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Annotated classification
- Order Diptera
- Size range 1 mm to 7.5 cm; wings, when present, number 2; hind wings reduced to halteres; sucking mouthparts; 125,000 species; worldwide distribution; diverse habitats and diets in both larvae and adults.
- Suborder Nematocera
- Antennae consist of scape, pedicel, and flagellum with numerous similar segments; maxillary palpi with more than 3 segments, often pendulous; anal cell of wing open; larvae usually with well-defined head, mandibles horizontally opposed.
- Family Tipulidae (crane flies)
- Elongated body, wings, legs; slow-flying; larvae in soil (leatherjackets), moss, rotten wood, mud, fresh water, littoral, marine.
- Family Mycetophilidae (fungus gnats)
- Fragile, slender; flit about in damp, shady places, among decaying vegetation.
- Family Sciaridae
- Similar to fungus gnats but more compact, more often indoors.
- Family Bibionidae (march flies in Northern Hemisphere)
- Compact, well-armoured flies; strong spurs on legs; often abundant on spring blossoms; larvae in soil, sometimes found in a tangled mass near roots of plants.
- Family Scatopsidae
- Similar to march flies, more often indoors.
- Family Cecidomyiidae (gall midges)
- Tiny flies seldom seen as adults; shapeless larvae burrow into plant tissues, cause formation of plant galls, and deform leaves, stems, and roots; some horticultural and agricultural pests.
- Family Psychodidae (moth flies)
- Tiny, with hairy wings; often seen singly in kitchens, on windows above sinks; some larvae numerous in sewage sedimentation tanks; larvae mostly aquatic.
- Family Phlebotomidae (sand flies)
- Closely related to Psychodidae; adult females suck blood, carry dermal and intestinal leishmaniasis and sandfly fever.
- Family Ceratopogonidae (biting midges)
- Tiny, often with spotted wings (e.g., Culicoides); adult females with irritating bite suck blood, carry some parasitic worms; Forcipomyia suck blood of insects.
- Family Chironomidae (nonbiting midges)
- Related to biting midges, but females do not suck blood; larvae aquatic; important fish food; adults swarm near water.
- Family Simuliidae (black flies)
- Also buffalo gnats; small, humpbacked, with short antennae; females suck blood, carry parasitic worms that cause “river blindness”; forms nodules under skin; larvae aquatic, filter feeders, attached to stones, underwater vegetation, or freshwater crustaceans.
- Family Culicidae (mosquitoes)
- Small; elongated; proboscis prominent; palpi often long; best recognized by scaly wings; many females suck blood, carry human diseases (Anophelini carry malaria; Culicini carry yellow fever, filariasis, dengue, viral encephalitis); larvae and pupae aquatic.
- Suborder Brachycera
- Flagellum of antennae nearly always fused into a compound 3rd segment, remaining diminutive segments form a stumpy “style” or bristle-like arista; anal cell of wing narrowed, nearly always closed on or before wing margin; palpi seldom with more than 3 segments, often 2 or 1, held forward (porrect); larvae usually with well-defined head, mandibles move vertically or parallel, cannot be opposed; adult escapes from pupa by a rectangular slit (“Orthorrhapha”).
- Family Stratiomyidae (soldier flies)
- Colourful flies, found resting on vegetation with wings closed; males sometimes dance in air; larvae sometimes elongate, aquatic, active, carnivorous (Stratiomys); others in decaying vegetation (Hermetia).
- Family Rhagionidae (snipe flies)
- Inconspicuous, usually rest on vegetation; some females (e.g., Symphoromyia) suck blood; most larvae in soil or in water (some Atherix females form egg-laying swarms); some make pits in dust, like ant lions (Vermileo).
- Family Pantophthalmidae
- Large, archaic flies, now found only in tropical forests of South America; wood-boring larval grubs sometimes damage commercial timber.
- Family Tabanidae (horseflies, deerflies; march flies in Australia)
- Squat flies with big heads, brilliantly coloured eyes; some females (Chrysops, Tabanus, Haematopota) suck blood, are livestock pests; many primitive genera feed only from flowers; larvae in mud or wet soil, either vegetarian (Chrysops) or carnivorous (Tabanus, Haematopota).
- Family Asilidae (robber flies)
- Adults catch other insects in flight, suck their blood; size varies from a few millimetres to 8 cm (longest of all flies); characteristic “moustache” of bristles probably protects eyes from damage by fly’s victim; larvae in soil or wood; eat any food.
- Family Bombyliidae (bee flies)
- Hairy, scaly; superficially resemble bees, hover over flowers in similar way; often brightly patterned, pattern destroyed by rubbing scales; larvae scavenge in bee and wasp nests or are parasitic (e.g., locust egg pods, tsetse pupae).
- Family Scenopinidae (window flies)
- Tiny black flies, on windows indoors; develop from larvae in carpets, feed on flea and clothes moth larvae; natural habitat, bird nests or similar dry debris.
- Family Therevidae (stiletto flies)
- Adults resemble Asilidae, but not predatory; larvae like Scenopinidae, elongated, worm-like, carnivorous but sometimes attack plant roots.
- Family Nemestrinidae
- Rather like Bombyliidae; larvae parasitic in grasshoppers, locusts, perhaps beetles; remarkable for beautiful hovering.
- Family Acroceridae (balloon flies)
- Grotesque; abdomen swollen, thorax small, head tiny; larvae parasitic in spiders.
- Family Dolichopodidae (long-legged flies)
- Tiny, metallic, bristly flies; large numbers sit on leaves in wet places; predatory on other insects; larvae like Empididae, elongated, with little external head structure, same habitats.
- Suborder Brachycera-Cyclorrhapha
- Usually shortened to Cyclorrhapha; characteristically form pupa inside last larval skin as a puparium; adult fly pushes off a circular cap, hence the name Cyclorrhapha; most families (Schizophora) with a ptilinum (membranous sac inside head), which emerges from a horseshoe-shaped ptilinal suture (identifies adult Schizophora) above antennae, is puffed in and out to help fly escape from puparium or soil or to inflate fly’s body; ptilinum atrophies and only ptilinal suture remains; a small group of Aschiza, without ptilinal suture, are recognized chiefly by their wing venation.
- Series Aschiza
- Family Lonchopteridae
- Little known; notable for parthenogenesis; few species; worldwide; sometimes abundant.
- Family Phoridae (coffin flies)
- Tiny flies sometimes numerous indoors; larvae live in any organic debris rich in protein or nitrogenous decay products and scavenge in nests of wasps, bees, ants, termites; breed in carrion; many adults wingless or with short wings (brachypterous).
- Family Pipunculidae
- Tiny flies; head spherical, noted for precise hovering; larvae parasitic in Homoptera.
- Family Platypezidae
- Small flies; peculiar legs; rarely seen; appear to dance in smoke of wood fires; larvae live in fungi.
- Family Syrphidae (hover flies)
- Vena spuria in wing runs between third and fourth veins; familiar everywhere; hover over flowers, settle on leaves; some larvae aquatic (“rat-tailed” maggots); larvae of many species feed on aphids on plant stems and leaves.
- Family Conopidae (thick-headed flies)
- Wasplike flies; larvae parasitic in bees and wasps; may be a separate evolutionary line.
- Series Schizophora
- All flies with a ptilinal suture in head; larvae with no external head structure, mouth hooks visible through cuticle, one pair of prothoracic spiracles and one pair of posterior spiracles, each with either three slits or a mass of small pores; larvae with fore end pointed and hind end truncate are called maggots; larvae with both ends blunt and fleshy, with bulges and tracts of spines, are called grubs.
- Section Acalyptrata
- Thoracic squamae (i.e., calypters that join base of wing to thorax) are small or evanescent; small soft-bodied flies; major families well established; placement of genera uncertain; families can be grouped according to food preferences of larvae.
- Flies breeding in vegetable compost and dung
- Family Lauxaniidae
- Larvae in decaying vegetable matter.
- Family Helomyzidae
- Like Lauxaniidae; most generalized of Acalyptrata.
- Family Dryomyzidae
- Like Lauxaniidae, but with wider range of food, including fungi; yellow flies often seen in winter.
- Family Chyromyiidae
- Yellow flies, 1 or 2 mm long; breed in debris of bird nests, mammal burrows, caves, cellars; seen singly on windows indoors.
- Family Celyphidae (beetle flies)
- Scutellum enormously enlarged until it covers both abdomen and wings when at rest; tropical dung breeding.
- Family Mormotomyiidae
- Contains one wingless, African species; looks like a spider; known from only one locality in Kenya; breeds in bat dung.
- Family Coelopidae (kelp flies, seaweed flies)
- Breed in wrack (i.e., heaps of decaying seaweed stranded on beaches) chiefly in temperate countries; adults of some species attracted by trichloroethylene; sometimes pests.
- Flies breeding in animal refuse, dung, carrion
- Family Sepsidae
- Small, black, roundhead flies, sometimes with spots at wing tips; may breed to infestation level in sewage sludge.
- Family Piophilidae (cheese skippers)
- Larvae in cheese, ham, cured meats, dried fruits, preserved skins and pelts; natural habitat in mummifying carrion; called “skippers” because larvae move both by crawling and “skipping” (i.e., gripping the tip of the abdomen with mouth hooks and flipping the body through a relatively long distance).
- Family Micropezidae
- Large, long-legged flies; often with conspicuously patterned, blue-black wings; spectacular in tropics.
- Family Sphaeroceridae
- Tiny, black-brown flies; first tarsal segments of hind legs swollen; abundant throughout world in dunglike materials; some members live in seaweed on beaches; many short-winged or wingless species.
- Family Sciomyzidae
- Aquatic larvae eat both living and dead snails; may be valuable as controlling agents for injurious snails.
- Family Milichiidae
- Breed in dung; adults attach to predatory insects and spiders and feed on them; called “insect jackals”; Madiza glabra sometimes numerous indoors.
- Family Carnidae
- Scavenge in nests and burrows. Adults of Carnus hemapterus scavenge among bird feathers, break off wings.
- Family Neottiophilidae
- Nest-breeding; larvae suck blood of nestling birds.
- Family Thyreophoridae
- Among the rarest of flies; larvae in dead bodies of large animals.
- Family Chamaemyiidae
- Predatory larvae; known as controlling agents of aphids.
- Family Braulidae (bee louse)
- Braula caeca, wingless fly, lives in beehives; larva feeds on wax and pollen stores; adult attaches to bee, may solicit nutritious saliva like other members of bee colony.
- Flies with plant-feeding larvae
- Family Ephydridae (shore flies)
- Transitional; wide range of larval habitats; no substance unpalatable for larvae (e.g., algae, sewage, excrement, carrion, urine, brine, hot springs, tar pools); carnivorous petroleum fly (Psilopa petrolei) lives in pools of crude petroleum seepage preying on trapped insects; many larvae feed in terrestrial and aquatic plants.
- Family Diopsidae (stalkeyed flies)
- Some larvae live in decaying plant tissue, others mine in living plants.
- Family Chloropidae (frit flies)
- Most important plant feeders; includes economic pests of cereal and other crops.
- Families Opomyzidae, Geomyzidae, Psilidae
- Small, usually yellow or grayish flies, plant feeders; Psila rosae, the carrot fly, an agricultural pest.
- Family Agromyzidae (leaf miners)
- Larvae feed in parenchymatous tissue of leaves, render epidermis transparent and produce either serpentine or “blotch” mines; rarely cause severe damage, but disfigure ornamental trees and shrubs.
- Flies with fruit-feeding larvae
- Family Trypetidae (large fruit flies)
- Form galls in certain flowers particularly Compositae; many Trypetidae larvae feed in living fruits, and ruin them; now worldwide distribution; economic damage by several members (e.g., the Mediterranean fruit fly Ceratitis capitata) has resulted in worldwide quarantine laws to regulate entry of fruit into countries.
- Family Drosophilidae (small fruit flies)
- Larvae in decaying and fermenting fruit or any sweet substance; includes Drosophila melanogaster, used in genetic studies.
A number of smaller families have been formed to accommodate genera closely related to the two above. Otitidae (Ortalidae) and Lonchaeidae are the most clearly defined. Others such as Ulidiidae, Pallopteridae, Phytalmidae, Camillidae, and Diastatidae are debatable.
- Section Calyptrata
- Characterized by large squamae (calypters that join base of wing to thorax); Scatophagidae are transitional.
- Family Scatophagidae (dung flies)
- Live around dung, other decaying materials; many also predacious as larvae and as adults.
- Family Muscidae (housefly and allies)
- Many species include the housefly; some larvae carnivorous, especially in third instar; breed in decaying vegetable matter or dung; larvae of Fannia, the “lesser housefly” like materials soaked in urine; economically important muscid larvae feed on plant stems and roots; subfamily (sometimes a separate family) Anthomyiinae contains dipteran plant pests; stable fly, Stomoxys, (biting proboscis in both sexes) may be placed in a separate family, Stomoxyidae; tsetse fly Glossina, confined to Africa, peculiar structurally and biologically, sometimes placed in the family Glossinidae, occurred in North America in the Miocene.
- Family Calliphoridae (blow flies)
- Some bristly flies with carrion-feeding maggots; common blow flies, Calliphora (bluebottles), feed as larvae in dead meat; Lucilia (greenbottles) sometimes attack living flesh; screw-worms (e.g., Cochliomyia, Callitroga) are dangerous feeders in living tissue.
- Family Cuterebridae
- Offshoot of Calliphoridae above; larvae are parasitic in rodents; one larva, Dermatobia hominis (human bot fly) also attacks man; eggs sometimes attached to mosquitoes and other biting flies and carried to their prospective prey.
- Family Oestridae (bots and warbles)
- Larvae live under skin, in nose, and in other head cavities of large mammals; includes the sheep nostril fly (Oestrus ovis), warble flies of cattle (Hypoderma bovis and other species).
- Family Gasterophilidae (horse bots)
- Larvae live in stomachs of horses, zebras, rhinos and elephants, attached to intestinal lining; relationship with other bot flies problematical; currently classified with other bot flies.
- Family Sarcophagidae (flesh flies)
- Large, gray and black; common around refuse dumps; larval habits diverse, in living or dead animal matter; many viviparous species.
- Family Tachinidae (tachinid flies)
- Ecologically important in balance of nature because larvae are parasites in other insects, spiders, woodlice, and centipedes; employed in biological control of pests.
- Section Pupipara
- Disputed group, families may merely be convergent in habit; lay living larvae, adults of both sexes feed exclusively on blood.
- Family Hippoboscidae (louse flies)
- Feed as adults on blood of mammals and birds; many fly, some have wings reduced or lost (e.g., sheep ked, Melophagus ovinus).
- Family Streblidae (bat flies)
- Distinct, rounded head, wings often functional but fly little; cling closely to host.
- Family Nycteribiidae (wingless bat flies)
- Always wingless; thorax weakened and de-sclerotized; live exclusively on bats; scarcely recognizable as flies.
Critical appraisal
Although there is general agreement concerning major groups of Diptera, disputes concerning relatively minor problems are not uncommon. After extensive study of relationships among families, probable lines of evolution within the order were traced in 1958. The order was surveyed according to the evidence of paleontology, and many fossil flies were illustrated in 1964; this resulted in subdividing the order into an unusually large number of families. Evolution of flesh-feeding maggots and classification and probable evolution of Oestridae were also being investigated. In later decades, many of the original groupings of dipterans (e.g., Nematocera) were recognized as being paraphyletic (unrelated).
Harold Oldroyd