Dolichopodidae

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Dolichopodidae
Temporal range: Cretaceous–Recent
Long legged fly.jpg
Chrysosoma sp.
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Diptera
Infraorder: Asilomorpha
Superfamily: Empidoidea
Family: Dolichopodidae
Latreille, 1809
Subfamilies

sensu stricto :

sensu lato:

Diversity
About 230 genera, more than 7,000 species
Synonyms

Dolichopidae

Dolichopodidae, the long-legged flies, are a large, cosmopolitan family of true flies with more than 7,000 described species in about 230 genera. The genus Dolichopus is the most speciose, with some 600 species.

Contents

Dolichopodidae generally are small flies with large, prominent eyes and a metallic cast to their appearance, though there is considerable variation among the species. Most have long legs, though some do not. In many species, the males have unusually large genitalia which are taxonomically useful in identifying species. Most adults are predatory on other small animals, though some may scavenge or act as kleptoparasites of spiders or other predators.

Austrosciapus connexus--a typical example of green Dolichopodidae Austrosciapus connexus (Green longlegged fly).jpg
Austrosciapus connexus—a typical example of green Dolichopodidae

Description

Dolichopodidae are a family of flies ranging in size from minute to medium-sized (1 mm to 9 mm). They have characteristically long and slender legs, though their leg length is not as striking as in families such as the Tipulidae. Their posture often is stilt-like standing high on their legs, with the body almost erect. In colour most species have a green-to-blue metallic lustre, but various other species are dull yellow, brown or black.

The frons in both sexes is broad. The eyes are separated on the frons of males, except in some species of Diaphorus and Chrysotus in which eyes touch above the antennal insertion. [5] On the heads of most species the ocellar bristles and outer vertical bristles are well developed. The face of some species is entire; in others it is divided into two sections: the epistoma and the clypeus. The largest antennal segment is the third; in most species it bears a long arista, which is apical in some species, dorsal in others. In most species the mouthparts are short and have a wide aperture as an adaptation for sucking small prey.

The legs are gracile and the tibiae usually bear long bristles. In some genera the legs are raptorial. In some species the tibiae of the males have modifications.

Dolichopodidae wing veins. This is one major type, with M1 bent and M2 present, though often incomplete. In another type, M2 is absent and M1 more or less straight Dolichopodidae wing veins-1.svg
Dolichopodidae wing veins. This is one major type, with M1 bent and M2 present, though often incomplete. In another type, M2 is absent and M1 more or less straight
Photograph of a Dolichopodidae wing, showing the R4+5 vein Dolichopodidae wing.png
Photograph of a Dolichopodidae wing, showing the R4+5 vein

The wings of most species are clear or tinged, but some species have wings that are patterned in strong colours or with distinct spots. There are three radial veins (R1, R2+3, R4+5). The medial vein M1+2 is simple or rarely furcate, as in the genus Sciapus . The anterior cross-vein is in the basal part of the wing. The posterior basal wing cell and the discoidal wing cell are always fused. The anal cell of the wing is always small. There are two veins branching from cross-vein DM-Cu in the direction of the wing margin; the upper one in some species curves strongly or forks into M1 and M2. R4+5 are simple, and costa ends near or at M1/M1+2, or continues along the wing margin. The point of origin of Rs is at or very close to h. [6]

Photograph of genetalia of a male Empis vitripennis. Note the asymmetrical tergite 8 on the left side of the epiandrium Male genetalia Dolichopodidae.png
Photograph of genetalia of a male Empis vitripennis. Note the asymmetrical tergite 8 on the left side of the epiandrium
Condylostylus patibulatus Longlegged Fly Dolichopodidae (Condylostylus patibulatus).jpg
Condylostylus patibulatus

The abdomen is elongate-conical or flat. The genitalia of the male often are free and borne on a petiole, with tergite 8 being asymmetrical, lying on the left side of the epandrium. They are also rotated dextrally between 90° and 180°, including segment 8 and sometimes segment 7, which makes them distinguishable from the family Hybotidae. [6] Males of most species have well developed gonopods of two or three lobes on the distal margin of the epandrium. The gonopods may fuse with the epandrium in genera such as Hydrophorus , Thrypticus and Argyra , or there may be a suture, as in the genera Porphyrops , Xiphandrium and Rhaphium . In some genera, such as Hypophyllus and Tachytrechus , the surstyli are well-developed as secondary outgrowths of the epandrium. In genera such as Tachytrechus, there are two pairs of surstyli—one proximal and one distal. The hypandrium in most species is a small sclerite, which may be asymmetrical as in the genera Porphyrops and Tachytrechus. Males of many species have highly developed cerci. Development of the phallus varies considerably between genera.

Dolichopodidae, cf. genus Medetera, typical of brown species inhabiting bark of damaged trees. Characteristic head-up, stilt-legged posture Dolichopodidae, Brion, Galicia (Spain).jpg
Dolichopodidae, cf. genus Medetera, typical of brown species inhabiting bark of damaged trees. Characteristic head-up, stilt-legged posture

Biology

Adults of the Dolichopodidae live largely in grassy places and shrubbery. The flies occur in a wide range of habitats, near water or in meadows, woodland edges and in gardens. Some groups are confined to wet places including sands on the banks of water bodies; examples include genera such as Porphyrops, Tachytrechus, Campsicnemus , and Teuchophorus . No truly aquatic species have been described, but many are semi-aquatic and live in or near water margins. A small number of species develop on the shores of saline inland bodies of water or the intertidal zone of seashores. An example of a species that develop close to water is P. nobilitatus, they can be found congregating around lakes and ponds. Other groups are found on trunks of trees damaged by bark beetles. Adults often are seen in a characteristic predatory posture standing high on their legs on the ground or on vegetation, tree trunks or rocks, and some species walk about on the surface of still water.

The adults are predators, feeding on small invertebrates including Collembola, aphids, and the larvae of Oligochaeta. Species of the genus Dolichopus commonly prey on the larvae of mosquitoes.

The larvae occupy a wide range of habitats. Many are predators of small invertebrates and generally live in moist environments such as soil, moist sand, or rotting organic matter. Genera such as Medetera live as predators under tree bark or in the tunnels of bark beetles. Larvae of the genus Thrypticus are unusual among Dolichopodidae, in that they are phytophagous and live in the stems of reeds and other monocots near water.[ citation needed ]

Behaviour

Foraging and nuptial behaviour of Poecilobothrus nobilitatus (video, 2m 58s)

Many studies have shown that Dolichopodidae give visual, rather than chemical or other signals during courtship. [7] The males of many species exhibit elaborate secondary sexual characters assumed to aid in species recognition during courtship. These characters include flaglike flattening of the arista and tarsi, strongly modified setae and projections of the tarsi, the prolongation and deformation of podomeres, orientated silvery pruinosity, and maculation or modification of the wings.

Evolution and systematics

Kleptoparasitic Microphor holosericeus (Microphorinae) feeding on captured prey of a spider

Dolichopodids are well represented in amber deposits throughout the world and the group has clearly been well distributed since the Cretaceous at the latest. Together with the Empididae they are the most advanced members of the Empidoidea. They represent the bulk of Empidoidea diversity, and include more than two-thirds of the known species in their superfamily.

Taxonomic interrelationships within the Dolichopodidae, and their delimitation from the Empididae, are not yet satisfactorily resolved. It is likely that many of the subfamilies currently within the Dolichopodidae will undergo drastic revision. [8]

An expanded concept of the family, Dolichopodidae sensu lato , includes the former family Micromorphidae as the subfamilies Microphorinae and Parathalassiinae, the latter forming a monophyletic group with the Dolichopodidae sensu stricto. [9] Alternatively, Grichanov (2011) proposes that Dolichopodidae s.l. should be treated as an epifamily Dolichopodoidae, containing Dolichopodidae, Microphoridae and the subfamily Parathalassiinae. [10]

Based on the most recent phylogenetic studies, the relationship between Dolichopodidae and other members of Empidoidea is as follows. The placement of Dolichopodidae is emphasized in bold formatting.

Atelestidae

Atelestinae

Nemedininae

Hybotidae

Bicellariinae

Hybotinae

Oedaleinae

Ocydromiinae

Trichininae

Tachydromiinae

Tachydromiini

Symballaphthalmini

Drapetini

Dolichopodidae

Ragadidae

Iteaphilinae

Ragadinae

Empididae

Clinocerinae

Brachystomatinae

Brachystomatini

Ceratomerini

Trichopezini

Empidinae

Chelipodini

Empidini

Hemerodromiini

Hilarini

Dolichopus ungulatus Dolichopus ungulatus (Dolichopodidae).tif
Dolichopus ungulatus

Identification

Species lists

See also

Sciapus sp. Longlegged fly - sciapus.jpg
Sciapus sp.

Footnotes

  1. Grootaert, P; Meuffels, H.J.G. (1997). "Dolichopodidae (Diptera) from Papua New Guinea. XV. Scepastopyga gen. nov. and the establishment of a new subfamily, the Achalcinae". J. Nat. Hist. 31 (10): 1587–1600. doi:10.1080/00222939700770841.
  2. 1 2 Yang, D.; Zhu, Y.; Wang, M.; Zhang, L. (2006). World Catalog of Dolichopodidae (Insecta: Diptera). Beijing: China Agricultural University Press. pp. 1–704. ISBN   9787811171020.
  3. Bickel, D. J. (1987). "Babindellinae, a new subfamily of Dolichopodidae (Diptera) from Australia, with a discussion of symmetry in the dipteran male postabdomen". Entomologica Scandinavica. 18: 97–103. doi:10.1163/187631287X00061. ISSN   1399-560X.
  4. Grichanov, I.Ya. (2018). "A new subfamily of Dolichopodidae (Diptera) for Tenuopus Curran, 1924 with description of new species from Tropical Africa" (PDF). Far Eastern Entomologist. 365: 1–25. doi: 10.25221/fee.365.1 . Archived from the original (PDF) on 24 October 2019. Retrieved 29 November 2018.
  5. Robinson, H. and JR Vockeroth. Dolichopodidae. JF McAlpine Manual of Nearctic Diptera vol. 1 1981. 625–639. Research Branch Agriculture Canada Monograph 27 Ottawa.
  6. 1 2 3 4 Wahlberg, Emma; Johanson, Kjell Arne (2018). "Molecular phylogenetics reveals novel relationships within Empidoidea (Diptera)". Systematic Entomology. 43 (4): 619–636. doi: 10.1111/syen.12297 . ISSN   1365-3113.
  7. E.g. Zimmer et al. (2003), Irwin (2007), Vikhrev (2007)
  8. Sinclair and Cumming (2006), Moulton and Wiegmann (2007)
  9. Sinclair, Bradley J.; Cumming, Jeffrey M. (2006). "The morphology, higher-level phylogeny and classification of the Empidoidea (Diptera)" (PDF). Zootaxa . 1180: 1–172. doi:10.11646/zootaxa.1180.1.1. ISBN   978-1-877407-80-2. Archived (PDF) from the original on 3 March 2016. Retrieved 20 December 2011.
  10. Grichanov, I. Ya. (2011). "An illustrated synopsis and keys to afrotropical genera of the epifamily Dolichopodoidae (Diptera: Empidoidea)". Priamus Supplement (24): 1–98.

Related Research Articles

<span class="mw-page-title-main">Asiloidea</span> Superfamily of flies

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.

<i>Argyra</i> Genus of flies

Argyra is a genus of flies in the family Dolichopodidae. The name "Argyra" comes from the Greek word for "silver".

<span class="mw-page-title-main">Empidoidea</span> Superfamily of flies

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, which are represented on all continents except Antarctica. They are known to have existed since the Jurassic period.

<span class="mw-page-title-main">Empididae</span> Family of flies

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.

<span class="mw-page-title-main">Hybotidae</span> Family of flies

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.

<span class="mw-page-title-main">Atelestidae</span> Family of flies

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.

<i>Hercostomus</i> Genus of flies

Hercostomus is a genus of flies in the family Dolichopodidae. It is a large genus, containing more than 483 species worldwide. Multiple studies have shown that Hercostomus is a polyphyletic assemblage of species.

Antyx is a genus of flies in the family Dolichopodidae from the Australasian realm. The genus was originally placed in the subfamily Sympycninae, but it was later found to be closer to the Neurigoninae and was placed as incertae sedis within the family. In the World Catalog of Dolichopodidae by Yang et al. (2006), the new subfamily Antyxinae was proposed, in which Antyx is the only genus. However, the validity of this new subfamily was later criticized by Sinclair et al. (2008). According to them, the subfamily's erection by Yang et al. (2006) was not justified by their phylogenetic analysis, and the genus would have been better placed as incertae sedis until a later phylogenetic study determines its placement.

<i>Lichtwardtia</i> Genus of flies

Lichtwardtia is a genus of flies in the family Dolichopodidae. In a phylogenetic analysis of the subfamily Dolichopodinae by Scott E. Brooks in 2005, Lichtwardtia is considered to be a synonym of Dolichopus, but other authors retain it as a valid genus. Before this, it was considered a possible subgenus of Pterostylus by Oleg Negrobov (1979).

<i>Sybistroma</i> Genus of flies

Sybistroma is a genus of flies in the family Dolichopodidae. It includes over 50 species, described mainly from the Palaearctic and Oriental realms. A single species is known from the Afrotropical realm. Until 2005, the genus was thought to be restricted to the Mediterranean in distribution, with five known species. It was recently expanded to include the former genera Hypophyllus, Ludovicius and Nordicornis, as well as some species of Hercostomus.

<span class="mw-page-title-main">Hydrophorinae</span> Subfamily of flies

Hydrophorinae is a subfamily of flies in the family Dolichopodidae. Several studies have found evidence that the subfamily in its current sense is polyphyletic.

<span class="mw-page-title-main">Dolichopodinae</span> Subfamily of flies

Dolichopodinae is a subfamily of flies in the family Dolichopodidae.

Kowmunginae is a subfamily of flies in the family Dolichopodidae. It was proposed in the World Catalog of Dolichopodidae by Yang et al. (2006) to include two genera previously placed as incertae sedis within the family. However, the validity of this new subfamily was later criticized by Sinclair et al. (2008). According to them, the subfamily's erection by Yang et al. (2006) was not justified by their phylogenetic analysis, and the genera included would have been better placed as incertae sedis until a later phylogenetic study determines their placement.

<span class="mw-page-title-main">Peloropeodinae</span> Subfamily of flies

Peloropeodinae is a subfamily of flies in the family Dolichopodidae. In some classifications, the genera of the subfamily are included in Sympycninae. According to Germann et al. (2011), the subfamily is polyphyletic.

Medeterella is a genus of flies in the family Dolichopodidae. It contains nine species formerly included in Medetera in the M. salomonis species group. The species are found in the Afrotropical, Oriental and Australasian regions. According to Naglis and Bickel (2012), it was unwarranted to establish a separate genus for this group of species.

<span class="mw-page-title-main">Parathalassiinae</span> Subfamily of flies

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.

Eothalassius is a genus of flies in the family Dolichopodidae. It contains four described species, from Southeast Asia, Papua New Guinea, Costa Rica and the Mediterranean. It also contains three undescribed species, one from New Caledonia and two from Japan.

<i>Microphor</i> Genus of flies

Microphor is a genus of flies in the family Dolichopodidae, subfamily Microphorinae.

<i>Microphorella</i> Genus of Dolichopodid flies

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.

References