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Phthiriidae Usiidae Systropodidae

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

Contents

1 Overview 2 Morphology

2.1 Adult 2.2 Larva

3 Biology 4 Zoogeography 5 Species lists 6 Systematics 7 Genera 8 Gallery 9 See also 10 References 11 External links

Overview[edit] The Bombyliidae
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
Lapeirousia
species with very long, narrow floral tubes. Unlike butterflies, bee flies hold their proboscis straight, and cannot retract it. 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
Bombyliidae
are opportunistic and will attack a variety of hosts. The Bombyliidae
Bombyliidae
include at least 4,500 described species, and certainly thousands more remain to be described. However, most species do not often appear in abundance, and compared to other major groups of pollinators they are much less likely to visit flowering plants in urban parks or suburban gardens. As a result, this is arguably one of the most poorly known families of insects relative to its species richness.

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[edit] Adult[edit] 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 usually 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. In other species patches of flattened hairs occur that can act as silvery, gilded or coppertone reflecting mirrors; these perhaps serve as visual signals in conspecific mate/rival recognition, or perhaps imitate reflecting surface particles on bare soils with high content of materials like quartz, mica or pyrite.

Exoprosopa
Exoprosopa
caliptera in Great Sand Dunes National Park, Colorado, US - note the silvery mirror stripes formed by patches of specialized hairs modified into reflecting scales

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. 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.

A male of Hyperalonia
Hyperalonia
morio patrolling a patch of vegetation near the visitor center of Quebrada de las Higueritas in Lujan, San Luis, Argentina

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
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

Hoverflies of the family Syrphidae
Syrphidae
often mimic Hymenoptera as well, and some syrphid species are hard to tell apart from Bombyliidae
Bombyliidae
at first glance, especially for bee fly species that lack a long probocis or long, thin legs. Such bombyliids can still be distinguished in the field by anatomical features such as: - They usually have an evenly curved or sloping face (hoverflies often have prominent bulges of the facial cuticle and/or beak- to knob-like facial projections). - The wings lack a "false rear edge" and often have large dark areas with sharp boundaries, or complex patterns of spots (hoverfly wings are often clear or have smooth gradients of tinting, and their veins merge posteriorly into a "false edge" rather than reaching the wing's true rear edge). - The abdomen and thorax hardly ever have large glossy areas formed by exposed cuticle (hoverflies often have glossy cuticular body surfaces). Larva[edit] 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[edit]

Xenox tigrinus
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. As with hoverflies, bee flies are capable of sudden acceleration or deceleration, all but momentum-free high-speed changes of direction, superb control of position while hovering in mid-air, as well as a characteristically cautious approach of a possible feeding or landing site. Bombyliids are often recognizable by their stocky shapes, by their hovering behavior, and for the particular length of their mouthparts and/or legs as they lean forward into flowers. Unlike hoverflies, which settle on the flower as do bees and other pollinating insects, those bee fly species which have a long proboscis generally feed while continuing to hover in the air, rather like Sphingidae, or while touching the flower with their front legs to stabilize their position - without fully landing or ceasing oscillation of the wings. Species with shorter proboscis do land and walk on flower heads, however, and can be much harder to distinguish from hoverflies in the field. As noted, many bee fly species spend regular time intervals at rest on or near the ground, while hoverflies hardly ever do so. It can therefore be informative to watch feeding individuals and see whether or not they move down to ground level after a few minutes. Close observation is often easier with feeding individuals than with flies on the ground, as the latter are especially quick to take flight at the first sight of moving silhouettes or approaching shadows. Mating behavior has only been observed in a handful of species. It can vary from fairly generic swarming or unsolicited mid-air interception, as is common in many Diptera, to courtship behavior involving a context-specific flight pattern and wingbeat pitch of the male, with or without repeated proboscis contact between male and female.[3] Males often seek out smaller or larger clearings on the ground, presumably in vicinity of flowering plants or host nesting habitats that are likely attractive to females. They can return to their chosen perch or patch after every feeding bout or after pursuit of other insects flying over, or they can instead survey their chosen territory while hovering one or more meters above the bare patch. Gravid females seek out nesting habitats of hosts, and can spend many minutes inspecting for example entrances of smaller burrows in soil. In some species this behavior consists of hovering and repeated split-second foreleg touches of soil near the edge of the burrow's entrance, presumably to detect biochemical clues about the burrow's constructor such as identity, recency of visiting etc. If a burrow passes scrutiny then the bee fly may proceed to land and insert its posterior abdomen into the soil, laying one or more eggs at the edge or in close vicinity to it. In nine subfamilies including the more frequently observable Bombyliinae and Anthracinae, the females often do not land at all during host burrow inspections, and will proceed to release their eggs from midair by quick flicks of the abdomen while hovering over the burrow's entrance. This remarkable behavior has earned such species the colloquial name of Bomber flies, it can be seen in Roy Kleuker's online video clip in YouTube.[4] Female flies with this remarkable oviposition strategy typically have a ventral storage structure known as a sand chamber on the posterior end of the abdomen, which is filled with sand grains gathered before egg laying.[5][6] These sand grains are used to coat each egg just before their aerial release, which is assumed to improve the female's aim as well as the egg's survival chances by adding weight, slowing down egg dehydration, masking biochemical cues that could trigger host behavior such as nest cleaning or abandonment - or a combination of all three.

Play media

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
Vespoidea
and Apoidea, beetles, other flies, and moths. Larvae of some species including Villa sp. feed on ova of Orthoptera. Bombylius major
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
Thyridanthrax
are parasitic pupae of tsetse flies. Villa morio is parasitic on the beneficial ichneumonid species Banchus femoralis. The larvae of Dipalta
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
Lepidophora
lepidocera a Nearctic ecozone
Nearctic ecozone
species

Zoogeography[edit] 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[edit]

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

A 4mm long female of Lepidanthrax in Cuyama Valley, California, showing the proportionally shorter wings and relatively larger head occurring in many of the smaller species in the family

Systematics[edit] 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[7] Boris Borisovitsch Rohdendorf (1974) dealt with the family in a separate superfamily (Bombyliidea), linking it to the superfamily of Asilidea.[8] Currently the close correlation either positions the bee-flies within the superfamily Asiloidea
Asiloidea
sensu Rohdendorf (Asilidea)or they are included with the families separated by Rohdendorf in the superfamily of Asiloidea.

Asiloidea  

 N.N. 

 ? Scenopinidae
Scenopinidae
and Therevidae

 ? Mydidae
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)[9] finally moved the genus Hilarimorpha into their own family (Hilarimorphidae). Zaitzev (1991)[10] moved the genus Mythicomyia and several other minor genera in the family Mythicomyiidae, Yeates (1992, 1994)[11] shifted the entire subfamily of Proratinae, with the exception of Apystomyia, into the family of Scenopinidae
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.[12] Phylogenetic analysis of CAD and 28S rDNA gene sequences supports monophyly of only eight subfamilies out of fifteen included in the study, with the Bombyliinae resolving as a highly polyphyletic group.[13] 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:

Poecilanthrax
Poecilanthrax
apache in Sheldon National Antelope Refuge, Nevada, US

Macrocondyla chorista in a grassy border in San Luis province, Argentina, illustrating less common features for Bombyliidae
Bombyliidae
such as a slender abdomen and white patches on the wings

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[edit]

Poecilanthrax
Poecilanthrax
eremicus nectaring on California Buckwheat near the visitor center of Devil's Punchbowl, Pearblossom, California

Pantarbes capito sunning in a dry wash in San Bernardino Mountains, California

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
Bombylius
Linnaeus, 1758 Brachyanax
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
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
Exhyalanthrax
Becker, 1916 Exoprosopa
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
Hemipenthes
Loew, 1869 Heteralonia Rondani, 1863 Heterostylum Macquart, 1848 Heterotropus Loew, 1873 Hyperalonia
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
Lepidophora
Westwood, 1835 Ligyra
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
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
Poecilanthrax
Osten Sacken, 1886 Poecilognathus
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
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
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[edit]

Two species of unidentified beeflies from Coimbatore, Tamil Nadu, India.

A bombyliid fly visiting a flower.

Bee
Bee
fly in Hampshire, United Kingdom
United Kingdom
conservatory .

Bee
Bee
fly landing on a flower

Exoprosopa
Exoprosopa
sp. feeding

Lepidophora
Lepidophora
on Bidens
Bidens
laevis

See also[edit]

List of soldierflies and allies of Great Britain

References[edit]

^ Alan Weaving; Mike Picker; Griffiths, Charles Llewellyn (2003). Field Guide to Insects of South Africa. New Holland Publishers, Ltd. ISBN 1-86872-713-0.  ^ a b Hull, Frank Montgomery, Bee
Bee
flies of the world: the genera of the family Bombyliidae
Bombyliidae
Washington, Smithsonian Institution Press 1973 ISBN 0-87474-131-9. Downloadable from: https://archive.org/details/beefliesofworl2861973hull ^ https://www.researchgate.net/publication/288683339_The_courtship_behavior_of_the_bee_fly_Meomyia_vetusta_Walker_Diptera_Bombyliidae Ferguson, D.J. and Yeates, D.K. 2013. The courtship behavior of the bee fly Meomyia vetusta Walker (Diptera: Bombyliidae). The Australian Entomologist 40, 89-92. ^ https://www.youtube.com/watch?v=whRkZ1BS1dE ^ Yeates David K. "The evolutionary pattern of host use in the Bombyliidae
Bombyliidae
(Diptera): a diverse family of parasitoid flies". Biological Journal of the Linnean Society. 60: 149–185. doi:10.1111/j.1095-8312.1997.tb01490.x.  ^ zoolstud.sinica.edu.tw/Journals/48.2/141.pdf Boesi, R., Polidori, C. and Andrietti, F. 2009. Searching for the Right Target: Oviposition and Feeding Behavior in Bombylius
Bombylius
Bee
Bee
Flies (Diptera: Bombyliidae). Zoological Studies 48: 141-150. ^ 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. ^ Boris B. Rohdendorf, Brian Hocking, Harold Oldroyd, George E. Ball. The Historical Development of Diptera. University of Alberta, 1974: 75-77. ISBN 088864003X. ^ Webb D.W., 1981 Hilarimorphidae. in: McAlpine J.F. (Ed.), Manual of Nearctic Diptera. Agriculture Canada, Ottawa, pp. 603-605. ^ Zaitzev, V.F., 1991 On the phylogeny and systematics of the dipteran superfamily Bombylioidea (Diptera). Entomol. Obozr. 70 [1991] : 716–36. ^ Yeates D.M. (1992). "Towards a monophyletic Bombyliidae
Bombyliidae
(Diptera): the removal of the Proratinae (Diptera: Scenopinidae)". American Museum Novitates. 3051: 1–30.  ^ Yeates & Lambkin, The Tree of Life, op. cit.. ^ http://currents.plos.org/treeoflife/article/overcoming-the-effects-of-rogue-taxa-2cn7m3af919c4-1/ Trautwein, M.D., Wiegmann, B.M. and Yeates, D.K. Overcoming the effects of rogue taxa: Evolutionary relationships of the bee flies. PLOS Currents Tree of Life. 2011 May 5 Edition 1.

Bowden, J.,1980 Family
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
Palaearctic
Region. Greathead & Evenhuis (Greathead, D.J., & N.L. Evenhuis, 1997. Family
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
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
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
Bombyliidae
(Diptera: Asiloidea). Bulletin of the American Museum of Natural History ; no. 219, 191 pp.

External links[edit]

Wikimedia Commons has media related to Bombyliidae.

Wikispecies
Wikispecies
has information related to Bombyliidae

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

v t e

Extant Diptera families

Kingdom: Animalia Phylum: Arthropoda Class: Insecta Subclass: Pterygota Infraclass: Neoptera Superorder: Endopterygota

Suborder Nematocera

Axymyiomorpha

Axymyiidae

Culicomorpha

Culicoidea

Dixidae
Dixidae
(meniscus midges) Corethrellidae
Corethrellidae
(frog-biting midges) Chaoboridae
Chaoboridae
(phantom midges) Culicidae (mosquitoes)

Chironomoidea

Thaumaleidae
Thaumaleidae
(solitary midges) Simuliidae (black flies) Ceratopogonidae
Ceratopogonidae
(biting midges) Chironomidae
Chironomidae
(non-biting midges)

Blephariceromorpha

Blephariceridae
Blephariceridae
(net-winged midges) Deuterophlebiidae (mountain midges) Nymphomyiidae

Bibionomorpha

Bibionoidea

Bibionidae
Bibionidae
(march flies, lovebugs)

Anisopodoidea

Anisopodidae
Anisopodidae
(wood gnats)

Sciaroidea (fungus gnats)

Bolitophilidae Diadocidiidae Ditomyiidae Keroplatidae Mycetophilidae Sciaridae
Sciaridae
(dark-winged fungus gnats) Cecidomyiidae
Cecidomyiidae
(gall midges)

Psychodomorpha

Scatopsoidea

Canthyloscelidae Perissommatidae Scatopsidae
Scatopsidae
(minute black scavenger flies, or dung midges)

Psychodoidea

Psychodidae (moth flies)

Ptychopteromorpha

Ptychopteridae
Ptychopteridae
(phantom crane flies) Tanyderidae (primitive crane flies)

Tipulomorpha

Trichoceroidea

Trichoceridae
Trichoceridae
(winter crane flies)

Tipuloidea

Pediciidae
Pediciidae
(hairy-eyed craneflies) Tipulidae (crane flies)

Suborder Brachycera

Asilomorpha

Asiloidea

Apioceridae
Apioceridae
(flower-loving flies) Apsilocephalidae Apystomyiidae Asilidae
Asilidae
(robber flies) Bombyliidae
Bombyliidae
(bee flies) Evocoidae Hilarimorphidae (hilarimorphid flies) Mydidae
Mydidae
(mydas flies) Mythicomyiidae Scenopinidae
Scenopinidae
(window flies) Therevidae
Therevidae
(stiletto flies)

Empidoidea

Atelestidae Hybotidae
Hybotidae
(dance flies) Dolichopodidae
Dolichopodidae
(long-legged flies) Empididae
Empididae
(dagger flies, balloon flies)

Nemestrinoidea

Acroceridae
Acroceridae
(small-headed flies) Nemestrinidae
Nemestrinidae
(tangle-veined flies)

Muscomorpha

Aschiza

Platypezoidea

Phoridae
Phoridae
(scuttle flies, coffin flies, humpbacked flies) Opetiidae
Opetiidae
(flat-footed flies) Ironomyiidae (ironic flies) Lonchopteridae
Lonchopteridae
(spear-winged flies) Platypezidae
Platypezidae
(flat-footed flies)

Syrphoidea

Syrphidae
Syrphidae
(hoverflies) Pipunculidae
Pipunculidae
(big-headed flies)

Schizophora

Acalyptratae

Conopoidea

Conopidae
Conopidae
(thick-headed flies)

Tephritoidea

Pallopteridae
Pallopteridae
(flutter flies) Piophilidae
Piophilidae
(cheese flies) Platystomatidae
Platystomatidae
(signal flies) Pyrgotidae Richardiidae Tephritidae
Tephritidae
(peacock flies) Ulidiidae
Ulidiidae
(picture-winged flies)

Nerioidea

Cypselosomatidae Micropezidae
Micropezidae
(stilt-legged flies) Neriidae
Neriidae
(cactus flies, banana stalk flies)

Diopsoidea

Diopsidae (stalk-eyed flies) Gobryidae Megamerinidae Nothybidae Psilidae
Psilidae
(rust flies) Somatiidae Strongylophthalmyiidae Syringogastridae Tanypezidae

Sciomyzoidea

Coelopidae
Coelopidae
(kelp flies) Dryomyzidae Helosciomyzidae Ropalomeridae Huttoninidae Heterocheilidae Phaeomyiidae Sepsidae
Sepsidae
(black scavenger flies) Sciomyzidae
Sciomyzidae
(marsh flies)

Sphaeroceroidea

Chyromyidae Heleomyzidae Sphaeroceridae
Sphaeroceridae
(small dung flies) Nannodastiidae

Lauxanioidea

Celyphidae
Celyphidae
(beetle-backed flies) Chamaemyiidae
Chamaemyiidae
(aphid flies) Lauxaniidae

Opomyzoidea

Agromyzidae
Agromyzidae
(leaf miner flies) Anthomyzidae Asteiidae Aulacigastridae (sap flies) Clusiidae
Clusiidae
(lekking, or druid flies) Fergusoninidae Marginidae Neminidae Neurochaetidae (upside-down flies) Odiniidae Opomyzidae Periscelididae Teratomyzidae Xenasteiidae

Ephydroidea

Camillidae Curtonotidae
Curtonotidae
(quasimodo flies) Diastatidae
Diastatidae
(bog flies) Ephydridae
Ephydridae
(shore flies) Drosophilidae
Drosophilidae
(vinegar and fruit flies)

Carnoidea

Acartophthalmidae Australimyzidae Braulidae
Braulidae
(bee lice) Canacidae
Canacidae
(beach flies) Carnidae Chloropidae
Chloropidae
(frit flies) Cryptochaetidae Inbiomyiidae Milichiidae
Milichiidae
(freeloader flies)

Lonchaeoidea

Cryptochetidae Lonchaeidae
Lonchaeidae
(lance flies)

Calyptratae

Muscoidea

Anthomyiidae
Anthomyiidae
(cabbage flies) Fanniidae
Fanniidae
(little house flies) Muscidae
Muscidae
(house flies, stable flies) Scathophagidae
Scathophagidae
(dung flies)

Oestroidea

Calliphoridae
Calliphoridae
(blow-flies: bluebottles, greenbottles) Mystacinobiidae (New Zealand batfly) Oestridae (botflies) Rhinophoridae Sarcophagidae (flesh flies) Tachinidae
Tachinidae
(tachina flies)

Hippoboscoidea

Glossinidae (tsetse flies) Hippoboscidae
Hippoboscidae
(louse flies) Mormotomyiidae
Mormotomyiidae
(frightful hairy fly) Nycteribiidae
Nycteribiidae
(bat flies) Streblidae
Streblidae
(bat flies)

Stratiomyomorpha

Stratiomyoidea

Pantophthalmidae
Pantophthalmidae
(timber flies) Stratiomyidae
Stratiomyidae
(soldier flies) Xylomyidae
Xylomyidae
(wood soldier flies)

Tabanomorpha

Rhagionoidea

Austroleptidae Bolbomyiidae Rhagionidae
Rhagionidae
(snipe flies)

Tabanoidea

Athericidae
Athericidae
(water snipe flies) Oreoleptidae Pelecorhynchidae Tabanidae (horse and deer flies)

Vermileonomorpha

Vermileonoidea

Vermileonidae

Xylophagomorpha

Xylophagoidea

Xylophagidae
Xylophagidae
(awl flies)

List of families of Diptera

Taxon identifiers

Wd: Q676390 ADW: Bombyliidae BugGuide: 185 EoL: 497 EPPO: 1BOMYF Fauna Europaea: 10889 Fossilworks: 138883 GBIF: 7285 ITIS: 134153 NCBI: 50674

Authority control

LCCN: sh85015482 BNF:

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