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''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 close resemblance to
bumblebee A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the genus ''Bombus'', part of Apidae, one of the bee families. This genus is the only extant group in the tribe Bombini, though a few extinct related gener ...
s and are often mistaken for them. ''B. major'' exhibits a unique flight behavior known as "yawing" and plays a role in general pollination, without preference of flower types. The fly does not bite, sting, or spread disease. However, the fly uses this mimicry of bumblebees to its own advantage, allowing close access to host solitary bee and wasp nests in order to deposit its eggs. After hatching, the larvae find their way into the nests to parasitically feed on the grubs.


Description

''B. major'' is part of the family Bombyliidae, with a reported 6000 species worldwide. The subfamily Bombyliinae contains approximately 1100 identified species. The genus ''
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'', ...
'' currently comprises around 450 described species. ''B. major'' can be found from April to June throughout
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
,
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
and some parts of
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...
, concentrated in the northern hemisphere. The species occurs across a variety of environments, from arid to moist.


Morphology

The adult body size varies from 6.3 to 12 mm in length and is considered a relatively medium-sized fly. The body color is dark, but is densely covered by a thick coat of lighter color hairs. The head is typically brown and black hairs, but the lower portion of the head is mostly white hairs. It has dark patches on the
anterior Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
half of the wings and long hairy legs that dangle while in flight. The dark wing span can range from 8.4 to 14 mm and has a dark brown edge. Their boldly patterned wings have a distinct dividing border through the horizontal middle between the dark and clear portions. Their antennae are typically very short and pointed. Additionally, the species has long legs and a long rigid proboscis found in the front of the head, which is used to feed on the nectar of flowers. The proboscis ranges from 5.5 to 7.5 mm in length. While its wings continue to beat, its front legs grip the flower and its long rigid beak is inserted to collect the nectar.''Insects'' , Collins Gem , Guide, 1986, page 114, Despite its fearsome appearance, the beak is quite harmless. Males are typically smaller than females. Movement of the bee is categorized by both hovering and darting between locations. The flies also emit a high-pitched buzz.


Mimicry

The species gets its name from its similarity in appearance to bumblebees. This mimicry is likely a defense mechanism against predators and also allows the species to get close to the nest of their hosts. However, there are several distinguishable factors. Bees have two pairs of wings, while ''B. major'' has only one pair. The body is more triangular shaped than the typical round oval shape of a bumblebee.


Life Cycle

''B. major'' is mostly seen in the spring, beginning to appear at the end of March and large numbers seen until the end of May, with the species being sighted into June. The fly is
holometabolous Holometabolism, also called complete metamorphosis, is a form of insect development which includes four life stages: egg, larva, pupa, and imago (or adult). Holometabolism is a synapomorphic trait of all insects in the superorder Endopterygot ...
. ''B. major'' has several host species, including the brood of solitary wasps and bees, particularly digging bees such as ''
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 monophyletic group that is difficult to split into more manageable divisions; currently, ''Andrena'' i ...
''. Egg deposition takes place by the female hovering above the entrance of a host insect nest, usually a solitary bee, and throwing down her eggs using a flicking movement.Boesi, R., Polidori, C. and Andrietti, F. 2009 — Searching for the Right Target: Oviposition and Feeding Behavior in Bombylius Bee Flies (Diptera:Bombyliidae).''Zool. Stud.'', 48:141-150. The larvae are hypermetamorphic
parasitoids In evolutionary ecology, a parasitoid is an organism that lives in close association with its host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionary strategies within parasi ...
which then feed on the food stored, as well as the young solitary bees or wasps. If the female is unable to flick her eggs near the nest, she plants them on flowers visited by the host insects. The developing larvae then make their way to the host nest or attach themselves to the bees or wasps to then be carried to the nest.


Egg

The parasitic eggs of ''B. major'' are produced in large numbers, however few will make it to the host insect burrow entrance. The female fly will dip down and coat her rear abdomen with dust that covers the eggs as they leave the female.


Larvae

Larvae live parasitically in the nests of various solitary bees and wasps. When the fly larva locates a host larva, it will consume it slowly, greatly increasing in size as it tightly holds onto the host, eventually becoming a pupa and overwintering.


Pupae

White larvae gradually turn into a yellowish brown pupa, with distinct mouthparts, wings, antennae, and legs.


Diet

The species acts as a nectar robber; this foraging behavior allows the species to feed on floral nectar and is an essential part of adult fly diets. This is facilitated by the characterizable long proboscis of the fly, which is horizontally inserted into the flower. This occurs as the fly continues to buzz in the air, without touching either the anthers or stigma of the flower. The fly also consumes pollen as part of its diet, with considerable differences between the sexes. Males and females visit the same range of flowers as a food source. Along with one other species, ''
Bombylius pygmaeus ''Bombylius pygmaeus'' is a species of bee flies in the family Bombyliidae The Bombyliidae are a family of flies, commonly known as bee flies. Adults generally feed on nectar and pollen, some being important pollinators. Larvae are mostly paras ...
'', ''B. major'' was observed to selectively (and almost exclusively) visit bluets at several North American sites despite the abundant presence of many other flowers. Of the other pollinators present, these flies were also the most frequent visitors to the flowers.


Female

In Russia, female ''B. major'' act in a narrow oligophagous manner, favoring a limited number of food sources, including pollen grains of Siberian squill (''
Scilla siberica ''Scilla siberica'', the Siberian squill or wood squill, is a species of flowering plant in the family Asparagaceae, native to southwestern Russia, the Caucasus, and Turkey. Despite its name, it is not native to Siberia. Description Growing to ...
''), lesser celandine (''
Ficaria verna ''Ficaria verna'' (formerly ''Ranunculus ficaria'' ), commonly known as lesser celandine or pilewort, is a low-growing, hairless perennial flowering plant in the buttercup family Ranunculaceae native to Europe and Western Asia. It has fleshy dar ...
''), and willow (''
Salix caprea ''Salix caprea'', known as goat willow, pussy willow or great sallow, is a common species of willow native to Europe and western and central Asia.Meikle, R. D. (1984). ''Willows and Poplars of Great Britain and Ireland''. BSBI Handbook 4. . Des ...
'') over dandelion (''
Taraxacum officinale ''Taraxacum officinale'', the dandelion or common dandelion, is a flowering herbaceous perennial plant of the dandelion genus in the family Asteraceae (syn. Compositae). The common dandelion is well known for its yellow flower heads that turn i ...
''), coltsfoot (''
Tussilago farfara ''Tussilago farfara'', commonly known as coltsfoot, is a plant in the tribe Senecioneae in the family Asteraceae, native to Europe and parts of western and central Asia. The name "tussilago" is derived from the Latin ''tussis'', meaning cough, an ...
''), and primrose (''
Primula vulgaris ''Primula vulgaris'', the common primrose, is a species of flowering plant in the family Primulaceae, native to western and southern Europe, northwest Africa, and parts of southwest Asia.''Flora Europaea'Primula vulgaris''/ref> The common name ...
'').


Male

In the same study, male ''B. major'' preferred plants similar to females, with the exception of dandelion. However, an examination of the gut showed a regularly lower amount of pollen than in females. Males consistently consume less quantities of pollen than females. Ingestion of pollen differs throughout a male life cycle, with more pollen being consumed in towards earlier stages. Overall, males had a preference for nectar and fed on significantly greater portions of nectar over pollen.


Behavior

There is continued research on the behavior of ''B. major'' in respect to environmental factors, mating, and human interaction.


Flight

It has been discovered that the fly is capable of a unique type of flight behavior, which was discovered with the use of a high speed camera. In this behavior, the flies are seen to rotate around a vertical axis as they fly (this action is known as “yawing”). However, it is still unknown what can cause this behavior to be triggered and what purpose it serves, but a proposed explanation includes mating habits.


Pollinator Role

The ''B. major'' bee-fly is a common, generalist floral pollinator, meaning that it does not give preference to one flower over another, instead pollinating a wide variety of plant families and species. The fly uses its proboscis to carry and transfer the pollen. The species is a dominant pollinator within its community, sometimes even pollinating up to two thirds of the local flowers. In addition, ''B. major'' will visit and pollinate plants that attract few other species. Some types of flowers, for example ''
Pulmonaria officinalis ''Pulmonaria officinalis'', common names lungwort, common lungwort, Mary's tears or Our Lady's milk drops, is a herbaceous rhizomatous evergreen perennial plant of the genus ''Pulmonaria'', belonging to the family Boraginaceae. Etymology The ge ...
,'' will be almost exclusively pollinated by ''B. major'', with other species contributing a negligible amount to that plants pollination. Some flower species, such as ''
Delphinium tricorne ''Delphinium tricorne'', known by the common names dwarf larkspur or spring larkspur, is a species of flowering plant in the Ranunculaceae (buttercup) family. It is native to the central and eastern United States, where it is the most common ''De ...
'', are even specifically adapted to the fly in terms of color, shape, and form. If given the choice, ''B. major'' will have a consistency in plant choice.


Flower Attraction

Long distance floral attraction is governed by optical sense, with color being the most important factor. The flies are typically more attracted to blue and violet colors, and occasionally yellow, over orange and pink. However, short distance floral attraction is based on the fly's olfactory sense.


Activity

The fly is mostly active during day hours when the weather conditions are warm and sunny. ''B. major'' is attracted to sunnier places and is more likely to pollinate these areas, with a larger average of flower visits in areas of higher amounts of sunshine. The fly will hide in the trees during the night and usually dart away from a cast shadow and occasionally hide in clean washing brought in fresh from the washing line and fly out causing unsettled behaviour in the discoverer.


References

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External links

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Photograph by Felipe vilarroya
{{Taxonbar, from=Q1549111 Bombyliidae Asilomorph flies of Europe Diptera of Asia Diptera of North America Insects of the United States Flies described in 1758 Taxa named by Carl Linnaeus Articles containing video clips