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 where Bittacomopha and Bittacomorphella (together forming the Bittacomorphinae) have an eighteen- or nineteen-segmented flagellum and unbranched M vein (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
    |    |--+--Boholdoya RJ93 [Boholdoyidae BLM02]
    |    |  |    |--B. alata Kovalev in Kalugina & Kovalev 1985 RJ93
    |    |  |    `--B. thoracica Kovalev 1990 RJ93
    |    |  `--Perissommatidae WT11
    |    |       |--Palaeoperissomma collessi Kovalev in Kalugina & Kovalev 1985 RJ93
    |    |       `--Perissomma WT11
    |    |            |--P. bellissima CM91
    |    |            |--P. fusca CM91
    |    |            `--P. mcalpinei WT11
    |    `--+--Bibionomorpha WT11
    |       `--Brachycera (see below for synonymy) WT11
    |            |  i. s.: Eophlebomyiidae BLM02
    |            |         Lithohypoderma S02
    |            |         Notodacus xanthodes B88
    |            |         Gonypes tipuloides G20
    |            |         Sicus lineatus G20
    |            |--Orthorrhapha WT11
    |            `--Eremoneura WT11
    |                 |--Empidoidea WT11
    |                 `--+--Cyclorrhapha WT11
    |                    `--Apystomyia [Apystomyiidae] WT11
    |                         `--A. elinguis WT11
    `--+--+--Psychodomorpha WT11
       |  `--Culicomorpha WT11
       `--+--+--Rhaetania Krzemiński & Krzemińska 2002 KK02 [Rhaetaniidae GE05]
          |  |    `--*R. dianae Krzemiński & Krzemińska 2002 KK02
          |  `--Tillyardiptera KK02 [Tillyardipteridae GE05]
          |       `--T. prima KK02
          `--+--Eoptychopteridae [Architendipedidae] GE05
             |    |--Bolboia mira S02
             |    |--Crenoptychoptera S02
             |    |--Leptychoptera GE05
             |    `--Eoptychopterina baisica Kalugina 1989 RJ93
             `--Ptychopteridae [Eolimbiidae, Eolimnobiidae, Eolimuobiidae, Ptychopteroidea] WT11
                  |  i. s.: Zhiganka S02
                  |         Eolimnobia geinitzi A81
                  |         Chlorolimnobia ostera Lin 1986 RJ93
                  |--Bittacomorphinae H09
                  |    |--Bittacomorpha clavipes A81
                  |    `--Bittacomorphella A81
                  |         |--B. jonesi A81
                  |         `--B. miocenica A81
                  `--Ptychopterinae H09
                       |--Ptychopterula Handlirsch 1909 H02
                       |--Sinoptychopterites Hong 2002 H02
                       |    `--*S. paludus Hong 2002 H02
                       `--Ptychoptera Meigen 1803 H02
                            |--P. albimana [=Tipula (Ptychoptera) albimana] G20
                            |--P. alexanderi H09
                            |--P. bellula Alexander 1937 A40
                            |--P. deleta A81
                            |--P. javensis O98
                            |--P. lenis A81
                            |--P. mesozoica Kalugina 1989 RJ93
                            |--P. quadrifasciata A81
                            `--P. sumatrensis O98

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

*Type species of generic name indicated


[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.

[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.

[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.

Last updated: 4 February 2021.

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