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 species from Ecuador. Colonomyia brasiliana sp.n. and Colonomyia freemani sp.n. are described respectively from southern 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 Mycetophiliformia 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 + Mycetophilidae). The topology within the Rangomaramidae is (Chiletrichinae subfam. n. (Heterotrichinae subfam. n. ((Rangomaraminae + Ohakuneinae subfam. n.))). The Chiletrichinae include the genera Kenyatricha, Rhynchoheterotricha, Insulatricha, Chiletricha, and Eratomyia n. gen. Heterotrichinae and Rangomaraminae are monotypic. The subfamily Ohakuneinae includes Ohakunea, Colonomyia, Cabamofa, and Rogambara. The positions of Freemanomyia, 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 Mycetophilidae in a clade are shown to be a combination of plesiomorphies and homoplasies.References
Amorim, D.S. (1993) A phylogenetic analysis of the basal groups of Bibionomorpha, with a critical reanalysis of the wing vein homology. Revista brasileira de Biologia, 52, 379–399.
Amorim, D.S. (1994) A new suprageneric classification of the Scatopsidae (Diptera: Psychodomorpha). Iheringia, Zoologia, 77,107–112.
Amorim, D.S. & Yeates, D. (2006) Pesky gnats: ridding dipteran classification of the “Nematocera”. Studia dipterologica, 13(1/2): 3-9.
Amorim, D.S. & Grimaldi, D. (2006) Valeseguyidae, a new family of Diptera in the Scatopsoidea, with a new genus in Cretaceous amber from Myanmar. Systematic Entomology, 31, 508–516.
Amorim, D.S., Bravo, F. & Collucci, E. (1996) Thoracic sclerites: additional information concerning the basal evolution of Diptera (Insecta). XX International Congress of Entomology, Proceedings, Abstract 01-250, Florence, Italy. p. 64.
Billberg, G.J. (1820) Enumeratio insectorum in Museo Gust. Joh. Billberg. Stockholm, pp. 1–138.
Blagoderov, V. (1993) Dipterans (Mesosciophilidae) from the lower Cretaceous of Transbaykal. Paleontological Journal, 27, 123–130.
Blagoderov, V. & Grimaldi D. (2004) Fossil Sciaroidea (Diptera) in Cretaceous ambers, exclusive of Cecidomyiidae, Sciaridae, and Keroplatidae. American Museum Novitates, 3433, 1–76.
Blaschke-Berthold, U. (1994) Anatomie und Phylogenie der Bibionomorpha (Insecta, Diptera). Bonner Zoologische Monographien, 34, 1–206.
Chandler, P. (2002) Heterotricha Loew and allied genera (Diptera: Sciaroidea): offshoots of the stem group of Mycetophilidae and/or Sciaridae? Annales de la Société Entomologique de France (n.s.), 38, 101–144.
Colless, D.H. (1963) New species of Ohakunea and related new genus with notes on the relationships of Heterotricha Loew (Diptera). Proceedings of the Linnean Society of New South Wales, 87, 303–308.
Colless, D.H. & McAlpine, D.K. (1991) 39. Diptera. In: Naumann I.D., Carne P.B., Lawrence J.F., Nielsen E.S., Spradbery J.P., Taylor R.W., Whitten M.J. & Littlejohn M.J. (Eds.), The Insects of Australia. A Textbook for Students and Research Workers, Vol. II, 2nd Edition. Cornell University Press, Ithaca, NY, pp. 717–786.
Edwards, F.W. (1925) Mycetophilidae and Bibionidae (Diptera) in the collections of the South African Museum. Annals of the South African Museum, 19, 601–616.
Edwards, F.W. (1933) New Neotropical Mycetophilidae (II) (Diptera). Revista de Entomologia (Rio de Janeiro), 3, 303–322.
Edwards, F.W. (1940) New Neotropical Mycetophilidae 9IV) (Diptera). Revista de Entomologia, 11, 440–465.
Freeman, P. (1951) Diptera of Patagonia and South Chile. Part III – Mycetophilidae. London: The British Museum (Natural History), 138 pp. plates I–XLIX.
Freeman, P. (1960) A new genus and species of Sciaridae (Diptera, Nematocera) from South Africa. Annals of the Natal Museum, 15, 75–77.
Gagné, R.J. (1981) Cecidomyiidae. In: McAlpine, J.F., Peterson, B.V., Shewell, G.E., Teskey, H.J., Vockeroth, J.R. & Wood, D.M. (Eds.), Manual of Nearctic Diptera. Volume 1. Monograph No. 27. Research Branch, Agriculture Canada, Ottawa, pp. 257–292.
Gagné, R.J. (1994) The Gall Midges of the Neotropical Region. Comstock Publishers, Ithaca and London, 360 pp.
Goloboff, P.A. (1993) Character Optimization and Calculation of Tree Lengths. Cladistics, 9, 433–436.
Grimaldi, D., Amorim, D.S. & Blagoderov, V. (2003) The Mesozoic family Archizelmiridae (Diptera: Insecta). Journal of Paleontology, 77, 368–381.
Hennig, W. (1954) Flügelgeäder und System der Dipteren unter Berücksichtigung der aus dem Mesozoicum beschriebenen Fossilien. Beiträge zur Entomologie, 4, 245–388.
Hennig, W. (1968) Kritische Bemerkungen über den Bau der Flügelwurzel bei den Dipteren und die Frage nach der Monophylie der Nematocera. Stuttgarter Beiträge zur Naturkunde, 193, 1–23.
Hennig, W. (1973) Diptera (Zweiflügler). Handbuch der Zoologie 4(2) 2/31. Berlin-New York: Walter de Gruyter, 335 pp.
Hippa, H. & Jaschhof, M. (2004) A re-evaluation of the genus Colonomyia Colless (Diptera, Sciaroidea), with the description of two new species. Insect Systematics and Evolution, 35, 335–352.
Hippa, H. & Vilkamaa, P. (2005) The genus Sciarotricha gen.n. (Sciaridae) and the phylogeny of recent and fossil Sciaroidea (Diptera). Insect Systematics and Evolution, 36, 121–144.
Hippa, H. & Vilkamaa, P. 2006. Phylogeny of the Sciaroidea: (Diptera): the implication of additional taxa and character data. Zootaxa, 1132, 63–68.
Jaschhof, M. (1998) Revision der “Lestremiinae” (Diptera, Cecidomyiidae, Lestremiinae) der Holarktis. Studia dipterologica Supplement, 4, 1–552.
Jaschhof, M. (2000) Catotrichinae subfam. n.: a re-examination of higher classification in gall midges (Diptera: Cecidomyiidae). Entomological Science, 3, 639–652.
Jaschhof, M. (2004a) The Heterotricha Loew group (Diptera: Sciaroidea) in New Zealand. Beiträge zur Entomologie, 54, 3–30.
Jaschhof, M. (2004b) Freemanomyia Jaschhof nom. nov., a new replacement name for Pterogymnus Freeman, 1951 (Diptera: Sciaroidea). Studia dipterologica, 10, 536.
Jaschhof, M. (2004c) Starkomyia gen.nov. from New Zealand and its implications for the phylogeny of the Sciaroidea (Diptera: Bibionomorpha). Studia dipterologica, 11, 63–74.
Jaschhof, M. (2005) Rogambara and Cabamofa, two new genera of enigmatic sciaroids from Costa Rica (Insecta: Diptera: Sciaroidea). Bonner zoologische Beiträge, 53, 323–332.
Jaschhof, M. & Didham, R.K. (2002) Rangomaramidae fam. nov. from New Zealand and implications for the phylogeny of the Sciaroidea (Diptera: Bibionomorpha). Studia dipterologica Supplement, 11, 1–60.
Jaschhof, M. & Hippa, H. (2003) Sciaroid but not sciarid: a review of the genus Ohakunea Tonnoir & Edwards, with the description of two new species (Insecta: Diptera: Bibionomorpha). Entomologische Abhandlungen, 60, 23–44.
Jaschhof, M., Jaschhof, C., Viklund, B. & Kallweit, U. (2005) [2006] On the morphology and systematic position of Sciarosoma borealis Chandler, based on new material from Fennoscandia (Diptera: Sciaroidea). Studia dipterologica, 12, 231–241.
Krzeminsky, W. & Krzeminska, E. (2003) Triassic Diptera: descriptions, revisions and phylogenetic relations. Acta Zoologica Cracoviensia, 46 (Supplement), 153–184.
Macquart, J. (1838) Insectes diptères nouveaux ou peu connus. Tome premier. –1re partie. Roret, Paris, pp. 1–207, 14 pls.
Malloch, J.R. (1917) A preliminary classification of Diptera, exclusive of Pupipara, based upon larval and pupal characters, with keys to imagines in certain families. Part I. Bulletin of the Illinois State Laboratory of Natural History, 12 (3), 161–409.
Matile, L. (1990) Recherches sur la systématique et l’évolution des Keroplatidae (Diptera, Mycetophiloidea). Mémoires du Muséum national d’Histoire naturelle Paris (A), 148, 1–682.
Matile, L. (1997) Phylogeny and evolution of the larval diet in the Sciaroidea (Diptera, Bibionomorpha) since the Mesozoic. In: Grandcolas, P. (Ed.), The origin of biodiversity in insects: phylogenetic tests of evolutionary scenarios. Mémoires du Muséum national d’Histoire naturelle Paris, 173, 273–303.
Matile, L. & Duret, JP. (1994) Le genre Colonomyia Colless découvert en région Néotropicale (Diptera, Mycetophiloidea). Revue Francaise d'Entomologie (n.s.), 16, 143–147.
McAlpine, J.F. (1981) Morphology and terminology. In: McAlpine, J.F., Peterson, B.V., Shewell, G.E., Teskey, H.J., Vockeroth, J.R. & Wood, D.M. (Eds.), Manual of Nearctic Diptera. Volume 1. Monograph No. 27. Research Branch, Agriculture Canada, Ottawa, pp. 9–63.
McAlpine, J.F., Peterson, B.V., Shewell, G.E., Teskey, H.J., Vockeroth, J.R. & Wood, D. M. (1981) Introduction. Manual of Nearctic Diptera. Volume 1. Monograph No. 27. Research Branch, Agriculture Canada, Ottawa, pp. 1–7.
Meunier, F. (1904) Monographie des Cecidomyidae, des Sciaridae, des Mycetophilidae et des Chironomidae de l'ambre de la Baltique [part]. Annales de la Societé Scientifique de Bruxelles (Mém.), 28, 12–92.
Munroe, D.D. (1974) The systematics, phylogeny, and zoogeography of Symmerus Walker and Australosymmerus Freeman (Diptera: Mycetophilidae: Ditomyiidae). Memoirs of the Entomological Society of Canada, 92, 1–183.
Newman, E. (1834). Attempted division of British insects into natural orders. Entomological Magazine, 2, 379–431.
Oosterbroek, P. & Courtney, G. (1995) Phylogeny of the nematocerous families of Diptera (Insecta). Zoological Journal of the Linnean Society, 115, 267–311.
Papavero, N. (1977) 19E Family Diadocidiidae. In: Papavero, N. (Ed.), A Catalogue of the Diptera of the Americas South of the United States. Museu de Zoologia da Universidade de São Paulo, pp. 1–3.
Papp, L. & Sevcik, J. (2005) Sciarokeroplatinae, a new subfamily of Keroplatidae (Diptera). Acta Zoologica Academiae Scientiarum Hungaricae, 51, 113–123.
Peterson, B.V. (1981) Anisopodidae. In: McAlpine, J.F., Peterson, B.V., Shewell, G.E., Teskey, H.J., Vockeroth, J.R. & Wood, D.M. (Eds.), Manual of Nearctic Diptera. Volume 1. Monograph No. 27. Research Branch, Agriculture Canada, Ottawa, pp. 305–312
Pinto, L.G. & Amorim, D.S. (2000) Bibionidae (Diptera: Bibionomorpha). Morfologia e Análise Filogenetica. [Bibionidae (Diptera: Bibionomorpha). Morphology and Phylogenetic Analysis.]. Holos Editora, Ribeirão Preto, pp. 1–98. [In Portuguese].
Rindal, E. & Søli, G.E.E. (2006) Phylogeny of the subfamily Mycetophilinae (Diptera: Mycetophilidae). Zootaxa,1302, 43–59.
Rohdendorf, B.B. (1962) [Order Diptera. Flies]. In: Rohdendorf, B.B. (Ed.), [Fundamentals of Paleontology]. [Volume 9] [English translation in Rohdendorf, 1991]
Rohdendorf, B.B. (1991) Order Diptera. Two-winged insects. In: Rohdendorf, B.B. (Ed.), Fundamentals of Paleontology. Volume 9. Arthropoda, Tracheata, Chelicerata. Washington, D.C.: Smithsonian Libraries and National Science Foundation [English translation of Rohdendorf, B.B. 1962.]
Rondani, C. (1856) Dipterologiae Italicae prodromus. Vol: I. Genera Italica … A. Stoschi, Parmae [= Parma], pp 1–226 + [2].
Saigusa, T. (1973) A new genus and species of the Ditomyiinae from Celebes. Sieboldia, Acta biologica, 4, 217–224.
Shaw, F.R. (1948) A contribution to the phylogeny of the Mycetophilidae. Annals of the Entomological Society of America, 61, 189–199.
Shaw, F.R. (1953) A review of some of the more important contributions to our knowledge of the systematic relationships of the Sciaridae (Diptera). Proceedings of the Hawaiian Entomological Society, 15, 25–32.
Shaw, F.R. & Shaw, M.M. (1951) Relationships of certain genera of fungus-gnats of the family Mycetophilidae. Smithsonian Miscellaneous Collection, 117(2), 3–23.
Skuse, F.A.A. (1888) Diptera of Australia. Part I. Proceedings of the Linnean Society of New South Wales, (2) 3, 17–145, pls. 2, 3.
Söli, G.E.E. (1997) The adult morphology of Mycetophilidae (s. str.), with a tentative phylogeny of the family (Diptera, Sciaroidea). Entomologica Scandinavica Supplement, 50, 5–55.
Swofford, D.L. (2003) PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4. Sinauer Associates, Sunderland, Massachusetts.
Tonnoir, A.L. & Edwards, F.W. (1927) New Zealand fungus-gnats (Diptera, Mycetophilidae). Transactions of the New Zealand Institute, 57, 747–878, plates 58–80.
Tonnoir, A. L. (1929) Australian Mycetophilidae. Synopsis of the genera. Proceedings of the Linnean Society of New South Wales, 54, 584–614.
Tozoni, S.H.S. (1998) Sistemática Filogenética dos Mycetophilidae (Diptera: Bibionomorpha). Ph.D. thesis, Universidade Federal do Paraná.
Tuomikoski, R. (1961) Zur Systematik der Bibionomorpha (Dipt.) I. Anisopodidae und Protorhypidae. Annales Entomologici Fennici, 27, 65–69.
Tuomikoski, R. (1966) Systematic position of Lygistorrhina Skuse (Diptera, Mycetophiloidea). Annales Entomologici Fennici, 32, 254–260.
Vilkamaa, P. & Hippa, H. (2004) The genus Xenosciara gen. n. and the phylogeny of the Sciaridae (Diptera). Zootaxa, 699, 1–24.
Vilkamaa, P. (2000) Phylogeny of Prosciara Frey and related genera (Diptera: Sciaridae). Systematic Entomology, 25, 47–72.
Vockeroth, R. (1981) Mycetophilidae. In: McAlpine, J.F., Peterson, B.V., Shewell, G.E., Teskey, H.J., Vockeroth, J.R. & Wood, D.M. (Eds.), Manual of Nearctic Diptera. Volume 1. Monograph No. 27. Research Branch, Agriculture Canada, Ottawa, pp. 223–246.
Wood, D.M. & Borkent, A. (1989) Phylogeny and classification of the Nematocera. In: McAlpine, J.F., Peterson, B.V., Shewell, G.E., Teskey, H.J., Vockeroth, J.R. & Wood, D.M. (Eds.), Manual of Nearctic Diptera. Volume 1. Monograph No. 27. Research Branch, Agriculture Canada, Ottawa, pp. 1333–1370.
Woodley, N.E. (1989) Phylogeny and classification of "orthorrhaphous" Brachycera. In: McAlpine, J.F., Peterson, B.V., Shewell, G.E., Teskey, H.J., Vockeroth, J.R. & Wood, D.M. (Eds.), Manual of Nearctic Diptera. Volume 1. Monograph No. 27. Research Branch, Agriculture Canada, Ottawa, pp. 1371–1395.