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Hymenoptera, Braconidae), with a Molecular Characterization of Hidden Fennoscandian Species Diversity Zootaxa 3084: 1–95 (2011) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Monograph ZOOTAXA Copyright © 2011 · Magnolia Press ISSN 1175-5334 (online edition) ZOOTAXA 3084 Revision of the Western Palearctic Meteorini (Hymenoptera, Braconidae), with a molecular characterization of hidden Fennoscandian species diversity JULIA STIGENBERG & FREDRIK RONQUIST Department of Zoology, Stockholm University and Department of Entomology, Swedish Museum of Natural History, Box 50007, SE-10405 Stockholm, Sweden; e-mail: [email protected] Department of Biodiversity Informatics, Swedish Museum of Natural History, Box 50007, SE-10405 Stockholm, Sweden; e-mail: [email protected] Magnolia Press Auckland, New Zealand Accepted by J. Jennings: 29 Aug. 2011; published: 28 Oct. 2011 JULIA STIGENBERG & FREDRIK RONQUIST Revision of the Western Palearctic Meteorini (Hymenoptera, Braconidae), with a molecular characterization of hidden Fennoscandian species diversity (Zootaxa 3084) 95 pp.; 30 cm. 28 Oct. 2011 ISBN 978-1-86977-817-0 (paperback) ISBN 978-1-86977-818-7 (Online edition) FIRST PUBLISHED IN 2011 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] http://www.mapress.com/zootaxa/ © 2011 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) 2 · Zootaxa 3084 © 2011 Magnolia Press STIGENBERG & RONQUIST Table of contents Abstract . 3 Introduction . 4 Material and methods . 7 Studied Material . 7 Terminology and Illustrations . 8 Species descriptions. 8 Morphological Phylogenetic Analysis. 8 Molecular Analysis . 8 Analysis of Species Circumscription . 10 Results . 10 Morphological Phylogenetic Analysis. 10 Molecular and Combined Phylogenetic Analyses. 13 Species Circumscription . 13 Evolution of Host Preferences . 19 Discussion . 19 Phylogeny . 19 Species Circumscription . 24 Evolution of Host Preferences . 25 Taxonomic Treatment with Keys. 26 The Tribe Meteorini Cresson 1887 . 26 Key to European genera of Meteorini. 26 Key to European species of Zele . 26 Key to European species of Meteorus . 26 Genus Zele Curtis . 40 Genus Meteorus Haliday. 43 Acknowledgements . 75 References . 85 Appendix I—Synonymic list of species . 90 Appendix II—Voucher specimens and GenBank accession numbers. 92 Abstract The tribe Meteorini includes two genera, Meteorus and Zele, which are koinobiont endoparasitoids of larval Lepidoptera and Coleoptera. More than 300 species are known, about one fifth of which occur in the Western Palearctic. Here, we re- vise the Western Palearctic species, based partly on traditional approaches and partly on molecular analysis of recent Swedish and Finnish material. For the analyses of phylogenetic relationships and cryptic species diversity, we coded 17 morphological characters and sequenced two markers, 28S D2 (649 bp) and CO1 (665 bp). More than 1 970 specimens representing 54 species of Meteorus Haliday and 5 species of Zele Curtis were studied; of these, 177 specimens represent- ing 41 species were sequenced. Seven new species are described, all from the Fennoscandian material: Meteorus artocer- cus sp. nov., M. densipilosus sp. nov., M. eklundi sp. nov., M. longipilosus sp. nov., M. sibyllae sp. nov., M. stenomastax sp. nov., and M. subtilisulcus sp. nov. Four new synonyms are introduced: Z. chlorophthalmus (Spinola 1808), syn. nov. for M. pallidus (Nees 1812), M. punctifrons Thomson 1895, syn. nov. for M. varinervis (Tobias 1986), M. melanostictus Capron 1887, syn. nov for M. monachae (Tobias 1986), and M. tenellus Marshall 1887, syn. nov. for M. boreus (Tobias 1986). Meteorus tenellus is removed from synonymy with M. cinctellus. Sequence analysis indicated the presence of at least 12 additional cryptic species but these cannot be separated morphologically at this point and, therefore, we do not describe them here. The phylogenetic results suggest that Zele should be included within Meteorus but we refrain from formal changes of the generic classification until more comprehensive phylogenetic analyses of the tribe can be complet- ed. A key to the known Western Palearctic species is presented. Key words: parasitic wasps, molecular taxonomy, cryptic species, general mixed Yule coalescent, evolution, host prefer- ences, identification keys REVISION OF THE WESTERN PALEARCTIC METEORINI Zootaxa 3084 © 2011 Magnolia Press · 3 Introduction The Meteorini (Cresson 1887) are small to medium size (2–10 mm) euphorine braconids, classified into two gen- era: the species-rich and cosmopolitan Meteorus Haliday (1832) and the small Holarctic and Neotropical Zele Cur- tis (1835). After accounting for the new species described herein, the two genera together include 337 species in the world (308 Meteorus and 29 Zele, Yu et al. 2005, Shaw et al. 2009, Aguirre et al. 2010, 2011), 59 of which occur in the Western Palearctic (54 Meteorus and 5 Zele) (Appendix I). Unlike other euphorine braconids that largely attack adult insects, the Meteorini target Lepidoptera and Coleoptera larvae. The host larvae are typically hidden under bark, in wood, or in fungi (Maeto 1990a, Shaw et al. 1991). Most meteorines are solitary parasitoids but some species are gregarious. They attack their hosts in the same way as most other euphorines, namely by bending the abdomen under the thorax and pushing the ovipositor out so that it projects forwards, in front of the wasp. They seem to preferably attack mobile host larvae (Parker 1931, Tobias 1966). Oviposition occurs in young host larvae, while they are exposed, and development is completed before the host enters the pupal stage. Members of Meteorus are either univoltine or multivoltine. Some species produce two complete generations upon one brood of host larvae, the first being upon very young individuals and the second upon those that are nearly mature (Clausen 1940). The solitary suspended cocoon made by some Meteorus larvae is unique amongst the Braconidae and only the Campopleginae amongst the Ichneumonidae have members that construct similar cocoons. No other hymenopterans are known to form suspended cocoons (Zitani 2003). The suspended cocoons are fully exposed whilst the non-suspended cocoons often are placed in a concealed environment. Members of Zele seem to be solitary parasitoids. In Britain, at least one species is noted to be bivoltine whilst others are univoltine (Shaw & Huddleston 1991). They overwinter as prepupae in a fusiform and felted cocoon in the pupation chamber of the host. Zele adults are predominantly crepuscular or nocturnal and are also attracted to light (Huddleston and Gauld 1988). In both genera, the adult emerges from the cocoon by cutting the cap off of its anterior end. The circumscription of the Meteorini has been a subject of controversy for some time (van Achterberg 1979, Huddleston 1980, Zitani 2003). Mason (1973), in his paper on the identity of Zele (treated by him under the junior synonym Zemiotes Förster), proposed that the Meteorini formed an unnatural assemblage and that Zele was more closely related to Homolobus Förster (subfam. Homolobinae) than to Meteorus. Mason based this hypothesis on the following similarities shared between Homolobus and Zele: (1) radiellan cell broadening apically and divided by a transverse vein; (2) tarsal claws with a large basal tooth; (3) apical half or more of abdominal tergites 3-6 uni- formly hairy, never with only a single, subapical, transverse row on each tergite; and (4) the cocoon covered with fine, fluffy witish silk and never suspended on a cord or covered in coarse brown silk as in Meteorus. In his revi- sion of the subfamily Zelinae, van Achterberg (1979) redefined Zele and moved it to the tribe Meteorini of the sub- family Euphorinae. He thoroughly studied the similarities between Zele and Homolobus listed by Mason (1973), as well as other.
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