PJOTR OOSTERBROEK & Br. THEOWALD Institute of Taxonomie Zoology, Amsterdam PHYLOGENY OF THE TIPULOIDEA BASED ON CHARACTERS OF LARVAE AND PUPAE (DIPTERA, NEMATOCERA) with an index to the literature except Tipulidae Oosterbroek, P. & Br. Theowald, 1991. Phylogeny of the Tipuloidea based on characters of larvae and pupae (Diptera, Nematocera), with an index to the literature except Tipulidae. -Tijdschrift voor Entomologie 134: 211-267, figs. 1-180. [ISSN 0040-7496]. Published 18 December 1991. Phylogenetic relationships of the Tipuloidea (the families Limoniidae, Cylindrotomidae and Tipulidae) are discussed on the basis of larval and pupal characters. In comparing outgroups, special attention is given to the synapomorphies listed by Wood & Borkent (1989) for the infraorder Psychodomorpha. In total 105 characters of larvae and pupae are evaluated. A phylogeny is presented for the Tipuloidea as well as for the genera of the subfamilies Eriopterinae, Hexatominae, Pediciinae and Limoniinae. An index to the literature on the immature stages except Tipulidae is added. Correspondence: Dr. P. Oosterbroek, University of Amsterdam, Institute of Taxonomie Zoology, Department of Entomology, Plantage Middenlaan 64, 1018 DH Amsterdam, The Netherlands. Keywords. -Phylogeny, Tipuloidea, Psychodomorpha, Nematocera, larvae, pupae. Contents 1. Introduction 211 2. Methods 212 3. Outgroups 213 4. Tipuloidea 217 5. Eriopterinae 232 6. Hexatominae 238 7. Pediciinae 242 8. Limoniinae 245 9. Remaining genera 245 10. Index to the literature 252 1 1 . Acknowledgements 260 12. References 260 i. Introduction The primary concern of this paper are the char-acters of larvae and pupae of Tipuloidea and their contribution to our understanding of phylogenetic relationships among and within the major groups. Translation of the available information into a for-mal classification is considered premature for sev-eral reasons. Characters of the adults are not yet evaluated phylogenetically. Moreover, immature stages are known for representative species of about 150 genera and subgenera, mainly from the Northern Hemisphere, which is about one-third of the recognized genera and subgenera worldwide. Progress in our knowledge of the immature stages of Southern Hemisphere taxa is very important and might well influence certain aspects of the postulated relationships. Tipuloidea as discussed in this paper is identical to: (a) the superfamily Tipuloidea sensu Hennig ( 1973), comprising the families Limoniidae, Cylin-drotomidae and Tipulidae, (b) the family Tipulidae s.l. of most North American and British workers, who usually recognize one family with three sub-families, (c) the infraorder Tipulomorpha sensu Wood & Borkent (1989). The family Limoniidae is, with about 10,000 described species, one of the largest families of Diptera. Five subfamilies are usually recognized. Immature stages of one subfamily, the Lechriinae (three genera with a limited number of species in Africa, SE Asia and Australia) are not known. The other four subfamilies, Eriopterinae, Hexatomi-nae, Pediciinae and Limoniinae, are discussed 211
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