ZOOTAXA

857

Systematics and phylogeny of (Diptera: )

SCOTT E. BROOKS

Magnolia Press Auckland, New Zealand SCOTT E. BROOKS Systematics and phylogeny of Dolichopodinae (Diptera: Dolichopodidae) (Zootaxa 857) 158 pp.; 30 cm. 16 Feb. 2005 ISBN 1-877354-84-8 (Paperback) ISBN 1-877354-85-6 (Online edition)

FIRST PUBLISHED IN 2005 BY Magnolia Press P.O. Box 41383 Auckland 1030 New Zealand e-mail: [email protected] http://www.mapress.com/zootaxa/

© 2005 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) Zootaxa 857: 1–158 (2005) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ ZOOTAXA 857 Copyright © 2005 Magnolia Press ISSN 1175-5334 (online edition)

Systematics and phylogeny of Dolichopodinae (Diptera: Dolichopodidae)

SCOTT E. BROOKS Department of Natural Resource Sciences, McGill University, Ste-Anne-de-Bellevue, Quebec, CANADA; Present Address: Diptera Unit, Canadian National Collection of , Invertebrate Biodiversity, Agriculture and Agri-Food Canada, K.W. Neatby Building, 960 Carling Avenue, Ottawa, Ontario, K1A 0C6, CANADA; email: [email protected]

Table of contents

Abstract ...... 4 Introduction ...... 5 Materials and Methods ...... 8 Results and Discussion ...... 11 Subfamily Dolichopodinae ...... 26 Preliminary key to world genera of Dolichopodinae ...... 27 Genera of Dolichopodinae ...... 34 Genus Allohercostomus Yang, Saigusa & Masunaga ...... 34 Genus Anasyntormon Parent ...... 37 Genus Argyrochlamys Lamb ...... 38 Genus Cheiromyia Dyte ...... 43 Genus Latreille ...... 46 New Genus A ...... 56 Genus Loew ...... 60 Genus Loew ...... 64 Genus Metaparaclius Becker ...... 74 Genus Muscidideicus Becker ...... 77 Genus Latreille ...... 80 Genus Loew ...... 83 Genus Parahercostomus Yang, Saigusa & Masunaga ...... 89 Genus Loew ...... 92 Genus Platyopsis Parent ...... 100 Genus Mik ...... 102 Genus Prohercostomus Grichanov ...... 107 Genus Stenopygium Becker ...... 109 Genus Meigen ...... 112 Genus Haliday ...... 120 Genera Removed from Dolichopodinae ...... 132 Genus Colobocerus Parent ...... 132 Genus Katangaia Parent ...... 133

Accepted by N.L. Evenhuis: 12 Jan. 2005; published: 16 Feb. 2005 3 ZOOTAXA Genus Pseudohercostomus Stackelberg ...... 134 857 Genus Vetimicrotes Dyte ...... 137 Acknowledgments ...... 137 References ...... 139 Appendix 1. Characters and character codings used in the analysis ...... 150 Appendix 2. Character state matrix for Dolichopodinae analysis ...... 156 Appendix 3. List of abbreviations used in figures ...... 158

Abstract

The phylogenetic relationships of the subfamily Dolichopodinae was investigated based on the examination of over 340 species of Dolichopodinae from all zoogeographic regions. Sixty-five exemplar species were included in a cladistic analysis based on 74 morphological characters of adult specimens. Twenty genera are recognized in the Dolichopodinae: Allohercostomus Yang, Sai- gusa & Masunaga, Anasyntormon Parent, Argyrochlamys Lamb, Cheiromyia Dyte, Dolichopus Latreille, Gymnopternus Loew, Hercostomus Loew, Metaparaclius Becker, Muscidideicus Becker, Ortochile Latreille, Paraclius Loew, Parahercostomus Yang, Saigusa & Masunaga, Pelastoneurus Loew, Platyopsis Parent, Poecilobothrus Mik, Prohercostomus Grichanov, Stenopygium Becker, Sybistroma Meigen, Tachytrechus Haliday, and New Genus A. Eleven genera are newly synony- mized: Halaiba Parent (=Argyrochlamys Lamb); Lichtwardtia Enderlein (=Dolichopus Latreille); Phalacrosoma Becker (=Hercostomus Loew); Steleopyga Grootaert & Meuffels (=Hercostomus Loew); Proarchus Aldrich (=Pelastoneurus Loew); Sarcionus Aldrich (=Pelastoneurus Loew); Mik (=Poecilobothrus Mik); Ludovicius Rondani (=Sybistroma Meigen); Nodicornis Rondani (=Sybistroma Meigen); Gonioneurum Becker (=Tachytrechus Haliday); Syntomoneurum Becker (=Tachytrechus Haliday). The following new generic combinations are established: Argyro- chlamys breviseta (Parent), Argyrochlamys cavicola (Parent), Cheiromyia maculipennis (Van Duzee), Dolichopus angulicornis (Grichanov), Dolichopus clypeatus (Grichanov), Dolichopus emelyanovi (Grichanov), Dolichopus fractinervis (Parent), Dolichopus hilgerae (Grichanov), Doli- chopus hollisi (Grichanov), Dolichopus maculatus (Parent), Dolichopus minusculus (Parent), Doli- chopus mironovi (Grichanov), Dolichopus nigrifacies (Grichanov), Dolichopus nigrotorquatus (Parent), Dolichopus nikolaevae (Grichanov), Dolichopus sukharevae (Grichanov), Dolichopus tikhonovi (Grichanov), Hercostomus amoenus (Becker), Hercostomus argyreus (Wei & Lui), Her- costomus briarea (Wei & Lui), Hercostomus dactylocera (Grootaert & Meuffels), Hercostomus ful- gidipes (Becker), Hercostomus hubeiensis (Yang), Hercostomus imperfectus (Becker), Hercostomus postiseta (Yang & Saigusa), Hercostomus zygolipes (Grootaert & Meuffels), Poecilo- bothrus aberrans (Loew), Poecilobothrus chrysozygos (Wiedemann), Prohercostomus bickeli (Evenhuis), Prohercostomus interceptus (Meunier), Prohercostomus intremulus (Meunier), Proher- costomus meunierianus (Evenhuis), Prohercostomus monotonus (Meunier), Prohercostomus nego- tiosus (Meunier), Prohercostomus notabilis (Meunier), Prohercostomus noxialis (Meunier), Prohercostomus vulgaris (Meunier), Stenopygium punctipennis (Say), Sybistroma acutatus (Yang), Sybistroma apicicrassus (Yang & Saigusa), Sybistroma apicilarius (Yang), Sybistroma biaristatus (Yang), Sybistroma biniger (Yang & Saigusa), Sybistroma bogoria (Grichanov), Sybistroma brevi- digitatus (Yang & Saigusa), Sybistroma crinicauda (Zetterstedt), Sybistroma curvatus (Yang), Sybistroma digitiformis (Yang, Yang & Li), Sybistroma dorsalis (Yang), Sybistroma emeishanus (Yang), Sybistroma eucerus (Loew), Sybistroma fanjingshanus (Yang, Grootaert & Song), Sybis- troma flavus (Yang), Sybistroma golanicus (Grichanov), Sybistroma henanus (Yang), Sybistroma

4 © 2005 Magnolia Press BROOKS impar (Rondani), Sybistroma incisus (Yang), Sybistroma inornatus (Loew), Sybistroma israelensis ZOOTAXA (Grichanov), Sybistroma longiaristatus (Yang & Saigusa), Sybistroma longidigitatus (Yang & Sai- 857 gusa), Sybistroma lorifer (Mik), Sybistroma luteicornis (Parent), Sybistroma miricornis (Parent), Sybistroma neixianganus (Yang), Sybistroma qinlingensis (Yang & Saigusa), Sybistroma sciophil- lus (Loew), Sybistroma sheni (Yang & Saigusa), Sybistroma sichuanensis (Yang), Sybistroma sinaiensis (Grichanov), Sybistroma spectabilis (Parent), Sybistroma sphenopterus (Loew), Sybis- troma transcaucasius (Stackelberg), Sybistroma yunnanensis (Yang), Tachytrechus alatus (Becker), Tachytrechus analis (Parent), Tachytrechus beckeri (Parent), Tachytrechus giganteus (Brooks), Tachytrechus varus (Becker). Pelastoneurus lineatus de Meijere, 1916, junior secondary homonym of Pelastoneurus lineatus Aldrich, 1896, is given the new replacement name Pelastoneurus neolin- eatus nom. nov. Four genera are excluded from the subfamily including Colobocerus Parent, Katangaia Parent, Pseudohercostomus Stackelberg and Vetimicrotes Dyte. A key to the world gen- era of Dolichopodinae is provided.

Key words: Dolichopodidae, Dolichopodinae, phylogeny, classification, morphology

Introduction

Dolichopodidae are one of the most abundant, widespread and diverse families of Diptera. Recent estimates indicate that there are over 6600 described species in 200 genera (Grich- anov 1999b and unpublished data). Dolichopodids are found in all terrestrial habitats from forests to agricultural fields; however, they are particularly diverse and abundant in wet habitats such as humid forests and shores of water bodies. Both larvae and adults of most species are predators on a variety of small insects and other invertebrates. Over the last century the Dolichopodidae has been variously divided into subfamilies both regionally and at a world scale (Lioy 1863–1864; Schiner 1864; Aldrich 1905; Kertész 1909; Becker 1917–1918, 1922a; Robinson 1970a, 1970b; Ulrich 1981; Negrobov 1986). The Dolichopodinae are one of the most diverse subfamilies of Dolichopodidae. Doli- chopodines occur worldwide and currently there are about 1700 described species, com- prising approximately 25 percent of the species in the entire family. Depending on which of the previously proposed classifications is followed, the number of genera included in the subfamily ranges from about 23 to 37. Most of the dolichopodine diversity is divided among six large genera including Dolichopus Latreille, with about 600 species, Hercosto- mus Loew, with about 470 species (not including species of Gymnopternus Loew, which are sometimes included within Hercostomus in Old World classifications), Tachytrechus Haliday, with about 140 species, and Paraclius Loew, Gymnopternus and Pelastoneurus Loew, with over 100 species each. These genera together comprise about 90% of described dolichopodine species. Dolichopodines are recognized by the possession of a patch of setae on the dorsal surface of the antennal scape, one or more distinct anterior or anterodorsal preapical setae on the mid and hind femora, male abdominal tergite 6 usually bare, and by the typically large, pedunculate genital capsule of males, which projects for- ward under the preceding abdominal segments.

DOLICHOPODINAE © 2005 Magnolia Press 5