Phylogenetic Systematics of the Milichiidae (Diptera, Schizophora)
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IRINA BRAKE Phylogenetic systematics of the Milichiidae (Diptera, Schizopbora) Entomologica scandinavica Supplement No. 57 now: Insect Systematics & Evolution J1:ntomologica scandinavica Supplements also supplement to: Insect Systematics & Evolution An international journal of systematic entomology owned by the Scandinavian Society of Entomology and sponsored by the national councils for natural science research in Denmark, Finland, Norway and Sweden. Editor-ill-chief: Dr. N. M\'lller Andersen, Zoological Museum, University of Copenhagen, Denmark. Managing editor: Dr. Lennart Cederholm, Museum of Zoology, University of Lund, Sweden. Editorial advisers: Dr. Gary A. P. Gibson, Centre for Land and BioI. Resources Research, Ottawa, Canada. Dr. Jyrki Muona, Oulanka BioI. Station, University of Oulu, Finland. Dr. E. Schmidt Nielsen, CSIRO, Division of Entomology, Canberra, Australia. Dr. Norman 1. Platnick, American Museum of Natural History, New York, U.S.A. (:oUllcii of the Scandinavian Society of Entomology Dr. Ilcnrik Enghoff, Entomologisk Forening, Copenhagen, Denmark. Prof. Marlin Meinander, (Chairman), Entomologiska Foreningen, Helsinki, Finland. Prof. I,auritz Somme, Norsk Entomologisk Forening, Oslo, Norway. Dr. Per Douwes, Entomologiska Sallskapet, Lund, Sweden. (;eneral aims J<:n[(lll1ologica scandinavica Supplement is issued by the Scandinavian Society of Entomology for original papers on systematics and taxonomic entomology. They appear at itTegular intervals, normally 2-3 vols. per year. Standing orders are accepted. Normally, the authors are required to meet the costs a/publication. Editorial office: Vastervang 28, S-247 34 S. Sandby, Sweden. Phone +46/(0)46-579 69. +46/(0)46-57969. E-mail:[email protected] Distributor: Scandinavian Entomology Ltd., Vastervang 28, S-247 34 S. Sandby, Sweden. Fax +46/(0)46-579 69. A catalogue of all publications is found on Internet: http://www.bm.lu.se~zool/ ZooMus.html Secondary Journals: EnL scand. Supplement is covered by Zoo!. Record, Ent. Abstr., Review of App!. Enl. (selectively) and CUlTent Contents. © Entomologica scandinavica Supplement 2000 Entomologica Scandinavica Supplement 57 Phylogenetic systematics of the Milichiidae (Diptera, Schizophora) IRINA BRAKE 2 Irina Brake ENT. SCAND. SUPPL. 57 (2000) Contents 1. Introduction .................................................................................................................................................... 3 1.1. Historical review of research on the Milichiidae .................................................................................. 4 1.2. Historical review of the relationship between the Milichiidae, Camidae and Chloropidae ................. 5 1.3. Biology ............................................................................................................................................. 5 1.4. Geographic distribution ......................................................................................................................... 8 2. Materials and methods .................................................................................................................................... 8 3. Cladistic analysis .......................................................................................................................................... 14 3.1. Characters used for cladistic analysis ................................................................................................. 14 3.2. Cladogram ........................................................................................................................................... 17 4. Morphology and character transfOlmations .................................................................................................. 17 4.1. Preliminary remarks ............................................................................................................................ 17 4.2. Head .................................................................................................................................................... 19 4.2.1. Postfrons .................................................................................................................................... 19 4.2.2. Prefrons ...................................................................................................................................... 27 4.2.3. Compound eyes .......................................................................................................................... 29 4.2.4. Proboscis .................................................................................................................................... 29 4.3. Thorax ................................................................................................................................................. 34 4.3.1. Prothorax .................................................................................................................................... 34 4.3.2. Mesothorax ................................................................................................................................ 39 4.3.3. Wings ......................................................................................................................................... 39 4.3.4. Legs ........................................................................................................................................... 42 4.4. Abdomen ........................................................................................................................................... 44 4.4.1. Male preabdomen ...................................................................................................................... 44 4.4.2. Male pregenital segments .......................................................................................................... 46 4.4.3. Male genitalia ............................................................................................................................ 47 4.4.4. Male reproductive system .......................................................................................................... 61 4.4.5. Female ovipositor ...................................................................................................................... 61 4.4.6. Female reproductive system ...................................................................................................... 65 4.5. Immature stages .................................................................................................................................. 72 4.5.1. Eggs ........................................................................................................................................... 72 4.5.2. Larvae and puparia ..................................................................................................................... 74 4.6. Stem-species pattem of Milichiidae .................................................................................................... 74 5. Phylogeny of the Milichiidae ....................................................................................................................... 75 5.1. Phyllomyzinae ..................................................................................................................................... 75 5.2. Madizinae ........................................................................................................................................... 77 5.3. Milichiinae .......................................................................................................................................... 78 5.4. Minimum age of the Phyllomyzinae, Madizinae, and Milichiinae ..................................................... 79 5.5. Evolutionary "trends" in the biology of the Milichiidae ..................................................................... 80 6. Phylogenetic position of the Milichiidae within the Schizophora ................................................................ 82 6.1. Comparison with traditional classifications ........................................................................................ 82 7. Acknowledgements ...................................................................................................................................... 83 8. References .................................................................................................................................................... 84 9. Appendix. Key to genera .............................................................................................................................. 89 10. Appendix. Catalogue of world Milichiidae ................................................................................................ 92 References ................................................................................................................................................ 109 Index ......................................................................................................................................................... 113 The illustration on the coverof a Litometopa sp. 1. is made by the author. Phylogenetic systematics of the Milichiidae (Diptera, Schizophora) II(INA BRAKE Brake, I.: Phylogenetic systematics of the Milichiidae (Diptera, Schizophora). Ent.scand. Supp!. 57:1-120. Lund, Sweden 2000-12-15. ISSN 0105-3574. A phylogenetic tree based on adult characters is proposed