The 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.
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). 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 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. 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.
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 insects relative to its species richness.
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.
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.
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.
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 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).
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.
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. 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. 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. 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, 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.
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.
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 Boris Borisovitsch Rohdendorf (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). Zaitzev (1991) moved the genus Mythicomyia and several other minor genera in the family Mythicomyiidae, Yeates (1992, 1994) 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. 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:
Anthracinae is a subfamily of bee flies in the family Bombyliidae. There are more than 80 genera and 2,000 described species in Anthracinae.Anthrax (fly)
Anthrax is a genus of bombyliid flies, commonly known as "bee-flies" due to their resemblance to bees. Most are dull black flies, and are usually small to medium in size, 4–20 millimetres (0.2–0.8 in), and many species have striking wing patterns.Anthrax is a very large genus. While worldwide in distribution, most species are from the Palaearctic and Afrotropic regions. The genus includes species parasitic on tiger beetles – an unusual trait among the bee-flies. A. anthrax larvae parasitize bees. Many North American species parasitize solitary wasps.The type species is Musca morio Linnaeus, 1758, later found to be a misidentification of Musca anthrax Schrank, 1781.Asiloidea
The Asiloidea comprise a very large superfamily insects in the order Diptera, the true flies. It has a cosmopolitan distribution, occurring worldwide, with many species living in dry, sandy habitat types. It includes the family Bombyliidae, the bee flies, which are parasitoids, and the Asilidae, the robber flies, which are predators of other insects. Members of the other families are mainly flower visitors as adults and predators as larvae.It is not entirely clear that this superfamily is monophyletic. It is closely related to the Empidoidea and the Cyclorrhapha.Bombyliinae
Bombyliinae is a subfamily of bee flies in the family Bombyliidae. There are about 65 genera and more than 1000 described species in Bombyliinae.Bombylius
Bombylius is a large genus of flies belonging to the family Bombyliidae. They are known as the bee-flies, due to their striking resemblance to bees and bumblebees, and are distributed worldwide. One species of the genus, Bombylius major, is widely distributed throughout the northern hemisphere and is very well known.Conophorini
Conophorini is a tribe of bee flies in the family Bombyliidae. There are about 11 genera and 125 described species in Conophorini.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.Exoprosopa
Exoprosopa is a large cosmopolitan genus of flies belonging to the family Bombyliidae (bee-flies), with over 325 described species. The genus parasitizes a wide range of insects, including locust and larvae of wasps.Geron (fly)
Geron is a genus of bee flies (insects in the family Bombyliidae). There are at least 180 described species in Geron.Hilarimorphidae
The Hilarimorphidae or hilarimorphid flies are a family of Diptera. They are placed in the superfamily Asiloidea, though some considerable doubt exists, but the consensus is that they are most closely related to the Bombyliidae. Most species are nearctic.Lepidanthrax
Lepidanthrax is a genus of bee flies in the family Bombyliidae. There are at least 50 described species in Lepidanthrax.Mythicomyiidae
Mythicomyiidae, commonly called mythicomyiids, are very tiny 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 the tropics, but genera such as Cephalodromia and Platypygus are known from these regions. Many of these "microbombyliids" have a humpbacked thorax (as in the Acroceridae) and lack the dense vestiture common in the Bombyliidae. Mythicomyiids have until recently not had much attention in the literature. Their small size has caused them to be missed when collecting. Yellow pan trapping and fine-mesh netting in Malaise and aerial sweep nets has resulted in a number of undescribed species from many parts of the world. A high diversity of both genera and species exists for this family in Africa, especially northern and southern portions. About 350 species are known (most in the genus Mythicomyia Coquillett). Hundreds more await description.
Because of their extremely small size and curious body shapes, some genera have been at times placed in the Acroceridae or Empididae. Originally, taxa were placed in the subfamily Mythicomyiinae in the Empididae. Later, they were transferred to the Bombyliidae, where mythicomyiids have long been treated. Zaitzev (1991) was the first to give characters warranting raising the group to family level. Subsequent workers have followed Zaitzev's lead and treat the group as a separate family. The family is separated from the Bombyliidae by the unbranched wing vein R4+5 (branched in Bombyliidae), the extremely reduced or absent maxillary palpi (present in Bombyliidae), wings held together over the abdomen at rest (held at an angle in Bombyliidae), and the abdominal spiracles being placed in the terga (placed in the pleural membrane in Bombyliidae). Augmenting the morphological characters, it is also a much older lineage than any known Bombyliidae, dating from as far back as the Middle Jurassic (Palaeoplatypygus Kovalev; Callovian: 163–168 mya) with other genera known from the Cretaceous (Procyrtosia Hennig and Proplatypygus Zaitzev). Bombyliidae, though, are not known from any older fossil material than Eocene Baltic amber deposits (Lutetian to Rupelian: 30–52 million years ago).Paravilla
Paravilla is a genus of bee flies (insects in the family Bombyliidae). There are at least 50 described species in Paravilla.Poecilanthrax
Poecilanthrax is a large, primarily Nearctic genus of flies belonging to the family Bombyliidae (bee-flies).
These are large to large robust flies with a body length of 8–14 mm. They have large elongate pictured wings, with 2 or 3 submarginal cells and no vein M2. The abdomen is moderately lengthened, with 7 visible tergites. The front tarsi are reduced, and the tibia smooth. The larvae feed on the moth larva of members of the family Noctuidae. Adults feed on pollen.Systropus
Systropus is a genus of bee flies (insects in the family Bombyliidae). There are about 16 described species in Systropus.Toxophora
Toxophora is a genus of flies belonging to the family Bombyliidae (bee-flies). There are 47 described species, distributed throughout the world, although they are most abundant in Southwestern United States and western Mediterranean. World catalog of bee flies (Diptera: Bombyliidae)
They are strange, stout, robust flies with a hunchbacked form, with a body length of 6–12 mm and wings 4 to 7.5mm. Most species are black with banding or spots.Larvae feed in the nest of wasps of the genus Odynerus.Ectoparasitoids in nests of solitary bees & wasps Research.net. Yeates, D. The evolutionary pattern of host use in the Bombyliidae (Diptera): a diverse family of parasitoid fliesVilla (fly)
Villa is a genus of flies belonging to the bee-fly family (Bombyliidae). They range in size from 5 to 17 millimetres (0.20 to 0.67 in), and have typically rounded heads. The males of some species have a brilliant mat of silvery patagial scales. About 270 Villa species are found on all continents except Antarctica. They can be distinguished from similar genera (Hemipenthes) by their wing venation.Villini
Villini is a tribe of bee flies in the family Bombyliidae. There are about 14 genera and at least 100 described species in Villini.Xenox
Xenox is a genus of bee flies (insects in the family Bombyliidae). There are about five described species in Xenox, all of which parasitize bees in the genus Xylocopa as larvae.
Extant Diptera families