Empidoidea | |
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Empis tessellata | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Diptera |
Infraorder: | Asilomorpha |
Superfamily: | Empidoidea |
Families | |
see text |
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]
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]
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]
Based on the most recent phylogenetic studies, [3] the internal classification of Empidoidea is as follows:
The Asiloidea comprise a very large superfamily insects in the order Diptera, the true flies. It has a cosmopolitan distribution, occurring worldwide. It includes the family Bombyliidae, the bee flies, which are parasitoids, and the Asilidae, the robber flies, which are predators of other insects.
The Lonchopteridae are a family of small (2–5 mm), slender, yellow to brownish-black Diptera, occurring all over the world. Their common name refers to their pointed wings, which have a distinct venation. Many are parthenogenic; males are very rare, however, at least in North American species, and have a somewhat different venation than do the females.
Dolichopodidae, the long-legged flies, are a large, cosmopolitan family of true flies with more than 8,000 described species in about 250 genera. The genus Dolichopus is the most speciose, with some 600 species.
Empididae is a family of flies with over 3,000 described species occurring worldwide in all the biogeographic realms but the majority are found in the Holarctic. They are mainly predatory flies like most of their relatives in the Empidoidea, and exhibit a wide range of forms but are generally small to medium-sized, non-metallic and rather bristly.
Hybotidae, the typical dance flies, are a family of true flies. They belong to the superfamily Empidoidea and were formerly included in the Empididae as a subfamily.
Atelestidae is a family of flies in the superfamily Empidoidea. The four genera were placed in a separate family in 1983; they were formerly either in Platypezidae or considered incertae sedis. While they are doubtless the most basal of the living Empidoidea, the monophyly of the family is not fully proven. The genus Nemedina seems to represent a most ancient lineage among the entire superfamily, while Meghyperus is probably not monophyletic in its present delimitation, and it is liable to be split up eventually, with some species being placed elsewhere. In 2010, the genus Alavesia, previously only known from Cretaceous fossils, was found alive in Namibia, subsequent species were also described from Brazil.
Trichopezinae are a subfamily of empidoid flies. They are mainly predatory flies like most of their relatives, and generally small to medium-sized, long-legged and large-eyed.
Apystomyia is a genus of flies in the family Apystomyiidae. The genus contains the single living Apystomyiidae species, Apystomyia elinguis, which is primarily found in California. Details of its life history are largely unknown. The extinct genus Hilarimorphites is known from the Cretaceous Burmese and New Jersey ambers. Formerly placed in the Asiloidea, molecular phylogenetic studies in 2010 placed the genus unambiguously as a sister of the Cyclorrhapha within the clade Eremoneura.
Tachydromiinae is a subfamily of hybotid flies widespread in the world.
Brachystomatinae is a subfamily of flies belonging to the family Empididae.
Clinocerinae is a subfamily of flies belonging to the family Empididae.
Apystomyiidae is a small family of flies containing the living genus Apystomyia and the extinct genera Apystomimus and Hilarimorphites. The single living Apystomyiidae species, Apystomyia elinguis, is native to California. Species of Hilarimorphites have been described from Mid to late Cretaceous Burmese and New Jersey ambers, while the single Apystomimus species is from the Late Jurassic of Kazakhstan.
Platypalpus candicans is a species of fly in the family Hybotidae. It is found in the Palearctic.
Parathalassiinae is a subfamily of flies in the family Dolichopodidae. It is part of an extended concept of the family, Dolichopodidae sensu lato, and forms a monophyletic group with Dolichopodidae sensu stricto. It was once placed provisionally in the subfamily Microphorinae as the tribe Parathalassiini.
Chersodromia is a genus of flies in the family Hybotidae.
Microphorinae is a subfamily of flies in the family Dolichopodidae. It is part of an expanded concept of the family, Dolichopodidae sensu lato, though it was previously considered a family of its own.
Ragadidae is a family of true flies in the superfamily Empidoidea. It was formerly considered a lower taxon, but was published as a new subfamily within Empididae in 2016. Since then, it has been classified as the sister group to Empididae, and has been elevated to family level based on the genetic differences which separate it from Empididae.
Microphorella is a genus of flies in the family Dolichopodidae. It is currently considered both paraphyletic and polyphyletic, and several species groups may need to be recognised as subgenera or genera.
Eremoneura is a clade of flies within the Brachycera that includes the Empidoidea and the Cyclorrhapha and is a sister of the Asilomorpha. They are thought to have evolved around the Mesozoic. The group includes fossils described in the genus Chimeromyia from 125 million year old amber which show both empidoid and cyclorrhaphan characters. The monotypic family Apystomyiidae has also been placed within the Eremoneura as a sister of the Cyclorrhapha.