Bombyliidae

Bombyliidae
Bombylius major
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Diptera
Suborder:	Brachycera
Infraorder:	Asilomorpha
Superfamily:	Asiloidea
Family:	Bombyliidae Latreille, 1802
Subfamilies
Anthracinae Antoniinae Bombyliinae Crocidiinae Cythereinae Ecliminae Heterotropinae Lomatiinae Mariobezziinae Oligodraninae Oniromyiinae Phthiriinae Tomomyzinae Toxophorinae Usiinae Xenoprosopinae
Synonyms
Phthiriidae Usiidae Systropodidae

The Bombyliidae are a family of flies. Their common name is bee flies. Adults generally feed on nectar and pollen, some being important pollinators. Larvae generally are parasitoids of other insects.

Overview

The Bombyliidae are a large family of flies comprising hundreds of genera, but the lifecycles of most species are known poorly, or not at all. They range in size from very small (2 mm in length) to very large for flies (wingspan of some 40 mm).^[1][2] When at rest, many species hold their wings at a characteristic "swept back" angle. Adults generally feed on nectar and pollen, some being important pollinators, often with spectacularly long probosces adapted to plants such as Lapeirousia species with very long, narrow floral tubes. Unlike butterflies, bee flies hold their proboscises straight, and cannot retract them. In parts of East Anglia, locals refer to them as beewhals, thanks to their tusk-like appendages. Many Bombyliidae superficially resemble bees and accordingly the prevalent common name for a member of the family is bee fly.^[2] Possibly the resemblance is Batesian mimicry, affording the adults some protection from predators.

The larval stages are predators or parasitoids of the eggs and larvae of other insects. The adult females usually deposit eggs in the vicinity of possible hosts, quite often in the burrows of beetles or wasps/solitary bees. Although insect parasitoids usually are fairly host-specific, often highly host-specific, some Bombyliidae are opportunistic and will attack a variety of hosts.

While the Bombyliidae include a large number of species in great variety, most species do not often appear in abundance, and for its size this is one of the most poorly known families of insects. At least 4,500 species are described, and certainly thousands yet remain to be described.

Bombyliidae, India: Note the bright bands of coloured hair, the long and thin legs and upright posture, the "delta wings", the proboscis, and the forward-pointing antennae.

Morphology

Adult

Although the morphology of beeflies varies in detail, adults of most bee flies are characterized by some morphological details that make recognition easy. The dimensions of the body vary, depending on the species, from 1.0 mm to 2.5 cm. The form is often compact and the integument is covered with dense and abundant hair. The livery is usually inconspicuous and colours such as brown, blackish- grey, and light colors like white or yellow predominate. Many species are mimics of Hymenoptera Apoidea.

The head is round, with a convex face, often holoptic in males. The antennae are of the type aristate composed of three to six segments, with the third segment larger than the others; the stylus is absent (antenna of three segments) or is composed of one to three flagellomeres (antenna of four to six segments). The mouthparts are modified for sucking and adapted for feeding on flowers. The length varies considerably: for example, the Anthracinae have short mouthparts, with the labium terminating in a large fleshy labellum, in Bombyliinae; in Phthiriinae, the tube is considerably longer, and in Bombyliinae more than four times the length of the head.

The legs are long and thin and the front legs are sometimes smaller and more slender than the middle and rear legs. Typically, they are provided with bristles at the apex of the tibiae, without empodia and, sometimes, also without pulvilli . The wings are transparent, often hyaline or evenly colored or with bands. The alula are well developed and in the rest position the wings are kept open and horizontal in a V shape revealing the sides of the abdomen.

The abdomen is generally short and wide, subglobose-shaped, cylindrical, or conical, composed of six to eight apparent uriti apparent. The remaining urites are part of the structure of the external genitalia. The abdomen of the females often ends with spinous processes, used in ovideposition. In Anthracinae and Bombyliinae, a diverticulum is present in the eighth urite, in which the eggs are mixed with sand before being deposited.

The wing venation, although variable within the family, has some common characteristics that can be summarized basically in the particular morphology of the branches of the radial sector and the reduction of the forking of the media. The costa is spread over the entire margin and the subcosta is long, often ending on the distal half of the costal margin.The radius is almost always divided into four branches, with fusion of the branches R 2 and R 3, and is characterized by the sinuosity of the end portions of the branches of the radial sector. The venation presents a marked simplification compared to other Asiloidea and, in general, to other lower Brachycera. M 1 is always present and converges on the margin or, sometimes, of R 5. M 2 is present and reaches the margin, or is absent. M 3 is always absent and merged with M 4. The discal cell is usually present. The branch M 3 +4 is separated from the discal cell at the distal posterior vertex, so the mid-cubital connects directly to the posterior margin of the discal cell.The cubital and anal veins are complete and end separately on the margin or converge joining for a short distance Consequently, the cell cup may be open or closed.

• 
  Wing venation type 1 Bombylius
• 
  Wing venation type 2 Anthrax
• 
  Wing venation type 3 Usiinae

Larva

The larvae of most bee flies are of two types. Those of the first type are elongated and cylindrical in shape and have a metapneustic or amphipneustic tracheal system, provided with a pair of abdominal spiracles and, possibly, a thoracic pair. Those of the second type are stubby and eucephalic and have one pair of spiracles positioned in the abdomen.

Biology

Xenox tigrinus mating

Adults favour sunny conditions and dry, often sandy or rocky areas. They have powerful wings and are found typically in flight over flowers or resting on the bare ground exposed to the sun (watch video) They significantly contribute to cross pollination of plants, becoming the main pollinators of some plant species of desert environments. Unlike the majority of glyciphagous dipterans, the bee flies feed on pollen (from which they meet their protein requirements). A similar trophic behavior occurs among the hoverflies, another important family of Diptera pollinators. They are often recognizable by their stocky shapes, by their hovering behavior, and for the particular length of their mouthparts as they lean forward into flowers. Unlike hoverflies, which settle on the flower as do bees and other pollinating insects, the bee flies behave like Sphingidae and generally feed hovering in the air and using their front legs to stabilize their position.

Villa sp. gathering sand grains

Despite the high number of species of this family, the biology of juveniles of most species is poorly understood. The postembryonic development is of the type hypermetamorphic, with parasitoid or hyperparasitoid larvae. Exceptions are the larvae of Heterotropinae, whose biology is similar to that of other Asiloidea, with predatory larvae that do not undergo hypermetamorphosis. Hosts of bee flies belong to different orders of insects, but mostly are among the holometabolous orders. Among these are Hymenoptera, in particular the superfamilies of Vespoidea and Apoidea, beetles, other flies, and moths. Larvae of some species including Villa sp. feed on ova of Orthoptera. Bombylius major larvae are parasitic on solitary bees including Andrena. Anthrax anale is a parasite of tiger beetle larvae, and A. trifasciata is a parasite of the wall bee. Several African species of Villa and Thyridanthrax are parasitic pupae of tsetse flies. Villa morio is parasitic on the beneficial ichneumonid species Banchus femoralis. The larvae of Dipalta are parasitic on antlions.

The behavior of known forms is similar to that of the larvae of Nemestrinoidea: the first instar larva of is a planidium while the other stages have a parasitic habitus. The eggs are laid usually in a future host or at the nest where the host develops. The planidium enters the nest and undergoes changes before starting to feed.

Lepidophora lepidocera a Nearctic ecozone species

Zoogeography

The family is worldwide (Palearctic ecozone, Nearctic ecozone, Afrotropic ecozone, Neotropic ecozone, Australasian ecozone, Oceania ecozone, Indomalaya ecozone), but has the greatest biodiversity in tropical and subtropical arid climates. In Europe, 335 species are distributed among 53 genera.

Species lists

• West Palaearctic including Russia
• Australasian/Oceanian
• Nearctic
• Japan
• World list

Systematics

The systematics of bee flies are the most uncertain of any family of lower Brachycera. Willi Hennig(1973) placed the bee flies in the superfamily of Nemestrinoidea, on the basis of analogies in the behaviour of the larvae, positioning the superfamily in Tabanomorpha inside the infraorder Homoeodactyla^[3] Boris Borisovitsch Rohdendorf (1974) dealt with the family in a separate superfamily (Bombyliidea), linking it to the superfamily of Asilidea.^[4] Currently the close correlation either positions the bee-flies within the superfamily Asiloidea sensu Rohdendorf (Asilidea)or they are included with the families separated by Rohdendorf in the superfamily of Asiloidea. Currently, the close correlation either positions the bee-flies within the superfamily Asiloidea sensu Rohdendorf (Asilidea) or they are included with the families separated by Rohdendorf in the superfamily of Asiloidea.

Asiloidea	N.N.   ? Scenopinidae and Therevidae  ? Mydidae and Apioceridae  ? Asilidae  Bombyliidae

Clade showing relationship of Asiloidea

The internal systematic of bee-flies is uncertain. In the past, 31 subfamilies were well defined, but the family is thought to be polyphyletic (sensu lato). In the 1980s and '90s, the family has undergone several revisions: Webb (1981)^[5] finally moved the genus Hilarimorpha into their own family (Hilarimorphidae). Zaitzev (1991)^[6] moved the genus Mythicomyia and several other minor genera in the family Mythicomyiidae, Yeates (1992, 1994)^[7] shifted the entire subfamily of Proratinae, with the exception of Apystomyia, into the family of Scenopinidae and subsequently the genus Apystomyia into the family Hilarimorphidae. Nagatomi & Liu (1994) moved Apystomyia into a family of their own (Apystomyiidae. After these revisions, the bee flies sensu stricto have a greater morphological homogeneity, but the monophyly of the family still remains dubious and, consequently, the internal classification.^[8]

Overall, the family includes about 4700 described species, distributed among 230 genera. The internal arrangement varies according to the source, according to the different frameworks the authors attribute to tribes and subfamilies. To divide the family, often this scheme is used:

• Anthracinae
  • Anthracini
  • Aphoebantini
  • Exoprosopini
  • Plesiocerini
  • Villini
  • Xeramoebini
• Antoniinae
• Bombyliinae
  • Acrophthalmydini
  • Bombyliini
  • Conophorini
  • Dischistini
  • Eclimini
  • Lordotini
• Crocidiinae
• Cythereinae
• Lomatiinae
  • Comptosiini
  • Lomatiini
• Lordotinae
• Mariobezziinae
• Oligodraninae
• Oniromyiinae
• Phthiriinae
  • Phthiriini
  • Poecilognathini
• Tomomyzinae
• Toxophorinae
  • Gerontini
  • Systropodini
  • Toxophorini
• Usiinae
  • Apolysini
  • Usiini
• Xenoprosopinae

Genera

Acanthogeron Bezzi, 1925 Acreophthiria Evenhuis, 1986 Acreotrichus Macquart, 1840 Acrophthalmyda Bigot, 1858 Adelidea Macquart, 1840 Adelogenys Hesse, 1938 Aldrichia Coquillett, 1894 Alepidophora Cockerell, 1909 Aleucosia Edwards, 1934 Alomatia Cockerell, 1914 Amictites Hennig, 1966 Amictus Wiedemann, 1817 Amphicosmus Coquillett, 1891 Anastoechus Osten-Sacken, 1877 Anisotamia Macquart, 1840 Anthrax Scopoli, 1763 Antonia Loew, 1856 Antoniaustralia Becker, 1913 Apatomyza Wiedemann, 1820 Aphoebantus Loew, 1872 Apolysis Loew, 1860 Astrophanes Osten Sacken, 1877 Atrichochira Hesse, 1956 Australiphthiria Evenhuis, 1986 Australoechus Greathead, 1995 Balaana Lambkin & Yeates, 2003 Beckerellus Greathead, 1995 Bombomyia Greathead, 1995 Bombylella Greathead, 1995 Bombylisoma Rondani, 1856 Bombylius Linnaeus, 1758 Brachyanax Evenhuis, 1981 Brachydemia Hull, 1973 Bromoglycis Hull, 1971 Brychosoma Hull, 1973 Bryodemina Hull, 1973 Cacoplox Hull, 1970 Caecanthrax Greathead, 1981 Callostoma Macquart, 1840 Callynthrophora Schiner, 1868 Canariellum Strand, 1928 Chalcochiton Loew, 1844 Choristus Walker, 1852 Chrysanthrax Osten Sacken, 1886 Colossoptera Hull, 1973 Comptosia Macquart, 1840 Conomyza Hesse, 1956 Cononedys Hermann, 1907 Conophorina Becker, 1920 Conophorus Meigen, 1803 Corsomyza Wiedemann, 1820 Coryprosopa Hesse, 1956 Crocidium Loew, 1860 Cryomyia Hull, 1973 Cyananthrax Painter, 1959 Cyllenia Latreille, 1802 Cyrtomyia Bigot, 1892 Cytherea Fabricius, 1794 Cyx Evenhuis, 1993 Dasypalpus Macquart, 1840 Desmatomyia Williston, 1895 Desmatoneura Williston, 1895 Deusopora Hull, 1971 Diatropomma Bowden, 1962 Dicranoclista Bezzi, 1924 Diochanthrax Hall, 1975 Dipalta Osten Sacken, 1877 Diplocampta Schiner, 1868 Dischistus Loew, 1855 Docidomyia White, 1916 Doddosia Edwards, 1934 Dolichomyia Wiedemann, 1830 Doliogethys Hesse, 1938 Eclimus Loew, 1844 Edmundiella Becker, 1915 Efflatounia Bezzi, 1925 Enica Macquart, 1834 Epacmoides Hesse, 1956 Epacmus Osten Sacken, 1886 Eremyia Greathead, 1996

Eristalopsis Evenhuis, 1985 Eucessia Coquillett, 1886 Euchariomyia Bigot, 1888 Euprepina Hull, 1971 Eurycarenus Loew, 1860 Euryphthiria Evenhuis, 1986 Eusurbus Roberts, 1929 Exechohypopion Evenhuis, 1991 Exepacmus Coquillett, 1894 Exhyalanthrax Becker, 1916 Exoprosopa Macquart, 1840 Geminaria Coquillett, 1894 Geron Meigen, 1820 Glaesamictus Hennig, 1966 Gnumyia Bezzi, 1921 Gonarthrus Bezzi, 1921 Gyrocraspedum Becker, 1913 Hallidia Hull, 1970 Hemipenthes Loew, 1869 Heteralonia Rondani, 1863 Heterostylum Macquart, 1848 Heterotropus Loew, 1873 Hyperalonia Rondani, 1863 Hyperusia Bezzi, 1921 Inyo Hall & Evenhuis, 1987 Isocnemus Bezzi, 1924 Kapu Lambkin & Yeates, 2003 Karakumia Paramonov, 1927 Laminanthrax Greathead, 1967 Larrpana Lambkin & Yeates, 2003 Laurella Hull, 1971 Legnotomyia Bezzi, 1902 Lepidanthrax Osten Sacken, 1886 Lepidochlanus Hesse, 1938 Lepidophora Westwood, 1835 Ligyra Newman, 1841 Litorhina Bowden, 1975 Lomatia Meigen, 1822 Lordotus Loew, 1863 Macrocondyla Rondani, 1863 Mallophthiria Edwards, 1930 Mancia Coquillett, 1886 Mandella Evenhuis, 1983 Mariobezzia Becker, 1913 Marleyimyia Hesse, 1956 Marmosoma White, 1916 Megapalpus Macquart, 1834 Megaphthiria Hall, 1976 Melanderella Cockerell, 1909 Meomyia Evenhuis, 1983 Metacosmus Coquillett, 1891 Micomitra Bowden, 1964 Munjua Lambkin & Yeates, 2003 Muscatheres Evenhuis, 1986 Muwarna Lambkin & Yeates, 2003 Myonema Roberts, 1929 Neacreotrichus Cockerell, 1917 Nectaropota Philippi, 1865 Neobombylodes Evenhuis, 1978 Neodiplocampta Curran, 1934 Neodischistus Painter, 1933 Neosardus Roberts, 1929 Nomalonia Rondani, 1863 Nothoschistus Bowden, 1985 Notolomatia Greathead, 1998 Oestranthrax Bezzi, 1921 Oestrimyza Hull, 1973 Ogcodocera Macquart, 1840 Oligodranes Loew, 1844 Oncodosia Edwards, 1937 Oniromyia Bezzi, 1921 Othniomyia Hesse, 1938 Pachyanthrax François, 1964 Pachysystropus Cockerell, 1909 Palaeoamictus Meunier, 1916 Palaeogeron Meunier, 1915 Palintonus François, 1964 Palirika Lambkin & Yeates, 2003 Pantarbes Osten Sacken, 1877

Pantostomus Bezzi, 1921 Paracorsomyza Hennig, 1966 Paradiplocampta Hall, 1975 Parachistus Greathead, 1980 Paracosmus Osten Sacken, 1877 Parageron Paramonov, 1929 Paranthrax Bigot, 1876 Parasysteochus Hall, 1976 Paratoxophora Engel, 1936 Paravilla Painter, 1933 Parisus Walker, 1852 Perengueyimyia Bigot, 1886 Petrorossia Bezzi, 1908 Phthiria Meigen, 1803 Pilosia Hull, 1973 Pipunculopsis Bezzi, 1925 Platamomyia Brèthes, 1925 Plesiocera Macquart, 1840 Poecilanthrax Osten Sacken, 1886 Poecilognathus Jaennicke, 1867 Praecytherea Théobald, 1937 Prorachthes Loew, 1868 Prorostoma Hesse, 1956 Prothaplocnemis Bezzi, 1925 Pseudopenthes Roberts, 1928 Pteraulacodes Hesse, 1956 Pteraulax Bezzi, 1921 Pterobates Bezzi, 1921 Pusilla Paramonov, 1954 Pygocona Hull, 1973 Relictiphthiria Evenhuis, 1986 Rhynchanthrax Painter, 1933 Satyramoeba Sack, 1909 Semiramis Becker, 1913 Semistoechus Hall, 1976 Sericosoma Macquart, 1850 Sericothrix Hall, 1976 Sericusia Edwards, 1937 Sinaia Becker, 1916 Sisyromyia White, 1916 Sisyrophanus Karsch, 1886 Sosiomyia Bezzi, 1921 Sparnopolius Loew, 1855 Sphenoidoptera Williston, 1901 Spogostylum Macquart, 1840 Staurostichus Hull, 1973 Stomylomyia Bigot, 1888 Stonyx Osten Sacken, 1886 Synthesia Bezzi, 1921 Systoechus Loew, 1855 Systropus Wiedemann, 1820 Thevenetimyia Bigot, 1875 Thraxan Yeates & Lambkin, 1998 Thyridanthrax Osten Sacken, 1886 Tillyardomyia Tonnoir, 1927 Timiomyia Evenhuis, 1978 Tithonomyia Evenhuis, 1984 Tmemophlebia Evenhuis, 1986 Tomomyza Wiedemann, 1820 Tovlinius Zaitzev, 1979 Toxophora Meigen, 1803 Triplasius Loew, 1855 Triploechus Edwards, 1937 Turkmeniella Paramonov, 1940 Usia Latreille, 1802 Veribubo Evenhuis, 1978 Verrallites Cockerell, 1913 Villa Lioy, 1864 Villoestrus Paramonov, 1931 Walkeromyia Paramonov, 1934 Wurda Lambkin & Yeates, 2003 Xenoprosopa Hesse, 1956 Xenox Evenhuis, 1984 Xerachistus Greathead, 1995 Xeramoeba Hesse, 1956 Ylasoia Speiser, 1920 Zaclava Hull, 1973 Zinnomyia Hesse, 1955 Zyxmyia Bowden, 1960

Gallery

• 
  Two species of unidentified beeflies from Coimbatore, Tamil Nadu, India.
• 
  A bombyliid fly visiting a flower.
• 
  Bee fly in Hampshire, United Kingdom conservatory .
• 
  Bee fly landing on a flower
• 
  Exoprosopa sp. feeding
• 
  A bombyliid fly on Bidens laevis

See also

• List of soldierflies and allies of Great Britain

References

1. Alan Weaving; Mike Picker; Griffiths, Charles Llewellyn (2003). Field Guide to Insects of South Africa. New Holland Publishers, Ltd. ISBN 1-86872-713-0.
2. Hull, Frank Montgomery, Bee flies of the world: the genera of the family Bombyliidae Washington, Smithsonian Institution Press 1973 ISBN 0-87474-131-9. Downloadable from: https://archive.org/details/beefliesofworl2861973hull
3. Willi Hennig, 1973. Diptera (Zweiflüger). In J.G. Helmcke, D. Starck, H. Vermuth Hanbuch der Zoologie, Eine Naturgeschichte der Stämme des Tierreiches. IV. Band: Arthropoda - 2- Hälfte: Insecta. 2. Teil: Spezielles. Berlin, De Gruyter, 1973. pp. 1-337. ISBN 311004689X.
4. Boris B. Rohdendorf, Brian Hocking, Harold Oldroyd, George E. Ball. The Historical Development of Diptera. University of Alberta, 1974: 75-77. ISBN 088864003X.
5. Webb D.W., 1981 Hilarimorphidae. in: McAlpine J.F. (Ed.), Manual of Nearctic Diptera. Agriculture Canada, Ottawa, pp. 603-605.
6. Zaitzev, V.F. , 1991 On the phylogeny and systematics of the dipteran superfamily Bombylioidea (Diptera). Entomol. Obozr. 70 [1991] : 716–36.
7. Yeates, D.M., 1992 Towards a monophyletic Bombyliidae (Diptera): the removal of the Proratinae (Diptera: Scenopinidae). American Museum Novitates 3051: 1-30.
8. Yeates & Lambkin, The Tree of Life, op. cit..

• Bowden, J.,1980 Family Bombyliidae. pp. 381–430. In R.W. Crosskey (ed.), Catalogue of the Diptera of the Afrotropical Region, 1437 pp., London: British Museum (Natural History)
• Engel, E.O., 1932-1937. Bombyliidae. In: Die Fliegen der paläarktischen Region 4(3) ( Erwin Lindner, ed.): 1-619, pl. 1-15. E. Schweizerbart, Stuttgart.). Old and outdated, not easy to get and expensive but some of the only keys to taxa in the Palaearctic Region.
• Greathead & Evenhuis (Greathead, D.J., & N.L. Evenhuis, 1997. Family Bombyliidae. In: Contributions to a manual of Palaearctic Diptera Volume 2 (L. Papp & B. Darvas, eds.): 487-512. Science Herald, Budapest.) provide a key to the Palaearctic genera and (may) give references to available generic revisions.
• Evenhuis, N.L. 1991. Catalog of genus-group names of bee flies (Diptera: Bombyliidae) Bishop Museum Bulletin of Entomology 5: 1–105.
• Evenhuis, N.L. & Greathead, D.J. 1999. World catalog of bee flies (Diptera: Bombyliidae). Backhuys Publishers, Leiden, 756 pp. online
• Hull, F.M. 1973. Bee flies of the world. The genera of the family Bombyliidae.Washington (Smithsonian Institution Press) 687 pp. Keys subfamilies, genera (many generic placements superseded by Evenhuis & Greathead, 1999).
• Yeates, David K. 1994. The cladistics and classification of the Bombyliidae (Diptera: Asiloidea). Bulletin of the American Museum of Natural History ; no. 219, 191 pp.

External links

• Media related to Bombyliidae at Wikimedia Commons
• Image Gallery from Diptera.info
• Images from Encyclopaedia of Life
• BioLib Bombyliidae taxon tree
• Bombyliidae (Bee Flies) by David K. Yeates and Christine L. Lambkin in the Tree of Life web project. Consulted March 28, 2007.
• Wing venation
• Bombyliidae Fauna Europaea