Neoneura

Phantom crane fly Ptychoptera quadrifasciata, copyright John F. Carr.

Belongs within: Diptera.
Contains: Bibionomorpha, Orthorrhapha, Empidoidea, Cyclorrhapha, Psychodomorpha, Culicomorpha.

The Neoneura are a clade containing the majority of flies except for the basal tipulomorphs. They are characterised by having vein 1A distally reduced and not reaching the margin of the wing, and the arolium replaced by an empodium (Blagoderov et al. 2002). Basal lineages in the Neoneura include the Ptychopteridae, phantom crane flies, a group of long-legged flies with aquatic larvae found mostly in cooler regions. Ptychopterids are distinguished from other flies by the presence of a basal appendage on the halter, the prehalter. Three genera of ptychopterids are recognised worldwide: members of the genus Ptychoptera have thirteen segments in the antennal flagellum and a branched M vein in the wing whereas Bittacomopha and Bittacomorphella (together forming the Bittacomorphinae) have an eighteen- or nineteen-segmented flagellum and unbranched M vein. Bittacomorphella possess macrotrichia in the distal radial and medial cells of the wing, whereas Bittacomorpha lack such macrotrichia and have the first tarsomeres of all legs dilated (Alexander 1981).

A molecular analysis of fly phlogeny by Wiegmann et al. (2011) separated a basal division between clades containing the Ptychopteridae, Psychodomorpha and Culicomorpha on one side and Perissomatidae, Bibionomorpha and Brachycera on the other. The latter clade is known as the Neodiptera and is also united by features of the neck region sclerites and musculature. The Perissomatidae are a family of small flies found in Australia and South America that have the eyes completely divided into separate dorsal and ventral components. The Brachycera are a well-supported clade of flies distinguished by, among other features, their compacted antennae with distal flagellomeres modified into a slender style or hairlike arista. Within the Brachycera, Wiegmann et al. (2011) supported two basal clades, the Orthorrhapha and Eremoneura. The Eremoneura include the Cyclorrhapha and the long-legged empidoids; eremoneurans are united by the possession of three larval instars, larval mandibles with a single article, and adults with ocellar setae, one-segmented maxillary palps, and the cell cup in the wing closed apically or absent (Grimaldi & Engel 2005). They also include the relict Apystomyia elinguis, a small, superficially bombyliid-like fly found in California.

<==Neoneura [Ptychopteromorpha]GE05
    |--Neodiptera [Axymyiiformia, Perissommatoidea]WT11
    |    |--+--BoholdoyaRJ93 [BoholdoyidaeBLM02]
    |    |  |    |--B. alata Kovalev in Kalugina & Kovalev 1985RJ93
    |    |  |    `--B. thoracica Kovalev 1990RJ93
    |    |  `--PerissommatidaeWT11
    |    |       |--Palaeoperissomma collessi Kovalev in Kalugina & Kovalev 1985RJ93
    |    |       `--PerissommaWT11
    |    |            |--P. bellissimaCM91
    |    |            |--P. fuscaCM91
    |    |            `--P. mcalpineiWT11
    |    `--+--BibionomorphaWT11
    |       `--Brachycera (see below for synonymy)WT11
    |            |  i. s.: EophlebomyiidaeBLM02
    |            |         LithohypodermaS02
    |            |         Notodacus xanthodesB88
    |            |         Gonypes tipuloidesG20
    |            |--OrthorrhaphaWT11
    |            `--EremoneuraWT11
    |                 |--EmpidoideaWT11
    |                 `--+--CyclorrhaphaWT11
    |                    `--Apystomyia Melander 1950WT11, H72 [Apystomyiidae]
    |                         `--*A. elinguisH72
    `--+--+--PsychodomorphaWT11
       |  `--CulicomorphaWT11
       `--+--+--Rhaetania Krzemiński & Krzemińska 2002KK02 [RhaetaniidaeGE05]
          |  |    `--*R. dianae Krzemiński & Krzemińska 2002KK02
          |  `--TillyardipteraKK02 [TillyardipteridaeGE05]
          |       `--T. primaKK02
          `--+--Eoptychopteridae [Architendipedidae]GE05
             |    |--Bolboia miraS02
             |    |--CrenoptychopteraS02
             |    |--LeptychopteraGE05
             |    `--Eoptychopterina baisica Kalugina 1989RJ93
             `--Ptychopteridae [Eolimbiidae, Eolimnobiidae, Eolimuobiidae, Ptychopteroidea]WT11
                  |  i. s.: ZhigankaS02
                  |         Eolimnobia geinitziA81
                  |         Chlorolimnobia ostera Lin 1986RJ93
                  |--BittacomorphinaeH09
                  |    |--Bittacomorpha clavipesA81
                  |    `--BittacomorphellaA81
                  |         |--B. jonesiA81
                  |         `--B. miocenicaA81
                  `--PtychopterinaeH09
                       |--Ptychopterula Handlirsch 1909H02
                       |--Sinoptychopterites Hong 2002H02
                       |    `--*S. paludus Hong 2002H02
                       `--Ptychoptera Meigen 1803H02
                            |--P. albimana [=Tipula (Ptychoptera) albimana]G20
                            |--P. alexanderiH09
                            |--P. bellula Alexander 1937A40
                            |--P. contaminataR26
                            |--P. deletaA81
                            |--P. javensisO98
                            |--P. lenisA81
                            |--P. mesozoica Kalugina 1989RJ93
                            |--P. quadrifasciataA81
                            `--P. sumatrensisO98

Brachycera [Asilici, Asilomorpha, Dolichopodes, Heterodactyla, Heterotropinae, Nemestrinoidea, Raptatores, Siphunculini, Tabanii]WT11

*Type species of generic name indicated

References

[A40] Alexander, C. P. 1940. New or little-known Tipulidae from eastern China. Notes d’Entomologie Chinoise 8 (1): 1–28.

[A81] Alexander, C. P. 1981. Ptychopteridae. In: McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth & D. S. Wood (eds) Manual of Nearctic Diptera vol. 1 pp. 325–328. Research Branch, Agriculture Canada.

[BLM02] Blagoderov, V. A., E. D. Lukashevich & M. B. Mostovski. 2002. Order Diptera Linné, 1758. The true flies (=Muscida Laicharting, 1781). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 227–240. Kluwer Academic Publishers: Dordrecht.

[B88] Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

[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).

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[H72] Hall, J. C. 1972. New North American Heterotropinae (Diptera: Bombyliidae). Pan-Pacific Entomologist 48 (1): 37–50.

[H09] Hancock, E. G. 2009. Ptychopteridae (phantom crane 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. 1 pp. 357–358. NRC Research Press: Ottawa.

[H02] Hong Y. 2002. Amber Insect of China. Beijing Scientific and Technological Publishing House.

[KK02] Krzemiński, W., & E. Krzemińska. 2002. Rhaetaniidae, a new family of the Diptera from the Upper Triassic of Great Britain (Diptera: Nematocera). Annales Zoologici 52 (2): 211–213.

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

[R26] Risso, A. 1826. Histoire naturelle des principales productions de l’Europe méridionale et particulièrement de celles des environs de Nice et des Alpes maritimes vol. 5. F.-G. Levrault: Paris.

[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.

[S02] Sinitshenkova, N. D. 2002. Ecological history of the aquatic insects. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 388–426. Kluwer Academic Publishers: Dordrecht.

[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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