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{{Short description|Family of flies}}
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The '''Bombyliidae''' are a [[family (biology)|family]] of [[fly|flies]]. Their common name are '''bee flies''' or '''humbleflies'''. Adults generally feed on [[nectar]] and [[pollen]], some being important [[pollinator]]s. Larvae generally are [[parasitoid]]s of other insects.
The '''Bombyliidae''' are a [[family (biology)|family]] of [[fly|flies]], commonly known as '''bee flies'''. Some are colloquially known as '''bomber flies'''. Adults generally feed on [[nectar]] and [[pollen]], some being important [[pollinator]]s. Larvae are mostly [[parasitoid]]s of other insects.


== Overview ==
== Overview ==
The '''Bombyliidae''' are a large [[family (biology)|family]] of [[fly|flies]] comprising hundreds of [[genus|genera]], but the lifecycles of most species are known poorly, or not at all. They range in size from very small (2&nbsp;mm in length) to very large for flies (wingspan of some 40&nbsp;mm).<ref name="isbn1-86872-713-0">{{cite book |author1=Alan Weaving |author2=Mike Picker |author3=Griffiths, Charles Llewellyn |title=Field Guide to Insects of South Africa |publisher=New Holland Publishers, Ltd |location= |year=2003 |pages= |isbn=1-86872-713-0 |oclc= |doi= |accessdate=}}</ref><ref name="HullFM">Hull, Frank Montgomery, Bee flies of the world: the genera of the family Bombyliidae Washington, Smithsonian Institution Press 1973 {{ISBN|0-87474-131-9}}. Downloadable from: https://archive.org/details/beefliesofworl2861973hull</ref> When at rest, many species hold their wings at a characteristic "swept back" angle. Adults generally feed on [[nectar]] and [[pollen]], some being important [[pollinator]]s, 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 [[bee]]s and accordingly the prevalent common name for a member of the family is '''bee fly'''.<ref name="HullFM"/> Possibly the resemblance is [[Batesian mimicry]], affording the adults some protection from [[predator]]s.
The '''Bombyliidae''' are a large [[family (biology)|family]] of [[fly|flies]] comprising hundreds of [[genus|genera]], but the life cycles of most species are known poorly, or not at all. They range in size from very small (2&nbsp;mm in length) to very large for flies (wingspan of some 40&nbsp;mm).<ref name="isbn1-86872-713-0">{{cite book |author1=Alan Weaving |author2=Mike Picker |author3=Griffiths, Charles Llewellyn |title=Field Guide to Insects of South Africa |publisher=New Holland Publishers, Ltd |year=2003 |isbn=1-86872-713-0 }}</ref><ref name="HullFM">Hull, Frank Montgomery, Bee flies of the world: the genera of the family Bombyliidae Washington, Smithsonian Institution Press 1973 {{ISBN|0-87474-131-9}}. Downloadable from: https://archive.org/details/beefliesofworl2861973hull</ref> When at rest, many species hold their wings at a characteristic "swept back" angle. Adults generally feed on [[nectar]] and [[pollen]], some being important [[pollinator]]s, often with spectacularly long proboscises 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. Many Bombyliidae superficially resemble [[bee]]s and accordingly the prevalent common name for a member of the family is '''bee fly'''.<ref name="HullFM"/> Possibly the resemblance is [[Batesian mimicry]], affording the adults some protection from [[predator]]s.


The larval stages are predators or [[parasitoid]]s of the eggs and [[larva]]e of other insects. The adult females usually deposit eggs in the vicinity of possible hosts, quite often in the burrows of [[beetle]]s or [[wasp]]s/solitary bees. Although insect parasitoids usually are fairly host-specific, often highly host-specific, some Bombyliidae are [[opportunistic feeder|opportunistic]] and will attack a variety of hosts.
The larval stages are predators or [[parasitoid]]s of the eggs and [[larva]]e of other insects. The adult females usually deposit eggs in the vicinity of possible hosts, quite often in the burrows of [[beetle]]s or [[wasp]]s/solitary bees. Although insect parasitoids usually are fairly host-specific, often highly host-specific, some Bombyliidae are [[opportunistic feeder|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 [[insect]]s relative to its species richness. The family has a patchy fossil record, with species being known from a handful of localities,<ref>{{Cite journal|last=Wedmann|first=Sonja|last2=Yeates|first2=David K.|title=Eocene Records of Bee Flies (Insecta, Diptera, Bombyliidae, Comptosia): Their Palaeobiogeographic Implications and Remarks on the Evolutionary History of Bombyliids|date=2008-01-17|journal=Palaeontology|language=en|volume=51|issue=1|pages=231–240|doi=10.1111/j.1475-4983.2007.00745.x}}</ref> the oldest known species are known from the Middle Cretaceous [[Burmese amber]], around 99 million years old.<ref>{{Cite journal|last=Ye|first=Xiuna|last2=Yao|first2=Gang|last3=Shih|first3=Chungkun|last4=Ren|first4=Dong|last5=Wang|first5=Yongjie|date=August 2019|title=New bee flies from the mid-Cretaceous Myanmar amber (Brachycera: Asiloidea: Bombyliidae)|journal=Cretaceous Research|language=en|volume=100|pages=5–13|doi=10.1016/j.cretres.2019.03.026}}</ref>
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]]s relative to its species richness. The family has a patchy fossil record, with species being known from a handful of localities,<ref>{{Cite journal|last1=Wedmann|first1=Sonja|last2=Yeates|first2=David K.|title=Eocene Records of Bee Flies (Insecta, Diptera, Bombyliidae, Comptosia): Their Palaeobiogeographic Implications and Remarks on the Evolutionary History of Bombyliids|date=2008-01-17|journal=Palaeontology|language=en|volume=51|issue=1|pages=231–240|doi=10.1111/j.1475-4983.2007.00745.x|s2cid=128576916 |doi-access=free|bibcode=2008Palgy..51..231W }}</ref> the oldest known species are known from the Middle Cretaceous [[Burmese amber]], around 99 million years old.<ref>{{Cite journal|last1=Ye|first1=Xiuna|last2=Yao|first2=Gang|last3=Shih|first3=Chungkun|last4=Ren|first4=Dong|last5=Wang|first5=Yongjie|date=August 2019|title=New bee flies from the mid-Cretaceous Myanmar amber (Brachycera: Asiloidea: Bombyliidae)|journal=Cretaceous Research|language=en|volume=100|pages=5–13|doi=10.1016/j.cretres.2019.03.026|bibcode=2019CrRes.100....5Y |s2cid=135365568 }}</ref>


[[Image:Bombyliidae 3 by kadavoor.jpg|thumb|230px|''[[Euchariomyia dives]]'', a Bombyliidae in 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.]]
[[Image:Bombyliidae 3 by kadavoor.jpg|thumb|230px|''[[Euchariomyia dives]]'', a Bombyliidae in 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==
==Morphology==
{{Unreferenced section|date=August 2024}}


===Adult===
===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&nbsp;mm to 2.5&nbsp;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 [[Batesian mimicry|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.
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&nbsp;mm to 2.5&nbsp;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 [[Batesian mimicry|mimics]] of [[Apoidea|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.


[[File:1500exoprosopacaliptera DSC6698 DxO.jpg|thumb|''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]]
[[File:1500exoprosopacaliptera DSC6698 DxO.jpg|thumb|''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 [[arista (biology)|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 [[Arthropod mouthparts#Labium|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 head is round, with a convex face, often [[holoptic]] in males. The antennae are of the type [[arista (biology)|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 [[Arthropod mouthparts#Labium|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 [[Glossary of entomology terms|empodia]] and, sometimes, also without [[pulvilli]] . The wings are transparent, often hyaline or evenly colored or with bands. The [[Calypter|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 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 [[Glossary of entomology terms|empodia]] and, sometimes, also without [[pulvilli]]. The wings are transparent, often hyaline or evenly colored or with bands. The [[Calypter|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 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.
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.
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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:
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).
* 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 ([[Syrphidae#Description|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 wings lack a "false rear edge" and often have large dark areas with sharp boundaries, or complex patterns of spots ([[Syrphidae#Description|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 abdomen and thorax hardly ever have large glossy areas formed by exposed cuticle (hoverflies often have glossy cuticular body surfaces).


===Larva===
===Larva===
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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.
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.<ref>{{cite web|url=https://www.youtube.com/watch?v=whRkZ1BS1dE|title=Gewone wolzwever Bombylius major, eieren droppend in een kolonie zandbijen|last=Roy Kleukers|date=22 June 2013|publisher=|via=YouTube}}</ref> 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.<ref>{{cite journal|doi=10.1111/j.1095-8312.1997.tb01490.x |volume=60 |issue=2 |title=The evolutionary pattern of host use in the Bombyliidae (Diptera): a diverse family of parasitoid flies |journal=Biological Journal of the Linnean Society |pages=149–185 |author=Yeates David K|year=1997 |doi-access=free }}</ref><ref>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'' Bee Flies (Diptera: Bombyliidae). Zoological Studies 48: 141-150.</ref> 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.
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.<ref>Archived at [https://ghostarchive.org/varchive/youtube/20211212/whRkZ1BS1dE Ghostarchive]{{cbignore}} and the [https://web.archive.org/web/20200518181356/https://www.youtube.com/watch?v=whRkZ1BS1dE Wayback Machine]{{cbignore}}: {{cite web|url=https://www.youtube.com/watch?v=whRkZ1BS1dE|title=Gewone wolzwever Bombylius major, eieren droppend in een kolonie zandbijen|last=Roy Kleukers|date=22 June 2013|via=YouTube}}{{cbignore}}</ref> 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.<ref>{{cite journal|doi=10.1111/j.1095-8312.1997.tb01490.x |volume=60 |issue=2 |title=The evolutionary pattern of host use in the Bombyliidae (Diptera): a diverse family of parasitoid flies |journal=Biological Journal of the Linnean Society |pages=149–185 |author=Yeates David K|year=1997 |doi-access=free }}</ref><ref>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'' Bee Flies (Diptera: Bombyliidae). Zoological Studies 48: 141-150.</ref> 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.


[[File:Villa sp - 2012-10-16.webm|thumb|thumbtime=0:28|''[[Villa (genus)|Villa]]'' sp. gathering sand grains]]
[[File:Villa sp - 2012-10-16.webm|thumb|thumbtime=0:28|''[[Villa (genus)|Villa]]'' sp. gathering sand grains]]
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==Species lists==
==Species lists==
[[File:1500lepidanthraxgirl DSC1876 DxO.jpg|thumb|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]]
[[File:1500lepidanthraxgirl DSC1876 DxO.jpg|thumb|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]]
*[http://www.faunaeur.org/full_results.php?id=10889 West Palaearctic including Russia]
*[https://web.archive.org/web/20051015232841/http://www.faunaeur.org/full_results.php?id=10889 West Palaearctic including Russia]
*[http://hbs.bishopmuseum.org/aocat/bombyliidae.html Australasian/Oceanian]
*[http://hbs.bishopmuseum.org/aocat/bombyliidae.html Australasian/Oceanian]
*[http://www.nearctica.com/nomina/main.htm Nearctic]
*[http://www.nearctica.com/nomina/main.htm Nearctic]
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[[File:2000macrocondyla chorista DSC6649 DxO.jpg|thumb|''[[Macrocondyla chorista]]'' in a grassy border in San Luis province, Argentina, illustrating less common features for Bombyliidae such as a slender abdomen and white patches on the wings]]
[[File:2000macrocondyla chorista DSC6649 DxO.jpg|thumb|''[[Macrocondyla chorista]]'' in a grassy border in San Luis province, Argentina, illustrating less common features for Bombyliidae such as a slender abdomen and white patches on the wings]]


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<ref name="Willi Hennig" >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}}.</ref>
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<ref name="Willi Hennig" >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}}.</ref>
[[Boris Borisovitsch Rohdendorf]] (1974) dealt with the family in a separate superfamily (Bombyliidea), linking it to the superfamily of Asilidea.<ref name=" Boris B. Rohdendorf" >Boris B. Rohdendorf, Brian Hocking, Harold Oldroyd, George E. Ball. The Historical Development of Diptera. University of Alberta, 1974: 75-77. {{ISBN|088864003X}}.</ref> 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.
[[Boris Rohdendorf]] (1974) dealt with the family in a separate superfamily (Bombyliidea), linking it to the superfamily of Asilidea.<ref name=" Boris B. Rohdendorf" >Boris B. Rohdendorf, Brian Hocking, Harold Oldroyd, George E. Ball. The Historical Development of Diptera. University of Alberta, 1974: 75-77. {{ISBN|088864003X}}.</ref> 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.


{{Clade
{{Clade
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}}


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)<ref name="Webb D.W. " >Webb D.W., 1981 Hilarimorphidae. in: McAlpine J.F. (Ed.), ''Manual of Nearctic Diptera''. Agriculture Canada, Ottawa, pp. 603-605.</ref> finally moved the genus ''Hilarimorpha'' into their own family ([[Hilarimorphidae]]). Zaitzev (1991)<ref name="Zaitzev, V.F. " >Zaitzev, V.F., 1991 On the phylogeny and systematics of the dipteran superfamily Bombylioidea (Diptera). ''Entomol. Obozr''. 70 [1991] : 716–36.</ref> moved the genus ''Mythicomyia'' and several other minor genera in the family [[Mythicomyiidae]], Yeates (1992, 1994)<ref name="Yeates, D.M." >{{cite journal |author=Yeates D.M. |year=1992 |title=Towards a monophyletic Bombyliidae (Diptera): the removal of the Proratinae (Diptera: Scenopinidae) |url=|journal=American Museum Novitates |volume=3051 |issue=|pages=1–30 }}</ref> 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.<ref name="Yeates & Lambkin" >Yeates & Lambkin, The Tree of Life, op. cit..</ref> 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.<ref>{{Cite journal |url=http://currents.plos.org/treeoflife/article/overcoming-the-effects-of-rogue-taxa-2cn7m3af919c4-1/ |doi=10.1371/currents.RRN1233|pmid=21686308|pmc=3088465|title=Overcoming the effects of rogue taxa: Evolutionary relationships of the bee flies|journal= PLOS Currents|volume=3|pages=RRN1233|year=2011|last1=Trautwein|first1=Michelle D.|last2=Wiegmann|first2=Brian M.|last3=Yeates|first3=David K.}}</ref>
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)<ref name="Webb D.W. " >Webb D.W., 1981 Hilarimorphidae. in: McAlpine J.F. (Ed.), ''Manual of Nearctic Diptera''. Agriculture Canada, Ottawa, pp. 603-605.</ref> finally moved the genus ''Hilarimorpha'' into their own family ([[Hilarimorphidae]]). Zaitzev (1991)<ref name="Zaitzev, V.F. " >Zaitzev, V.F., 1991 On the phylogeny and systematics of the dipteran superfamily Bombylioidea (Diptera). ''Entomol. Obozr''. 70 [1991] : 716–36.</ref> moved the genus ''Mythicomyia'' and several other minor genera in the family [[Mythicomyiidae]], Yeates (1992, 1994)<ref name="Yeates, D.M." >{{cite journal |author=Yeates D.M. |year=1992 |title=Towards a monophyletic Bombyliidae (Diptera): the removal of the Proratinae (Diptera: Scenopinidae) |journal=American Museum Novitates |issue=3051 |pages=1–30 }}</ref> 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.<ref name="Yeates & Lambkin" >Yeates & Lambkin, The Tree of Life, op. cit.</ref> 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.<ref>{{Cite journal |url=http://currents.plos.org/treeoflife/article/overcoming-the-effects-of-rogue-taxa-2cn7m3af919c4-1/ |doi=10.1371/currents.RRN1233|pmid=21686308|pmc=3088465|title=Overcoming the effects of rogue taxa: Evolutionary relationships of the bee flies|journal= PLOS Currents|volume=3|pages=RRN1233|year=2011|last1=Trautwein|first1=Michelle D.|last2=Wiegmann|first2=Brian M.|last3=Yeates|first3=David K. |doi-broken-date=1 November 2024 |doi-access=free }}</ref>


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:<ref name=Evenhuis2015/>
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:<ref name=Evenhuis2015/>
Line 137: Line 144:
**[[Aphoebantini]]
**[[Aphoebantini]]
**[[Exoprosopini]]
**[[Exoprosopini]]
**[[Plesiocerini]]
**[[Prorostomatini]]
**[[Villini]]
**[[Villini]]
**[[Xeramoebini]]
**[[Xeramoebini]]
Line 187: Line 194:
*''[[Amictus (fly)|Amictus]]'' <small>[[Christian Rudolph Wilhelm Wiedemann|Wiedemann]], 1817</small>
*''[[Amictus (fly)|Amictus]]'' <small>[[Christian Rudolph Wilhelm Wiedemann|Wiedemann]], 1817</small>
*''[[Amphicosmus]]'' <small>[[Daniel William Coquillett|Coquillett]], 1891</small>
*''[[Amphicosmus]]'' <small>[[Daniel William Coquillett|Coquillett]], 1891</small>
*''[[Anastoechus]]'' <small>[[Karl Robert Osten-Sacken|Osten Sacken]], 1877</small>
*''[[Anastoechus]]'' <small>[[Carl Robert Osten-Sacken|Osten Sacken]], 1877</small>
*''[[Anisotamia]]'' <small>[[Pierre-Justin-Marie Macquart|Macquart]], 1840</small>
*''[[Anisotamia]]'' <small>[[Pierre-Justin-Marie Macquart|Macquart]], 1840</small>
*''[[Anthrax (fly)|Anthrax]]'' <small>[[Giovanni Antonio Scopoli|Scopoli]], 1763</small>
*''[[Anthrax (fly)|Anthrax]]'' <small>[[Giovanni Antonio Scopoli|Scopoli]], 1763</small>
*''[[Antonia (insect)|Antonia]]'' <small>[[Hermann Loew|Loew]], 1856</small>
*''[[Antonia (fly)|Antonia]]'' <small>[[Hermann Loew|Loew]], 1856</small>
*''[[Antoniaustralia]]'' <small>[[Theodor Becker|Becker]], 1913</small>
*''[[Antoniaustralia]]'' <small>[[Theodor Becker|Becker]], 1913</small>
*''[[Apatomyza]]'' <small>[[Christian Rudolph Wilhelm Wiedemann|Wiedemann]], 1820</small>
*''[[Apatomyza]]'' <small>[[Christian Rudolph Wilhelm Wiedemann|Wiedemann]], 1820</small>
*''[[Aphoebantus]]'' <small>[[Hermann Loew|Loew]], 1872</small>
*''[[Aphoebantus]]'' <small>[[Hermann Loew|Loew]], 1872</small>
*''[[Apolysis (fly)|Apolysis]]'' <small>[[Hermann Loew|Loew]], 1860</small>
*''[[Apolysis (fly)|Apolysis]]'' <small>[[Hermann Loew|Loew]], 1860</small>
*''[[Astrophanes]]'' <small>[[Karl Robert Osten-Sacken|Osten Sacken]], 1877</small>
*''[[Astrophanes]]'' <small>[[Carl Robert Osten-Sacken|Osten Sacken]], 1877</small>
*''[[Atrichochira]]'' <small>Hesse, 1956</small>
*''[[Atrichochira]]'' <small>Hesse, 1956</small>
*''[[Australiphthiria]]'' <small>Evenhuis, 1986</small>
*''[[Australiphthiria]]'' <small>Evenhuis, 1986</small>
Line 203: Line 210:
*''[[Bombomyia]]'' <small>Greathead, 1995</small>
*''[[Bombomyia]]'' <small>Greathead, 1995</small>
*''[[Bombylella]]'' <small>Greathead, 1995</small>
*''[[Bombylella]]'' <small>Greathead, 1995</small>
*''[[Bombylisoma]]'' <small>[[Camillo Róndani|Róndani]], 1856</small>
*''[[Bombylisoma]]'' <small>[[Camillo Rondani|Rondani]], 1856</small>
*''[[Bombylius]]'' <small>[[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]], 1758</small>
*''[[Bombylius]]'' <small>[[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]], 1758</small>
*''[[Brachyanax]]'' <small>Evenhuis, 1981</small>
*''[[Brachyanax]]'' <small>Evenhuis, 1981</small>
Line 213: Line 220:
*''[[Caecanthrax]]'' <small>Greathead, 1981</small>
*''[[Caecanthrax]]'' <small>Greathead, 1981</small>
*''[[Callostoma]]'' <small>[[Pierre-Justin-Marie Macquart|Macquart]], 1840</small>
*''[[Callostoma]]'' <small>[[Pierre-Justin-Marie Macquart|Macquart]], 1840</small>
*''[[Callynthrophora]]'' <small>[[Ignaz Rudolph Schiner|Schiner]], 1868</small>
*''[[Callynthrophora]]'' <small>[[Ignaz Rudolph Schiner|Schiner]], 1868</small><ref name="Schiner1868">{{cite book |last1=Schiner |first1=I.R. |title=Diptera. vi In [Wullerstorf-Urbair, B. von (in charge)], Reise der osterreichischen Fregatte Novara. Zool. 2(1)B. |date=1868 |publisher=K. Gerold's Sohn |location=Wien |pages=388pp., 4 pls}}</ref>
*''[[Canariellum]]'' <small>Strand, 1928</small>
*''[[Canariellum]]'' <small>Strand, 1928</small>
*''[[Chalcochiton]]'' <small>[[Hermann Loew|Loew]], 1844</small>
*''[[Chalcochiton]]'' <small>[[Hermann Loew|Loew]], 1844</small>
*''[[Choristus]]'' <small>[[Francis Walker (entomologist)|Walker]], 1852</small>
*''[[Choristus]]'' <small>[[Francis Walker (entomologist)|Walker]], 1852</small>
*''[[Chrysanthrax]]'' <small>[[Karl Robert Osten-Sacken|Osten Sacken]], 1886</small>
*''[[Chrysanthrax]]'' <small>[[Carl Robert Osten-Sacken|Osten Sacken]], 1886</small>
*''[[Colossoptera]]'' <small>Hull, 1973</small>
*''[[Colossoptera]]'' <small>Hull, 1973</small>
*''[[Comptosia]]'' <small>[[Pierre-Justin-Marie Macquart|Macquart]], 1840</small>
*''[[Comptosia]]'' <small>[[Pierre-Justin-Marie Macquart|Macquart]], 1840</small>
Line 240: Line 247:
*''[[Dicranoclista]]'' <small>[[Mario Bezzi|Bezzi]], 1924</small>
*''[[Dicranoclista]]'' <small>[[Mario Bezzi|Bezzi]], 1924</small>
*''[[Diochanthrax]]'' <small>Hall, 1975</small>
*''[[Diochanthrax]]'' <small>Hall, 1975</small>
*''[[Dipalta]]'' <small>[[Karl Robert Osten-Sacken|Osten Sacken]], 1877</small>
*''[[Dipalta]]'' <small>[[Carl Robert Osten-Sacken|Osten Sacken]], 1877</small>
*''[[Diplocampta]]'' <small>[[Ignaz Rudolph Schiner|Schiner]], 1868</small>
*''[[Diplocampta]]'' <small>[[Ignaz Rudolph Schiner|Schiner]], 1868</small><ref name="Schiner1868"/>
*''[[Dischistus]]'' <small>[[Hermann Loew|Loew]], 1855</small>
*''[[Dischistus]]'' <small>[[Hermann Loew|Loew]], 1855</small>
*''[[Docidomyia]]'' <small>White, 1916</small>
*''[[Docidomyia]]'' <small>White, 1916</small>
Line 252: Line 259:
*''[[Enica]]'' <small>[[Pierre-Justin-Marie Macquart|Macquart]], 1834</small>
*''[[Enica]]'' <small>[[Pierre-Justin-Marie Macquart|Macquart]], 1834</small>
*''[[Epacmoides]]'' <small>Hesse, 1956</small>
*''[[Epacmoides]]'' <small>Hesse, 1956</small>
*''[[Epacmus]]'' <small>[[Karl Robert Osten-Sacken|Osten Sacken]], 1886</small>
*''[[Epacmus]]'' <small>[[Carl Robert Osten-Sacken|Osten Sacken]], 1886</small>
*''[[Eremyia]]'' <small>Greathead, 1996\</small>
*''[[Eremyia]]'' <small>Greathead, 1996\</small>
*''[[Eristalopsis]]'' <small>Evenhuis, 1985</small>
*''[[Eristalopsis]]'' <small>Evenhuis, 1985</small>
Line 266: Line 273:
*''[[Exoprosopa]]'' <small>[[Pierre-Justin-Marie Macquart|Macquart]], 1840</small>
*''[[Exoprosopa]]'' <small>[[Pierre-Justin-Marie Macquart|Macquart]], 1840</small>
*''[[Geminaria (fly)|Geminaria]]'' <small>[[Daniel William Coquillett|Coquillett]], 1894</small>
*''[[Geminaria (fly)|Geminaria]]'' <small>[[Daniel William Coquillett|Coquillett]], 1894</small>
*''[[Geron (insect)|Geron]]'' <small>[[Johann Wilhelm Meigen|Meigen]], 1820</small>
*''[[Geron (fly)|Geron]]'' <small>[[Johann Wilhelm Meigen|Meigen]], 1820</small>
*''[[Glaesamictus]]'' <small>[[Willi Hennig|Hennig]], 1966</small>
*''[[Glaesamictus]]'' <small>[[Willi Hennig|Hennig]], 1966</small>
*''[[Gnumyia]]'' <small>[[Mario Bezzi|Bezzi]], 1921</small>
*''[[Gnumyia]]'' <small>[[Mario Bezzi|Bezzi]], 1921</small>
Line 273: Line 280:
*''[[Hallidia]]'' <small>Hull, 1970</small>
*''[[Hallidia]]'' <small>Hull, 1970</small>
*''[[Hemipenthes]]'' <small>[[Hermann Loew|Loew]], 1869</small>
*''[[Hemipenthes]]'' <small>[[Hermann Loew|Loew]], 1869</small>
*''[[Heteralonia]]'' <small>[[Camillo Róndani|Róndani]], 1863</small>
*''[[Heteralonia]]'' <small>[[Camillo Rondani|Rondani]], 1863</small>
*''[[Heterostylum]]'' <small>[[Pierre-Justin-Marie Macquart|Macquart]], 1848</small>
*''[[Heterostylum]]'' <small>[[Pierre-Justin-Marie Macquart|Macquart]], 1848</small>
*''[[Heterotropus]]'' <small>[[Hermann Loew|Loew]], 1873</small>
*''[[Heterotropus]]'' <small>[[Hermann Loew|Loew]], 1873</small>
*''[[Hyperalonia]]'' <small>[[Camillo Róndani|Róndani]], 1863</small>
*''[[Hyperalonia]]'' <small>[[Camillo Rondani|Rondani]], 1863</small>
*''[[Hyperusia]]'' <small>[[Mario Bezzi|Bezzi]], 1921</small>
*''[[Hyperusia]]'' <small>[[Mario Bezzi|Bezzi]], 1921</small>
*''[[Inyo (genus)|Inyo]]'' <small>Hall & Evenhuis, 1987</small>
*''[[Inyo (fly)|Inyo]]'' <small>Hall & Evenhuis, 1987</small>
*''[[Isocnemus]]'' <small>[[Mario Bezzi|Bezzi]], 1924</small>
*''[[Isocnemus]]'' <small>[[Mario Bezzi|Bezzi]], 1924</small>
*''[[Kapu (genus)|Kapu]]'' <small>Lambkin & Yeates, 2003</small>
*''[[Kapu (fly)|Kapu]]'' <small>Lambkin & Yeates, 2003</small>
*''[[Karakumia]]'' <small>Paramonov, 1927</small>
*''[[Karakumia]]'' <small>Paramonov, 1927</small>
*''[[Laminanthrax]]'' <small>Greathead, 1967</small>
*''[[Laminanthrax]]'' <small>Greathead, 1967</small>
Line 286: Line 293:
*''[[Laurella]]'' <small>Hull, 1971</small>
*''[[Laurella]]'' <small>Hull, 1971</small>
*''[[Legnotomyia]]'' <small>[[Mario Bezzi|Bezzi]], 1902</small>
*''[[Legnotomyia]]'' <small>[[Mario Bezzi|Bezzi]], 1902</small>
*''[[Lepidanthrax]]'' <small>[[Karl Robert Osten-Sacken|Osten Sacken]], 1886</small>
*''[[Lepidanthrax]]'' <small>[[Carl Robert Osten-Sacken|Osten Sacken]], 1886</small>
*''[[Lepidochlanus]]'' <small>Hesse, 1938</small>
*''[[Lepidochlanus]]'' <small>Hesse, 1938</small>
*''[[Lepidophora]]'' <small>[[John Obadiah Westwood|Westwood]], 1835</small>
*''[[Lepidophora]]'' <small>[[John Obadiah Westwood|Westwood]], 1835</small>
Line 293: Line 300:
*''[[Lomatia belzebul|Lomatia]]'' <small>[[Johann Wilhelm Meigen|Meigen]], 1822</small>
*''[[Lomatia belzebul|Lomatia]]'' <small>[[Johann Wilhelm Meigen|Meigen]], 1822</small>
*''[[Lordotus]]'' <small>[[Hermann Loew|Loew]], 1863</small>
*''[[Lordotus]]'' <small>[[Hermann Loew|Loew]], 1863</small>
*''[[Macrocondyla]]'' <small>[[Camillo Róndani|Róndani]], 1863</small>
*''[[Macrocondyla]]'' <small>[[Camillo Rondani|Rondani]], 1863</small>
*''[[Mallophthiria]]'' <small>[[Frederick Wallace Edwards|Edwards]], 1930</small>
*''[[Mallophthiria]]'' <small>[[Frederick Wallace Edwards|Edwards]], 1930</small>
*''[[Mancia (genus)|Mancia]]'' <small>[[Daniel William Coquillett|Coquillett]], 1886</small>
*''[[Mancia (fly)|Mancia]]'' <small>[[Daniel William Coquillett|Coquillett]], 1886</small>
*''[[Mandella (genus)|Mandella]]'' <small>Evenhuis, 1983</small>
*''[[Mandella (fly)|Mandella]]'' <small>Evenhuis, 1983</small>
*''[[Mariobezzia]]'' <small>[[Theodor Becker|Becker]], 1913</small>
*''[[Mariobezzia]]'' <small>[[Theodor Becker|Becker]], 1913</small>
*''[[Marleyimyia]]'' <small>Hesse, 1956</small>
*''[[Marleyimyia]]'' <small>Hesse, 1956</small>
Line 316: Line 323:
*''[[Neodischistus]]'' <small>Painter, 1933</small>
*''[[Neodischistus]]'' <small>Painter, 1933</small>
*''[[Neosardus]]'' <small>Roberts, 1929</small>
*''[[Neosardus]]'' <small>Roberts, 1929</small>
*''[[Nomalonia]]'' <small>[[Camillo Róndani|Róndani]], 1863</small>
*''[[Nomalonia]]'' <small>[[Camillo Rondani|Rondani]], 1863</small>
*''[[Nothoschistus]]'' <small>Bowden, 1985</small>
*''[[Nothoschistus]]'' <small>Bowden, 1985</small>
*''[[Notolomatia]]'' <small>Greathead, 1998</small>
*''[[Notolomatia]]'' <small>Greathead, 1998</small>
Line 332: Line 339:
*''[[Palintonus]]'' <small>François, 1964</small>
*''[[Palintonus]]'' <small>François, 1964</small>
*''[[Palirika]]'' <small>Lambkin & Yeates, 2003</small>
*''[[Palirika]]'' <small>Lambkin & Yeates, 2003</small>
*''[[Pantarbes]]'' <small>[[Karl Robert Osten-Sacken|Osten Sacken]], 1877</small>
*''[[Pantarbes]]'' <small>[[Carl Robert Osten-Sacken|Osten Sacken]], 1877</small>
*''[[Pantostomus]]'' <small>[[Mario Bezzi|Bezzi]], 1921</small>
*''[[Pantostomus]]'' <small>[[Mario Bezzi|Bezzi]], 1921</small>
*''[[Paracorsomyza]]'' <small>[[Willi Hennig|Hennig]], 1966</small>
*''[[Paracorsomyza]]'' <small>[[Willi Hennig|Hennig]], 1966</small>
*''[[Paradiplocampta]]'' <small>Hall, 1975</small>
*''[[Paradiplocampta]]'' <small>Hall, 1975</small>
*''[[Parachistus]]'' <small>Greathead, 1980</small>
*''[[Parachistus]]'' <small>Greathead, 1980</small>
*''[[Paracosmus]]'' <small>[[Karl Robert Osten-Sacken|Osten Sacken]], 1877</small>
*''[[Paracosmus]]'' <small>[[Carl Robert Osten-Sacken|Osten Sacken]], 1877</small>
*''[[Parageron]]'' <small>Paramonov, 1929</small>
*''[[Parageron]]'' <small>Paramonov, 1929</small>
*''[[Paramonovius]]'' <small>Li & Yeates, 2018</small>
*''[[Paramonovius]]'' <small>Li & Yeates, 2018</small>
Line 352: Line 359:
*''[[Platamomyia]]'' <small>Brèthes, 1925</small>
*''[[Platamomyia]]'' <small>Brèthes, 1925</small>
*''[[Plesiocera]]'' <small>[[Pierre-Justin-Marie Macquart|Macquart]], 1840</small>
*''[[Plesiocera]]'' <small>[[Pierre-Justin-Marie Macquart|Macquart]], 1840</small>
*''[[Poecilanthrax]]'' <small>[[Karl Robert Osten-Sacken|Osten Sacken]], 1886</small>
*''[[Poecilanthrax]]'' <small>[[Carl Robert Osten-Sacken|Osten Sacken]], 1886</small>
*''[[Poecilognathus]]'' <small>[[Johann Friedrich Jaennicke|Jaennicke]], 1867</small>
*''[[Poecilognathus]]'' <small>[[Johann Friedrich Jaennicke|Jaennicke]], 1867</small>
*''[[Praecytherea]]'' <small>Théobald, 1937</small>
*''[[Praecytherea]]'' <small>Théobald, 1937</small>
Line 381: Line 388:
*''[[Staurostichus]]'' <small>Hull, 1973</small>
*''[[Staurostichus]]'' <small>Hull, 1973</small>
*''[[Stomylomyia]]'' <small>[[Jacques-Marie-Frangile Bigot|Bigot]], 1888</small>
*''[[Stomylomyia]]'' <small>[[Jacques-Marie-Frangile Bigot|Bigot]], 1888</small>
*''[[Stonyx]]'' <small>[[Karl Robert Osten-Sacken|Osten Sacken]], 1886</small>
*''[[Stonyx]]'' <small>[[Carl Robert Osten-Sacken|Osten Sacken]], 1886</small>
*''[[Synthesia (fly)|Synthesia]]'' <small>[[Mario Bezzi|Bezzi]], 1921</small>
*''[[Synthesia (fly)|Synthesia]]'' <small>[[Mario Bezzi|Bezzi]], 1921</small>
*''[[Systoechus]]'' <small>[[Hermann Loew|Loew]], 1855</small>
*''[[Systoechus]]'' <small>[[Hermann Loew|Loew]], 1855</small>
Line 387: Line 394:
*''[[Thevenetimyia]]'' <small>[[Jacques-Marie-Frangile Bigot|Bigot]], 1875</small>
*''[[Thevenetimyia]]'' <small>[[Jacques-Marie-Frangile Bigot|Bigot]], 1875</small>
*''[[Thraxan]]'' <small>Yeates & Lambkin, 1998</small>
*''[[Thraxan]]'' <small>Yeates & Lambkin, 1998</small>
*''[[Thyridanthrax]]'' <small>[[Karl Robert Osten-Sacken|Osten Sacken]], 1886</small>
*''[[Thyridanthrax]]'' <small>[[Carl Robert Osten-Sacken|Osten Sacken]], 1886</small>
*''[[Tillyardomyia]]'' <small>Tonnoir, 1927</small>
*''[[Tillyardomyia]]'' <small>Tonnoir, 1927</small>
*''[[Timiomyia]]'' <small>Evenhuis, 1978</small>
*''[[Timiomyia]]'' <small>Evenhuis, 1978</small>
Line 401: Line 408:
*''[[Veribubo]]'' <small>Evenhuis, 1978</small>
*''[[Veribubo]]'' <small>Evenhuis, 1978</small>
*''[[Verrallites]]'' <small>Cockerell, 1913</small>
*''[[Verrallites]]'' <small>Cockerell, 1913</small>
*''[[Villa (genus)|Villa]]'' <small>Lioy, 1864</small>
*''[[Villa (fly)|Villa]]'' <small>Lioy, 1864</small>
*''[[Villoestrus]]'' <small>Paramonov, 1931</small>
*''[[Villoestrus]]'' <small>Paramonov, 1931</small>
*''[[Walkeromyia]]'' <small>Paramonov, 1934</small>
*''[[Walkeromyia]]'' <small>Paramonov, 1934</small>
Line 424: Line 431:
Image:Bombyliid 7294.jpg|''[[Lepidophora]]'' on ''[[Bidens|Bidens laevis]]''
Image:Bombyliid 7294.jpg|''[[Lepidophora]]'' on ''[[Bidens|Bidens laevis]]''
</gallery>
</gallery>

==In Popular Culture==
* The [[Pokémon]] [[Ribombee]], introduced in [[Pokémon Sun and Moon]], is a Bug and Fairy type with a design heavily based on a beefly.


== References ==
== References ==
Line 429: Line 439:
<ref name=Evenhuis2015>
<ref name=Evenhuis2015>
{{Cite web
{{Cite web
| accessdate = 2018-12-31
| access-date = 2018-12-31
| title = World catalog of bee flies (Diptera: Bombyliidae)
| title = World catalog of bee flies (Diptera: Bombyliidae)
| date = 2015
| date = 2015
Line 440: Line 450:
*Engel, E.O., 1932-1937. ''Bombyliidae''. In: ''Die Fliegen der paläarktischen Region'' 4(3) ( [[Erwin Lindner]], ed.): 1-619, pl. 1-15. E. Schweizerbart, Stuttgart.). Old and outdated, not easy to get and expensive but some of the only keys to taxa in the [[Palaearctic]] Region.
*Engel, E.O., 1932-1937. ''Bombyliidae''. In: ''Die Fliegen der paläarktischen Region'' 4(3) ( [[Erwin Lindner]], ed.): 1-619, pl. 1-15. E. Schweizerbart, Stuttgart.). Old and outdated, not easy to get and expensive but some of the only keys to taxa in the [[Palaearctic]] Region.
*Greathead & Evenhuis (Greathead, D.J., & N.L. Evenhuis, 1997. Family Bombyliidae. In: ''Contributions to a manual of Palaearctic Diptera'' Volume 2 (L. Papp & B. Darvas, eds.): 487-512. Science Herald, Budapest.) provide a key to the Palaearctic genera and (may) give references to available generic revisions.
*Greathead & Evenhuis (Greathead, D.J., & N.L. Evenhuis, 1997. Family Bombyliidae. In: ''Contributions to a manual of Palaearctic Diptera'' Volume 2 (L. Papp & B. Darvas, eds.): 487-512. Science Herald, Budapest.) provide a key to the Palaearctic genera and (may) give references to available generic revisions.
*{{cite journal |author=Evenhuis N.L. |year=1991 |title=Catalog of genus-group names of bee flies (Diptera: Bombyliidae) |url=|journal=Bishop Museum Bulletins in Entomology |volume=5 |issue=|pages=1–105 }}
*{{cite journal |author=Evenhuis N.L. |year=1991 |title=Catalog of genus-group names of bee flies (Diptera: Bombyliidae) |journal=Bishop Museum Bulletins in Entomology |volume=5 |pages=1–105 }}
*Evenhuis, N.L. & Greathead, D.J. 1999. World catalog of bee flies (Diptera: Bombyliidae). Backhuys Publishers, Leiden, 756 pp. [http://hbs.bishopmuseum.org/bombcat/ online]
*Evenhuis, N.L. & Greathead, D.J. 1999. World catalog of bee flies (Diptera: Bombyliidae). Backhuys Publishers, Leiden, 756 pp. [http://hbs.bishopmuseum.org/bombcat/ online]
*Hull, F.M. 1973. ''Bee flies of the world. The genera of the family Bombyliidae''.Washington (Smithsonian Institution Press) 687 pp. Keys subfamilies, genera (many generic placements superseded by Evenhuis & Greathead, 1999).
*Hull, F.M. 1973. ''Bee flies of the world. The genera of the family Bombyliidae''.Washington (Smithsonian Institution Press) 687 pp. Keys subfamilies, genera (many generic placements superseded by Evenhuis & Greathead, 1999).
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[[Category:Brachycera families]]
[[Category:Brachycera families]]
[[Category:Pollinator flies]]
[[Category:Insects used as insect pest control agents]]
[[Category:Insects used as insect pest control agents]]
[[Category:Articles containing video clips]]
[[Category:Articles containing video clips]]

Latest revision as of 18:36, 6 November 2024

Bombyliidae
Temporal range: Cenomanian–Recent
Bombylius major
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Diptera
Superfamily: Asiloidea
Family: Bombyliidae
Latreille, 1802
Subfamilies
Synonyms

Phthiriidae
Usiidae
Systropodidae

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 pollinators. Larvae are mostly parasitoids of other insects.

Overview

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The Bombyliidae are a large family of flies comprising hundreds of genera, but the life cycles 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 proboscises 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. Many Bombyliidae superficially resemble bees and accordingly the prevalent common name for a member of the family is bee fly.[2] Possibly the resemblance is Batesian mimicry, affording the adults some protection from predators.

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

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. The family has a patchy fossil record, with species being known from a handful of localities,[3] the oldest known species are known from the Middle Cretaceous Burmese amber, around 99 million years old.[4]

Euchariomyia dives, a Bombyliidae in 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

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Adult

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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.

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 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 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.

A male of 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 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 1 Bombylius
  • Wing venation type 2 Anthrax
    Wing venation type 2 Anthrax
  • Wing venation type 3 Usiinae
    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

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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

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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.[5] 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.[6] 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.[7][8] 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.

Villa sp. gathering sand grains
Lepidophora lepidocera, a Nearctic realm species

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.[9]

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.

Zoogeography

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The family is worldwide (Palearctic realm, Nearctic realm, Afrotropical realm, Neotropical realm, Australasian realm, Oceanian realm, Indomalayan realm), but has the greatest biodiversity in tropical and subtropical arid climates. In Europe, 335 species are distributed among 53 genera.

Species lists

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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

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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 such as a slender abdomen and white patches on the wings

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[10] Boris Rohdendorf (1974) dealt with the family in a separate superfamily (Bombyliidea), linking it to the superfamily of Asilidea.[11] 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.

Clade showing relationships within Asiloidea
Asiloidea  
 N.N. 

 Bombyliidae

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)[12] finally moved the genus Hilarimorpha into their own family (Hilarimorphidae). Zaitzev (1991)[13] moved the genus Mythicomyia and several other minor genera in the family Mythicomyiidae, Yeates (1992, 1994)[14] 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.[15] 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.[16]

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:[17]

Genera

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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
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References

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  1. ^ Alan Weaving; Mike Picker; Griffiths, Charles Llewellyn (2003). Field Guide to Insects of South Africa. New Holland Publishers, Ltd. ISBN 1-86872-713-0.
  2. ^ a b Hull, Frank Montgomery, Bee flies of the world: the genera of the family Bombyliidae Washington, Smithsonian Institution Press 1973 ISBN 0-87474-131-9. Downloadable from: https://archive.org/details/beefliesofworl2861973hull
  3. ^ Wedmann, Sonja; Yeates, David K. (2008-01-17). "Eocene Records of Bee Flies (Insecta, Diptera, Bombyliidae, Comptosia): Their Palaeobiogeographic Implications and Remarks on the Evolutionary History of Bombyliids". Palaeontology. 51 (1): 231–240. Bibcode:2008Palgy..51..231W. doi:10.1111/j.1475-4983.2007.00745.x. S2CID 128576916.
  4. ^ Ye, Xiuna; Yao, Gang; Shih, Chungkun; Ren, Dong; Wang, Yongjie (August 2019). "New bee flies from the mid-Cretaceous Myanmar amber (Brachycera: Asiloidea: Bombyliidae)". Cretaceous Research. 100: 5–13. Bibcode:2019CrRes.100....5Y. doi:10.1016/j.cretres.2019.03.026. S2CID 135365568.
  5. ^ 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.
  6. ^ Archived at Ghostarchive and the Wayback Machine: Roy Kleukers (22 June 2013). "Gewone wolzwever Bombylius major, eieren droppend in een kolonie zandbijen" – via YouTube.
  7. ^ Yeates David K (1997). "The evolutionary pattern of host use in the Bombyliidae (Diptera): a diverse family of parasitoid flies". Biological Journal of the Linnean Society. 60 (2): 149–185. doi:10.1111/j.1095-8312.1997.tb01490.x.
  8. ^ 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 Bee Flies (Diptera: Bombyliidae). Zoological Studies 48: 141-150.
  9. ^ The evolutionary pattern of host use in the Bombyliidae (Diptera): a diverse family of parasitoid flies
  10. ^ 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.
  11. ^ Boris B. Rohdendorf, Brian Hocking, Harold Oldroyd, George E. Ball. The Historical Development of Diptera. University of Alberta, 1974: 75-77. ISBN 088864003X.
  12. ^ Webb D.W., 1981 Hilarimorphidae. in: McAlpine J.F. (Ed.), Manual of Nearctic Diptera. Agriculture Canada, Ottawa, pp. 603-605.
  13. ^ Zaitzev, V.F., 1991 On the phylogeny and systematics of the dipteran superfamily Bombylioidea (Diptera). Entomol. Obozr. 70 [1991] : 716–36.
  14. ^ Yeates D.M. (1992). "Towards a monophyletic Bombyliidae (Diptera): the removal of the Proratinae (Diptera: Scenopinidae)". American Museum Novitates (3051): 1–30.
  15. ^ Yeates & Lambkin, The Tree of Life, op. cit.
  16. ^ Trautwein, Michelle D.; Wiegmann, Brian M.; Yeates, David K. (2011). "Overcoming the effects of rogue taxa: Evolutionary relationships of the bee flies". PLOS Currents. 3: RRN1233. doi:10.1371/currents.RRN1233 (inactive 1 November 2024). PMC 3088465. PMID 21686308.{{cite journal}}: CS1 maint: DOI inactive as of November 2024 (link)
  17. ^ Evenhuis, N.L.; Greathead, D.J. (2015). "World catalog of bee flies (Diptera: Bombyliidae)". Retrieved 2018-12-31.
  18. ^ a b Schiner, I.R. (1868). Diptera. vi In [Wullerstorf-Urbair, B. von (in charge)], Reise der osterreichischen Fregatte Novara. Zool. 2(1)B. Wien: K. Gerold's Sohn. pp. 388pp., 4 pls.
  • Bowden, J.,1980 Family Bombyliidae. pp. 381–430. In R.W. Crosskey (ed.), Catalogue of the Diptera of the Afrotropical Region, 1437 pp., London: British Museum (Natural History)
  • Engel, E.O., 1932-1937. Bombyliidae. In: Die Fliegen der paläarktischen Region 4(3) ( Erwin Lindner, ed.): 1-619, pl. 1-15. E. Schweizerbart, Stuttgart.). Old and outdated, not easy to get and expensive but some of the only keys to taxa in the Palaearctic Region.
  • Greathead & Evenhuis (Greathead, D.J., & N.L. Evenhuis, 1997. Family Bombyliidae. In: Contributions to a manual of Palaearctic Diptera Volume 2 (L. Papp & B. Darvas, eds.): 487-512. Science Herald, Budapest.) provide a key to the Palaearctic genera and (may) give references to available generic revisions.
  • Evenhuis N.L. (1991). "Catalog of genus-group names of bee flies (Diptera: Bombyliidae)". Bishop Museum Bulletins in Entomology. 5: 1–105.
  • Evenhuis, N.L. & Greathead, D.J. 1999. World catalog of bee flies (Diptera: Bombyliidae). Backhuys Publishers, Leiden, 756 pp. online
  • Hull, F.M. 1973. Bee flies of the world. The genera of the family Bombyliidae.Washington (Smithsonian Institution Press) 687 pp. Keys subfamilies, genera (many generic placements superseded by Evenhuis & Greathead, 1999).
  • Yeates, David K. 1994. The cladistics and classification of the Bombyliidae (Diptera: Asiloidea). Bulletin of the American Museum of Natural History ; no. 219, 191 pp.
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