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{{short description|Superfamily of flies}}
{{Taxobox
{{Automatic taxobox|taxon=Empidoidea|authority=|image=Tanzfliege Dance fly.jpg|image_caption=''[[Empis tessellata]]''|subdivision_ranks=[[Family (biology)|Families]]|subdivision=see [[#Systematics|text]]}}
| name = Empidoidea
[[File:Male and female genetalia in Empidoidea.jpg|thumb|Pictures of posterior part of abdomen. (A–E) male genitalia in dorsal view, ''Atelestus pulicarius'' (A), ''Neurigona suturalis'' (B), ''Empis vitripennis'' (C), ''Hybos grossipes'' (D), ''Ragas unica'' (E); (F, G) male genitalia in dorsal view, ''Clinocera nivalis'' (F), ''E. vitripennis'' (G); (H–J), female genitalia in dorsal view, ''C. nivalis'' (H), ''E. vitripennis'' (I), ''Trichopeza longicornis'' (J).]]
| image = Empis sp.jpg
[[File:Wing structures of several Empidoidea species.png|thumb|Photographs of right wing of several Empidoidea species. Abbreviations: h, humeral crossvein; R<sub>s</sub>, radial sector; r‐m, radial‐medial crossvein; M<sub>1</sub>, first medial vein; M<sub>1+2</sub>, first and second medial vein (unbranched); R<sub>4</sub>, fourth radial vein; R<sub>5</sub>, fifth radial vein; R<sub>4+5</sub>, fourth and fifth radial vein (unbranched); A<sub>1</sub>, anal vein; CuA<sub>2</sub>, second anterior branch of cubital vein; bm‐cu, basal medial‐cubital crossvein; dm, discal medial cell; bm, basal medial cell; cu''p'', posterior cubital cell (anal cell).]]
| image_width = 220px
The '''Empidoidea''' are a large [[monophyletic]] [[Taxonomic rank|superfamily]] of [[Diptera|true flies]], the [[sister taxon]] to the [[Muscomorpha]] ([[Cyclorrhapha]]). These two groups are sometimes united in the unranked taxon [[Eremoneura]]. There are some 10,000 known [[species]] within Empidoidea,<ref name="m+w2007">Moulton & Wiegmann (2007)</ref> which are represented on all continents except Antarctica. They are known to have existed since the Jurassic period.<ref name="g+e2005">Grimaldi & Engel (2005)</ref>
| image_caption = ''[[Empis]]'' sp.
| regnum = [[Animal]]ia
| phylum = [[Arthropod]]a
| classis = [[Insect]]a
| ordo = [[Diptera]]
| subordo = [[Brachycera]]
| infraordo = [[Asilomorpha]]
| superfamilia = '''Empidoidea'''
| superfamilia_authority =
| subdivision_ranks = [[Family (biology)|Families]]
| subdivision =
4, see [[#Systematics|text]]
}}


Empidoidea has been subject to much debate regarding its phylogeny. Based on morphology alone, three major hypotheses had been proposed until 2007 and seemed to be consensus for some time—however, in 2018, Wahlberg & Johanson published the most current phylogeny supported by extensive genetic data, changing the relationship between quite a few families and tribes.<ref name="w+j2018">Wahlberg & Johanson (2018)</ref>
The '''Empidoidea''' are a large [[monophyletic]] [[superfamily]] of [[Diptera|true flies]], the [[sister taxon]] to the [[Muscomorpha]] (= "[[Cyclorrhapha]]"). These two groups are sometimes united in the unranked taxon [[Eremoneura]]. There are some 10,000 known [[species]] in the Empidoidea<ref name = m+w2007>Moulton & Wiegmann (2007)</ref>.


==Description==
The majority of these insects are [[predator]]y, often with large [[compound eye]]s (sometimes covering almost the entire surface of the head), and tend to be associated with moist, [[temperate]] habitats. Many have a well-developed [[anal lobe]] in the wings, and/or a small but distinct [[anal cell]].
The majority of these insects are [[predator]]y, often with large [[compound eye]]s (sometimes covering almost the entire surface of the head), and tend to be associated with moist, [[temperate]] habitats. Many have a well-developed [[anal lobe]] in the wings, and/or a small but distinct [[anal cell]].

Several species within Empidoidea are of particular interest to ethologists and ecologists, due to elaborate mating rituals and life histories involving freshwater.<ref name="w+j2018" />[[File:Empidoidea.jpg|thumb|''Empidoidea sp.''|alt=|none]]


==Systematics==
==Systematics==
The most familiar [[Family (biology)|families]] in the group are the [[Empididae]] and [[Dolichopodidae]], both of which occur worldwide and contain thousands of [[species]]. The smaller families were formerly included in the Empididae, but have since been accorded family status<ref>Sinclair & Cumming (2006)</ref>. The [[Brachystomatinae]] and [[Microphoridae]] are sometimes elevated to full-blown families too, but this would apparently make the Empididae and Dolichopodidae [[paraphyletic]] and thus they are retained as subfamilies here.
The most familiar [[Family (biology)|families]] in the group are the [[Empididae]] and [[Dolichopodidae]], both of which occur worldwide and contain thousands of [[species]]. The smaller families were formerly included in the Empididae, but have since been accorded family status.<ref>Sinclair & Cumming (2006)</ref> The monophyly of most of these groups have since been confirmed by subsequent studies.<ref name="w+j2018" /> The [[Brachystomatinae]] and [[Microphorinae]] are sometimes elevated to full family status, too, but this would apparently make the Empididae and Dolichopodidae [[paraphyletic]] and thus they are retained as subfamilies here. [[Atelestidae]] forming the sister group to the rest of Empidoidea is a unanimous result in all studies that have utilized molecular data.<ref name="w+j2018" />


The families are:<ref name="m+w2007" /><ref name="w+j2018" />
It seems that the present systematic treatment essentially reflects [[phylogeny]] rather well; there are some of the more [[basal (evoution)|basal]] [[taxa]] in each lineage which are of more uncertain position in [[cladistic]] analyses, but at least as an interim solution, the four-family approach seems most warranted. Arranged in phylogenetic sequence, the families are:<ref name = m+w2007 />

* [[Atelestidae]]
* [[Hybotidae]]
*[[Atelestidae]]
*[[Hybotidae]] &ndash; dance flies
* [[Dolichopodidae]] (including Microphoridae)
* [[Empididae]] (including Brachystomatinae)
*[[Dolichopodidae]] &ndash; long-legged flies (including Microphoridae)
*[[Empididae]] &ndash; dagger flies and balloon flies (including Brachystomatidae)
*[[Homalocnemiidae]]
*[[Oreogetonidae]]
*[[Ragadidae]]

Based on the most recent phylogenetic studies,<ref name=w+j2018/> the internal classification of Empidoidea is as follows:

{{clade
|label1=[[Atelestidae]]
|1={{clade
|1=[[Atelestinae]]
|2=[[Nemedininae]]
}}
|2={{clade
|label1=[[Hybotidae]]
|1={{clade
|1=[[Bicellariinae]]
|2={{clade
|label1=[[Tachydromiinae]]
|1={{clade
|1=[[Drapetini]]
|2={{clade
|1=[[Symballaphthalmini]]
|2=[[Tachydromiini]]
}}
}}
|2={{clade
|1=[[Trichininae]]
|2={{clade
|1=[[Ocydromiinae]]
|2={{clade
|1=[[Hybotinae]]
|2=[[Oedaleinae]]
}}
}}
}}
}}
}}
|2={{clade
|1=[[Dolichopodidae]]
|2={{clade
|label1=[[Ragadidae]]
|1={{clade
|1=[[Iteaphilinae]]
|2=[[Ragadinae]]
}}
|label2=[[Empididae]]
|2={{clade
|1=[[Clinocerinae]]
|2={{clade
|label1=[[Brachystomatinae]]
|1={{clade
|1=[[Brachystomatini]]
|2=[[Ceratomerini]]
|3=[[Trichopezini]]
}}
|label2=[[Empidinae]]
|2={{clade
|1={{clade
|1=[[Chelipodini]]
|2=[[Empidini]]
}}
|2={{clade
|1=[[Hemerodromiini]]
|2=[[Hilarini]]
}}
}}
}}
}}
}}
}}
}}
}}


==Footnotes==
==Footnotes==
Line 34: Line 98:


==References==
==References==
* {{aut|Moulton, J.K. & Wiegmann, B.M.}} (2007): The phylogenetic relationships of flies in the superfamily Empidoidea (Insecta: Diptera). ''[[Molecular Phylogenetics and Evolution|Mol. Phylogenet. Evol.]]'' '''43'''(3): 701-713. <small>{{doi|10.1016/j.ympev.2007.02.029}}</small> (HTML abstract)
* {{aut|Moulton, J.K. & Wiegmann, B.M.}} (2007): The phylogenetic relationships of flies in the superfamily Empidoidea (Insecta: Diptera). ''[[Molecular Phylogenetics and Evolution|Mol. Phylogenet. Evol.]]'' '''43'''(3): 701–713. <small>{{doi|10.1016/j.ympev.2007.02.029}} {{PMID|17468014}}</small> (HTML abstract)
*{{aut|Grimaldi, D. & Engel, M.S.}} (2005): Evolution of the Insects. Cambridge University Press, Cambridge.
* {{aut|Sinclair, B.J. & Cumming, J.M.}} (2006): The morphology, higher-level phylogeny and classification of the Empidoidea (Diptera). ''[[Zootaxa]]'' '''1180''': 1-172
* {{aut|Sinclair, B.J. & Cumming, J.M.}} (2006): The morphology, higher-level phylogeny and classification of the Empidoidea (Diptera). ''[[Zootaxa]]'' '''1180''': 1–172. [http://www.uoguelph.ca/nadsfly/Doid/Doidclass/SinclairCumming2006.pdf PDF fulltext]

*{{aut|Wahlberg, E. & Johanson, K.A.}} (2018): Molecular phylogenetics reveals novel relationships within Empidoidea (Diptera). ''[[Systematic Entomology]]'' '''43'''(4): 619–636. <small>{{doi|10.1111/syen.12297}} [[File:CC-BY icon.svg|50px]] Material was copied from this source, which is available under a [https://creativecommons.org/licenses/by/4.0/ Creative Commons Attribution 4.0 International License].</small>

{{fly-stub}}


{{Diptera|2}}
[[Category:Flies]]
{{Taxonbar|from=Q1644617}}


[[no:Empidoidea]]
[[Category:Empidoidea| ]]
[[Category:Diptera superfamilies]]

Latest revision as of 22:38, 2 November 2024

Empidoidea
Empis tessellata
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Diptera
Infraorder: Asilomorpha
Superfamily: Empidoidea
Families

see text

Pictures of posterior part of abdomen. (A–E) male genitalia in dorsal view, Atelestus pulicarius (A), Neurigona suturalis (B), Empis vitripennis (C), Hybos grossipes (D), Ragas unica (E); (F, G) male genitalia in dorsal view, Clinocera nivalis (F), E. vitripennis (G); (H–J), female genitalia in dorsal view, C. nivalis (H), E. vitripennis (I), Trichopeza longicornis (J).
Photographs of right wing of several Empidoidea species. Abbreviations: h, humeral crossvein; Rs, radial sector; r‐m, radial‐medial crossvein; M1, first medial vein; M1+2, first and second medial vein (unbranched); R4, fourth radial vein; R5, fifth radial vein; R4+5, fourth and fifth radial vein (unbranched); A1, anal vein; CuA2, second anterior branch of cubital vein; bm‐cu, basal medial‐cubital crossvein; dm, discal medial cell; bm, basal medial cell; cup, posterior cubital cell (anal cell).

The Empidoidea are a large monophyletic superfamily of true flies, the sister taxon to the Muscomorpha (Cyclorrhapha). These two groups are sometimes united in the unranked taxon Eremoneura. There are some 10,000 known species within Empidoidea,[1] which are represented on all continents except Antarctica. They are known to have existed since the Jurassic period.[2]

Empidoidea has been subject to much debate regarding its phylogeny. Based on morphology alone, three major hypotheses had been proposed until 2007 and seemed to be consensus for some time—however, in 2018, Wahlberg & Johanson published the most current phylogeny supported by extensive genetic data, changing the relationship between quite a few families and tribes.[3]

Description

[edit]

The majority of these insects are predatory, often with large compound eyes (sometimes covering almost the entire surface of the head), and tend to be associated with moist, temperate habitats. Many have a well-developed anal lobe in the wings, and/or a small but distinct anal cell.

Several species within Empidoidea are of particular interest to ethologists and ecologists, due to elaborate mating rituals and life histories involving freshwater.[3]

Empidoidea sp.

Systematics

[edit]

The most familiar families in the group are the Empididae and Dolichopodidae, both of which occur worldwide and contain thousands of species. The smaller families were formerly included in the Empididae, but have since been accorded family status.[4] The monophyly of most of these groups have since been confirmed by subsequent studies.[3] The Brachystomatinae and Microphorinae are sometimes elevated to full family status, too, but this would apparently make the Empididae and Dolichopodidae paraphyletic and thus they are retained as subfamilies here. Atelestidae forming the sister group to the rest of Empidoidea is a unanimous result in all studies that have utilized molecular data.[3]

The families are:[1][3]

Based on the most recent phylogenetic studies,[3] the internal classification of Empidoidea is as follows:

Atelestidae

Footnotes

[edit]
  1. ^ a b Moulton & Wiegmann (2007)
  2. ^ Grimaldi & Engel (2005)
  3. ^ a b c d e f Wahlberg & Johanson (2018)
  4. ^ Sinclair & Cumming (2006)

References

[edit]
  • Moulton, J.K. & Wiegmann, B.M. (2007): The phylogenetic relationships of flies in the superfamily Empidoidea (Insecta: Diptera). Mol. Phylogenet. Evol. 43(3): 701–713. doi:10.1016/j.ympev.2007.02.029 PMID 17468014 (HTML abstract)
  • Grimaldi, D. & Engel, M.S. (2005): Evolution of the Insects. Cambridge University Press, Cambridge.
  • Sinclair, B.J. & Cumming, J.M. (2006): The morphology, higher-level phylogeny and classification of the Empidoidea (Diptera). Zootaxa 1180: 1–172. PDF fulltext
  • Wahlberg, E. & Johanson, K.A. (2018): Molecular phylogenetics reveals novel relationships within Empidoidea (Diptera). Systematic Entomology 43(4): 619–636. doi:10.1111/syen.12297 Material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
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