Schizophora

 Syringogaster sp., from here.

Belongs within: Cyclorrhapha.
Synapomorphies: Odiniidae, Periscelididae, Carnidae, Agromyzidae, Sphaeroceridae, Ephydroidea, Calyptratae, Tephritoidea, Chyromyidae, Opomyzidae, Sepsidae, Chloropidae, Milichiidae, Diopsidae, Canacidae, Nerioidea, Aulacigastridae, Tanypezoidea, Heleomyzidae, Sciomyzoidea, Lauxanioidea, Ropalomeridae, Asteiidae, Psilidae, Clusiidae, Pallopteridae.

The Schizophora is a major clade of flies distinguished by the presence of the ptilinum, an eversible membrane in the head that is filled with fluid to break the adult fly out of the pupal case. For the remainder of the fly’s life, the ptilinum is held within the head but its presence can still be traced by a ptilinal fissure crossing between the eyes above the antennal insertions. Schizophorans have often been divided for convenience between the Calyptratae and Acalyptratae, with the lower calypter of the wing typically being more developed in the former, though ‘Acalyptratae’ have long been recognised as an artificial group. The relationships of many ‘acalyptrate’ families have only recently begun to be resolved and many families are difficult to distinguish without close examination. The fossil record suggests an explosive radiation of the Schizophora in the early Tertiary (Wiegmann et al. 2011).

Examples of smaller families not covered on separate pages are the Megamerinidae, a group of slender, elongate flies known from Eurasia. Acartophthalmus is a European genus of small grey or black flies with a pubescent arista. The Neminidae, nobody flies, are minute flies found in the Old World tropics, with a proclinate fronto-orbital bristle present and parallel or divergent postocellar bristles. Inbiomyia is a recently described genus of Neotropical flies characterised by a shortened head with a remarkably long arista on the antenna. Somatia, another Neotropical endemic, includes mostly black and yellow flies with an anteromedially produced frons, neck-like pronotum and broadened abdomen. Margo is an African genus of small, shining flies with porrect antennae and relatively long legs. The Nannodastiidae are very small, dull-coloured flies that are mostly found along beaches in warmer regions of the world. The Teratomyzidae are narrow-winged flies of the Oriental, Australian and Neotropical regions that are mostly found on ferns in cool forests (Colless & McAlpine 1991). Heteromyza is a Palaearctic genus of somewhat long-legged flies historically placed within the Heleomyzidae. The Xenasteiidae are minute flies with reduced wing venation found on coastal vegetation along tropical and subtropical shores of the Old World (Oosterbroek 1998). The mostly Australian genus Fergusonina has larvae that live in galls on Myrtaceae in association with the nematode genus Fergusobia (Colless & McAlpine 1991). Syringogaster includes small, ant-mimicking flies found in lowland tropical forests of the Americas (Marshall & Buck 2010). The Neurochaetidae are elongate, cursorial flies found in warm regions of the Old World that maintain a head-downwards orientation on vertical surfaces (Oosterbroek 1998).

Characters (from Grimaldi & Engel 2005): Ptilinum present, adult emerging from puparium via circular sclerite; adult with specialised structure (cardia) between fore- and mid gut.

<==Schizophora (see below for synonymy)
    |--+--+--OdiniidaeWT11
    |  |  `--+--+--PeriscelididaeWT11
    |  |     |  `--CarnidaeWT11
    |  |     `--+--AgromyzidaeWT11
    |  |        `--SphaeroceridaeWT11
    |  `--+--EphydroideaWT11
    |     `--CalyptrataeWT11
    `--+--MegamerinidaeWT11
       |    |--TexaraO98
       |    |--Megamerina doliumO98
       |    `--Palaeotanypeza spinosaRJ93
       `--+--+--TephritoideaWT11
          |  `--+--ChyromyidaeWT11
          |     `--+--OpomyzidaeWT11
          |        `--+--SepsidaeWT11
          |           `--AcartophthalmidaeWT11
          |                |--Acartophthalmites tertiariaRJ93
          |                `--AcartophthalmusM87
          |                     |--A. bicolor Oldenberg 1910C-T92
          |                     `--A. nigrinus (Zetterstedt 1848)C-T92
          `--+--+--+--+--+--ChloropidaeWT11
             |  |  |  |  `--MilichiidaeWT11
             |  |  |  `--SomatiaLM10 [SomatiidaeWT11]
             |  |  |       |--S. aestivaLM10
             |  |  |       |--S. schildiLM10
             |  |  |       `--S. sophistonLM10
             |  |  `--+--DiopsidaeWT11
             |  |     `--+--Margo [Marginidae]WT11
             |  |        `--+--CanacidaeWT11
             |  |           `--Nannodastiidae [Nannodastiinae]WT11
             |  |                |--Azorastia mediterraneaM10
             |  |                `--NannodastiaM10
             |  |                     |--N. atlanticaM10
             |  |                     `--N. horniM10
             |  `--+--+--NerioideaWT11
             |     |  `--AulacigastridaeWT11
             |     `--+--InbiomyiaB10 [InbiomyiidaeWT11]
             |        |    |--I. anemosyrisB10
             |        |    |--I. exulB10
             |        |    |--I. mcalpineorumB10
             |        |    |--I. scoliostylusB10
             |        |    `--I. zeugodontaB10
             |        `--NeminidaeWT11
             |             |--NingulusRM10
             |             |--Nemula Freidberg 1994RM10
             |             |    `--N. longaristaWT11
             |             `--NemoRM10
             |                  |--N. arbelosO98
             |                  |--N. centrisetaCM91
             |                  `--N. corticeusO98
             `--+--TanypezoideaWT11
                `--+--+--+--HeleomyzidaeWT11
                   |  |  `--TeratomyzidaeWT11
                   |  |       |--TeratomyzaWT11
                   |  |       |--Camur williiBW09
                   |  |       `--TeratopteraCM91
                   |  `--+--+--SciomyzoideaWT11
                   |     |  `--LauxanioideaWT11
                   |     `--HeteromyzaWT11 [incl. ThelidaGP87; Heteromyzidae]
                   |          |--H. atricornisWT11
                   |          |--‘Heteromyiella’ miocenica [=Thelida miocenica]GP87
                   |          `--H. oculataGP87
                   `--+--+--RopalomeridaeWT11
                      |  `--+--+--Australimyza [Australimyzidae, Australimyzinae]WT11
                      |     |  `--Xenasteiidae [Tunisimyiidae]WT11
                      |     |       |--Xenasteia shalamWT11
                      |     |       `--TunisimyiaO98
                      |     `--+--AsteiidaeWT11
                      |        `--Fergusonina [Fergusoninidae]WT11
                      |             |--F. carteriS00
                      |             |--F. nicholsoniN70
                      |             |--F. syzygiiS00
                      |             `--F. turneriWT11
                      `--+--+--PsilidaeWT11
                         |  `--SyringogasterMB10 [SyringogastridaeWT11]
                         |       |--S. brunneaMB10
                         |       |--S. brunneinaMB10
                         |       |--S. figurataMB10
                         |       |--S. rufaMB10
                         |       `--S. subnearcticaMB10
                         `--+--ClusiidaeWT11
                            `--+--PallopteridaeWT11
                               `--NeurochaetidaeWT11
                                    |--NothoasteiaCM91
                                    |--AnthoclusiaRJ93
                                    |    |--A. gephyreaRJ93
                                    |    `--A. remotinerviaP92
                                    `--NeurochaetaO98
                                         |--N. inversaCM91
                                         `--N. parvicepsO98
Schizophora incertae sedis:
  Nothybus [Nothybidae]O98
    `--N. biguttatusO98
  ParaleucopidaeW10
    |--MallochianamyiaW10
    |--SchizostomyiaW10
    `--ParaleucopisW10
         |--P. corvinaW10
         `--P. mexicanaW10
  GobryaCM91 [GobryidaeO98]
  Bromophila caffra Macq. 1846 [=Dichromyia caffra; incl. Scatophaga zamid Walker 1849]R01

Schizophora [Acalypterae, Acalyptrata, Agromyzoidea, Agromyzoinea, Anthomyzoidea, Asteioidea, Asteioinea, Carnoidea, Chloropoidea, Clusioinea, Conopsariae, Diopsoidea, Heleomyzoidea, Holometopa, Lonchaeoidea, Muscaria, Nothyboidea, Opomyzoidea, Opomyzoidea, Pallopteroidea, Sphaeroceroidea]

*Type species of generic name indicated

References

[B10] Buck, M. 2010. Inbiomyiidae (inbiomyiid flies). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 2 pp. 1105–1108. NRC Research Press: Ottawa.

[BW09] Buck, M., N. E. Woodley, A. Borkent, D. M. Wood, T. Pape, J. R. Vockeroth, V. Michelsen & S. A. Marshall. 2009. Key to Diptera families—adults. In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 1 pp. 95–156. NRC Research Press: Ottawa.

[C-T92] Carles-Tolrá, M. 1992. New and interesting records of Diptera Acalyptrata from Spain. Part I: Acartophthalmidae, Opomyzidae, Anthomyzidae, Asteiidae, Carnidae, Tethinidae, Milichiidae and Cryptochetidae. Bull. Annls Soc. R. Belge Ent. 128: 343–353.

[CM91] Colless, D. H., & D. K. McAlpine. 1991. Diptera (flies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2nd ed. vol. 2 pp. 717–786. Melbourne University Press: Carlton (Victoria).

[GP87] Gill, G. D., & B. V. Peterson. 1987. Heleomyzidae. In: McAlpine, J. F. (ed.) Manual of Nearctic Diptera vol. 2 pp. 973–980. Research Branch, Agriculture Canada.

[LM10] Lonsdale, O., & D. K. McAlpine. 2010. Somatiidae (somatiid flies). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 2 pp. 833–835. NRC Research Press: Ottawa.

[MB10] Marshall, S. A., & M. Buck. 2010. Syringogastridae (syringogastrid flies). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 2 pp. 843–846. NRC Research Press: Ottawa.

[M10] Mathis, W. N. 2010. Nannodastiidae (nannodastiid beach flies). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 2 pp. 1235–1239. NRC Research Press: Ottawa.

[M87] McAlpine, J. F. 1987. Acartophthalmidae. In: McAlpine, J. F. (ed.) Manual of Nearctic Diptera vol. 2 pp. 859–861. Research Branch, Agriculture Canada.

[N70] Norris, K. R. 1970. General biology. In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 107–140. Melbourne University Press.

[O98] Oosterbroek, P. 1998. The Families of Diptera of the Malay Archipelago. Brill: Leiden.

[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.

[R01] Ricardo, G. 1901. Notes on Diptera from South Africa (concluded). Annals and Magazine of Natural History, series 7, 7: 89–110.

[RJ93] Ross, A. J., & E. A. Jarzembowski. 1993. Arthropoda (Hexapoda; Insecta). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 363–426. Chapman & Hall: London.

[RM10] Rung, A., & W. N. Mathis. 2010. Aulacigastridae (aulacigastrid flies). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 2 pp. 1083–1085. NRC Research Press: Ottawa.

[S00] Siddiqi, M. R. 2000. Tylenchida: Parasites of plants and insects 2nd ed. CABI Publishing: Wallingford (UK).

[W10] Wheeler, T. A. 2010. Paraleucopidae (paraleucopid flies). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 2 pp. 1009–1011. NRC Research Press: Ottawa.

[WT11] Wiegmann, B. M., M. D. Trautwein, I. S. Winkler, N. B. Barr, J.-W. Kim, C. Lambkin, M. A. Bertone, B. K. Cassel, K. M. Bayless, A. M. Heimberg, B. M. Wheeler, K. J. Peterson, T. Pape, B. J. Sinclair, J. H. Skevington, V. Blagoderov, J. Caravas, S. N. Kutty, U. Schmidt-Ott, G. E. Kampmeier, F. C. Thompson, D. A. Grimaldi, A. T. Beckenbach, G. W. Courtney, M. Friedrich, R. Meier & D. K. Yeates. 2011. Episodic radiations in the fly tree of life. Proceedings of the National Academy of Sciences of the USA 108 (14): 5690–5695.

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