Empididae - Empis Tessellata
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__NOTOC__ Empididae is a family of flies with over 3,000 described
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
occurring worldwide in all the
biogeographic realm A biogeographic realm or ecozone is the broadest biogeographic division of Earth's land surface, based on distributional patterns of terrestrial organisms. They are subdivided into bioregions, which are further subdivided into ecoregions. De ...
s but the majority are found in the
Holarctic The Holarctic realm is a biogeographic realm that comprises the majority of habitats found throughout the continents in the Northern Hemisphere. It corresponds to the floristic Boreal Kingdom. It includes both the Nearctic zoogeographical region ...
. They are mainly
predatory Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
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, but the dance flies proper, formerly included herein, are now considered a separate family
Hybotidae 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. Some, such as '' Tachydromia'', are predators that run around on the bark of t ...
.


Description

For terms see Morphology of Diptera. Empididae are small to medium-sized flies, rarely large (1.0 to 15.0mm.). 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. 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 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. Some, such as '' Tachydromia'', are predators that run around on the bark of t ...
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 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.Sinclair (2016) Since then, it has been classified as the sister group to E ...
and
Atelestidae 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 (which are not even particularly closely related) or considered ''incertae sedis'' ...
by having a prosternum that is fused with the proepisternum, which forms a precoxal bridge. 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.


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 Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...
(including other empidids),
Hemiptera Hemiptera (; ) is an order (biology), order of insects, commonly called true bugs, comprising over 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, Reduviidae, assassin bugs, Cimex, bed bugs, and shield bugs. ...
, Homoptera,
Lepidoptera Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...
, Trichoptera, Thysanoptera,
Hymenoptera Hymenoptera is a large order (biology), order of insects, comprising the sawfly, sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are Par ...
,
Neuroptera The insect order Neuroptera, or net-winged insects, includes the lacewings, mantidflies, antlions, and their relatives. The order consists of some 6,000 species. Neuroptera can be grouped together with the Megaloptera and Raphidioptera in th ...
,
Plecoptera Plecoptera is an order of insects, commonly known as stoneflies. Some 3,500 species are described worldwide, with new species still being discovered. Stoneflies are found worldwide, except Antarctica. Stoneflies are believed to be one of the mo ...
, Ephemeroptera,
Coleoptera Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
,
Collembola Springtails (Collembola) form the largest of the three lineages of modern hexapods that are no longer considered insects (the other two are the Protura and Diplura). Although the three orders are sometimes grouped together in a class called Ento ...
, and
Acari Mites are small arachnids (eight-legged arthropods). Mites span two large orders of arachnids, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari, but genetic analysis does not show clear evid ...
. 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
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
an species ''
Hilara maura ''Hilara maura'' is a species of dance fly, in the fly family Empididae __NOTOC__ 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. ...
'', have an elaborate
courtship Courtship is the period wherein some couples get to know each other prior to a possible marriage. Courtship traditionally may begin after a betrothal and may conclude with the celebration of marriage. A courtship may be an informal and private m ...
ritual in which the male wraps a prey item in
silk Silk is a natural protein fiber, some forms of which can be woven into textiles. The protein fiber of silk is composed mainly of fibroin and is produced by certain insect larvae to form cocoons. The best-known silk is obtained from the coc ...
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 America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
n 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 Amber is fossilized tree resin that has been appreciated for its color and natural beauty since Neolithic times. Much valued from antiquity to the present as a gemstone, amber is made into a variety of decorative objects."Amber" (2004). In Ma ...
deposits and the family certainly seems to have been well established by the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
period at the latest. Two groups formerly placed here as
subfamilies In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...
are now generally regarded as separate families in the Empidoidea:
Atelestidae 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 (which are not even particularly closely related) or considered ''incertae sedis'' ...
and
Hybotidae 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. Some, such as '' Tachydromia'', are predators that run around on the bark of t ...
. The Brachystomatidae are also sometimes separated as a distinct family, but this seems to be in error. The
Microphorinae 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. Genera The subfamily includes seven gene ...
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).Moulton & Wiegmann (2007) Among the subfamilies currently placed herein, not all are confirmed to be
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
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 Brachystomatinae is a subfamily of flies belonging to the family Empididae. (2018): Molecular phylogenetics reveals novel relationships within Empidoidea (Diptera). ''Systematic Entomology'' 43(4): 619–636. Taxonomy Until 2006, Brachystomatina ...
,
Empidinae Empidinae 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. Genera *'' Afroempis'' Smith ...
and
Hemerodromiinae Hemerodromiinae are a worldwide group of predatory flies with raptorial forelegs.Plant, A.R. 2011: Hemerodromiinae (Diptera: Empididae): a tentative phylogeny and biogeographical discussion. ''Systematic entomology'', 36(1): 83-103. Genera *'' ...
however seem to be natural groups of closest relatives in their entirety, and the
Clinocerinae Clinocerinae is a subfamily of flies belonging to the family Empididae. Genera *'' Afroclinocera'' Sinclair, 1999 *'' Asymphyloptera'' Collin, 1933 *'' Bergenstammia'' Mik, 1881 *'' Clinocera'' Meigen, 1803 *'' Dolichocephala'' Macquart, 182 ...
apparently are for the largest part. Based on the most recent phylogenetic studies, the relationship between Empididae and other members of Empidoidea is as follows. The placement of Empididae is emphasized in bold formatting.


Species Lists


West Palaearctic including RussiaJapan

World list


Identification

*Chvála, M. (1994) ''The Empidoidea (Diptera) of Fennoscandia and Denmark''. III. Genus Empis. Leiden: Brill 29. *Collin, J.E. (1961) ''British Flies''. VI. Empididae. Cambridge: Cambridge University Press. * Engel, E. O., 1938-1954 Empididae. In:
Lindner Lindner is a German surname, which may refer to: * Lindner family, American business family based in Cincinnati, Ohio * Lindner (agricultural machinery manufacturer), Austrian family company * Arlon Lindner (1935–2021), American businessman and p ...
, E. (Ed.). ''Die Fliegen der Paläarktischen Region'', 4, 28, 1-400. *
Frey Freyr (Old Norse: 'Lord'), sometimes anglicized as Frey, is a widely attested god in Norse mythology, associated with kingship, fertility, peace, and weather. Freyr, sometimes referred to as Yngvi-Freyr, was especially associated with Sweden and ...
, R. 1954- 1956 Empididae. In: Lindner, E. (Ed.). ''Die Fliegen der Paläarktischen Region'' 4, 28, 400-639. *Gorodkov, K. B. and Kovalev V.G. Family Empididae in Bei-Bienko, G. Ya, 1988 ''Keys to the insects of the European Part of the USSR'' Volume 5 (Diptera) Part 2 English edition. Keys to Palaearctic species but now needs revision .


Gallery

Image:Empis.borealis.jpg, ''
Empis borealis ''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 ...
'' Image:Empis.ciliata.jpg, ''
Empis ciliata ''Empis ciliata'', the black dance fly, is a species of dance fly, in the fly family Empididae. It is included in the subgenus ''Euempis''. Distribution This species can be found in western and southern Europe (Czech Republic, France, Germany, I ...
'' Image:Empis.livida.with.prey.jpg, ''
Empis livida ''Empis livida'' is a species of dance fly, in the fly family Empididae. It is included in the subgenus ''Kritempis'' of the genus ''Empis''. Males range from , females . The male's abdomen is brownish, and his wings appear faintly brown and clo ...
'' with prey Image:Empis.opaca.jpg, '' Empis opaca'' Image:Empis.pennipes.female.2.jpg, '' Empis pennipes'' Image:Empis.stercorea.jpg, '' Empis stercorea'' Image:Empis tesselata male (aka).jpg, ''
Empis tessellata ''Empis tessellata'' is a species of dance fly, in the fly family Empididae. It is included in the subgenus ''Euempis''. Distribution The species is distributed in most of Europe from the Mediterranean to Scandinavia, in the east to the Middle E ...
'' File:Empis tessallata (AF) -right.jpg, ''
Empis tessellata ''Empis tessellata'' is a species of dance fly, in the fly family Empididae. It is included in the subgenus ''Euempis''. Distribution The species is distributed in most of Europe from the Mediterranean to Scandinavia, in the east to the Middle E ...
'' Image:Empis.variegata.jpg, '' Empis variegata'' Image:Hilara litorea.jpg, '' Hilara litorea'' Image:Rhamphomyia.crassirostris.jpg, ''
Rhamphomyia crassirostris ''Rhamphomyia crassirostris'' is a species of dance flies, in the fly family Empididae. It is found in most of Europe, east to Poland and Hungary. It is absent from Italy and the Balkan Peninsula The Balkans ( ), also known as the Balk ...
'' Image:Rhamphomyia.lamellata.jpg, ''
Rhamphomyia lamellata ''Rhamphomyia lamellata'' is a species of dance flies, in the fly family Empididae. It is included in the subgenus ''Holoclera'' of the genus ''Rhamphomyia''. It is found in Great Britain and Ireland, Germany, Switzerland, the Czech Republic, Slo ...
'' Image:Rhamphomyia.marginata.female.jpg, ''
Rhamphomyia marginata ''Rhamphomyia marginata'' is a species of dance flies, in the fly family Empididae. It is found in Europe, from Great Britain east to Romania and from Fennoscandia south to France, Austria and Hungary Hungary ( hu, Magyarország ) is a l ...
'' Image:Rhamphomyia.sulcata.jpg, ''
Rhamphomyia sulcata ''Rhamphomyia sulcata'' is a species of dance flies, in the fly family Empididae. It is included in the subgenus ''Rhamphomyia''. It is found in most of Europe, except the Balkan Peninsula The Balkans ( ), also known as the Balkan Pen ...
'' Image:Danseflue.jpg, Empididae sp. Image:Empis trigramma.tif, ''
Empis trigramma ''Empis trigramma'' is a species of dance flies, in the fly family Empididae. The thorax with yellowish sides, with three broad dark stripes. Abdomen The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of ...
'' Image:Empis tessellata 2.tif, ''
Empis tessellata ''Empis tessellata'' is a species of dance fly, in the fly family Empididae. It is included in the subgenus ''Euempis''. Distribution The species is distributed in most of Europe from the Mediterranean to Scandinavia, in the east to the Middle E ...
''


Footnotes


References

* (1986): ''Collins Guide to the Insects of Britain and Western Europe''. * * *K. G. V. Smith, 1989 An introduction to the immature stages of British Flies. Diptera Larvae, with notes on eggs, puparia and pupae.'' Handbooks for the Identification of British Insects'' Vol 10 Part 14
pdf
download manual (two parts Main text and figures index) *


External links

* * *
Empididae __NOTOC__ 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 Empido ...
In Italian
Family Empididae at EOL
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Diptera.info
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Key to the British genera of Empididae
{{Taxonbar, from=Q1326176 Brachycera families Articles containing video clips Extant Cretaceous first appearances