate above, the small fringes intermixed with long dorsal bristles. d. Head rather dark grey dusted on occiput, eyes meeting for a long distance on frons, upper ommatidia very enlarged. Upper half of occiput with a row of forwardly-curved thin postocular bristly-hairs, behind them with two irregular rows of short black occipital hairs, those below neck becoming paler. A pair of long ocellar bristles as long as upper postocular bristles. Antennae black, style only slightly shorter than rest of 3rd segment. Palpi black, with 2 or 3 fine small black hairs apically. Labrum black, slightly longer than head is high. Thorax rather dark grey dusted, mesonotum uniformly grey and unstriped in anterior view, viewed from above and from behind with 3 blackish stripes along the lines of bristles, the stripes distinct because of the very pale grey interspaces. Acrostichal and dorsocentral bristles black, biserial, arranged as in E. nigripes; both rather long anteriorly but absent on anterior quarter of mesonotum, posteriorly longer and bristle-like. Large marginal bristles prominent: a humeral (with several small bristly-hairs anteriorly), a posthumeral, 2-3 notopleurals, 2 supra-alars, a postalar, and 4

known only from Skane and then from 3 males tak-

scutellars, outer pair not very much shorter than

en by H. Andersson on 28 June 1963 at Nb., Ranea, H6gs6n. Commoner in Finland, from Al, Ab and

the inner pair. All hairs on prothorax and metapleural bristles black.

N in the south, northwards to Om, Ok, ObS and

Wings clear, costa and radial veins blackish, other veins less distinct, faintly brownish. A rather fine black costal bristle. All veins complete, reach-

iae with a row of minute dark bristles dorsally,

those on hind tibiae obviously double-rowed and more distinct. Hind femora laterally very compressed, nearly twice as deep as the slender hind tibiae. All tibiae and tarsi with circlets of very short bristles, basitarsi on all pairs long and slender, covered with minute hairs, following tarsal segments much shorter, 2nd segment on all legs less than half length of corresponding basitarsus. Abdomen thinly grey dusted, somewhat subshining, pubescence very short and mostly pale. Cerci very long and thin. Length: body 2.8-4.2 mm, wing 3.2-3.6 mm. The male is very closely allied to E. nigripes, with which it is compared in the above description. Because of its simple legs without pennation or flattened hairs, female needs comparison with the

female of E. acinerea. The latter is a larger species, with very brown wings and long, slender, duller

grey hind femora, which are as deep and long as hind tibiae.

Distribution. Widely distributed but uncommon, in isolated localities far towards the north, approximately

to 65-66° N: for instance,

ODN. The species has been misidentified by all dipterists in Scandinavia, including Frey (see Chvala 1991), and Lundbeck, who took a series of 5 d and

9 @ of E. laminata over a period of 5 days between 6 to 20 June 1909 on Amager at Copenhagen and identified the males as E. vernalis and the females as E. cinerea. I have identified 56 specimens (31 ¢

ing wing-margin. Squamae translucent white, with dark margin and pale fringes, halteres pale yellow. Legs black, somewhat subshining, all hairs and bristles black, including those on fore coxae ante-

riorly. Fore femora covered with short adpressed

113

Figs 186-190. Male genitalia of Empis (s.str.) gootisp.n., paratype (CZ, Bohemia, Davle).—186: postabdomen; 187: 7th and 8th tergum in dorsal view; 188: lateral lamella; 189: dorsal lamella in lateral view; 190: aedeagus. Scale: 0.2 mm.

hairs except for a few longish hairs antero- and posteroventrally before tip. Mid femora with a row of densely set fine hairs anterodorsally, the hairs hardly half as long as femur is deep except for several longer bristly-hairs at base, similar dense pubescence also anteriorly; ventrally with a double row of bristles, 2 or 3 pairs at base very long, as long as those in anterior row at middle, about twice as long as femur is deep, and posteriorly with much shorter tiny bristly-hairs. Hind femora with fine hairs above, anterodorsally with several longish bristly-hairs, similar bristling also anteroventrally on apical half. Fore tibiae with soft brownish pile beneath, dorsally with rather short adpressed hairs and a single longer bristle beyond middle, posterodorsally with a row of rather long bristly-hairs about one-and-a-half times as long as tibia is deep,

and a terminal circlet of short bristles. Mid tibiae

114

usually with 3 long dorsal bristles (about 3 times as long as tibia is deep) in each third, and an even longer preapical bristle, ventrally with a row of anterior bristles on apical two-thirds (at least as long as tibia is deep), posteroventral bristly-hairs much shorter. Hind tibiae gradually and evenly dilated towards tip, dorsally in addition to short adpressed bristling with about 8 pairs of bristly-hairs becoming longer in apical third (longer than tibia is deep), ventrally with numerous rather long adpressed bristles. All segments on anterior four tarsi with terminal circlets of rather short bristles (generally about as long as 3rd segment), 2nd segment scarcely more than half length of corresponding basitarsus. Hind tarsi with even shorter terminal circlets, basitarsus long, cylindrical, hard-

ly as stout as tibia at tip, 2nd segment half as long and slightly narrower.

Abdomen dark grey dusted, more blackish when viewed from behind. All hairs on dorsum black, sides of anterior six segments densely whitish pubescent and sterna with long pale straggling bristly-hairs. 7th tergum subshining black, posterior margin produced into a small swelling-like tubercle at sides. no sublateral excisions as in E. nigripes or E. laminata. 8th tergum polished black, narrow, laterally with a finger-like tubercle pointing forwards, the tubercle very narrowed at tip and its swollen base with about 5 rather long black bristles. Genitalia with a slender aedeagus as in E. bicuspidata, but dorsal lamellae narrow and covered

with minute fine hairs; lateral lamellae larger, covered with rather long bristly-hairs and its outer tip more produced. 8th sternum with a row of long curved marginal bristles. Length: body 3.0-3.8mm, wing 3.3-3.8 mm, holotype body 3.5 mm, wing 3.8 mm. 2. Eyes separated by a broad grey frons with three pairs of long dark downcurved orbital bristly-hairs on lower half. Occipital bristles shorter than in male, labrum black, nearly twice as long as head is high. Thorax rather dark grey dusted, mesonotum with a similar pattern: uniformly grey and unstriped in anterior view, but viewed from behind prescutellar depression and the stripes between lines of bristles very light grey dusted, making the 3 dark stripes along the lines of bristles strikingly blackish. All hairs and bristles on thorax black, including the fine hairs on prothorax. Wings very faintly brownish, almost clear, stigma distinct, brown. Halteres pale, squamae extensively darkened, fringes pale. Legs black, front pair almost bare except for a short dorsal ciliation on femora and tibiae. Mid femora with short pennate fringes anterodorsally on the whole length and ventrally on apical two-thirds, mid tibiae with minute ordinary hairs only. Hind femora with two rows of adpressed pennation above except at tip, the posterior pennate bristles much shorter, ventrally on apical two-thirds with pennate fringes shorter than femur is deep. Hind tibiae with short, almost ordi-

nary (barely flattened) bristly-hairs along the whole length on dorsum and ventrally on basal half, latter with a tendency to be flattened at base; dorsum of tibia in addition with 3 long bristly-hairs

rior view, subshining when viewed from behind, all pubescence short and black, even at sides; the hairs

on anterior segments at sides with a tendency to be brown, not pale from any point of view. Cerci very long, slender. Length: body 3.2-4.0 mm, wing 3.3-4.0 mm. The male of FE. gooti sp.n. has an apically narrowed aedeagus similar to that in E. bicuspidata, but the characteristic tubercle at the sides of 8th tergum is very different, smaller and narrowly pointed forward only. Furthermore, the genital lamellae are differently shaped, the abdomen has entirely black hairs on dorsum, and there are further differences in the leg bristling. The black hairing at sides of prosternum distinguishes both sexes from all other species of the E. nigripes-complex. The female also differs in the practically non-pennate hind tibiae and almost entirely black pubescent abdomen; on the basis ofthe latter, females of E. gooti come very close to E. melanotricha and its allies, to species with a uniformly black pubescent abdomen. Derivatio nominis. The species is named in honour of the Dipterist Mr V.S. van der Goot, Amster-

dam, who found this new species in the Netherlands and who recognised it as a distinct species. Holotype d: Slovakia or., Cejkov, 13.5.1965, leg. M. Chvala; in the author’s collection at the Charles University, Prague. Paratypes: Slovakia or., Latorica most, Leles, 15.5.1965 3 5 and 1 2, leg. M. Chvala; Bohemia centr., Radotin (Sulava), 16.5.1965,5-¢

and 1 @,

7.1971, 2 2, leg. J.Macek; Bohemia centr., Davle,

11.5.1985, 1 d, 1.6.1985 2 3, leg. J. BouSkova. The Netherlands, St. Pietersberg Lbg., 15.5.1988, 2 d, leg. B. Van Aartsen. In the author’s collection; the Dutch specimens in the University Museum, Amsterdam. Note. I have also seen 2 females taken by R. Da-

nielsson on 28 May 1988 at Sk., Dalby, O. Molle, Sweden, together with a long series of both sexes of E. nigripes, which are almost certainly females of E. gooti. The only difference seems to be a suggestion of pale hairs at the extreme sides of the Ist tergum, and one female has narrowly flattened hairs also on the dorsum of hind tibia. Since no

in each quarter, almost asjong as tibia is deep, and

males were collected, the two females are not in-

another pair of similar preapical bristles. All tibiae and tarsal segments with circlets of very short bris_ tles at tip. Fore basitarsus very long, 3 times as long as following 2nd tarsal segment, basitarsi on posterior two pairs almost as long but the following 2nd segment longer, almost half as long as corresponding basitarsus. Abdomen dark grey dusted in ante-

cluded in the type series. The same applies to six further females collected by me at several localities in southern Bohemia (Veseli n.Luznici, Ruda 19.5.1966, Zlata Koruna, Rajov 22.5.1978, Alber

20.-26.6.1977): these have not been included in the type series as no associated males were captured, though they belong almost certainly to E. gooti.

115

Distribution. Rare species, known in Scandinavia

only from the two females from Swedish Skane. — Known only from the Netherlands and central Europe (Bohemia, Slovakia), from May to July.

39. Empis s.str. bicuspidata Collin, 1927 Figs 191-195. Empis bicuspidata Collin, 1927:22 (as subsp. of E. pennaria Fallén). Very closely resembling E. nigripes in both sexes. The male differs chiefly in the apically slender and narrowed aedeagus, in the bifurcate tubercle on 8th tergum, which is remarkably bifid, and in the differently arranged bristles on mid femora beneath. Female hind tibiae fringed dorsally with uniform pennate bristles. 6. Head as in E. nigripes but the curved postocular bristles on occiput above apparently finer and smaller, not as prominent as a pair of ocellar

bristles. On the other hand, lower part below neck with longer and more numerous pale hairs. Thorax grey dusted, mesonotal stripes much more like those in E. /aminata: the stripes are not at all brownish, very indistinct in anterior view, and alternating with two dark stripes between the rows of bristles. Thoracic bristling as in E. nigripes, acrostichal bristles also absent on anterior part of mesonotum, and all less prominent than in E. lami-

nata. Metapleural fan varying in colour from entirely pale to individuals of the same local population with metapleural bristles predominantly black. Four scutellar bristles, with outer pair rather long. Wings, halteres and squamae as in E. nigripes. Legs not entirely black, with a tendency to be translucent dark brown. Otherwise, structure and bristling almost the same as in E. nigripes, the only substantial difference being the arrangement of the ventral bristles on mid femora, where the bris-

tles in a double row are almost equally long and

Figs 191-195. Male genitalia of Empis (s.str.) bicuspidata Collin (SF, N: Esbo, Kolmpera). — 191: postabdomen; 192: 7th and 8th tergum in dorsal view; 193: lateral lamella; 194: dorsal lamella in lateral view; 195: aedeagus. Scale: 0.2 mm.

116

regularly spaced on basal half of femur, and not clustered as in E. nigripes. Abdomen rather densely grey dusted, duller

E. bicuspidata: the radial fork is often incomplete or entirely missing.

black when

Distribution. Widely distributed throughout Scandinavia, including the extreme north, in Norway to

viewed

from

behind,

covered

with

whitish hairs even on dorsum, but the longish bristly-hairs on posterior margins on all terga very darkened or nearly black. 7th tergum grey dusted, not distinctly excised as in E. nigripes or E. laminata, but with a small polished tubercle on each side below. 8th tergum polished black, lateral tubercles prominent, conically bi-pointed, one process pointing anteriorly above 7th tergum, the other posteriorly. Genitalia with a short, basally stout but apically very narrowed aedeagus, upper lamellae rather larger and covered with short hairs, lateral lamellae more ovate (not produced on outer upper margin) and covered with only a few, rather short brownish bristly-hairs. Length: body 2.5-3.7 mm, wing 3—4.1 mm. 2. Very much like female of EF. nigripes, differing as follows: Mesonotum practically unstriped

TRi, in Sweden to Lu.Lpm., and in Finland to Le and Li; much less common towards the south, as

for instance only 2 ¢ are known from Denmark (one taken by Hansen in 1881 at Frederikshavn, the second in 1964 by O. Martin and B.V. Petersen on Bornholm) and only 4 ¢ and1 2 from Swedish Skane. Altogether 277 specimens have been examined (183 ¢ and 94 @), on dates ranging from 27 May to 6 August. Rather an early summer species, mainly in the second half of June and in July, anda few records from early August are all from the extreme north (Nb., Lu.Lpm.), but there is one male in the Lund University Collection taken by Roth in Skane (S. Sandby, Kungsmarken) allegedly on 28 April. — Rather a boreal species in distribution, recorded from England and Scotland, the Nether-

when viewed from in front, the blackish stripes

lands, and south only to central Europe, to Germa-

(not brownish) distinct in posterior view only. Metapleural fan generally entirely pale, very rarely with one or two additional black bristles. Wings faintly brownish, posterior half often almost clear. Mid femora with short pennation dorsally and on

ny, the former Czechoslovakia and east to central European Russia as far as the Moscow region. Eyerywhere uncommon; in temperate Europe especially in the second half of May and in June.

apical half beneath as in E. nigripes, but hind femora with longer pennation on apical half beneath,

40. Empis s.str. nuntia Meigen, 1838

that in apical third almost as long as femur is deep; dorsal pennation on the contrary perhaps shorter, especially the fringes in posterodorsal row smaller. Hind tibiae above with more uniform pennate fringes, apparently longer than in E. nigripes, no ordinary bristles intermixed and, if some longer bristles present on dorsum, then always more or less flattened. Abdomen paler pubescent, especially discs of terga with exclusively short pale hairs, some longer dark to almost black hairs on posterior margins. Length: body 2.8-3.8 mm, wing 3.2-3.7 mm. The male of EF. bicuspidata has a short, apically very narrowed aedeagus, as in E. gooti, but may be easily recognised by paler hairing and by the prominent bi-pointed lateral tubercle on 8th tergum with points directed forwards and backwards. The female is difficult to separate from that of EF. nigripes, and the only useful differences seem to be the indistinctly striped mesonotum, the mostly pale metapleural fan and abdominal pubescence, the longer pennation ventrally on hind femora before tip, and in particular the uniform pennation on hind tibiae above. As in other species of this complex, there is a frequent venation aberration in

Figs 196-200. Empis nuntia Meigen, 1838:85.

Medium-sized (about 3-4 mm), dull grey dusted species with pale halteres, resembling E. nigripes but legs brownish, mesonotum viewed from in front with two darker stripes between the lines of bristles, metapleural fan and abdominal pubescence completely pale. 8th tergum simple in male, without tubercles, aedeagus slender, with a loop before tip. Hind tibiae. in female short pennate above, without additional bristles.

3. Occiput densely grey dusted, with rather fine black bristly-hairs above, with longer whitish hairs below. Antennae black, style not very much short-

er than 3rd segment. Labrum about 1.5 times as long as head is high. Thorax dull, rather brownish-grey dusted on mesonotum, pleura not paler. Mesonotum with two darker (blackish-grey) narrow stripes between the lines of bristles in anterior view, viewed

from above with a dark stripe along acrostichal bristles and another broader stripe on each side laterally of dorsocentral bristles. Large bristles black, in full number: a humeral, a smaller intrahumeral, a posthumeral, 3 notopleurals, one small

ay

and one strong supra-alar, a postalar, and 2 pairs of scutellar bristles, unlike EF. nigripes outer pair always very small and fine, at least less than half as long as inner pair. Acrostichal and dorsocentral bristles rather fine, black, former biserial, latter uniserial anteriorly, biserial about middle and ending in 2 strong prescutellar pairs. Prothoracic pubescence pale, pronotum rarely with darkened bristles; metapleural fan completely pale, no additional black bristles as in E. nigripes. Wings clear or slightly yellowish to light brownish clouded, veins distinct except for fine medial veins, anal vein complete. A very small, fine, black costal bristle. Squamae pale with concolorous fringes, halteres yellowish. Legs slender, distinctly brownish, almost yellowish-brown on femora and tibiae at base, only hind tibiae slightly evenly dilated apically, hind basitarsus as deep as tibia at tip or hind femur at about middle. Fore femora practically bare, tibiae with small dark posterodorsal bristly-hairs shorter than tibia is deep, ventrally with soft brownish pubescence present also on basitarsus; basal three

tarsal segments with long black bristles in terminal circlets. Mid femora with short hairs above, beneath with a double row of long black bristles, those on basal half about twice as long as femur is deep, the bristles in anterior row regularly spaced even at base, not ‘brush-like’ as in E. nigripes. Mid tibiae with 2 long black anterodorsal bristles on basal half, a smaller one at base, a longer one before middle, and another preapical one the longest; ventrally with short adpressed bristling, the bristles before middle about as long as tibia is deep. Mid basitarsus with longish black anterodorsal bristles (as long as 3rd tarsal segment), otherwise terminal circlets very short. Hind femora with black anterodorsal bristles longer on basal third, becoming shorter and more dorsal apically, posteriorly near base with long pale hairs, and ventrally with darker hair-like bristles, about 4 bristles beyond middle the longest, as long as femur is deep. Hind tibiae bristled above, about 8 bristles in antero- and posterodorsal rows nearly as long as tibia is deep, intermixed with shorter hairing, ventrally with short adpressed bristly-hairs. Hind basitarsus

Figs 196-200. Male genitalia of Empis (s.str.) nuntia Meigen (CZ, Bohemia, Davle).—196: postabdomen; 197: 6th-8th terga in dorsal view, diagrammatic; 198: lateral lamella; 199: dorsal lamellae in anterodorsal view, lateral lamella dotted; 200:

aedeagus with hypandrium. Scale: 0.2 mm.

118

with similar more adpressed bristling as on tibia, with short black spines ventrally. Abdomen dulled by greyish dust, densely pale pubescent, longer at sides and on basal segments,

sterna with long straggling pale bristly-hairs, and even long bristles on posterior margin of 8th sternum and on genitalia pale. Unlike E. nigripes and its allies, 8th tergum without tubercles or protuberances, but apical two terga (7th and 8th) distinctly produced laterally on posterior margin. Genitalia small, lateral lamellae triangular in shape, with dark short hairs above, rather as on the

mostly concealed dorsal lamellae. Aedeagus slender, light brownish,

Denmark at Jydelejet with an abbreviated vein M, as in Coptophlebia. Synonymy. For synonymy and type revisions, see Collin (1961:528). However, E. baldensis Strobl (described as a variety of E. pennaria Fallén and for a long time thought to be identical with E. nuntia), which was described by Strobl (1899) from a single holotype female from Mte Baldo, Italy (not Spain, as erroneously stated by Collin), is a good species closely related to E. pennipes, E. procera Loew and E. nitidiventris Loew, a group of species with conspicuously long labrum and strongly pennate legs in female.

before tip with an outward

loop, tip not broadened; ventral lamella projecting posteriorly, shining, and shielding base of aedeagus from below. Length: body 3-4.2 mm, wing 44.5 mm. 2. The main differential characters as in male, including the mesonotal pattern and the distinctly brownish legs, strikingly paler than in other related species, but wings more brownish and abdominal pubescence short. Legs pennate as in E. nigripes, but hind femora above with shorter pennation in one row only, and hind tibiae above, on the contrary, with longer and more uniform pennation (though still shorter than tibia is deep) and with no longer simple bristles intermixed except for a diverging dorsal pair at tip; similar pennate fringes also beneath on basal half. Hind femora with rather long pennate fringes ventrally on apical half, those before tip not very much shorter than femur is deep, but mid femora with only short flattened bristles on the whole length above, and a few similar bristles beneath just before tip. Fore legs and mid tibiae with simple short bristly-hairs, almost bare. Length: body 3-4.3 mm, wing 3.2-4.2 mm. E. nuntia belongs to the very difficult species complex of E. nigripes but can be recognised by the distinctly lighter brownish legs, the two darker stripes on mesonotum between the lines of bristles clearly visible in anterior view, and the very small outer pair of scutellar bristles. The male has a rather slender aedeagus with a subapical loop, thus resembling the smaller-sized E. staegeri or E. planetica. ira: As in E. nigripes, frequent venation aberrations occur in FE. nuntia. Collin (1961) mentioned a female from England, resembling Aclonempis, with no indication ofa radial fork on either wing, exactly the same as in a female taken by B. Van Aartsen in the Netherlands at St. Pietersberg, and I have seen

a male taken by S. Andersen and V. Michelsen in

Distribution. A species with a typical southern distribution in Scandinavia, known to me from 37 6

and 10 2 only from Denmark and Swedish Skane. All specimens were captured during a brief period from 18 May to 14 June and, apart from a female taken by Ammitzboll at Vanga Nytorp (Skane) on 20 May 1919, all specimens have been collected recently, between 1981 and 1989. As Collin (1961) recorded it as a very common species from all parts of Great Britain between 28 April and 10 July, it seems as if FE. nuntia has only recently expanded into new areas in northern Europe. — Also the Netherlands, but south only to central parts of Europe, to Switzerland and Hungary. In central temperate Europe, in the Czech countries, a very common species everywhere and almost as common as E. nigripes, in May and June. Biology. In the same biotopes as E. nigripes, adults are also frequent on flowers throughout the whole day but their epigamic behaviour is very different (Chvala 1990). The swarming and mating activity of E. nuntia was described by Smith (1952) and by Laurence (1959) who observed successful mating without prey transfer. E. nuntia produces male swarms, sometimes of males carrying prey (usually small Nematocera), but more often males swarm without prey and the entering females copulate readily with preyless males. In this species, apparently, the prey functions neither as a necessary food for females nor as a signal for mating, as part of an ethological isolating mechanism.

41. Empis s.str. chioptera Meigen, 1804 Figs 106-107, 120, 201-203.

Empis Empis 1790 Empis

chioptera Meigen, 1804:221. leucoptera Macquart, 1823:161 (not Gmelin, and Meigen, 1804). funesta Meigen, 1838:84.

119

Not very small (2.44 mm) dull greyish species with black halteres, and with whitish hairs on prothorax, humeri, notopleural depression and abdomen, metapleural fan pale. Male with fore basitarsi swollen and milky-white wings, female with brownish wings and posterior four femora with flattened bristles. 3d. Upper ommatidia above antennae very enlarged, occiput greyish dusted though almost shining black when viewed from above, covered with forwardly-curved black bristles that become finer and whitish below neck. Antennae black, style unusually short, about one-quarter length of 3rd segment, palpi black. Proboscis not very long, only slightly longer than head is high, head rather large and closely attached to thorax. Thorax densely grey dusted in anterior view, somewhat bluish-grey, but subshining black when viewed

from

above,

and

with

distinct

blacker

stripes along the lines of bristles when viewed from behind. Acrostichal bristles hair-like, black, narrowly biserial and slightly diverging, dorsocentrals longer and more bristle-like, irregularly triserial at middle, posteriorly uniserial and becoming longer, ending in a pair of strong prescutellar bristles of the same size as other black marginal bristles, scutellum with one strong pair. Humeri, notopleural depression anteriorly, and prothorax (pronotum, episterna and sides of prosternum) covered with fine whitish hairs. Metapleural fan unicolorous whitish, no dark bristles intermixed.

Wings milky-white, veins whitish and inconspicuous except for yellowish veins R, and R,,;, and blackish apical part of costa beyond vein R,. Radial fork (Fig. 106) open, very obtuse, discal cell truncate, anal vein complete even if fine, and axillary excision acute. Squamae whitish with concolorous fringes, halteres blackish. Legs dark brown to blackish, dulled by greyish pollen, pubescence and fine bristles mostly whitish. All femora and tibiae rather slender, but fore basitarsus distinctly swollen (usually very broad and flattened in dried specimens), at least as long as rest of tarsus, hind basitarsus obviously less swollen but equally long. Fore femora with short pale hairs and longer posteroventral pubescence ending in 3 long bristly-hairs before tip, fore tibiae with long, densely set, very fine posterodorsal hairs, continuing also on basitarsus. Posterior four femora with a double row of long whitish bristlyhairs ventrally, longer than femora are deep, on

and also with fine dense pubescence half as long as these, ventrally with two rows of densely set black bristly-hairs, longer around middle. Hind tibiae dorsally with several very fine long bristles ending in preapical diverging pairs, intermixed with shorter hairs. All basitarsi somewhat spinose beneath, and first three tarsal segments with circlets of not very long, black apical bristles. Abdomen black, finely whitish-grey pollinose, covered with entirely whitish hairs that become longer at sides and on venter, 8th sternum with long white hind-marginal hairs. Genitalia small and closed, lamellae very small and covered with entirely fine whitish hairs; lateral lamella blunttipped, dorsal lamella broadly excised, the lower larger lobe merging with side lamella. Aedeagus brownish, uniformly stout, concealed within lamellae, its lower half shielded by enlarged polished hypandrium (ventral lamella). Length: body 2.2-3.8 mm, wing 2.5-3.6 mm. ?. All hairs and bristles on body shorter, wings distinctly broadened, brownish clouded, costal margin darker brown, and all veins distinct, brownish. Legs covered with short hairs, rather slender, but fore basitarsus somewhat dilated, broader than tibia at tip. Mid and hind femora with short pennate fringes both dorsally and ventrally, on mid femora ventrally rather on apical half only, and hind femora with several longer pennate bristles anteroventrally before tip. Fore and hind tibiae fringed above, covered with short but somewhat stouter bristles than elsewhere. Abdomen uniformly dark grey, pubescence short and wholly pale. Length: body 2.64 mm, wing 3-3.7 mm. E. chioptera varies in size but may easily be distinguished from all the other species of this group by the entirely whitish pubescence on abdomen and thoracic metapleura, and by the presence of fine pale hairs on prothorax, humeri, and notopleural depression. Very rarely there are 3 or 4 scutellar bristles, but not infrequently there are venation aberrations with radial fork partly or entirely lost, or even vein M, abbreviated as found in Cop-

tophlebia. For synonymy, see Collin (1961:538) and Syrovatka and Chvala (1986:237) under E. funesta Meigen. Distribution. A characteristically southern species in Scandinavia, of the distribution-pattern ‘B’ defined in the section on Zoogeography. Common in Denmark and south Sweden to Smaland, in Nor-

hind femur more distinct on apical half only, but

way to VE, in Finland

also with three black anterodorsal bristles before tip. Mid tibiae dorsally usually with 5 very long (3 times as long as tibia is deep) but fine dark bristles,

northwards to Kb, also in Russian Karelia. More than 300 specimens examined (196 ¢ and 126 2),

120

from isolated localities

mainly from southern Scandinavia, on dates rang-

Figs 201-203. Male genitalia of Empis (s.str.) chioptera Meigen (CZ, Bohemia, Jabkenice). — 201: postabdomen; 202: lamellae in lateral view; 203: aedeagus with hypandrium. Scale: 0.2 mm.

ing from 4 May to 27 June, but mainly in May and the beginning of June. — Northern and central parts of Europe, absent in the south, the southern boundary of its distribution in Switzerland, Aus-

with

black

halteres,

thoracic

hairs and bristles

black, but abdominal pubescence whitish. Wings whitish in male, brownish in female with almost

Biology. One of the earliest spring species, often on flowering willow catkins together with Empis (Platyptera) borealis, the later specimens on blossoming blackthorn and hawthorn. Only a few incomplete records on swarming and mating activity are available, by Parmenter (1951) and Smith (1952); copulation is obviously connected with

clear apex, anal vein complete, reaching wing-margin. Legs not distinctly pennate in female, hind femora convex and clothed with flattened hairs above. ; 3. Occiput subshining black, covered with black bristly-hairs above that become longer and slightly paler below. Antennae black, style very long, nearly as long as 3rd segment. Proboscis and palpi blackish, labrum about 1.5 times as long as head is high. Thorax polished black, though somewhat very finely greyish pollinose when viewed from in front. Large thoracic bristles black: a humeral, with a

prey presentation.

smaller outer bristle and additional small hairs, 2

tria, Slovenia and northern Croatia, east to central

European

Russia.

Collin (1961) recorded

it in

Great Britain from 18 April, and this matches its

occurrence in central Europe.

42. Empis s.str. prodromus Loew, 1867 Figs 121-123, 204-206.

smaller posthumerals, 3 strong notopleurals, 2 finer supra-alars, a strong postalar, and 2 (usually crossing) strong scutellar bristles. Acrostichal and dorsocentral bristles biserial, distinctly diverging,

Empis prodromus Loew, 1867:54, 42.

rather small and hair-like, latter ending in 3 long

Small (about 2.5-3.5 mm) shining black species

prescutellar pairs. Pronotum usually with black bristly-hairs, but episterna and sides of proster-

IZ

num with greyish hairs. Metapleural fan usually black, sometimes with additional pale bristlyhairs, rarely almost entirely pale. Wings rather whitish, not milky-white as in E. chioptera, but not entirely clear. Costa blackish, veins R, and R,,,; more or less darkened, otherwise veins whitish and inconspicuous. Radial fork very open, vein R, straight and almost at right-angles to vein R,, axillary excision almost right-angled. Squamae dirty white with pale fringes, halteres blackish. Legs almost subshining black, covered with entirely brown or black hairs or bristles. Fore and hind basitarsi slightly dilated, broader than tips of corresponding tibiae. Fore legs with short hairs, only tibiae posteriorly with rather long dense pubescence about as long as tibia is deep. Mid femora ventrally with two rows of long bristles, those about middle much longer than femur is deep, mid tibiae with 3 very long anterodorsal bristles placed at base, at middle, and at tip, ventrally with 2 or 3

shorter pairs of diverging bristles. Hind femora with 2 conspicuously long anteroventral bristles in apical third, and 2 much shorter similar adpressed bristly-hairs anterodorsally. Hind tibiae with rather long reclinate hairs ventrally, and several long dorsal bristly-hairs continuing also on basitarsus. Basal two (or three on fore legs) tarsal segments with circlets of long apical bristles, and all basitarsi with spine-like bristles beneath. Abdomen subshining black, covered with whit-

ish hairs, rather short on dorsum, much longer at sides, sterna with only fine minute hairs. Genitalia in their ground-plan of the E. chioptera-type, upper lamella rather large with both lobes broad, ventral lamella bare, covering most of basal part of aedeagus. Length: body 2.5-3.2 mm, wing 2.6-3.1 mm. 2. Very black and rather shining, abdominal pubescence all short and with a tendency to be brownish on posterior segments. Wings faintly brownish clouded on costal half, apical half paler, prac-

Figs 204-206. Male genitalia of Empis (s.str.) prodromus Loew (CZ, Moravia, Lednice). — 204: postabdomen; 205: lamellae in lateral view; 206: aedeagus with hypandrium. Scale: 0.2 mm.

122

tically clear, all veins darkened and more distinct except for pale M, and M.,. Legs rather short and broadened, especially femora, mid femora and hind tibiae with rather long black ciliae above, shorter ciliation also on fore and mid tibiae posterodorsally. Hind femora distinctly laterally compressed, straight below and with only a short fine ciliation except for 2 or 3 longer anteroventral bristles at tip, dorsum very convex and covered with rather long (though scarcely half as long as femur is deep) flattened bristly-hairs. Length: body 2.6-3.5 mm, wing 2.6-3.2 mm. Because of its whitish abdominal pubescence, E. prodromus runs in the key to E. dasyprocta but is undoubtedly more closely related to E. caudatula; the latter is also a small, rather shining, black species but it can be immediately recognised by its black abdominal pubescence, shorter antennal style, the unusual long posterior setae on male genitalia, and by the ventrally pennate hind femora in female. The central European E. pseudoprodromus Collin is a distinctly greyer species with shorter antennal style, much more bristled mid tibiae in male, and pennate hind femora in female. A lecto-

type for E. prodromus was designated by Syrovat-

ka (1991). Distribution. Common in southern and southeastern parts of Scandinavia, in Finland northwards approximately to 63° (Tb, Kb), in Sweden one isolated capture of a male in As.Lpm. (Asele) on 28 June 1961 by Andersson and Larsson. Common in Denmark and in the south of Sweden, unknown to me from Norway. Altogether 172 specimens (85 d and 87 2) have been examined; often misidentified as E. chioptera in Scandinavian collections. A typical spring species, known to me on dates from 30 May to 17 July, but mainly in June, but there is one very exceptional late capture from Denmark, a male taken by H.J. Hansen at Frede-

rikshavn (NEJ) on7 August 1901. — From England in the west and Russian Karelia in the east south to central temperate Europe (Austria, northern Italy, Hungary and Moldavia); a common species especially in warm lowland biotopes on dates from 12 ‘May to 26 June. The adults are common on flowers, especially on Geranium.

E. prodromus but halteres rather brownish, male with more bristled legs and an unusually long bristled hypandrium (ventral lamella); female legs longer and more

slender, hind femora also with

long flattened bristles beneath. d. Occiput dull blackish-grey,

upper bristly-

hairs black, lower part below neck with whitish

hairs. Antennae black, style long and slender, almost as long as 3rd segment. Otherwise as in E. prodromus. Thorax subshining blackish when viewed from above, but rather densely dark grey dusted in anterior and posterior views. Bristling on thorax as in E. prodromus but metapleural fan usually brownish, often

completely

black

in more

southern

populations, and extensively pale in specimens towards the north. Wings clear, not whitish, costa and vein R, extensively darkened, vein R,,; brownish, other veins whitish and inconspicuous including the anal vein that reaches the wing-margin. Squamae whitish with concolorous fringes, halteres rather dark brown (not black), stalk often extensively yellowish. Legs blackish-brown to almost black, rather finely grey pollinose. Fore and hind basitarsi slightly swollen. Fore femora bare except for several long anteroventral hairs which become longer before tip, about as long as femur is deep at that point, fore tibiae finely long pubescent posteriorly, the hairs becoming longer on apical half, and similar pubescence also on basitarsus. Mid femora with long fine bristly-hairs anteroventrally that become longer on apical half (nearly twice as long as femur is deep), several similar bristly-hairs posteriorly about middle; the anteroventral bristles on mid femur confined to 3 bristles in apical third of femur. Mid tibiae with 4-5 long anterodorsal bristles on basal half (more than twice as long as tibia is deep) and another long preapical bristle, ventrally with more numerous shorter bristles on basal half arranged in diverging pairs. Hind tibiae with numerous long fine hair-like bristles dorsally that are distinctly longer than tibia is deep (also present on basitarsus) and ventrally with short pubescence except for longer bristly-hairs on basal half. Basal three tarsal segments on all legs with circlets of apical bristles, basitarsi beneath with rather dense

43. Empis s.str. dasyprocta Loew, 1867 Figs 124-125, 207-209. Empis dasyprocta Loew, 1867: 55, 46. Small, finely grey dusted species with blackish halteres and pale abdominal pubescence, resembling

short bristles. Abdomen subshining black, terga finely grey dusted when viewed from above, covered with fine whitish hairs that become longer at sides of all segments and on hind margin of last segment. Genitalia very conspicuous, with the long black bristled ventral lamella (hypandrium) shielding a

123

large portion of the aedeagus at base; lamellae small, covered with short fine black pubescence,

aedeagus usually free above, yellowish, slender at tip.

done in the description above. I have studied the type specimens from Langenau (Germany), which are in the Loew Collection in Berlin; a lectotype was designated by Syrovatka (1991).

Length: body 2.5-3.3 mm, wing 3-3.5 mm. 2. Everywhere with shorter hairs and bristles,

abdomen only short pale pubescent, wings almost uniformly and very indistinctly brownish-grey clouded, practically clear, and all veins much more distinct. Legs obviously longer and more slender than in E. prodromus, femora not broadened and not so flattened. Anterior two pairs almost bare,

Distribution. Western and central parts of Europe,

along the Atlantic coast from Spain northwards to the Netherlands, where it is not at all rare, and its occurrence in Denmark and Sweden is to be ex-

pected. Not known from England; common in lowland central Europe on dates ranging from 6 July to 12 August, but mainly in July. A typical summer

without fringes or ciliation; hind femora only very slightly convex on both sides, and both above and

species, like E. aestiva.

below clothed with rather long flattened bristles,

44. Empis s.str. caudatula Loew, 1867

those below distinct on apical half only. Hind tibiae not very slender, covered with minute simple hairs except for several short flattened bristles near base beneath. Abdomen subshining, very finely greyish pollinose. Length: body 2.6-3.6 mm, wing 3-3.5 mm. In the Scandinavian fauna, E. dasyprocta needs comparison mainly with E. prodromus, as has been

Figs 126, 210-213.

Empis caudatula Loew, 1867:35, 33.

Small (2.5-3.5 mm) subshining black species with 4scutellar bristles, black halteres, and black meta-

pleural fan and abdominal

pubescence.

Wings

milky-white in male, brownish towards base in fe-

Figs 207-209. Male genitalia of Empis (s.str.) dasyprocta Loew (NL, Cadier). — 207: postabdomen; 208: lamellae in lateral view; 209: aedeagus with hypandrium. Scale: 0.2 mm.

124

male, anal vein complete. Male with an unusual

posterior pair of long crossing setae on genitalia, female with only posterior four femora short pennate. 3d. Eyes with upper ommatidia very enlarged, occiput black, greyish dusted from some points of view, with forwardly-curved rather stout black bristles

above

that become

finer, straight, and

somewhat brownish but not pale below. Antennae black, style about half as long as 3rd segment. Labrum not very much longer than head is high. Thorax almost shining black when viewed from above, distinctly brownish grey pollinose in anterior and posterior views, pleura duller grey. All thoracic hairs and bristles black. Large bristles not conspicuous: a humeral, a posthumeral, 3 notopleurals, one larger and one smaller supra-alar, a postalar, and 4 scutellar bristles, outer pair much smaller (at most half as long) and finer. Acrostichal and dorsocentral bristles narrowly biserial, fine and hair-like, diverging, anteriorly equally small but posteriorly dorsocentrals becoming longer and stouter, ending in 3 long pairs. All hairs

on prothorax black, metapleural fan uniformly black. Wings milky-white, veins whitish except for almost blackish costa and darkened veins R, and R,,;, anal vein complete. Squamae dirty grey with black margin and black fringes, halteres blackish. Legs black, not very long but slender, fore and hind basitarsi slightly swollen, latter obviously less dilated, about as deep as hind tibia at tip. Fore legs with only short hairs except for several longer bristly-hairs on femora posteroventrally at tip, and rather long, dense posterior pubescence on tibiae and basitarsi, the hairs at least as long as tibiae are

deep. Mid femora with several longer bristly-hairs ventrally, those anteroventrally on apical half to third the longest, hind femora with 4—5 similar bristles before tip, and 2—3 shorter bristly-hairs anterodorsally. Mid tibiae with four anterodorsal bristles regularly spaced in each quarter, the last one forming part of preapical circlet, all scarcely twice as long as tibia is deep, ventrally with more numerous but much shorter reclinate bristly-hairs. Hind tibiae with short hairs, dorsally with several fine brist-

Figs 210-213. Male genitalia of Empis (s.str.) caudatula Loew (CZ, Bohemia, Albev). — 210: postabdomen; 211: lamellae in lateral view; 212: the same in anterodorsal view, aedeagus at middle; 213: aedeagus with hypandrium. Scale: 0.2 mm.

125

ly-hairs arranged in pairs, the last preapical pair the longest, a similar apical pair also present on basitarsi. Basal three segments of all tarsi with short bristles in apical circlets, and basitarsi short

but only Laurence (1950) has observed mating in aerial swarms: males were carrying prey of about their own size, mostly small nematoceran flies.

bristled beneath, not spinose.

45. Empis s.str. aestiva Loew, 1867

Abdomen finely grey pollinose though distinctly almost shining black in dorsal view, pubescence black, short on dorsum, longer and denser at sides, venter with fine hairs. Genitalia very much as in E. chioptera, but lateral lamellae blunt-tipped and with a long apical seta (both setae are unusual as they cross posteriorly when viewed from above), and all genital pubescence blackish. Length: body 2.5-3.4 mm, wing 2.8-3.3 mm. 2. Resembling male but wings brownish clouded on basal third, clear towards tip, and all veins brownish. Hind basitarsi slender but fore basitarsus slightly thickened, as deep as tibia at tip, and all legs covered with simple short hairs except for posterior four femora, which are short pennate on both sides. Length: body 2.5-3.5 mm, wing 2.6-3.3 mm. E. caudatula is the only species in Scandinavia of the taxonomically difficult group of small species with black halteres possessing the combination of black abdominal pubescence and complete anal vein. The male is very characteristic in having the long crossing posterior setae on genitalia, and the female in having non-pennate posterior tibiae (strongly pennate in E. aestiva). The species has often been misidentified as E. chioptera, for instance by Lundbeck (1910) in Denmark. I have seen one male taken by R. Danielsson on 18 May 1989 in southern Sweden (Sk., Dalby, Rogle) with radial fork completely missing on both wings as in Aclonempis, and even vein M, was abbreviated as in Coptophlebia. A lectotype for E. caudatula was designated by Syrovatka (1991). Distribution. A species of southern distribution in Scandinavia, known so far only from Denmark and southern Norway and Sweden, the northern-

most limit being Ostfold and Vestfold in Norway and the Swedish county Narke. However, it is very common in the south, and a total of 142 d and 100 ? has been examined. A typical spring species, on dates from 11 May to 9 July, but mainly in June. — Collin (1961) recorded it from England in May and June; rather common in temperate central Europe, south to northern France, Austria and northern ar-

eas of the former Yugoslavia. In central Europe a typical lowland species known to me from a short period in spring, from 14 May to 18 June. Biology. Adults are frequently swept from flowers,

126

Figs 127, 214-217. Empis aestiva Loew, 1867:54, 39.

Small (2.3-3.5 mm) shining black species with 2 scutellar bristles, black halteres, and black hairs and bristles on body and legs. Wings clear in male, brownish in female, anal vein incomplete. Posterior four femora and tibiae strongly pennate in female. 3d. Eyes with upper ommatidia very enlarged on a large area above antennae. Occiput at least subshining black, not grey from any point-of-view, at most dull black, and all bristly-hairs blackish. Antennal style about half as long as 3rd segment. Palpi black, with a long black terminal bristle as long as palpus. Proboscis short, labrum not very much longer than head is high. Thorax shining black when viewed from above, from other angles at most very finely dark grey pollinose, even pleura somewhat subshining. All thoracic hairs and bristles black, large marginal bristles in full number, but only 2 long scutellar bristles. Acrostichal and dorsocentral bristles biserial, not very long, hair-like, diverging anteriorly, on posterior half of mesonotum acrostichals absent and dorsocentral bristles uniserial, becoming longer and bristle-like, posterior 2 or 3 pairs equal to other strong marginal bristles. All hairs on prothorax, and metapleural fan, black. Wings clear, costa, subcosta and radial veins blackish-brown, otherwise veins pale and rather indistinct, anal vein abbreviated, not reaching

wing-margin. Squamae greyish with black fringes, halteres blackish. Legs black to blackish-brown, rather long and slender in comparison with species of the E. chioptera-group, especially mid legs slender. Fore basitarsus small, as deep as the slender tibia, and short-

er than next two tarsal segments together, covered with short hairs. Hind tibiae dilated towards tip and basitarsus distinctly swollen, at least as stout as

tibia at tip, dorsally with long black bristles and with an outstanding long bristle at middle. Fore femora without distinct bristles, for tibiae with posterodorsal dense hairing at most as long as tibia is deep, posteriorly with short, soft, dense pubescence. Mid femora with a double row of rather dense bristly-hairs ventrally, hardly as long as femur is deep, near base both antero- and postero-

215 Figs 214-217. Male genitalia of Empis (s.str.) aestiva Loew (CZ, Bohemia, Jistebnice).—214: postabdomen; 215: lamellae in

lateral view; 216: dorsal lamella in anterodorsal view, lateral lamella dotted; 217: aedeagus with hypandrium. Scale: 0.2 mm.

dorsally with 2 or 3 very long bristly-hairs, and with several not very long outstanding bristles dorsally near

base,

otherwise

dorsum

with

shorter

ad-

vein abbreviated). Legs subshining blackishbrown, rather slender, and fore legs practically bare. Posterior four femora and hind tibiae broadly pennate on both sides, mid femora beneath on apical half only and hind tibiae only on basal half; hind tibiae with two additional diverging bristles dorsally at tip. Mid tibiae dorsally with only short flattened hairs, ventrally fringed. Abdomen almost shining black, covered with sparse, minute black hairs.

pressed bristly-hairs. Mid tibiae above with one or two very long, rather anterodorsal bristles at base and in basal third (very rarely 3 bristles), about 3 times as long as tibia is deep, apical bristle very short, and ventrally tibiae clothed with only short reclinate bristly-hairs. Hind legs covered with rather long but fine bristly-hairs, especially on femora ventrally and on tibiae dorsally, latter with several stronger bristles intermixed which are also present on dorsum of basitarsus. Terminal circlets of bristles on all tarsi unusually short. Abdomen subshining black, pubescence black, longer at sides of basal segments. Genitalia with lateral lamellae small, rectangular and_ blunttipped, covered with minute dark hairs, the slender, apically pointed and spinose dorsal lamellae concealed within lateral lamellae. Aedeagus very narrowed apically, thin at tip, and often clearly visible above lamellae, the broadened base only narrowly shielded by bare ventral lamella. Length: body 2.5-3.5 mm, wing 2.6-3.4 mm. 2. Wings brownish leaving tip and posterior

E. aestiva is a characteristic small black species with black halteres and black abdominal pubescence, abbreviated anal vein, male with mid tibiae with only two long dorsal bristles in basal third, and female with strongly pennate legs. E. aestiva is more closely related to E. (Coptophlebia) vitripennis than to any other species of the E. chiopteragroup. I have seen a male in the Lund University Museum collections collected by R. Danielsson on 1 July 1989 in Smaland (Torsas, Siggesbo) with an abbreviated vein M, on both wings. In such a case, the different bristling on the legs is the only useful diagnostic character, especially the small preapical dorsal bristle on mid tibia. A lectotype for E. aesti-

margin clear, all veins distinct, brown

va was designated by Syrovatka (1991).

(but anal

Length: body 2.3-3.5 mm, wing 2.6-3.4 mm.

127

Distribution. Very common in Denmark and southern Sweden, north to Vg. and Og., also on Oland and Gotland; not recorded from Finland; in Norway approximately to 60° N, to AK and HOy. More than 650 specimens have been examined (395 6 and 273 2), on.dates from 17 June to 23 August, but mainly in July. - From Great Britain and the Baltic coast to the central areas of Europe; known to me in the south only from Austria and Slovenia, but in the Palaearctic Collection in Helsinki there is a long series of obviously conspecific specimens collected by R. Frey on 22 to 24 May 1938 on the Azores (S. Miguel Furnas), recorded by Frey (1945). Collin (1961) recorded E. aestiva as a very common species throughout Great Britain over along period from the beginning of May (!) to 23 September, but I have records from temperate Europe only from 3 June to 21 August. A typical summer species, everywhere mainly in July.

Biology. The swarming and mating behaviour remains unknown but the adults are very common on flowers, especially on Chrysanthemum and Cirsium, preferring marshy biotopes, shaded places and forest margins. The unusually strongly pennate legs of the female suggest obligatory mating in aerial aggregations.

46. Empis s.str. praevia Collin, 1927 Figs 128-129, 218-222. Empis praevia Collin, 1927:23.

Small (about 3 mm) subshining black species with black halteres, abdominal pubescence black and anal vein incomplete. Small thoracic hairs (acrostichal and dorsocentral) very long, hair-like. Fore tibiae in male with long bristles above and tarsi with circlets of very long terminal bristles, female legs without pennation or flattened hairs. 3d. Eyes with upper ommatidia very enlarged; occiput blackish, covered with long, forwardlycurved, densely set black bristles, becoming shorter and finer below but still black. Antennae with 3rd segment rather elongate, style more than half length of 3rd segment. Palpi black, covered with a few long black bristly-hairs at tip. Labrum rather short, only slightly longer than head is high. Thorax mainly shining black when viewed from above, thinly dark grey to brownish-grey pollinose in anterior and posterior views. All hairs and bristles black and conspicuously long, but rather thin, including a humeral, 2 posthumerals, 3 notopleurals, a supra-alar, a longer postalar, and usually only 2 stronger scutellar bristles. Acrostichal bris-

128

tles narrowly 2-serial, dorsocentrals irregularly 3serial, all hair-like, long and erect, distinctly longer than antennal style or the space between acrostichal and dorsocentral bristles, latter becoming uniserial behind and as long as other marginal bristles. Sides of prosternum and episterna with numerous long black hair-like bristles, pronotum with a row of shorter stouter bristles. Metapleural fan long, black. Wings clear with brown veins, only the upper two veins from discal cell (M, and M,) pale and inconspicuous, a distinct brownish costal stigma, and a long black costal bristle. Anal vein abbreviated. Squamae hyaline, with long black ciliae, halteres blackish. Legs black to brownish-black, finely pollinose and almost subshining, rather long and slender, only hind tibiae dilated towards tip and hind basitarsus swollen, as deep as tibia at tip, next two segments also stouter. Fore femora with short hairs, fore tibiae with soft dense brownish pubescence posteriorly, posterodorsally with numerous long bristly-hairs that are much longer than tibia is deep, tarsi slender and with rather short hairs, but

conspicuous long bristles in apical circlets, the bristles at least as long as the following segment is long, those on 3rd segment much longer (Fig. 129). Mid femora ventrally with two rows of long thin bristles at base, those in posterior row the longest (about 3 times as long as femur is deep), becoming shorter and more densely set towards tip. Mid tibiae with 2 to 4 very long anterodorsal bristles on basal half, some shorter bristles posterodorsally (4-6 bristles about twice as long as tibia is deep spread over its whole length), ventrally with numerous shorter bristles (about as long as tibia is deep) arranged in two rows, and a long anterodorsal preapical bristle, also present on basitarsus; circlets of terminal bristles on following tarsal segments shorter. Hind legs covered with long but fine bristles, longer on femora anterodorsally and anteroventrally towards tip, about 5 dorsal pairs on tibiae and 3 similar pairs on dorsum of basitarsus longer than tibia or basitarsus is deep; ventral bristling of basitarsus shorter, as long as tarsus is deep. Abdomen shining black, although dulled by fine greyish pollen from some points-of-view, pubescence black, short on dorsum, longer and denser at sides of anterior segments, venter almost bare. Genitalia very conspicuous because of the long narrow lateral lamellae pointing upwards, dorsal lamellae small and concealed, consequently aedeagus clearly visible, shielded at base by the enlarged polished ventral lamella as far as the swelling-like broadening at middle, tip of aedeagus

Figs 218-222. Male genitalia of Empis (s.str.) praevia Collin (GB, England, Surrey: Bookham). — 218: postabdomen; 219: lamellae in lateral view; 220: dorsal lamella in anterodorsal view, lateral lamella dotted; 221: tip of hypandrium from the inner side; 222: aedeagus. Scale: 0.2 mm.

slightly recurved, not narrowed and thin as in E. aestiva or E. levis. Length: body 2.6-3.2 mm, wing 3-3.2 mm. 2. Not studied in detail by me, but I have seen females in the London Natural History Museum that agree with the short diagnosis given by Collin (1961). Thoracic bristling longer than in females of related species and bristling of legs very distinctive, in general agreeing with that of male but shorter, without pennation or flattened hairs. Wings almost clear, only very faintly brownish. E. praevia needs special comparison only with E. levis; however, males of E. levis have quite dif-

ferently shaped genitalia and tarsi without long preapical circlets, and females have the legs practically bare.

Distribution. A western European species distributed along the Atlantic coast from France northwards to England. Although not recorded from Belgium

and

the Netherlands,

it should

occur

there, and may also be found in Denmark or southern Sweden. My records are only from May, as given by Collin (1961) and as are specimens collected by A.C. Pont, on 25 May 1969 at Bookham Com~ mon (Surrey).

47. Empis s.str. levis Loew, 1873 Figs 130, 223-226.

Empis levis Loew, 1873:40. Empis laevis: authors (error, misspelling).

Very small (2-3 mm), rather shining black species with black halteres and black abdominal pubescence. Resembling EF. praevia but male fore tarsi without apical circlets of long bristles, and female legs almost bare. 6. Eyes with upper ommatidia very enlarged above the deep antennal excision. Occiput subshining black, not dull grey from any point-ofview, covered with not very numerous black bristly-hairs even below neck. Antennae with 3rd segment elongate, style not very much shorter; palpi black, at tip with one long and several shorter black bristly-hairs. Labrum blackish, short, only slightly longer than head is high. Thorax subshining black when viewed from above, finely dark grey to brownish grey pollinose when viewed from in front and from behind, all hairs and bristles black. Large marginal bristles in full number, long and coarse, acrostichal bristles small, narrowly biserial and hair-like, dorsocen-

trals distinctly longer, obviously alternating in one row at least anteriorly and behind, but anterior

129

224 Figs 223-226. Male genitalia of Empis (s.str.) levis Loew (CZ, Bohemia, Praha-Radotin).— 223: postabdomen; 224: lamel-

lae in lateral view; 225: dorsal lamella in anterodorsal view, lateral lamella dotted; 226: aedeagus. Scale: 0.2 mm.

bristles shorter than in E. praevia, scarcely as long as antennal style, ending posteriorly in several long prescutellar pairs; all bristles shorter than in E. praevia but also less numerous and more widely spaced. Four scutellar bristles, outer pair rather long, more than half as long as inner pair, sometimes with an additional minute hair between them. All hairs on prothorax black, sides of pro-

sternum with one or two longer bristly-hairs and several much smaller ones, metapleural fan black. Wings very clear, iridescent, costa and radial veins blackish, other veins fine but darkened and clearly distinct. No costal bristle, costal stigma faintly brownish, anal vein abbreviated. Axillary excision clearly right-angled. Squamae dirty grey and with long black fringes, halteres black. Legs rather long and slender, hind tibiae dilated apically and basitarsus swollen, longer than the following three tarsal segments combined. Fore femora with short hairs except for several longer anteroventral bristles at tip, fore tibiae with 3 long

black dorsal bristles much longer than tibia is deep, posterodorsally with about four regularly spaced slightly shorter bristles, and with soft, dense, somewhat brownish pubescence posteriorly. Fore tarsi

slender and covered with only short adpressed bristly-hairs, no terminal circlets. Mid femora with unusually strong black ventral bristles in 4 to 5 diverging pairs on basal half, intermixed with shorter finer bristly-hairs that are also present on apical

130

half of femur. Mid tibiae usually with 2 very long anterodorsal bristles in basal third, followed by somewhat shorter bristles arranged more posterodorsally, and another anterodorsal preapical bristle, all intermixed with dense shorter dorsal hairing, ventrally with dense reclinate bristles about as long as tibia is deep, arranged in two rows. Hind femora anteroventrally towards tip with fine bristles about as long as femur is deep, and a single anterodorsal bristly-hair in apical quarter. Hind tibiae rather densely long pubescent above, with about 4 pairs of long diverging dorsal bristles longer than tibia is deep, ventrally with short adpressed bristling; hind basitarsus with short bristles above, one longer bristle at middle and at tip not longer than basitarsus is deep, ventrally short spinose. Tarsi of posterior two pairs without long bristles in apical circlets. Abdomen subshining black, very finely light grey pollinose in some lights, and in dried specimens very characteristically downcurved in apical half like a sickle. Pubescence black, longer and dense at sides, sterna translucent brownish and al-

most bare. Genitalia with small ventral lamella shielding only extreme base of aedeagus, lateral lamellae short black pubescent, pointed apically, dorsal lamellae spinose and practically concealed. Aedeagus polished yellowish-brown, very narrowed beyond a bulbous swelling, the very slender

tip free and pointing backwards above apex of abdomen. Length: body 2.3-2.6 mm, wing 2.6-3 mm. ?. Bristling on thorax as in male, abdominal pubescence shorter and sparse. Wings clouded brown, distinctly darker than in related species,

only extreme apex slightly lighter. Legs rather long and slender, and unusually bare: femora practically without hairs, finely greyish pollinose, and all tibiae posterodorsally with a row of minute dark hairs. Hind tibiae with one or two small bristles above at tip, and with similar small bristles also on hind basitarsus above at middle and at tip, and all

basitarsi with very short dense bristles beneath. Length: body 2-3 mm, wing 2.5-3.3 mm. E. levis needs to be compared only with E. praevia so far as the northern and Scandinavian faunas

All the main diagnostic characters as in Empis s.str.: eyes meeting on frons in males with upper ommatidia very enlarged; short 3rd antennal segment with rather long style; very long slender proboscis with narrow sclerotised labella at tip; similarly bristled thorax and legs, with frequent pennation in females; and male genitalia with the same ground-plan structure. Wings clear, milky-white or brown clouded as in Empis s.str., with the main characters the same including a broad obtuse radial fork and discal cell truncate, but the upper vein (M,) from discal cell abbreviated, not reaching the wing-margin. However, there are several groups of non-Palaearctic species with two upper veins from the discal cell abbreviated (veins M, and M,), as in Kritempis, which are generally classified within Coptophlebia (see Smith 1971).

are concerned, as E. aestiva has only short dorsal

bristles on fore tibiae and less bristled mid tibiae, and the legs in females are broadly pennate. I have studied the types of E. levis, and a lectotype was designated by Syrovatka (1991). Distribution. FE. levis was described from the Romanian Carpathians (Baile Herculane), and has not been correctly recognised until now: it has generally been misidentified in central Europe as the closely related FE. pusio Egger, even as recently as the Palaearctic Catalogue (Chvala and Wagner 1989). However, it has been found to be very common locally in the lowlands of central Europe, northwards to the Netherlands where it seems to be very common in suitable biotopes such as Wrakelberg (Mal.Val). Not recorded from England, but its occurrence in Denmark or Sweden seems likely. My records are from 26 May to 2 July; a typical spring species, especially common in June.

Subgenus Coptophlebia Bezzi, 1909 Coptophlebia Bezzi, 1909:100 (as subgenus of Empis). ~Type-species: Empis hyalipennis Fallén, 1816

(orig. des.). Small (about 2.5-3.5 mm, with larger species rarely up to 5 mm long), black, more or less greyish dusted species, remarkably similar to species of Empis s.str., but the upper vein from discal cell (vein M,) abbreviated and not reaching the wingmargin.

Phylogeny and classification. There are no reasonable grounds other than practical convenience for separating Coptophlebia subgenerically from Empis s.str. The Palaearctic Coptophlebia species, as that group has been generally accepted in recent years, are an unnatural artificial group of species sharing one common character, the abbreviated vein M,, and are treated as a separate group for practical purposes only. As discussed in the section on phylogeny in the Introduction, I am aware of that the ‘subgeneric’ classification presented here is a purely practical one and follows the long-established division of Empis, as not better alternative will be available until the world fauna of Empis and allied genera has been thoroughly studied. However, it is also clear that the majority of Coptophlebia species, together with Empis s.str., form a monophyletic group within Empis, and that Rhamphomyia subgenus Aclonempis should also be included in this group. When describing Coptophlebia, Bezzi (1909) designated Empis hyalipennis Fallén as type-species, but from many points of view (absence of acrostichal bristles, structure of male genitalia) this is undoubtedly very different from most other ‘Coptophlebia’. However, current interpretation of this ‘subgenus’ agrees with Bezzi’s (1909) original concept as his diagnosis agrees in all details with Empis s.str., except for ‘vierte und gew6hnlich auch die sechste Langsader abgekiirzt’. Even the other characters given in his diagnosis, such as ‘acrostichals biserial, only rarely absent’ and ‘genitalia small, open or closed’, clearly show that he

did not base this subgenus on E. hyalipennis and its allies alone. As stated by Collin (1961), ‘it might reasonably be argued that the absence of acrostichals is a character of greater fundamental impor-

131

Fig. 227. Empis (Coptophlebia) hyalipennis Fallén, male. Total length: 3-3.3 mm (A. Veltman del.).

tance than a small difference in the venation’, and in this case the name Coptophlebia should be used only for asmall group of three species, E. hyalipennis Fallén, E. leptomorion Bezzi and E. dasychira Mik. I agree with Collin that the convenient character of the abbreviated vein M, and the descriptive name of Coptophlebia used in its present sense would appear to be the most useful proposition (Collin 1961: 542). The current concept of Coptophlebia actually includes several natural groups of species. Apart from the E. hyalipennis-group, E. vitripennis and

cell’, and further examples have been cited here

its allies are closely related to E. s.str. aestiva, anda

under several Empis s.str. species, for example E. chioptera, E. albopilosa and E. aestiva; the latter species causes particular difficulties because of its close resemblance to E. (Coptophlebia) vitripen-

group of species close to E. albinervis is more closely related to E. s.str. chioptera or E. caudatula than to any other Coptophlebia species. The abbreviated vein M, is the only diagnostic character

nis. On the other hand, I have seen several specimens of the central European E. (Coptophlebia) abbreviata Loew with vein M, complete on both wings and reaching the wing-margin as in Empis

132 /

unifying the smaller-sized Empis species into one unit called Coptophlebia. However, although the abbreviated vein M, seems to be a useful character for separating Coptophlebia, it is also anot uncommon venation aberration even in the subgenus Empis s.str. Collin (1961) mentioned several examples of the ‘Coptophlebid aberration of the upper vein from discal

Figs 228-230. Male mid leg in anterior view of Empis subgenus Coptophlebia. — 228: E. albinervis; 229: E. vitripennis; 230: E. volucris. Scale: 0.5 mm.

s.str. An extra cross-vein connecting the upper two veins M, and M, behind the discal cell, forming another small square-shaped cell, is another not uncommon venation variation of Coptophlebia species. E. hyalipennis and its allies are undoubtedly the most specialised forms within Coptophlebia, characterised by a set of apomorphous characters in the male genitalia, which are of very complicated structure, and are also the only species-group within Empis s.str. and Coptophlebia in the Palaearctic region where the acrostichal bristles have been lost.

do not fit into any recent ‘subgenus’, especially in tropical Africa. 19 species of Coptophlebia have been described from the Palaearctic region: 15 are European or at least eastern

Mediterranean,

and 4 species are

known from Japan. Nine species occur in temperate and cool Europe, and six of these are treated here as they are distributed northwards at least to the coasts of the North and Baltic Seas, approximately to 53° N. However, this number of known

Distribution. As currently defined, Coptophlebia is almost worldwide in distribution, known with certainty fromthe. Holarctic, Afrotropical and Oriental regions. However, many taxa assigned to

Coptophlebia species is far from final as I have seen several new and still undescribed species from the central and southern parts of Europe, including the Caucasus. On the other hand, I did not find any Coptophlebia in Middle Asia, in the area between the Caspian Sea and China, although Empis s.str. is richly represented there by a large number of species that are often very abundant in the

Coptophlebia from subtropical and tropical areas

mountainous areas of Uzbekistan, Turkmenistan,

are very different from the diagnosis given above and should be included in other ‘subgenera’; many may even represent quite distinctive groups that

Tadzhikistan and Kazakhstan.

Biology. The adults are often found on flowers and are sometimes very common on Cirsium or Chry-

133

Figs 231-233. Empis (Coptophlebia) pilimana Loew. — 231: male mid leg in anterior view; 232: male hind leg in anterior view; 233: thoracic prosternum, diagrammatic. Scale: 0.5 mm, 0.2 mm in Fig. 233.

santhemum, often with several specimens of both sexes sucking at one flower. Precise data on the epigamic behaviour, swarming and mating activity are not available, but males have often been caught with prey (another insect) and it is believed that aerial swarming and mating in species-specific synorchesic aggregations are a common habit, as in other Empis s.str. species. The majority of Coptophlebia species are typical species of summer occurrence, often on the wing over a long period; none is a spring species, as is obligatory for Empis s.str. Some of them, for example E. vitripennis or E. hyalipennis, belong to a very small group of Empis species with a very late summer and even autumn flight-period.

Key to Coptophlebia species 1 —

_Halteres yellowish, abdominal pubescence paledi...% » ake ber eetule Nw Halteres blackish, abdominal pubescence generally black, rarely (E. vitripennis) pale

2(1) Thoracic bristles pale, acrostichals absent. Anal vein complete, reaching wing-margin. Male genitalia with dorsal lamellae unusually

spinose, hind femora in female pennate. About 3-3.5 mm long ... 48. E. hyalipennis — Thoracic bristles black, acrostichals present. Anal vein incomplete, ending before wingMarin qin 5SAe eee eee 3) 3(2) Mesonotum subshining black when viewed from above. Wings milky-white in d , creamywhite in 2. d: 7th tergum simple. @: legs not pennate, with only fine fringes on hind femur at tip. Smaller, about 25-3mm ........... SEORL EMS FISTS 49. E. impennis — Mesonotum duller grey. Wings clear in d, yellowish to yellowish-brown in ° (with a tendency for the anal vein to be complete). d: 7th tergum with a dorsal projection. 2: hind femora and tibiae pennate. Larger, about 33 Somme eee thee 50. E. albinervis 4(1) Sides of prosternum with a tuft of about 5S to 8 short but strong black spinose bristles. Wings clear, veins dark except for the fine medial

veins. d: mid tibia and basitarsus with very long dorsal bristly-hairs, fore basitarsus densely long pubescent posterodorsally. @: legs almost bare, posterior four femora with fine anteroventral and anterodorsal flattened haircwet: Gioieie i SR aus 53. E. pilimana —

Sides of prosternum with only a few, fine, MM te dais: pa deye teee hee sits itiees 5)

5(4) Anal vein incomplete. Abdominal pubescence pale, or at least brownish on basal segments in male. Wings clear in ¢, brownish in @, all veins brown and distinct. ¢: mid tibia

with 1-4 anterodorsal bristles on basal third to half, and another one at tip; aedeagus yellowish, stout at base. @: fore tibiae and basi-

—

tarsus not pennate, mid and hind legs with pennation. 2 scutellar bristles ............. tection: Getewieeris wasn 51. E. vitripennis Anal vein usually reaching wing-margin, though faint at middle. Abdominal pubescence brown to blackish. Wings clear in both sexes, veins M, and M, whitish and inconspicuous. 3d: mid tibia with 5-6 long bristles along the whole length dorsally; aedeagus blackish, slender at base. 2: legs pennate, also on fore tibiae and basitarsi above. 4 scutellar bristles 13 Of ORC OEE Ae eee 52. E. volucris

48. Empis (Coptophlebia) hyalipennis Fallén, 1816 Figs 227, 234-238. Empis hyalipennis Fallén, 1816:21.

Greyish, about 3-3.5 mm, with incomplete vein M, and very long labrum, about 2.5 times as long as head is high. All thoracic bristles pale, acrostichals absent. Male genitalia very complex, hind femora in female pennate. 3. Eyes meeting on frons with upper ommatidia very enlarged. Occiput dull dark grey, covered with long, fine, bristly-hairs that are black above and whitish on more than lower half. Antennae black, basal segment not very much longer than 2nd, 3rd segment rather short, the somewhat stout ‘style about two-thirds as long as 3rd segment. Proboscis very long, resembling that of E. pennipes, labrum much more than twice as long as head is high. 3 Thorax blackish-grey or almost subshining when viewed from above, but much paler and uniformly light grey dusted in anterior view. All hairs and bristles pale, yellowish. Acrostichals absent, dorsocentrals uniserial anteriorly, biserial at about middle and diverging, rather long and hair-like, as

long as antennal style, ending in one very long prescutellar pair that is equal in length to the other large marginal bristles. Usually one pair of long scutellar bristles with a very small hair each side. All prothoracic hairs fine and pale, including somewhat smaller hairs in a row on pronotum. Metapleural fan long, pale yellow. Wings completely clear, veins dark, but vein M, abbreviated and like M, fine and much less distinct, as well as the complete anal vein. A distinct light brown costal stigma; no costal bristle. Squamae and halteres pale yellow. Legs long and slender, black, but more or less translucent brownish, only finely greyish pollinose and subshining (except the dull coxae). Fore basitarsus elongate and slightly swollen, broader than tip of tibia, the following tarsal segments very slender. Hind tibiae slightly dilated apically, basitarsus as deep as tibia at its broadest part, and about as long as the next three tarsal segments combined, latter also partly dilated, not as slender as on front pair. Posterior four femora with very long fine pale outstanding hairs antero- and posteroventrally, hind femora also posteriorly, dorsum with short reclinate hairs. Fore tibiae with short dense pubescence, longer posterodorsally, mid tibiae with two very long black bristles anterodorsally on basal half, one equally long ventral one in basal third, often another shorter one at about middle, and apical half of tibia ventrally with a row of rather adpressed black bristles about as long as tibia is deep (much longer in E. leptomorion). All tibiae with short preapical bristles, but mid basitarsus with an unusual long black anterior bristle near base and another at tip. Hind tibiae finely bristled, the longest bristly-hairs in anterodorsal row, and similar equal single bristles at middle posterodorsally and posteroventrally. Tarsal segments on front pair with rather long circlets of terminal bristles, but

unlike E. leptomorion the bristles are-not longer than the next segment. Hind basitarsus with a tendency to be spinose ventrally. Abdomen lighter grey than thorax, rather densely dusted and rather long pubescent especially at sides of anterior segments, and with long straggling bristly-hairs on venter; all hairs pale yellowish. Genitalia very complex, dorsal lamella and the upper broader part of lateral lamella at tip short black spinose, the spines distinctly flattened, and the lower very slender and pointed process of lateral lamella with 2 very long bristles dorsally before tip. 8th sternum enlarged, cornet-like, shielding genitalia from below, aedeagus short and very thin apically. Length: body 3-3.3 mm, wing 3.2-3.6 mm.

135

2. Very much like male but frons broad, as wide as face, dark grey dusted. Wings faintly brownishgrey clouded, not clear as in male. Legs with short pubescence or fine bristling, but hind femora pennate on the whole length-above, the pennation becoming progressively shorter towards tip, longer pennate ventrally but on apical half only, both flattened bristles about as long as femur is deep. Hind tibiae very slender on basal half, slightly widening apically, everywhere with only short dark pubescence. Abdomen dull light grey, covered with short pale hairs. Length: body 2.8-3.8 mm, wing 3.3-3.8mm. With its yellowsih thoracic bristles, absent acrostichals, long slender legs, very long proboscis and unusually complex male genitalia, E. hyalipennis can hardly be confused with any other northern Coptophlebia species. The closely related central European E. leptomorion Bezzi and E. dasychira Mik have longer bristles in general, with different-

ly bristled and shaped legs in male, female legs not pennate, and with slight differences in the male genitalia.

Note. For synonymy, see Collin (1961:551). Empis proxima Meigen, 1838, described from Belgium (Liége), is hardly a Coptophlebia species and has been listed in the Palaearctic Catalogue (Chvala and Wagner 1989) among the unrecognised species of Empis s.lato. In 1985 I revised the type specimens in the Fallén Collection in Stockholm, and can confirm Collin’s (1961) notes: a male and two females have been correctly recognised and are all conspecific. A lectotype was not designated but, if necessary, the male should be selected. Distribution. Throughout Scandinavia, but rather rare everywhere; I have examined only 24 ¢ and 24 2 over a wide area from the south to approximately 68° N, in Norway to TRi, in Sweden to Nb.

Figs 234-238. Male genitalia of E. (Coptophlebia) hyalipennis Fallén (CZ, Bohemia, Jistebnice). — 234: postabdomen; 235: 7th and 8th tergum in dorsal view; 236: lateral lamella; 237: dorsal lamella in anterodorsal view; 238: aedeagal complex. Scale: 0.2 mm.

136

and Lu.Lpm., in Finland only to Tb and Sb. There are further specimens from Norway in T. Jonassen’s Collection at Sjernargy (included in the distribution table through the courtesy of T. Jonas-

in notopleural depression, also on pronotum, at sides of prosternum and on episterna. Metapleural fan short but dense, whitish. Wings milky-white, only costa dark, subcosta

sen), and I have seena further2 d and5

labelled

and radial veins yellowish-brown, otherwise veins

‘Norwegen’ in the Becker Collection in Berlin. In July and August, on dates ranging from 15 July to 30 August, in July mainly in the south, but there isa single female taken by Tord Tjeder in Sweden (Dir., Leksund) on 15 September 1971. A typical late summer species. — Widely distributed but ev-

whitish and inconspicuous. Vein M, and anal vein abbreviated, radial fork very ‘open’, vein R, almost at right-angles to vein R,. Squamae and halteres pale yellow.

erywhere uncommon,

from Scotland and Wales,

and NW Russia through the central parts of Europe south to the submontane and montane regions of Spain and Romania. In temperate Europe in July and August; in rather shaded forest biotopes at higher altitudes, together with the closely related but undoubtedly much more common E. leptomorion Bezzi.

49. Empis (Coptophlebia) impennis Strobl, 1902 Figs 239-242. Empis impennis Strobl, 1902:470, form of E. albinervis). Empis melaena Bezzi, 1908:394.

1904:529

(as

Very small (2.3-3.1 mm) subshining black species with milky-white wings in male, creamy-white in female, veins M, and Cu abbreviated.

Thoracic

bristles black but metapleural fan and abdominal pubescence whitish, halteres pale. Legs short, simple, not distinctly bristled in male and not pennate in female. 3. Upper ommatidia very enlarged, occiput dark grey, covered with short fine dark hair-like bristles which become longer and whitish below. Antennae black, basal two segments small, about as long as deep, 3rd segment broad at base, pointed apically, scarcely 3 times as long as deep, but style not very much shorter. Palpi very small, black, devoid of hairs; labrum about one-and-a-half times

as long as head is high, blackish and very slender. Thorax almost shining black when viewed from above, finely greyish pollinose in anterior view, pleura more dulled by greyish dust. Large marginal bristles black and not very long (a humeral, 2-3 notopleurals, a postalar and 2 scutellars); acrostichal and dorsocentral bristles very small and fine, blackish to brownish, former almost uniserial, al-

ternating in one row and distinctly diverging, like the irregularly bi- to triserial dorsocentral bristles ending in one prescutellar pair as long as scutellar bristles. Some fine pale hairs on humeri, anteriorly

Legs black to brownish-black, short and simple,

all pubescence and bristling short and fine. Only mid femora beneath with a double row of widely spaced black bristles, longer than femur is deep; similar ventral bristles on hind femora much shorter and finer, including a single anterodorsal bristle before tip. Mid tibiae with two diverging pairs of dark bristles (slightly longer than tibia is deep) in basal third and beyond middle, and another pair at tip; ventrally with several similar bristles at about middle becoming shorter towards tips. Dorsal bristly-hairs on hind tibiae short, barely as long as tibia is deep. Tarsi with short circlets of black preapical bristles, those on front pair the longest. Abdomen more greyish pollinose than thorax, though still somewhat subshining. Pubescence pale, longer at sides of anterior segments and on sterna. Genitalia small, dorsal lamellae deeply excised and, like the small, somewhat triangularshaped lateral lamellae, covered

with fine black

bristles posteriorly. Ventral lamella small, blacker, partly shielding the short and rather stout yellowish aedeagus. Length: body 2.3-2.8 mm, wing 2.8-3.0 mm. 2. Frons dark grey dusted, as wide as face, clypeus polished black. Thorax with all hairs and bristles shorter and finer, wings creamy-white, not at all clear, with all veins brown and much more dis-

tinct. Legs short pubescent, no distinct bristles or pennate hairs, hind femora beneath almost straight, dorsally slightly convex, and clothed with small black reclinate fringes towards tip above, posteroventrally similarly fringed towards tip. Abdomen subshining, covered with sparse, minute, uniform pale hairs. Length: body 2.3-3.1 mm, wing 2.5-3.0 mm. E. impennis is the smallest Coptophlebia species in Europe, known for a long time as E. melaena Bezzi thanks to Collin’s (1927) brief re-description and separation of it from the closely related E. albinervis. Both these species, having black thoracic bristles, pale metapleural fan, whitish abdominal pubescence, yellowish halteres and incomplete anal vein, have no other close allies in the European fauna. Synonymy. The species was first recognised by

1S:

Figs 239-242. Male genitalia of E. (Coptophlebia) impennis Strobl (NL, Kunrade, Kunder Berg). —239: postabdomen; 240: lamellae in lateral view; 241: dorsal lamella in anterodorsal view; 242: aedeagus with hypandrium. Scale: 0.2 mm.

Strobl (1902), who distinguished two forms (impennis and pennata) of Empis albinervis, based on the different bristling of the legs in females. Since Meigen’s name E. albinervis was restricted by Collin (1961) to the species with pennate legs in female, the smaller species with legs not pennate in female must receive the first available name, which is Empis impennis. The synonymy Empis impennis Strobl, 1902 = Empis melaena Bezzi, 1908, was published in the Palaearctic Catalogue (Chvala and Wagner 1989), but the reason for it is given here for the first time.

Distribution. E. impennis has been known for a long time as E. melaena, and all the published records have been under the latter name. It has not yet been found in Scandinavia but it should occur there, as it is known from England, the Netherlands, and through Germany and temperate Europe south to southern France and Croatia (Split). A spring and early summer species, known to me on dates ranging from 3 June to 29 July, but mainly

in June, and I have seen specimens labelled as August but without exact day.

50. Empis (Coptophlebia) albinervis Meigen, 1822 Figs 228, 243-245.

Empis albinervis Meigen, 1822:26. Empis pennata Strobl, 1902:470, 1904:529 (as form of E. albinervis), not Schrank, 1781.

Somewhat larger and greyer than E. impennis, halteres yellowish, and abdominal pubescence rather long, whitish. Tarsal segments on front and hind legs slightly dilated. Male with an unusual projection on 7th tergum above, legs in female pennate. 3. Eyes with upper ommatidia anteriorly very conspicuously enlarged, occiput grey dusted, with small, fine, dark hairs above that become longer

and whitish below. Antennae black, basal segments short, with a tendency to be brownish; 3rd segment rather short and broad at base, onion-

shaped and pointed apically, style very slender and

138

nearly as long as 3rd segment. Palpi small, yellowish brown towards tip or tip almost whitish. Proboscis rather short, labrum brownish-black, hardly

1.5 times as long as head is high. Thorax rather densely dark grey dusted. Mesonotum lighter grey dusted in anterior view, concolorous with pleura. Marginal bristles black, thin and not very long, in full number including a posthumeral and one or two supra-alars, scutellum with 2 bristles. Acrostichal bristles narrowly biserial, diverging, black and very small, hair-like, dorsocentrals slightly longer but equally fine, biserial but more widely spaced, ending in 3 longer pairs and a log final prescutellar pair. Humeri, notopleural depression, pronotum, sides of prosternum and episterna clothed with fine whitish hairs. Metapleural fan whitish-yellow. Wings clear with anterior veins brown, but veins

M and Cu whitish including discal cell; vein M, and anal vein incomplete, radial fork very open, almost 90°. Squamae and halteres pale yellow. Legs black to brownish-black, finely grey pollinose, slender, but hind tibiae slightly dilated to-

wards tip, and fore and hind tarsi with all segments slightly swollen; fore basitarsus wider than tibia at tip and about as long as the next three segments together, hind metatarsus even longer. Fore legs almost bare, only tibiae with dense longer pubescence posteriorly. Mid femora with long ventral bristly-hairs, those in anteroventral row, and about three similar bristles posteroventrally towards base, at least twice as long as femur is deep; hind femora with similar bristles anteroventrally before tip and anterodorsally (or almost anteriorly) on apical half. Mid tibiae with 3 long anterodorsal bristles placed near base, at middle, and at tip, ventrally on basal half with rather long adpressed bristles in two rows. Hind tibae with several not very long hair-like bristles dorsally, and with a dense, long pubescence ventrally. All tarsal segments with apical circlets of black bristles, especially long on all basitarsi, which are almost spinose beneath, less distinctly so on fore basitarsus; mid basitarsus

with a long black anterodorsal bristle at tip. Abdomen more subshining black, less greyish pollinose than thorax, rather densely whitish pu—st

be !

' 5

i \

Figs 243-245. Male genitalia of E. (Coptophlebia) albinervis Meigen (CZ, Bohemia, Jistebnice).— 243: postabdomen; 244: lamellae in lateral view; 245: aedeagus with hypandrium. Scale: 0.2 mm.

139

bescent, longer at sides and on venter. 7th tergum with a large dorsal projection pointing slightly backwards. Genitalia with both lamellae very small, finely short pubescent, but ventral lamella (hypandrium) long, tubular and polished brown, covering basal half of the rather short yellowish aedeagus. Length: body 2.9-3.5 mm, wing 3.2-3.8 mm. 2. Generally darker grey than male, with all thoracic bristles and abdominal pubescence shorter. Frons not very broad, narrower than face, dark grey. Wings yellowish-brown clouded at least at base and along costal margin, apex of wing and posterior margin broadly clear; veins brownish, much more distinct, anal vein though incomplete often visible at wing-margin. Legs short pennate on both sides on hind femora and tibiae, tibiae ventrally on basal half only, mid femora short pennate on the whole length above but ventrally at tip only, mid tibiae and basitarsus dorsally with flattened bristly-hairs, similar but finer flattened fringes also on fore tibiae and basal two tarsal segments dorsally. Fore tarsi slightly dilated but posterior four tarsi slender, hind femora somewhat laterally compressed. Length: body 2.8-3.5 mm, wing 3-3.4 mm. Resembling E. impennis, but easily distinguished by the larger size, the striking dorsal projection on 7th tergum in male, and pennate legs in female. Further differential characters are the slightly dilated tarsal segments on front and hind legs in male, and the yellowish tips of the palpi. Synonymy. For details of the synonymy, see under E. impennis. The form pennata described by Strobl (1902) for females of E. albinervis with pennate legs is actually the true a/binervis described by Meigen and becomes a junior synonym of Empis albinervis Meigen. Distribution. Rare in Scandinavia, so far found only at two localities in Norway, in AK, Baerum: Ostgya by F. Midtgaard, 6 d and 7 @ (10 June-1 September 1984), and Bg, Rgyken: Hyggen by L.O. Hansen, 4 ¢ and1 2 (June and July 1989), see also Jonassen (1992); and in Skane in Sweden, 1 2 by H. Andersson on 21 July 1985 at Trollenads. A species of southern distribution, its occurrence in Denmark has not been confirmed by recent specimens; one male collected and recorded by Lund-

beck (1910) from Funen (Middelfart, 18 July 1907) has apparently been lost, but the description proves that it was correctly identified by Lundbeck. — Great Britain according to Collin (1961), commoner in the southern half of England, but re-

corded also in Wales and Ireland. Widely distrib-

140

uted in central and southern parts of Europe; known in the south from France including Corsica, Italy, east to Romania. In temperate Europe, and also in England and the Netherlands, over a long period from the beginning of June to the end of August, but mainly in July; in the south (Corsica) as early as May.

51. Empis (Coptophlebia) vitripennis Meigen, 1822 Figs 229, 246-250. Empis vitripennis Meigen, 1822:25. Empis nana Macquart, 1823:161. Empis turbida Meigen, 1838:86.

Moderately large (2.8-3.7 mm) subshining black species with abbreviated vein M, and anal vein, halteres blackish and base of abdomen with pale pubescence. Wings with all veins contrastingly dark, clear in male, brownish in female. Posterior four legs in female pennate. 3 . Occiput dark grey, covered with rather long black bristly-hairs which become greyish below. Antennae black, basal segments very small, 3rd segment broad at base, pointed apically, about 2.5 times as long as deep at base, style very short, barely one-third as long as 3rd segment. Palpi very small, black. Proboscis long, labrum practically black and nearly twice as long as head is high. Thorax almost shining black when viewed from above, uniformly grey pollinose in anterior view, and viewed from behind with three broad black stripes anteriorly along the lines of bristles. Marginal bristles black, in full number and not very prominent, 2 scutellar marginal bristles. Acrostichal and dorsocentral bristles biserial, only indistinctly diverging, rather long, nearly as long as upper occipital bristles but more hair-like, black. Pronotum and prothoracic episterna with black bristly-hairs, those on sides of prosternum paler. Metapleural fan black, bristles whitish at tip. Wings clear with all veins (except anal vein which is in the form of an indistinct fold visible at base only) contrasting dark brown, including abbreviated vein M,. Radial fork very open, vein R, straight and almost upright. Squamae dark dirty grey with very dense long black fringes, halteres blackish. Legs brownish-black in ground-colour, rather long and slender, only hind tibiae dilated apically and hind basitarsus swollen and enlarged, about as long as rest of tarsus, the following tarsal segments almost slender, and fore tarsi not at all dilated. Bristling very much as in E. albinervis, especially

Figs 246-250. Male genitalia of E. (Coptophlebia) vitripennis Meigen (GB, England, Dorset: Studland). — 246: postabdomen; 247: lateral lamella; 248: dorsal lamella in anterodorsal view, lateral lamella dotted; 249: the same in lateral view;

250: aedeagus with hypandrium. Scale: 0.2 mm.

that of mid tibiae and posterior femora of the same ground-plan. Fore femora clothed beneath with short hair-like bristles, similar dense bristling on fore tibiae above, but tibia with denser soft pubescence posteriorly, also present on basitarsus. Posterior four femora with double rows of long fine ventral bristles, much longer than femora are deep, those on mid femur posteroventrally to-

ten brownish and covered with short hairs, otherwise pubescence pale, at least light brownish on basal segments, longer at sides, posteriorly with mainly black hairs; longer black bristles on posterior margin of the enlarged, polished black 8th sternum. Genitalia with lateral lamellae elongate, long ovate and shining black, with upper hind tip produced; narrow upper lamellae closely attached to

wards base, and on hind femur anteriorly and ante-

lateral lamellae. Ventral lamella very small, and

roventrally towards tip, the longest. Mid femora with indistinct whitish pollen beneath. Mid tibiae with usually 2 (rarely up to 4) strong black bristles above and another similarly long bristle at tip, ventrally with two rows of more adpressed black bristly-hairs, longest at about middle, those in anteroventral row longer. Hind tibiae with short adpressed bristling ventrally, with irregularly-placed

consequently the yellowish aedeagus clearly visible on its broadened basal half. Length: body 2.8-3.5 mm, wing 2.8-3.2 mm 2. Obviously more greyish than male, frons dull blackish-grey and very broad, as wide as face. Thoracic bristles slightly shorter, and acrostichal and dorsocentral bristles more diverging. Wings faintly brownish-grey clouded, veins dark brown. Legs

long and short bristles above, also present on basi-

blackish-brown in ground-colour, all femora and

tarsus. Basal two segments on all tarsi with circlets of short preapical bristles, basitarsi more or less spinose beneath, and mid basitarsus with long apical bristle dorsally, not much shorter than similar strong tibial bristles. Abdomen subshining black on terga, sterna of-

also tibiae on posterior two pairs somewhat broadened and laterally compressed, tarsi slender. Posterior four femora and tibiae strongly pennate on both sides, pennation almost as long as corresponding femora and tibiae are deep, hind tibiae beneath pennate on basal half only, and mid tibiae

141

with shorter pennation ventrally at base. Fore legs with short ordinary hairs and bristles, without pennation or flattened hairs, at most tibia and basitarsus finely fringed dorsally. Hind basitarsus with short hairs only. Abdomen subshining, everywhere covered with short, rather sparse fine pale or light brownish hairs. Length: body 2.8-3.7 mm, wing 3-3.3 mm. E. vitripennis belongs to a group of Coptophlebia species with black halteres and only fine hairs at the sides of prosternum (actually prothoracic epimera). Its only ally in northern Europe is E. volucris, but there are two or three further related but still undescribed species in temperate and southern Europe. However, E. vitripennis is also closely related to Empis s.str. aestiva, sharing many features in the bristling of legs in both sexes and structure of the male genitalia. The abbreviated vein M,, although a dubious character in view of its

known variability, some small details in the bristling of legs, and slight differences in the male genitalia are the only available differential features. For synonymy, see Collin (1961:544). Distribution. A species of southern distribution in Scandinavia, known to me from Denmark and southern Sweden (Sk., Hall., Sm.); 48 ¢ and 42 @

have been examined. Jonassen (letter of 21. Dec. 1992) has also found it in southern Norway, In B@,

R¢yken: Kinnartangen, 4 August to 8 September, 4d and1 2, andin VE, Vale: Langgya, 2 August to 1 September 1991, 1 2, in both cases collected in Malaise traps by L.O. Hansen. A typical late summer or even autumn species, occurring in August and September, in Scandinavia ranging from 1 August to 20 Sepbember. — England, Belgium, the Netherlands, central and SW Europe (Spain, France), but unknown from the central and east Mediterranean or SE Europe. The record from the Azores (Frey 1945) needs verification as there are further closely related species in southern Europe. In temperate Europe rarely in July, mainly in August and September as in the north.

52. Empis (Coptophlebia) volucris Wiedemann, 1822 Figs 230, 251-255.

fore tibiae pennate in female. Four scutellar bris-. tles. 3d. Eyes with upper ommatidia very enlarged above. Antennae black with style distinctly longer than in E. vitripennis, about two-thirds as long as 3rd segment. Labrum polished black, slender and rather short, less than 1.5 times as long as head is high. Thorax subshining black, more greyish dusted in anterior and posterior views, unlike E. vitripennis unstriped even when viewed from behind. All hairs and bristles black, dorsocentral bristles more distinctly diverging, and scutellum with a small hair-like bristle each side of the strong inner pair. Metapleural bristles completely black. Wings clear, veins on costal margin (C, Sc, R) dark brown, but other veins including abbreviated

vein M, whitish and inconspicuous, radial fork very broad. Squamae dark grey with blackish margin and densely-placed long black fringes, halteres blackish. Legs rather shining brownish-black, mainly with long black bristles, hind tibiae and hind tarsi dilated and, unlike E. vitripennis, fore basitarsus also slightly swollen. Fore femora practically bare, fore tibiae with a row of posterodorsal bristly-hairs about as long as tibia is deep, and shorter dense pubescence posteriorly. Mid femora besides a double row of long ventral bristles and dense whitish soft pile beneath, with an anterodorsal row of long somewhat reclinate bristles which become longer towards base. Mid tibiae with long bristles, dorsally with 5 to 6 evenly spaced long bristles at least 3 times as long as tibia is deep, the apical bristle the longest, ventrally with a double row of denselyplaced reclinate bristles, longest at middle. Hind femora with short fine bristles beneath, becoming

longer towards tip (but still shorter than femur is deep), several still shorter bristles anterodorsally before tip, the very fine hair-like bristles anteriorly at base the longest. Hind tibiae with short reclinate bristles beneath, dorsally with several pairs of outstanding long bristles, at least twice as long as tibia is deep, intermixed with shorter bristly-hairs, and with similar bristling also on hind basitarsus above. Tarsal segments without circlets of long apical bristles, but basitarsi spinose ventrally, very strongly so on hind legs and, unlike E. vitripennis, mid basi-

Empis volucris Wiedemann in Meigen, 1822:23.

A species with black halteres M,, resembling E. vitripennis robust species, with longer wings clear in both sexes with Aedeagus in male blackish

142

and abbreviated vein but generally a larger bristles everywhere, medial veins whitish. and slender at base,

tarsus obviously more dilated and without a long preapical bristle (Fig. 230). Abdomen shining black, pubescence uniformly black to blackish-brown, not pale, short, longer at sides of anterior segments. Genitalia small, black,

rather finely black pubescent; lateral lamellae short triangular in shape, dorsal lamellae nar-

:

\:AlXY“ \\

253

Se

Ni

Ww

\

SS

a

252

Figs 251-255. Male genitalia of E. (Coptophlebia) volucris Wiedemann (CZ, Bohemia, Bor near Skuteé). — 251: postabdomen; 252: lateral lamella; 253: dorsal lamella in anterodorsal view, lateral lamella dotted; 254: the same in lateral view;

255: aedeagus with hypandrium. Scale: 0.2 mm.

rowed as in E. vitripennis but clearly visible above, not concealed. Aedeagus slender along its whole length, blackish, ventral lamella small and conse-

quently aedeagus clearly visible on its basal half and even at tip above lamellae. Length: body 3.2-3.6 mm, wing 3-3.5 mm. 2. Resembling male but all hairs and bristles shorter on thorax and abdomen, frons as wide as

face. Wings clear as in male, with medial veins whitish and inconspicuous, but anal vein usually reaching margin, though inconspicuous at middle. Posterior four legs with similar pennation to that in E. vitripennis, but hind femora on both sides and hind tibiae dorsally more broadly pennate, and short pennate fringes also on fore tibiae and basi-

tarsi above. Hind tibiae posterodorsally with a row of about 6 to 8 simple bristles, those on basitarsus shorter and placed more dorsally, besides a very long single posterodorsal black bristle at middle. Length: body 3.4-3.8 mm, wing 3.2-3.6 mm. E. volucris is compared in the above description

with the only other related northern species, E. vitripennis. The two species were first correctly separated by Collin (1961:545), who also gave the full synonymy. The southern E. pavesii Bezzi is another closely related species belonging to this

group. Distribution. E. volucris has been found in the southern parts of England (Collin 1961; Allen 1987) and, although it has not yet been recorded from the Netherlands or along the Baltic cost, its

occurrence in Denmark or southern Sweden is to be expected. A widely distributed though overlooked and often misidentified species, rather common in central Europe, known throughout the Mediterranean, from Portugal and Spain in the west to Crete (Kalemelyhia Lasithion, leg. A.Fjeldsi) in the east. Mainly in June and July, in the Mediterranean as early as May.

143

53. Empis (Coptophlebia) pilimana Loew, 1869 Figs 231-233, 256-259. Empis pilimana Loew, 1869:86. Empis holoneura Strobl;1910:65 (as var. of E. pilimana).

Small (2.5—3.2 mm), subshining black species with black halteres, abbreviated vein M,, sides of prothoracic sternum with a tuft of small black spinose bristles, and abdominal pubescence blackish. Legs in male densely clothed with very long woolly hairs, femora fringed in female. 6. Eyes with upper ommatidia not very enlarged, about twice as large above as those below: occiput blackish, covered with uniformly black long bristly-hairs even on lower part below neck. Antennae black, basal segments short, 3rd segment broad at base, abruptly pointed on apical half, about 2.5 times as long as deep at base, style almost as long. Palpi black. Labrum_ blackishbrown, slender, about 1.5 times as long as head is high. Thorax shining black when viewed from above, rather thinly brownish-grey dusted in anterior and posterior views, and then concolorous with grey pleura. All thoracic hairs and bristles black, including small hairs on prothorax and metapleural fan. Large bristles long but thin: a humeral, 2 posthumerals, numerous merging notopleural and supraalar bristles, a postalar, and one pair of scutellars, exceptionally with a minute hair on each side. Acrostichal and dorsocentral bristles long, hairlike, at least as long as antennal style, former narrowly biserial and diverging, latter uniserial anteriorly and posteriorly, ending in 3 long prescutellar pairs, at about middle more broadly biserial and diverging like the acrostichals. Sides of prosternum with a tuft of about 5 to 8 strong, downcurved spinose bristles. Wings clear, veins distinct and almost blackish, except for very whitish veins M,, M, and the veins closing discal cell from above and behind, and the very abbreviated anal vein. Radial fork largely open as usual in Coptophlebia, but axillary excision unusually obtuse, about 90°, although axillary lobe very large. Squamae (with concolorous fringes) and halteres blackish. Legs with very characteristic long woolly pubescence, black to blackish-brown, with black hairs

and bristles. Fore basitarsi slightly swollen, a little wider than tip of tibia, next tarsal segment slightly dilated, not quite slender. Hind tibiae slightly broadened at tip, but basitarsus and next two seg-

144

ments distinctly swollen. Fore femora almost bare except for rather short ventral hairs, fore tibiae with long posterodorsal bristly-hairs (the longest about 3 times as long as tibia is deep), continuing dorsally onto basal three tarsal segments. Mid and hind legs as in Figs 231 and 232, femora with very long and widely-spaced thin bristles ventrally, and both tibiae and tarsi on both sides with long dense hairing. Abdomen almost shining black with longish black hairs at sides and on venter, anterior segments densely pubescent. Genitalia small, lateral lamellae rounded, with two long hairs at tip, dorsal lamellae elongate, duller and microscopically short dark pubescent. Ventral lamella small, cornet-like, shining, and shielding only the extreme base of the not very stout yellowish aedeagus. Length: body 2.8-3.2 mm, wing 3.1-3.5 mm (lectotype body 2.8 mm, wing 3.2 mm). 2. All hairs and bristles on thorax and abdomen shorter, especially acrostichal and dorsocentral bristles very short, both evenly biserial and diverging. Frons dark grey dusted, as wide as face. Wings clear as in male, with large brownish costal stigma and all veins contrasting darker, even the abbreviated vein M,; anal vein very fine, incomplete. Legs rather short but not stout, everywhere with only short hairs and bristles, fore femora almost bare and all tibiae dorsally with very short adpressed hairs. Posterior four femora anterodorsally along the whole length and anteroventrally on apical third to quarter with long flattened hairs (no real pennation), the hairs shorter than femora are deep. Hind tibiae dorsally with a preapical diverging pair of bristles, about as long as tibia is deep, and another single anterodorsal bristle on hind basitarsus at middle. Length: body 2.9-3.1 mm, wing 2.6-3 mm. E. pilimana is the only northern European species with a tuft of black spinose bristles at the sides of the prosternum, and it needs to be compared only with the more central European E. abbreviata, as is done in the description of the latter below. Lectotype designation. There are 2 d and2 2 in the Loew Collection in Berlin labelled as E. pilimana. One male labelled ‘1/6 Prg/10461 is in perfect condition and is apparently from the vicinity of Guben (? Germany), as stated by Loew (1869). One pair is labelled “Aug.56, Kittel’ (also in good condition), undoubtedly sent to Loew by Prof. Kittel. The second female has an illegible locality label, but is probably another syntype. The firstmentioned male (No. 10461) is herewith designatedas lectotype of Empis pilimana Loew, and it was labelled as such by me in 1986.

Figs 256-259. Male genitalia of E. (Coptophlebia) pilimana Loew (CZ, Bohemia, Starkoé near Nachod). — 256: postabdomen; 257: lamellae in lateral view; 258: dorsal lamella in anterodorsal view, lateral lamella dotted; 259: aedeagus with

hypandrium. Scale: 0.2 mm.

Distribution. Unknown from Great Britain, but recorded on several occasions from the Netherlands and its occurrence in Denmark or southern Sweden is quite possible. A species of the temperate and cooler parts of Europe, fairly common in central Europe, south to Switzerland (Engadine) and the Romanian Carpathians (Sinaia), preferring higher altitudes in warmer areas. Flying over a long period, from the beginning of June to the end of August, the adults locally common on flowers.

Empis (Coptophlebia) abbreviata Loew, 1869 A species closely related to E. pilimana, having the same main key characters including the tuft of _ black spinose bristles on prosternum at sides. Males of E. abbreviata have shorter and less bristled legs, in particular mid tibiae usually with 3 to 5 (rarely up to 10) long dorsal bristles, and mid basitarsus with only short hairs, like the fore basitar-

sus. Legs in female obviously longer and slender, posterior four femora without flattened hairs, but hind femora with an anteroventral row of simple bristles that are not much shorter than femur is deep. Vein M, is sometimes complete in E. abbreviata (at least on one side), reaching the wing-margin as in Empis s.str.; in such cases, males run in the

key to Empis s.str. praevia and females to Empis s.str. levis. Lectotype designation. Described by Loew (1869) with no locality given, but it is presumed that the specimens were collected somewhere in Germany (Chvala and Wagner 1989). There are three conspecific males in the Loew Collection in Berlin under E. abbreviata: one is labelled ‘11/6 67, Europa, coll.Loew, 10460’, with broken genitalia;

the second, labelled 11/6 67, Europa’, is in perfect condition, and was labelled by me in 1986 and is herewith designated as lectotype of Empis abbreviata Loew; the third, labelled ‘16/6 67, Europa’, has been dissected, and the macerated genitalia

are on a small mount pinned with the specimen; it

145

has a complete vein M, on the left wing, as mentioned by Loew (1869:85). Length of lectotype: body 2.8 mm, wing 3.2 mm. E. abbreviata is a central European species, distributed southwards to mountainous areas of northern Italy and Romania (Carpathians), from June to end of August.

Subgenus Xanthempis Bezzi, 1909 Xanthempis Bezzi, 1909:88 (as subgenus of Empis). Type-species: Empis stercorea Linné, 1761 (orig. des.).

Generally large, from 4 to 8 mm long, yellow species with small eyes that are broadly separated on frons in both sexes, conspicuously long antennae, and body with reduced but very specialised bristling. Head very characteristically shaped, elongate posteriorly with occiput very convex, bearing two more or less irregular circlets of black occipital bristles. Eyes smaller than is usual in the genus Empis, broadly separated on frons, and with all ommatidia equally small in both sexes. Face usually even broader than frons, and jowls below eyes very deep. Antennae very long and slender, basal segment generally about 4 to 5 times as long as deep (except for the phylogenetically more primitive forms such as E. scutellata and E. laetabilis), covered with short, fine bristly-hairs; 2nd segment almost globular, as long as deep, with a preapical circlet of minute bristles; 3rd segment the longest, slender and barely broader than basal segment, together with the short terminal style always much longer than rest of antenna. Palpi small, usually concealed in a large mouth-cavity and scarcely visible, clothed with several minute hairs. Proboscis long and rather strong, labrum more than twice as long as head is high and almost uniform in all recent species; labella rather short, soft and stout. Thorax almost bare, devoid of small hairs of uni-

form pubescence, but with a very specialised and species-specific distinctive chaetotaxy. In general, dorsocentral setae uniserial and acrostichals absent, though in some species of the EF. /utea-complex the dorsocentral bristles may be more numerous and then irregular, merging posteriorly with additonal irregular setae, evidently acrostichal bristles. Otherwise dorsocentral bristles always

146

regularly uniserial, but varying in length, number and position, even on the left and right sides of mesonotum. Large bristles often very prominent, in full number represented by humeral, posthumeral, notopleural, supra-alar, postalar and scutellar bristles, sometimes with additional small hairs. Metapleural fan composed of only a few bristles that vary in length. Prothorax large, prosternum (partly fused with episterna) with minute fine hairs at sides, but pronotum strongly developed and bilobed, bearing several strong stubby spines on both sides. Thorax completely yellow without any pattern (E. /utea-complex), or mesonotum with specific patterns of dark stripes to almost uniformly blackish-grey; in non-Scandinavian species, thorax rarely almost completely black, mostly in alpine species. Wings large, conspicuously elongated and practically clear, stigma (if present) yellowish and very faint, veins contrasting thick and blackish. Axillary lobe only slightly developed and axillary excision very obtuse. Radial fork broadly ‘open’, obtuse, vein R, often almost at right-angles to R; and only slightly curved, though in general distinctly Sshaped. Legs simple, long and slender, femora and tarsi never dilated or thickened, pubescence very short and inconspicuous, femora beneath covered with soft dense pile, usually whitish on front legs, blackish on posterior two pairs. No distinct bristles except for circlets of preapical bristly-hairs on femora and tibiae, and very short dorsal bristles on tibiae that vary in number, size and position even in the same specimen. Abdomen covered with rather sparse short hairs, hind-marginal bristles distinct on the last (8th) pregenital segment, and less distinct bristles also present on Ist tergum, other terga with longer hairs on posterior margin, if any are differentiated. Male genitalia with a large number of very conspicuous species-specific distinctive characters on lamellae and aedeagus, though of simple structure in ground-plan. Hypandrium (ventral lamella) always small, membraneous anteriorly below and shielding only extreme base of aedeagus; aedeagus usually very simple, with a characteristically shaped sheath-like opening, rarely long or specifically bent or curved. Genital lamellae very small and simple in primitive forms, but in most species dorsal lamellae of very complicated structure and with specific bristling. Female abdomen narrowed apically, last two segments elongate, and cerci very long and slender.

Phylogeny and classification.

Xanthempis appears

i

Ky Abey ka

J 3

Fig. 260. Empis (Xanthempis) stercorea L., male. Total length: 5.2-7.5 mm (A. Veltman del.).

to be a natural monophyletic group within Empis s.lat., well-defined in.a very broad sense even if Rhamphomyia is considered to be generically problematic, as there are no allied forms without a

radial fork on wing or with a tendency for its reduction. It is undoubtedly a group of very specialised forms, sharing many characters in an apomorphous condition such as the horizontally elongate

head with posteriorly produced occiput, the very specialised chaetotaxy and entire lack of pubescence or hairing. Moreover, it is the only group of Empis so far known with a very specialised ethology, where the mating activity has been transferred from the air to a solid substrate, and where mating

in swarms, with parallel prey presentation, has been completely abandoned. The long separation

147

of this group of species even in its biology is supported by the very specialised displays that have been developed in the course of a long evolutionary process for better species-specific short-range recognition on the ground, as described in the section on biology. ‘ The almost completely yellow species with a rather short basal antennal segment, simple male genitalia and a trace of acrostichal bristles on mesonotum (E. scutellata, E. lutea-complex) are regarded as the primitive forms of the subgenus. The dark pattern that may be dominant on the head and thorax is considered to be only an ecological adaptation (cryptic colouration), without phylogenetic significance. The more specialised forms within the subgenus are those species with small eyes and broad jowls below, long antennae, male genitalia of more complicated structure, and reduced sensory bristling on body and legs. E. digramma is believed to be one of the most specialised species. The subgenus Argyrandrus Bezzi, with a single western Palaearctic species (E. dispar Scholtz), superficially resembles Xanthempis, especially in the yellow-coloured body, the very similar specialised bristling on thorax, and the legs devoid of bristles. However, in Argyrandrus the head is closely set on the thorax, with the occiput not at all produced posteriorly, the males are holoptic with upper ommatidia considerably enlarged, and the antennae (with short basal segment as in E. scutellata) have the 3rd segment ofa different structure. Even if it is accepted that these two subgenera had a common ancestor, these somatic differences indicate a long phylogenetic separation. Although nothing is known of the epigamic behaviour of Empis dispar, the structure of the eyes in the male indicates swarming behaviour, as is usual in other groups of Empis. Collin (1961) synonymised Argyrandrus with Xanthempis, but the reasons that he gave seem to be unsatisfactory and not well-founded. Distribution. Of the 25 species of Xanthempis so far described from the Palaearctic region, only 8 have been found in Scandinavia, though a further three may well occur there. Most of the Scandinavian species are of the so-called ‘southern’ distribution pattern, which means that the northern limit of their distribution is in the warmer southern parts of Scandinavia demarcated approximately by latitude 60° N. The only exception is E. stercorea, undoubtedly the commonest Xanthempis species in Scandinavia, which is widely distributed

throughout the whole area. All the other species of Xanthempis belong to zoogeographic distribution-

148

patterns A or B, as defined in the section on Zoogeography. Of the 25 described species of Xanthempis, only two occur in Japan and the others are European in distribution, with at least a further five undescribed species known to me from the central and southern parts of Europe. In general, Xanthempis is a rather boreal group of very specialised Empis species, as all the southern and Mediterranean species seem to occur in the mountains, or at least in colder mountainous areas. All species, even in the south, occur in the second half of April or in May; they are usually spring and early summer species, and none of them (except perhaps E. lutea) can be regarded as a late summer or autumn species. No Xanthempis species are known from the Canary Islands and North Africa, nor I have seen any specimens from Middle Asia.

Biology. Most of the species are restricted in their occurrence to shaded, sheltered and rather moist biotopes; they are common on low herbage in deciduous forests, or on shrubs and low vegetation

along side streams. They also commonly inhabit wet ruderal biotopes near man’s dwellings, such as parks and patches of nettles. The adults do not fly very much, and if disturbed they fly for a short distance and then settle again in the vegetation. Although they have large wings they are sluggish fliers, the females in particular. Their feeding and mating habits have been transferred to solid substrates, on the leaves of trees and bushes and on low vegetation. Larvae live in the soil, as adults are very commonly found in soil emergence traps, but Collin (1961) cited Belling who had bred E. trigramma from larvae found in April in an old cow-pat from the previous year. It is clear that the zoophagous Empis larvae were preying upon other saprophagous animals living in the cow-pat. The adults of both sexes have preserved both predation and nectar feeding in parallel throughout their whole lives, sources of protein-rich food and of sweet fluid substances. Mating, which has been secondarily transferred to the ground, is not connected with parallel predatory activity, the transfer of prey by male to female, as in other secondarily purely nectar-feeding Empis species. Mating in aerial swarms has been abandoned in Xanthempis species, except perhaps in the phylogenetically very old E. scutellata (this species still has rather large eyes and a narrower frons in male), as Hamm (1933) has observed small male swarms where females entered and copulated with males. It may even be a relict, generally aban-

doned, exceptional behaviour which appears from time to time in otherwise non-swarming populations, as for instance in the rhagionid genus Chrysopilus. No other records or observations on mating swarms in Xanthempis are available. The very rare and exceptional observations on small swarms or individual hovering males of some other species, as reported for instance by Collin (1961:455) for E. trigramma, are apparently of no importance for bringing together the different sexes, and they may be regarded as relict swarms (Chvala 1980) or as territorial behaviour (Pajunen 1980); hunting swarms do not come into consideration in such a ‘terrestrial’ group. The common epigamic behaviour of Xanthempis species is very exceptional within the genus Empis. It was first recorded by Poulton (1913) and fully described by Hamm (1933), and closely resembles the terrestrial displays known in many acalyptrates and some Mecoptera, such as scorpion-flies. The males perform a species-specific courtship on the vegetation or more often directly on the ground, with a very characteristic display that consists of the interrupted vibration of the wings with a parallel circling round the receptive conspecific female; she remains stationary, performs the same rapid vibrations of the wings, and is undoubtedly the initiator of the mating process, attracting the attention of nearby males. This performance climaxes with copulation, probably initiated by tactile signals as the male approaches the female. This is a very unusual habit in the family Empididae and is undoubtedly a highly specialised courtship that indicates a very different evolution in Xanthempis, leading to a very specific ethological isolation.

Key to Xanthempis species (Eight species of Xanthempis have been found in Scandinavia, but a further eight are included in the Key to avoid possible misidentifications of species occurring in the cooler parts of Europe.) 1

—

Ist antennal segment very short, slightly longer than 2nd segment. Thorax yellow, mesonotum densely light grey dusted, rarely almost greyish, with a very narrow, linear darker median stripe; supra-alar bristle present. Small, about 3.5-5 mm .... 54. E. scutellata {st antennal segment long, at least 3 times as long as deep, much longer than 2nd segment

2(1) Mesonotum uniformly yellow, unstriped (E. LULCG=STOUD) OER eae ete oe 3 — Mesonotum with dark stripes, or completely blackish eae Te 7 3(2) Supra-alar bristle present. Generally smaller species, about 4-5 mm long ............ 4 — Supra-alar bristle absent. Generally larger species, about 5-6 mm long. Dorsocentral bristles regularly uniserial, acrostichals abSENIMe eee oe ee oe eae eee 6 4(3) Dorsocentral bristles irregularly biserial, acrostichals present (except E. lutea 3). Posthumeral bristle absent, 1 or 2 notopleural bristles. Occiput uniformly yellow, ocellar tubercleidarkened ae see eae cote ae 5) — Dorsocentral bristles uniserial, acrostichals absent. Posthumeral bristle present, 3 notopleurals. Occiput with a black patch behind ocelli. Frons and occiput silvery dusted. ¢: aedeagus bent at right-angles before tip .... RES ans ee MAES 2 chat E. rohaceki 5(4) Upper part of frons and occiput shining yellow; 1 strong notopleural bristle. d: acrostichal bristles absent; aedeagus evenly arched, broad at base; dorsal lamella simple, lateral

lamella with a long narrow projection behind BaP ort rane dadecadat ha ahielrs 56. E. lutea — Frons and occiput silvery dusted; 2 notopleural bristles. d: acrostichal bristles present, even if fine; aedeagus slender and straight, directed upwards, and with a loop before tip; dorsal lamella bifurcate, lateral lamella witha short triangular-shaped projection ........ SE es Ae b. E. albifrons 6(3) Mesonotum dull, greyish dusted, 2 strong notopleural bristles. Occiput yellow with a faint greyish patch behind ocelli, abdominal pubescence whitish. Antennae shorter, Ist segment 3 times as long as deep. d: genitalia small, lamellae simple ..... 55. E. laetabilis — Mesonotum shining yellow, 1 strong notopleural bristle. Occiput largely black above, abdominal pubescence black. Antennae longer, Ist segment 4—5 times as long as deep. d: genitalia with dorsal lamella deeply exCISEAHANE 9 VE GIG. BE Bie 57. E. concolor 7(2) Mesonotum with a single dark median stripe which may be very narrow and deep black, or broadened or greyer, occupying whole area between dorsocentral bristles; no supra-alar. Generally larger species, over 5 mm long (E. stercorea-group); if in doubt, a species with vestigial thoracic bristles (E. digramma) belongs to the following section ........... 8

—

Mesonotum with two or more mesonotum completely blackish

stripes, or ....... 13

8(7) Prothoracic spiracle black. Mesonotum shin-

—

ing yellow with a narrow, contrasting black median stripe occupying at most one-third of the space between.dorsocentral bristles. Abdominal pubescence black ............. 9 Prothoracic spiracle yellow. Mesonotum at least finely greyish pollinose, median stripe broadened, or with a grey shadow at sides. Abdominal pubescence pale........... 10

9(8) 3: lateral lamella of genitalia concave above,

aedeagus bent beyond middle, shorter apical part as stout as basal part. 2: fore and hind tibiae with uniformly small hairs, no small dorsal bristles a" ==. = 58. E. stercorea — 6: lateral lamella convex above, aedeagus bent at middle, basal part with a swelling, apical part as long as, but much more slender than, basal part. 2: fore and often also hind tibiae with 1 or 2 small black bristles above . Qui ha Seven Se ee ee segs 59. E. aemula 10(8) Mesonotum with a narrow black median stripe as in E. stercorea, but margins right up to dorsocentral bristles greyish. Occiput yellow, with a narrow dark median strip-like Patchiivds cere se Rome E. loewiana — Mesonotum with a broad blackish grey median stripe occupying most of the space between dorsocentral bristles (Fig. 289) .... saihied oetash BOs fatate a6 marae Bases 11 11(10) Dorsocentral bristles outside the dark median stripe, situated on a yellow background (Fig. 289). A strong posthumeral bristle, 1notopleural. Occiput above neck mostly black. d: genitalia with three strip-like projections on dorsal lamella behind ......... RE 38 35.5 CO eo 60. E. univittata — Dorsocentral bristles on the margin of the dark median stripe, situated on a black background (Fig. 293). A small, hair-like posthumeral bristle, at least 2 notopleurals. Occiput with a black central patch above neck, postocular margins broadly yellow ..... 12 12(11) Pronotum with 3 to 6 black spines on each side, scutellum with one pair of large bristles. All tibiae with the usual short hairs beneath sot te: Bears sow oadhews E. aequalis — Pronotum with 10 to 12 strong black spines on each side, scutellum with 2 pairs of large bristles. Anterior four tibiae with very long antero- and posteroventral bristly-hairs, hind tib-

13(7) Mesonotum with 2 dark, narrow, longitudinal lines on a broad greyish median stripe. Thoracic bristles very reduced, minute, dorsocentrals invisible. Squamae and abdomen almost bare. Head yellow, occiput with only a small polished black patch behind ocelli. Prothoracic spiracle yellow. Large, 5-8 mm long’) :aeR ee ee. SEER 64. E. digramma — Mesonotum with 3 or4 more or less confluent stripes, or mesonotum mostly blackish. Thoracic bristles well developed, dorsocentrals hair-like but distinct. Squamae fringed, abdomen more or less distinctly pubescent ... 14 14(13) Mesonotum with 3 or 4, more or less con-

fluent, dark stripes, lateral margins (at least humeri, notopleural depression and postalar calli) yellowish. Thoracic pleura yellow in ground-colour, often with dark patches (E. trigramma-complex) ...............-. 15 — Thorax including pleura completely greyishblack (notin NW: Europe)se-e 2 See 25 oweee ou (E. semicinerea-complex) 15(14) Supra-alar bristle present. Mesonotum with 3 well-separated black stripes, dorsocentral bristles placed on the yellow interspaces. Occiput behind ocelli largely subshining black. Thoracic pleura uniformly yellow, prothoracic spiracles blackish. Large species, S-8 mm long 2teeesdsaue- Seen: 61. E. trigramma — Supra-alar bristle absent. Dark mesonotal stripes confluent, dorsocentral bristles on a dark background. Occiput dull blackish-grey above neck. Thoracic pleura with grey or dark patches, sometimes less visible .... 16 16(15) Smaller, about

—

5mm. Mesonotum

with a

single, broad, darker, median stripe on a dark grey background when viewed from in front; | notopleural bristle. Frons blackish-grey. Prothoracic spiracles blackish to brown; humeri contrasting yellowish, posthumeral region'blackishi@=.4--e= = 22- 62. E. punctata Larger, 68mm. Mesonotum lighter grey dusted in anterior view, with two narrow black lines between dorsocentral bristles; 2 notopleural bristles. Frons yellowish. Prothoracic spiracles yellow; humeri and posthumeral region yellowish ... 63. E. testacea

54. Empis (Xanthempis) scutellata Curtis, 1835 Figs 261-264.

iae with similar posteroventral bristles, and

all tibiae with long dense pubescence beneath Re ee ee . laeta

150

Empis scutellata Curtis, 1824: plate 18 (nomen nudum).

Empis scutellata Curtis, 1835: plate 18. Empis pallens von Roser, 1840:53. Empis parvula Egger, 1860:344. Small (about 4-5 mm) yellow, finely light grey pollinose species with black prothoracic spiracle anda narrow, dark, posteriorly widening median stripe. Basal antennal segments almost equally short, yellow; a long black supra-alar bristle. 3. Head rather closely set upon thorax, yellow in ground-colour, more or less densely light grey to silvery grey dusted. Frons about as deep as the distance between posterior ocelli, slightly widening below, face much broader, short, almost parallel-

sided. Eyes large, reaching to mouth opening, cheeks not visible. Occiput with two rows of rather coarse, forwardly-directed, black bristles, one behind postocular margin, the second behind middle of occiput, the rest of occiput behind the second

row of bristles distinctly yellowish. One pair of rather long black posterior ocellar bristles and a minute pair of vertical bristles. Antennae (Fig. 262) yellow on basal segments, Ist segment unusually short, barely longer than 2nd segment; 3rd segment black except for paler base, long-cylindrical, apical style rather stout. Proboscis yellow, rather

stout, twice as long as head is high. Palpi minute, brownish. Thorax dull yellow, thinly silvery dusted. Mesonotum with a very slender dark median stripe, more or less enlarged posteriorly, widening into a triangle before scutellum in prescutellar depression, and occupying at least base of scutellum. Metanotum extensively dark, prothoracic spiracle black. All thoracic bristles black, rather thin but

long: a long humeral with several smaller bristles anteriorly, a very long posthumeral, 2 scarcely shorter notopleurals, asmaller intrahumeral about as long as the row of 6 to 10 dorsocentral bristles (often a different number on each side) which becomes longer posteriorly; a long supra-alar (witha smaller bristle anteriorly), one equally long postalar, and a pair of scutellar bristles, usually with a minute hair on each side. Pronotum with a row of 6 to 8 small stubby spines, metapleural fan composed of 4 to 5 thin bristly-hairs, accompanied by several minute hairs. Wings clear with dark veins, a small costal bris-

tle, stigma indistinct. Squamae pale with fine dark brownish fringes, halteres whitish-yellow. Legs subshining yellow with apical two or three tarsal segments darkened. Pubescence short, blackish, except for longer preapical bristles on

Figs 261-264. E. (Xanthempis) scutellata Curtis, male (F, Provence, Montagne d. Lure). — 261: head and thorax in dorsal view; 262: antenna; 263: postabdomen; 264: aedeagus with hypandrium. Scale: 0.2 mm, 0.5 mm in Fig. 261.

151

femora and tibiae: all tibiae with several short dorsal bristly-hairs, those on fore tibiae (usually 3 bnistles) the longest, about as long as tibia is deep. Abdomen dirty yellow, somewhat subshining, although distinctly thinly silvery dusted in anterior view; pubescence short, black, hind-marginal bristly-hairs distinct. Genitalia small, yellow in ground-colour, covered with short black hairs; lateral lamellae triangular in shape and distinctly black pubescent above and at tip. Aedeagus short and rather stouter towards base, tip recurved. Length: body 3.74.8 mm, wing 5.6-6.5 mm. 2. Resembling male but dorsocentral bristles apparently weaker and smaller, and abdomen with less distinct hind-marsginal bristles. Colour of thorax clearly more variable than in male; the mesonotum may be almost uniformly grey, and sometimes with grey patches on pleura above coxae. Length: body 4.6-5.2 mm, wing 5.3-6.3 mm. With its short basal antennal segment and large eyes, E. scutellata can hardly be misidentified so far the European fauna is concerned. Rather a variable species in the colour of head and mesonotum, the central stripe on mesonotum is very light srey and indistinct in very pale specimens. In many respects, E. scutellata resembles the central European Empis (Argyrandrus) dispar Scholtz, but the latter has holoptic eyes in the male with upper ommatidia considerably enlarged, a single character that justifies its subgeneric separation. F. scutellata was described from England (Coomb Wood) by Curtis (1835): for the synonymy, see Collin (1961).

Distribution. Widely distributed in Europe, from England and Ireland and the Netherlands through temperate central Europe south to France (Provence). It has not yet been found in southern Scan-

dinavia, but its occurrence at least in Denmark is very possible. Collin (1961) gave a flight period in England from 10 May to 16 June, which corresponds almost exactly with that known from other parts of Europe —7 May to 14 June. Biology. Not at all a common species, preferring rather warmer and open biotopes. Copulating pairs without prey have often been observed, but Hamm (1933) described small male swarms in the early evening at a height of about 2.5 to 3.5 m (8-12 feet), the males flying rapidly to and fro and females entering the swarm. Copulation took place immediately in the air, and the pairs then settled on the ground. This behaviour is very unlike other species of Xanthempis, and it may be regarded asa

very primitive mode of mating in the air without prey transfer, a primitive behaviour aban-

doned long ago by other Xanthempis species. Hamm (1933) also mentioned a curious habit in females of E. scutellata, namely the production of a globule of transparent fluid extruded from the abdomen and imbibed by her through the proboscis immediately after copulation, a habit fully described by Hamm (1933) for E. trigramma, a typical Xanthempis species with characteristic epigamic displays on the ground.

55. Empis (Xanthempis) laetabilis Collin, 1926

Figs 265-268.

Empis (Xanthempis) laetabilis Collin, 1926:237.

Resembling EF. concolor but mesonotum dulled by greyish dust, 2 strong notopleural bristles, posthumeral bristle minute. Basal antennal segment decidedly shorter than usual, about 3 times as long as deep, often brownish. Abdomen whitish pubescent, and male genitalia small, simple. é. Head yellow, frons and face dull, covered by greyish dust, clypeus polished; face broader than frons. Occiput dulled by silver-grey dusting, generally very yellow, with an indefinite greyish shadow behind ocelli, sometimes a distinct greyish patch extending right up to neck and often widening at sides at about middle. A pair of fine black ocellar bristles, and numerous, not very much longer but stronger black bristles irregularly arranged on the upper half of occiput above neck. Antennae black, though basal segments often greyish-brown or almost light brown; Ist segment shorter than is usual in other species of Xanthempis (except for E. scutellata), about 3 times as long as deep, and together with the short 2nd segment about half as long as 3rd segment including style. Palpi very small, yellow, clothed with several fine dark hairs. Labrum yellow, only extreme tip darkened. Thorax uniformly yellow and rather dull because of a coating of grey pollen, mesonotum usually darker yellow, leaving humeri and postalar calli (sometimes also scutellum) more clearly yellow. Thoracic bristles black and rather sparse; without _ supra-alar or acrostichal bristles; dorsocentrals uniserial, about 8 to 9 bristles in a row, fine and hair-like, becoming longer posteriorly, last pair in prescutellar depression the longest, though not as long as other large bristles: a humeral (with additional minute hairs anteriorly), 2 notopleurals (sometimes a third hair-like bristle anteriorly), a postalar and one pair of scutellar bristles; posthumeral bristle very minute, at most as long as the fine anterior dorsocentrals. Metapleural fan com-

152 >

« 2

posed usually of 2 to 3 not very long bristly-hairs, surrounded by additional small hairs. Pronotum with 8 to 10 irregular black bristles each side, varying both in number and size. Prothoracic spiracle pale, notopleural line usually whitish.

spine-like bristles. Tergum and sternum on 8th segment separated (fused in other species of the E. lutea-complex). Length: body 44.8 mm, wing 5.2-6.3 mm. ?. More robust and larger than male, abdominal

Wings almost clear, veins dark, yellowish at ex-

pubescence finer and shorter, not so dense. Fore

treme base. A small black costal bristle. Squamae pale yellow, fringes black, halteres almost whitish. Legs pale yellow on coxae, otherwise rather dirty yellow because of the short, rather dense black pubescence, tarsi almost black on apical three or four segments. All femora covered with fine dense pubescence beneath, very sparsely on fore femora, but the usual whitish microscopic pile not developed. Tibiae dorsally with several (1 to 3) small black bristly-hairs, not longer than tibiae are

femora with whitish pile beneath. Abdomen pointed apically, 8th sternum and long slender cerci blackish. Length: body 5.5—6.5 mm, wing 6-6.3 mm. With its rather short basal antennal segment, discrete tergum and sternum on 8th abdominal segment, hind-marginal bristles on terga not differentiated, and very simple male genitalia, E. /aetabilis is in many respects a very primitive form with many characters in a plesiomorphous state, and representing an ancient type of the subgenus. In cool and temperate Europe E. laetabilis needs comparison only with E. concolor; it can be easily distinguished from the latter by the less darkened occiput, shorter first antennal segment, greyish pollinose dull mesonotum with 2 large notopleural

deep, mid and hind tibiae often without

dorsal

bristles. Abdomen rather subshining yellow, terga more or less darkened at least along anterior margins, or discs entirely dark. Pubescence very pale, dense and rather long, hind-marginal bristles not differentiated, though there are longer dark posterior hairs on first and last (8th) terga. Genitalia small, both lamellae simple, with short dark pubescence along margins, hypandrium (ventral lamella) very membraneous below, at tip with 3 to 4 small black

266

bristles (Collin, 1961: 458, erroneously referred to 1

bristle) and very minute posthumeral bristle, pale abdominal pubescence, and very small, simple male genitalia. However, there are several very similar and still undescribed species in the moun-

268

Figs 265-268. E. (Xanthempis) laetabilis Collin, male (N, SFi: Luster). — 265: antenna; 266: postabdomen; 267: aedeagus with tip of hypandrium in lateral view; 268: the same in posterior view, lateral lamella dotted. Scale: 0.2 mm.

153

tains of temperate Europe (Alps, Carpathians, Caucasus), as well as species described from Spain (E. algecirasensis Strobl, E. nevadensis Chvala) and from the Caucasus (E. caucasica Bezzi). Collin (1961:459) compared E. /aetabilis with E. aequalis Loew and E. laeta Loew, but these two species have a dark median stripe on mesonotum, as do E. loewiana Bezzi and E. unistriata Becker, and will therefore be discussed under E. univittata Loew. Distribution. A very rare species in Scandinavia, so far known only from four widely-scattered localities from Finland to Norway north to 62°; not known from Denmark. In Finland found in Al, Jomala (1 d, R. Frey), in Sweden in Sk., Maltesholm (1 5, 6 June 1985, H. Andersson) and Tollarp (1 2, 16 June 1924, O. Ringdahl), but in Norway (SFi, Lustev) G. Soli took a series of 7 d and 6 2 ina

Malaise trap on 23-24 June 1988. — A rare species everywhere, known outside Scandinavia from a few specimens taken in Scotland, and in central Europe in the Swiss Alps and in Slovakia during a short period at the end of May and in June.

56. Empis (Xanthempis) lutea Meigen, 1804 Figs 269-272. Empis lutea Meigen, 1804:228. Empis fulva Macquart, 1823:160. Empis ochracea Curtis, 1824: plate 18 (nomen nudum). Empis ochracea Curtis, 1835: plate 18. Generally small (4-5 mm) somewhat shining yellow species, mesonotum without dark pattern, completley yellow, supra-alar bristle present. Dorsocentral bristles irregularly biserial, small and fine in male, in the form of numerous small black spines in female, acrostichals absent in male, irregularly straggling in female. 3. Head completely yellow, leaving only ocellar swelling contrasting black or at least darkened. Frons broad, polished, though covered with silver dust just above antennae, as is most of the equally broad face, clypeus polished yellow. Vertex and upper half of occiput shining, yellow. A pair of rather small, fine, black ocellar bristles, occipital bristles (arranged in two circlets) not much longer but stouter. Antennae black, basal segments often brownish, the small 2nd segment even contrasting

yellowish; lst segment very long and slender, about 5 times as long as deep, covered with minute

154

black hairs similar to the preapical hairs on 2nd segment. Palpi small, yellowish, covered with several fine dark hairs. Proboscis yellow. Thorax uniformly yellow including mesonotum, the latter very finely covered by greyish pollen, almost subshining, thoracic pleura more dusted. All thoracic bristles black, rather fine and inconspicuous. The strong bristles confined to one notopleural, one postalar, and one pair of scutellar bristles. The humeral, posthumeral and supra-alar much smaller, and dorsocentral bristles arranged in two irregular rows, fine and hair-like, including the last prescutellar pair, irregular even in size. With several small black stubby spine-like bristles on humeri (about 3), in the notopleural depression

(at least 5), and about 4 to 5 similar small bristles each side on pronotum. Metapleural fan composed of 2 to 3 small black bristles similar in size to the larger dorsocentral bristles. Prosternum at sides with several very small fine pale hairs, also present on the upper anterior part of fore coxae. Prothoracic spiracle yellow. Wings clear with dark veins, though they are yellowish at base of wing, and the small fine costal bristle is hardly differentiated from the other costal bristling. Squamae and halteres pale yellowish, former with indistinct pale fringes. Legs yellow with tips of tibiae and whole of tarsi darkened, basitarsi often dirty yellow. Pubescence short, black; posterior four femora with fine dense hairing beneath, fore femora ventrally practically bare, clothed with only fine pale microscopic pile. All tibiae with irregular black bristly-hairs dorsally about as longas tibiae are deep, the bristles varying both in number (from 1 to 5) and in position. Abdomen subshining yellow in ground-colour, often brownish or darkened in dry specimens, covered with minute pale hairs. Basal three terga and 8th segment with longer black hind-marginal bristles. Genitalia with simple dorsal lamellae, clothed above with short black pubescence. Lateral lamellae with a conspicuously long, slender outer projection which is black bristled on upper margin and at tip. Aedeagus rather short, evenly curved, broader at base and gradually narrowed apically. Length: body 3.5-5.5 mm, wing 4.7-6.2 mm. 2. Differing from male in the bristling of mesonotum; dorsocentral bristles more numerous, shorter and more spine-like, distributed down the middle of mesonotum and over its entire width in front of scutellar depression; acrostichals also present and hardly differentiated from dorsocentral bristles. Other larger thoracic bristles as in male, including a small final prescutellar pair of dorsocentral bristles, and 2 or 3 small metapleural bris-

Figs 269-272. E. (Xanthempis) lutea Meigen (CZ, Moravia, Lednice). — 269: male thorax in dorsal view; 270: male postabdomen; 271: aedeagus with hypandrium; 272: female head and thorax in dorsal view. Scale: 0.2 mm.

tles. Mesonotum with a tendency to be more silvery dusted, not as subshining as in male. Abdomen pointed, cerci black, long and slender. Length: body 4—5.7 mm, wing 5-5.6 mm. E. lutea is a characteristic species in the northern European fauna because of its small size (body generally less than 5 mm long) and uniformly yellow thorax with a distinct supra-alar bristle. There are, however, several closely related and superficially similar species in temperate Europe that _have often been misidentified as E. lutea but apparently do not occur north of latitude 51°. In the Zoologisches Museum Berlin, for instance, there

are more than 150 specimens under Empis lutea Meigen (collectionsof Loew, Becker and Duda) which include a mixture of several superficially similar species, including Empis concolor Verrall. To avoid further misidentifications, these closely related species are discussed here, and one is de-

scribed as new to science.

The species of the Empis lutea-complex (E. lutea, E. rohaceki sp.n., E. albifrons Bezzi, and one undescribed species from the Swiss Alps with an indistinct dark median stripe) are characterised by their rather small size, usually yellow mesonotum with a tendency to have more numerous small dorsocentral bristles mixed with acrostichal bristles, and 8th segment in male ring-like and with tergum and sternum not differentiated. Distribution. A rare species of southern distribution in Scandinavia, so far known only from three Danish localities. All the records are old: H.J. Hansen took 1 ¢ in August 1880 at Brenderup (F) and 3 6 at Horsens (EJ); the third record is from Hou (NEJ), where W. Lundbeck took 24 ¢ and6 @ on six days between 6 and 25 July 1922. No recent data are available. —- A widely distributed species throughout Europe, from England and _ the Netherlands through central parts south to Spain,

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Albania and Romania, but not recorded from Russia or the Ukraine. A common species in central temperate Europe over a long period from 19 May (Bohemia) to 16 September, but more abundant in summer, and even in southern Europe more often in July. Mainly in sheltered lowland biotopes, practically absent in mountains, though in the Italian Alps recorded up to 1,300 m.

Empis (Xanthempis) albifrons Bezzi, 1909 Figs 273 -276. Empis nana Loew, 1867:21 (not Macquart, 1823). Empis (Xanthempis) albifrons Bezzi, 1909:91 (new name for Empis nana Loew, not Macquart).

Resembling E. Jutea but frons and upper part of occiput silvery pollinose, two strong notopleural bristles, acrostichals present even in male, and metapleural fan composed of more numerous bristly-hairs. Male genitalia very distinctive, aedeagus with a loop before tip. 6. Head uniformly yellow, frons and face very pale (sometimes darkened in dry specimens) and clothed with silvery-grey pollen, its appearance

then rather whitish. Ocellar tubercle slightly darkened (not as black as in E. Jutea), vertex and upper part of occiput, right up to the lower circlet of black bristles, densely silvery-grey dusted. Antennae as in E. lutea but completely black. Labrum yellow, tip largely blackish. Thorax uniformly yellow. Mesonotum subshining when viewed from above, finely greyish pollinose in anterior view. Thoracic bristling similar to that of E. lutea but with two notopleural bristles, the second upper anterior often almost as long as the lower one, at least half as long, and always very distinct from other much smaller and thinner side hairs, including a small posthumeral. Dorsocentral bristles irregularly biserial, hair-like, very much as in E. lutea including the last prescutellar pair, but usually with 2 pairs of equally fine acrostichal bristles at about middle of mesonotum. Notopleural line blackish. Metapleural bristles longer and weaker than in E. /utea, more numerous, usually 4

to 5 bristly-hairs with additional small dark hairs, together forming a true metapleural fan. Prothoracic spiracle yellow. Wings and legs as in E. /utea, but tarsi extensively darkened, often completely black even on basitarsi.

Figs 273-276. Male genitalia of E. (Xanthempis) albifrons Bezzi (SLK, Mala Fatra Mts, Bystfi¢ka). — 273: postabdomen;

274: dorsal lamella in anterodorsal view; 275: the same in lateral view; 276: aedeagus with hypandrium. Scale: 0.2 mm.

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Abdomen almost shining yellow, finely pale pubescent, dark hind-marginal hairs present on anterior three terga and on 8th segment. Genitalia very distinctive, lateral lamellae with the outer projection much shorter, and its upper hind margin (above the projection) with very dense short black bristles; dorsal lamellae long bifurcate near base. Aedeagus rather slender, pointing upwards, and with a distinct, apically broadened loop before a thread-like tip. Length: body 4.2-4.5 mm, wing 5.3-6 mm. 2. Mesonotum and thoracic pleura more silvery-grey pollinose. Bristling as in female of E. /utea, with numerous short dorsocentral and acrostichal bristles at about middle of mesonotum, posthumeral bristle practically absent as in E. lutea, but with a second upper anterior notopleural bristle as in male. Supra-alar bristle rather long, about as long as a humeral, a postalar and a pair of scutellar bristles. Tarsi contrasting black. Abdomen pointed, terga often narrowly darkening posteriorly in dry specimens, cerci very long and slender, dark grey. Length: body 4-6 mm, wing 5.3-6 mm. E. albifrons is often misidentified as E. lutea, with which it is compared in the above redescription. Lectotype designation. Empis nana (preoccupied, and renamed as E. albifrons) was described

by Loew (1867:21) as from ‘Styria et Carinthia’, Austria. There are 3 d and2 2 in the Loew Collection in Berlin, but only the males are conspecific and belong to the species redescribed above. The male labelled ‘Miirz. aug 55, 10515’ (either Miirz River or Miirzsteg in Steiermark) is herewith designated as the lectotype of Empis nana Loew, 1867 (a synonym of Empis albifrons Bezzi) and was labelled accordingly in 1981. Distribution. A species of temperate Europe, it seems unlikely to occur in Denmark or Scandinavia. Rather uncommon in central parts of Europe, commoner at higher altitudes, especially in Slovakia, Austria, Serbia and even Bulgaria (Vitoscha), from the middle of June to August.

Empis (Xanthempis) rohaceki sp.n. Figs 277-279. Rather a smaller species, generally up to 5 mm long, in the E. /utea-complex, with uniformly yellow, finely pollinose mesonotum; no acrostichal bristles, dorsocentrals regularly uniserial, supraalar bristle present. 3. Head yellow to reddish-yellow in ground-col-

our, distinctly though thinly silvery dusted, oral margin narrowly polished. Frons broad and almost parallel-sided, no frontal or orbital hairs or bristles, face as broad as frons, slightly widening out below. A pair of small forwardly-curved posterior ocellar bristles. Occiput with a more or less distinct black rhombus-shaped patch behind ocelli, and with two almost regular transverse rows of black spine-like bristles, the upper one at the broadest part of the dark patch, the lower one at the lower tip of the patch. Antennae black, lst segment about 3 to 4 times as long as deep, 2nd segment as long as deep, both covered with minute dark bristly-hairs; 3rd segment long and slender, distinctly longer than basal two segments together. Proboscis yellowish, labrum black at extreme tip. Palpi short ovate, yellowish, and with one or two minute

dark hairs at tip. Thorax uniformly yellow, rarely somewhat reddish-yellow, mesonotum without dark pattern and notopleural line pale. Both mesonotum and pleura slightly subshining, finely covered with a weak coating of silvery-grey dust. All thoracic bristles black: a long humeral (with 2 or 3 minute hairs anteriorly), one equally long posthumeral, 2 notopleurals (the upper slightly shorter), a supra-alar, a postalar and 1 pair of converging scutellar bristles, sometimes with 1 or 2 small hairs on each side. Acrostichal bristles absent, dorsocentrals uniserial, about 5 to 7 in a row, the first (? an intrahumeral) and the last pair on front margin of prescutellar depression the longest, and with another almost equally long pair in prescutellar depression in front of scutellum. Pronotum with about 6 spinelike bristles each side, metapleura with a tuft of about 5 to 10 rather hair-like bristles. Prothoracic spiracle pale yellow. Wings with dark veins becoming yellow at base (as is the whole of the incomplete Sc), a black costal bristle. Squamae pale with black fringes, halteres almost whitish. Legs long and slender, yellow, all tibiae darkened towards tip and tarsi practically black. Femora short dark pubescent, beneath with a dense coating of microscopic pile, almost whitish on fore femora. All tibiae with small black preapical bristles and additonal distinct dorsal bristles varying both in number and position; those on fore tibiae obviously the strongest, at least as long as tibia is deep, usually 3 to 5, either as single bristles or as slightly diverging pairs. Abdomen yellowish, dorsum often partly brownish and almost shining in some lights, all segments covered with fine, short dark hairs which become

paler ventrally. Lateral lamellae of genitalia witha

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short triangular-shaped projection, with distinct black bristles on upper margin, dorsal lamellae simple, not bifurcate. Aedeagus slender, pointing straight up, but bent before tip and curved backwards, its terminal part concealed within dorsal lamellae. Length: body 4-5 mm, wing 5.3-6.3 mm, holotype body 4.2 mm, wing 5.3 mm. 2. Resembling male in all respects, even the mesonotal bristling completely identical (very different in female of EF. /utea). Abdomen narrowed

apically, though similarly pubescent, last segment and long slender cerci almost black. Length: body 3.8-5S.5 mm, wing 5-5.6 mm. Empis rohaceki sp.n. needs comparison with E. lutea and E. albifrons in particular, which are smaller-sized species with uniformly yellow mesonotum and a supra-alar bristle. However, both these species have smaller and more numerous bristles on mesonotum; dorsocentrals are irregularly biserial in males, numerous in females, and on posterior half of mesonotum there are also irregular acrostichal bristles (absent in male of E. lu-

tea). There are also remarkable differences in the

male genitalia: lateral lamellae with a long slender conical projection in E. /utea, and aedeagus evenly curved, stout at base; E. albifrons has the dorsal

lamellae deeply bifurcate and aedeagus pointing upwards, with a loop before tip. Other northern species with a uniformly yellow unstriped mesonotum (£. concolor, E. laetabilis) are generally larger-sized (up to 6 mm), and without supra-alar bristle. Derivatio nominis. The species is named in honour of the Dipterist Dr Jindrich Rohaéek, Opava, who collected most of the specimens of this new species. Holotype ¢: Slovakia or., Nova Sedlica env., Stuzica reserve (sweeping undergrowth of mixed forest), 5.6.1985, leg. J. Rohatek;

in the Silesian

Museum, Opava. Paratypes: Same locality and data, 13.6.19862 ¢, 10.6.1990 1 2, 9.7.1990 1 d and 2 Q, 11.7.19901 3 and 2 2,10.6.19911 2; same locality, (sweeping undergrowth of deciduous forest) 9.6.1988 1 Q, (sweeping over boggy meadow) 15.6.1991 1 2; Slo-

Figs 277-279. E. (Xanthempis) rohaceki sp.n., male (SLK, Stuzica).—277: head and thorax of holotype in dorsal view; 278: postabdomen of paratype; 279: aedeagus with hypandrium of the same. Scale: 0.2 mm.

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vakia or., Stak¢in env., distr. Humenné (sweeping undergrowth of deciduous forest) 5.6.1984 1 @, 6.6.1985 1 3; all leg. J. Rohdéek and in the Silesian Museum, Opava, with 3 pairs in author’s collection. Romania, E. Carpathians, Cimpulung Mts., 6.6.1956 1 3, Cesna-Tesna, 1.7.19771 2, both leg. I. Ceianu and in coll. I. Ceianu, Bucarest. Distribution. Evidently a Carpathian species, known so far from the Slovak West Carpathians and the Romanian South Carpathians, on dates ranging from 5 June to 11 July.

57. Empis (Xanthempis) concolor Verrall, 1872 Figs 280-283. Empis concolor Verrall, 1872:283. Empis occipitalis Bezzi, 1909:91 (MS name).

Somewhat larger, generally about 5 (¢) to6mm (2). Mesonotum uniformly almost shining yellow, no supra-alar bristle, only one strong notopleural, and dorsocentral bristles mostly fine, hair-like. Occiput largely black above. Male genitalia with dorsal lamella deeply excised. 3d. Head yellow in ground-colour, the equally broad frons and face usually densely light grey dusted, but often darkened and frons sometimes almost blackish above. Occiput subshining yellow on a narrow strip just above neck and below, upper two-thirds largely blackish and finely grey pollinose. Ocellar tubercle black. A pair of ocellar bristles fine, black, scarcely longer than the stronger occipital bristles which are arranged in two circlets, those in upper row irregular. Antennae black, long and slender, basal segment long, about 4 to 5 times as long as deep and occasionally translucent brownish towards tip. Palpi yellow, very small, almost invisible. Labrum yellow with black-

ish tip, but often whole apical half extensively darkened. Thorax unicolorous yellow, mesonotum somewhat darker yellow and shining, without distinct dusting, pleura paler yellow and covered with fine greyish dusting. Bristling sparse, black, and rather fine. The only strong bristle is the upper posterior

both about as long as humeral or postalar but finer. Metapleural fan composed usually of 2 rather strong bristles surrounded by several not very much shorter but weaker bristly-hairs. Pronotum with about 6 short black spine-like bristles on both sides. Also with several minute dark hairs on humeri and along lower edge of notopleural depression, notopleural line usually black, rarely almost white. Prothoracic spiracle pale. Wings faintly yellowish-brown clouded, in some lights almost clear and iridescent, veins dark, yellowish at base. A short black costal bristle. Squamae almost whitish with black fringes, halteres dirty yellow. Legs long and slender, yellow, only extreme tips of all tibiae, and all of tarsi, blackish. Fore femora with a whitish microscopic pile beneath, posterior four femora with very short but dense fine ventral pubescence. Fore tibiae with usually two, closelyset, short, black bristles dorsally in apical third; with a similar, perhaps longer, bristle dorsally on hind tibiae, mid tibiae without bristles except for a circlet of short preapical spine-like bristles. Abdomen rather shining yellow, terga often with an irregular darker pattern along anterior margin and along median line. Abdomen clothed with short dark hairs, those hind-marginal hairs on anterior two terga and on 8th segment longer. Rarely abdomen extensively darkened in dry specimens. Genitalia very characteristic, having deeply cleft dorsal lamellae. The larger upper projection finger-like and very darkened, the smaller lower one pale, and armed with a strong black bristle. Lateral lamellae long, triangular in shape, upper margin and tip covered with black bristles; aedeagus rather short, stout and almost straight, concealed within lamellae. Unlike the species of the E. lutea-complex, the 8th tergum is in the form of a discrete sclerite in E. concolor. Length: body 4.3-5.5 mm, wing 5.5—6.6 mm. ?. Resembling the male in all details but generally larger. Abdomen long pointed, pubescence shorter, blackish as in male, though sometimes

a postalar and one pair of scutellars, sometimes with a small hair each side) much smaller and finer. Supra-alar and acrostichal bristles absent; dorso-

dark brown in some lights. Cerci black, very long and slender. Length: body 5—6.6 mm, wing 6-6.5 mm. E. concolor needs comparison with E. laetabilis in particular, but may be distinguished by the long antennae, the subshining mesonotum, the presence of only one strong notopleural bristle, the blackish pubescent abdomen, and the quite dis-

centrals uniserial, about 6 in a row and very small,

tinctive male genitalia, which are very small and

becoming slightly longer posteriorly, ending in a longer pair on frontal margin of prescutellar de-

simple in E. laetabilis.

pression, and another one in front of scutellum,

Distribution. According to Collin (1961), not un-

notopleural, the others (a humeral, a posthumeral,

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Figs 280-283. E. (Xanthempis) concolor Verrall, male (SLK, Mala Fatra Mts, Bystfi¢ka). — 280: head and thorax in dorsal view; 281: postabdomen; 282: dorsal lamella in anterodorsal view, with a tip of aedeagus at middle; 283: aedeagus with hypandrium. Scale: 0.2 mm.

common almost throughout Great Britain in the second half of June and in July, and, although not yet recorded from Belgium or the Netherlands, its occurrence in the southern parts of Scandinavia and in Denmark is very likely. A widely distributed species on the continent, through the central parts of Europe south to Spain and Romania; the latest record of its occurrence is 18 August from Switzerland (Jura).

58. Empis (Xanthempis) stercorea Linné, 1761 Figs 4, 7, 9, 260, 284-286. Empis stercorea Linné, 1761:247. Asilus ferrugineus Scopoli, 1763:364. Empis pertinax Harris, 1780:150.

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Empis rapax Pallas in Wiedemann, 1818:25. Empis stercoraria: Macquart, 1827: 117 (lapsus). Empis stigma Meigen, 1838:88. Large species, body up to 9 mm long, mesonotum subshining yellow with a contrasting black narrow median stripe, prothoracic spiracle black; no supra-alar bristle and only one notopleural. Male genitalia with lateral lamellae concave above, and

tibiae in female without short dorsal bristles. 3. Head yellow with a distinct rhomboid black patch on occiput behind ocelli, almost reaching pronotum. Frons broad, silvery dusted, though shining above around ocellar tubercle. Face broader than frons and dulled by silver pollen, upper edge of mouth cavity polished, clypeus dusted. Occiput with two circlets of rather long strong black

bristles, those in upper row irregular, all becoming finer and hair-like below. One pair of almost equally long ocellar bristles pointing forwards, and another one or two pairs of minute hairs posteriorly. Minute dark hairs also at sides of frons. Antennae very long, black, basal segment slender, about 4 to 5 times as long as deep; this and the very short 2nd segment with a tendency to be brownish-grey. Palpi yellow, very small, clothed with minute dark hairs. Proboscis long, yellow, labrum black at extreme tip and about 2.5 times as long as head is high. Thorax somewhat reddish-yellow on mesonotum, very finely greyish dusted, subshining in some lights, with a narrow, deep black, median stripe somewhat broadening behind on scutellum and metanotum. Pleura more yellowish, and with denser greyish dusting. All thoracic hairs and bristles black, no acrostichal bristles; dorsocentrals uniserial, about 6 to 10 in a row, fine and hair-like, ending

in a pair of large bristles on anterior margin of prescutellar depression, and another similar pair in front of scutellum. No supra-alar bristle, one notopleural bristle the strongest, other large bristles smaller: a humeral (with about 4 dark hairs anteriorly), a posthumeral, a postalar, and 1 pair of inner scutellars, with a smaller bristly-hair on each

side. Notopleural depression with several small dark hairs along lower margin above the pale notopleural line; pronotum with numerous (about 10) short spine-like bristles on each side; metapleural fan composed of about 6 rather long bristles and about

the same

number

of additional

smaller hairs. Prothoracic spiracle black. Wings almost clear,with distinctly contrasting dark veins (yellowish at extreme base) and a distinct brownish costal stigma. Squamae pale with black hair-like fringes, halteres whitish-yellow. Legs yellow, covered with short dark hairs, with-

out distinct bristles except for preapical ones on femora and tibiae. Tarsi darkened, blackish towards tip. Femora with rather dense fine black pubescence ventrally on posterior four pairs, fore femora with whitish pile beneath. Abdomen subshining yellow with a distinct dark median stripe on dorsum, more or less widening out at posterior margins of terga, and with a similar dark pattern along side margins of all terga. Pubescence short, pale, hind-marginal bristles not well differentiated, but those on first and last pregenital segment (8th) darker and more prominent. Genitalia yellow, lateral lamellae triangular in shape, upper margin concave, and small dorsal lamellae with a small point on hind margin below bearing along thin bristle. Aedeagus with an angu-

lar bend behind middle, apical shorter part equally stout, with a large sheath-like opening. Length: body 5.2-7.5 mm, wing 6.6—9 mm. ?. Resembling male in all details except for sexual differences, but generally larger, abdomen very narrowed apically. Tibiae without dorsal bristles, everywhere with short adpressed pubescence, fore femora beneath with microscopic whitish pile as in male. Length: body 6.8—9.2 mm, wing 7—8.3 mm. Because of the black prothoracic spiracle and the narrow deep black median stripe on mesonotum, E. stercorea needs to be compared only with E. aemula in the Scandinavian fauna, as has been done in the description of the latter. Distribution. The commonest species of Xanthempis in the northern parts of Europe; more than 500 specimens have been examined from all parts of Scandinavia, northwards to and including Lapland. From the end of May to July, on dates ranging from 14 May (EJ, Denmark) to 4 August (Vrm., Sweden); there is one female in the Copenhagen Zoological University Museum from S¢gnderborg (EJ, Denmark) collected by Wiistnei on 11 September 1902, which is either an error or a very exceptional second generation. —- A common species throughout Europe, though not as common in temperate Europe as E. trigramma. From the beginning of May very rarely to the end of July and the beginning of August, mainly in May and June. Biology. A typical species of sheltered biotopes, on bushes and low herbage, often on patches of nettles, with both sexes preying on other insects as recorded by Parmenter (1951) and Tuomikoski (1952). In May 1979 I observed the copulation of E. stercorea on low herbage in a deciduous forest near Gombasek (SE Slovakia): males were performing a special ‘dance’ round the females on the ground before copulation, starting at a distance of about 15 cm, with a rapid vibration of the wings and very characteristic movements by both sexes. A similar display was described by Hamm (1933) for E. tri-

gramma,

59. Empis (Xanthempis) aemula Loew, 1873 Figs 287-288. Empis aemula Loew, 1873:228.

Resembling E. stercorea very closely, but male genitalia with lateral lamellae convex above and aedeagus much more slender on its longer apical

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Figs 284-286. E. (Xanthempis) stercorea L., male (SLK, Gombasek near Roznava).— 284: head and thorax in dorsal view; 285: postabdomen; 286: aedeagus with hypandrium. Scale: 0.5 mm.

part behind angular bend, and female with 1 or 2 small bristles on fore tibiae in apical third above. é¢. Head very much as in E. stercorea with frons subshining above, its lower part above antennae and face dulled by greyish dust. Vertex and occiput with the same bristling; the usually velvety-black patch behind ocelli varying in size and intensity, but the shape always rhombus-like. Antennae, palpi and labrum as in E. stercorea. Thorax rather reddish-yellow on mesonotum, though humeri, notopleural depression, postalar calli and sides of scutellum more clearly yellow, like the more greyish dusted pleura. Median stripe almost velvety-black, narrow as in E. stercorea, sometimes varying in shape: either rather broadened, in the central part of mesonotum occupying more than one-third of the space between dorsocentral bristles and still broadening out anteriorly

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and posteriorly; or very narrow and somewhat less distinct between

humeri and front of scutellum,

continuing on to scutellum as a median line, or scutellum mostly black except for narrow side-margins. All hairs and bristles as in E. stercorea, the single notopleural bristle the longest and stoutest, other large bristles not very much smaller. Metapleural fan with numerous black bristly-hairs as in E. stercorea, and prothoracic spiracle also black. Wings and halteres as in E. stercorea; legs similar in colour, tarsi hardly ‘blacker’ as given by Collin (1961:463). The colour of tarsi varying as in E. stercorea, from specimens with tarsi entirely black to pale specimens with basal tarsal segments brownish. Fore femora with microscopic whitish pile beneath, posterior four femora with short dense pu-

bescence ventrally. Fore tibiae usually with 2 short (about as long as tibia is deep) black bristles in ap-

287

288

Figs 287-288. Male genitalia of E. (Xanthempis) aemula Loew (DK, NWJ: Rydhave, Vinderup). — 287: postabdomen; 288: aedeagus with hypandrium. Scale: 0.5 mm.

ical third, arranged dorsally closely one above the other, sometimes also with 1 or 2 similar small bris-

tles on hind tibiae above. Abdomen almost shining yellow, though silverygrey pollinose in some lights and then dulled, short black pubescent, with a similar black pattern to that in E. stercorea, sometimes whole dorsum of abdomen completely black. Genitalia very distinctive because of the convex upper margin of lateral lamellae and the larger projection on dorsal lamellae behind bearing a long bristle. Dorsal lamellae posteriorly, and hind upper margin of lateral lamellae, with distinctly longer black bristly-hairs than in E. stercorea. Aedeagus much more slender on a longer apical part behind the angular bend, and with a distinct inner swelling before the bend. Length: body 6.3—7.3 mm, wing 8-9 mm. 2. Resembling male in all respects except for sexual differences. Fore tibiae with 2 (rarely one) short black bristles above as in male, and similar bristles usually also on hind tibiae. Length: body 7-8.2 mm, wing 7.5-8.3 mm. E. aemula is very closely related to E. stercorea, and females are particularly difficult to distinguish in these two species. Collin (1961) gave several distinguishing characters for E. aemula, but they all seem to be very questionable as they suggest a range of variation, including a larger size for females of FE. aemula (7-9.5 mm) and a smaller size

for E. stercorea (5.5-8 mm according to Collin). The sole reliable distinguishing character for the female of E. aemula is very probably the presence of small black bristles on fore tibiae above, whereas the male is easily distinguished by the very differently shaped male genitalia. Note. E. aemula was described by Loew (1873) from Poland, ‘Galizien (Grzegorzek)’, but there is neither a type specimen nor any label with the name ‘aemula’ in the Loew Collection in Berlin. The types appear to be lost, and I have followed Collin (1961) who differentiated E. aemula from E. stercorea. Distribution. A rare species of southern distribution in Scandinavia, known only from 4 ¢ and5 2 taken in Denmark (LFM, Lolland and NWJ, Rydhave, Vinderup) and the south of Sweden (Skane and Sm., Torsas and Bergkvara), on dates ranging from 13 to 18 June. — Widely distributed in Europe, from Great Britain and southern Scandinavia through central Europe south to northern Romania (Mt. Rodnei 1,400 m); in temperate Europe at rather higher altitudes, on dates ranging from 19 May to 23 July. It seems to be rather uncommon everywhere, though many records of E. stercorea may well refer to E. aemula.

163

Biology. More often in submontane regions in central Europe, on herbage along side streams and in wet shaded biotopes. No data on the epigamic behaviour are available, but both males and females are frequently found on.flowers, sucking nectar or preying on other insects.

60. Empis (Xanthempis) univittata Loew, 1867

Figs 289-291.

Empis stercorea L. var. b Zetterstedt, 1842:378. Empis univittata Loew, 1867:20.

Large species, about 5-6 mm, with a yellow prothoracic spiracle and a broad blackish median stripe on mesonotum that occupies almost the whole area between dorsocentral bristles. Occiput largely black above; no supra-alar bristle, 1 long notopleural and 2 scutellars. 3. Head yellow in ground-colour but frons often darkened, greyish dusted, sometimes almost blackish-grey, face and clypeus yellow but very densely silvery-grey dusted, leaving only upper margin of the mouth-cavity shining. Occiput almost black above neck, finely greyish dusted, rarely upper postocular margins very narrowly yellowish; lower part of occiput below neck yellow. One rather long but weak pair of ocellar bristles with a minute pair behind, upper part of occiput with two somewhat irregular circlets of black, rather long and strong bristles; below neck with sparse, small pale hairs. Antennae long, black, Ist segment at least 4 times as long as deep, sometimes translucent brownish on inner side. Palpi small, yellow, with a few minute dark hairs at tip. Labrum long, yellow, apically darkened. Thorax yellow, somewhat subshining, but in some lights dulled by a dense dusting of silver-grey pollen. Mesonotum with a single broad blackish to blackish-grey clearly defined median stripe, occupying almost the whole area between dorsocentral bristles, which are inserted on the yellow background outside the mid-stripe (in FE. aequalis and E. laeta, dorsocentral bristles are inserted on the black backgrond of the mid-stripe). Dorsocentral bristles uniserial, small and fine, hair-like, about 6

hairs in a row, scarcely longer posteriorly but ending in two strong pairs, one on anterior margin of prescutellar depression, the second in front of scutellum. Large thoracic bristles black, very long but not strong: a humeral (usually with another small hair anteriorly), a posthumeral, a notopleural (undoubtedly the longest bristle), a postalar, 2 pairs of prescutellar dorsocentrals, and one pair of scutel-

164

lars, with a much smaller hair laterally; no supraalar bristle. Pronotum with a few (4-6) short black spinose bristles on each side; metapleural fan composed of 4 to 5 rather long black bristly-hairs and a larger number of fine smaller hairs. Prosternum with a few minute pale hairs at sides. Wings clear with dark veins (yellow at base of wing), a small black costal bristle. Squamae pale with fine black fringes, halteres pale yellow. Legs yellow, with extreme tips of tibiae darkened, tarsi almost entirely black. Pubescence short, dark, posterior four femora densely short black pubescent beneath, fore femora with whitish ventral pile. Bristling of legs very much as in E. aemula, all femora and tibiae with circlets of short black preapical bristles, fore tibiae usually with 2 close-set dorsal bristles in apical third (one above each other), hind tibiae usually with a single dorsal bristle, and mid tibiae with ony short adpressed hairs. Abdomen shining yellow in ground-colour with a dark dorsal pattern consisting of blackish anterior and lateral margins on each tergum, anterior dark stripes often widened into triangles below right up to posterior margin. Pubescence short, pale, except for blackish hind-marginal bristlyhairs on first and the last pregenital (8th) segment. Genitalia with lateral lamellae triangular in shape, convex above and pointed and upturned apically, upper margin and tip with dense black bristling. Dorsal lamellae with two strip-like projections posteriorly, the smaller lower one with a long hairlike bristle, the tip of the upper larger one and tip of lamella blackish coloured and with short black bristles. Aedeagus with a slight bend at base, and another more distinct one just before the long sheath-like opening. Length: body 4.5—5.8 mm, wing 6—7 mm, lectotype body 5.1 mm, wing 6.5 mm. 2. Resembling male in all details, but generally larger because of the apically narrowed abdomen. Bristling on legs as in male, abdomen rather densely and very pale pubescent, shining, cerci blackish and very long. Length: body 5.2-7.3 mm, wing 6.5—7.2 mm. E. univittata is sometimes misidentified as E. stercorea, but in the light of its yellow prothoracic spiracle, whitish pubescent abdomen, and broad dark median stripe on mesonotum, it needs to be compared with E. aequalis, E. laeta, and partly also with E. loewiana, as has been done in the Key and in the additional notes below. Engel (1943:292), in Lindner’s ‘Die Fliegen’, correctly recognised this species and even the male genitalia were correctly illustrated, but his statements that the basal anten-

Figs 289-293. E. (Xanthempis) univittata Loew, male (DK, LFM: Lolland, Fuglsang Park). — 289: thorax in dorsal view; 290: postabdomen; 291: aedeagus. E. (Xanthempis) loewiana Bezzi (lectotype male). — 292: postabdomen. E. (Xanthempis) aequalis Loew (CZ, Bohemia, Davle). — 293: thorax in dorsal view. Scale 0.5 mm.

nal segments were ‘braungelb’ and the thorax ‘slanzendgelb’ are somewhat misleading. Lectotype designation. Loew (1867) described this species from ‘Europe media and septentrionalis’. In the Loew Collection in Berlin there are 6 3

Scandinavia, extending approximately to 60° N. Very rare in Finland, with 2 2 taken by R. Frey at Karislojo (AB), but fairly common in southern Sweden north to Vrm., and throughout Denmark; not recorded from Norway. Altogether 18 ¢ and 17

and 1 2 under ‘Empis univittata m.’, all conspecific

° have been examined from Denmark, and 33 3

and belonging to the species described above, from the localities (where given) of ‘Charl.br.’ (i.e. Charlottebrun, now in southern Poland), ‘Langenau’, ‘Carniolia Terglau’ and ‘Schnub.’. One male

and 24 2 from Sweden, on dates ranging from 30 May to 17 July. — Central and eastern parts of Europe, south to Romania, from the middle of May to the end of July, everywhere rather uncommon. Unknown from Great Britain and the western parts of the continent along the Atlantic coast.

labelled ‘Charl.br. Jul.57, 10512’ is herewith desig-

nated as lectotype of Empis univittata Loew and was labelled by me as such in 1981.

Additional

Distribution. A species of southern distribution in

notes

on

Empis

univittata, with re-

marks on three closely related central European

165

species that may also occur further north in the cooler parts of Europe: 1. Empis (Xanthempis) loewiana Bezzi, 1909 Lectotype designation. Empis loewiana was a new name proposed by Bezzi (1909) for Empis dimidiata Loew, 1867, preoccupied by Meigen in 1835 (now in Leptempis). E. dimidiata was described by Loew (1867:19) from both sexes from ‘Germania australis’. There is a single male in the Loew Collection in Berlin, undoubtedly a syntype, labelled ‘10504, TYPUS’

and ‘dimidiatus’, subse-

quently identified by Oldenberg as a Loew type and labelled ‘Loewiana Bezzi’. The male was labelled by me in 1981, and is herewith designated, as lectotype of Empis dimidiata Loew (= E. loewiana Bezzi). The specimen has the head gummed to a card, and the legs (except coxae and trochanters) and left wing are lost. Diagnosis. E. loewiana is a large species (lectotype body without head 5.3 mm, wing 7.3 mm), with an extensively yellow occiput with only a narrow strip-like dark patch (or line) between ocelli and neck. Thorax yellow, finely silvery-grey dusted, subshining from some angles; mesonotum with a narrow dark median stripe originating from a brown patch above neck and continuing to base of scutellum, then broadening and less distinct on metanotum and, as a polished brown stripe (widening posteriorly on each tergum), also on abdomen. The median mesonotal stripe has a greyish shadow at sides, and the uniserial, rather fine dorsocentral bristles (becoming longer and stronger posteriorly) are inserted right on the greyish and yellow border. Large thoracic bristles in full number, including 1 notopleural and 1 pair of scutellars, but no supra-alar bristle. Prothoracic spiracle yellow, squamae pale with long black fringes. Abdomen shining, pubescence fine, pale. Male genitalia (Fig. 292) very characteristic because of the broad, almost rectangular and apically blunt lateral lamellae, dorsal lamellae with two almost equally large lobes above, and a narrow upcurved projection behind. Aedeagus very much as in E. univittata. The genitalia of E. lJoewiana illustrated by Engel (1943, Fig. 180) are actually of E. /aeta, as pointed out by Collin (1961:459). 2. Empis (Xanthempis) aequalis Loew, 1867 Lectotype designation. Described by Loew (1867) from ‘Germania’. There are 3 d and 6 @ in the Loew Collection in Berlin under Empis aequalis, but not all are syntypes: the probable type localities are ‘Suffr, G6rz, Waldenb., Schneeb., Charl.br. and Bergun’. A male labelled ‘Suffr

166

10513’ and ‘Empis aequalis m.’ was labelled by me in 1981, and is herewith designated, as lectotype of Empis aequalis Loew. Diagnosis. Differing from E. univittata by the partly black occiput above neck, with a dark patch leaving the broad postocular margins and sides yellow; frons yellow. Thorax yellow, rather densely silvery-grey dusted; mesonotum with a broad, greyish-black median stripe occupying the whole area between (and including) dorsocentral bristles, which are inserted on the dark margins of the stripe (Fig. 293). Large thoracic bristles black, not very prominent: a humeral, a smaller hair-like posthumeral, 2-3 long notopleurals (!), a postalar, 2 to 3 pairs of long prescutellar dorsocentrals, and only one pair of scutellars, rarely with a minute hair on each side. Metapleural fan composed of about 4 to S fine, dark and pale bristly-hairs. Squamae with rather long, fine pale fringes; Bezzi (1909:89) separated E. aequalis and E. laeta in his key as having ‘Schiippchen am Rande ganz kurz und sparsam behaart’, as opposed to ‘Schiippchen dicht und lang bewimpert, die Wimpern wenigstens so lang wie das Schiippchen breit’ for all other Xanthempis species except E. digramma-—a very dubious differential character. Legs yellow with dark tarsi as in other related species, all tibiae with 1 to 3 short dorsal bristles but, unlike E. /aeta,

tibiae very short pubescent beneath. The male genitalia were illustrated by Engel (1943), showing conspicuously long rectangular lateral lamellae, but the broadly ovate dorsal lamella has a small narrow dorsal excision at about middle. 3. Empis (Xanthempis) laeta Loew, 1869 Holotype identification. Loew (1869) described this species on p. 82, but on p. 81 there is a note that ‘Zeller am 30 Mai ein Mannchen bei Preth fing’. Although there are 3 d and1 @ under ‘Empis laeta m.’ in the Loew Collection in Berlin labelled “TYPUS’, only the male labelled ‘30/5 67, Coll. H. Loew, Typus’ can be the holotype of Empis laeta Loew, and I labelled it accordingly in 1981. Diagnosis. Very much like E. aequalis, but both head (except frons and central parts of occiput) and thorax more shining, pronotum with more numerous (10-12) spinose bristles on each side (3 to 6 smaller spines in E. aequalis), scutellum with at least 2 pairs of large bristles, usually with 2 smaller pairs at sides (altogether 8 scutellar bristles). Legs as in E. aequalis, but anterior four tibiae with very long antero- and posteroventral bristly-hairs that are much longer than tibiae are deep, hind tibiae with similar bristly-hairs posteroventrally, and all tibiae with additional long, dense ventral pubes-

cence, about as long as tibiae are deep. Male genitalia with lateral lamellae shorter and more narrowly produced apically, dorsal lamellae ovate, convex above; Engel (1943) illustrated the genitalia of E. laeta on Fig. 180 as E. loewiana, though the lateral lamellae are not as blunt-tipped as shown by Engel.

much smaller in size (about 4mm), EF. wnistriata

eral and supra-alar bristles. Sides of mesonotum including humeri, whole of notopleural depression, and postalar calli largely yellow; scutellum entirely blackish, metanotum only narrowly yellowish at sides. Large thoracic bristles black: a humeral (with 1 or 2 small black hairs anteriorly), a similar long posthumeral, 2 notopleurais (the lower one longer and usually another small black bristle anteriorly), a supra-alar, a postalar (with another small hair anteriorly) and a pair of scutellar bristles, often with a small bristly-hair at side. Dor-

Becker, 1887, and E. kuntzei Becker, 1910. The for-

socentral bristles fine and hair-like, about 5 to 7

mer was described from the Swiss Alps (St. Moritz) from a single holotype female which has evidently been lost as it was not found in the Becker Collection by Bezzi (1909:92), who took it at that time for a form of EF. /utea with a black median stripe. E. kuntzei was described from Corsica; it has a black prothoracic spiracle, black occiput, no supra-alar bristle, and was arranged in the Becker Collection in Berlin under E. lutea.

bristles in a row, ending in 2 pairs of long bristles, one on the edge of prescutellar depression, the second in front of scutellum. Pronotum with numerous (about 10), small, black, stubby spines each side, metapleural fan composed of 3 to 5 rather long black bristles and several additional smaller paler hairs. Prothoracic spiracle contrasting blackish. Wings almost clear with distinct dark veins, yellow at extreme base, a small black costal bristle. Squamae whitish-yellow with brownish fringes, halteres pale yellow. Legs yellow, tibiae hardly darker at tip, but tarsi darkened and apical segments almost black. Fore femora with whitish microscopic pile beneath, posterior four femora with short, dense, rather brownish pubescence below. All tibiae with several short black bristles dorsally, often spread along the whole length (and even on mid tibia), the bristles not longer than tibiae are deep. Abdomen yellow, rather subshining, with a distinct dark pattern on terga consisting of median triangles and narrow side-margins. Pubescence pale, rather short but dense, even the longer hind-marginal hairs light brownish, only those on 8th segment blackish. Genitalia very conspicuous because of the very long and thin aedeagus, formed by the extremely long and thin sheath-like opening. Lamellae simple, with rather dense fine black pubescence on margins, dorsal lamellae pointing upwards to almost backwards. Length: body 4.8-7.2 mm, wing 5.8-8.2 mm. 2. Resembling male in all details except for sexual differences. Abdominal pubescence shorter and quite pale, cerci long, dark brown to blackish. Length: body 5.8-8.3 mm, wing 6—7.5 mm. E. trigramma is a rather variable species in its size, which is not an unusual feature in this subgenus, although the majority of specimens is about 6 mm long, and in the colour of thorax, especially in the shape and size of the dark mesonotal stripes. In very dark specimens the pale interspaces between the black mesonotal stripes are very narrow

Note. There are two further species with a single narrow black median stripe on mesonotum,

both

61. Empis (Xanthempis) trigramma Wiedemann, 1822 Figs 294-296. Empis trigramma Wiedemann in Meigen, 1822:38. Large species, generally about 6-8 mm, mesonotum with three broad blackish longitudinal stripes, supra-alar bristle present. Prothoracic spiracle blackish. Aedeagus in male very long and thin, apical upcurved part thread-like. 3d. Head yellow in ground-colour; frons, face and narrow jowls below eyes dulled by greyish dust, but upper part of frons close to ocelli and clypeus polished. Upper part of occiput above neck largely black, almost polished behind ocelli when viewed from in front, in posterior view wrinkled

and dulled. The dark upper part of occiput with numerous irregularly-set, short, black, spine-like bristles that are forwardly curved, not arranged in the usual circlets; a pair of ocellar bristles as long. Antennae completely black and very long, Ist segment 4 to 5 times as long as deep, 3rd segment about twice as long as basal segment. Palpi yellow, not very small or invisible as usual, ventrally with several minute pale hairs. Labrum yellowish, long,

more than twice’as long as head is high. Thorax yellow in ground-colour, rather densely silvery-grey dusted even on pleura. Mesonotum with 3 broad black stripes separated by more or less wide yellow stripes down the lines of dorsocentral bristles; lateral dark stripes shorter and narrower, demarcated on the outside by posthum-

167

Figs 294-296. E. (Xanthempis) trigramma Meigen, male (CZ, Bohemia, Davle). — 294: thorax in dorsal view; 295: postabdomen; 296: aedeagus with hypandrium and lateral lamella. Scale: 0.5 mm.

and rather yellowish-grey. On the other hand, specimens from eastern parts of Europe, especially from the vicinity of Moscow, are generally much paler: the yellow colour of the mesonotum is dominant, and the yellow stripes along the lines of the dorsocentral bristles are almost as broad as, or even

broader

than,

the three

blackish

stripes.

Other characters, including the structure of the male genitalia, show that these eastern populations are undoubtedly conspecific. However, there is a closely related, smaller, undescribed species with distinctive male genitalia known to me from the Caucasus. For the synonymy of E. trigramma and notes on types, see Collin (1961:467).

Distribution. Although it is a common species in Great Britain according to Collin (1961) and is

168

known in large numbers for numerous localities in England and Scotland, it is an unexpectedly southern species in Scandinavia with a distribution confined to a small area covering Denmark and the

south of Sweden, including Oland, north to Ostergotland. It is, however, a common species here, and more than 250 specimens have been examined, on dates ranging from 21 April to 3 July. In the Copenhagen collection there is again a very questionable record, 3 2 taken by Wiistnei at Sénderborg (SJ) on 9 April 1904; such an early date seems to be unlikely. — Widely distributed in Europe, in the south from Portugal to the Ukraine and east to central European Russia, but absent in NE Europe including the Baltic states, Finland and NW Russia. Undoubtedly the commonest Xanthempis species in temperate Europe, on dates ranging

from 24 April to 27 July, but mainly in May and only sporadically by June.

Biology. The very characteristic epigamic behaviour was fully described by Hamm (1933) and has been confirmed by subsequent authors. As such it has been accepted as a fundamental pattern of behaviour common to most Xanthempis species. The ‘dancing’ of solitary males and females on the ground lasts from one to several minutes and the time spent in copulation is about 4 to 6 minutes. Often in large numbers on shrubs and low vegetation in sheltered rather moist biotopes.

62. Empis (Xanthempis) punctata Meigen, 1804 Figs 297-299. Empis punctata Meigen, 1804:227. Empis dorsalis Curtis, 1824: plate 18 (nomen nudum). Empis ignota Meigen, 1830:338.

Empis dorsalis Curtis, 1835: plate 18. Rather smaller, generally about 5 mm, head mostly blackish-grey and mesonotum with three completely coalescent dark stripes, leaving only very narrow margins and humeri yellowish; no supraalar bristle. Thoracic pleura yellow with distinct grey patches on sterno- and hypopleura. 6. Head black in ground-colour, rather densely silvery-grey pollinose, clypeus and narrow jowls below eyes translucent yellowish, in paler specimens face and lower part of occiput above mouth also brownish. Upper edge of mouth-cavity polished black. Occiput with two almost regular circlets of black forwardly-curved bristles, anterior

pair of ocellar bristles almost as long but weaker; posterior pair very small. Antennae black, Ist segment slender, about 4 times as long as deep, but 3rd segment conspicuously long, at least twice as long (including short style) as basal two segments together. Palpi not as small as usual in this subgenus, clearly visible; short-ovate and brownish, covered

with silver pile, and 2 or 3 dark hairs at tip. Pro-boscis long, labrum more than twice as long as head is high, yellowish-brown on basal part, very darkened on apical half, or almost entirely blackish in dark-coloured specimens. Thorax viewed from above almost entirely blackish-grey, with a duller black median stripe and two shorter and somewhat broader stripes at sides, in anterior view the lateral stripes invisible. The yellow ground-colour present on humeri, in notopleural depression, and on postalar calli, in

darker specimens the yellow parts on dorsum of thorax almost brownish. Thoracic pleura yellow, densely silvery pollinose, and with distinct blackish-grey patches on lower part of sterno- and hypopleura, a less conspicuous patch also on mesopleura. Large thoracic bristles black: a humeral, a posthumeral (equally long), a very long notopleural (with a small hair anteriorly), no supra-alar, a postalar (with another small hair anteriorly), and 2 pairs of scutellars, outer pair smaller; no acrostichal bristles, dorsocentrals regularly uniserial, small and hair-like, about 6 bristles in a row, ending

in one very strong pair on anterior edge of prescutellar depression, and another smaller pair in front of scutellum. Metapleural fan black and very distinct, about 4 long black bristles surrounded by a number of smaller bristly-hairs; pronotum with 5 to 8 rather long, black stubby spines on each side. Prothoracic spiracle blackish or almost brownish. Wings faintly brownish clouded with distinct dark veins, paler at extreme base only. Squamae whitish to brownish, with light brownish fringes, halteres pale yellow. Legs yellow, tips of tibiae scarcely darker but tarsi very extensively blackish. Fore femora with whitish pile beneath, posterior four femora ven-

trally with the usual, very short, soft, dense, dark pubescence. All tibiae with a single to several irregularly-placed short black dorsal bristles, much shorter than the tibiae are deep. Abdomen subshining, yellow in ground-colour, with a distinct black pattern consisting of amore or less broadened dorsal stripe and narrow lateral margins of terga. Pubescence pale, rather short but dense, only the longer hind-marginal bristly-hairs on 8th sternum blackish. Genitalia with lateral lamellae triangular in shape, dorsal lamellae with two blunt, dorsal, apically darkened and short-pubescent processes, posteriorly near base with another projection bearing a long black bristle. Aedeagus simple, not very long and almost equally slender throughout, the apical sheath-like opening not very long. Length: body 4.2-5.3 mm, wing 5.5—6.7 mm. 2. Closely resembling male but generally paler, face, lower part of occiput and labrum often very yellow, and the pale colour on humeri, notopleural depression and postalar calli more contrasting yellow. Bristles on tibiae usually more numerous and longer, those on fore tibiae generally as long as tibia is deep. Abdomen very pointed, cerci black and very long. Length: body 4.8-6 mm, wing 5.6-6 mm. Because of its almost uniformly blackish-grey mesonotum combined with its rather smaller size,

169

Figs 297-299. EF. (Xanthempis) punctata Meigen, male (CZ, Bohemia, Mariadnské Lazné).—297: thorax in dorsal view; 298: postabdomen; 299: aedeagus. Scale: 0.5 mm.

E. punctata cannot be mistaken for any other northern European species. The closely related species of the E. semicinerea-complex (E. semicinerea Loew, E. styriaca Strobl and E. testiculata Bezzi) are confined to warmer areas of central and southern Europe. For the type specimens and synonymy of EF. punctata, see Collin (1961:461).

Poulton (1913), Gruhl (1924) and Smith (1952); mating takes place on the ground without prey.

63. Empis (Xanthempis) testacea Fabricius, 1805

Figs 300-301.

Empis testacea Fabricius, 1805:141.

Distribution. A species of rather southern distribution in Scandinavia, northwards to approximately 60° N, but only common in Denmark and the extreme south of Sweden; I have examined about 140 specimens. Not known from Norway or Sweden north of Dalarna, and there is an isolated area of distribution in Finnish Karelia australis, apparently a small isolated population of continental NW Russian origin. On dates ranging from 21 May to 14 July, but mainly in June; there is again a female in the Copenhagen Museum collections taken by Wiistnei at Madeskov (SJ) on 15 August 1889, another very questionable date. — Widely distributed throughout Europe except for the southeast parts of the Black Sea region; a typical spring and early summer species, on dates from 10 May to 21 July. Uncommon everywhere, both in lowlands and in the mountains.

Biology. The predaceous activity was described by

170

Very large, robust species, 6-8 mm long, mesonotum almost uniformly light grey dusted leaving only broad margins yellow, with 3 to 4 dark stripes (depending on the angle of view), the two narrow middle ones abbreviated behind. No supra-alar bristle, 2 notopleurals, prothoracic spiracle yellow. 3. Head yellowish on face, clypeus and lower part of occiput below neck, frons usually brown with narrow orbital margins black, vertex and at least upper third of occiput completely black in colour and densely grey dusted. Frons with minute dark marginal hairs, anterior pair of ocellar bristles black, pointing forwards and very long, not much shorter than Ist antennal segment. Upper part of occiput with dense black bristles, the bristles arranged in three irregular rows, those in the hind row the longest (but still shorter than anterior ocellar bristles). Antennae black, at most basal segment very indefinitely brownish beneath; Ist segment at least 4 times as long as deep; 3rd seg-

ment long and slender, without style distinctly longer than basal two segments together. Palpi small, yellow, covered with minute dark hairs to-

wards tip and finely silvery pollinose. Labrum about twice as long as head is high, yellow, darkened towards tip. Thorax almost uniformly black in colour and densely light grey dusted on mesonotum, leaving broad margins yellow, including humeri and inner posthumeral region, notopleural depression, anda wide area round postalar calli. Scutellum with margins broadly yellow, pronotum yellow except for blackish median patch. Thoracic pleura yellow, thinly silvery-grey pollinose, often with more or less distinct blackish-grey patches on pleural sclerites, at least on sternopleura. Mesonotum viewed from in front lighter grey dusted and with 2 narrow blackish stripes between dorsocentral bristles, in dorsal view the mid-lines disappearing on anterior edge of prescutellar depression, and with 2 further broad black lateral stripes; in posterior view with 3 stripes visible on mesonotum, as in E. trigramma, the two median lines completely disappearing and with a new, single, median stripe reaching the very darkened (in this view) prescutellar depression posteriorly. All bristles black: a strong humeral

(with 1 to 4 small bristles anteriorly), a strong posthumeral, 2 still larger notopleurals (often with several small bristly-hairs anteriorly below), a postalar, and usually 2 pairs of scutellar bristles, outer pair much smaller; no supra-alar or acrostichal bristles; dorsocentrals uniserial, usually alternat-

ing 1 strong bristle and 1 small hair in a row, and ending in 2 strong pairs in prescutellar depression. Pronotum with about 5 rather long black spines each side and with additional smaller stubby spines; metapleural fan composed of about 6 strong black bristles and about the same number of smaller bristly-hairs. Prothoracic spiracle yellow. Wings clear with distinct black veins, yellow at extreme base, subcosta pale throughout. Squamae light brownish with numerous small black fringes arranged in two rows, halteres yellow. Legs yellow but all tarsi almost completely contrasting black, tips of tibiae more or less darkened. Fore femora

with whitish pile beneath

as usual,

posterior four femora short black pubescent beneath. Tibiae with a few short dorsal bristles, usually1or 2 small bristles on fore tibia in apical third, and similar bristles on hind tibia at about middle. Abdomen subshining yellow, often completely yellow without any dark pattern, or at most witha

Figs 300-301. E. (Xanthempis) testacea F., male (NL, Slenaken). — 300: thorax in dorsal view; 301: postabdomen. Scale: 0.5 mm.

ii

narrow dark median line. Pubescence short and sparse, pale except for blackish hind-marginal bristly-hairs on 1st tergum and 8th segment. Genitalia yellow except for the darkened upper process of dorsal lamellae, beneath this with a small projection bearing a long bristle. Lateral lamellae almost rectangular, aedeagus evenly bowed, simple, the sheath-like opening not very long. Length: body 6.2—7 mm, wing 8.3-9 mm. 2. Resembling male in all details except for sexual differences, but tibiae often with more numerous short dorsal bristles, also present on mid tibiae. Abdomen long-pointed, cerci blackish, very long and slender. Length: body 6.5-8.3 mm, wing 7.5-8.6 mm. E. testacea is a very characteristic species because of its large size and the extensively dull light grey mesonotum with dark stripes that alternate from different points of view. E. trigramma has dorsocentral bristles (of equal length) on the yellowish interspaces, and prothoracic spiracle blackish. There is, however, another undescribed and very closely related species known to me from Spain, but this has differently shaped male genitalia, and the dark mesonotal pattern, very much as in E. digramma, is constant and with the stripes not coalescing.

Empis diagramma Meigen: Chvala and Wagner, 1989: 275 (misprint).

64. Empis (Xanthempis) digramma

Large yellow species, about 5-8 mm long, with small eyes and wide jowls below eyes, all thoracic bristles stubby and inconspicuous. Mesonotum with two narrow dark lines on a light grey background along the line of the minute dorsocentral bristles, mesonotal margins broadly yellow. 3. Head yellow, dulled by silver dust. Frons very broad, almost polished above, widening out below, but face still broader. Ocellar tubercle and a narrow patch behind the tubercle polished black, surrounded by a larger, dull, dark greyish patch. Upper part of occiput covered with scattered, small, black, forwardly-curved stubby spines, lower part practically bare. Antennae black, basal segments often translucent brownish or even yellowishbrown; Ist segment long, nearly 3 times as long as the globular 2nd segment; 3rd segment (without short style) at least as long as basal two segments together. Palpi yellow, covered with minute black hairs, very small, concealed in the large mouthcavity. Labrum yellow, at least twice as long as head is high. Thorax yellow in ground-colour, mesonotum with 2 narrow blackish grey lines on a light grey background; the greyish pollinose median stripe of about the same width posteriorly as scutellum, narrowing anteriorly, confined to the space between the two dark lines. All thoracic bristles exceedingly short and inconspicuous. In addition toa few minute black dots on humeri, at sides of prosternum, and in notopleural depression, there is a very small humeral bristle, a longer posterior notopleural and a postalar, and 2 minute scutellar bristles. Acrostichal bristles absent, dorsocentrals in the form of very small dark hairs in a row inserted on the black narrow line, very inconspicuous throughout, as far as scutellum. Pronotum with about 8 to 10 small black stubby spines on each side on a produced lateral lobe; about 5 similar small spines, together with some small additional black dots forming the metapleural ‘fan’. Prothoracic spiracle yellow. Wings almost clear, with distinct black veins that become yellowish at extreme base of wing, subcosta completely yellowish. Squamae pale with mi-

Meigen, 1835

nute whitish hairs, halteres yellow.

Note. E. testacea was described by Fabricius (1805) as ‘in Austria Dom. de Megerle’, but in the Fabricius Collection in Copenhagen only a single wing is preserved, as noted by Zimsen (1964). It is believed that the species redescribed above is a correct interpretation of Fabricius’ species. There is a conspecific series of 4 d and 12 9 in the Becker Collection in Berlin under this name, from Spain and France. Distribution. Not yet found in Scandinavia or Denmark, nor in Great Britain, but its occurrence in the Netherlands (Slenaken, leg. B. Van Aartsen) illustrates a distribution from SW Europe along the Atlantic coast as far as NW Europe. Locally common in Spain and France, including Andorra in the Pyrenees, on dates ranging from 23 April (Spain) to 8 July.

Figs 302-305.

Empis punctata: Fabricius, 1805 and Meigen, 1830, not Empis punctata Meigen, 1804. Empis digramma Meigen in Gistl, 1835:69. Empis bilineata Loew, 1867:20.

172

Legs yellow, tarsi black towards tip, at least hind basitarsi dirty yellow. Pubescence short, black, without distinct bristles except for circlets of short black preapical bristles on all femora and tibiae, fore tibiae with 1 or 2 small black bristles on apical

Figs 302-305. E. (Xanthempis) digramma Meigen, male (SLK, Mala Fatra Mts, Bystri¢ka). —302: head and thorax in dorsal

view; 303: head in lateral view; 304: postabdomen; 305: aedeagus. Scale: 0.5 mm. half above, and usually a similar small bristle on

hind tibiae in apical third. Abdomen dull yellow, almost devoid of bristles and hairs, dorsum with a more or less distinct blackish median stripe. Genitalia with small dorsal lamellae brownish, clothed with densely-set fine

dark hairs; lateral lamellae yellow, produced posteriorly and covered with dense, mostly yellow bristly-hairs along upper margin and at tip. Aedeagus very stout at base, slender apically, with a curious loop at about middle. Length: body 5.5—7 mm, wing 6.6-8.3 mm. 2. Resembling the male in all details. Abdomen pointed, 8th sternum with a short terminal process,

its tip, as well as the long slender cerci, blackish. Length: body 5-8.3 mm, wing 7-8.3 mm. With its small eyes and deep jowls below, the very characteristic mesonotal pattern consisting of 2 narrow dark stripes along the lines of dorsocentral bristles, and the very reduced bristling on head and

thorax, E. digramma has no close ally in the northern and central parts of Europe. Note. No types or type locality were specified by Meigen (1935), but the environs of Munich, Germany, was given in the Palaearctic- Catalogue (Chvala and Wagner 1989). Collin (1961: 469) examined a single male under ‘digramma’ in the Meigen Collection in Paris that agrees with the description, given above.

Distribution. A species of southern distribution in Scandinavia,

rather

common

in Denmark

(75

specimens examined) and southern Sweden, in Skane, Smaland and Oland (54 specimens examined); in Finland, R. Frey took 7 ¢ and 4 @ at Jomala on the Aland islands; not yet found in Nor-

way. Mainly in May and in the first half of June, on dates ranging from 7 May to 3 July. — Widely distributed but never very common in Europe; locally common in suitable biotopes in central Europe, in

173

the mountains in warmer regions (Romania, Albania), though not recorded from the south-west and from eastern parts of Europe. Flight period as in Scandinavia, in May and June.

Subgenus LissempisBezzi, 1909 Lissempis Bezzi, 1909:99 (as subgenus of Empis). Type-species: Empis nigritarsis Meigen, 1804 (orig. des.).

A group of uniform, rather small (about 4mm), black species with long slender antennae and narrow wings, axillary angle very obtuse. Pubescence fine, sparse, no spinose bristles, all bristles on body and legs rather weak, hair-like. Head distinctly higher than broad, occiput not

convex, both sexes more or less dichoptic with all ommatidia equally small. Eyes sometimes only very narrowly separated on frons and even face very narrow; a slight sexual dimorphism in structure of head present in only the type-species, though in other species the degree of eye separation is identical in males and females. Ocellar bristles and occipital pubescence weak. Antennae very long and slender, basal two segments with sparse, minute hairs; 2nd segment globular, Ist segment usually more slender and varying in length, sometimes as long as 2nd segment; 3rd segment always long and slender, much longer (without style) than basal two segments together, style short, bristle-like. Palpi small, slender, at most with a few tiny hairs at tip. Proboscis shorter than is usual in Empis, labrum only slightly longer than head is high, heavily

Fig. 306. Empis (Lissempis) nigritarsis Meigen, male. Total length: 3.6-4.3 mm (A. Veltman del.).

174

sclerotised, slender and pointed, labium with small, soft ovate labellae covered with sparse hairs. Thorax with weakly developed simple pronotum, humeri and postalar calli, but scutellum large and prominent. Prosternum large, almost bare, only sides (epimera) with several long hairs. All thoracic bristles hair-like, practically no pubescence, and in most species mesonotum almost bare, acrostichal and dorsocentral bristles at most regularly biserial and always fine, or completely absent in some southern species; metapleural bristly-hairs very long, sometimes metapleural fan composed of only a few bristles, but these are weak and very long. Large marginal bristles on mesonotum including those on humeri and scutellum always fine, often reduced or practically absent. Wings slender, in particular axillary lobe weakly developed and axillary angle obtuse, in some species axillary lobe, and consequently axillary angle, completely absent, and wings very hemerodromiine-like. No costal bristle, radial fork rather longer

and acute, not ‘open’ and Empis-like; vein Cu (anal vein) more or less disappearing in apical half, but all veins from discal cell reaching wing-margin. Legs in general long and slender, though not always simple, especially in males. Hind tibiae apically and tarsi of hind legs often swollen, or even (in Mediterranean species) mid femora stouter or posterior tibiae curved, variously excavated and unusually bristled, but legs never spinose and even long bristles rather hair-like. Abdomen long-cylindrical, often sickle-shaped and downcurved in both sexes, covered with sparse fine hairs (especially terga at sides) but no distinct hind-marginal bristles. Male genitalia very simple, consisting of finely pubescent rounded lateral lamellae, dorsal lamellae small and usually concealed within lateral lamellae. Hypandrium (ventral lamella) small, shielding only the base of the short straight aedeagus, usually concealed within the rounded lateral lamellae as well. Female abdomen pointed apically, 7th and 8th segments somewhat laterally compressed, cerci long, slender. Phylogeny and classification. The subgenus Lissempis is without doubt a distinct and homogeneous group of very specialised Empis species. It shows several apomorphous characters with Xanth-

empis, such as those in the antennae and wings, but otherwise these two groups have undoubtedly evolved in two quite different evolutionary directions (lineages). On the other hand, Lissempis species very closely resemble the species of Rhamphomyia subgenus Lundstroemiella and, as stated by

Collin (1961:451), the main character separating Lissempis from Lundstroemiella appears to be the presence of the radial fork on the wing. It should be noted that the wing-venation is very constant in both Lissempis and Lundstroemiella, that there are no venation aberrations (as for instance in Empis s.str.and Aclonempis), and that Lissempis species always have the radial fork completely developed. The subgenus Lissempis is clearly well-separated from all the other Palaearctic groups (subgenera) of Empis, but it is difficult to judge whether Lundstroemiella is the sister-group of Lissempis or whether their close resemblance is the result of parallel development. The structure of the male genitalia in Lundstroemiella, with a long slender aedeagus as in the type-species Rhamphomyia hybotina Linné, does not represent the ground-plan condition, as Rhamphomyia magellensis Frey for instance has simple male genitalia with a short aedeagus very much as in Empis nigritarsis. In my opinion, the main difference lies in the structure of the proboscis, which is very distinctive, long and differently shaped in the purely nectar-feeding Lundstroemiella species. This may indicate a long and separate evolution, with different biology and feeding habits in these two groups. Distribution. Species of Lissempis are known with certainty only from the western part of the Palaearctic region, with the centre of distribution in the Mediterranean region including North Africa. Altogether 5 species of Lissempis are known to me: four are typical Mediterranean species, with two new species awaiting description in addition to E. liosoma Bezzi and E. cuneipennis Bezzi; only the type-species, E. nigritarsis, is known from the temperate and cooler parts of Europe.

Biology. Nothing is known about the feeding habits and mating behaviour of Lissempis species because of their rather rare occurrence. The south European species are known from single specimens only, and even E. nigritarsis never occurs in large numbers. It is believed that species of Lissempis, unlike those of Lundstroemiella, are nectar

feeders and predators in parallel, as is suggested by the relatively short labrum and the undoubtedly functional, long, stylet-like maxillary laciniae. The structure of the legs in both sexes, the ‘hybotidlike’ enlargement of some parts of the legs for grasping and holding and not only for sexual activity, supports this hypothesis. The structure of eyes, which in both sexes are rather narrowly dichoptic with all the ommatidia

175

equally small, suggests that Lissempis species may well belong to the non-swarming group of empidids (Chvala 1976:359), with their mating activity transferred from the air to the ground, as in Xanthempis.

65. Empis (Lissempis) nigritarsis Meigen, 1804 Figs 5, 8, 10, 306-310. Empis nigritarsis Meigen, 1804:221. Empis crassipes Meigen, 1822:33 (not Schrank, 1781). Empis nitidicollis Curtis, 1831:214 (nomen nudum). ? Hilara atritarsis Oldenberg, 1924:232 (MS name of von Roser).

Small (about 4mm) shining black species with long black antennae, and fine pale biserial acrostichal and dorsocentral bristles. Legs mostly yellow, hind tibae apically and hind tarsi contrasting black and swollen: fore basitarsi and mid legs in male with very long pale bristly-hairs. S. Frons polished black, very linear, about as wide as antennal style. Face slightly broader, about as wide as Ist antennal segment, shining black but somewhat rugose. A pair of very long dark hairlike ocellar bristles: occiput with much shorter hairs (dark above, pale below), black in groundcolour but thinly grey pollinose. Antennae black, very long and slender; Ist segment slightly longer than deep; 2nd segment broader and almost globular; 3rd segment very long and almost equally slender, at least 3 times as long as basal two segments together, style very short, bristle-like. La-

brum yellowish-brown, about as long as head is high. Thorax very shining black on mesonotum and scutellum, prescutellar depression dull grey, pleura greyish dusted. All hairs and weak bristles pale; acrostichal and dorsocentral bristles hair-like, about as long as antennal style, both in two closelyplaced rows, latter diverging and becoming longer posteriorly. Marginal bristles not differentiated, humeri and notopleural depression with numerous rather longer pale hairs, though one notopleural distinctly longer. Scutellum with a pair of long, apparently coarse bristly-hairs, but both ‘bristles’ formed by two coalescing and equally long pale hairs. Metapleural fan composed of more than 10 very long pale bristly-hairs with some additional smaller hairs. Wings clear with distinct blackish-brown veins and a brown ovate costal stigma at tip of vein R,. Axillary angle very obtuse and anal vein disappearing halfway to wing-margin. Squamae pale with concolorous fringes, halteres yellowish. Legs yellow, leaving base of fore coxae, posterior four coxae and trochanters, and anterior four tarsi towards tip blackish, as well as tip of the apically broadened hind tibiae and entire hind tarsi. Hind basitarsus enlarged and swollen, as deep as tip of hind tibia, and about as long as rest of hind tarsus; 2nd tarsal segment still distinctly dilated. Fore femora and tibiae almost bare, but fore tarsi with very long pale outstanding bristly-hairs on dorsum of basal two segments, basitarsus at base with additional very long (not very much shorter than segment is long) sloping bristly-hairs. Erect and very long bristly-hairs also on basal half of mid tibiae, both dorsally and ventrally, hind femora with sev-

Figs 307-310. Male genitalia of E. (Lissempis) nigritarsis Meigen (F, Rambouillet).— 307: postabdomen; 308; dorsal lamella in anterodorsal view; 309: tip of dorsal lamella in lateral view; 310: aedeagus with hypandrium. Scale: 0.2 mm.

176

eral long bristly-hairs before tip, and hind tibiae dorsally; the longest bristly-hairs on both sides of mid tibia and on basal two segments of fore tarsi. Abdomen brilliantly shining black, dorsum almost bare, but sides of terga and venter clothed with long pale hairs. Genitalia small and very simple, lateral lamellae rounded, pale marginal hairs becoming longer posteriorly and darker on lower edge; dorsal lamellae simple, broadened apically and completely concealed within lateral lamellae. Aedeagus short and rather stout, usually only the tip of the somewhat arrow-like opening visible above lamellae. Length: body 3.64.3 mm, wing 3.6—4.2 mm. ?. Frons broader, about as wide as 3rd antennal segment, polished black as in male; face also slightly broader, as wide as frons and similarly rugose. Legs with short hairs everywhere, without the long weak bristling present in male, and hind legs less dilated on tibiae and tarsi. Abdomen very shining black, covered with only minute, sparse, pale hairs. Length: body 44.3 mm, wing 3.64 mm. Meigen’s types of Empis nigritarsis in Paris were examined by Collin (1961), and there is no doubt that the species is correctly recognised. Because of its mostly yellow legs with the hind pair dilated, the

pale hairing on the shining mesonotum, the presence of acrostichal and dorsocentral bristles, and the rather well-developed axillary lobe on wings, E. nigritarsis can be easily separated from all the other European species of Lissempis.

Distribution. A species with a southern distribution in Scandinavia, known to me in Denmark only from Jutland and the smaller Danish islands (LFM, B), and in Sweden only from the island of Oland. Rather rare; only 16 ¢ and 19 2 have been examined, from a long period from 9 May to 23 July. — Collin (1961) recorded it in Great Britain, from Only six counties in southern England north to Nottingham, as an uncommon

and overlooked

species ‘because it apparently is to be found for only a short period at the end of May and the beginning of June’. This does not agree with the captures in Denmark and Sweden, and I myself have seen a female taken by E.A. Fonseca in England (N. Somerset, Leigh Woods) as early as 27 April 1957. Not yet found in the Netherlands, but widely distributed through the central parts of Europe to the Mediterranean, from Spain east to the Greek island Samos. Everywhere rather a rare species.

a

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449-454. — 1991: Empididae (Diptera) of Finland: The Empis subgenera Empis and Coptophlebia. — Ent. fenn., 2: 79-86. Chvala, M. et R. Wagner, 1989: Family Empididae, in: Sods, A. et L. Papp (eds), Catalogue of Palaearctic Diptera, 6: 228-336, Akadémiai Kiad6, Budapest.

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mologist’s mon. Mag., 62: 146-159, 185-190, 213-219, 231-237. — 1927: Notes on the Empididae (Diptera) with additions and corrections to the British List. — Ibid., 63: 20-29, 61-67, 93-98. — 1961: Empididae, in: British Flies, 6, 782 pp., Cambridge. —1963: Empis staegeria new species of Empididae (Dipt.) from Denmark. — Ent. Meddr, 32: 171-172. Curtis, J., 1824-1839: British Entomology, London.

1(1824): pls 1-50, 12(1835): pls 530-577. — 1829-1831: A Guide to an Arrangement of British Insects, London, columns 161—256(1831).

Emeis, W., 1970: Zur Verbreitung und Okologie der Empididen (Ins. Dipt.) in Schleswig-Holstein. — Schr. Naturw. Ver. Schlesw.-Holst., 40: 79-96. Engel, E.O., 1941-1946: Gattung Xanthempis Bezzi, Gattung Empis s.lat., in: Lindner, E., Die Fliegen der Palaearktischen Region, 28 Empididae, IV, 4, pp. 193-280

(1941), 281-320 (1943), 321-384 (1946), Stuttgart. Fabricius, J.C., 1787: Mantissa insectorum, 2, 382 pp., Hafniae. — 1794: Entomologia systematica emendata et aucta, 4, 472 pp., Hafniae. — 1798: Supplementum entomologiae systematicae, 572 pp., Hafniae. — 1805: Systema antliatorum secundum ordines, genera, species, 373 + 30 pp., Brunsvigae. Fallén, C.F., 1816: Empidiae Sveciae, 17-34 pp., Lundae. Fourcroy de, A.F, 1785: Catalogus Insectorumque in Agro Parisiensi reperiuntur. Ent. parisiensis, 1: 1-231,

2: 233-544. Frey, R., 1913: Zur Kenntnis der Dipterenfauna Finlands. II. Empididae. — Acta Soc. Fauna Flora fenn., 37 (3): 1-89. — 1945: Tiergeographische Studien tber die Dipterenfauna der Azoren. I. Verzeichnis der bisher von den Azoren bekannten Dipteren. — Commentat. biol., 8(10): 1-114. — 1953: Studien iiber ostasiatische Dipteren. I. Die Gattung Empis L. — Notul. ent., 33: 29-57. —1954: Gattung Empis L., in: Lindner, E., Die Fliegen der Palaearktischen Region, 28 Empididae, IV, 4, pp. 385— 422, Stuttgart. Gistl von, J.N.F.X., 1848: Naturgeschichte des Thierreichs fiir hGhere Schulen. 216 pp., Stuttgart. Goot van der, V.S., 1989: De dansvliegen van het geslacht Empis in Nederland (Diptera: Empididae). — Ent. Ber., Amst., 49: 173-184. Gruhl, K., 1924: Paarungsgewohnheiten der Dipteren. — Z. Wiss. Zool., 122: 205-280. — 1955: Neue Beobachtungen an Schwarm- und Tanzge-

sellschaften der Dipteren (Dipt.). — Dt. ent. Z., (N.F.)

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A.H., 1908: Observations on Empis livida, L. —

Entomologist’s mon. Mag., 19: 181-184. — 1909: Further observations on the Empidinae. — Ibid.,

zweifliigeligen Insecten, von Johann Wilhelm Meigen. Vol. 3, 320 pp., Halle. Lundbeck, W., 1910: Empididae, in: Diptera Danica, genera and species of flies hitherto found in Denmark. Vol. 3, 324 pp., Copenhagen. Macquart, J., 1823: Monographie des insectes Diptéres de la famille des Empides, observés dans le nord-ouest de la France. —- Mem. Soc. Sci. Agric. Lille, 1822: 137—

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165.

— 1933: The epigamic behaviour and courtship of three species of Empididae. — Ibid., 69: 113-117. Harris, M., [1780]: An exposition of English insects with curious observations and remarks wherein each insect is particularly described, its parts and properties considered, the different sexes distinguished, and the natural history faithfully related. Decad 5: 139-166, London. Hobby, B.M., 1932: The epigamic behaviour of the male Empis opaca, F. (Dipt., Empididae). — Proc. R. ent. Soc. Lond. (B), 6: 67-68. Hobby, B.M. et K.G.V. Smith, 1961: The bionomics

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Empis tessellata F. (Dipt., Empididae). — Entomologist’s mon. Mag., 97: 2-10. —1962: The bionomics of Empis opaca Mg. (Dipt., Empididae). — Ibid., 97(1961): 204-208. Jonassen, T., 1987: New Norwegian Empididae (s.str.) (Dipt.). — Fauna norv., Ser. B, 34: 37-40. — 1988: Empidoidea (Dipt.) new to the Norwegian fauna. — Ibid., 35: 71-76. — 1992: Further Empidoidea (Dipt.) new to the Norwegian fauna. — Ibid., 39: 73-75. Laurence, B.R., 1950: Predators and prey. — Beds. Nat., 4(1949): 27-30. —1959: Swarming in six species of Empididae (Diptera). — Entomologist’s Rec. J. Var., 71: 37-39.

—1992: The two forms of Empis tessellata (Diptera, Empididae). — Dipterists Digest, 12: 1-2. Linné, C., 1758: Systema naturae per regna tria naturae.

Ed. 10, Vol. 1, Pt. 2, 824 pp., Holmiae. —1761: Fauna Svecica sistens animalia sveciae regni. Ed. 2 (revised), 578 pp., Holmiae. Loew, H., 1865: Ueber einige bei Kutais Imeretien gefangene Dipteren. — Berl. ent. Z., 9: 234-242. — 1867: Ueber diejenigen mit Empis chioptera Meig. verwandten Arten, welche dunkle Schwinger haben. — Ibid., 11: 25-62. — 1867: Nachtragliche Bemerkungen zu den Empis-Arten aus den Verwandtschaftskreisen der E. stercorea und chioptera. Ibid., 11: 157-166.

— 1869: Ueber einige Empis-Arten, welche zu den im XI. . Bande besprochenen Verwandtschaftskreisen geh6éren. Ibid., 13: 65-94. — 1869: Beschreibungen europidischer Dipteren. Systematische Beschreibung der bekannten europdischen

zweifliigeligen Insecten, von Johann Wilhelm Meigen. Vol. 1, 310 pp., Halle.

— 1873: Diptera nova, in Pannonia inferiori et in confinibus Daciae regionibus a Ferd. Kowarzio capta. — Berl. ent. Z., 17: 33-52.

— 1873: Beschreibungen europdischer Dipteren. Systematische Beschreibung der bekannten europaischen

— 1827: Insectes Diptéres du Nord de la France. Platypézines, Dolichopodes, Empides, Hybotides. Ibid., 1827:

1-159. — 1834: Histoire naturelle des Insectes. Dipteéres. Vol. 1, 578 pp., Paris. — 1838: Diptéres exotiques nouveaux ou peu connus. —

Mém. Soc. Sci. Agric. Lille, 1(2): 5-207. Meigen, J.W., 1803: Versuch einer neuen Gattungseintheilung der europaischen zweifliigeligen Insekten. — Magazin Insektenk., 2: 259-281.

— 1804: Klassifikazion und Beschreibung der europaischen zweifliigeligen Insekten (Diptera Linn.). Erster Band, 1: XXVIII + 152, 2: VI+ 153-314 pp., Braunschweig. — 1820-1838: Systematische Beschreibung der bekannten europdischen zweifliigeligen Insekten. 3(1822): X+

416 pp., 6(1830): [TV + 401 pp., 7(1838): XII + 434 pp., Hamm. — 1835: Neue Arten von Diptera aus der Umgegend von Miinchen, bekannt und beschrieben von Meigen, aufgefunden von Dr. J. Waltl, Professor der Naturgeschichte in Passau. Jn: Gistl, J. (ed.): Faunus, 2: 66-72. Meijere, de J.C.H., 1918: Neue hollandische Dipteren. —

Tijdschr. Ent., 61: 128-141. —

1935:

Ueber

zwei

neue

hollandische

Empididen

(Dipt.). — Ibid., 28: 126-128. Meyer, H. et B. Deydemann, 1990: Faunistisch-dkologische Untersuchungen an Dolichopodiden und Empididen (Diptera-Dolichopodidae u. Empididae, Hybotidae) in Kiisten- und Binnenlandbiotopen Schles-

wig-Holsteins. — Faun.-Okol. Mitt., 6: 147-172. Oldenberg, L., 1924: Die Empididen v. Rosers in Stuttgart (Dipt.). — Dt. ent. Z., 1924: 226-236. Pajunen, V.I., 1980: A note on the connection between swarming and territorial behaviour in insects. — Annls ent. fenn., 46: 53-55. Parmenter, L., 1951: Notes on the genus Empis (Dipt.,

Empididae) in Britain. — Entomologist’s mon. Mag.,

87: 41-44. Poulton, E.B., 1913: Empidae and their prey in relation to

courtship. — Ibid., 49: 177-180. Roser, C. von, 1840: Erster Nachtrag zu dem in Jahre 1834

bekannt gemachten Verzeichnisse in Wirttemberg vorkommender zweifliigeliger Insekten. — KorrespBIl. wiirtt. landw. Ver. Stuttg., (N.S.) 17(1): 49-64. Schrank, F. de P., 1803: Fauna Boica. Durchgedachte Ge-

schichte der in Baiern einheimischen und zahmen Thiere. Landshut, 3(1), 272 pp. Scopoli, J.A., 1763: Entomologia carniolica exhibens in-

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Vindobonae.

179

Smith, K.G.V., 1949: Some observations on Empis livida, Lin. (Dip., Empididae), with notes on prey. Entomologist’s Rec. J. Var., 61: 39-42.

~1952: Observations on the prey and predacious habits of various Empididae (Diptera). —J. Soc. Brit. Ent.,4:90— 93; — 1971: The sub-Saharan species of Rhamphomyia and Coptophlebia with abbreviated discal venation (Diptera: Empididae). — Ann. Natal Mus., 21: 77-96. Stephens, J.F., 1829: A systematic catalogue of British insects: Being an attempt to arrange all the hitherto discovered indigenous insects in accordance with their natural affinities. Vol. 2, 388 pp., London. Strobl, G., 1880: Dipterologische Funde um Seitenstetten. Ein Beitrag zur Fauna Nieder-Osterreichs. — Programm des k.k. Ober-Gymnasiums der Benedictiner zu Seitenstetten, 1880: 1-65. — 1893: Die Dipteren von Steiermark. I. — Mitt. naturw. Ver. Steierm., 29(1892): 1-199. — 1898: Fauna diptera Bosne, Hercegovine i Dalmacie. — Glasn. zemalj. Mus. Bosni Herceg., 10: 387-466, 561 616 (in Serbian). — 1899: Spanische Dipteren. — Wien. ent. Ztg., 18: 12-83. — 1902: Novi prilozi fauni diptera balkanskov poluostrova. —Glasn. zemalj. Mus. Bosni Herceg., 14: 461-517 (in Serbian).

— 1904: Neue Beitrage zur Dipterenfauna der Balkanhalbinsel. — Wiss. Mitt. Bosn. Herzeg., 9: 519-581 (German translation of the paper published in 1902). —1910: Die Dipteren von Steiermark. II. Nachtrag. — Mitt. naturw. Ver. Steierm., 46(1909): 45-292. Sullivan, R.T., 1981: Insect swarming and mating. — Fla Ent., 64: 44-65. Svensson, B.G. et E. Petersson, 1987: Sex-role reversed courtship behaviour, sexual dimorphism and nuptial gift in the dance fly, Empis borealis (L.). — Annls zool.

fenn., 24: 323-334. —1988: Non-random mating in the dance fly Empis borealis: The importance of male choice. — Ethology, 79: 307-316. Svensson, B.G., E. Petersson et E. Forsgren, 1989: Why

do males of the dance fly Empis borealis refuse to

180

mate? The importance of female age and size. — J. Insect Behavior, 2: 387-395. Svensson, B.G., E. Petersson et M. Frisk, 1990: Nuptial gift size prolongs copulation duration in the dance fly Empis borealis. — Ecol. Ent., 15: 225-229. Syrovatka, O., 1980: Revision of the subgenus Empis s.str. (= Pterempis Bezzi) (Diptera: Empididae) in Czechoslovakia. — Acta Univ. Carol.-Biol., 1977: 423—

427. — 1991: Revision of H. Loew’s and T. Becker’s types of Empis s.str. species (Diptera, Empididae) in the Berlin and St Petersburg Museums. — Mitt. Zool. Mus. Berl.,

67: 225-278. Syrovatka, O. et M. Chvala, 1986: Revision of J.W. Meigen’s types of Empis s.str. (Diptera: Empididae) of the Paris Museum, with an appendix of Macquart’s species. — Vést. és. Spoleé. zool., 50: 231-239. Tuomikoski, R., 1939: Beobachtungen tiber das Schwarmen und die Kopulation einiger Empididen (Dipt.). — Anns ent. fenn., 5: 1-30. 1952: Uber die Nahrung der Empididen-Imagines

(Dipt.) in Finland. — Ibid., 18: 170-181. Ulrich, H., 1972: Zur Anatomie des Empididen-Hypopygiums (Diptera). — Ver6ff. zool. StSamml. Miinch., 16: 1-28. Verrall, G.H., 1872: On the species of Empis allied to E. stercorea Linn. including one new species to science. — Entomologist’s mon. Mag., 8: 281-284. Villers, de C.J., 1789: Caroli Linnaei entomologia, faunae

sueciae descriptionibus aucta. Vol. 3,657 pp., Lugduni. Waters, S.B., 1989: A Cretaceous dance fly (Diptera: Empididae) from Botswana. — Syst. Ent., 14: 233-241. Wiedemann, C.R.W., 1818: Aus Pallas dipterologischen Nachlasse. — Zool. Mag., Kiel, 1(2): 1-39. Zetterstedt, J.W., 1838: Insecta Lapponica. Diptera, pp. 477-868, Lipsiae. — 1842-1859: Diptera Scandinaviae. Disposita et descrip-

ta, 1(1842): 1-440, 12(1855): 4547-4942, 13(1859): 4943— 6190, Lund. Zimsen, E., 1964: The type material of I.C. Fabricius, 656 pp., Munksgaard, Copenhagen.

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