Tethinidae

Dasyrhicnoessa sp., from PaDIL.

Belongs within: Ephydroidea.

The Tethinidae are a group of small to medium-sized flies usually associated with coastal marine or saline habitats. They may be distinguished from similar flies by the absence of well-developed genal setae, the presence of a single costal break and a complete cell cup in the wing, and the absence of a preapical dorsal seta on the tibiae (Mathis 2010).

Characters (from Mathis 2010): Small to medium-sized flies, body length 1.6–4 mm; frequently invested with pale yellowish to brown microtomentum. With one to four orbital and zero to four fronto-orbital setae; postocellar setae convergent or absent. Vibrissal seta various, if present usually weak. With one proepisternal seta; one proepimeral seta; anepisternum with one or more setae and some setulae posteriorly. Precoxal bridge present. Wing hyaline to infuscate or pale yellow; C with Sc break only; cell cup present but small; A₁ weakened apically, not reaching wing margin. Tibiae lacking preapical dorsal seta. Male terminalia with epandrium bearing two surstyli ventrally; anterior surstylus distinctive; posterior surstylus generally strongly setulose and lobe-like; phallapodeme long, slender; ejaculatory apodeme usually large; phallus usually elongate, ribbon-like, sinuous, subcylindrical, with a more or less dense ventral pubescence, often with several microscopic papillae. Female terminalia with two sclerotised spermathecae; cercus subcylindrical or compressed, sometimes bearing strong, spinelike setulae (pseudacanthophorites); tergites 7 and 7 mostly with characteristic pigmented areas; epiproct generally small, bearing pair of setulae dorsally on apical third; hypoproct large.

<==Tethinidae M10a
    |  i. s.: Listriomastax M10a
    |         Macrocanace V87
    |           |--M. australis V87
    |           `--M. littorea V87
    |--Suffomyia M10b [Zaleinae M10a]
    |--Horaismoptera V87 [Horaismopterinae M10a]
    |    `--H. hennigi V87
    |--Apetaenus [Apetaeninae] M10a
    |    |--A. littoreus M10a
    |    `--A. watsoni M10a
    |--Pelomyiinae M10a
    |    |--+--Pelomyiella mallochi M10a
    |    |  `--Neopelomyia rostrata M10a
    |    `--+--Pelomyia M10a
    |       |    |--P. coronata M10a
    |       |    `--P. occidentalis M10a
    |       `--Masoniella M10a
    |            |--M. advena M10a
    |            |--M. argentinaensis M10a
    |            `--M. richardsi M10a
    `--Tethininae M10a
         |--Afrotethina M04
         |--Plesiotethina Munari 2000 M04
         |    `--*P. australis Munari 2000 M04
         |--Thitena Munari 2004 M04
         |    `--*T. cadaverina Munari 2004 M04
         |--Sigaloethina Munari 2004 M04
         |    `--*S. phaia Munari 2004 M04
         |--Pseudorhicnoessa Malloch 1914 [incl. Macrotethina Malloch 1935] M04
         |    |--*P. spinipes Malloch 1914 [incl. Tethina (*Macrotethina) tibiseta Malloch 1935, P. tibiseta] M04
         |    `--P. rattii M10a
         |--Tethina Haliday in Curtis 1837 M04 [incl. Phycomyza V87]
         |    |  i. s.: T. albosetulosa M10a
         |    |         T. flavigenis M10a
         |    |         T. grisea M10a
         |    |         T. hirsuta Munari 2000 M04
         |    |         T. horripilans M10a
         |    |         T. nigriseta Malloch 1924 [=Rhicnoessa nigriseta] M04
         |    |         T. pallidiseta Malloch 1935 M04
         |    |         T. parvula M10a
         |    |--*T. (Tethina) illota (Haliday 1838) [=Opomyza (*Tethina) illota] M04
         |    `--T. (Rhicnoessa) C-T92
         |         |--T. (R.) alboguttata Strobl 1900 C-T92
         |         |--T. (R.) ochracea (Hendel 1913) C-T92 (see below for synonymy)
         |         `--T. (R.) simplex Collin 1966 C-T92
         `--Dasyrhicnoessa Hendel 1934 M04
              |  i. s.: D. serratula Malloch 1935 (n. d.) M04
              |--*D. fulva (Hendel 1913) [=Rhicnoessa fulva] M04
              |--D. aquila Munari 2002 M04
              |--D. atripes Munari 2004 M04
              |--D. bicolor Munari 2002 M04
              |--D. boninensis Sasakawa 1995 M04
              |--D. clandestina Munari 2002 M04
              |--D. ferruginea (Lamb 1914) [=Rhicnoessa ferruginea, Tethina ferruginea] M04
              |--D. fulvescens Malloch 1935 M04
              |--D. humilis Munari 2004 M04
              |--D. insularis (Aldrich 1931) (see below for synonymy) M04
              |--D. macalpinei Munari 2004 M04
              |--D. mathisi Munari 2002 M04
              |--D. ostentatrix Munari 2004 M04
              |--D. pallida Munari 2004 M04
              |--D. platypes Sasakawa 1986 M04
              |--D. sexseriata (Hendel 1913) (see below for synonymy) M04
              |--D. tripunctata Sasakawa 1974 [incl. D. phyllodes Sasakawa 1995] M04
              |--D. vockerothi Hardy & Delfinado 1980 [incl. D. occidentalis Munari 1986] M04
              |--D. yoshiyasui Sasakawa 1986 M04
              `--D. ciliata group M04
                   |--D. ciliata Munari 2004 M04
                   `--+--D. longisetosa Munari 2004 M04
                      `--D. priapus Munari 2004 M04

Dasyrhicnoessa insularis (Aldrich 1931) [=Tethina insularis, Rhicnoessa insularis; incl. Dasyrhicnoessa freidbergi Munari 1994, Tethina lasiophthalma Malloch 1933, D. lasiophthalma] M04

Dasyrhicnoessa sexseriata (Hendel 1913) [=Rhicnoessa sexstriata, Tethina sexstriata; incl. Dasyrhicnoessa asymbasia Sasakawa 1995] M04

Tethina (Rhicnoessa) ochracea (Hendel 1913) C-T92 [=Rhicnoessa ochracea M04; incl. T. canzonerii Munari 1981 M04, T. chilensis Malloch 1934 M04, Rhicnoessa texana Malloch 1913 M04, T. texana M04]

*Type species of generic name indicated

REFERENCES

[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.

[M10a] Mathis, W. N. 2010a. Tethinidae (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. 1109–1114. NRC Research Press: Ottawa.

[M10b] Mathis, W. N. 2010b. Canacidae (surf 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. 1115–1119. NRC Research Press: Ottawa.

[M04] Munari, L. 2004. Beach flies (Diptera: Tethinidae: Tethininae) from Australia and Papua New Guinea, with descriptions of two new genera and ten new species. Records of the Australian Museum 56: 29–56.

[V87] Vockeroth, J. R. 1987. Tethinidae. In: McAlpine, J. F. (ed.) Manual of Nearctic Diptera vol. 2 pp. 1073–1078. Research Branch, Agriculture Canada.

Last updated: 11 July 2021.

Drosophila

Drosophila busckii, from Wikimedia Commons.

Belongs within: Drosophilidae.
Contains: Drosophila (Drosophila), Drosophila (Sophophora).

Drosophila is a diverse, cosmopolitan genus of vinegar flies; the monophyly of the genus as generally recognised is currently under debate. Many species have become widely dispersed outside their native ranges as a result of human activity. One of these is southeast Asian native D. busckii, a small, spotted species that breeds in decaying matter (Grimaldi 2010). This species is distinguished by the presence of longitudinal stripes on the pleurae and the absence of preapical setae on the second and third tibiae.

Characters (from Grimaldi 2010): Arista plumose; facial carina large, often with flat surface; eyes pubescent; acrostichals in six or more rows; two pairs of dorsocentral setae; prescutellar setae absent; two katepisternal setae present of unequal lengths; wing tip rounded; crossveins distant.

<==Drosophila Fallén 1823 (see below for synonymy) SDF04
    |--D. (Dorsilopha) busckii RV05
    `--+--D. (Drosophila) RV05
       `--D. (Sophophora) RV05

Drosophila incertae sedis:
  D. acutilabella S00
  D. artificialis D37
  D. auraria KVV96
  D. berryi W87
  D. colorata M90
  D. earlei D37
  D. fenestrarum RD77
  D. grimshawi WT11
  D. guttifera M47
  D. heteroneura C96
  ‘*Tanygastrella’ hypopygialis Duda 1924 SDF04
  D. macrospina D51
    |--D. m. macrospina D51
    |--D. m. limpiensis D51
    `--D. m. ohioensis D51
  D. megagenys W91
  ‘*Adrosophila’ minuta Séguy 1938 SDF04
  ‘*Spuriostyloptera’ multipunctata Duda 1923 SDF04
  D. nasutoides LG91
  D. neorepleta D51
  D. nigrospiracula S85
  D. notha W91
  D. onychophora G10
  D. orientacea GJR97
  D. planitibia G91
  D. prosaltans D51
  D. quadrilineata de Meijere 1911 [=*Chaetodrosophilella quadrilineata] SDF04
  D. recens S00
  D. rubida W70
  D. silvestris VD94
  D. sturtevanti D51
  D. subarctica CE79
  D. suboccidentalis S00
  D. sulcata D37
  D. transversa D37
  D. uvarum B88
  D. vibrissina D37
  D. wheeleri G84
  D. (Paradrosophila Duda 1923) SDF04
    `--D. (*P.) pictipennis Kertész 1901 SDF04
  D. (Phalodoris) lebanonensis CE79
  D. (Pholadoris) T68
    |--D. (P.) lativittata T68
    |--D. (P.) pattersoni T68
    `--D. (P.) victoria T68
  D. (Spinulophila Duda 1923) SDF04
    `--D. (*S.) signata (Duda 1923) [=*Spinulophila signata] SDF04

Drosophila Fallén 1823 [incl. Adrosophila Séguy 1938, Chaetodrosophilella Duda 1923, Chaetodrosophila (l. c.), Spuriostyloptera Duda 1923, Tanygastrella Duda 1924] SDF04

*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).

[C96] Campbell, N. A. 1996. Biology 4^th ed. The Benjamin/Cummings Publishing Company, Inc.: Menlo Park (California).

[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.

[D37] Dobzhansky, T. 1937. Genetics and the Origin of Species. Columbia University Press: New York.

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

[GJR97] Giordano, R., J. J. Jackson & H. M. Robertson. 1997. The role of Wolbachia bacteria in reproductive incompatibilities and hybrid zones of Diabrotica beetles and Gryllus crickets. Proceedings of the National Academy of Sciences of the USA 94: 11439–11444.

[G84] Gosling, E. M. 1984. The systematic status of Mytilus galloprovincialis in western Europe: a review. Malacologia 25 (2): 551–568.

[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.

[G91] Groves, C. P. 1991. A Theory of Human and Primate Evolution revised ed. Clarendon Press: Oxford.

[KVV96] Kroon, A., R. L. Voenendaal & A. Veerman. 1996. Does the spider mite Tetranychus urticae Koch (Tetranychidae) respond to photoperiod in a quantitative manner? In: Mitchell, R., D. J. Horn, G. R. Needham & W. C. Welbourn (eds) Acarology IX vol. 1. Proceedings pp. 5–7. Ohio Biological Survey: Columbus (Ohio).

[LG91] Li, W.-H., & D. Graur. 1991. Fundamentals of Molecular Evolution. Sinauer: Sunderland (MA).

[M47] Mayr, E. 1947. Ecological factors in speciation. Evolution 1: 263–288.

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

[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.

[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.

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

[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.

[S85] Slatkin, M. 1985. Gene flow in natural populations. Annual Review of Ecology and Systematics 16: 393-430.

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

[VD94] Vogler, A. P., & R. Desalle. 1994. Diagnosing units of conservation management. Conservation Biology 8 (2): 354–363.

[W87] Wheeler, M. R. 1987. Drosophilidae. In: McAlpine, J. F. (ed.) Manual of Nearctic Diptera vol. 2 pp. 1011–1018. Research Branch, Agriculture Canada.

[W70] White, M. J. D. 1970. Cytogenetics. In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 72–82. Melbourne University Press.

[W91] Whitten, M. J. 1991. Australian insects in scientific research. In: CSIRO. The Insects of Australia: A textbook for students and research workers vol. 1 pp. 236–251. Melbourne University Press: Carlton (Victoria).

[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: 24 April 2022.

Diastatidae

Diastata fuscula, copyright Dick Belgers.

Belongs within: Ephydroidea.

The Diastatidae are a group of small, drosophilid-like flies found primarily among lush, low vegetation, often in relatively damp locations. The classification of this group has historically been disputed with Diastata alternatively placed within the Ephydridae, in which case the remaining genera are treated as the Campichoetidae (Mathis 2010).

Characters (from Mathis 2010): Adults small, greyish-brown, body length 2.5–4 mm; usually with patterned wings and weakly spinose costa. Postocellar setae convergent; both inner and outer verticals present; ocellar and vibrissal setae well developed; two well-developed fronto-orbital setae present, proclinate seta laterad of reclinate seta present. Antenna decumbent; arista elongate, macropubescent or short plumose. Anepisternum setose or bare; katepisternum bearing twelve setae, posterior seta stronger. C extending to M; C with either subcostal break only or both humeral and subcostal breaks; Sc incomplete, not reaching C, often fusing with R₁ distally; R₁ bare; both CuA₂ and A₁ present; cell cup closed, crossvein bm-cu present. Legs moderatly elongate, slender. Abdominal segments 1–5 unmodified; male with segments 6–8 reduced, although tergite 7 well developed, sternite 7 absent; abdominal spiracles (6 or 7) in membrane.

<==Diastatidae [Campichoetidae] M10
    |--Euthychaeta M10
    |--Pareuthychaeta M10
    |    |--P. electrica M87
    |    `--P. minuta M87
    |--Campichoeta M10
    |    |--C. basalis [incl. Thryptocheta tristis] M10
    |    |--C. latigena M10
    |    `--C. punctum M87
    `--Diastata M10
         |--D. argentina M10
         |--D. costaricensis M10
         |--D. dimidiata M10
         |--D. fuscula WT11
         `--D. vagans M10

*Type species of generic name indicated

REFERENCES

[M10] Mathis, W. N. 2010. Diastatidae (diastatid 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. 1207–1210. NRC Research Press: Ottawa.

[M87] McAlpine, J. F. 1987. Diastatidae. In: McAlpine, J. F. (ed.) Manual of Nearctic Diptera vol. 2 pp. 1019–1022. 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.

Last updated: 11 July 2021.

Drosophila (Sophophora)

Drosophila melanogaster, copyright Ryszard.

Belongs within: Drosophila.

Drosophila subgenus Sophophora is a genus of vinegar flies most diverse in central Africa and southeast Asia, with a number of species (included the widely studied D. melanogaster) introduced worldwide (Grimaldi 2010).

<==Drosophila (Sophophora) G10
    |--+--D. (S.) obscura RV05
    |  `--D. (S.) pseudoobscura RV05
    `--+--D. (S.) erecta RV05
       |--D. (S.) yakuba RV05
       `--+--D. (S.) melanogaster Meigen 1830 RV05, GE05
          `--+--D. (S.) mauritiana GE05
             |--D. (S.) sechellia H99, GE05
             `--D. (S.) simulans RV05

Drosophila (Sophophora) incertae sedis:
  D. (S.) affinis CE79
  D. (S.) algonquin CE79
  D. (S.) ambigua CE79
  D. (S.) ananassae CE79
  D. (S.) athabasca CE79
  D. (S.) azteca CE79
  D. (S.) bipectinata CE79
  D. (S.) birchii CE79
  D. (S.) bocqueti H99
  D. (S.) burlai H99
  D. (S.) capricorni CE79
  D. (S.) elegans H99
  D. (S.) emarginata T68
  D. (S.) equinoxialis CE79
  D. (S.) eugracilis H99
  D. (S.) fumipennis CE79
  D. (S.) guanche CE79
  D. (S.) malerkotliana G10
  D. (S.) miranda CE79
  D. (S.) narragansett CE79
  D. (S.) nebulosa CE79
  D. (S.) neocordata T68
  D. (S.) neoelliptica T68
  D. (S.) orena CE79
  D. (S.) paulistorum T68
  D. (S.) pavlovskiana CE79
  D. (S.) persimilis CE79
  D. (S.) populi T68
  D. (S.) pseudobocainensis T68
  D. (S.) saltans G10
  D. (S.) subobscura CE79
  D. (S.) sucinea T68
  D. (S.) simulans H99
  D. (S.) teissieri CE79
  D. (S.) tolteca CE79
  D. (S.) tropicalis CE79
  D. (S.) vulcana H99
  D. (S.) willistoni G10

*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.

[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.

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

[H99] Houck, M. A. 1999. Mites as potential horizontal gene transfer agents. In: Needham, G. R., R. Mitchell, D. J. Horn & W. C. Welbourn (eds) Acarology IX vol. 2. Symposia pp. 7–14. Ohio Biological Survey: Columbus (Ohio).

[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.

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

Gymnomyzinae

Placopsidella grandis, from the National Museum of Natural History.

Belongs within: Ephydroidea.

The Gymnomyzinae are a group of shore flies with pectinate aristae and small prescutellar acrostichal setae. Members include the cosmopolitan predatory genus Ochthera in which the fore legs are raptorial with the femur enlarged and the tibia bearing a large apical spur (Mathis 2010).

Characters (from Mathis 2010): Arista pectinate; reclinate fronto-orbital seta usually inserted slightly anterior to larger, proclinate fronto-orbital seta; median facial area and lower facial margin without setae; facial setae inserted in more or less vertical series, parallel with parafacial setae; prescutellar acrostichal setae small (about one-half to two-thirds length of posterior dorsocentral seta), inserted close together (distance between about one-half that between either prescutellar and posterior dorsocentral seta on the same side), behind or aligned with intra-alar seta.

<==Gymnomyzinae M10
    |--Ochthera Latreille 1802 L02 [Ochtherini M10]
    |    |--O. manicata [=Musca manicata] L02
    |    `--O. mantis [=Musca mantis] L02
    |--Hecamede [Hecamedini] M10
    |    |--H. albicans M10
    |    `--H. brasiliensis M10
    |--Atissini M10
    |    |--Ptilomyia mabelae M10
    |    `--Atissa M10
    |         |--A. atlantica M10
    |         `--A. pygmaea WMV87
    |--Gastropsini M10
    |    |--Beckeriella M10
    |    `--Gastrops M10
    |         |--G. nebulosus M10
    |         `--G. niger M10
    |--Discocerinini M10
    |    |--Discocerina obscurella M10
    |    |--Gymnoclasiopa M10
    |    |--Pectinifer aeneus M10
    |    |--Ditrichophora M10
    |    |--Polytrichophora agens M10
    |    |--Hydrochasma leucoproctum M10
    |    |--Lamproclasiopa M10
    |    `--Orasiopa mera M10
    |--Lipochaetini M10
    |    |--Glenanthe interior M10
    |    |--Paraglenanthe bahamensis M10, WMV87
    |    |--Lipochaeta M10
    |    |    |--L. ranica M10
    |    |    `--L. slossonae M10
    |    |--Diphuia M10
    |    |    |--D. anomala M10
    |    |    `--D. nitida M10
    |    `--Allotrichoma M10
    |         |--A. (Pseudohecamede) abdominale M10 [=Pseudohecamede abdominalis WMV87]
    |         `--A. simplex M10
    `--Gymnomyzini M10
         |--Cerometopum lacunosum M10
         |--Mosillus M10
         |    |--M. cellaris [=Micropeza (Mosillus) cellaris] G20
         |    |--M. stegmaieri M10
         |    `--M. tibialis M10
         |--Athyroglossa M10
         |    |--A. glabra M10
         |    |--A. glaphyropus M10
         |    `--A. granulosa M10
         `--Placopsidella M10
              |--P. cynocephala M10
              |--P. grandis M10
              `--P. marquesana M10

*Type species of generic name indicated

REFERENCES

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

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

[M10] Mathis, W. N. 2010. Ephydridae (shore 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. 1211–1233. NRC Research Press: Ottawa.

[WMV87] Wirth, W. W., W. N. Mathis & J. R. Vockeroth. 1987. Ephydridae. In: McAlpine, J. F. (ed.) Manual of Nearctic Diptera vol. 2 pp. 1027–1047. Research Branch, Agriculture Canada.

Last updated: 11 July 2021.

Ephydrinae

Alkali flies Ephydra hians, copyright Judy Gallagher.

Belongs within: Ephydroidea.

The Ephydrinae are a group of shore flies with lateroclinate fronto-orbital setae, and a large, cavernous subcranial cavity. The subcranial cavity is larger and the median facial area and lower facial margin are setose in members of the tribes Scatellini and Ephydrini, whereas Parydra and Dagini often have a smaller subcranial cavity and the median facial area and lower facial margin are bare (Mathis 2010).

Characters (from Mathis 2010): Fronto-orbital setae lateroclinate, usually conspicuous; median facial area and lower facial margin setose, the latter often with long setae (if not, costa short, extended to R₄₊₅); insertions of facial series of setae convergent above; subcranial cavity large and cavernous.

<==Ephydrinae M10
    |  i. s.: Setacera pacifica WMV87
    |         Coenia palustris WMV87, WT11
    |         Paracoenia WMV87
    |           |--P. (Paracoenia) bisetosa WMV87
    |           `--P. (Thiomyia) quatei WMV87
    |         Calocoenia platypelta WMV87
    |         Philotelma alaskense WMV87
    |--Parydra [Parydrini] M10
    |    |--P. (Parydra) aquila WMV87
    |    |--P. fossarum M10
    |    |--P. nigripes M10
    |    `--P. (Chaetoapnaea) parasocia WMV87
    |--Scatellini M10
    |    |--Scatophila cribrata M10
    |    |--Limnellia huachuca M10
    |    |--Haloscatella muria [=Lamproscatella muria] M10
    |    |--Neoscatella vittithorax WMV87, M83
    |    `--Scatella M10
    |         |--S. marinensis M10
    |         |--S. stagnalis P09
    |         `--S. triseta WMV87
    |--Dagini M10
    |    |--Dagus rostratus M10
    |    |--Physemops nemorosus M10
    |    |--Brachydeutera M10
    |    |    |--B. argentata M10
    |    |    |--B. longipes M10
    |    |    `--B. neotropica M10
    |    `--Diedrops M10
    |         |--D. hitchcocki M10
    |         |--D. roldanorum M10
    |         `--D. steineri M10
    `--Ephydrini M10
         |--Dimecoenia spinosa M10
         |--Cirrula M10
         |    |--C. austrina M10
         |    `--C. gigantea WMV87
         `--Ephydra M10
              |  i. s.: E. buresi WMV87
              |         E. californica RD77
              |         E. gracilis M10
              |         E. mexicana M10
              |         E. millibrae M10
              |         E. packardi M10
              |--E. (Ephydra) M10
              |    |--E. (E.) riparia M10
              |    `--E. (E.) subopaca WMV87
              |--E. (Halephydra) cinerea WMV87
              `--E. (Hydropyrus) hians M10

*Type species of generic name indicated

REFERENCES

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

[M10] Mathis, W. N. 2010. Ephydridae (shore 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. 1211–1233. NRC Research Press: Ottawa.

[P09] Pape, T. 2009. Economic importance of 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. 65–77. NRC Research Press: Ottawa.

[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.

[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.

[WMV87] Wirth, W. W., W. N. Mathis & J. R. Vockeroth. 1987. Ephydridae. In: McAlpine, J. F. (ed.) Manual of Nearctic Diptera vol. 2 pp. 1027–1047. Research Branch, Agriculture Canada.

Last updated: 10 July 2021.

Discomyzinae

Psilopa polita, copyright Christophe Quintin.

Belongs within: Ephydroidea.

The Discomyzinae are a group of shore flies with pectinate aristae and large prescutellar acrostichal bristles. They are divided between the Psilopini, in which the face is generally smooth and vein R₄₊₅ is dorsally bare, and Discomyzini, in which the face may be coarsely sculpted and vein R₄₊₅ generally bear two to four setulae dorsally near the base (Mathis 2010).

Characters (from Mathis 2010): Arista pectinate; reclinate fronto-orbital seta usually inserted behind larger proclinate fronto-orbital seta; median facial area and lower facial margin without setae; facial setae inserted in more or less vertical series, parallel with parafacial setae; prescutellar acrostichal setae large (subequal to posterior dorsocentral seta), inserted widely apart (distance between them subequal to that between either prescutellar and the posterior dorsocentral seta on the same side), usually in front of intra-alar seta.

<==Discomyzinae [Psilopinae] M10
    |  i. s.: Platygymnopa helicis WMV87
    |         Trimerinoides adfinis WMV87
    |         Trimerina madizans WMV87
    |         Pelignus salinus WMV87
    |         Nesopsilopa WMV87
    |         Diclasiopa lacteipennis WMV87
    |         Hecamedoides glaucellus WMV87
    |         Asmeringa lindsleyi WMV87
    |         Pelignellus subnudus WMV87
    |--Discomyzini M10
    |    |--Discomyza M10
    |    |--Clasiopella uncinata M10
    |    |--Mimapsilopa M10
    |    |    |--M. cressoni M10
    |    |    `--M. metatarsata M10
    |    |--Helaeomyia petrolei M10, WMV87
    |    |--Paratissa semilutea M10
    |    |--Guttipsilopa M10
    |    `--Rhysophora M10
    |         |--R. ardeoceras M10
    |         |--R. griseola M10
    |         `--R. robusta M10
    `--Psilopini M10
         |--Cressonomyia aciculata M10
         |--Peltopsilopa M10
         |--Ceropsilopa coquilletti M10
         |--Clanoneurum americanum M10, WMV87
         |--Leptopsilopa varipes M10
         `--Psilopa WT11
              |--P. flavida M10
              |--P. girschneri M10
              |--P. metallica M83
              |--P. petrolei WB37
              |--P. polita WT11
              `--P. pulchripes M10

*Type species of generic name indicated

REFERENCES

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

[M10] Mathis, W. N. 2010. Ephydridae (shore 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. 1211–1233. NRC Research Press: Ottawa.

[WB37] Walton, W. R., & F. C. Bishopp. 1937. Dr. L. O. Howard and the Entomological Society of Washington. Proceedings of the Entomological Society of Washington 39 (6): 121–132.

[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.

[WMV87] Wirth, W. W., W. N. Mathis & J. R. Vockeroth. 1987. Ephydridae. In: McAlpine, J. F. (ed.) Manual of Nearctic Diptera vol. 2 pp. 1027–1047. Research Branch, Agriculture Canada.

Last updated: 10 July 2021.

Ephydroidea

Cryptochetum buccatum, copyright Andre Burgers.

Belongs within: Schizophora.
Contains: Tethinidae, Drosophilidae, Diastatidae, Gymnomyzinae, Ephydrinae, Discomyzinae, Hydrelliinae.

The Ephydroidea are a clade of mostly small flies supported by molecular data, members of which possess a single pair of proclinate fronto-orbital bristles and precoxal bridges. Members include the Ephydridae, shore flies, a group of flies with a sclerotised, often protruding face commonly associated with aquatic or semiaquatic habitats (Mathis 2010). The Braulidae, beelice, are minute, wingless flies that live clinging to bees and subsist by inducing the host to regurgitate food (Brown & Thompson 2010). The Cryptochetidae are stout, metallic blue-black flies lacking an arista on the antenna that are endoparasitic as larvae on scale insects (McAlpine & Thompson 2010).

<==Ephydroidea [Drosophiloidea] WT11
    |  i. s.: Tethinidae M90
    |--+--+--Drosophilidae WT11
    |  |  `--+--Braulidae [Brauloidea] WT11
    |  |     |    |--Megabraula BT10
    |  |     |    `--Braula BT10
    |  |     |         |--B. coeca BT10
    |  |     |         |--B. hansruttneri BT10
    |  |     |         `--B. schmitzi BT10
    |  |     `--Cryptochetidae [Cryptochaetidae] WT11
    |  |          |--Librella demetrius MT10
    |  |          |--Phanerochaetum tuxeni MT10
    |  |          `--Cryptochetum MT10
    |  |               |  i. s.: C. iceryae MT10
    |  |               |         C. monophlebi CM91
    |  |               |         C. yokohama MT10
    |  |               `--C. (Cryptochetum) C-T92
    |  |                    |--C. (C.) buccatum Hendel 1933 C-T92
    |  |                    `--C. (C.) jorgepastori Cadahia 1984 C-T92
    |  `--+--Camillidae WT11
    |     |    |--Katacamilla cavernicola M87b
    |     |    |--Protocamilla succini M87b
    |     |    |--Afrocamilla W10
    |     |    |    |--A. arizonica W10
    |     |    |    `--A. bispinosa W10
    |     |    `--Camilla W10
    |     |         |--C. arnaudi W10
    |     |         `--C. glabra W10
    |     `--+--Diastatidae WT11
    |        `--Curtonotidae [Cyrtonotinae] WT11
    |             |--Cyrtona albomacula MK-SK10, M87a
    |             |--Axinota pictiventris MK-SK10, CM91
    |             `--Curtonotum MK-SK10
    |                  |--C. angustipennis O98
    |                  |--C. anus M87a
    |                  |--C. helvum MK-SK10
    |                  `--C. vulpinum MK-SK10
    `--Ephydridae [Parydrinae] WT11
         |  i. s.: Papuama H03
         |         Chlorichaeta tuberculosa A71
         |         Teichomyza fusca RD77
         |         Hyadinini M10
         |           |--Lytogaster granulosus M10
         |           `--Hyadina M10
         |                |--H. albovenosa M10
         |                |--H. binotata M10
         |                |--H. bulbosa M10
         |                `--H. pallipes M10
         |         Risa [Risidae] M10
         |         Gymnomyzinae M10
         |         Ilytheinae M10
         |           |--Philygrini M10
         |           |    |--Garifuna sinuata M10
         |           |    |--Philygria debilis M10
         |           |    `--Nostima approximata M10
         |           `--Ilytheini M10
         |                |--Zeros flavipes M10
         |                `--Ilythea M10
         |                     |--I. caniceps M10
         |                     `--I. spilota M10
         |         Pelina WMV87
         |           |--P. biloba WMV87
         |           `--P. truncatula WMV87
         |         Microlytogaster extera WMV87
         |         Callinapaea aldrichi WMV87
         |         Pseudohyadina longicornis WMV87
         |         Rhinonapaea metallica WMV87
         |         Eutaenionotum guttipennis WMV87
         |         Axysta WMV87
         |         Rhynchopsilopa O98
         |--Ephydrinae WT11
         `--+--Discomyzinae WT11
            `--Hydrelliinae WT11

*Type species of generic name indicated

REFERENCES

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

[BT10] Brown, B. V., & F. C. Thompson. 2010. Braulidae (beelice). 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. 1097–1099. 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).

[H03] Heads, M. 2003. Ericaceae in Malesia: vicariance biogeography, terrane tectonics and ecology. Telopea 10 (1): 311–449.

[MK-SK10] Marshall, S. A., A. H. Kirk-Spriggs & J. Klymko. 2010. Curtonotidae (hunchbacked 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. 1189–1192. NRC Research Press: Ottawa.

[M10] Mathis, W. N. 2010. Ephydridae (shore 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. 1211–1233. NRC Research Press: Ottawa.

[M87a] McAlpine, J. F. 1987a. Curtonotidae. In: McAlpine, J. F. (ed.) Manual of Nearctic Diptera vol. 2 pp. 1007–1010. Research Branch, Agriculture Canada.

[M87b] McAlpine, J. F. 1987b. Camillidae. In: McAlpine, J. F. (ed.) Manual of Nearctic Diptera vol. 2 pp. 1023–1025. 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.

[MT10] McAlpine, J. F., & F. C. Thompson. 2010. Cryptochetidae (cryptochetid 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. 1121–1123. NRC Research Press: Ottawa.

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

[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.

[W10] Wheeler, T. A. 2010. Camillidae (camillid 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. 1193–1195. 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.

[WMV87] Wirth, W. W., W. N. Mathis & J. R. Vockeroth. 1987. Ephydridae. In: McAlpine, J. F. (ed.) Manual of Nearctic Diptera vol. 2 pp. 1027–1047. Research Branch, Agriculture Canada.

Last updated: 11 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.