Cyclorrhapha

Big-headed fly Eudorylas sp., photographed by Nikola Rahmé.


Belongs within: Brachycera.
Contains: Syrphidae, Schizophora, Phoridae.

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. The earliest cyclorrhaphans are known from the Cretaceous (Grimaldi & Engel 2005).

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pupariation in ecdysed cuticle of last larval instar; arista with two small basal segments; proboscis retractile, withdrawn at rest into large oral cavity; female terminalia without acanthophorite spines.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, Muscomorpha]
    |--Chimeromyia GE05
    `--+--Lonchopterites prisca GE05
       `--+--Syrphoidea M90
          |    |--Syrphidae WT11
          |    `--+--+--Cretophormia fowleri GE05
          |       |  `--Schizophora WT11
          |       `--Pipunculidae [Dorilaidae] WT11
          |            |--Chalarus A71
          |            |--Alloneura A71
          |            |--Verrallia aucta A71
          |            |--Dorylomorpha A71
          |            |--Nephrocerus A71
          |            |--Dasydorylas filiformis Kehlmaier 2005 F05
          |            |--Cephalops WT11
          |            |    |--C. longistylis WT11
          |            |    `--C. mauritianus A71
          |            |--Eudorylas F05
          |            |    |--E. auctus Kehlmaier 2005 F05
          |            |    |--E. ibericus Kehlmaier 2005 F05
          |            |    |--E. venturai Kehlmaier 2005 F05
          |            |    `--E. zonatus A71
          |            `--Pipunculus B88
          |                 |--P. ater M90
          |                 |--P. campestris A71
          |                 |--P. cinerascens B88
          |                 `--P. fuscus M90
          `--Platypezoidea WT11
               |--+--Lonchopteromorpha asetocella GE05
               |  `--Lonchoptera M83 [Lonchopteridae WT11, Lonchopteroidea]
               |       |--L. furcata M83
               |       `--L. uniseta WT11
               `--+--Opetiala shatalkani GE05
                  `--+--Opetiidae WT11
                     |    |--Lithopetia hirsuta GE05
                     |    `--Opetia nigra GE05
                     `--+--+--Electrosania cretica GE05
                        |  `--Platypezidae WT11
                        |       |--Maritulus GE05
                        |       |--Parnassos GE05
                        |       |--Proplatypeza GE05
                        |       |--Lebambromyia acrai GE05
                        |       |--Promittor GE05
                        |       |--Oloplatypeza GE05
                        |       |--Paraplatypeza velutina WT11
                        |       |--Plesioclythia agarici M90
                        |       |--Polypivora M90
                        |       `--Platypeza consobrina M90
                        `--+--Ironomyiidae WT11
                           |    |--Paleopetia laiyangensis GE05
                           |    |--Hermaeomyia GE05
                           |    |--Eridomyia GE05
                           |    |--Cretonomyia pristina GE05
                           |    |--Sinolesta [Sinolestinae] GE05
                           |    `--Ironomyia [Ironomyiinae] GE05
                           |         `--I. nigromaculata WT11
                           `--+--Archisciada lebanensis GE05
                              `--+--Phoridae WT11
                                 `--Sciadoceridae GE05
                                      |--Sciadocera rufomaculata GE05
                                      `--Archiphora GE05
                                           |--A. patagonica GE05
                                           |--A. pria GE05
                                           `--A. robusta GE05

*Type species of generic name indicated

REFERENCES

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

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

[F05] Fernández, J. 2005. Noticia de nuevos táxones para la ciencia en el ámbito Íbero-Balear y Macaronésico. Nuevos táxones animales descritos en la península Ibérica y Macaronesia desde 1994 (IX). Graellsia 61 (2): 261-282.

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

[M83] Martin, N. A. 1983. Miscellaneous observations on a pasture fauna: An annotated species list. DSIR Entomology Division Report 3: 1-98.

[M90] McAlpine, J. F. 1990. Insecta: Diptera adults. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 1211-1252. John Wiley & Sones: New York.

[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.

Schizophora

Geomyza tripunctata, from here.


Belongs within: Cyclorrhapha.
Synapomorphies: Ephydroidea, Calyptratae, Agromyzidae, Sphaeroceridae, Tephritoidea, Nerioidea, Diopsidae, Chloropidae, Sciomyzoidea, Lauxanioidea, Heleomyzidae.

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. Schizophorans have often been divided for convenience between the Calyptrata and Acalyptrata (defined primarily by the presence or absence of an enlarged calypter at the base of the wing), though 'Acalyptrata' have long been recognised as an artificial group. However, the relationships of many 'acalyptrate' families have only recently begun to be resolved.

Many families of acalyptrate flies are difficult to distinguish without close examination. The Odiniidae are small flies closely associated as adults with trees; their larvae, where known, are often associated with wood- and plant-boring insects such as beetles or caterpillars. The Carnidae are saprophagous as larvae and in many species as adults, but adults of Carnus hemapterus are ectoparasitic on nestling birds, generally losing their wings through breakage.

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
    |     |    |--Neoalticomerus seamansi WT11
    |     |    |--Odinia betulae M81b
    |     |    `--Traginops irroratus [=T. irrorata] BW09
    |     `--+--+--Agromyzidae WT11
    |        |  `--Sphaeroceridae WT11
    |        `--+--Carnidae WT11
    |           |    |--Carnus hemapterus A71
    |           |    |--Meoneura obscurella CW09
    |           |    `--Hemeromyia anthracina KP10
    |           `--Periscelididae WT11
    |                |  i. s.: Periscelis BW09
    |                |           |--P. annulata T81a
    |                |           `--P. flinti BW09
    |                |         Scutops BW09
    |                `--Stenomicrinae BW09
    |                     |--Stenomicra BW09
    |                     `--Planinasus BW09
    `--+--Megamerina [Megamerinidae] WT11
       `--+--+--Tephritoidea WT11
          |  `--+--Chyromyidae WT11
          |     |    |--Gymnochiromyia concolor WT11, M81b
          |     |    |--Chyromya flava M81b
          |     |    `--Aphaniosoma M81b
          |     `--+--Opomyzidae WT11
          |        |    |--Opomyza WT11
          |        |    |    |--O. florum WT11
          |        |    |    `--O. petrei T81b
          |        |    `--Geomyza WT11
          |        |         |--G. apicalis K-P87
          |        |         |--G. balachowskyi T81b
          |        |         |--G. lurida M90
          |        |         `--G. tripunctata WT11
          |        `--+--Acartophthalmus [Acartophthalmidae] WT11
          |           |    `--A. nigrinus WT11
          |           `--Sepsidae [Sepsinae] WT11
          |                |--Orygma luctuosum M90, KP10
          |                |--Nemopoda cylindrica K01
          |                |--Saltella scutellaris K01
          |                |--Palaeosepsis BW09
          |                |--Archisepsis BW09
          |                `--Sepsis WT11
          |                     |--S. cynipsea WT11
          |                     |--S. neocynipsea T90
          |                     `--S. punctum M90
          `--+--+--+--+--Inbiomyia [Inbiomyiidae] WT11
             |  |  |  |    `--I. mcalpineorum WT11
             |  |  |  `--Nemula [Neminidae] WT11
             |  |  |       `--N. longarista WT11
             |  |  `--+--Nerioidea WT11
             |  |     `--Aulacigastridae WT11
             |  |          |--Cyamops nebulosa WT11
             |  |          `--Aulacigaster BW09
             |  |               |--A. leucopeza M81b
             |  |               `--A. neoleucopeza BW09
             |  `--+--+--Diopsidae WT11
             |     |  `--+--Margo [Marginidae] WT11
             |     |     `--+--Nannodastiidae WT11
             |     |        |    |--Azorastia mediterranea WT11
             |     |        |    `--Nannodastia horni BW09
             |     |        `--Canacidae WT11
             |     |             |--Procanace dianneae WT11
             |     |             |--Tethinosoma fulvifrons WT11
             |     |             |--Canacea macateei BW09
             |     |             |--Canaceoides nudatus M81b
             |     |             `--Nocticanace BW09
             |     `--+--Somatia [Somatiidae] WT11
             |        |    `--S. aestiva WT11
             |        `--+--Chloropidae WT11
             |           `--Milichiidae WT11
             |                |--Paramyia nitens WT11
             |                |--Desmometopa m-nigrum M90
             |                |--Madiza glabra M90
             |                |--Leptometopa halteralis M81a
             |                |--Phyllomyza securicornis BW09
             |                |--Aldrichiomyza agromyzina BW09
             |                |--Eusiphona mira BW09
             |                |--Phloeomyia indecora BW09
             |                `--Milichiella BW09
             `--+--+--Strongylophthalmyia [Strongylophthalmyiidae] WT11
                |  |    |--S. angustipennis M90
                |  |    `--S. pengellyi WT11
                |  `--Tanypezidae [Tanypezinae] WT11
                |       |--Neotanypeza elegans BW09
                |       |--Tanypeza longimana M90
                |       `--Calobata cibaria K01
                `--+--+--+--Heteromyza atricornis WT11
                   |  |  `--+--Sciomyzoidea WT11
                   |  |     `--Lauxanioidea WT11
                   |  `--+--Heleomyzidae WT11
                   |     `--Teratomyzidae WT11
                   |          |--Teratomyza WT11
                   |          `--Camur willii BW09
                   `--+--+--Ropalomeridae WT11
                      |  |    |--Willistoniella pleuropunctata WT11
                      |  |    |--Ropalomera femorata M81b
                      |  |    `--Lenkokroeberia BW09
                      |  `--+--+--Australimyza [Australimyzidae] WT11
                      |     |  `--Xenasteia [Xenasteiidae] WT11
                      |     |       `--X. shalam WT11
                      |     `--+--Fergusonina [Fergusoninidae] WT11
                      |        |    `--F. turneri WT11
                      |        `--Asteiidae WT11
                      |             |--Asteia amoena WT11
                      |             |--Leiomyza laevigata M81b
                      |             |--Phlebosotera setipalpis BW09
                      |             |--Loewimyia BW09
                      |             `--Sigaloessa BW09
                      `--+--+--Syringogaster [Syringogastridae] WT11
                         |  |    |--S. brunnea BW09
                         |  |    `--S. rufa BW09
                         |  `--Psilidae [Psilinae] WT11
                         |       |--Chamaepsila hennigi WT11
                         |       |--Loxocera cylindrica T81b
                         |       |--Chyliza BW09
                         |       `--Psila M90
                         |            |--P. gracilis K01
                         |            |--P. nigra K01
                         |            |--P. pallida K01
                         |            `--P. rosae M90
                         `--+--Clusiidae WT11
                            |    |--Clusia lateralis WT11
                            |    |--Clusiodes albimanus M81b
                            |    |--Chaetoclusia bakeri M81b
                            |    `--Sobarocephala BW09
                            |         |--S. flaviseta BW09
                            |         `--S. picta BW09
                            `--+--Neurochaeta [Neurochaetidae] WT11
                               `--Pallopteridae WT11
                                    |--Eurygnathomyia M81a
                                    |--Temnosira subarcuata M81b
                                    `--Palloptera WT11
                                         |--P. claripennis M81b
                                         |--P. superba M81b
                                         `--P. umbellatarum WT11

Schizophora incertae sedis:
  Mormotomyia [Mormotomyiidae] GE05
    `--M. hirsuta GE05
  Borborinae K01
    |--Heteroptera acutangula K01
    `--Borborus K01
         |--B. equinus K01
         |--B. oeneus Macquart 1849 E12
         `--B. vitripennis K01
  Nothybidae WBW09
  Paraleucopidae BW09
    |--Paraleucopis corvina BW09
    `--Mallochianamyia BW09

Schizophora [Acalypterae, Acalyptrata, Agromyzoinea, Asteioinea, Carnoidea, Clusioinea, Diopsoidea, Holometopa, Opomyzoidea, Opomyzoidea, Sphaeroceroidea] WT11

*Type species of generic name indicated

REFERENCES

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

[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. Ottawa, NRC Research Press.

[CW09] Cumming, J. M., & D. M. Wood. 2009. Adult morphology and terminology. 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. 9-50. Ottawa, NRC Research Press.

[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.

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

[K01] Kertész, K. 1901. Legyek [Dipteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy], Band II. 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.

[K-P87] Kukalová-Peck, J. 1987. New Carboniferous Diplura, Monura, and Thysanura, the hexapod ground plan, and the role of thoracic side lobes in the origin of wings (Insecta). Canadian Journal of Zoology 65: 2327-2345.

[KP10] Kutty, S. N., T. Pape, B. M. Wiegmann & R. Meier. 2010. Molecular phylogeny of the Calyptratae (Diptera: Cyclorrhapha) with an emphasis on the superfamily Oestroidea and the position of Mystacinobiidae and McAlpine's fly. Systematic Entomology 35: 614-635.

[M81a] McAlpine, J. F. 1981a. 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.

[M81b] McAlpine, J. F. 1981b. Key to families—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. 89-124. Research Branch, Agriculture Canada.

[M90] McAlpine, J. F. 1990. Insecta: Diptera adults. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 1211-1252. John Wiley & Sones: New York.

[T81a] Teskey, H. J. 1981a. Morphology and terminology—larvae. 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. 65-88. Research Branch, Agriculture Canada.

[T81b] Teskey, H. J. 1981b. Key to families—larvae. 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. 125-147. Research Branch, Agriculture Canada.

[T90] Teskey, H. J. 1990. Insecta: Diptera larvae. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 1253-1276. John Wiley & Sones: New York.

[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.

[WBW09] Woodley, N. E., A. Borkent & T. A. Wheeler. 2009. Phylogeny of the Diptera. 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. 79-94. Ottawa, NRC Research Press.

Lauxaniidae

Luzonomyza sp., from here.


Belongs within: Schizophora.

The Lauxaniidae are a family of flies whose larvae mostly live in fallen leaves and other rotting vegetation.

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
    |  i. s.: Noonamyiini P07
    |         Paranomina Hendel 1907 P07
    |         Lyciella decipiens WT11
    |         Minettia flaveola WT11
    |--Homoneura [Homoneurinae] P07
    `--Trigonometopini [Lauxaniinae] P07
         |--Kerteszomyia Malloch 1929 P07
         |    `--*K. maculifrons (Malloch 1929) [=Trigonometopus maculifrons] P07
         |--Trigonometopus Macquart 1835 P07
         |    |--*T. frontalis (Meigen 1830) [=Tetanocera frontalis] P07
         |    |--T. canus De Meijere 1916 P07
         |    `--T. eborifacies Shatalkin 1997 P07
         |--Neotrigonometopus Malloch 1928 P07
         |    |--*N. fuscifrons (Malloch 1926) [=Trigonometopus fuscifrons] P07
         |    |--N. albibasis (Malloch 1928) [=Trigonometopus (Neotrigonometopus) albibasis] P07
         |    `--N. breviceps (Hennig 1948) P07
         |--Trigonometopsis Malloch 1925 P07
         |    |--*T. binotata (Thomson 1869) [=Oxyrhina binotata] P07
         |    |--T. abnormis Shatalkin 1999 P07
         |    `--T. punctipennis Malloch 1929 P07
         |--Protrigonometopus Hendel 1938 P07
         |    |--*P. maculifrons Hendel 1938 P07
         |    |--P. ornatus Papp 2007 P07
         |    |--P. sexlituris (Shatalkin 1992) [=Sapromyxa sexlituris] P07
         |    `--P. shatalkini Papp 2007 P07
         |--Diplochasma Knab 1914 P07
         |    |--D. (Diplochasma) P07
         |    |    |--*D. (D.) monochaeta (Hendel 1909) [=Trigonometopus monochaeta] P07
         |    |    `--D. (D.) australis Papp 2007 P07
         |    `--D. (Sauteromyia Malloch 1927) P07
         |         |--D. (*S.) alboapicata (Malloch 1927) [=*Sauteromyia alboapicata] P07
         |         |--D. (S.) albomarginata Papp 2008 P08
         |         |--D. (S.) aphaniosignata Papp 2008 P08
         |         |--D. (S.) clara Papp 2008 P08
         |         |--D. (S.) occidentalis Papp 2008 P08
         |         |--D. (S.) punctata Papp 2008 P08
         |         |--D. (S.) shewelli Papp 2008 P08
         |         |--D. (S.) stuckenbergi Papp 2008 P08
         |         `--D. (S.) variegata Papp 2008 P08
         |--Shaltalkinella Papp 2007 P07
         |    |--*S. marginata Papp 2007 P07
         |    |--S. deceptor (Malloch 1927) [=Sapromyza deceptor] P07
         |    |--S. okinawanus (Shatalkin 1998) [=Protrigonometopus okinawanus, Trigonometopsis okinawanus] P07
         |    |--S. punctipennis Papp 2007 P07
         |    |--S. ronkayi Papp 2007 P07
         |    `--S. thailandica Papp 2007 P07
         |--Luzonomyza Malloch 1929 P07
         |    |--*L. bakeri (Bezzi 1913) [=Trigonometopus bakeri] P07
         |    |--L. japonica (Sasakawa 2002) [=Trigonometopus (Luzonomyia) japonica] P07
         |    |--L. nigripalpis (Shatalkin 1997) [=Trigonometopus (Tetroxyrhina) nigripalpis] P07
         |    |--L. pseudoforficula Papp 2007 P07
         |    |--L. sasakawai Papp 2007 P07
         |    |--L. sinica Shatalkin 1998 P07
         |    `--L. vietnamensis Papp 2007 P07
         |--Maquilingia Malloch 1929 P07
         |    |--*M. hirticeps Malloch 1929 P07
         |    |--M. biroi Papp 2007 P07
         |    |--M. facialis Malloch 1929 P07
         |    |--M. malaita Curran 1936 P07
         |    |--M. matema Curran 1936 P07
         |    |--M. serristylis Sasakawa 2003 P07
         |    `--M. thaii Papp 2007 P07
         `--Tetroxyrhina Hendel 1938 P07
              |--*T. submaculipennis (Malloch 1927) [=Trigonometopus submaculipennis] P07
              |--T. albiseta (Bezzi 1913) [=Trigonometopus albiseta] P07
              |--T. alboapicalis (Shatalkin 1997) [=Trigonometopus alboapicalis] P07
              |--T. brevicornis (De Meijere 1911) [=Trigonometopus brevicornis] P07
              |--T. brunneicosta (Malloch 1927) [=Trigonometopus brunneicosta] P07
              |--T. cuneata (Shatalkin 1997) [=Trigonometopus cuneatus] P07
              |--T. forficula (Shatalkin 1997) [=Trigonometopus forficula] P07
              |--T. gressitti (Sasakawa 2002) [=Trigonometopus gressitti] P07
              |--T. interrupta (Sasakawa 2005) [=Luzonomyza (Tetroxyrhina) interrupta] P07
              |--T. peregovitsi Papp 2007 P07
              |--T. sauteri (Hendel 1912) [=Trigonometopus sauteri, Diplochasma sauteri] P07
              |--T. tinctipennis (De Meijere 1924) [=Trigonometopus tinctipennis] P07
              `--T. zeylanica (Senior-White 1921) [=Trigonometopus zeylanicus] P07

*Type species of generic name indicated

REFERENCES

Colless, D. H., & D. K. McAlpine. 1991. Diptera. In: CSIRO. The Insects of Australia, pp. 717-786. Melbourne University Press.

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

[P08] Papp, L. 2008. Afrotropical species of the genus Diplochasma, subgenus Sauteromyia (Diptera: Lauxaniidae). Annales Historico-Naturales Musei Nationalis Hungarici 100: 307-330.

[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.

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.

Characters (from Colless & McAlpine 1991): Mid-coxal prong absent; proboscis usually very slender, often much longer than head; palp small or absent; vein R1 bare; cell CuP acute distally, usually long.

Conopidae
    |--Leopoldius coronatus A71
    |--Myopa picta A71
    |--Conops flavipes A71
    |--Physoconops A71
    |--Physocephala WT11
    |    |--P. marginata WT11
    |    `--P. vittata A71
    |--Sicus A71
    |--Zodion A71
    |--Thecophora A71
    `--Dalmannia A71

*Type species of generic name indicated

REFERENCES

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

Colless, D. H., & D. K. McAlpine. 1991. Diptera. In: CSIRO. The Insects of Australia, pp. 717-786. Melbourne University Press.

[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.

Agromyzidae

Serpentine leafminer Liriomyza brassicae, from here.


Belongs within: Schizophora.

The Agromyzidae are a group of mostly small or minute flies, the larvae of which form mines or galls in plants. As a result, some species are economically significant pests, such as the vegetable leafminer Liriomyza sativae and the American serpentine leafminer L. trifolii.

Characters (from Colless & McAlpine 1991): Postvertical bristles divergent; lower fronto-orbital bristles incurved; costa not broken near hm; R1 not setulose; mesopleural bristle present; tibiae without preapical dorsal bristles; female with abdominal segment 7 enlarged, entirely sclerotised.

<==Agromyzidae
    |--Phytoliriomyza cyathea Spencer 1976 M03
    |--Phytobia Z02
    |--Ceradontha australis [=C. denticornis] M83
    |--Chromatomyia horticola B88
    |--Cerodonta dorsalis WT11
    |--Ophiomyia B88
    |    |--O. phaseoli B88
    |    `--O. solanicola B88
    |--Agromyza B88
    |    |--A. angulata G17
    |    |--A. demeijerei A71
    |    `--A. pseudoreptans M90
    |--Phytomyza B88
    |    |--P. atricornis B88
    |    |--P. clematidicola B88
    |    |--P. cytisi A71
    |    |--P. ilicicola WT11
    |    |--P. ilicis A71
    |    `--P. plantaginis M83
    `--Liriomyza B88
         |--L. brassicae B88
         |--L. chenopodii M83
         |--L. helichrysi B88
         |--L. huidobrensis MHG04
         |--L. sativae B88
         `--L. trifolii H91

*Type species of generic name indicated

REFERENCES

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

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

Colless, D. H., & D. K. McAlpine. 1991. Diptera. In: CSIRO. The Insects of Australia, pp. 717-786. Melbourne University Press.

[G17] Girault, A. A. 1917. Descriptiones hymenopterorum chalcidoidicarum variorum cum observationibus. III. Privately published (reprinted Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 102-111).

[H91] Hokkanen, H. M. T. 1991. Trap cropping in pest management. Annual Review of Entomology 36: 119-138.

[M83] Martin, N. A. 1983. Miscellaneous observations on a pasture fauna: An annotated species list. DSIR Entomology Division Report 3: 1-98.

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

[MHG04] Maynard, G. V., J. G. Hamilton & J. F. Grimshaw. 2004. Quarantine - phytosanitary, sanitary and incursion management: an Australian entomological perspective. Australian Journal of Entomology 43 (3): 318-328.

[M90] McAlpine, J. F. 1990. Insecta: Diptera adults. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 1211-1252. John Wiley & Sones: New York.

[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.

[Z02] Zherikhin, V. V. 2002. Insect trace fossils. In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 303-324. Kluwer Academic Publishers: Dordrecht.

Drosophila (Drosophila)

Drosophila virilis, photographed by Darren J. Obbard.


Belongs within: Drosophilidae.

Drosophila subgenus Drosophila is one of the major recognised subdivisions of the large genus Drosophila. Phylogenetic analysis suggests that some taxa recognised as separate genera are nested within this subgenus (Robe et al. 2005).

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 (Drosophila)
    |--+--+--D. (D.) nannoptera RV05
    |  |  `--D. (D.) pallidipennis RV05
    |  |       |--D. p. pallidipennis D51
    |  |       `--D. p. centralis D51
    |  `--+--+--D. (D.) robusta RV05
    |     |  `--D. (D.) virilis RV05
    |     `--+--+--+--D. (D.) annulimana RV05
    |        |  |  `--D. (D.) aracatacas RV05
    |        |  `--+--D. (D.) cestri RV05
    |        |     |--D. (D.) flavopilosa RV05
    |        |     `--D. (D.) incompta RV05
    |        `--+--D. (D.) canalinea RV05
    |           `--+--+--D. (D.) camargoi RV05
    |              |  `--D. (D.) dreyfusi RV05
    |              `--+--+--D. (D.) mercatorum RV05
    |                 |  `--+--+--D. (D.) eohydei RV05
    |                 |     |  `--D. (D.) hydei RV05
    |                 |     `--+--D. (D.) buzzati RV05
    |                 |        `--+--D. (D.) borborema RV05
    |                 |           `--D. (D.) serido 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. pictiventris RV05
          `--+--D. (D.) immigrans Sturtevant 1921 RV05, SDF04
             `--+--+--D. (D.) phalerata RV05
                |  `--D. (D.) quinaria RV05
                `--+--+--*D. (D.) funebris (Fabricius 1787) SDF04, RV05, SDF04 [=Musca funebris SDF04]
                   |  `--+--D. (D.) putrida RV05
                   |     `--D. (D.) testacea RV05
                   `--+--D. (D.) nappae [incl. D. angustibucca] 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. (D.) griseolineata RV05
                               |        `--D. (D.) maculifrons RV05
                               `--+--D. (D.) mediostriata [incl. D. crocina] RV05
                                  `--+--D. (D.) ornatifrons RV05
                                     `--+--D. (D.) guaru RV05
                                        `--D. (D.) subbadia RV05

Drosophila (Drosophila) incertae sedis:
  D. (D.) aldrichi CE79
  D. (D.) americana CE79
    |--D. a. americana CE79
    `--D. a. texana CE79
  D. (D.) analspina Singh & Negi 1995 SDF04
  D. (D.) arizonensis CE79
  D. (D.) bageshwarensis Singh, Dash & Fartyal 2004 SDF04
  D. (D.) bifurca CE79
  D. (D.) bishti Singh & Negi 1995 SDF04
  D. (D.) elongata Singh, Dash & Fartyal 2004 SDF04
  D. (D.) ezoana CE79
  D. (D.) flavomontana CE79
  D. (D.) fulvimacula CE79
  D. (D.) guaramunu RV05
  D. (D.) guarani RV05
  D. (D.) hexaspina Singh, Dash & Fartyal 2004 SDF04
  D. (D.) khansuensis Singh, Dash & Fartyal 2004 SDF04
  D. (D.) kunzei CE79
  D. (D.) lacertosa Okada 1956 SDF04
  D. (D.) lacicola CE79
  D. (D.) littoralis CE79
  D. (D.) lummei CE79
  D. (D.) medioimpressa RV05
  D. (D.) melanopalpa CE79
  D. (D.) meridiana CE79
  D. (D.) micromelanica CE79
  D. (D.) mohavensis CE79
  D. (D.) montana CE79
    |--D. m. montana CE79
    `--D. m. ovivororum CE79
  D. (D.) mulleri CE79
  D. (D.) nainitalensis Singh & Bhatt 1988 SDF04
  D. (D.) nasuta Lamb 1914 SDF04
  D. (D.) navajoa CE79
  D. (D.) neohydei CE79
  D. (D.) nigrohydei RV05
  D. (D.) novamexicana CE79
  D. (D.) pachea CE79
  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.) paranaensis CE79
  D. (D.) paunai Singh & Negi 1989 SDF04
  D. (D.) repleta Wollaston 1858 SDF04
  D. (D.) serrata Singh, Dash & Fartyal 2004 SDF04
  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
  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, 3rd ed. Columbia University Press: New York.

[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.