Abstract
The phylogenetic relationships of the chloropid tribe Elachipterini were analysed. Sixty-eight exemplar species and seven outgroup species were included in a cladistic analysis based on 76 morphological characters of adult specimens in order to test existing, non-phylogenetic, classifications of the tribe. Nine genera are recognized in the Elachipterini: Allomedeia Mlynarek & Wheeler, Alombus Becker, Anatrichus Loew, Ceratobarys Coquillett, Disciphus Becker, Elachiptera Macquart, Goniaspis Duda, Melanochaeta Bezzi and Sepsidoscinis Hendel. Myrmecosepsis Kertesz is synonymised with Anatrichus, and Togeciphus Nishijima and Cyrtomomyia Becker are synonymised with Elachiptera. Ceratobarys is removed from synonymy with Elachiptera and all Neotropical species and two Nearctic species previously assigned to Elachiptera are transferred to Ceratobarys. Melanochaeta is a valid genus; the type species Melanochaeta capreolus clusters with other species of Melanochaeta and not Oscinella. New combinations include Anatrichus hystrix (Kertesz, 1914) (Myrmecosepsis); Anatrichus taprobane (Andersson, 1977) (Myrmecosepsis), Ceratobarys attenuata (Adams, 1908) (Elachiptera); Ceratobarys cultrata (Wheeler & Forrest, 2002) (Elachiptera); Ceratobarys flavida (Williston, 1896) (Elachiptera); Ceratobarys melinifrons (Mlynarek & Wheeler, 2008) (Elachiptera); Ceratobarys fucosa (Mlynarek & Wheeler, 2008) (Elachiptera); Ceratobarys queposana (Mlynarek & Wheeler, 2008) (Elachiptera); Ceratobarys rubida (Becker, 1912) (Elachiptera); Ceratobarys sacculicornis (Enderlein, 1911) (Elachiptera); Ceratobarys willistoni (Sabrosky, 1948) (Elachiptera), Elachiptera ensifer (Sabrosky, 1951) (Cyrtomomyia); Elachiptera ericius (Kanmiya, 1983) (Togeciphus); Elachiptera katoi (Nishijima, 1955) (Togeciphus); Elachiptera maculinervis (Becker, 1910) (Cyrtomomyia); Elachiptera punctulata (Becker, 1912) (Cyrtomomyia); Elachiptera subelongata (Kanmiya, 1983) (Disciphus); Elachiptera truncatus (Liu & Yang, 2012) (Togeciphus); Elachiptera tuberculata (Adams, 1905) (Cyrtomomyia) and all the species that were placed in Lasiochaeta are returned to Melanochaeta. A key to genera of the tribe Elachipterini is provided and diagnoses are provided for all genera. The tribe is divided into two geographically distinct clades: the Anatrichus clade includes the Old World tropical genera Allomedeia, Alombus, Anatrichus, Disciphus and Sepsidoscinis; the Elachiptera clade includes the primarily Neotropical genera Goniaspis and Ceratobarys and the widespread, but primarily Holarctic, genera Elachiptera and Melanochaeta.
References
Andersson, H. (1977) Taxonomic and phylogenetic studies on Chloropidae (Diptera) with special reference to Old World genera. Entomologica Scandinavica supplementum, 8, 5–200.
Andersson, H. (1979) Problem vid kladistik analys av flugfamiljen Chloropidae. Entomologisk Tidskrift, 100, 180–187.
Beaulieu, F. & Wheeler, T.A. (2002) Insects (Diptera, Coleoptera, Lepidoptera) reared from wetland monocots (Cyperaceae, Poaceae, Typhaceae) in southern Quebec. Proceedings of the Entomological Society of Washington, 104, 300–308.
Becker, T. (1911) Chloropidae. Eine monographische Studie. III teil. die Indo-Australische Region. Annales Musei Nationalis Hungarici, 9, 35–170.
Becker, T. (1913) Chloropiden aus Abessynien. Annales Musei Nationalis Hungarici, 11, 147–167.
Becker, T. (1914) Diptères nouveaux récoltés par mm. Ch. Alluaud et R. Jeannel en Afrique Orientale. Annals of the Entomological Society of France, 83, 120–130.
Beschovski, V.L. & Krusteva, C. (1998) Two new Elachiptera species from Bulgaria (Insecta: Diptera: Chloropidae). Reichenbachia, 32, 329–332.
Bezzi, M. (1895) Contribuzioni alla fauna ditterologica italiana. I. Ditteri della Calabria. Bullettino Della Società Entomologica Italiana, 27, 39–78.
Bezzi, M. (1906) Noch einige neue Namen für Dipterengattungen. Zeitschrift für Systematische Hymenopterologie und Dipterologie, 6 (1), 49–55.
Buck, M. (2006) A new family and genus of acalypterate flies from the Neotropical region, with a phylogenetic analysis of Carnoidea family relationships (Diptera, Schizophora). Systematic Entomology, 31, 377–404.
https://doi.org/10.1111/j.1365-3113.2006.00328.xCherian, P.T. (2012) Review of the taxonomic status of some and revised key to all the genera of the tribe Elachipterini (Diptera: Oscinellinae) and description of a new species of Myrmecosepsis Becker from India. Biosystematica, 5, 13–20.
Cherian, P.T. (2012) Revision of the genus Elachiptera Macquart (Diptera: Chloropidae: Oscinellinae: Elachipterini) from India with description of three new species. Hexapoda (Insecta indica), 19, 1–13
Cherian, P.T. (2015) Revision of the genus Gampsocera Schiner (Diptera: Chloropidae: Oscinellinae) from India with descriptions of seven new species. Entomon, 39, 15–42
Coquillett, D.W. (1898) Notes and descriptions of Oscinidae (Diptera). Journal of the New York Entomological Society, 6, 44–49.
Corti, E. (1909) Contributo alla conoscenza del gruppo della “Crassisete” in Italia (Ditteri). Bullettino Della Società Entomologica Italiana, 40, 121–162.
Deeming, J.C. & Al-Dhafer, H.M. (2012) Chloropidae from the Arabian Peninsula (Diptera: Cyclorrhapha). Zoology in the Middle East, 58, 1–88.
Duda, O. (1930) Die neotropischen Chloropiden (Diptera). Folia Zoologica et Hydrobiologica, 2, 46–128.
Duda, O. (1931) Die neotropischen Chloropiden (Dipt.). 1. Fortsetzung: Nachtrag, Ergänzungen, Berichtigungen und Index. Folia Zoologica et Hydrobiologica, 3, 159–172.
Duda, O. (1932) 61. Chloropidae [part] In: Lindner, E. (Ed.), Die Fliegen der palaearktischen Region. Lieferung 64. 6 (1). Schweizerbart’sche Verlagsbuchhandlung, Stuttgart, pp. 1–48.
Evenhuis, N.L., Pape, N.L., Pont, A.C. & Thompson, F.C. (2007) BioSystematic Database of World Diptera. Version 10. Available from: http://www.diptera.org/ (accessed 15 January 2018)
Ferrar, P. (1987) A guide to the breeding habits and immature stages of Diptera Cyclorrhapha. Part 1. Brill/Scandinavian Science Press, Leiden, 478 pp.
Goloboff, P.A., Farris, J.S. & Nixon, K.C. (2008) TNT a free program for phylogenetic analysis. Cladistics, 24, 774–786.
https://doi.org/10.1111/j.1096-0031.2008.00217.xHendel, F. (1907) Nomina nova für mehrere Gattungen der acalyptraten Musciden. Wiener Entomologische Zeitung, 26, 98.
https://doi.org/10.5962/bhl.part.5119Hendel, F. (1914) Eine neue Gattung der Oscinellinae. Annales Musei Nationalis Hungarici, 12, 247–248.
ICZN (2014) Opinion 2336 (case 3576): Oscinella Becker, 1909 (Insecta, Diptera, Chloropidae): precedence reversed with Melanochaeta Bezzi, 1906 and Pachychaetina Hendel, 1907. Bulletin of Zoological Nomenclature, 71, 141–143.
https://doi.org/10.21805/bzn.v71i2.a3Johannsen, O.A. (1924) A new chloropid subgenus and species from New York. The Canadian Entomologist, 56, 89.
https://doi.org/10.4039/Ent5689-4Kanmiya, K. (1983) A systematic study of the Japanese Chloropidae (Diptera). Memoirs of the Entomological Society of Washington, 11, 3–370.
Kertesz, K. (1914) Myrmecosepsis, a new genus of Diptera with very reduced wings. Annales Musei Nationalis Hungarici, 12, 244–246.
Liu, X. & Yang, D. (2012) Togeciphus Nishijima and Neoloxotaenia Sabrosky (Diptera: Chloropidae) from China. Zootaxa, 3298, 17–29.
Liu, X., Nartshuk, E.M. & Yang, D. (2017) First report of wingless grassflies from China (Diptera: Chloropidae) with a key to the Chinese and Oriental species of Elachiptera. Zoosystematica Rossica, 26, 329–336
Loew, H. (1845) Dipterologische Beiträge [I]. In: [Einladung] zu der öffentlichen Prüfung der Schüler des Königlichen Friedrich-Wilhelms-Gymnasiums zu Posen am 17. März 1845. Königliche Hofdruckerei W. Decker, Posen, pp. i + 1–52, 1 tafel.
Loew, H. (1860) Bidrag till kännedomen om Afrikas Diptera [part]. Öfversigt af Kongl. Vetenskaps-Akademiens Förhandlingar, 17, 81–97.
Macquart, P.J.M. [Pierre Justin Marie, as M. = Monsieur)] (1835) Histoire naturelle des insectes. Diptères. Tome deuxième. In: Nouvelles suites à Buffon, formant, avec les oeuvres de cet auteur, un cours complet d'histoire naturelle. Collection accom-pagnée de Planches. Librairie encyclopédique de Roret, Paris, pp. i–iii + 1–710, pls. 13–24, pp. 1–8 + 1–2.
Maddison, W.P., Donoghue, M.J. & Maddison, D.R. (1984) Outgroup analysis and parsimony. Systematic Zoology, 33, 83–103.
https://doi.org/10.2307/2413134Maddison, D.R. & Maddison, W.P. (2005) MacClade 4: Analysis of phylogeny and character evolution. Version 4.08. Sunderland, Massachusetts; Available from: http://macclade.org (accessed 15 January 2018)
McAlpine, J.F. (1981) Morphology and terminology—adults. In: McAlpine, J.F. (Ed.), Manual of Nearctic Diptera. Vol. 1. Research Branch Agriculture Canada, Ottawa, Ontario, pp. 9–63.
Mlynarek, J.J. & Wheeler, T.A. (2008) Revision of the Costa Rican species of Elachiptera (Diptera: Chloropidae). Zootaxa, 1754, 41–51.
Mlynarek, J.J. & Wheeler, T.A. (2009) Revision of the Neotropical genus Goniaspis Duda (Diptera: Chloropidae). Zootaxa, 2033, 26–40.
Mlynarek, J.J. & Wheeler, T.A. (2010a) Camptopeltes, a new Neotropical genus of oscinelline Chloropidae (Diptera). The Canadian Entomologist, 142, 193–199.
https://doi.org/10.4039/n09-075Mlynarek, J.J. & Wheeler, T.A (2010b) Systematics of Allomedeia, a new genus of Afrotropical Chloropidae (Diptera). Annals of the Entomological Society of America, 103, 465–471.
https://doi.org/10.1603/AN09141Nartshuk, E.P. (1983) A system of the superfamily Chloropoidea (Diptera: Cyclorrhapha). Entomologicheskoe Obozrenie, 62, 638–648. [in Russian, English translation in Entomological Review, 62, 180–193]
Nartshuk, E.P. (2012) A check-list of the world genera of the family Chloropidae (Diptera, Cyclorrhapha, Muscomorpha). Zootaxa, 3267, 1–43.
Nartshuk, E.P. & von Tschirnhaus, M.V. (2012) New generic synonyms in the Chloropidae (Diptera, Acalyptratae), with additional taxonomic notes. Zootaxa, 3267, 44–54.
Nishijima, Y. (1954) Descriptions of a new genus and a new species of Chloropidae from Japan (Diptera). Insecta Matsumurana, 18, 84–85.
Paramonov, S.J. (1961) Notes on Australasian Diptera XXXII. Echiniidae—A new family of Diptera (Acalyptrata). Annals and Magazine of Natural History, Series 13, 4, 97–100.
https://doi.org/10.1080/00222936108655785Rondani, C. (1856) Dipterologiae Italicae prodromus. Vol. I. Genera Italica ordinis Dipterorum ordinatum disposita et dis-tincta et in familias et stirpes aggregata. A. Stoschi, Parmae [= Parma], 226 pp.
https://doi.org/10.5962/bhl.title.8160Sabrosky, C.W. (1941) An annotated list of genotypes of the Chloropidae of the world (Diptera). Annals of the Entomological Society of America, 34, 735–765.
https://doi.org/10.1093/aesa/34.4.735Sabrosky, C.W. (1951) Chloropidae. Ruwenzori Expedition 1934–35, 2, 711–828.
Sabrosky, C.W. (1980) Family Chloropidae. In: Crosskey, R.W. (Ed.), Catalogue of the Diptera of the Afrotropical Region. British Museum (Natural History), London, pp. 695–712.
Sabrosky, C.W. (1984) Check list of the Chloropidae of the world (Diptera). Unpublished manuscript, 21 pp. [draft]
Sabrosky, C.W. (1987) Chloropidae. In: McAlpine, J.F., Peterson, B.V., Shewell, G.E., Teskey, H.J., Vockeroth, J.R., & Wood, D.M. (Eds.), Manual of Nearctic Diptera. Vol. 2. Research Branch Agriculture Canada, Ottawa, pp. 1049–1067.
Sabrosky, C. & Valley, K. (1987) A new Elachiptera from salt marshes, with redescription of E. penita and partially revised key to nearctic Elachiptera (Diptera: Chloropidae). Proceedings of the Entomological Society of Washington, 89, 581–586.
Séguy, E. (1938) Mission scientifique de l'Omo. 4 (Zoologie). Diptera I. Nematocera et Brachycera. Mémoires du Muséum national d’Histoire naturelle, Nouvelle Série, 8, 319–380.
von Roser, K.L.F. (1840) III. Beiträge zur Vaterlandskunde. Erster Nachtrag zu dem im Jahre 1834 bekannt gemachen Verzeichnisse in Württemberg vorkommender zweiflügliger Insekten. Correspondenzblatt des Königlich Württemberischen Landwirthschaftlichen Vereins, 37, 49–64.
von Tschirnhaus, M. (2017) The taxonomy of species globally described in or formerly included in the genus Elachiptera and new combinations with Lasiochaeta and Gampsocera (Diptera: Chloropidae). Zoosystematica Rossica, 26, 337–368.
von Tschirnhaus, M. & Nartshuk, E.P. (2012) Case 3576. Oscinella Becker, 1909 (Insecta, Diptera, Chloropidae): proposed conservation by reversal of precendence with Melanochaeta Bezzi, 1906 and Pachychaetina Hendel, 1907. Bulletin of Zoological Nomenclature, 69 (1), 37–43.
https://doi.org/10.21805/bzn.v69i1.a13Wheeler, T.A. & Forrest, J. (2002) A new species of Elachiptera Macquart from the Galápagos Islands, Ecuador, and the taxonomic status of Ceratobarys (Diptera: Chloropidae). Zootaxa, 98, 1–9.
https://doi.org/10.11646/zootaxa.98.1.1Wiens, J.J. (1998) Does adding characters with missing data increase or decrease phylogenetic accuracy? Systematic Biology, 44, 501–514.
https://doi.org/10.1080/106351598260635Yeates, D.K. (1995) Groundplans and exemplars: paths to the tree of life. Cladistics, 11, 343–35.
https://doi.org/10.1111/j.1096-0031.1995.tb00094.xYang, C. & Yang, D. (1990) New species of Melanochaeta from China (Diptera: Chloropidae). Acta Agriculturae Universitatis Pekinensis, 16, 201–204.
Yang, D. & Yang, C.-K. (1990) A new species of Melanochaeta Bezzi (Diptera: Chloropidae) from Nei Mongol). Entomotaxonomia, 12, 308–310.
Yang, C.-K. & Yang, D. (1991) A new genus and four new species of Chloropidae from China (Diptera). Acta zootaxonomica sinica, 16, 471–475.
Yang, C.-K. & Yang, D. (1991) Eight new species of Melanochaeta from Yunnan, China (Diptera: Chloropidae). Acta Zootaxonomica Sinica, 16, 476–483.
Yang, D. & Yang, C. [2004 (2003)] Diptera: Chloropidae. In: Huang, B.K. (Ed.), Fu jian kun chong zhi di ba juan [Fauna of Insects in Fujian Province of China]. Vol. 8. Diptera—Mallophaga. Chinese Academy of Science, Fuzhou, pp. 527–534.