Bee-fly
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The Bombyliidae are a
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
of
flies Flies are insects of the Order (biology), order Diptera, the name being derived from the Ancient Greek, Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwin ...
, commonly known as bee flies. Some are colloquially known as bomber flies. Adults generally feed on
nectar Nectar is a viscous, sugar-rich liquid produced by Plant, plants in glands called nectaries, either within the flowers with which it attracts pollination, pollinating animals, or by extrafloral nectaries, which provide a nutrient source to an ...
and
pollen Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...
, some being important
pollinator A pollinator is an animal that moves pollen from the male anther of a flower to the female carpel, stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains. Insects are ...
s. Larvae are mostly
parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
s of other insects.


Overview

The Bombyliidae are a large
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
of
flies Flies are insects of the Order (biology), order Diptera, the name being derived from the Ancient Greek, Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwin ...
comprising hundreds of
genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
, but the life cycles of most species are poorly known, or not at all. Their size varies between species ranging from 2 mm long to a 40 mm
wingspan The wingspan (or just span) of a bird or an airplane is the distance from one wingtip to the opposite wingtip. For example, the Boeing 777–200 has a wingspan of , and a wandering albatross (''Diomedea exulans'') caught in 1965 had a wingsp ...
making them some of the largest flies. When at rest, many species hold their wings at a characteristic "swept back" angle. Adults generally feed on
nectar Nectar is a viscous, sugar-rich liquid produced by Plant, plants in glands called nectaries, either within the flowers with which it attracts pollination, pollinating animals, or by extrafloral nectaries, which provide a nutrient source to an ...
and
pollen Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...
, some being important
pollinator A pollinator is an animal that moves pollen from the male anther of a flower to the female carpel, stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains. Insects are ...
s, often with spectacularly long proboscises adapted to plants such as ''
Lapeirousia ''Lapeirousia'' is a genus in the plant family Iridaceae. It is endemic to sub-Saharan Africa, about a third of the species occurring in fynbos. Origin of genus name The genus ''Lapeirousia'' was described by Pierre André Pourret in Mém. Aca ...
'' species with very long, narrow floral tubes. Unlike butterflies, bee flies hold their proboscis straight, and cannot retract it. Many Bombyliidae superficially resemble
bee Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfamil ...
s and accordingly the prevalent common name for a member of the family is bee fly. Possibly the resemblance is
Batesian mimicry Batesian mimicry is a form of mimicry where a harmless species has evolved to imitate the warning signals of a harmful species directed at a predator of them both. It is named after the English naturalist Henry Walter Bates, who worked on butt ...
, affording the adults some protection from
predator Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
s. The larval stages are predators or
parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
s of the eggs and
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
e of other insects. The adult females usually deposit eggs in the vicinity of possible hosts, quite often in the burrows of
beetle Beetles are insects that form the Taxonomic rank, order Coleoptera (), in the superorder Holometabola. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 40 ...
s,
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder ...
s, or solitary bees. Although insect parasitoids usually are fairly host-specific, often highly host-specific, some Bombyliidae are
opportunistic 300px, ''Opportunity Seized, Opportunity Missed'', engraving by Theodoor Galle, 1605 Opportunism is the practice of taking advantage of circumstances — with little regard for principles or with what the consequences are for others. Opport ...
and will attack a variety of hosts. The 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
insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
s relative to its species richness. The family has a patchy fossil record, with species being known from a handful of localities, the oldest known species are known from the Middle Cretaceous
Burmese amber Burmese amber, also known as Burmite or Kachin amber, is amber from the Hukawng Valley in northern Myanmar. The amber is dated to around 100 million years ago, during the latest Albian to earliest Cenomanian ages of the mid-Cretaceous period. Th ...
, around 99 million years old.


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 usually covered with dense and abundant hair. The coloration is usually inconspicuous and colours such as brown, blackish- grey, and light colors like white or yellow predominate. Many species are mimics of apoid Hymenoptera. In other species patches of flattened hairs occur that can act as silvery, gilded or copper-tone 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. The head is round, with a convex face, often
holoptic Holoptic refers to one of the ways in which the arthropod eye develops, particularly the eyes of various species of insects. Unlike dichoptic and cycloptic eyes, holoptic eyes meet along the median dorsal line of the head, in many species near ...
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 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 Pulvilli are soft, cushionlike pads on the feet of insects and other arthropods, such as the housefly and ixodid ticks. They are located at the base of the claws (#2 in the figure at right). The pulvilli function as an adhesive system. Their stic ...
. The wings are transparent, often hyaline or evenly colored or with bands. The
alula The alula , or bastard wing, (plural ''alulae'') is a small projection on the anterior edge of the wing of modern birds and a few non-avian dinosaurs. The word is Latin and means "winglet"; it is the diminutive of ''ala'', meaning "wing". The a ...
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 urites. 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. Wing shape between species varies, with noticeable difference in shaped and
venation Venation may refer to: * Venation (botany), the arrangement of veins in leaves * Wing venation, the arrangement of veins in insect wings See also * * Vernation Vernation or leafing is the formation of new leaves or fronds. In plant anatomy, it ...
patterns. In Bombylius, differences in the position of the veins such as CuA and A1 are found between species and also vary between the sexes. Females have slightly larger wings than males that aid in better stability during flight and enable more efficient movement, especially in feeding and laying eggs. The difference in size gives females the ability to have longer flight periods. 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. Image:Bombyliidae wing veins-1.svg, Wing venation type 1 ''Bombylius'' Image:Bombyliidae wing veins-2.svg, Wing venation type 2 ''Anthrax'' Image:Bombyliidae wing veins-3.svg, Wing venation type 3 Usiinae Hoverflies of the family Syrphidae often mimic Hymenoptera as well, and some syrphid species are hard to tell apart from Bombyliidae at first glance, especially for bee fly species that lack a long proboscis 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

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

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
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They significantly contribute to cross pollination of plants, becoming the main pollinators of some plant species of desert environments. Unlike the majority of nectar-feeding flies, the bee flies feed on pollen which provides essential proteins for their diet. A similar trophic behavior occurs among the
hoverflies Hoverflies, also called flower flies or syrphids, make up the insect family (biology), family Syrphidae. As their common name suggests, they are often seen Hover (behaviour), hovering or nectaring at flowers; the adults of many species feed main ...
, another important family of Diptera pollinators.


Pollination and Feeding Behavior

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 and Reproduction

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 Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advance ...
, 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. 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. This remarkable behavior has earned some species the nickname "bomber flies", and involves the female hovering steadily above the entrance to a
burrow file:Chipmunk-burrow (exits).jpg, An eastern chipmunk at the entrance of its burrow A burrow is a hole or tunnel excavated into the ground by an animal to construct a space suitable for habitation or temporary refuge, or as a byproduct of Animal lo ...
before releasing her eggs midair with a sharp flick of the abdomen. Just prior to release, each egg is coated in sand taken from a storage chamber at the tip of the abdomen, where grains had been collected earlier during ground visits. The coating has multiple methods to help increase the chance of successful
parasitism Parasitism is a close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The en ...
. For example, the coating adds weight for better aim, protects the egg from drying out, and masking its scent from the
host A host is a person responsible for guests at an event or for providing hospitality during it. Host may also refer to: Places * Host, Pennsylvania, a village in Berks County * Host Island, in the Wilhelm Archipelago, Antarctica People * ...
insect. 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 behavior has earned such species the colloquial name of Bomber flies. Females with this remarkable
oviposition The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typica ...
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. 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. 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 Hypermetamorphosis, or heteromorphosis, is a term used mainly in entomology; it refers to a class of variants of holometabolism, that is to say, complete insect metamorphosis. Hypermetamorphosis is exceptional in that some instars, usually larva ...
, 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 is a superfamily of wasps in the order Hymenoptera. Vespoidea includes wasps with a large variety of lifestyles including eusocial, social, and solitary habits, predators, scavengers, parasitoids, and some herbivores. Descriptio ...
and
Apoidea The superfamily Apoidea is a major group (of over 30 000 species) within the Hymenoptera, which includes two traditionally recognized lineages, the "sphecoid" wasps, and the bees. Molecular phylogeny demonstrates that the bees arose from ...
, beetles, other flies, and moths. Larvae of some species including ''
Villa A villa is a type of house that was originally an ancient Roman upper class country house that provided an escape from urban life. Since its origins in the Roman villa, the idea and function of a villa have evolved considerably. After the f ...
'' sp. feed on ova of
Orthoptera Orthoptera () is an order of insects that comprises the grasshoppers, locusts, and crickets, including closely related insects, such as the bush crickets or katydids and wētā. The order is subdivided into two suborders: Caelifera – gras ...
. ''
Bombylius major ''Bombylius major'' (commonly named the large bee-fly, the dark-edged bee-fly or the greater bee fly) is a parasitic bee mimic fly. ''B. major'' is the most common type of fly within the ''Bombylius'' genus. The fly derives its name from its clos ...
'' larvae are parasitic on solitary bees including ''
Andrena ''Andrena'' is a genus of bees in the family Andrenidae. With over 1,500 species, it is one of the largest genera of animals. It is a strongly Monophyly, monophyletic group that is difficult to split into more manageable divisions; currently, '' ...
''. ''
Anthrax anale Anthrax is an infection caused by the bacterium ''Bacillus anthracis'' or ''Bacillus cereus'' biovar ''anthracis''. Infection typically occurs by contact with the skin, inhalation, or intestinal absorption. Symptom onset occurs between one ...
'' is a parasite of
tiger beetle Tiger beetles are a family of beetles, Cicindelidae, known for their aggressive predatory habits and running speed. The fastest known species of tiger beetle, '' Rivacindela hudsoni'', can run at a speed of , or about 125 body lengths per second. ...
larvae, and ''A. trifasciata'' is a parasite of the wall bee. Several African species of ''
Villa A villa is a type of house that was originally an ancient Roman upper class country house that provided an escape from urban life. Since its origins in the Roman villa, the idea and function of a villa have evolved considerably. After the f ...
'' and ''
Thyridanthrax ''Thyridanthrax'' is a genus of bee flies in the family Bombyliidae. There are about 50 described species in the genus ''Thyridanthrax'', found across Eurasia Eurasia ( , ) is a continental area on Earth, comprising all of Europe and Asia ...
'' are parasitic pupae of
tsetse flies Tsetse ( , or ) (sometimes spelled tzetze; also known as tik-tik flies) are large, biting flies that inhabit much of tropical Africa. Tsetse flies include all the species in the genus ''Glossina'', which are placed in their own family, Glos ...
. '' Villa morio'' is parasitic on the beneficial
ichneumonid The Ichneumonidae, also known as ichneumon wasps, ichneumonid wasps, ichneumonids, or Darwin wasps, are a family of parasitoid wasps of the insect order Hymenoptera. They are one of the most diverse groups within the Hymenoptera with roughly 25,0 ...
species ''
Banchus femoralis ''Banchus'' is a genus of parasitoid wasps belonging to the family Ichneumonidae. The genus was first described by Fabricius in 1798. The species of this genus are found in Europe, North America, and Asia. Species: * ''Banchus dilatatorius'' ...
''. The larvae of ''
Dipalta ''Dipalta'' is a North American genus of bee flies in the family Bombyliidae. There are two described species of ''Dipalta''. The genus is closely related to ''Villa''. Description These medium-sized flies have conical faces, and have three ma ...
'' are parasitic on
antlion The antlions are a group of about 2,000 species of insect in the neuropteran family (biology), family Myrmeleontidae. They are known for the predation, predatory habits of their larvae, which mostly dig pits to trap passing ants or other prey. ...
s. The behavior of known forms is similar to that of the larvae of
Nemestrinoidea Nemestrinoidea is a small, monophyletic superfamily of flies, whose relationship to the other Brachycera is uncertain; they are sometimes grouped with the Tabanomorpha rather than the Asilomorpha. They are presently considered to be the sister ta ...
: the first instar larva of is a
planidium A planidium is a specialized form of insect larva seen in the first-instar of a few families of insects that have parasitoidal ways of life. They are usually flattened, highly sclerotized (hardened), and quite mobile. The function of the plani ...
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.


Zoogeography

The family is worldwide (
Palearctic realm The Palearctic or Palaearctic is a biogeographic realm of the Earth, the largest of eight. Confined almost entirely to the Eastern Hemisphere, it stretches across Europe and Asia, north of the foothills of the Himalayas, and North Africa. The ...
,
Nearctic realm The Nearctic realm is one of the eight biogeographic realms constituting the Earth's land surface. The Nearctic realm covers most of North America, including Greenland, Central Florida, and the highlands of Mexico. The parts of North America t ...
,
Afrotropical realm The Afrotropical realm is one of the Earth's eight biogeographic realms. It includes Sub-Saharan Africa, the southern Arabian Peninsula, the island of Madagascar, and the islands of the western Indian Ocean. It was formerly known as the Ethiopi ...
,
Neotropical realm The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropics, tropical Ecoregion#Terrestrial, terrestrial ecoregions of the Americas and the entire South American temperat ...
,
Australasian realm The Australasian realm is one of eight biogeographic realms that is coincident with, but not (by some definitions) the same as, the geographical region of Australasia. The realm includes Australia, the island of New Guinea (comprising Papua Ne ...
,
Oceanian realm The Oceanian realm is one of the eight biogeographic realms and is unique in not including any continental land mass. It has the smallest land area of any of the List of terrestrial ecoregions (WWF), WWF realms. This realm includes the islands ...
,
Indomalayan realm The Indomalayan realm is one of the eight biogeographic realms. It extends across most of South and Southeast Asia and into the southern parts of East Asia. Also called the Oriental realm by biogeographers, Indomalaya spreads all over the Ind ...
), but they are most diverse in tropical and subtropical arid climates.


Species lists


West Palaearctic including RussiaJapanWorld list


Systematics

The systematics of bee flies are the most uncertain of any family of lower Brachycera.
Willi Hennig Emil Hans Willi Hennig (20 April 1913 – 5 November 1976) was a German biologist and zoologist who is considered the founder of phylogenetic systematics, otherwise known as cladistics. In 1945 as a prisoner of war, Hennig began work on his th ...
(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
Boris Rohdendorf Boris Borisovich Rohdendorf (, 12 July 1904, Saint Petersburg – 21 November 1977, Moscow) was a Soviet entomologist and curator at the Zoological Museum at the Moscow University. He attained the position of head of the Laboratory of Arthropods, ...
(1974) dealt with the family in a separate superfamily (Bombyliidea), linking it to the superfamily of Asilidea. 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. 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) finally moved the genus ''Hilarimorpha'' into their own family (
Hilarimorphidae The Hilarimorphidae or hilarimorphid flies are a Family (biology), family of Fly, Diptera. They are of uncertain placement and may be related to the Acroceridae. Most species are Nearctic realm, Nearctic. Species Genus ''Cretahilarimorpha'' Mys ...
). Zaitzev (1991) moved the genus ''Mythicomyia'' and several other minor genera in the family
Mythicomyiidae Mythicomyiidae, commonly called mythicomyiids, are very tiny Fly, flies (0.5–5.0 mm) found throughout most parts of the world, especially desert and semi-desert regions, except the highest altitudes and latitudes. They are not as common in ...
, Yeates (1992, 1994) shifted the entire subfamily of Proratinae, with the exception of ''Apystomyia'', into the family of
Scenopinidae The Scenopinidae or window flies are a small (about 400 described species) family of flies (Diptera), distributed worldwide. In buildings, they are often taken at windows, hence the common name window flies. The two species with cosmopolitan dis ...
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.Yeates & Lambkin, The Tree of Life, op. cit. 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. Overall, the family includes about 4700 described species, distributed among 270 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:


Genera

*'' Acanthogeron'' Bezzi, 1925 *'' Acreophthiria'' Evenhuis, 1986 *'' Acreotrichus'' Macquart, 1840 *'' Acrophthalmyda''
Bigot Discrimination is the process of making unfair or prejudicial distinctions between people based on the groups, classes, or other categories to which they belong or are perceived to belong, such as race, gender, age, class, religion, or sexu ...
, 1858
*'' Adelidea'' Macquart, 1840 *'' Adelogenys'' Hesse, 1938 *''
Aldrichia ''Aldrichia'' is a North American genus of bee flies in the family Bombyliidae The Bombyliidae are a family of flies, commonly known as bee flies. Some are colloquially known as bomber flies. Adults generally feed on nectar and pollen, some be ...
'' 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 Anthrax is an infection caused by the bacterium '' Bacillus anthracis'' or ''Bacillus cereus'' biovar ''anthracis''. Infection typically occurs by contact with the skin, inhalation, or intestinal absorption. Symptom onset occurs between one ...
''
Scopoli Giovanni Antonio Scopoli (sometimes Latinized as Johannes Antonius Scopolius) (3 June 1723 – 8 May 1788) was an Italian physician and naturalist. His biographer Otto Guglia named him the "first anational European" and the "Linnaeus of the Au ...
, 1763
*''
Antonia Antonia may refer to: People * Antonia (name), including a list of people with the name * Antonia gens, a Roman family, any woman of the gens was named ''Antonia'' * Antônia (footballer) * Antônia Melo Entertainment * '' Antonia's Line'', o ...
'' Loew, 1856 *'' Antoniaustralia''
Becker Becker () is one of the German-language surnames, along with Bäcker and Baecker, that derive from the root, which refers to baking. The surname began as a name for a baker (and thus his family). In northern Germany, it can also derive from th ...
, 1913
*'' Apatomyza'' Wiedemann, 1820 *'' Aphoebantus'' Loew, 1872 *''
Apolysis Apolysis ( "discharge, lit. absolution") is the separation of the cuticle from the epidermis in arthropods and related groups (Ecdysozoa). Since the cuticle of these animals is also the skeletal support of the body and is inelastic, it is shed dur ...
'' 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 Rondani is an Italian surname. Notable people having this name include: * Camillo Rondani, (1808–1879), an Italian entomologist noted for his studies of Diptera * Giovanni Maria Francesco Rondani Giovanni Maria Francesco Rondani (15 July 1490 ...
, 1856
*'' Bombylius''
Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
,
1758 Events January–March * January 1 – Swedish biologist Carl Linnaeus (Carl von Linné) publishes in Stockholm the first volume (''Animalia'') of the 10th edition of ''Systema Naturae'', the starting point of modern zoologic ...
, 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 Walker or The Walker may refer to: People *Walker (given name) *Walker (surname) *Walker (Brazilian footballer) (born 1982), Brazilian footballer Places In the United States *Walker, Arizona, in Yavapai County *Walker, Mono County, California * ...
, 1852
*''
Chrysanthrax ''Chrysanthrax'' is a genus of bee flies in the family Bombyliidae, found in North and South America. Species *'' Chrysanthrax adumbrata'' ( Coquillett, 1887) *'' Chrysanthrax albicomus'' Tabet & Hall, 1987 *'' Chrysanthrax altus'' (Tucker, 19 ...
'' Osten Sacken, 1886 *'' Colossoptera'' Hull, 1973 *'' Comptosia'' Macquart, 1840 *'' Conomyza'' Hesse, 1956 *'' Cononedys'' Hermann, 1907 *'' Conophorina''
Becker Becker () is one of the German-language surnames, along with Bäcker and Baecker, that derive from the root, which refers to baking. The surname began as a name for a baker (and thus his family). In northern Germany, it can also derive from th ...
, 1920
*''
Conophorus ''Conophorus'' is a genus of bee flies in the family Bombyliidae. There are at least 16 described species in ''Conophorus'' in the United States, and 67 total worldwide. Species *'' C. atratulus'' ( Loew, 1872) *'' C. auratus'' Priddy, 1954 *' ...
'' Meigen, 1803 *'' Corsomyza'' Wiedemann, 1820 *'' Coryprosopa'' Hesse, 1956 *'' Crocidium'' Loew, 1860 *'' Cryomyia'' Hull, 1973 *'' Cyananthrax'' Painter, 1959 *'' Cyllenia'' Latreille, 1802 *'' Cyrtomyia''
Bigot Discrimination is the process of making unfair or prejudicial distinctions between people based on the groups, classes, or other categories to which they belong or are perceived to belong, such as race, gender, age, class, religion, or sexu ...
, 1892
*'' Cytherea''
Fabricius Fabricius (, ) is a surname. Notable people with the surname include: *people from the Ancient Roman Fabricia gens, gens Fabricia: **Gaius Fabricius Luscinus, the first of the Fabricii to move to Rome * Johann Goldsmid (1587–1616), known by his ...
, 1794
*'' Cyx'' Evenhuis, 1993 *'' Dasypalpus'' Macquart, 1840 *'' Desmatomyia'' Williston, 1895 *''
Desmatoneura ''Desmatoneura'' is a genus of bee flies in the family Bombyliidae. There are about 18 described species in ''Desmatoneura''. Species These 18 species belong to the genus ''Desmatoneura'': * ''Desmatoneura aegypticola'' (Paramonov, 1935) * ''Des ...
'' Williston, 1895 *'' Deusopora'' Hull, 1971 *'' Diatropomma'' Bowden, 1962 *'' Dicranoclista'' Bezzi, 1924 *'' Diochanthrax'' Hall, 1975 *''
Dipalta ''Dipalta'' is a North American genus of bee flies in the family Bombyliidae. There are two described species of ''Dipalta''. The genus is closely related to ''Villa''. Description These medium-sized flies have conical faces, and have three ma ...
'' 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 Becker () is one of the German-language surnames, along with Bäcker and Baecker, that derive from the root, which refers to baking. The surname began as a name for a baker (and thus his family). In northern Germany, it can also derive from th ...
, 1915
*'' Efflatounia'' Bezzi, 1925 *'' Enica'' Macquart, 1834 *'' Epacmoides'' Hesse, 1956 *'' Epacmus'' Osten Sacken, 1886 *'' Eremyia'' Greathead, 1996\ *'' Eristalopsis'' Evenhuis, 1985 *'' Eucessia'' Coquillett, 1886 *''
Euchariomyia ''Euchariomyia'' is a monotypic genus of the subfamily Bombyliinae. The only species is ''Euchariomyia dives''. Taxonomy ''Euchariomyia dives'' is described by French entomologist Jacques-Marie-Frangile Bigot in 1888. This species is highly var ...
''
Bigot Discrimination is the process of making unfair or prejudicial distinctions between people based on the groups, classes, or other categories to which they belong or are perceived to belong, such as race, gender, age, class, religion, or sexu ...
, 1888
*'' Euprepina'' Hull, 1971 *'' Eurycarenus'' Loew, 1860 *'' Euryphthiria'' Evenhuis, 1986 *'' Eusurbus'' Roberts, 1929 *'' Exechohypopion'' Evenhuis, 1991 *'' Exepacmus'' Coquillett, 1894 *''
Exhyalanthrax ''Exhyalanthrax'' is a small genus of Bombyliidae, bombyliid Fly, flies. Bombyliids are commonly known as ''bee flies'' due to their resemblance to bees. ''Exhyalanthrax'' are found in the Afrotropical realm and the Palearctic realm. Exhyalanthra ...
''
Becker Becker () is one of the German-language surnames, along with Bäcker and Baecker, that derive from the root, which refers to baking. The surname began as a name for a baker (and thus his family). In northern Germany, it can also derive from th ...
, 1916
*'' Exoprosopa'' Macquart, 1840 *'' Geminaria'' Coquillett, 1894 *'' Geron'' Meigen, 1820 *'' Glaesamictus'' Hennig, 1966 *'' Gnumyia'' Bezzi, 1921 *'' Gonarthrus'' Bezzi, 1921 *'' Gyrocraspedum''
Becker Becker () is one of the German-language surnames, along with Bäcker and Baecker, that derive from the root, which refers to baking. The surname began as a name for a baker (and thus his family). In northern Germany, it can also derive from th ...
, 1913
*'' Hallidia'' Hull, 1970 *''
Hemipenthes ''Hemipenthes'' is a large genus of fly, flies belonging to the family Bombyliidae (bee-flies). There are many described species, distributed throughout the Holarctic, Holarctic realm. These are small to large robust flies with a body length of 5 ...
'' Loew, 1869 *'' Heteralonia''
Rondani Rondani is an Italian surname. Notable people having this name include: * Camillo Rondani, (1808–1879), an Italian entomologist noted for his studies of Diptera * Giovanni Maria Francesco Rondani Giovanni Maria Francesco Rondani (15 July 1490 ...
, 1863
*'' Heterostylum'' Macquart, 1848 *''
Heterotropus ''Heterotropus'' is a genus of bee flies in the family Bombyliidae The Bombyliidae are a family of flies, commonly known as bee flies. Some are colloquially known as bomber flies. Adults generally feed on nectar and pollen, some being importa ...
'' Loew, 1873 *'' Hyperalonia''
Rondani Rondani is an Italian surname. Notable people having this name include: * Camillo Rondani, (1808–1879), an Italian entomologist noted for his studies of Diptera * Giovanni Maria Francesco Rondani Giovanni Maria Francesco Rondani (15 July 1490 ...
, 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 ''Lepidanthrax'' is a genus of bee flies in the family Bombyliidae. There are at least 50 described species in ''Lepidanthrax''. The genus is primarily North America, North and Central American, but one species is recorded from the Galapagos Islan ...
'' Osten Sacken, 1886 *'' Lepidochlanus'' Hesse, 1938 *'' Lepidophora'' Westwood, 1835 *''
Ligyra ''Ligyra'' is a genus of bee flies in the Bombyliidae The Bombyliidae are a family of flies, commonly known as bee flies. Some are colloquially known as bomber flies. Adults generally feed on nectar and pollen, some being important pollinato ...
''
Newman Newman is a surname of Germanic Anglo-Saxon origins. Newman is the modern English form of the name used in Great Britain and among people of British ancestry around the world (as is 'Numan'), while Neumann (with variant spellings) is used in Ger ...
, 1841
*'' Litorhina'' Bowden, 1975 *'' Lomatia'' Meigen, 1822 *'' Lordotus'' Loew, 1863 *'' Macrocondyla''
Rondani Rondani is an Italian surname. Notable people having this name include: * Camillo Rondani, (1808–1879), an Italian entomologist noted for his studies of Diptera * Giovanni Maria Francesco Rondani Giovanni Maria Francesco Rondani (15 July 1490 ...
, 1863
*'' Mallophthiria'' Edwards, 1930 *'' Mancia'' Coquillett, 1886 *'' Mandella'' Evenhuis, 1983 *'' Mariobezzia''
Becker Becker () is one of the German-language surnames, along with Bäcker and Baecker, that derive from the root, which refers to baking. The surname began as a name for a baker (and thus his family). In northern Germany, it can also derive from th ...
, 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'' Rodolfo Amando Philippi, Philippi, 1865 *''Neobombylodes'' Evenhuis, 1978 *''Neodiplocampta'' Charles Howard Curran, Curran, 1934 *''Neodischistus'' Painter, 1933 *''Neosardus'' Roberts, 1929 *''Nomalonia''
Rondani Rondani is an Italian surname. Notable people having this name include: * Camillo Rondani, (1808–1879), an Italian entomologist noted for his studies of Diptera * Giovanni Maria Francesco Rondani Giovanni Maria Francesco Rondani (15 July 1490 ...
, 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 *''Paramonovius'' Li & Yeates, 2018 *''Paranthrax''
Bigot Discrimination is the process of making unfair or prejudicial distinctions between people based on the groups, classes, or other categories to which they belong or are perceived to belong, such as race, gender, age, class, religion, or sexu ...
, 1876
*''Parasysteochus'' Hall, 1976 *''Paratoxophora'' Engel, 1936 *''Paravilla'' Painter, 1933 *''Parisus''
Walker Walker or The Walker may refer to: People *Walker (given name) *Walker (surname) *Walker (Brazilian footballer) (born 1982), Brazilian footballer Places In the United States *Walker, Arizona, in Yavapai County *Walker, Mono County, California * ...
, 1852
*''Perengueyimyia''
Bigot Discrimination is the process of making unfair or prejudicial distinctions between people based on the groups, classes, or other categories to which they belong or are perceived to belong, such as race, gender, age, class, religion, or sexu ...
, 1886
*''Petrorossia'' Bezzi, 1908 *''Phthiria'' Meigen, 1803 *''Pilosia'' Hull, 1973 *''Pipunculopsis'' Bezzi, 1925 *''Platamomyia'' Brèthes, 1925 *''Plesiocera'' Macquart, 1840 *''Poecilanthrax'' Osten Sacken, 1886 *''Poecilognathus'' Johann Friedrich Jaennicke, 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 (fly), Semiramis''
Becker Becker () is one of the German-language surnames, along with Bäcker and Baecker, that derive from the root, which refers to baking. The surname began as a name for a baker (and thus his family). In northern Germany, it can also derive from th ...
, 1913
*''Semistoechus'' Hall, 1976 *''Sericosoma'' Macquart, 1850 *''Sericothrix'' Hall, 1976 *''Sericusia'' Edwards, 1937 *''Sinaia (fly), Sinaia''
Becker Becker () is one of the German-language surnames, along with Bäcker and Baecker, that derive from the root, which refers to baking. The surname began as a name for a baker (and thus his family). In northern Germany, it can also derive from th ...
, 1916
*''Sisyromyia'' White, 1916 *''Sisyrophanus'' Karsch, 1886 *''Sosiomyia'' Bezzi, 1921 *''Sparnopolius'' Loew, 1855 *''Sphenoidoptera'' Williston, 1901 *''Spogostylum'' Macquart, 1840 *''Staurostichus'' Hull, 1973 *''Stomylomyia''
Bigot Discrimination is the process of making unfair or prejudicial distinctions between people based on the groups, classes, or other categories to which they belong or are perceived to belong, such as race, gender, age, class, religion, or sexu ...
, 1888
*''Stonyx'' Osten Sacken, 1886 *''Synthesia (fly), Synthesia'' Bezzi, 1921 *''Systoechus'' Loew, 1855 *''Systropus'' Wiedemann, 1820 *''Thevenetimyia''
Bigot Discrimination is the process of making unfair or prejudicial distinctions between people based on the groups, classes, or other categories to which they belong or are perceived to belong, such as race, gender, age, class, religion, or sexu ...
, 1875
*''Thraxan'' Yeates & Lambkin, 1998 *''
Thyridanthrax ''Thyridanthrax'' is a genus of bee flies in the family Bombyliidae. There are about 50 described species in the genus ''Thyridanthrax'', found across Eurasia Eurasia ( , ) is a continental area on Earth, comprising all of Europe and Asia ...
'' 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 (fly), 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'' Paul Gustav Eduard Speiser, Speiser, 1920 *''Zaclava'' Hull, 1973 *''Zinnomyia'' Hesse, 1955 *''Zyxmyia'' Bowden, 1960


Gallery

Image:Unidentified Beeflies.jpg, Two species of unidentified beeflies from Coimbatore, Tamil Nadu, India. Image:12Sep Ft Union National Monument 44-1.jpg, A bombyliid fly visiting a flower. Image:Bee Fly x 3 - 2012-4.jpg, Bee fly in Hampshire, United Kingdom conservatory . Image:FlowerBee2011.png, Bee fly landing on a flower Image:Bee fly feeding02.jpg, '' Exoprosopa'' sp. feeding Image:Bombyliid 7294.jpg, '' Lepidophora'' on ''Bidens, Bidens laevis''


References


Further reading

* * * * * * *


External links


Image Gallery from Diptera.info

Bombyliidae
(Bee Flies) by David K. Yeates and Christine L. Lambkin in the Tree of Life web project. Consulted March 28, 2007.
Wing venation
{{Authority control Bombyliidae, Brachycera families Insects used as insect pest control agents Articles containing video clips Taxa named by Pierre André Latreille Brachycera