Asilidae

Leptogaster cylindrica, copyright Fritz Geller-Grimm.

Belongs within: Asiloidea.
Contains: Dioctriinae, Stenopogoninae, Laphriinae, Dasypogoninae, Apocleinae, Asilinae.

The Asilidae, robber flies, are a group of predatory flies characterised by a moustache of bristles across the lower margin of the face. They are divided between several subfamilies including the Leptogastrinae which are slender-bodied robber flies lacking alulae or pulvilli. The Trigonomiminae are generally small flies with large 'goggle-eyes' and lacking strong bristles on the thorax. The cosmopolitan, primarily arboreal Ommatius has a long, plumose apical flagellomere on the antenna and closed cell r₁ in the wing (Fisher 2009).

Characters (from Wood 1981): Small (3 mm) to very large (over 50 mm) flies, averaging 9-15 mm in length, long and slender to short, robust, and bee-like. Head with compound eyes large, usually widely separated, equally so in both sexes; eye facets enlarged medially; eye extending dorsally above level of vertex, causing vertex to appear excavated when viewed anteriorly; ocellar tubercle prominent; median ocellus enlarged; vestiture of ocellar tubercle of fine pile, often mixed with bristles. Face extensively developed, often convexly protuberant (gibbous); facial protuberance, or gibbosity, with a characteristic group of stout anteroventrally directed hairs or bristles, the mystax, arranged as a cluster on lower part of face. Clypeus reduced, concave or flat, folded posteroventrally below lower facial margin. Antenna held erect; scape and pedicel each usually with long or short pile, often with stout bristles; first flagellomere elongate or oval, pubescent or tomentose but without pile or bristles, usually longer than either scape or pedicel. Bent apical or subapical stylus usually present on first flagellomere, typically two-segmented, occasionally one-segmented, apparently absent in Laphriinae. A minute spine, often in a pit, present at apex of antenna on either first or succeeding flagellomeres. Mouthparts adapted for stabbing and sucking prey. Mandible absent; stabbing function performed by needle-like hypopharynx that injects a paralyzing saliva. Proboscis elongate, heavily sclerotized, formed from ventromedial fusion of prementum and labella as continuous tube open at apex and dorsomedially, enclosing maxillae and hypopharynx; tip of labella pointed or rounded, usually surrounded with sensory hairs; ventral surface of labella and most of prementum pilose. Palpus usually slender or clavate, setose. Thorax with notopleural, supra-alar, and postalar bristles almost always present; dorsocentral bristles usually present. Scutellum usually pruinose but sometimes shiny, almost always with hairs or bristles along its margin especially at apex; katatergite almost always with a row or group of bristles. Legs raptorial, stout, frequently with numerous stour bristles; all tibiae with several stout bristles at apices. Wing venation with R always four-branched, with R₂₊₃ unbranched. Abdomen with dight segments usually present in both male and female, with remaining two segments incorporated into terminalia. Larva elongate, subcylindrical to somewhat dorsoventrally flattened, often tapering at each end; colour white to yellowish, sometimes with a few fine longitudinal streaks. Head capsule much narrower than prothorax, usually exerted, directed ventrally. Nine abdominal segments apparently present, with eighth and ninth partly fused; length of segments increasing toward seventh. Respiratory system functionally amphipneustic, although vestigial spiracles are present on first seven abdominal segments.

<==Asilidae F09
    |--+--+--Stenopogoninae BT04
    |  |  `--+--Laphriinae BT04
    |  |     `--Dasypogoninae BT04
    |  `--+--+--Apocleinae BT04
    |     |  `--+--Asilinae BT04
    |     |     `--Ommatius F09 [Ommatiinae BT04, Ommatiini]
    |     |          |--O. mackayi ZS10
    |     |          |--O. megacephalus F09
    |     |          `--O. tibialis W81
    |     `--Trigonomiminae [Damalinae, Damalini] F09
    |          |--Haplopogon W81
    |          |--Bromleyus flavidorsus W81
    |          |--Holcocephala BT04
    |          |    |--H. abdominalis BT04
    |          |    `--H. affinis F09
    |          `--Orrhodops F09
    |               |--O. americanus F09
    |               `--O. occidentalis F09
    `--Leptogastrinae [Leptogastridae, Leptogastrini] BT04
         |--Psilonyx annulatus F09 [=Leptogaster (Psilonyx) annulatus W81]
         |--Beameromyia bifida F09
         |--Eurhabdus zephyreus F09
         |--*Tipulogaster glabrata F09 [=Leptogaster (Tipulogaster) glabrata W81]
         |--Apachekolos F09
         |    |--A. crinitus F09 [=Leptogaster (Apachekolos) crinita W81]
         |    `--A. scapularis F09
         |--Schildia F09
         |    |--*S. microthorax F09
         |    |--S. guatemalae F09
         |    `--S. zonae F09
         |--Leptopteromyia F09
         |    |--L. americana F09
         |    |--L. argentinae Martin 1972 M72
         |    |--L. gracilis F09
         |    |--L. mexicanae F09
         |    `--L. peruae F09
         `--Leptogaster F09
              |--*L. cylindrica F09
              |--L. flavipes F09
              |--L. nigricornis K01
              |--L. spinitarsis F09
              `--L. titanus [=Tipulogaster titanus] F09

Asilidae incertae sedis:
  Araripogon axelrodi Grimaldi 1990 RJ93
  Dioctriinae F09
  Stizochimus salinator ZS10
  Brachyrhopala ZS10
  Deromyia discolor BM76
  Coleomyia setigera W81
  Itolia maculata W81
  Sintoria Hull 1962 W72
    |--*S. emeralda Hull 1962 W72
    |--S. cazieri Wilcox 1972 W72
    |--S. cyanea Wilcox 1972 W72
    |--S. lagunae Wilcox 1972 W72
    |--S. mojavae Wilcox 1972 W72
    |--S. pappi Wilcox 1972 W72
    `--S. rossi Wilcox 1972 W72
  Hodophylax W81
  Parataracticus W81
  Willistonina bilineata W81
  Nannocyrtopogon neoculatus W81
  Metapogon gilvipes W81
  Callinicus W81
    |--C. calcaneus L72
    `--C. pollenius W81
  Backomyia limpidipennis W81
  Pritchardomyia vespoides W81
  Lophonotus punctipennis K01
  Eutolmus sedakoffi K01
  Phinocus BW09
  Habropogon [Eremocneminae] J61
    |--H. appendiculatus J61
    |--H. exquisitus J61
    `--H. striatus J61
  Selidopogon [Acanthocneminae] J61
    |--S. crassus J61
    |--S. diadema J61
    `--S. octonotatus J61
  Codula limbipennis C91
  Chrysopogon crabroniformis C91
  Borapisma O98
  Cenochromyia O98
  Clariola O98
  Dichaetothyrea O98
  Epaphroditus O98
  Ophionomima O98
  Protoloewinella keilbachi Schumann 1984 P92

*Type species of generic name indicated

REFERENCES

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

[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.

[BT04] Bybee, S. M., S. D. Taylor, C. R. Nelson & M. F. Whiting. 2004. A phylogeny of robber flies (Diptera: Asilidae) at the subfamilial level: molecular evidence. Molecular Phylogenetics and Evolution 30: 789–797.

[C91] CSIRO. 1991. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 1. Melbourne University Press: Carlton (Victoria).

[F09] Fisher, E. M. 2009. Asilidae (robber flies, assassin flies, moscas cazadoras, moscas ladronas). 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. 585–632. NRC Research Press: Ottawa.

[J61] Janssens, E. 1961. Sur quelques asilides mediterranees. Boll. Mus. Civ. Venezia 14: 29–36.

[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.

[L72] Linsley, E. G. 1972. The robber fly Callinicus calcaneus (Loew) as a predator on Andrena omninigra Viereck (Diptera: Asilidae; Hymenoptera: Andrenidae). Pan-Pacific Entomologist 48 (2): 94–96.

[M72] Martin, C. H. 1972. A new species of Leptopteromyia (Diptera: Leptogastridae). Pan-Pacific Entomologist 48 (4): 270.

[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.

[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.

[W72] Wilcox, J. 1972. The genus Sintoria Hull (Diptera: Asilidae). Pan-Pacific Entomologist 48 (1): 51–58.

[W81] Wood, G. C. 1981. Asilidae. 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. 549–573. Research Branch, Agriculture Canada.

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

Last updated: 17 April 2022.

Asilinae

Asilus crabroniformis, copyright António A. Gonçalves.

Belongs within: Asilidae.

The Asilinae are a group of robber flies with a long, bare stylus on the antenna. They are the most diverse subfamily of robber flies in temperate regions of the world but are less diverse in the tropics (Fisher 2009). The larger number of species are assigned to the genus Asilus, including fossils going back to the Eocene (Wood 1981).

Characters (from Wood 1981, as Asilini): Apex of antenna with long slender bristle-like stylus; antennal stylus bare. Palpus one-segmented. Postmetacoxal area membranous. Wing with R₂₊₃ joining R₁ proximal to end of R₁, with cell r thus separated from wing margin; crossvein m-cu absent, and veins M₃ and CuA₁ joined together.

<==Asilinae [Asilini] F09
    |--Asilopsis W81
    |--Senoprosopis W81
    |--Promachella pilosa W81
    |--Promachina trapezoidalis W81
    |--Mallophorina frustra W81
    |--Proctacanthella cacopiloga W81
    |--Rhadiurgus variabilis W81
    |--Polacantha (Echinitropis) xanthocera F09
    |--Antipalus varipes F09
    |--Anarmostus iopterus F09
    |--Prolatiforceps anonyma F09
    |--Talamancasilus clavatus F09
    |--Threnia F09
    |--Pamponerus germanicus J61
    |--Acanthopleura brunnipes J61
    |--Colepia CM91
    |--Zosteria CM91
    |--Blepharotes CM91
    |--Neoaratus hercules CM70
    |--Heligmonevra O98
    |--Michotamia O98
    |--Cerdistus CM91 [incl. Neomochtherus W81]
    |    |--C. elegans J61
    |    `--‘Neomochtherus’ ochriventris J61
    |--Philonicus F09
    |    |--P. albiceps F09
    |    `--P. arizonensis BT04
    |--Wyliea F09
    |    |--W. chrysauges F09
    |    `--W. mydas F09
    |--Neoitamus W81
    |    |--N. cyanurus V09
    |    `--N. orphne W81
    |--Dysmachus J61
    |    |--D. dasynotus J61
    |    `--D. forcipula K01
    |--Machimus F09 [incl. Tolmerus W81]
    |    |--M. achterbergi Tomasovic 2004 F05
    |    |--M. gertschi F09
    |    |--M. paropus F09 [=Asilus (Machimus) paropus W81]
    |    |--M. perplexus J61
    |    |--M. rusticus J61
    |    `--M. soikai Janssens 1961 J61
    `--Asilus Linnaeus 1758 L58
         |--A. ater Linnaeus 1758 L58
         |--A. aureus L02
         |--A. auriannulatus W81
         |--A. barbarus Linnaeus 1758 L58
         |--A. bipartitus Macquart 1847 E12
         |--A. blantoni W81
         |--A. crabroniformis Linnaeus 1758 L58
         |--A. cyanurus R13
         |--A. flavopilosus Macquart 1847 E12
         |--A. forcipatus Linnaeus 1758 L58
         |--A. germanicus Linnaeus 1758 L58
         |--A. gibbosus Linnaeus 1758 L58
         |--A. gilvus Linnaeus 1758 L58
         |--A. hirtipes Macquart 1847 E12
         |--A. marginatus Linnaeus 1758 L58
         |--A. maurus Linnaeus 1758 L58
         |--A. morio Linnaeus 1758 L58
         |--A. oelandicus Linnaeus 1758 L58
         |--A. periscelis Macquart 1847 E12
         |--A. teutonus L02
         |--A. tingitanus Boisduval 1835 B35
         `--A. tipuloides Linnaeus 1758 L58

*Type species of generic name indicated

REFERENCES

[B35] Boisduval, J. B. 1835. Voyage de Découvertes de l’Astrolabe. Exécuté par ordre du Roi, pendant les années 1826–1827–1828–1829, sous le commandement de M. J. Dumont d'Urville. Faune entomologique de l'océan Pacifique, avec l'illustration des insectes nouveaux recueillis pendant le voyage vol. 2. Coléoptères et autres ordres. J. Tastu: Paris.

[BT04] Bybee, S. M., S. D. Taylor, C. R. Nelson & M. F. Whiting. 2004. A phylogeny of robber flies (Diptera: Asilidae) at the subfamilial level: molecular evidence. Molecular Phylogenetics and Evolution 30: 789–797.

[CM70] Colless, D. H., & D. K. McAlpine. 1970. Diptera (flies). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 656–740. Melbourne University Press.

[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.

[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.

[F09] Fisher, E. M. 2009. Asilidae (robber flies, assassin flies, moscas cazadoras, moscas ladronas). 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. 585–632. NRC Research Press: Ottawa.

[J61] Janssens, E. 1961. Sur quelques asilides mediterranees. Boll. Mus. Civ. Venezia 14: 29–36.

[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.

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

[R13] Reuter, O. M. 1913. Lebensgewohnheiten und Instinkte der Insekten bis zum Erwachen der sozialen Instinkte. R. Friedländer & Sohn: Berlin.

[V09] Verdcourt, B. (ed.) 2009. Additions to the wild fauna and flora of the Royal Botanic Gardens, Kew. XXVI. Miscellaneous records. Kew Bulletin 64 (1): 183–194.

[W81] Wood, G. C. 1981. Asilidae. 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. 549–573. Research Branch, Agriculture Canada.

Last updated: 3 January 2021.

Laphriinae

Andrenosoma fulvicaudum, copyright Tom Murray.

Belongs within: Asilidae.
Contains: Atomosini.

The Laphriinae are a group of robber flies lacking a conspicuous stylus on the antenna (Wood 1981). Members of the genus Laphria are particularly large, hairy robber flies that mimic bees in appearance. The Andrenosomatini are characterised by a dorsoventrally strongly flattened proboscis and flat, lamellate palpi, and females bear a cylindrical anal tube formed from the cerci (Fisher 2009).

Characters (from Wood 1981): Antenna blunt apically, without stylus, or with one or two abruptly tapered short microsegments. Palpus two-segmented. Wing with R₂₊₃ joining R₁ proximal to end of R₁, with cell r thus separated from wing margin; crossvein m-cu present; thus apex of cell bm closed by three veins. Male terminalia rotated 180° permanently.

<==Laphriinae F09
    |  i. s.: Smeryngolaphria [Dasytrichini] F09
    |           `--S. numitor F09
    |--+--Atomosini BT04
    |  `--Laphriini [Dasylechiini] BT04
    |       |--Maira BT04
    |       |--Dasylechia W81
    |       |--Orthogonis stygia W81
    |       |--Ichneumolaphria fascipennis F09
    |       |--*Phellopteron farri F09
    |       |--Rhopalogaster F09
    |       |    |--R. araujoi F09
    |       |    `--R. micronyx F09
    |       |--Lampria F09
    |       |    |--L. aurifex F09
    |       |    |--L. clavipes F09
    |       |    `--L. rubriventris W81
    |       `--Laphria BT04
    |            |--L. bomboides Macquart 1847 E12
    |            |--L. coerulea Meigen in Boisduval 1835 B35
    |            |--L. colorata Boisduval 1835 B35
    |            |--L. flava G20
    |            |--L. gilva K01
    |            |--L. hirta C91
    |            |--L. sadales W81
    |            `--L. thoracica CW09
    `--Andrenosomatini F09
         |--Cerotainiops F09
         |--Exodasyllis F09
         |--Neophoneus [Neophoneini] F09
         |    `--N. amandus F09
         |--Pilica F09
         |    |--*P. formidolosa F09
         |    `--P. erythrogaster F09
         |--Dasandra F09
         |    |--*D. ayalai F09
         |    |--D. hespenheidei F09
         |    `--D. zanutoi F09
         |--Dasyllis F09
         |    |--D. corallia F09
         |    |--D. funebris F09
         |    |--D. haemorrhoa F09
         |    `--D. phoenicogaster F09
         |--Eurycheilis F09
         |    |--E. bromleyi F09
         |    |--E. collini F09
         |    |--E. eufriesiae F09
         |    `--E. nigra F09
         `--Andrenosoma BT04
              |--*A. (Andrenosoma) atrum F09
              |--A. cinereum F09
              |--A. fulvicaudum F09
              |--A. (Pogonosoma) ridingsi W81
              |--A. sarcophagum F09
              `--A. xanthocnemum F09

*Type species of generic name indicated

REFERENCES

[B35] Boisduval, J. B. 1835. Voyage de Découvertes de l’Astrolabe. Exécuté par ordre du Roi, pendant les années 1826–1827–1828–1829, sous le commandement de M. J. Dumont d'Urville. Faune entomologique de l'océan Pacifique, avec l'illustration des insectes nouveaux recueillis pendant le voyage vol. 2. Coléoptères et autres ordres. J. Tastu: Paris.

[BT04] Bybee, S. M., S. D. Taylor, C. R. Nelson & M. F. Whiting. 2004. A phylogeny of robber flies (Diptera: Asilidae) at the subfamilial level: molecular evidence. Molecular Phylogenetics and Evolution 30: 789–797.

[C91] CSIRO. 1991. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 1. Melbourne University Press: Carlton (Victoria).

[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. NRC Research Press: Ottawa.

[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.

[F09] Fisher, E. M. 2009. Asilidae (robber flies, assassin flies, moscas cazadoras, moscas ladronas). 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. 585–632. NRC Research Press: Ottawa.

[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.

[W81] Wood, G. C. 1981. Asilidae. 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. 549–573. Research Branch, Agriculture Canada.

Last updated: 17 April 2022.

Stratiomyidae

Barred snout Nemotelus uliginosus, copyright David Gould.

Belongs within: Stratiomyomorpha.
Contains: Stratiomyini, Oxycerini, Sarginae, Clitellariinae, Pachygastrinae.

The Stratiomyidae, soldier flies, are an ecologically diverse group of often colourful flies. Larvae of the Stratiomyinae and Nemotelinae are aquatic whereas larvae of other subfamilies are terrestrial, mostly living in association with decaying plant or animal matter. As adults, the Chiromyzinae are markedly sexually dimorphic, with females much larger than males, and have atrophied mouthparts, an unarmed scutellum and an elongate abdomen. The Beridinae have a similarly elongate abdomen but retain functional mouthparts and usually have six or more marginal spines or tubercles on the scutellum (Woodley 2009).

Characters (from James 1981): Slender to quite robust flies, 2-18 mm long. Colour usually yellow, green, blue, or black, sometimes metallic, frequently with variegated color patterns particularly on abdomen. Bristles not developed, though body, especially thorax, sometimes densely pilose. Head usually as broad as thorax but sometimes broader, hemispherical to spherical. Occipital (postocular) orbit often well-developed, especially in female. Eyes ranging from bare to densely hairy, widely separated in female, more narrowly separated or contiguous in male, each eye in male often divided into an upper area comprising large facets and a lower area of small facets; ocelli present. Antenna inserted at or below middle of head; scape and pedicel unmodified except in respect to elongation; flagellum consisting of five to eight flagellomeres. Proboscis usually fleshy, sometimes atrophied, sometimes elongated, never adapted for sucking blood; palpus one- or two-segmented, not conspicuous. Thorax rectangular-oval in dorsal outline, sometimes elongated. Scutellum sometimes with one to three pairs of apical spines. Wing well-developed (rarely absent); C not reaching beyond apex of wing; stronger veins crowded anteriorly; branches of R always meeting wing margin well before wing apex; Rs usually arising from R₁ shortly before base of cell d or dm; R₄ short, often indistinct or wanting; veins toward posterior margin of wing weak, tending to evanesce. Legs simple, at most with certain segrnents or tarsomeres inflated or elongated, never densely pilose; fore coxa never unusually elongated; pulvilli and empodia pad-like; tibial spurs only rarely developed. Abdomen composed of five to eight visible segments, with the remaining segments telescoped into visible ones or forming terminalia; outline from dorsal view varying from almost round to elongate-oval, sometimes petiolate or spatulate; vestiture moderate, never dense. Male terminalia usually small, rather simple; surstylus simple and ventral; aedeagus two- or three-pronged; cercus simple. Terminal female segments forming at most a simple weakly developed ovipositor. Larva elongated, flattened, composed of head capsule and ll body segments; integument shagreened from deposit of calcium carbonate plates; head capsule well-developed, although broadly open posteroventrally, retracted more than half its length within thorax, but not functionally retractile; anterior exposed portion elongated and narrow except for ocular prominences, broadened posteriorly; mandible and maxilla more or less completely fused, operating vertically; maxillary palpus distinct; no free pharyngeal skeleton within head capsule; antenna usually located anterolaterally. Prothorax broader than head, with remainder of body either parallel-sided to near apex or attaining maximum width at abdominal segment 2; prominent lateral spiracles present on prothorax; lateral spiracles sometimes present on metathorax and on abdominal segments l-6 or 1-7, but small and probably not functional; posterior spiracles functional, located in transverse cleft on terminal abdominal segment; prolegs lacking, although ventral hooks sometimes present, especially on penultimate or antepenultimate segment, or both. Pupation occurring within last larval integument.

<==Stratiomyidae [Stratiomyiidae, Stratiomyriidae] W09
    |--+--Parhadrestia [Parhadrestiinae] GC99
    |  `--Cretaceogaster Teskey 1971 GC99
    |       `--*C. pygmaeus Teskey 1971 GC99
    `--+--Pachygastrinae GC99
       |--Chiromyzinae GC99
       |    |--Chiromyza vittata W09
       |    |--Boreoides subulatus CM91
       |    `--Inopus J81 [incl. Altermetoponia CM74]
       |         `--I. rubriceps (Macquart 1847) H82 [=Altermetoponia rubriceps CM70]
       `--Beridinae GC99
            |--Allognosta J81
            |--Actina viridis J81
            |--Arcuavena W09
            |--Heteracanthia ruficornis W09
            |--Beridella W09
            |--Berismyia W09
            |--Paraberismyia W09
            |--Oplachantha W09
            `--Beris Latreille 1802 L02
                 |--*B. sexdentata [=Stratiomys sexdentata] L02
                 |--B. annulifera J81
                 |--B. clavipes [=Xylophagus (Beris) clavipes] G20
                 `--B. fuscipes W09

Stratiomyidae incertae sedis:
  Barbiellinia Bezzi 1922 K03
    `--*B. hirta Bezzi 1922 K03
  Montsecia martinezdelclosi GE05
  Syndipnomyia auricincta C91
  Lysozus W09
  Pedinocera W09
  Exaireta spiniger CM91
  Neoexaireta spinigera CM70
  Evaza O98
  Saldubella O98
  Geosargus [Geosarginae] RD77
  Chrysochlorina [Chrysochlorininae] W09
    |--C. haterius W09
    `--C. quadrilineata J81
  Hermetiinae [Hermetiini] W09
    |--Chaetohermetia apicalis W09
    `--Hermetia W09
         |--H. albitarsis W09
         |--H. flavipes W09
         `--H. illucens [=Xylophagus (Hermetia) illucens] G20
  Stratiomyinae W09
    |--Stratiomyini W09
    |--Oxycerini W09
    `--Prosopochrysini [Myxosargini] W09
         |--Nothomyia parvicornis W09
         `--Myxosargus W09
              |--M. knowltoni W09
              `--M. nigricornis W09
  Raphiocerinae W09
    |--Raphiocera armata W09
    |--Analcoceroides longicornis W09
    |--Brachythrix dileucostigma W09
    |--Anisoscapus chlorovittatus W09
    |--Neanalcocerus hortulanus W09
    |--Phanerozus pampinus W09
    |--Heptozus hansoni W09
    |--Dolichothrix melanothorax W09
    `--Histiodroma caloptera [=Raphiocera caloptera] W09
  Nemotelinae W09
    |--Akronia frontosa J81
    |--Brachycara W09
    |    |--B. maculata W09
    |    `--B. slossonae W09
    `--Nemotelus W09
         |--N. albirostris W09
         |--N. canadensis W09
         |--N. (Camptopelta) glaber W09
         |--N. kansensis J81
         |--N. signatus K01
         |--N. trinotatus E66
         |--N. uliginosus L02
         |--N. varius K01
         `--N. zichyi Kertész 1901 K01
  Sarginae W09
  Clitellariinae W09
  Cacosis P92
  Hermetiella P92
  Eulalia evansi E66

*Type species of generic name indicated

REFERENCES

[CM70] Colless, D. H., & D. K. McAlpine. 1970. Diptera (flies). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 656–740. Melbourne University Press.

[CM74] Colless, D. H., & D. K. McAlpine. 1974. Diptera (flies). In: CSIRO. The Insects of Australia: A textbook for students and research workers. Supplement 1974 pp. 91–96. Melbourne University Press.

[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).

[C91] CSIRO. 1991. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 1. Melbourne University Press: Carlton (Victoria).

[E66] Evans, H. E. 1966. The Comparative Ethology and Evolution of the Sand Wasps. Harvard University Press: Cambridge (Massachusetts).

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

[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.

[H82] Helmore, D. W. 1982. Drawings of New Zealand insects. Bulletin of the Entomological Society of New Zealand 8: 1–52.

[J81] James, M. T. 1981. Stratiomyidae. 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. 497–511. Research Branch, Agriculture Canada.

[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.

[K03] Kury, A. B. 2003. Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Ibérica de Aracnología, volumen especial monográfico 1: 1–337.

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

[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.

[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.

[W09] Woodley, N. E. 2009. Stratiomyidae (soldier flies). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 1 pp. 521–549. NRC Research Press: Ottawa.

Last updated: 17 April 2022.

Therevidae

Phycus frommeri, copyright Spencer Entomological Collection.

Belongs within: Asiloidea.
Contains: Agapophytinae, Therevinae.

The Therevidae, stiletto flies, are a group of often fast-flying flies that are predatory as larvae but not as adults. Major subgroups include the Phycinae, found in Africa and the Americas (Irwin & Lyneborg 1981).

Characters (from Irwin & Lyneborg 1981): Slender to moderately thick-bodied flies; length, excluding antennae, 2.5-15 mm. Background color light yellow to black. Head hemispherical, not depressed at vertex. Eyes dichoptic in female, holoptic in most males, usually without hairs in both sexes. Frons of female wide, reaching ocellar tubercle; frons of male usually small, acutely to broadly triangular, pruinose to bare. Three prominent ocelli set at vertex or slightly anterior to vertex. Antenna three-segmented; scape usually setose, especially ventrally; pedicel usually with ring or two of short setae; first flagellomere without setae, or with setae usually confined to basal third; stylus comprising one or two flagellomeres and a terminal or subterminal bristle, set apically or subapically on basal flagellomere. Face often pruinose; lateral area hairy or not; gena hairy or not; occiput finely pruinose; postocular setae usually present though often weak. Palpus one- or two-segmented, usually hairy and not prominent, set beside proboscis in subcranial cavity; proboscis slightly longer than palpus, usually carried within subcranial cavity. Thorax with scutum varying from nearly square to elongate rectangular when viewed from above. Scutellum prominent, often without hairs, but almost always pruinose and often tomentose. Pleuron variously pilose; upper portion usually densely pruinose. Chaetotaxy with one pair postalar setae; one to six pairs notopleural setae, or more; one or two pairs supra-alar setae; usually from zero to two pairs dorsocentral setae, but three or more in a few species; zero to three, rarely four, pairs scutellar setae. Legs usually fairly long and slender; hindleg longer than other two. Fore coxa with none to several setae on anterior surface; all femora bare to heavily setose especially postventrally, often with long scale-like pile along dorsal surface; tibiae and tarsi setulose in definite longitudinal rows; fore tibia lacking setae anteroventrally. Five tarsomeres present; first tarsomere longest; claw with two pulvilli and seta-like central empodium, or with empodium absent. Wing venation generally uniform; R₁ setose or not; R₄ elongate, often sinuate; cell d elongate, with M₁, M₂ and M₃ arising from apex; CuA₁ not meeting posterior margin of cell d; crossvein m-cu present; cell cup closed behind; cell m₃ open or closed. Pterostigma usually well-developed. Calypter well-developed. Halter large and well-developed. Abdomen usually convex to flattened dorsally and tapering at apex, sometimes laterally compressed, always with eight well-developed pregenital segments. Fine silvery pollen often adorning abdomen. Tomentum and pile present or absent, usually longer and denser on male. Female terminalia with large conspicuous sternite 8; sternite 9, or genital fork, completely internal, with two lateral sclerites fused posteriorly to sclerotized bar containing membranous central sheath through which pass ducts of spermathecae and accessory glands; three usually unsclerotised spermathecae and two accessory glands present; tergite 8 generally unmodified; tergite 9 usually a single sclerite, generally with extended lateral margins fused to posterolateral margins of genital fork; tergite l0 always divided, often fused with tergite 9; digging-anchoring spines often present posterodorsally and anterolaterally; sternites l0 and ll (hypoproct, subanal plate) usually a single plate, usually heavily sclerotised; cerci disc-shaped. Male terminalia with sclerites forming sternite 8 and tergite 8 unmodified to narrowly constricted; tergite 9 (epandrium) acting as large covering for terminalia, often with surstyli or lateral surstylar-like lobes of various sizes and shapes; hypoproct and cerci borne apically; gonocoxites often extending as broad to narrow lobe well beyond insertion of gonostylus, usually with one or more other appendages; aedeagus consisting of a short to long variously twisted distiphallus, an elongate or vestigial dorsal aedeagal apodeme, a simple or forked ventral apodeme, and a variously shaped ejaculatory apodeme. Larva elongate, cylindrical, tapering anteriorly toward head and apex of abdomen; head small, with cranium well-sclerotised and divided into two portions; anterior portion exposed and strongly tapered anteriorly toward mouthparts; posterior portion a long apically spatulate metacephalic rod, flexibly articulated to posterior dorsal margin of anterior cranial portion and projecting into thorax; mouthparts comprising median slender tapered labrum flanked by curved pointed mandibles, and laciniae and palpi of maxillae, with elements of labium ventrally; labium consisting of large postmentum closing ventral part of cranial cavity, a fused pair of labial palpi usually bearing several pairs of setae, and prementum anteriorly; cranium exteriorly with several sensory cells both dorsally and ventrally, with antennae set in crescent-shaped cups on anterodorsal surface, with one pair of elongate dorsal setae and two pairs of elongate ventral setae, with heavily sclerotized postmentum ventrally, and dorsally with some weakly sclerotized portions called white areas; two pairs of heavy tentorial arms present internally. Thorax with pair of dorsolateral setae on each segment; anterior spiracle distinct, with two or three spiracular openings on specimens examined; abdominal segments l-8 secondarily constricted, giving appearance of two segments each, which with three thoracic segments appear as 19 segments in all; posterior spiracles on antipenultimate segment, apparently with eight spiracular openings; terminal segment ending in pair of retractable finger-like prolegs.

<==Therevidae
    |--+--+--Nanexila gracilis WY01
    |  |  `--+--Agapophytinae WY01
    |  |     `--Therevinae WY01
    |  `--+--Ectinorhynchus WY01
    |     `--Taenogerella WY01
    |          |--T. elizabethae WY01
    |          `--T. platina WY01
    `--Phycinae IL81
         |--Parapherocera wilcoxi IL81
         |--Schlingeria IL81
         |--Ataenogera brevicornis IL81, GW09
         |--Ruppellia vagabunda IL81
         |--Pherocera IL81
         |    |--P. albilateralis GW09
         |    `--P. flavipes IL81
         `--Phycus WY01
              |--P. frommeri GW09
              |--P. frontalis GW09
              |--P. kroberi WT11
              `--P. niger WY01

Therevidae incertae sedis:
  Rhagiophryne bianalis BLM02
  Taenogera luteola WY01, ZS10
  Neodialineura WY01
  Acrosathe IL81
    |--A. novella WT11
    `--A. pacifica IL81
  Megalinga GW09
    |--M. bolbocera GW09
    `--M. insignata IL81
  Nebritus pellucidus IL81
  Litolinga acuta IL81
  Pallicephala IL81
    |--P. pachyceras GW09
    `--P. willistoni IL81
  Rhagioforma GW09
    |--R. maculipennis IL81
    `--R. schmidti GW09
  Lyneborgia IL81
  Chromolepida IL81
  Eothereva simplex IL81
  Arenigena semitaria GW09
  Lysilinga IL81
    |--L. crassiseta GW09
    |--L. dolichophalla GW09
    `--L. pilifrons GW09
  Dialineura gorodkovi IL81
  Pandivirilia bussi IL81
  Ammonaios niveus IL81
  Tabuda IL81
  Tabudamima melanophleba IL81
  Spiriverpa candidata IL81
  Viriliricta IL81
  Dichoglena IL81
  Penniverpa GW09
    |--P. epidema GW09
    |--P. evani GW09
    |--P. festina IL81
    `--P. multisetosa GW09
  Xestomyzinae GW09
    |--Xestomyza IL81
    `--Henicomyia hubbardii GW09
  Microthereva GW09
  Peralia GW09
  Entesia GW09
  Melanothereva GW09
  Ptilotophallos GW09
  Apenniverpa GW09
  Pachyrrhiza GW09
  Distostylus GW09
  Spinalobus GW09
  Insulatitan GW09
  Incoxoverpa GW09
  Protothereva GW09
  Winthemmyia GW09
  Amplisegmentum GW09
  Notiothereva GW09
  Nigranitida GW09
  Elcaribe GW09
  Argolepida GW09
  Cliorismia GW09
  Lindneria dicosta GW09

*Type species of generic name indicated

REFERENCES

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

[GW09] Gaimari, S. D., & D. W. Webb. 2009. Therevidae (stiletto flies). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 1 pp. 633–647. NRC Research Press: Ottawa.

[IL81] Irwin, M. E., & L. Lyneborg. 1981. Therevidae. 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. 513–523. 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.

[WY01] Winterton, S. L., L. Yang, B. M. Wiegmann & D. K. Yeates. 2001. Phylogenetic revision of Agapophytinae subf. n. (Diptera: Therevidae) based on molecular and morphological evidence. Systematic Entomology 26: 173–211.

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

Last updated: 15 May 2018.

Tabanidae

Splayed deer fly Chrysops caecutiens, copyright Hectonichus.

Belongs within: Tabanomorpha.
Contains: Diachlorini, Tabanini, Pangoniini, Scionini, Bouvieromyiini.

The Tabanidae, horse flies, are a group of medium-sized to large flies, females of which commonly feed on vertebrate blood. Males and females of this family are readily distinguished by the eyes being holoptic in males but distinctly separated in females.

Characters (from Pechuman & Teskey 1981): Moderate to large flies, 6-30 mm long, without bristles. Head large; eyes large, often brightly patterned, holoptic in male, separated by frons in female. Antenna porrect and composed of scape, pedicel, and flagellum; flagellum usually consisting of a larger basal portion and terminal annulations usually numbering four to eight but sometimes as few as two. Palpus composed of two segments; basal palpal segment usually rather small; other palpal segment usually rather long and curved downward in female, often inflated near base. Proboscis stout and rigid; mandible and maxilla of female often styliform and adapted for piercing, sometimes absent in species that are not bloodsuckers. Thorax large; notopleural lobe prominent; scutellum without spines or macrotrichia. Legs rather stout; apical spurs present on mid tibia, absent on fore tibia, present or absent on hind tibia; tarsi each with three pads formed by pulvilliform empodium located between two pulvilli. Wing venation with as many as 11 veins reaching margin (Sc, four branches of R, three branches of M, CuA₁, CuA₂, A₁), C extending around wing, cells br, bm, and d conspicuous, and cell cup usually closed near wing margin; calypter large. Abdomen broad, with seven segments usually visible; terminalia of both sexes usually inconspicuous; male with gonocoxites fused with sternite 9, with gonostylus single or partially divided, aedeagus and associated pair of slender filamentous recurved aedeagal tines enclosed in a sheath apparently derived from parameres, tergite 9 entire or divided, tergite l0 absent, cercus flattened, round to oval or somewhat pointed; female usually with tergite l0 divided, with sternite 8 shield-shaped, and with cercus one-segmented. Larva fusiform or more or less expanded posteriorly or anteriorly, comprising 11 body segments and completely retractile head capsule; head capsule elongate, subcylindrical, tapering anteriorly, features including semitubular cranium, paired tentorial arms attached anteriorly to inner dorsal surfaces of cranium and appearing to be free posteriorly, three-segmented antennae, median down-curved labrum, mandibular-maxillary sclerites including maxillary palpi, and labium closing head capsule ventrally; mandibles each with central canal for excretion of paralytic enzyme, linked with subdorsal brushes of spines that are erected to anchor head within host when mandibles strike downward; body without projecting appendages on thoracic segments, with annular rings of small tubercles and crenulate frills or with prolegs on abdominal segments; respiratory system functionally metapneustic.

<==Tabanidae B09
    |  i. s.: Japenoides BLM02
    |         Mesopangonius RL09
    |         Pangonia A71
    |           |--P. lasiophtalma Boisduval 1835 B35
    |           `--P. longirostris A71
    |--Scepsidinae [Scepsidini] M54
    |    |--Adersia oestroides M54
    |    |--Scepsis nivalis M54
    |    |--Braunsiomyia M54
    |    `--Lesneus M54
    |--Tabaninae PT81
    |    |--Diachlorini PT81
    |    |--Tabanini PT81
    |    `--Haematopotini PT81
    |         |--Heptatoma M54
    |         |--Hippocentrum M54
    |         `--Haematopota PT81
    |              |--H. americana PT81
    |              `--H. pluvialis WT11
    |--Pangoniinae PT81
    |    |  i. s.: Palaepangonius GE05
    |    |         Eopangonius GE05
    |    |         Scaptiella aperta M54
    |    |--Pangoniini PT81
    |    |--Scionini PT81
    |    `--Philolichini M54
    |         |--Philoliche M54
    |         |--Ommatiosteres M54
    |         |--Metaphara M54
    |         |--Nuceria M54
    |         |--Stenophara M54
    |         |--Dorcaloemus compactus M54
    |         |--Phara M54
    |         |--Subpangonia M54
    |         `--Buplex M54
    |              |--B. albifacies M54
    |              `--B. suavus M54
    `--Chrysopsinae PT81
         |--Bouvieromyiini PT81
         |--Rhinomyzini M54
         |    |--Tabanocella denticornis M54
         |    |--Orgizomyia M54
         |    |--Thriambeutes M54
         |    |--Sphecodemyia M54
         |    |--Gastroxides ater M54
         |    |--Rhinomyza M54
         |    |--Thaumastocera M54
         |    `--Guyona M54
         `--Chrysopsini PT81
              |--Nemorius M54
              |--Melissomorpha indiana M54
              |--Neochrysops globosus PT81
              |--Silvius PT81
              |    |  i. s.: S. vituli M54
              |    |--S. (Assipala) ceras PT81
              |    |--S. (Griseosilvius) B09
              |    |    |--S. (G.) atitlanensis B09
              |    |    `--S. (G.) quadrivittatus PT81
              |    `--S. (Zeuximyia) philipi PT81
              `--Chrysops B09
                   |--C. caecutiens M54
                   |--C. cincticornis PT81
                   |--C. dimidiata A71
                   |--C. discalis A71
                   |--C. excitans PT81
                   |--C. furcatus PT81
                   |--C. (Liochrysops) hyalinus PT81
                   |--C. marmoratus K01
                   |--C. pikei PT81
                   |--C. scalaratus B09
                   `--C. silacea A71

*Type species of generic name indicated

REFERENCES

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

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

[B35] Boisduval, J. B. 1835. Voyage de Découvertes de l’Astrolabe. Exécuté par ordre du Roi, pendant les années 1826–1827–1828–1829, sous le commandement de M. J. Dumont d'Urville. Faune entomologique de l'océan Pacifique, avec l'illustration des insectes nouveaux recueillis pendant le voyage vol. 2. Coléoptères et autres ordres. J. Tastu: Paris.

[B09] Burger, J. F. 2009. Tabanidae (horse flies, deer flies, tabanos). 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. 495–507. NRC Research Press: Ottawa.

[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] 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.

[M54] Mackerras, I. M. 1954. The classification and distribution of Tabanidae (Diptera). I. General review. Australian Journal of Zoology 2 (3): 431–454.

[PT81] Pechuman, L. L., & H. J. Teskey. 1981. Tabanidae. 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. 463–478. Research Branch, Agriculture Canada.

[RL09] Ren, D., C. C. Labandeira, J. A. Santiago-Blay, A. Rasnitsyn, C. Shih, A. Bashkuev, M. A. V. Logan, C. L. Hotton & D. Dilcher. 2009. A probable pollination mode before angiosperms: Eurasian, long-proboscid scorpionflies. Science 326: 840–847.

[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: 13 May 2018.

Stratiomyini

Odontomyia sp., copyright Muhammad Mahdi Karim.

Belongs within: Stratiomyidae.

The Stratiomyini are a group of soldier flies with the antenna lacking an apical stylus, and vein A₁ in the wing straight. They may be divided between tribes Stratiomyini sensu stricto and Odontomyiini based on whether the antenna has five (Stratiomyini) or six (Odontomyiini) flagellomeres (James 1981) but this division is not recognised below, following Woodley (2009). Members of the Stratiomyini often resemble bees or wasps and may frequent flowers as adults (Woodley 2009).

Characters (from James 1981, as Stratiomyini + Odontomyiini): Antennal flagellum with five or six flagellomeres, not aristate. Scutellum with spines (sometimes minute). Wing with crossvein m-cu present; cell d present; A₁ not at all sinuous. Abdomen at least three-quarters as wide as long, usually patterned.

<==Stratiomyini [Odontomyiini] W09
    |--Psellidotus johnsoni [=Labostigmina johnsoni] W09
    |--Panamamyia silbergliedi W09
    |--Stratiomyella nana W09
    |--Chloromelas W09
    |--Zuerchermyia [=Zuercheria (preoc.)] W09
    |    `--Z. bequaerti W09
    |--Anoplodonta W09
    |    |--A. fratella W09
    |    `--A. nigrirostris J81
    |--Promeranisa W09
    |    |--P. nasuta W09
    |    `--P. varipes W09
    |--Hoplitimyia W09
    |    |--H. constans W09
    |    |--H. inbioensis W09
    |    `--H. subalba W09
    |--Hedriodiscus W09
    |    |--H. binotatus W09
    |    |--H. dorsalis W09
    |    |--H. trivittatus W09
    |    `--H. truquii W09
    |--Stratiomys W09
    |    |--S. badia M90
    |    |--S. barbata W09
    |    |--S. chamaeleon L02
    |    |--S. constricta W09
    |    |--S. convexa W09
    |    |--S. longicornis [=Hirtea longicornis] L02
    |    |--S. norma W09
    |    |--S. potamida I92
    |    `--S. strigata L02
    `--Odontomyia W09
         |--O. cincta W09
         |--O. discolorata W09
         |--O. (Odontomyia) furcata [=Stratiomys (O.) furcata] G20
         |--O. hydroleon K01
         |--O. (Catatasina) interrupta J81
         |--O. limbata Macquart 1847 E12
         |--O. ornata K01
         |--O. pilimana W09
         |--O. scutellata CM91
         |--O. (Odontomyiina) virgo J81
         `--O. viridula K01

*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).

[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.

[I92] Imes, R. 1992. The Practical Entomologist. Aurum Press: London.

[J81] James, M. T. 1981. Stratiomyidae. 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. 497–511. Research Branch, Agriculture Canada.

[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.

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

[W09] Woodley, N. E. 2009. Stratiomyidae (soldier flies). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 1 pp. 521–549. NRC Research Press: Ottawa.

Last updated: 1 January 2021.

Ceratopogonidae

Female Leptoconops, copyright Alice Abela.

Belongs within: Culicomorpha.
Contains: Forcipomyia, Culicoidini, Ceratopogonini, Sphaeromiini.

The Ceratopogonidae, biting midges, are a cosmopolitan group of small flies, a number of species of which have females that feed on vertebrate blood. Four subfamilies are recognised, the most divergent of which includes the genus Leptoconops which has eyes widely separated on top of the head (vs closely approximated in other subfamilies) and lacks the crossvein r-m in the wing (Downes & Wirth 1981).

Characters (from Downes & Wirth 1981): Small flies, 1–6 mm long, slender to moderately robust. Antenna with 11–13 flagellomeres, showing marked sexual dimorphism; female with five distal flagellomeres elongated and differing from proximal flagellomeres in many details; male with pedicel greatly enlarged and containing a specialised Johnston's organ, flagellomeres 1–8 usually bearing whorls of long setae giving characteristic plumose appearance. Proboscis about as long as head; palpus slightly longer. Female with biting and sucking mouthparts, consisting of a rigid labrum, blade-like mandibles and laciniae, and a hypopharynx, each typically with apical armature; females in non-biting species and all males with proboscis weaker and armature reduced. Palpus five-segmented; segments 1 and 2 sometimes poorly developed; segment 3 with sensory organ enclosed in a pit or on surface; segments 4 and 5 occasionally fused. Wings held horizontally at rest one above the other, narrower in male than in female; apex evenly rounded. Venation characterized by compact radial system lying closely behind costal margin and meeting margin before apex of wing, often near or even before the midpoint. R₁ and Rs typically short, with Rs branching into R₂₊₃, and R₄₊₅; R₂₊₃ running forward and rejoining R₁ to form cell r₁ (usually termed first radial cell); R₄₊₅ meeting C farther out to form cell r₂₊₃; one or both of these cells sometimes lost either by closure (approximation of upper and lower veins) or by becoming confluent through loss of R₂₊₃. Strong and characteristic crossvein r-m present. Posterior veins relatively weak; M₂ sometimes incomplete basally or absent; crossvein m-cu absent; cubital fork well-developed; CuP and A weak, not reaching wing margin. Legs moderately long; claws usually similar, average in size; male with small claws that are usually bifid at tips. Abdomen l0-segmented; spiracles present on segments 2–7; segment 1 somewhat reduced; tergites 1 and 2 sometimes fused. Sternite 8 in female with slightly bilobate hind margin extending below opening of spermathecal duct; segment 9 with small tergite fused laterally to narrow band-like sternite, sternite usually dividing in midline around spermathecal opening; segment l0 small; cercus rounded, on small dorsolateral basal plate. Segment 8 in male with normal tergite and sternite; tergite 9 usually enlarged and prolonged backward to cover other elements of terminalia; gonopod two-segmented, conspicuous; aedeagus extending back from sternite 9 and articulated laterally with base of gonocoxite, often triangular with sclerotized lateral arms or ventral plate; paired parameres usually present above aedeagus. Early stages usually found in moist or aquatic habitats; larva apneustic, often elongate in form, and a strong swimmer; head usually sclerotized; mandible strong, toothed, not apposed; pharyngeal apparatus conspicuous internally, with two strongly diverging arms and series of combs apparently serving to reduce and sort food; short collar between head and thorax; three thoracic and nine abdominal segments well-defined; no functional spiracles.

<==Ceratopogonidae [Heleidae]
    |--Dasyhelea BSG09 [Dasyheleinae DW81]
    |    |--D. antiqua S02
    |    |--D. cincta BSG09
    |    `--D. pseudoincisurata DW81
    |--Leptoconops BSG09 [Leptoconopinae DW81]
    |    |  i. s.: L. bequaerti BSG09
    |    |--L. (Leptoconops) torrens DW81
    |    |--L. (Brachyconops) californiensis DW81
    |    `--L. (Megaconops) floridensis DW81
    |--Forcipomyiinae DW81
    |    |--Forcipomyia DW81
    |    `--Atrichopogon DW81
    |         |--A. (Atrichopogon) levis DW81
    |         `--A. polydactylus DW81
    `--Ceratopogoninae DW81
         |--Culicoidini DW81
         |--Ceratopogonini DW81
         |--Sphaeromiini DW81
         |--Stenoxenini DW81
         |    |--Stenoxenus DW81
         |    |    |--S. coomani DW81
         |    |    `--S. johnsoni DW81
         |    `--Paryphoconus DW81
         |         |--P. angustipennis DW81
         |         `--P. sonorensis DW81
         |--Heteromyiini DW81
         |    |--Heteromyia fasciata DW81
         |    |--Neurohelea nigra DW81
         |    |--Neurobezzia granulosa DW81
         |    |--Clinohelea bimaculata DW81
         |    `--Pellucidomyia DW81
         |         |--P. ugandae BSG09
         |         `--P. wirthi DW81
         |--Palpomyiini DW81
         |    |--Pachyhelea pachymera DW81
         |    |--Bezzia DW81
         |    |    |--B. nobilis BSG09
         |    |    `--B. setulosa DW81
         |    |--Palpomyia BSG09
         |    |    |--P. lacustris BSG09
         |    |    |--P. lineata DW81
         |    |    `--P. plebeja BSG09
         |    `--Phaenobezzia DW81
         |         |--P. maya BSG09
         |         |--P. opaca DW81
         |         `--P. pistiae DW81
         `--Stilobezziini DW81
              |--Echinohelea lanei DW81
              |--Serromyia femorata DW81
              |--Parabezzia DW81
              |    |--P. bystraki BSG09
              |    `--P. petiolata DW81
              |--Monohelea DW81
              |    |--M. hieroglyphica DW81
              |    `--M. (Schizohelea) leucopeza DW81
              `--Stilobezzia DW81
                   |--S. antennalis DW81
                   |--S. (Eukraiohelea) elegantula DW81
                   `--S. (Neostilobezzia) lutea DW81

Ceratopogonidae incertae sedis:
  Austroconops GE05
    |--A. fossilis GE05
    `--A. macmillani GE05
  Fittkauhelea amazonica BSG09
  Cacaohelea youngi BSG09
  Baeohelea nana BSG09
  Leptohelea micronyx BSG09
  Nannohelea bourioni BSG09
  Schizonyxhelea forattinii BSG09
  Allohelea BSG09
    |--A. johannseni BSG09
    `--A. neotropica BSG09
  Parastilobezzia leei BSG09
  Downeshelea stonei BSG09
  Baeodasymyia dominicana BSG09
  Amerohelea BSG09
    |--A. frontispina BSG09
    `--A. galindoi BSG09
  Clastrieromyia dycei BSG09
  Spinellihelea BSG09
  Macrurohelea CM91
  Acanthohelea CM91
  Xenohelea CM91
  Dibezzia CM91
  Archiaustroconops alavensis Szadziewski & Arillo 1998 PR13
  Styloconops CM70

*Type species of generic name indicated

REFERENCES

[BSG09] Borkent, A., G. R. Spinelli & W. L. Grogan Jr. 2009. Ceratopogonidae (biting midges, purrujas). 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. 407–435. NRC Research Press: Ottawa.

[CM70] Colless, D. H., & D. K. McAlpine. 1970. Diptera (flies). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 656–740. Melbourne University Press.

[DW81] Downes, J. A., & W. W. Wirth. 1981. Ceratopogonidae. 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. 393–421. Research Branch, Agriculture Canada.

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

[PR13] Peris, D., & E. Ruzzier. 2013. A new tribe, new genus, and new species of Mordellidae (Coleoptera: Tenebrionoidea) from the Early Cretaceous amber of Spain. Cretaceous Research 45: 1–6.

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

Last updated: 7 May 2018.