Cyclorrhapha

Lonchoptera lutea, copyright Saxifraga.

Belongs within: Neoneura.
Contains: Syrphoidea, Schizophora, Platypezoidea.

The Cyclorrhapha are a clade of flies in which the larvae have developed the familiar 'maggot' form, with the head desclerotised and invaginated into the body. Representatives include Lonchoptera, a cosmopolitan genus of small, slender flies with apically pointed wings. The biology of Lonchoptera is largely unknown though adults may sometimes be found in large numbers in humid habitats (Petersen 1987).

The earliest cyclorrhaphans are known from the Cretaceous (Grimaldi & Engel 2005). Possible examples include Chimeromyia, a genus of slender flies known from Lebanese amber. The name of this genus refers to its members combining features of the Cyclorrhapha with those of its living sister group, the Empidoidea. Lonchopterites prisca, also from Lebanese amber, resembles modern Lonchoptera but has a terminal rather than subapical antennal arista and two pairs of fronto-orbital setae (Grimaldi & Cumming 1996).

Synapomorphies (from Grimaldi & Engel 2005): Face with pair or more of thick vibrissae; abdominal plaques lost in adults; embryos not developing in amnion; larvae saprophagous 'maggots' with maxilla absent, head desclerotised and invaginated; pupariation in ecdysed cuticle of last larval instar; arista with two small basal segments; proboscis retractile, withdrawn at rest into large oral cavity; female terminalia without acanthophorite spines.

<==Cyclorrhapha [Aschiza, Muscides, Muscomorpha, Myionomorpha]
    |  i. s.: Neria cibaria WJ18
    |--Chimeromyia Grimaldi & Cumming 1999 GE05, GC99
    |    |--*C. intriguea Grimaldi & Cumming 1999 GC99
    |    |--C. acuta Grimaldi & Cumming 1999 GC99
    |    `--C. reducta Grimaldi & Cumming 1999 GC99
    `--+--Lonchopterites Grimaldi & Cumming 1999 GE05, GC99
       |    `--*L. prisca Grimaldi & Cumming 1999 GC99
       `--+--+--Syrphoidea RS10
          |  `--Schizophora RS10
          `--+--Platypezoidea WT11
             `--Lonchopteridae [Acroptera, Anatriata, Lonchopteroidea, Lonchopteromorpha, Musidoridae] WT11
                  |--Lonchopteromorpha Grimaldi & Cumming 1999 GC99
                  |    `--*L. asetocella Grimaldi & Cumming 1999 GC99
                  `--Lonchoptera Meigen 1803 [incl. Musidora Meigen 1800 (nom. inv.)] P87
                       |--L. bifurcata BW09 [incl. L. dubia CM91, L. furcata O98]
                       |--L. borealis P87
                       |--L. fallax P87
                       |--L. lutea P87
                       |--L. occidentalis P87
                       |--L. tristis P87
                       `--L. uniseta P87

*Type species of generic name indicated

REFERENCES

[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.

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

[GC99] Grimaldi, D., & J. Cumming. 1999. Brachyceran Diptera in Cretaceous ambers and Mesozoic diversification of the Eremoneura. Bulletin of the American Museum of Natural History 239: 1–124.

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

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

[P87] Peterson, B. V. 1987. Lonchopteridae. In: McAlpine, J. F. (ed.) Manual of Nearctic Diptera vol. 2 pp. 675–680. Research Branch, Agriculture Canada.

[RS10] Rafael, J. A., & J. H. Skevington. 2010. Pipunculidae (big-headed 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. 793–803. NRC Research Press: Ottawa.

[WJ18] Wahlberg, E., & K. A. Johanson. 2018. Molecular phylogenetics reveals novel relationships within Empidoidea (Diptera). Systematic Entomology 43: 619–636.

[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: 18 July 2021.

Schizophora

Syringogaster sp., from here.

Belongs within: Cyclorrhapha.
Synapomorphies: Ephydroidea, Calyptratae, Odiniidae, Periscelididae, Carnidae, Agromyzidae, Sphaeroceridae, 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 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.

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)
    |--+--+--Ephydroidea WT11
    |  |  `--Calyptratae WT11
    |  `--+--Odiniidae WT11
    |     `--+--+--Periscelididae WT11
    |        |  `--Carnidae WT11
    |        `--+--Agromyzidae WT11
    |           `--Sphaeroceridae WT11
    `--+--Megamerinidae WT11
       |    |--Texara O98
       |    |--Megamerina dolium O98
       |    `--Palaeotanypeza spinosa RJ93
       `--+--+--Tephritoidea WT11
          |  `--+--Chyromyidae WT11
          |     `--+--Opomyzidae WT11
          |        `--+--Sepsidae WT11
          |           `--Acartophthalmidae WT11
          |                |--Acartophthalmites tertiaria RJ93
          |                `--Acartophthalmus M87
          |                     |--A. bicolor Oldenberg 1910 C-T92
          |                     `--A. nigrinus (Zetterstedt 1848) C-T92
          `--+--+--+--+--+--Chloropidae WT11
             |  |  |  |  `--Milichiidae WT11
             |  |  |  `--Somatia LM10 [Somatiidae WT11]
             |  |  |       |--S. aestiva LM10
             |  |  |       |--S. schildi LM10
             |  |  |       `--S. sophiston LM10
             |  |  `--+--Diopsidae WT11
             |  |     `--+--Margo [Marginidae] WT11
             |  |        `--+--Canacidae WT11
             |  |           `--Nannodastiidae [Nannodastiinae] WT11
             |  |                |--Azorastia mediterranea M10
             |  |                `--Nannodastia M10
             |  |                     |--N. atlantica M10
             |  |                     `--N. horni M10
             |  `--+--+--Nerioidea WT11
             |     |  `--Aulacigastridae WT11
             |     `--+--Inbiomyia B10 [Inbiomyiidae WT11]
             |        |    |--I. anemosyris B10
             |        |    |--I. exul B10
             |        |    |--I. mcalpineorum B10
             |        |    |--I. scoliostylus B10
             |        |    `--I. zeugodonta B10
             |        `--Neminidae WT11
             |             |--Ningulus RM10
             |             |--Nemula Freidberg 1994 RM10
             |             |    `--N. longarista WT11
             |             `--Nemo RM10
             |                  |--N. arbelos O98
             |                  |--N. centriseta CM91
             |                  `--N. corticeus O98
             `--+--Tanypezoidea WT11
                `--+--+--+--Heleomyzidae WT11
                   |  |  `--Teratomyzidae WT11
                   |  |       |--Teratomyza WT11
                   |  |       |--Camur willii BW09
                   |  |       `--Teratoptera CM91
                   |  `--+--+--Sciomyzoidea WT11
                   |     |  `--Lauxanioidea WT11
                   |     `--Heteromyza WT11 [incl. Thelida GP87; Heteromyzidae]
                   |          |--H. atricornis WT11
                   |          |--‘Heteromyiella’ miocenica [=Thelida miocenica] GP87
                   |          `--H. oculata GP87
                   `--+--+--Ropalomeridae WT11
                      |  `--+--+--Australimyza [Australimyzidae, Australimyzinae] WT11
                      |     |  `--Xenasteiidae [Tunisimyiidae] WT11
                      |     |       |--Xenasteia shalam WT11
                      |     |       `--Tunisimyia O98
                      |     `--+--Asteiidae WT11
                      |        `--Fergusonina [Fergusoninidae] WT11
                      |             |--F. carteri S00
                      |             |--F. nicholsoni N70
                      |             |--F. syzygii S00
                      |             `--F. turneri WT11
                      `--+--+--Psilidae WT11
                         |  `--Syringogaster MB10 [Syringogastridae WT11]
                         |       |--S. brunnea MB10
                         |       |--S. brunneina MB10
                         |       |--S. figurata MB10
                         |       |--S. rufa MB10
                         |       `--S. subnearctica MB10
                         `--+--Clusiidae WT11
                            `--+--Pallopteridae WT11
                               `--Neurochaetidae WT11
                                    |--Nothoasteia CM91
                                    |--Anthoclusia RJ93
                                    |    |--A. gephyrea RJ93
                                    |    `--A. remotinervia P92
                                    `--Neurochaeta O98
                                         |--N. inversa CM91
                                         `--N. parviceps O98

Schizophora incertae sedis:
  Nothybus [Nothybidae] O98
    `--N. biguttatus O98
  Paraleucopidae W10
    |--Mallochianamyia W10
    |--Schizostomyia W10
    `--Paraleucopis W10
         |--P. corvina W10
         `--P. mexicana W10
  Gobrya CM91 [Gobryidae O98]
  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 2^nd 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 2^nd 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.

Last updated: 12 July 2021.

Lauxaniidae

Sapromyza sp., copyright frogprince.

Belongs within: Lauxanioidea.
Contains: Trigonometopini.

The Lauxaniidae are a family of flies whose larvae mostly live in fallen leaves and other rotting vegetation. Several genera include nidicolous species, feeding on decaying matter in bird nests. Adults graze on fungi growing on fallen leaves (Gaimari & Silva 2010).

The internal classification of the family is somewhat poorly established. The Eurychoromyiinae have previously been recognised as a separate family due to a reduction in setation but examination at high magnification has found that a number of 'lost' setae are present but greatly reduced. The Homoneurinae are characterised by wings with costal setulae reaching the apex of vein R₄₊₅ whereas in Lauxaniinae they end shortly past vein R₂₊₃; however, it seems likely that the 'Lauxaniinae' are paraphyletic to the Homoneurinae. Similarly, certain near-cosmopolitan genera such as Sapromyza and Homoneura may prove to comprise multiple distinct lineages when further revised (Gaimari & Silva 2010).

Characters (from Colless & McAlpine 1991): One or two fronto-orbital bristles; costa unbroken; Sc complete; Cu+1A discontinued well before margine; vein 2A generally visible beyond alula; anepisternite with one or two mesopleural bristles; tibiae with preapical dorsal bristles; sternite 1 well developed, sclerotised.

<==Lauxaniidae [Sapromyzidae]
    |--Lauxaniinae GS10
    |    |  i. s.: Lauxania GS10
    |    |           |--L. aenea RD77
    |    |           `--L. cylindricornis [=Micropeza (Lauxania) cylindricornis] G20
    |    `--Trigonometopini P07
    |--Sapromyza GS10 [Sapromyzinae K01]
    |    |--S. flavimana CM91
    |    |--S. frontalis GS10
    |    |--S. lupulina K01
    |    |--S. nigrans GS10
    |    |--S. picticornis GS10
    |    |--S. rotundicornis S87
    |    |--S. slossonae GS10
    |    |--S. sororia GS10
    |    `--S. vinnula GS10
    |--Homoneurinae GS10
    |    |--Trypetisoma GS10
    |    |    |--T. sticticum GS10
    |    |    `--T. zacatecasense GS10
    |    `--Homoneura GS10
    |         |  i. s.: H. bispina GS10
    |         |         H. proximella CM91
    |         |--H. (Homoneura) philadelphica GS10
    |         `--H. (Tarsohomoneura) GS10
    |              |--H. (T.) americana GS10
    |              `--H. (T.) ornatipes GS10
    `--Eurychoromyiinae [Eurychoromyiidae] GS10
         |--Eurychoromyia GS10
         |--Tauridion shewelli GS10
         |--Choryeuromyia senisma GS10
         |--Eurystratiomyia GS10
         |--Roryeuchomyia GS10
         |--Euryhendelimyia GS10
         `--Physegeniopsis GS10
              |--P. albeto GS10
              |--P. ankhoidea GS10
              `--P. hadrocara GS10

Lauxaniidae incertae sedis:
  Noonamyia O98 [Noonamyiini P07]
  Paranomina Hendel 1907 P07
  Lyciella S87
    |--L. decipiens WT11
    `--L. discolor S87
  Cerastocara ZS10
  Cephaloconus ZS10
  Poecilohetarus ZS10
  Neogriphoneura GS10
    |--N. laevifrons GS10
    |--N. sordida GS10
    |--N. striatifrons GS10
    |--N. striga GS10
    |--N. tertia GS10
    `--N. timida GS10
  Siphonophysa albipes GS10
  Neodeceia GS10
    |--N. cineracea GS10
    `--N. exul GS10
  Hypagoga GS10
  Oncodometopus umbrosus GS10
  Tricholauxania GS10
  Pseudocalliope GS10
    |--P. eucephala GS10
    |--P. flaviceps GS10
    `--P. longicornis S87
  Poecilominettia macula GS10
  Meiosimyza GS10
  Calliopum quadrisetosum GS10
  Pseudogriphoneura GS10
  Camptoprosopella borealis GS10
  Poecilolycia serrata GS10
  Melanomyza GS10
    |--M. (Melanomyza) GS10
    |    |--M. (M.) gracilipes GS10
    |    `--M. (M.) proana GS10
    `--M. (Lauxaniella) GS10
         |--‘Lauxaniella’ femoralis M81
         `--M. (L.) manuleata GS10
  Trivialia GS10
    |--T. celeste GS10
    |--T. fuscocapitata GS10
    |--T. nigroapicata GS10
    `--T. puella GS10
  Xenochaetina GS10
    |--X. flavipennis GS10
    |--X. muscaria GS10
    `--X. pallida GS10
  Physoclypeus GS10
    |--P. coquilletti GS10
    |--P. farinosus GS10
    |--P. hendeli GS10
    |--P. unimaculatus GS10
    `--P. zebrinus GS10
  Xeniconeura costalis GS10
  Trisapromyza vittigera GS10
  Allominettia GS10
    |--A. approximata GS10
    |--A. assimilis GS10
    `--A. geniseta GS10
  Mallochomyza citreifrons GS10
  Steganolauxania latipennis GS10
  Cephalella fumipennis GS10
  Triconopsis GS10
  Asilostoma GS10
    |--A. atriceps GS10
    |--A. flavifacies GS10
    |--A. pallipes GS10
    `--A. palpalis GS10
  Pachyopella GS10
    |--P. flavida GS10
    `--P. ornata GS10
  Stenolauxania GS10
    |--S. nubecula GS10
    `--S. striata GS10
  Minettia [Minettiinae] GS10
    |--M. aibonito GS10
    |--M. flaveola GS10
    |--M. hubbardii GS10
    `--M. lupulina GS10
  Xenopterella GS10
    |--X. beameri GS10
    `--X. obliqua GS10
  Baliopteridion GS10
  Hirtodeceia GS10
  Paracestrotus GS10
  Deutominettia GS10
    |--D. bimaculata GS10
    `--D. geniseta S87
  Scutominettia GS10
    |--S. innuba GS10
    `--S. mallochi GS10
  Griphoneura GS10
  Chaetominettia GS10
    |--C. corollae GS10
    `--C. spinitibia GS10
  Dryomyzothea GS10
  Setulina GS10
    |--S. geminata GS10
    |--S. interjecta GS10
    |--S. prima GS10
    `--S. stata GS10
  Chaetocoelia GS10
    |--C. caloptera GS10
    |--C. distinctissima GS10
    `--C. pallans GS10
  Dryosapromyza GS10
  Sciosapromyza GS10
    |--S. coronata GS10
    |--S. ficulnea GS10
    `--S. tinctinervis GS10
  Pseudominettia platypeza GS10
  Agriphoneura GS10
  Eriurgus GS10
  Ocellominettia GS10
  Celypholauxania GS10
  Procrita pectinata GS10
  Blepharolauxania GS10
  Platygraphium penicillatum GS10
  Freyia nigrita GS10
  Physegenua GS10
  Neoxangelina ornata GS10
  Minilauxania bulbifacies GS10
  Ritaemyia GS10
  Scutolauxania GS10
  Neopachycerina GS10
  Gibbolauxania GS10
    |--G. aliena GS10
    `--G. lateralis GS10
  Deceia GS10
  Marmarodeceia marmorata GS10
  Neominettia contigua GS10
  Depressa striatipennis CM91
  ‘Lonchaea’ senescens S87
  Eolausanites Hong 2002 H02
    `--*E. ellipticus (Hong 1981) [=Trypaneoides ellipticus] H02
  Chamaelauxania P92
  Hemilauxania P92

*Type species of generic name indicated

REFERENCES

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

[GS10] Gaimari, S. D., & V. C. Silva. 2010. Lauxaniidae (lauxaniid 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. 971–995. NRC Research Press: Ottawa.

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

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

[K01] Kertész, K. 1901. Legyek [Dipteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 179–201. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[M81] McAlpine, J. F. 1981. Morphology and terminology—adults. 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. 9–63. Research Branch, Agriculture Canada.

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

[P07] Papp, L. 2007. A review of the Old World Trigonometopini Becker (Diptera: Lauxaniidae). Annales Historico-Naturales Musei Nationalis Hungarici 99: 129–169.

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

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms' General Textbook of Entomology 10^th ed. vol. 2. Classification and Biology. Chapman and Hall: London.

[S87] Shewell, G. E. 1987. Lauxaniidae. In: McAlpine, J. F. (ed.) Manual of Nearctic Diptera vol. 2 pp. 951–964. Research Branch, Agriculture Canada.

[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.

[ZS10] Zborowski, P., & R. Storey. 2010. A Field Guide to Insects in Australia 3^rd ed. Reed New Holland: Sydney.

Last updated: 9 January 2021.

Homalorhagida

Pycnophyes calmani, from here.

Belongs within: Ecdysozoa.

The Homalorhagida are a group of marine kinorhynchs distinguished by their triangular cross-section.

Characters (from Higgins 1986): 6-8 neck plates; distinctly triangular in cross-section; spines reduced.

Homalorhagida [Homalorhagae]
    |--Neocentrophyes Higgins 1969 [Neocentrophyidae] H86
    |    |--N. intermedius Higgins 1969 H86
    |    `--N. satyai Higgins 1969 H86
    `--Pycnophyidae H86
         |--Kinorhynchus Sheremetevskij 1974 H86
         |    |--K. anomalus Lang 1953 H86
         |    |--K. apotomus Higgins 1983 H86
         |    |--K. belizensis Higgins 1983 H86
         |    |--K. cataphractus Higgins 1961 H86
         |    |--K. deirophorus Higgins 1983 H86
         |    |--K. distentus Higgins 1983 H86
         |    |--K. erismatus Higgins 1983 H86
         |    |--K. fimbriatus Higgins 1982 H86
         |    |--K. giganteus (Zelinka 1908) H86
         |    |--K. ilyocryptus Higgins 1961 H86
         |    |--K. langi (Higgins 1964) H86
         |    |--K. mainensis (Blake 1930) H86
         |    |--K. paraneapolitanus Sheremetevskij 1974 H86
         |    |--K. phyllotropis Brown & Higgins 1983 H86
         |    |--K. spinosus (Lang 1949) H86
         |    |--K. stenopygus Higgins 1983 H86
         |    `--K. trisetosus Higgins 1983 H86
         `--Pycnophyes Zelinka 1907 H86
              |--P. beaufortensis Higins 1964 H86
              |--P. calmani Southern 1914 H86
              |--P. carinatus Zelinka 1928 H86
              |--P. chiliensis Lang 1953 H86
              |--P. communis Zelinka 1908 H86
              |--P. corrugatus Higgins 1983 H86
              |--P. dentatus (Reinhard 1881) H86
              |--P. echinoderoides Zelinka 1928 H86
              |--P. ecphantor Higgins 1983 H86
              |--P. egyptensis Higgins 1966 H86
              |--P. emarginatus Higgins 1983 H86
              |--P. flagellatus H86
              |--P. flaveolatus Zelinka 1928 H86
              |--P. frequens Blake 1930 H86
              |--P. iniorhaptus Higins 1983 H86
              |--P. kielensis Zelinka 1928 H86
              |--P. longicornis Higgins 1983 H86
              |--P. maximus Reimer 1963 H86
              |--P. odhneri Lang 1953 H86
              |--P. ponticus (Reinhard 1881) H86
              |--P. robustus Zelinka 1928 H86
              |--P. rugosus Zelinka 1928 H86
              |--P. sanjuanensis Higgins 1961 H86
              |--P. sculptus Lang 1949 H86
              `--P. zelinkaei Southern 1914 H86

*Type species of generic name indicated

REFERENCES

[H86] Higgins, R. P. 1986. Kinorhyncha. In Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) (L. Botosaneanu, ed.) pp. 110-118. E. J. Brill / Dr. W. Backhuys: Leiden.

Cyclorhagida

Images of Campyloderes cf. vanhoeffeni from Neuhaus & Sørensen (2012). Original caption: "SEM (A–C, F–H) and DIC images (D, E, J, K) of male (C, J) and female (A, B, D–H, K) Campyloderes cf. vanhoeffeni from the Galápagos area (A, E), Antarctica (B, H) and the Faroe Islands (C, D, F, G, J, K). (A, B) Habitus from left side (A) and ventral side (B). Arrow heads mark sensory spots. (C) Mouth cone in surface view, individual outer oral styles absent. (D, E) Optical section of mouth cone of specimens ZMUC KIN-87 (D) and ZMB 5967i (E). Inner oral styles recognizable in specimen from Galápagos area (E) but not in Faroe specimen (D). Pharynx with alternating circular and radial muscle cells in (E). (F, G). Tip of mouth cone (F) and detail of fused outer oral styles (G). (H) Frontal view of placids. (J, K) Head and placids of specimens ZMUC KIN-95 (J) and KIN-94 (K) in ventral (J) and dorsal view (K). Scale bar in A and B 200 μm, in C and H 20 μm, in D 30 μm (valid for D, E), in F 5 μm, in G 2 μm, in J 30 μm (valid for J, K)."

Belongs within: Ecdysozoa.
Contains: Echinoderes.

The Cyclorhagida is the larger of the two major groups of kinorhynchs, small worm-like animals found mostly in marine sediments.

Characters (from Higgins 1986): 14-16 neck plates; generally round to oval-triangular; usually with numerous lateral middorsal and occasionally midterminal spines.

Cyclorhagida
    |--Cateria Gerlach 1956 [Cateriidae, Cryptorhagae] H86
    |    |--C. gerlachi Higgins 1968 H86
    |    |--C. styx Gerlach 1956 H86
    |    `--C. submersa Gerlach 1969 H86
    |--Semnoderidae [Conchorhagae] H86
    |    |--Sphenoderes Higgins 1969 H86
    |    |    `--S. indicus Higgins 1969 H86
    |    `--Semnoderes Zelinka 1907 H86
    |         |--S. armiger Zelinka 1928 H86
    |         |--S. pacificus Higgins 1967 H86
    |         `--S. ponticus Băcescu & Băcescu 1956 H86
    `--Cyclorhagae H86
         |--Echinoderes H86
         |--Condyloderes Higgins 1969 [Condyloderidae] H86
         |    |--C. multispinosus (McIntyre 1962) H86
         |    `--C. paradoxus Higgins 1969 H86
         `--Centroderidae H86
              |--Centroderes Zelinka 1896 H86
              |    |--C. eisigii Zelinka 1928 H86
              |    `--C. spinosus (Reinhard 1881) H86
              `--Campyloderes Zelinka 1913 H86
                   |--C. adherens Nyholm 1947 H86
                   |--C. macquariae Johnston 1938 H86
                   `--C. vanhoeffeni Zelinka 1913 H86

*Type species of generic name indicated

REFERENCES

[H86] Higgins, R. P. 1986. Kinorhyncha. In Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) (L. Botosaneanu, ed.) pp. 110-118. E. J. Brill / Dr. W. Backhuys: Leiden.

Neuhaus, B., & M. V. Sørensen. In press, 2012. Populations of Campyloderes sp. (Kinorhyncha, Cyclorhagida): one global species with significant morphological variation? Zoologischer Anzeiger.

Conopidae

Physocephala vittata, photographed by Pietro Niolu.

Belongs within: Sciomyzoidea.

The Conopidae is a family of flies of which most members are mimics of wasps. Larvae are endoparasites of other insects. Living conopids are divided between three subfamilies, Dalmanniinae, Myopinae and Conopinae (a fourth subfamily, Stylogastrinae, is probably not monophyletic with other conopids), with a further subfamily Palaeomyopinae known from the fossil record (Skevington et al. 2010). Members of the subfamily Conopinae have an apical three-segmented stylus on the antenna and a basally constricted abdomen. The Myopinae and Dalmanniinae possess a dorsal two- or three-segmented arista on the antenna. The ovipositor is short in Myopinae but elongate in Dalmannia. Within Conopinae, the genus Physocephala has crossvein r-m on the wing well beyond cell dm, and lacks ocelli or bristles on the proepisternum; Conopini have r-m at most a little past dm and have at least one bristle on the proepisternum (Smith & Peterson 1987).

Characters (from Smith & Peterson 1987): Rather bare and elongate flies, 3.0-20.0 mm long, usually pruinose, and colored black and yellow, reddish brown, or blackish, often with striking resemblance to solitary wasps and other Hymenoptera. Head broad; proboscis usually long, singly or doubly geniculate. Wing spotted or unmarked, or with a band along costal margin; cell r₄₊₅ closed or nearly so.

<==Conopidae [Archischiza, Conopoidea]
    |--Palaeomyopa SP87 [Palaeomyopinae STC10]
    |    `--P. tertiaria SP87
    |--Dalmannia SP87 [Dalmanniinae STC10]
    |    |--D. nigriceps SP87
    |    `--D. punctata SP87
    |--Myopinae STC10
    |    |--Robertsonomyia palpalis SP87
    |    |--Thecophora SP87
    |    |    |--T. nigripes STC10
    |    |    `--T. occidensis STC10
    |    |--Zodion Latreille 1802 SP87, L02
    |    |    |--*Z. cinereum L02 [=Conops (Zodion) cinereus G20, Myopa cinerea L02]
    |    |    |--Z. abitus SP87
    |    |    |--Z. americanum STC10
    |    |    |--Z. fulvifrons SP87
    |    |    |--Z. intermedium SP87
    |    |    |--Z. obliquefasciatum STC10
    |    |    `--Z. perlongum STC10
    |    `--Myopa STC10
    |         |--M. buccata SP87
    |         |--M. fasciata STC10
    |         |--M. ferruginea [=Conops (Myopa) ferrugineus] G20
    |         |--M. picta A71
    |         |--M. rubida SP87
    |         `--M. variegata K01
    `--Conopinae STC10
         |  i. s.: Pleurocerinella O98
         |--Physocephala STC10 [Physocephalini SP87]
         |    |--P. bimarginipennis SP87
         |    |--P. chrysorrhoea BM76
         |    |--P. flavipes SP87
         |    |--P. furcillata STC10
         |    |--P. gigas O98
         |    |--P. inhabilis STC10
         |    |--P. marginata SP87
         |    |--P. rufipes K01
         |    |--P. sagittaria SP87
         |    |--P. texana STC10
         |    `--P. vittata BM76
         `--Conopini SP87
              |--Physoconops SP87
              |    |--P. brachyrhynchus STC10
              |    |--P. bulbirostris STC10
              |    |--P. fronto STC10
              |    |--P. ornatifrons STC10
              |    `--P. (Gyroconops) sylvosus STC10
              `--Conops Linnaeus 1758 L58
                   |--C. aculeatus G20
                   |--C. algira Macquart 1847 E12
                   |--C. (Asiconops) bermudensis SP87
                   |--C. buccata Linnaeus 1758 L58
                   |--C. flavipes Linnaeus 1758 L58
                   |--C. fuscipennis Macquart 1847 E12
                   |--C. geminatus STC10
                   |--C. macrocephala Linnaeus 1758 L58
                   |--C. nobilis STC10
                   |--C. rostrata Linnaeus 1758 L58
                   |--C. rufiventris Macquart 1847 E12
                   `--C. splendidus CM91

Conopidae incertae sedis:
  Leopoldius coronatus A71
  Oncomyia K01
    |--O. distincta K01
    `--O. sundevalli K01
  *Parazodion schmidti STC10
  Tropidomyia bimaculata STC10
  Myopella Robineau-Desvoidy 1853 M10
  Microconops CM91
  Palaeosicus loewi SP87
  Zodiomyia O98
  Australoconops splendidus O98
  Poliomyia recta RJ93
  Sicus lineatus SP87, G20

*Type species of generic name indicated

REFERENCES

[A71] Askew, R. R. 1971. Parasitic Insects. Heinemann Educational Books: London.

[BM76] Bohart, R. M., & A. S. Menke. 1976. Sphecid Wasps of the World. University of California Press: Berkeley.

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

[E12] Evenhuis, N. L. 2012. Publication and dating of the Exploration Scientifique de l’Algérie: Histoire Naturelle des Animaux Articulés (1846–1849) by Pierre Hippolyte Lucas. Zootaxa 3448: 1–61.

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

[K01] Kertész, K. 1901. Legyek [Dipteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 179–201. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[L58] Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii: Holmiae.

[M10] Michelsen, V. 2010. Anthomyiidae (anthomyiid 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. 1271–1276. NRC Research Press: Ottawa.

[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.

[STC10] Skevington, J. H., F. C. Thompson & S. Camras. 2010. Conopidae (thick-headed 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. 847–855. NRC Research Press: Ottawa.

[SP87] Smith, K. G. V., & B. V. Peterson. 1987. Conopidae. In: McAlpine, J. F. (ed.) Manual of Nearctic Diptera vol. 2 pp. 749–756. Research Branch, Agriculture Canada.

Last updated: 29 June 2021.

Agromyzidae

Cambium miner Phytobia sp., copyright Rami P.

Belongs within: Schizophora.
Contains: Agromyza, Melanagromyza, Phytomyza, Cerodontha, Liriomyza.

The Agromyzidae are a group of mostly small or minute flies, the larvae of which form mines or galls in plants. Females have a well-sclerotised ovipositor that is used to insert eggs into plant tissue. As a result of damage from oviposition and larval feeding, some species are economically significant pests, particularly those such as the vegetable leaf miner Liriomyza sativae that may attack a wide range of hosts. The majority of species, however, restrict themselves to a single host species or closely related host species only (Boucher 2010). The fossil record of the family is limited; however, feeding channels in wood similar to those formed by the modern genus Phytobia are known from the Upper Miocene (Spencer 1987).

Agromyzids are divided between the subfamilies Agromyzinae, in which the subcostal vein fuses with vein R₁ before reaching the costa, and Phytomyzinae, in which the two veins usually reach the costa independently (some Phytobia species have the subcostal vein as in Agromyzinae). The subfamilies are also distinguished by the larval cephalopharyngeal skeleton having two upper arms in Agromyzinae but only one in Phytomyzinae (Boucher 2010).

Characters (from Boucher 2010): Minute to medium-sized flies (wing length 0.9–6.5 mm, usually 2–3 mm). Often yellow and/or black, brown or gray, few have some metallic greenish, bluish or coppery coloration. Head with first flagellomere small, round, or sometimes enlarged, elongate, or with anterodorsal projection or spine and with well-developed dorsobasal arista. Fronto-orbital and ocellar bristles trong; postocellar bristles divergent; fronto-orbital setulae in one or more rows, usually reclinate but sometimes proclinate, upright or absent; vibrissa well developed, some males with vibrissal fasciculus. Scutum with two to five dorsocentral bristles present but many genera in Phytomyzinae with three postsutural and one presutural dorsocentral bristles; acrostichals present (in up to ten rows) or absent. Costal break present near end of R₁, humeral break absent; Sc incomplete distally or reaching C as linear fold, independently of R₁, or Sc complete and fusing with R₁ before reaching costa; CuA₂ and A₁ present, forming small cell cup; A₁ not reaching wing margin. Abdomen tapered, composed of six visible segments. Females with abdominal segment 7 modified in large conical nonretractile oviscape.

<==Agromyzidae
    |  i. s.: Ceratomyza lateralis K01
    |         Selachops G10
    |         Aulagromyza B10
    |           |--A. nitida B10
    |           `--A. orbitalis B10
    |         Xeniomyza S87
    |         Penetagromyza S87
    |         Gymnophytomyza S87
    |         Pseudoliriomyza cordiae O98
    |--Agromyzinae B10
    |    |--Agromyza B10
    |    |--Tropicomyia Spencer 1973 S87
    |    |--Hexomyza S87
    |    |--Melanagromyza B10
    |    |--Ophiomyia B10
    |    |    |--O. aberrans S87
    |    |    |--O. centrosematis O98
    |    |    |--O. labiatarum S87
    |    |    |--O. lantanae B10
    |    |    |--O. maura B10
    |    |    |--O. phaseoli B10
    |    |    `--O. solanicola B88
    |    `--Japanagromyza B10
    |         |--J. jamaicensis B10
    |         |--J. maculata [=Geratomyza maculata] B10
    |         |--J. nebulifera B10
    |         |--J. orbitalis B10
    |         |--J. phaseoli B10
    |         |--J. polygoni B10
    |         `--J. spadix [=Geratomyza spadix] B10
    `--Phytomyzinae B10
         |--Phytomyza B10
         |--Haplopeodes B10
         |--Galiomyza B10
         |--Cerodontha B10
         |--Liriomyza B10
         |--Metopomyza S87
         |--Napomyza lateralis S87
         |--Paraphytomyza S87
         |    |--P. nitida S87
         |    `--P. orbitalis S87
         |--Pseudonapomyza B10
         |    |--P. asiatica B10
         |    |--P. cingulata B88
         |    `--P. lacteipennis B10
         |--Nemorimyza B10
         |    |--N. maculosa B10
         |    |--N. posticata B10
         |    `--N. ranchograndensis B10
         |--Amauromyza B10 [incl. Campanulomyza S87, Melanophytobia S87]
         |    |--A. abnormalis B10
         |    |--A. boliviensis B10
         |    |--A. (Trilobomyza) flavifrons S87
         |    |--A. karli S87
         |    `--A. (Catalpomyza) pleuralis S87
         |--Calycomyza B10
         |    |--C. devia B10
         |    |--C. illustris B10
         |    |--C. menthae S87
         |    |--C. meridiana B10
         |    `--C. obscura B10
         |--Phytoliriomyza B10 [incl. Lemurimyza S87, Nesomyza S87, Pteridomyza S87]
         |    |--P. conjunctimontis B10
         |    |--P. costaricensis B10
         |    |--P. cyathea Spencer 1976 M03
         |    |--P. hilarella S87
         |    |--P. leechi B10
         |    `--P. pacifica S87
         `--Phytobia B10
              |--P. coylesi S87
              |--P. guatemalensis B10
              |--P. kallima B10
              |--P. mentula B10
              |--P. pipinna B10
              |--P. rabelloi B10
              `--P. unica B10

*Type species of generic name indicated

REFERENCES

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

[B10] Boucher, S. 2010. Agromyzidae (leaf-mining 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. 1057–1071. NRC Research Press: Ottawa.

[G10] Gaimari, S. D. 2010. Odiniidae (odiniid 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. 1049–1055. NRC Research Press: Ottawa.

[K01] Kertész, K. 1901. Legyek [Dipteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 179–201. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[M03] Martin, N. 2003. Changes in abundance of the silver fern leaf miner. Weta 25: 31–32.

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

[S87] Spencer, K. A. 1987. Agromyzidae. In: McAlpine, J. F. (ed.) Manual of Nearctic Diptera vol. 2 pp. 869–879. Research Branch, Agriculture Canada.

Last updated: 2 July 2021.

Drosophila (Drosophila)

Drosophila virilis, photographed by Darren J. Obbard.

Belongs within: Drosophila.

Drosophila subgenus Drosophila is one of the major recognised subdivisions of the large genus Drosophila. Two major subdivisions have been recognised within the subgenus as the virilis-repleta and quinaria-tripunctata sections; a phylogenetic study by Robe et al. (2005) supported the monophyly of the former as well as a clade including most (but not all) species studied of the latter. The same phylogenetic analysis also suggested that some lineages recognised as distinct genera may be nested within the subgenus.

Characters (from Singh et al. 2004): Gena often broad; second oral bristle large; prescutellar acrostichals not, or barely, enlarged; propleural bristle absent; apical bands on anterior abdominal tergites, when present, usually interrupted in midline; usually rather large species.

<==Drosophila Fallén 1823 (Drosophila) SDF04
    |--+--+--D. nannoptera group CE79
    |  |  |    |--D. (D.) nannoptera RV05
    |  |  |    `--D. (D.) pachea CE79
    |  |  `--D. (D.) pallidipennis RV05
    |  |       |--D. p. pallidipennis D51
    |  |       `--D. p. centralis D51
    |  `--D. virilis-repleta section RV05
    |       |  i. s.: D. (D.) aracea T68
    |       |         D. (D.) bromeliae T68
    |       |         D. (D.) castanea T68
    |       |         D. (D.) micromelanica T68
    |       |         D. (D.) paramelanica T68
    |       |         D. (D.) peruviana T68
    |       |         D. (D.) serenensis T68
    |       |         D. (D.) tira T68
    |       |--+--D. (D.) robusta RV05
    |       |  `--D. virilis group CE79
    |       |       |--D. (D.) americana CE79
    |       |       |    |--D. a. americana CE79
    |       |       |    `--D. a. texana CE79
    |       |       |--D. (D.) borealis CE79
    |       |       |--D. (D.) ezoana CE79
    |       |       |--D. (D.) flavomontana CE79
    |       |       |--D. (D.) lacicola CE79
    |       |       |--D. (D.) littoralis CE79
    |       |       |--D. (D.) lummei CE79
    |       |       |--D. (D.) montana CE79
    |       |       |    |--D. m. montana CE79
    |       |       |    `--D. m. ovivororum CE79
    |       |       |--D. (D.) novamexicana CE79
    |       |       `--D. (D.) virilis RV05
    |       `--+--+--D. annulimana group RV05
    |          |  |    |--D. (D.) annulimana RV05
    |          |  |    `--D. (D.) aracatacas RV05
    |          |  `--D. flavopilosa group RV05
    |          |       |--D. (D.) cestri RV05
    |          |       |--D. (D.) flavopilosa RV05
    |          |       `--D. (D.) incompta RV05
    |          `--+--D. (D.) canalinea RV05
    |             `--+--D. dreyfusi group RV05
    |                |    |--D. (D.) camargoi RV05
    |                |    `--D. (D.) dreyfusi RV05
    |                `--+--D. repleta group RV05
    |                   |    |  i. s.: D. (D.) nigrohydei RV05
    |                   |    |         D. melanopalpa subgroup CE79
    |                   |    |           |--D. (D.) fulvimacula CE79
    |                   |    |           |--D. (D.) melanopalpa CE79
    |                   |    |           `--D. (D.) repleta Wollaston 1858 CE79, SDF04
    |                   |    |--D. mercatorum subgroup CE79
    |                   |    |    |--D. (D.) mercatorum RV05
    |                   |    |    `--D. (D.) paranaensis CE79
    |                   |    `--+--D. hydei subgroup CE79
    |                   |       |    |--D. (D.) bifurca CE79
    |                   |       |    |--D. (D.) eohydei CE79
    |                   |       |    |--D. (D.) hydei RV05
    |                   |       |    `--D. (D.) neohydei CE79
    |                   |       `--+--D. mulleri subgroup CE79
    |                   |          |    |--D. (D.) aldrichi CE79
    |                   |          |    |--D. (D.) arizonensis CE79
    |                   |          |    |--D. (D.) buzzatii RV05
    |                   |          |    |--D. (D.) meridiana CE79
    |                   |          |    |--D. (D.) mohavensis CE79
    |                   |          |    |--D. (D.) mulleri CE79
    |                   |          |    `--D. (D.) navajoa CE79
    |                   |          `--+--D. (D.) borborema RV05
    |                   |             `--D. (D.) serido RV05
    |                   `--D. mesophragmatica group RV05
    |                        |--D. (D.) viracochi RV05
    |                        `--+--+--D. (D.) gaucha RV05
    |                           |  `--D. (D.) pavani RV05
    |                           `--+--D. (D.) mesophragmatica RV05
    |                              `--+--D. (D.) brncici RV05
    |                                 `--D. (D.) gasici RV05
    `--+--+--Liodrosophila aerea RV05
       |  `--Zaprionus RV05
       |       |--Z. indianus RV05
       |       `--Z. tuberculatus RV05
       `--+--+--Samoaia leonensis RV05
          |  `--Hirtodrosophila Duda 1923 RV05, SDF04 [=Dasydrosophila Duda 1925 SDF04]
          |       |--*H. latifrontata (Fronta-Pessoa 1945) (see below for synonymy) SDF04
          |       |--H. batracida [=Drosophila (Hirtodrosophila) batracida] G10
          |       `--H. pictiventris RV05
          `--+--D. (D.) immigrans Sturtevant 1921 RV05, SDF04
             `--+--D. quinaria group RV05
                |    |--D. (D.) kunzei CE79
                |    |--D. (D.) phalerata RV05
                |    `--D. (D.) quinaria RV05
                `--+--+--*D. (D.) funebris (Fabricius 1787) SDF04, RV05, SDF04 [=Musca funebris SDF04]
                   |  `--D. testacea group RV05
                   |       |--D. (D.) putrida RV05
                   |       `--D. (D.) testacea RV05
                   `--+--D. (D.) nappae [incl. D. angustibucca] RV05
                      `--+--D. cardini group RV05
                         |    |--D. (D.) polymorpha RV05
                         |    `--+--D. (D.) cardini RV05
                         |       `--+--D. (D.) cardinoides RV05
                         |          `--D. (D.) neocardini RV05
                         `--+--+--D. (D.) tripunctata RV05
                            |  `--+--D. (D.) bandeirantorum RV05
                            |     `--D. (D.) mediopunctata RV05
                            `--+--+--D. (D.) calloptera RV05
                               |  `--+--D. (D.) paramediostriata RV05
                               |     `--D. guaramunu group RV05
                               |          |--D. (D.) griseolineata RV05
                               |          |--D. (D.) guaramunu RV05
                               |          `--D. (D.) maculifrons RV05
                               `--+--D. (D.) mediostriata [incl. D. crocina] RV05
                                  `--D. guarani group RV05
                                       |  i. s.: D. (D.) guarani RV05
                                       |--D. (D.) ornatifrons RV05
                                       `--+--D. (D.) guaru RV05
                                          `--D. (D.) subbadia RV05

Drosophila (Drosophila) incertae sedis:
  D. (D.) analspina Singh & Negi 1995 SDF04
  D. (D.) bageshwarensis Singh, Dash & Fartyal 2004 SDF04
  D. (D.) bishti Singh & Negi 1995 SDF04
  D. (D.) elongata Singh, Dash & Fartyal 2004 SDF04
  D. (D.) falleni T68
  D. (D.) hexaspina Singh, Dash & Fartyal 2004 SDF04
  D. (D.) hypocausta T68
  D. (D.) khansuensis Singh, Dash & Fartyal 2004 SDF04
  D. (D.) lacertosa Okada 1956 SDF04
  D. (D.) medioimpressa RV05
  D. (D.) nainitalensis Singh & Bhatt 1988 SDF04
  D. (D.) nasuta Lamb 1914 SDF04
  D. (D.) paharpaniensis Singh, Dash & Fartyal 2004 SDF04
  D. (D.) painii Singh & Negi 1995 SDF04
  D. (D.) paramarginata Singh, Dash & Fartyal 2004 SDF04
  D. (D.) paunai Singh & Negi 1989 SDF04
  D. (D.) procardinoides T68
  D. (D.) serrata Singh, Dash & Fartyal 2004 SDF04
  D. (D.) sticta T68
  D. (D.) sulfurigaster Duda 1923 SDF04
  D. (D.) surangensis Singh, Dash & Fartyal 2004 SDF04
  D. (D.) trisetosa Okada 1966 SDF04
  D. (D.) trizonata Okada 1966 SDF04
  D. (D.) uninubes T68
  0--D. (D.) mediopicta RV05
  `--+--+--D. (D.) cuaso RV05
     |  `--D. (D.) metzii RV05
     `--+--D. (D.) mediosignata RV05
        `--D. (D.) paraguayensis RV05

*Hirtodrosophila latifrontata (Fronta-Pessoa 1945) [=Drosophila carinata Duda 1923 non Grimshaw 1901, D. (*Hirtodrosophila) latifrontata] SDF04

*Type species of generic name indicated

REFERENCES

[CE79] Coyne, J. A., W. F. Eanes, J. A. M. Ramshaw & R. K. Koehn. 1979. Electrophoretic heterogeneity of α-glycerophosphate dehydrogenase among many species of Drosophila. Systematic Zoology 28 (2): 164–175.

[D51] Dobzhansky, T. 1951. Genetics and the Origin of Species 3^rd ed. Columbia University Press: New York.

[G10] Grimaldi, D. 2010. Drosophilidae (small fruit flies, pomace flies, vinegar 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. 1197–1206. NRC Research Press: Ottawa.

[RV05] Robe, L. J., V. L. S. Valente, M. Budnik & E. L. S. Loreto. 2005. Molecular phylogeny of the subgenus Drosophila (Diptera, Drosophilidae) with an emphasis on Neotropical species and groups: a nuclear versus mitochondrial gene approach. Molecular Phylogenetics and Evolution 36: 623–640.

[SDF04] Singh, B. K., S. Dash & R. S. Fartyal. 2004. Revision of the species of the subgenus Drosophila (Drosophila) of the Kumaon region, India, with the description of eight new species. Senckenbergiana Biologica 83 (2): 163–176.

[T68] Throckmorton, L. H. 1968. Concordance and discordance of taxonomic characters in Drosophila classification. Systematic Zoology 17 (4): 355–387.

Last updated: 2 December 2019.