ZOOTAXA

1535

Phylogeny of the Mycetophiliformia, with proposal of the subfamilies Heterotrichinae, Ohakuneinae, and Chiletrichinae for the Rangomaramidae (Diptera, )

DALTON DE SOUZA AMORIM & EIRIK RINDAL

Magnolia Press Auckland, New Zealand Dalton de Souza Amorim & Eirik Rindal Phylogeny of the Mycetophiliformia, with proposal of the subfamilies Heterotrichinae, Ohakuneinae, and Chiletrichinae for the Rangomaramidae (Diptera, Bibionomorpha) (Zootaxa 1535) 92 pp.; 30 cm. 30 July 2007 ISBN 978-1-86977-137-9 (paperback) ISBN 978-1-86977-138-6 (Online edition)

FIRST PUBLISHED IN 2007 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] http://www.mapress.com/zootaxa/

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

2 · Zootaxa 1535 © 2007 Magnolia Press AMORIM & RINDAL Zootaxa 1535: 1–92 (2007) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ ZOOTAXA Copyright © 2007 · Magnolia Press ISSN 1175-5334 (online edition)

Phylogeny of the Mycetophiliformia, with proposal of the subfamilies Heterotrichinae, Ohakuneinae, and Chiletrichinae for the Rangomaramidae (Diptera, Bibionomorpha)

DALTON DE SOUZA AMORIM1,3 & EIRIK RINDAL2 1Departamento de Biologia, FFCLRP, Universidade de São Paulo, Av. Bandeirantes 3900, 14.040-901 Ribeirão Preto SP Brazil. E-mail: [email protected] 2University of Oslo, Natural History Museum, Department of Zoology, P.O. Box 1172 Blindern, 0318 Oslo, Norway. E-mail: [email protected] 3Corresponding author

Table of contents

Abstract ...... 4 Introduction ...... 4 Unusual genera and relationships among the Mycetophiliformia ...... 4 Material ...... 7 Methods ...... 10 New taxa and redescriptions ...... 11 Rangomaramidae Jaschhof & Didham, 2002...... 11 Key for identification of the genera of Rangomaramidae ...... 12 Heterotrichinae, subfam. n...... 13 Chiletrichinae, subfam. n...... 13 Eratomyia, n. gen...... 14 Eratomyia magnifica, sp. n...... 14 Ohakuneinae, subfam. n...... 15 Colonomyia Colless ...... 15 Colonomyia brasiliana, sp. n...... 15 Colonomyia freemani, sp. n...... 16 Colonomyia sp...... 17 Ohakunea Tonnoir & Edwards ...... 18 Ohakunea chilensis Freeman ...... 18 Cabamofa Jaschhof ...... 18 Cabamofa mira Jaschhof ...... 18 Billberg ...... 19 Keroplatoidea Rondani ...... 19 Mycetophiloidea Newman ...... 20 Character analysis ...... 20 Mycetophiliformia major clade relationships ...... 42 A phylogenetic classification of the Mycetophiliformia ...... 47 Acknowledgements ...... 48 References ...... 48 Appendix 1. List of examined specimens ...... 51

Accepted by P. Adler: 16 Mar. 2007; published: 30 Jul. 2007 3 Abstract

A phylogenetic analysis of the Mycetophiliformia (= Sciaroidea) was performed to determine the relationships among its families and to place the following genera of uncertain position in the system: Heterotricha, Ohakunea, Colonomyia, Freemanomyia, Rhynchoheterotricha, Chiletricha, Afrotricha, Anisotricha, Kenyatricha, Nepaletricha, Sciarosoma, Sciaropota, Insulatricha, Cabamofa, Rogambara, and Starkomyia. Eratomyia n. gen. is described based on a new spe- cies from Ecuador. Colonomyia brasiliana sp.n. and Colonomyia freemani sp.n. are described respectively from south- ern Brazil and Chile. The male of Cabamofa mira Jaschhof is described for the first time. A total of 64 terminal taxa and 137 transformation series (with 202 characters) were included in the data matrix, with a number of new features from thoracic morphology. Willi Hennig’s 1973 system for the higher Bibionomorpha was adopted using the name Myceto- philiformia for the Sciaroidea. The Mycetophiliformia are monophyletic. The family Cecidomyiidae appears as the sister group of the remaining Mycetophiliformia, followed by the Sciaridae. In the preferred topology, the Rangomaramidae appear as the group sister of a clade consisting of (Ditomyiidae + Bolitophilidae + Diadocidiidae + Keroplatidae) and of (Lygistorrhinidae + ). The topology within the Rangomaramidae is (Chiletrichinae subfam. n. (Heterotri- chinae subfam. n. ((Rangomaraminae + Ohakuneinae subfam. n.))). The Chiletrichinae include the genera Kenyatricha, Rhynchoheterotricha, Insulatricha, Chiletricha, and Eratomyia n. gen. Heterotrichinae and Rangomaraminae are mono- typic. The subfamily Ohakuneinae includes Ohakunea, Colonomyia, Cabamofa, and Rogambara. The positions of Free- manomyia, Loicia, Taxicnemis, Sciaropota, Starkomyia, Anisotricha, Nepaletricha, and Sciarosoma are considered. Afrotricha might belong to the Sciaridae. The similarities used by many authors to gather the Sciaridae and Mycetophil- idae in a clade are shown to be a combination of plesiomorphies and homoplasies.

Key words: Diptera, phylogeny, Bibionomorpha, Mycetophiliformia, Sciaroidea

Introduction

Amorim & Yeates (2006) revised the classification of the ealier groups of Diptera, formally eliminating the “” and ranking its infraorders as suborders, among which are the Bibionomorpha. It is well accepted that most families of the Bibionomorpha together form a monophyletic group consisting of the Pachyneuridae, Bibionidae, Cecidomyiidae, Sciaridae, Rangomaramidae, Ditomyiidae, Bolitophilidae, Dia- docidiidae, Keroplatidae, Lygistorrhinidae, and Mycetophilidae. The inclusion in this group of the Anisopo- didae s. l., however, has been more questionable. Some authors, such as Tuomikoski (1961), have proposed a separate clade, the Anisopodomorpha, for the family. Wood & Borkent (1989) interpreted some similarities in the mouthparts of the larvae to be shared, derived features with the Psychodidae and other families in the Psy- chodomorpha, whereas Woodley (1989) and Oosterbroek & Courtney (1995) accepted the family as the sister group of the Brachycera. The Scatopsoidea earlier were considered by Hennig (1973) to belong to the Bibion- omorpha, and the Axymyiidae have been included in the group by most authors, but have been transferred to a separate taxon Axymyiomorpha. Whatever the included families, however, the monophyly of the group has never been clearly demonstrated. This paper addresses the relationships within a less inclusive sample of clades, usually referred to by Hennig (1954, 1973) as the Mycetophiliformia, or as the Sciaroidea by more recent authors (e.g., Wood & Borkent 1989, Chandler 2002, Hippa & Vilkamaa 2005, Jaschhof et al. 2005, Hippa & Vilkamaa 2006, Jaschhof 2006).

Unusual genera and relationships among the Mycetophiliformia

The phylogenetic relationships among families of the Mycetophiliformia have remained unsolved, despite the large recent effort after many decades of relatively minor interest on the subject. Not only is there conflict about the position of the Sciaridae and Cecidomyiidae in relation to other families of the group (Bolitophil- idae, Ditomyiidae, Diadocidiidae, Keroplatidae, Lygistorrhinidae, Mycetophilidae, and Rangomaramidae),

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