| Asilidae | |
|---|---|
_female_with_bee_prey_Babadag.jpg) | |
| Robber fly (Promachus leoninus) with honeybee prey | |
Scientific classification | |
| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Arthropoda |
| Class: | Insecta |
| Order: | Diptera |
| Suborder: | Brachycera |
| Infraorder: | Asilomorpha |
| Superfamily: | Asiloidea |
| Family: | Asilidae Latreille, 1802 |
| Subfamilies | |
TheAsilidae are therobber fly family, also calledassassin flies. They are powerfully built, bristlyflies with a short, stoutproboscis enclosing the sharp, suckinghypopharynx.[1][2] The name "robber flies" reflects their expert predatory habits; they feed mainly or exclusively on other insects and, as a rule, they wait in ambush and catch their prey in flight.
The Asilidae are a family in theorderDiptera, the true flies. The common name for members of the family is the robber flies. The Asilidae arecosmopolitan, with over 7000 describedspecies.Latreille was the authority for establishing the family in 1802.[3] The Asilidae, together withBombyliidae andTherevidae, are the most representative families of the superfamily ofAsiloidea and they form one of the most characteristic groups of the lowerBrachycera.
Robber flies have stout, spiny legs and three simple eyes (ocelli) in a characteristic depression on the tops of their head between their two largecompound eyes.[4] They also have a dense moustache of stiff bristles on the face; this is called themystax, a term derived from the Greekmystakos meaning "moustache" or "upper lip". The mystax has been suggested to afford some protection for the head and face when the flies deal with strugglingprey; various Asilidae prey on formidable species includingstinging Hymenoptera, powerfulgrasshoppers,dragonflies and even other Asilidae – practically anything of a suitable size. Some Asilidae do, however, specialize in smaller prey, and this is reflected in their moregracile build.
In general, the family attacks a very wide range of prey, including other flies,beetles,butterflies andmoths, variousbees,ants,dragonflies anddamselflies,ichneumon wasps,grasshoppers, and somespiders. They seem to do so irrespective of anyrepugnatorial chemicals the prey may have at their disposal.[5] When attacked, many Asilidae do not hesitate to defend themselves in turn using theirprobosces and may deliver intensely painful bites to humans if handled incautiously.
Theantennae are short, have three segments, and sometimes bear a bristle-like structure called anarista.
Though they are a very characteristic group for such a large family, the Asilidae may easily be confused with the related and less widely known familyTherevidae. Some points of contrast between the families include that thelabium in the Therevidae is not a piercing, predatory organ, but ends in two fleshylabella adapted to the sucking of liquid foods. Again, the Therevidae commonly have fluffysetae above the mouthparts, unlike the stiffchaetae comprising the mystax of the Asilidae. Furthermore, in the Asilidae the depression on thevertex between the eyes tends to be more obvious than in the Therevidae.
The fly attacks its prey by stabbing it with its short, strong proboscis, injecting the victim with saliva containingneurotoxic andproteolytic enzymes which very rapidly paralyze the victim and soon digest the insides; the fly then sucks the liquefied material through the proboscis.
Many Asilidae have long, taperingabdomens, sometimes with a sword-likeovipositor. Others, for instanceLaphria, are fat-bodiedbumblebee mimics.
Female robber flies deposit whitish-colored eggs on low-lying plants and grasses, or in crevices within soil, bark, or wood. Egg-laying habits depend on the species and their specific habitat; most species lay their eggs in masses, which are then covered with a chalky protective coating.
After hatching, robber fly larvae generally seem to live in soil, rotting wood, leaf mold, and similar materials, some being predatory and othersdetrivorous.[5] Larvae are also predacious, feeding on eggs, larvae, or other soft-bodied insects. Robber flies overwinter as larvae and pupate in the soil. Pupae migrate to the soil surface and emerge as adults, often leaving behind their pupal casing. Complete development ranges from one to three years, depending on species and environmental conditions.[6]
Adults are generally medium to large in size, with an average body width of 1 to 1.5 cm (0.39 to 0.59 in), but with a range of 3 mm (0.12 in) to more than 5 cm (2.0 in) in length. The shape is generally elongated, due to the conformation of the long taperingabdomen, however there are also compact species with broad abdomens. The integument is covered with thick hair, especially on thehead andthorax and liveries are often showy, with colors ranging from brown to black to grey, sometimes in contrast with other colors such as red and yellow. Frequently they areaposematic, imitating the livery of Hymenoptera.
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The head is free and mobile anddichoptic in both sexes and has threeocelli arranged in a characteristic depression formed by the elevation of thecompound eyes. This feature is clearly visible in the front view and is a morphological peculiarity of Asilidae. Theoccipital region has one or more rows of bristles aligned behind the posterior margin of the eye. Thefacial region has a convex profile with a characteristic dense bundle of bristles, called amystax. The mystax helps protect the head and face when the fly encounters prey bent on defense. Other bristles are arranged on the ocellar tubercle.
The antennae are of thearistate type, composed typically of five segments but sometimes from three to four, depending on the structure of the stylus. Thescape andpedicel are generally relatively short and hairy; the third segment (or first flagellomere) has an oval or oblong shape, is generally longer than the two basal segments, and bears a stylus generally composed of two segments, of which the basal is very short. In some asilids, the stylus can be monoarticolate or absent.
The mouthparts are short, being modified for piercing and sucking. The strongly sclerotized proboscis is composed of thelabium andmaxillae which form a food canal, the labrum and a piercing organ, thehypopharynx. The proboscis is either rounded in cross section or compressed laterally or dorsoventrally; it is usually stout and straight and is sometimes able to penetrate through the hard integument of Coleoptera. The maxillary palpi are at the base beside the labium, two-segmented in all Dasypogoninae or single segmented in Asilinae and Leptogastrinae.
The thorax is robust and compact. Unlike in other lower Brachycera, it bears long bristles (macrochaeta) useful astaxonomic characters. Bristles of this type are always present on thenotopleuron (notopleural bristles) and, in two series, on mesonotum (dorsocentral, supralar and postalar). Other bristles are present on the metanotum (dorsocentral bristles on the ventral episternum and at the apex of the mesoscutellum.
The legs are relatively long and strong, with many macrochaetes performing araptorial function. The wings are well developed, often relatively narrow for speedy flight; thealula is generally well developed, with the exception of the Leptogastrinae and part of Dasypogoninae. The venation is much as in theRhagionidae,Tabanidae, and Therevidae; the radial R is always four-branched, with R2+3 unbranched. Details of wing venation determine subfamilies and lower taxa. The wings are most often hyaline, but sometimes smoky or dark colored, or partly infuscated in many genera or completely darkened.The abdomen consists of six to eight visible segments preceding the genitalia in males, but the eighth segment is sometimes entirely or partially concealed, and terminal forming the ovipositor. It is long and narrow conical in most species, but wide, dorsoventrally flattened and short in bee mimics. In the Leptogastrinae, the abdomen is extremely long and slender. In some tribes, the male undergoes axial torsion of 180°.
The egg is hyaline or pigmented, of variable shape from spherical to oval, and up to 2 mm in length. The surface can be smooth or bearmicrosculptures, which are generally polygonal and visible only in the electron microscope.
Thelarva is apodous, cylindrical, and elongated, more or less flattened dorsoventrally and tapered at the cephalic and caudal ends. The colour is white or yellowish. The head is small, rugged, dark-pigmented and hypognathous, while the abdomen is composed of eight apparent urites, with the last two often fused and more or less reduced. The respiratory system is amphineustic, with two pairs ofspiracles, one thoracic and one abdominal. Also, rudimentary and nonfunctional stigmata occur in other abdominal segments.
The pupa is naked, as in the majority of the Orthorrhapha,exarate and therefore able to move.
The Asilidae arepredators, in both the juvenile and the adult stages, and feed on small arthropods, mainly insects. Although predatory forms in the adult stage are present in other taxonomic groups of Diptera, the Asilidae are the most representative for the number of species and for uniformity of feeding behavior (>7000 species, all of which are predatory). The combination of high biodiversity and high predatory activity leads to this family playing an important role in theecological stability of entomofauna.
Thelife cycle takes place in 1–3 years. The postembryonic development consists of four larval stages (instars) and one pupa. The larvae of the first instar differ from other stages in both ethology and trophic regime. The larvae of most known asilids live in the soil or in the case of some taxonomic groups, in rotting organic material, usually wood and the bark of dead trees.
With regards to feeding behavior, most of the literature describes Asilidae larvae asentomophagous, but doubts remain about the real nature of the trophic regime and its mechanisms. The entomophagy of some species had indeed been already hypothesized by some authors of the 19th century, based on the findings of larvae of asilids associated with larvae of other insects, but Melin (1923) asserted that, in reality, predation was occasional and secondary to the plant-based diet. More recent studies have confirmed the entomophagy of some asilids without extending this species' feeding behavior for the whole family. Less certain, however, is the mechanism of entomophagy: in general, the behavior is cited as predation, but for some species may be ectoparasitoids. Musso (1983)[7] described the feeding behavior of the larvae ofMachimus rusticus: the larvae of the first instar does not feed on insects, those of the second instar feed on secretions by larvae of beetles (and may cause death), while the larvae of the third and fourth instars actually behave like predators. In short, the feeding behavior of larval asilids can be intermediate between predation and ectoparasitism.
Much better known and described in detail is the behavior of adults. In general, predation in adults is concentrated in the hottest hours in open, sunny spaces, while at night, they take refuge in dense vegetation. The Asilidae are excellent flyers, and in most of the family, capture prey in flight. They are often seen stationed to ambush prey at strategic points. This behavior signifies that sight plays an essential role in the detection of prey and their capture.
The prey is caught with the tarsi and injected with a paralyzing saliva. The asilid pierces the integument of the prey with the prepharynx (hypopharynx) in preferential points of least resistance such as the eyes, the membranous area of transition between the head and thorax (neck) or between thorax and abdomen, or between the lastabdominal tergites. Puncture is followed by the injection of saliva, whose active components perform two functions:neurotoxins cause paralysis of the victim, while proteolytic enzymes lead to the breakup and liquefaction of internal tissues. In a short time, the predator is able to feed by sucking the internal fluids through the alimentary canal.
With regards to interspecific trophic relationships, a large number of reports exists on the prey captured by the Asilidae. Lavigne (2003) has developed a database comprising over 13,000 reports.[8] The prey of Asilidae are predominantly represented by other insects, mostly winged, but several cases in which they have attacked spiders have also been reported. Within the insects, orders that include the most frequent prey of asilids include a wide range of families within theColeoptera,Hymenoptera, otherDiptera,Hemiptera, andLepidoptera; prey belonging to various other orders (Odonata,Neuroptera,Isoptera,Thysanoptera,Blattodea, etc.) are also mentioned.
With regards to the specificity of the trophic relationship, Wood (1981)[2] mentions some studies in the literature on the subject. Some genera have been found to be monophagic, but more generally Asilidae are polyphagic, with behaviors that vary from narrow specialization[8] to broad prey choice.[2]
Other studies[9] have shown that the ratio between the size of the prey and the asilid varies from 1.8:1 to 3.7:1, with an average of 2.6:1. The ratio tends to increase with decreasing size of the predator.
Egg-laying takes place, according to the species, with three different behaviors that relate to the structure and the morphology of the abdomen. Females with an undifferentiated ovipositor release eggs randomly and independently from the substrate. In other cases, however, the abdomen bears a differentiated, specialized ovipositor to lay eggs in the soil or sand, or lay them in cavities within plant tissues.
Asilidae generally occur in habitats that are open, sunny, and dry, even arid. They favour open or scattered vegetation, and some species even frequent bare ground. Typical habitats includesavannah,forest steppe, opensteppe, semidesert,maquis shrubland, and related shrubland types such asfynbos andchaparral. Their biodiversity is lower in forested ecosystems, and where asilids do occur in such environments, they tend to concentrate in the glades and margins. In those conditions, the interrupted canopy leaves space for various species of shrubs and herbaceous plants suited to asilid styles of predation.
In general, the biology of the Asilidae is still poorly known, but various authors have studied the population distribution in particular regions and ecosystems. They have classified the behavioral patterns in terms of microenvironments, ecological, and trophic factors, showing how different species of Asilidae favour particular habitats suited to particular patterns of reproduction and predation. Specific studies show correlations between the floristic composition and predatory behaviour.[10]
Asilidae occur in allzoogeographical regions except Antarctica. In the Northern Hemisphere, some species are even adapted to tundra.Alpine species occur at altitudes exceeding 4000 meters/13,000 feet. However, the highest levels of biodiversity are in warm climates; tropical or subtropical and arid or semi-arid regions tend to have the greatest variety of species, followed by areas where rainfall is highly seasonal.
The Asilidae currently include over 7500 describedspecies in about 556genera.[11] Their taxonomy is still under study in the light of new specimens and cladistic analysis. The 14 accepted subfamilies are:[12]
The oldest known member of the family isAraripogon from the Early Cretaceous (Aptian)Crato Formation of Brazil.[13]
| Asiloidea |
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Clade showing relationship of Asiloidea
Carl Linnaeus (Linné; 1758), in the 10th edition ofSystema Naturae, erected the genusAsilus, including 11 species, and added four others in the 12th edition (1767).Asilus crabroniformis (1758) is the type of the genus. The rank of family is credited toLeach in Samouelle (1819).Johan Christian Fabricius in five publications dated from 1775 to 1805, erected the genusDamalis and described 76 exotic and European species.Wiedemann, in publications appearing between 1817 and 1830, described 235 species, many exotic.Johann Wilhelm Meigen in an early work of 1803 erected four genera, three of which now represent subfamilies. He also described many species inSystematische Beschreibung der bekannten europäischen zweiflügeligen Insekten (1800 to 1838). During the rest of the 19th century, significant contributions were made byHermann Loew in particular. Other prominent authors dealing with the Asilidae during the 19th century includedPierre-Justin-Marie Macquart,Francis Walker,Camillo Rondani, andJacques-Marie-Frangile Bigot.
