Zootaxa, a Taxonomic Revision of Australian Conopidae (Insecta

ZOOTAXA

2581

A taxonomic revision of Australian Conopidae (Insecta: Diptera)

MARGARET A SCHNEIDER School of Biological Sciences, The University of Queensland, Brisbane Qld 4072 Australia. e-mail: [email protected]

Magnolia Press Auckland, New Zealand

Accepted by D. Bickel: 1 May 2010; published: 31 Aug. 2010 MARGARET A SCHNEIDER A taxonomic revision of Australian Conopidae (Insecta: Diptera) (Zootaxa 2581) 246 pp.; 30 cm. 31 Aug. 2010 ISBN 978-1-86977-557-5 (paperback) ISBN 978-1-86977-558-2 (Online edition)

FIRST PUBLISHED IN 2010 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] http://www.mapress.com/zootaxa/

© 2010 Magnolia Press All rights reserved. No part of this publication may be reproduced, stored, transmitted or disseminated, in any form, or by any means, without prior written permission from the publisher, to whom all requests to reproduce copyright material should be directed in writing. This authorization does not extend to any other kind of copying, by any means, in any form, and for any purpose other than private research use.

ISSN 1175-5326 (Print edition) ISSN 1175-5334 (Online edition)

1. Introduction ...... 5 1.1 Diagnosis of the Conopidae ...... 5 1.2 Current status of conopid taxonomy ...... 5 1.2.1 Subfamily classification...... 5 1.2.2 Tribal classification ...... 7 1.2.3 Australian conopid taxonomy ...... 8 1.3 History of conopid systematics...... 8 1.4 Fossil history...... 11 1.5 Biology ...... 12 1.6 Aims of this study ...... 13

2. Materials and Methods...... 14 2.1 Methods ...... 14 2.2 Definitions and terminology ...... 14 2.3 Abbreviations ...... 16 2.4 Format of descriptions ...... 16

3. Anatomy ...... 17 3.1 Head ...... 18 3.1.1 Antennae...... 18 3.1.2 Mouthparts ...... 18 3.2 Thorax ...... 19 3.2.1 Chaetotaxy ...... 19 3.2.2 Wing ...... 20 3.2.3 Vestiture ...... 20 3.2.4 Legs ...... 20 3.3 Male postabdomen ...... 21 3.3.1 External anatomy ...... 21 3.3.2 Internal anatomy...... 21 3.4 Female postabdomen...... 21 3.4.1 External anatomy ...... 21 3.4.2 Ultrastructure of spicules ...... 22 3.4.3 Internal skeletal structures ...... 22 3.4.4 Spermathecae ...... 22 3.5 Intraspecific variation ...... 23

4. Alpha taxonomy ...... 23 4.1 Key to the subfamilies of extant Conopidae ...... 24 4.2 Myopinae ...... 24 4.2.1 Introduction ...... 24 4.2.2 Subfamily diagnosis ...... 24 4.2.3 Myopa Fabricius ...... 25 4.2.4 Thecophora Rondani ...... 25 4.3 Notoconopinae subfam. nov...... 26 4.3.1 Introduction ...... 26 4.3.2 Subfamily diagnosis ...... 27 4.3.3 Notoconops gen. nov...... 27 4.4 Stylogasterinae ...... 29 4.4.1 Introduction ...... 29 4.4.2 Subfamily diagnosis ...... 29 4.4.3 Stylogaster Macquart ...... 30 4.5 Conopinae ...... 31

A TAXONOMIC REVISION OF AUSTRALIAN CONOPIDAE Zootaxa 2581 © 2010 Magnolia Press · 3 4.5.1 Introduction ...... 31 4.5.2 Subfamily diagnosis ...... 31 4.5.3 Key to genera of Australian Conopinae ...... 32 4.5.4 Atrichoparia gen. nov...... 33 4.5.5 Australoconops Camras ...... 35 4.5.6 Camrasiconops gen. nov...... 78 4.5.7 Chrysidiomyia Kröber ...... 82 4.5.8 Conops Linnaeus ...... 90 4.5.9 Delkeskampomyia Kröber...... 107 4.5.10 Heteroconops Kröber ...... 108 4.5.11 Microconops Kröber...... 115 4.5.12 Neoconops Kröber ...... 122 4.5.13 Physocephala Schiner...... 126 4.5.14 Pleurocerina Macquart...... 133 4.5.15 Setosiconops gen. nov...... 157 4.5.16 Smartiomyia Kröber ...... 161 4.5.17 Stenoconops Kröber ...... 168 4.5.18 Tanyconops gen. nov...... 170

5. Biogeography ...... 174 5.1 World distribution of Conopidae...... 174 5.2 Levels of endemism ...... 174

6. Phylogenetic relationships ...... 175

7. Conclusion ...... 176

Acknowledgements ...... 176

References ...... 176

Appendix Catalogue of Australian Conopidae ...... 181

Abstract

A taxonomic revision of the Australian Conopidae is presented. Nineteen genera in four subfamilies are recognised. The Stylogasterinae are represented by the cosmopolitan genus Stylogaster Macquart and the Myopinae by Myopa Fabricius and Thecophora Rondani. A monotypic genus, Notoconops gen. nov., is described and placed in a new subfamily, Noto- conopinae. The remaining fifteen Australian genera are assigned to the Conopinae. Australoconops Camras, Chrysidio- myia Kröber, Conops Linnaeus, Delkeskampomyia Kröber, Heteroconops Kröber, Microconops Kröber, Neoconops Kröber, Pleurocerina Macquart, Physocephala Schiner, Smartiomyia Kröber and Stenoconops Kröber are redescribed. Four new conopine genera are described: Atrichoparia; Camrasiconops; Setosiconops; and Tanyconops, with the following resultant new combinations: Atrichoparia curticornis (Kröber) (from Heteroconops); Camrasiconops ater (Camras) and Ca. rufofemoris (Camras) (from Microconops); and Setosiconops robustus (Kröber) (from Neoconops). Callosicon- ops Kröber and Paraconops Kröber are regarded as junior synonyms of Chrysidiomyia and Pleurocerina respectively with the following resultant new combinations: Chrysidiomyia hirsuta (Kröber), Pleurocerina aristalis (Camras), P. longicornis (Kröber), P. nigrifacies (Kröber), P. similis (Kröber), P. turneri (Camras). A total of 100 species of Australian Conopidae are now recognised. No new species of Myopinae or Stylogasterinae are described. Representative species only of Atrichoparia, Camrasiconops, Heteroconops and Microconops are described. All species of the other 12 genera are described or redescribed. These include the following 45 new species,: Australoconops aglaos, A. aurantius, A. balteus, A. breviplatus, A. brunneus, A. camrasi, A. cantrelli, A. elegans, A. fulvitarsus, A. furvus, A. nebrias, A. neuter, A. pallorivittus, A. phaeomeros, A. ruficrus, A. vespoides; Chrysidiomyia rugifrons, C. setosa; Conops aureolus, C. badius, C. chvalai, C. sparsus; Heteroconops carnarvonensis; Neoconops bre- vistylus, N. glaber; Notoconops alexanderi, Pleurocerina aquila, P. brevis, P. chrysopyga, P. lamellata, P. lutea,

4 · Zootaxa 2581 © 2010 Magnolia Press SCHNEIDER P. luteiceps, P. occidua, P. saxatilis, P. scutellata, P. vespiformis; Setosiconops epixanthus, Se. similis, Smartiomyia arena, Sm. cerina, Sm. danielsi, Sm. macalpinei; Tanyconops longicaudus, T. luteus, T. ocellatus. Australoconops ocellatus (de Meijere) comb. nov. is resurrected from synonymy with A. aurosus (Newman). Conops demeijerei Kröber is con- sidered a junior synonym of C. seminiger de Meijere. A lectotype and paralectotypes are designated for Microconops fasciatus Kröber. Anatomical features used in descriptions are discussed and illustrated. Special attention has been given to the ultrastructure of spicules on female genital structures. These spicules show differences between genera and species. The high degree of endemism of the Australian fauna is discussed in relation to the world fauna and some preliminary thoughts on phylogenetic relationships are presented. A catalogue of the Australian Conopidae is included as an Appendix.

Key words: Diptera, Conopidae, Australian, Conopinae, Myopinae, Stylogasterinae, Notoconopinae

1. Introduction

The Conopidae constitute a moderately small family of schizophorous flies including about 800 described species. Conopids are distributed worldwide except for the polar regions and many Pacific islands. The major- ity of conopids are black with yellow or orange markings and bands and often bear remarkable resemblance to their typical hosts—wasps or bees. Larvae for which biology is known are internal parasites of other insects.

1.1 Diagnosis of the Conopidae

Conopids possess a ptilinum and a combination of apomorphic character states that sharply differentiate them from other Schizophora. The head is large and wider than the thorax; this characteristic indicative of the com- mon name “thick-headed flies” (Hennig 1973). Other apomorphic character states with respect to the ground- plan of the Acalyptrata include: occiput and vertex with short setae; antennal pedicel without a dorsal cleft; precoxal bridges absent; basisternum of prosternum with elongated posterolateral corners (Speight 1969); midcoxal prong absent; R1 bare; R4+5 and M strongly convergent apically and usually fused before wing mar- gin (exceptions occur in the Baltic amber species, Palaeomyopa tertiaria Meunier and in the new subfamily from Australia described below; these have cell r4+5 widely open); vena spuria distinct in many genera; and females usually with highly modified sternites 5 and 6. Other characters useful in defining many acalyptrate families occur in the plesiomorphic state in Conopi- dae: cephalic and thoracic bristles usually poorly differentiated; costa without breaks; Sc complete, not fused with R1; crossvein sc-r present near apex of Sc; vein A1+CuA2 long, reaching or nearly reaching wing mar- gin; males with aedeagal apodeme rod-like (J. McAlpine 1989).

1.2 Current status of conopid taxonomy

1.2.1 Subfamily classification

The subfamily classification of Conopidae is relatively stable with four widely-accepted subfamilies: Conopi- nae, Myopinae, Dalmanniinae and Stylogasterinae. All subfamilies are represented in most geographical regions. Exceptions are discussed in Section 5—Biogeography. Conopinae have a terminal antennal stylus; most have an elongate, strongly sclerotised, anteriorly-projecting haustellum that can reach three times the length of the head and is jointed or geniculate only at the base; labella are short, ovoid and somewhat pointed apically. The other three subfamilies are characterised by an aristate antenna and a second joint near the mid- dle of the haustellum enabling it to fold back under the head at rest. In Stylogasterinae, the haustellum is very