Zootaxa, Empidoidea (Diptera)
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ZOOTAXA 1180 The morphology, higher-level phylogeny and classification of the Empidoidea (Diptera) BRADLEY J. SINCLAIR & JEFFREY M. CUMMING Magnolia Press Auckland, New Zealand BRADLEY J. SINCLAIR & JEFFREY M. CUMMING The morphology, higher-level phylogeny and classification of the Empidoidea (Diptera) (Zootaxa 1180) 172 pp.; 30 cm. 21 Apr. 2006 ISBN 1-877407-79-8 (paperback) ISBN 1-877407-80-1 (Online edition) FIRST PUBLISHED IN 2006 BY Magnolia Press P.O. Box 41383 Auckland 1030 New Zealand e-mail: [email protected] http://www.mapress.com/zootaxa/ © 2006 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 1180: 1–172 (2006) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ ZOOTAXA 1180 Copyright © 2006 Magnolia Press ISSN 1175-5334 (online edition) The morphology, higher-level phylogeny and classification of the Empidoidea (Diptera) BRADLEY J. SINCLAIR1 & JEFFREY M. CUMMING2 1 Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany. E-mail: [email protected] 2 Invertebrate Biodiversity, Agriculture and Agri-Food Canada, C.E.F., Ottawa, ON, Canada K1A 0C6. E-mail: [email protected] Table of Contents Abstract ..............................................................................................................................................4 Introduction ........................................................................................................................................4 Materials and Methods.......................................................................................................................6 Materials......................................................................................................................................6 Cladistic analysis.........................................................................................................................7 Taxon sampling ...........................................................................................................................7 Monophyly of the Eremoneura ........................................................................................................15 Historical Review of the Phylogeny of the Empidoidea..................................................................16 Description of Characters in Cladistic Analysis .............................................................................20 Adult stage ................................................................................................................................21 Immature stages ........................................................................................................................58 Characters not included in cladistic analysis ............................................................................60 Results of the Cladistic Analysis......................................................................................................62 Higher-level relationships .........................................................................................................63 Lower-level relationships in Empidoidea .................................................................................66 Reflections on Feeding Habits and Ground Plan Condition ............................................................69 Proposed Classification of the Empidoidea .....................................................................................71 Key to the Families and Unplaced Genus Groups of Empidoidea...................................................82 Acknowledgments............................................................................................................................84 References ........................................................................................................................................85 Table 1. Exemplar taxa scored in the analysis .................................................................................98 Table 2. Character state matrix for cladistic analysis ....................................................................101 Table 3. Proposed classification of the Empidoidea ......................................................................103 Trees ...............................................................................................................................................104 Plates .............................................................................................................................................. 110 Accepted by N. Evenhuis: 25 Feb. 2006; published: 21 Apr. 2006 3 ZOOTAXA Abstract 1180 A cladistic analysis of the Empidoidea and basal lineages of the Cyclorrhapha, based on morphological characters, confirms the monophyly of both groups as well as that of the Eremoneura. The resulting final trees are used to revise the classification of the Empidoidea to include the following five families: Empididae, Hybotidae, Atelestidae (including Nemedininae n. subfam.), Brachystomatidae rev. stat. (comprising the subfamilies Brachystomatinae, Ceratomerinae and Trichopezinae), and Dolichopodidae s.lat. The family Microphoridae is not recognized, and the Microphorinae and Parathalassiinae are assigned to the Dolichopodidae s.lat. The Dolichopodidae s.str. includes 15 subfamilies that were previously recognized within the family. Within the Empidoidea we found support for Atelestidae as the sister group to the Hybotidae and for the monophyly of Parathalassiinae + Dolichopodidae s.str. The Empididae remains poorly defined and the genera Homalocnemis Philippi, Iteaphila Zetterstedt, Anthepiscopus Becker, and Oreogeton Schiner are classified as incertae sedis within the Empidoidea. In addition, the following higher taxa are proposed: Symballophthalmini n. tribe, Bicellariini n. tribe, Oedaleinae rev. stat., and Trichininae rev. stat., which are all assigned to the Hybotidae. The genus Sematopoda Collin is tentatively assigned to Trichopezinae, and Xanthodromia Saigusa is transferred from Hemerodromiinae to Brachystomatinae. All morphological characters are extensively discussed and illustrated, including details of the antennae, mouthparts, internal thoracic structures, wings, and male and female terminalia. In addition, a key to families and unplaced genus groups of the Empidoidea is provided. Feeding habits are also discussed in terms of the empidoid ground plan condition. Key words: dance flies, long-legged flies, Empidoidea, Empididae, Hybotidae, Atelestidae, Brachystomatidae, Dolichopodidae, phylogeny, cladistics, morphology, genitalia, mouthparts, new subfamily, new tribes Introduction An estimated 11,400 species are described in the Empidoidea, making it among the largest of higher categories of Diptera (Thompson 2005). Fossils with empidoid-like venation are known from the upper Jurassic (Mostovski 1999), with the empidoid subfamilies present by the early Cretaceous (Grimaldi 1999; Grimaldi & Cumming 1999). In fact, the Empidoidea are among the best known lineages from the Cretaceous (Grimaldi 1999). Divergence time estimates for the Empidoidea range between 144–163 MYA (Wiegmann et al. 2003). They occur worldwide (except Antarctica), with certain lineages being particularly abundant or more diverse in temperate latitudes. For example, we have identified several “empidoid hotspots” (exclusive of Dolichopodidae s.str.) based on total number of described endemic genera, among which are included, New Zealand (13 endemic genera), South Africa (9), southern South America (11) and western North America (9). The tremendous species diversity of the Empidoidea corresponds to an enormous morphological or structural diversity (Figs. 417–424), especially in the male genitalia. The 4 © 2006 Magnolia Press SINCLAIR & CUMMING modifications range from symmetrical, unrotated, relatively simple genitalia (somewhat ZOOTAXA similar to asiloids) to asymmetrical and rotated (with secondary fusion of various sclerites) 1180 and occasionally secondary symmetry, which has often caused confusion and difficulties in determining homology. This wonderful and complex genitalic diversity was a major impetus for our earlier studies on male genitalia (Sinclair et al. 1994; Cumming et al. 1995). There are also extensive female and male (primarily) secondary sexual characters found on all body parts, particularly on the legs. In addition, there is a tremendous range in size, from the very large Empis (Planempis) pan Frey (wing length nearly 12 mm) to the tiny genus Enlinia Aldrich (wing length 0.8 mm: see Robinson 1969). Empidoids breed in a variety of habitats, including running water (e.g., Wagner & Gathmann 1996), tidal zones, decaying wood (e.g., Meyer 2005), and moist soil. Adults are predators or flower visitors (feeding on nectar and/or pollen) and nearly all known larvae are predators (except Thrypticus Gerstäcker, secondarily phytophagous in stems of monocots). Investigations on the potential impact of predaceous empidoids acting as biocontrol agents on agricultural pests are only in the initial stages (see Cumming & Cooper