Empididae

Last updated

Dagger flies and balloon flies
Temporal range: Cretaceous - Recent
Empis livida (aka).jpg
Male Empis livida
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Diptera
Suborder: Brachycera
Infraorder: Asilomorpha
Superfamily: Empidoidea
Family: Empididae
Latreille, 1804
Subfamilies

Brachystomatinae
Ceratomerinae
Clinocerinae
Empidinae
Hemerodromiinae
Trichopezinae

Contents

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.

Common names for members of this family are dagger flies (referring to the sharp piercing mouthparts of some species) and balloon flies. The term "dance flies" is sometimes used for this family too, [1] but the dance flies proper, formerly included herein, are now considered a separate family Hybotidae.

Description

Empididae are small to medium-sized flies, rarely large (1.0 to 15.0 mm). The body is slender, or elongated and rarely thickset. The colour ranges from yellow to black, and they may be pollinose or lustrous, but never have a metallic gloss. The head is often small and rounded with relatively large eyes. The eyes of the male and (rarely) the female may be contiguous (holoptic). The antenna usually have three segments of which the third is the largest and bears a long or short, usually apical arista or style. The eyes often have an incision (notch) at the level of the antennae. The proboscis is often long and in several genera powerful and piercing. If the mouthparts are strongly elongate they project forward or downward toward the fore legs. Some species have short mouthparts. The legs are usually long and slender but often powerful and in some cases, the fore legs are raptorial, adapted to catching and holding prey. The wings are clear or partially tinged and, in some species, with a stigma spot, or with a distinct pattern. Radial vein R4+5 is often forked and the discal-medial cell (dm) is almost always present. The costa ends at or just beyond R4+5, or continues along the wing margin, and can be used as a characteristic to distinguish it from other families. [2] The venation of the wings in minute species is often simplified. The posterior part of the wings on the basal side of the anal vein bears the designation of axillary or anal lobe. In species with a highly developed axillary lobe, the margin may form an angle with the margin of the alula. This is termed the axillary angle, the size of which is often a good diagnostic feature. It can be distinguished from Dolichopodidae and Hybotidae by the presence of unrotated and symmetrical terminalia. The point of origin of Rs (radial sector) is at a distance from the humeral crossvein (h) equal to or longer than the length of h. Except for a few species, Empididae is easily distinguishable from the families Ragadidae and Atelestidae by having a prosternum that is fused with the proepisternum, which forms a precoxal bridge. [2]

The larvae have an incomplete head capsule (hemicephalic) which is often retracted into the thorax 8-segmented abdomen with paired parapods on abdominal segments 1‒7 or 1‒8. The posterior spiracles are quite widely separated and the anal segment is frequently lobed. The pupae is without a puparium.

Mating Empididae: Female consuming male's mating gift of an ant Empid flies Empididae mating 01.jpg
Mating Empididae: Female consuming male's mating gift of an ant

Biology

Adult empidids are found in a variety of forest habitats, on the leaves of plants, on tree trunks, aquatic vegetation and also in stream beds and seepage habitats. Some species are associated with open areas such as grasslands, agricultural fields, marshes, coastal zones and beaches. Adults capture arthropod prey, including other Diptera (including other empidids), Hemiptera, Homoptera, Lepidoptera, Trichoptera, Thysanoptera, Hymenoptera, Neuroptera, Plecoptera, Ephemeroptera, Coleoptera, Collembola, and Acari. Because of their predation they are important natural and biological control agents of various pest insect species and as a group with a vast species diversity they occupy a wide range of habitats. Some adult Empididae also visit flowers to obtain nectar. At least a few groups (Iteaphila, Anthepiscopus, Anthalia, Allanthalia and Euthyneura) obtain all their protein needs by feeding on pollen. Larvae are generally found in moist soil, rotten wood, dung, or in aquatic habitats and also appear to be predaceous on various arthropods, particularly other Diptera larvae.

Some Empididae, such as the European species Hilara maura , have an elaborate courtship ritual in which the male wraps a prey item in silk and presents it to the female to stimulate copulation. It was first noticed that these flies carried "nuptial gifts" of silk by Baron Karl von Osten-Sacken. Empididae show diverse mating systems, ranging from species in which males aggregate in mating swarms, and compete for choosing females to sex-role reversed species in which females do aggregate and compete for the attention of choosing males. In some species, such as the North American species Rhamphomyia longicuada , competition for the food provided by males is so strong that females have developed elaborate ornaments, including feathery "pennate" scales on their legs, darkened wings, and inflatable abdominal sacs that enhance their attractiveness.

Evolution and systematics

Empididae are well represented in amber deposits and the family certainly seems to have been well established by the Cretaceous period at the latest.

Two groups formerly placed here as subfamilies are now generally regarded as separate families in the Empidoidea: Atelestidae and Hybotidae. The Brachystomatidae are also sometimes separated as a distinct family, but this seems to be in error. The Microphorinae were long placed in the Empididae as a subfamily, then briefly classified as a distinct family, and are now considered a subfamily of the long-legged flies (Dolichopodidae). [3] [4]

Dolichocephala irrorata (Clinocerinae) in copula

Among the subfamilies currently placed herein, not all are confirmed to be monophyletic groups. Some rearrangements, in particular regarding the delimitation of Empididae versus Dolichopodidae – which together represent the bulk and the most advanced lineages of the Empidoidea – are likely to take place in the future. The Brachystomatinae, Empidinae and Hemerodromiinae however seem to be natural groups of closest relatives in their entirety, and the Clinocerinae apparently are for the largest part. [4]

Based on the most recent phylogenetic studies, [2] the relationship between Empididae and other members of Empidoidea is as follows. The placement of Empididae is emphasized in bold formatting.

Atelestidae
Hybotidae

Bicellariinae

Tachydromiinae

Tachydromiini

Symballaphthalmini

Drapetini

Dolichopodidae

Ragadidae
Empididae

Clinocerinae

Brachystomatinae

Brachystomatini

Ceratomerini

Trichopezini

Empidinae

Chelipodini

Empidini

Hemerodromiini

Hilarini


Identification

Footnotes

  1. ↑ Sjöberg, Fredrik (2014). The Fly Trap. Particular Books. p. 197. ISBN   978-1-846-14776-0.
  2. 1 2 3 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.
  3. ↑ Sinclair, Bradley J.; Cumming, J.M. (2006). "The morphology, higher-level phylogeny and classification of the Empidoidea (Diptera)" (PDF). Zootaxa . 1180. New Zealand: Magnolia Press: 1–172. ISSN   1175-5334 . Retrieved 2009-06-04.
  4. 1 2 Moulton & Wiegmann (2007)

Related Research Articles

<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">Dolichopodidae</span> Family of flies

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.

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

The Scenopinidae or window flies are a small family of flies (Diptera), distributed worldwide. In buildings, they are often taken at windows, hence the common name window flies.

<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>Empis livida</i> Species of fly

Empis livida is a species of fly in the Empididae family. It is included in the subgenus Kritempis of the genus Empis. Males range from 7.5 to 9.3 millimetres, females 7.5 to 10.2 millimetres. The male's abdomen is brownish and its wings appear faintly brown and clouded. The female's abdomen is gray and its wings are clear. E. livida lives in hedgerows, feeding on the nectar of several species of Rosaceae, several species of Asteraceae, and Heracleum sphondylium nectar; they also feed on other insects. They live all across temperate and Northern Europe, the only species with such a wide distribution. E. livida larvae are also carnivorous and live in damp soil and leaf litter. Adults fly in between April and July.

<i>Empis borealis</i> Species of fly

Empis borealis is a species of dance flies in the fly family Empididae. It is found in most of Europe, except the Balkan Peninsula, the Baltic Region and the Iberian Peninsula. The brown wings of the female are very broad, while the male fly’s wings are narrower.

<i>Empis tessellata</i> Species of fly

Empis tessellata is a species of dance fly, in the fly family Empididae. It is included in the subgenus Euempis.

<i>Empis ciliata</i> Species of fly

Empis ciliata, the black dance fly, is a species of dance fly, in the fly family Empididae. It is included in the subgenus Euempis.

Milan Chvála was a Czech entomologist who specialised in Diptera.

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

Empidinae, also called dance flies, are a subfamily of empidoid flies. They are mainly predatory flies like most of their relatives, and generally small to medium-sized. Most species are flower visitors and they can be effective pollinators.

<i>Bicellaria vana</i> Species of fly


Bicellaria vana is a species of fly in the family Hybotidae. It is found in the Palearctic.

<i>Platypalpus candicans</i> Species of fly

Platypalpus candicans is a species of fly in the family Hybotidae. It is found in the Palearctic.

Chersodromia is a genus of flies in the family Hybotidae.

<span class="mw-page-title-main">Ragadidae</span> Family of true flies in the superfamily Empidoidea

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.

Empis rohaceki is a species of fly in the family Empididae. It is included in the subgenus Xanthempis. It is found in the Palearctic.

Homalocnemis is a genus of flies which is placed in a family of its own, the Homalocnemidae. There are about seven species in the genus found in the Afrotropical, Neotropical, and Australasian regions, suggestive of a Gondwanan origin. The genus was formerly considered a primitive empidoid and placed variously in the Hybotidae or in the empidid subfamily Brachystomatinae. They are recognized by their wing venation which includes a long anal cell and a long basal segment of the antennal style.

Stilpon is a genus of flies in the family Hybotidae.

References

Species lists

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