Fly

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This article is about the flying insect. For other uses, see Fly (disambiguation), Flying (disambiguation), and Flies (disambiguation).
Fly
Temporal range: 245–0 Ma
Middle Triassic – Recent
Bessenbandzweefvlieg Vrouwtje (2).JPG
Syrphus ribesii, showing characteristic dipteran features: large eyes, small antennae, sucking mouthparts, single pair of flying wings, hindwings reduced to clublike halteres
Scientific classification e
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Superorder: Panorpida
(unranked): Antliophora
Order: Diptera
Linnaeus, 1758

True flies are insects of the order Diptera, the name being derived from the Greek di = two, and ptera = wings. Insects of this order use only a single pair of wings to fly, the hindwings being reduced to club-like balancing organs known as halteres. Diptera is a large order containing an estimated 1,000,000 species including horse-flies,[a] crane flies, hoverflies and others, although only about 150,000 species have been described.

Flies have a mobile head, with a pair of large compound eyes, and mouthparts designed for piercing and sucking (mosquitoes, black flies and robber flies), or for lapping and sucking in the other groups. Their wing arrangement gives them great manoeuvrability in flight, and claws and pads on their feet enable them to cling to smooth surfaces. Flies undergo complete metamorphosis; the eggs are laid on the larval food-source and the larvae, which lack true limbs, develop in a protected environment, often inside their source of their food. The pupa is a tough capsule from which the adult emerges when ready to do so; flies mostly have short lives as adults.

Diptera is one of the major insect orders and has high ecological and human importance. Fruit flies are used as model organisms in research, but less benignly, mosquitoes are vectors for malaria, dengue, West Nile fever, yellow fever, encephalitis, and other infectious diseases, and houseflies spread food-borne illnesses. Larger flies such as tsetse fly and screwworm cause significant economic harm to cattle. Blowfly larvae, known as gentles, and other dipteran larvae, known more generally as maggots, are used as fishing bait and as food for carnivorous animals. In medical debridement, they are used to clean wounds.

Taxonomy and phylogeny[edit]

External relationships[edit]

Dipterans are Endopterygota, insects that undergo radical metamorphosis. They belong to the Mecopterida, alongside the Mecoptera, Siphonaptera, Lepidoptera and Trichoptera.[3] The presence of a single pair of wings distinguishes most true flies from other insects with "fly" in their names. However, some true flies such as Hippoboscidae (louse flies) have become secondarily wingless.[4]

The cladogram is from the work of David K. Yeates and Brian Wiegmann.[5]

part of Endopterygota
Mecopterida
Antliophora

Diptera Common house fly, Musca domestica.jpg


Mecoptera (scorpionflies, hangingflies) Gunzesrieder Tal Insekt 3.jpg


Siphonaptera (fleas) Flea (251 01) Aphaniptera; total preparation.jpg




Trichoptera (caddisflies) Sericostoma.personatum.jpg


Lepidoptera (butterflies and moths) Tyria jacobaeae-lo.jpg




Hymenoptera (sawflies, wasps, ants, bees) AD2009Sep09 Vespula germanica 03.jpg


Fossil Nematoceran in Dominican amber. Sandfly, Lutzomyia adiketis (Psychodidae), Early Miocene, c. 20 million years ago

Internal relationships[edit]

Fossil Brachyceran in Baltic amber. Lower Eocene, c. 50 million years ago

The first true dipterans known are from the Middle Triassic (around 240 million years ago), and they became widespread during the Middle and Late Triassic.[6] The basal clades in the Diptera are the Deuterophlebiidae and Nymphomyiidae.[7] Three episodes of evolutionary radiation occurred. Many new species of lower Diptera developed in the Triassic, about 220 million years ago. Many lower Brachycera appeared in the Jurassic, some 180 million years ago. A third radiation took place among the Schizophora at the start of the Paleogene, 66 million years ago.[7]

Diptera is traditionally broken down into two suborders, Nematocera and Brachycera. The Nematocera are recognized by their elongated bodies and many-segmented, often feathery antennae as represented by mosquitoes and crane flies. The Brachycera have rounder bodies and much shorter antennae.[8][9]

The following cladogram is from the work of David K. Yeates, Rudolf Meier and Brian Wiegmann. It places the Brachycera within the Nematocera.[10]

Nematocera


Ptychopteromorpha (phantom and primitive crane-flies) Ptychoptera contaminata.jpg


Culicomorpha (mosquitoes) AnophelesGambiaemosquito.jpg




Blephariceromorpha Imago of Blepharicera fasciata as Asthenia fasciata in Westwood 1842, plate 94.png



Bibionomorpha (gnats) Bibio marci02.jpg



Psychodomorpha Clogmia Albipunctata or moth fly.jpg



Tipuloidea (crane flies) Tipula submarmorata, Abergwynant, North Wales, May 2015 (23422515666).jpg

Brachycera
Tab

Stratiomyomorpha Hermetia illucens Black soldier fly edit1.jpg



Xylophagomorpha (stink flies, etc) Stinkfliege Coenomyia ferruginea male.jpg


Tabanomorpha (horse flies, snipe flies, etc) Tabanus bromius01.jpg



Mus

Nemestrinoidea



Asiloidea (robber flies, bee flies, etc) Asilidae June 2011-1.jpg

Ere

Empidoidea (dance flies, etc) Empis.tessellata.male.jpg

Cyc

Aschiza (in part)



Phoroidea (flat-footed flies, etc) Polyporivora-picta-Platypezid-fly-20111015a.jpg



Syrphoidea (hoverflies) Mosca cernidora de la grosella.jpg

Sch
Cal

Hippoboscoidea (louse flies, etc) CrataerhinaPallida.jpg



Muscoidea (house flies, dung flies, etc) Musca domestica housefly.jpg


Oestroidea (blow flies, flesh flies, etc) Sarcophaga Bercaea2.jpg




Acalyptrata (marsh flies, etc) Marsh fly01.jpg














Abbreviations used in the cladogram:

Diversity[edit]

Flies are often abundant and are found in almost all terrestrial habitats in the world apart from Antarctica. They include many familiar insects such as house flies, blow flies, mosquitoes, gnats, black flies, midges and fruit flies. About 150,000 have been formally described and an even larger number has still to be studied intensively.[11][12] The suborder Nematocera include generally small, slender insects with long antennae such as mosquitoes, gnats, midges and crane-flies, while the Brachycera includes broader, more robust flies with short antennae.[13] There are estimated to be a total of about 19,000 species of Diptera in Europe, 22,000 in the Nearctic region, 20,000 in the Afrotropical region, 23,000 in the Oriental region and 19,000 in the Australasian region.[14]

Brachycera are ecologically very diverse, with many being predatory at the larval stage and some being parasitic. Animals parasitised include molluscs, woodlice, millipedes, insects, mammals,[14] and amphibians.[15] The larvae of Megaselia scalaris are almost omnivorous and consume such substances as paint and shoe polish.[16] The larvae of the shore flies (Ephydridae) survive in extreme environments including hot springs, geysers, saline pools, sulphur pools, septic tanks and even crude oil.[14] Adult hoverflies (Syrphidae) are well known for their mimicry and the larvae adopt diverse lifestyles including being inquiline scavengers inside the nests of social insects.[17] Some brachycerans are agricultural pests, some bite animals and humans and suck their blood, and some transmit diseases.[14]

Anatomy and morphology[edit]

See also: Morphology of Diptera and Biology of Diptera

Flies are adapted for aerial movement and typically have short and streamlined bodies. The first tagma of the fly, the head, bears the eyes, the antennae, and the mouthparts (the labrum, labium, mandible, and maxilla make up the mouthparts). The second tagma, the thorax, bears the wings and contains the flight muscles on the second segment, which is greatly enlarged; the first and third segments have been reduced to collar-like structures, and the third segment bears the halteres, which help to balance the insect during flight. The third tagma is the abdomen consisting of 11 segments, some of which may be fused, and with the 3 hindermost segments modified for reproduction.[18]

Head of a horse-fly showing large compound eyes and stout piercing mouthparts

Flies have a mobile head with a pair of large compound eyes on the sides of the head, and in most species, three small ocelli on the top. The compound eyes may be close together or widely separated, and in some instances are divided into a dorsal region and a ventral region, perhaps to assist in swarming behaviour. The antennae are well-developed but variable, being thread-like, feathery or comb-like in the different families. The mouthparts are adapted for piercing and sucking, as in the black flies, mosquitoes and robber flies, and for lapping and sucking as in many other groups.[18] Female horse-flies use knife-like mandibles and maxillae to make a cross-shaped incision in the host's skin and then lap up the blood that flows. The gut includes large diverticulae, allowing the insect to store small quantities of liquid after a meal.[19]

For visual course control, flies' optic flow field is analyzed by a set of motion-sensitive neurons.[20] A subset of these neurons is thought to be involved in using the optic flow to estimate the parameters of self-motion, such as yaw, roll, and sideward translation.[21] Other neurons are thought to be involved in analyzing the content of the visual scene itself, such as separating figures from the ground using motion parallax.[22][23] The H1 neuron is responsible for detecting horizontal motion across the entire visual field of the fly, allowing the fly to generate and guide stabilizing motor corrections midflight with respect to yaw.[24]

A cranefly, showing the hind wings reduced to drumstick-shaped halteres

Diptera have one pair of fore wings on the mesothorax and a pair of halteres, or reduced hind wings, on the metathorax. A further adaptation for flight is the reduction in number of the neural ganglia, and concentration of nerve tissue in the thorax, a feature that is most extreme in the highly derived Muscomorpha infraorder.[19] Some species of flies are exceptional in that they are secondarily flightless. The only other order of insects bearing a single pair of true, functional wings, in addition to any form of halteres, are the Strepsiptera. In contrast to the flies, the Strepsiptera bear their halteres on the mesothorax and their flight wings on the metathorax.[25] Each of the fly's six legs has a typical insect structure of coxa, trochanter, femur, tibia and tarsus, with the tarsus in most instances being subdivided into five tarsomeres.[18] At the tip of the limb is a pair of claws, and between these are cushion-like structures known as pulvilli which provide adhesion.[26]

The abdomen shows considerable variability among members of the order. It consists of eleven segments in primitive groups and ten segments in more derived groups, the tenth and eleventh segments having fused.[27] The last two or three segments are adapted for reproduction. Each segment is made up of a dorsal and a ventral sclerite, connected by an elastic membrane. In some females, the sclerites are rolled into a flexible, telescopic ovipositor.[18] For detailed anatomy, the Anatomical Atlas of Flies gives examples from the four major fly groups (Lower Diptera, Lower Brachycera, Acalyptrate, Calyptrate) representing different anatomical expressions.[28]

Life cycle and development[edit]

Mating anthomyiid flies

Diptera go through a complete metamorphosis (egg, larva, pupa, adult). In many flies, the larval stage is predominant and the adults may have a short life. Most dipteran larvae develop in protected environments; many are aquatic and others are found in moist places such as carrion, fruit, vegetable matter, fungi and, in the case of parasitic species, inside their hosts. They tend to have thin cuticles and become desiccated if exposed to the air. Apart from the Brachycera, most dipteran larvae have sclerotinised head capsules, which may be reduced to remnant mouth hooks; the Brachycera however have soft, gelatinized head capsules from which the sclerites are reduced or missing. Many of these larvae retract their heads into their thorax.[29][18]

Life cycle of stable fly Stomoxys calcitrans, showing eggs, 3 larval instars, pupa, and adult

Some other anatomical distinction exists between the larvae of the Nematocera and the Brachycera. Especially in the Brachycera, little demarcation is seen between the thorax and abdomen, though the demarcation may be visible in many Nematocera, such as mosquitoes; in the Brachycera, the head of the larva is not clearly distinguishable from the rest of the body, and few, if any, sclerites are present. Informally, such brachyceran larvae are called maggots,[30] but the term is not technical and often applied indifferently to fly larvae or insect larvae in general. The eyes and antennae of brachyceran larvae are reduced or absent, and the abdomen also lacks appendages such as cerci. This lack of features is an adaptation to food such as carrion, decaying detritus, or host tissues surrounding endoparasites.[19] Nematoceran larvae generally have well-developed eyes and antennae, while those of Brachyceran larvae are reduced or modified.[31]

Dipteran larvae have no jointed, "true legs",[29] but some dipteran larvae, such as species of Simuliidae, Tabanidae and Vermileonidae, have prolegs adapted to hold onto a substrate in flowing water, host tissues or prey.[32] The majority of dipterans are oviparous and lay batches of eggs, but some species are ovoviviparous, the larvae starting development inside the eggs before they hatch. In Hylemya strigosa the larva moults to the second instar before hatching, and in Termitoxenia, it hatches as the third instar larva and almost immediately pupates.[33] The tsetse fly exhibits adenotrophic viviparity; a single fertilised egg is retained in the oviduct and the developing larva feeds on glandular secretions. When fully grown, the female finds a spot with soft soil and the larva works its way out of the oviduct, buries itself and pupates.[34]

The pupae take various forms. In some groups, particularly the Nematocera, the pupa is intermediate between the larval and adult form; these pupae are described as "obtect", having the future appendages visible as structures that adhere to the pupal body. The outer surface of the pupa may be leathery and bear spines, respiratory features or locomotory paddles. In other groups, described as "coarctate", the appendages are not visible. In these, the outer surface is a puparium, formed from the last larval skin, and the actual pupa is concealed within. When the adult insect is ready to emerge from this tough, desiccation-resistant capsule, it inflates a balloon-like structure on its head, and forces its way out.[18]

The adult stage is usually short, its function only to mate and lay eggs. The genitalia of female flies are rotated to a varying degree from the position found in other insects. In some flies, this is a temporary rotation during mating, but in others, it is a permanent torsion of the organs that occurs during the pupal stage. This torsion may lead to the anus being below the genitals, or, in the case of 360° torsion, to the sperm duct being wrapped around the gut and the external organs being in their usual position. When flies mate, the male initially flies on top of the female, facing in the same direction, but then turns around to face in the opposite direction. This forces the male to lie on his back for his genitalia to remain engaged with those of the female, or the torsion of the male genitals allows the male to mate while remaining upright. This leads to flies having more reproduction abilities than most insects, and at a much quicker rate. Flies occur in large populations due to their ability to mate effectively and in a short period of time during the mating season.[19]

Interactions with humans[edit]

In culture[edit]

Further information: Insects in culture
Petrus Christus's 1446 painting Portrait of a Carthusian has a fly painted on a trompe l'oeil frame.

In the traditional Navajo religion, Big Fly is an important spirit being.[35][36][37]

An Anopheles stephensi mosquito drinking human blood. The species carries malaria.

Economic importance[edit]

Dipterans are an important group of insects and have a considerable impact on the environment. Some leaf-miner flies (Agromyzidae), fruit flies (Tephritidae and Drosophilidae) and gall midges (Cecidomyiidae) are pests of agricultural crops; others such as tsetse flies, screwworm and botflies (Oestridae) attack livestock, causing wounds, spreading disease, and creating significant economic harm. Still others such as mosquitoes (Culicidae), blackflies (Simuliidae) and drain flies (Psychodidae) impact on human health, acting as vectors of major tropical diseases. Among these, Anopheles mosquitoes transmit malaria, filariasis, and arboviruses; Aedes aegypti mosquitoes carry dengue fever and the Zika virus; blackflies carry river blindness; sand flies carry leishmaniasis. Other dipterans are a nuisance to humans, especially when present in large numbers; these include houseflies, which contaminate food and spread food-borne illnesses; the biting midges and sandflies (Ceratopogonidae) and the houseflies and stable flies (Muscidae).[18]

Many dipterans serve useful roles. Houseflies, blowflies and fungus gnats (Mycetophilidae) are scavengers and aid in decomposition. Robber flies (Asilidae), tachinids (Tachinidae) and dagger flies and balloon flies (Empididae) are predators and parasitoids of other insects, helping to control a variety of pests. Many dipterans such as bee flies (Bombyliidae) and hoverflies (Syrphidae) are pollinators.[18]

Uses[edit]

Diptera in research: Drosophila melanogaster fruit fly larvae being bred in tubes in a genetics laboratory

Drosophila melanogaster, a fruit fly, has long been used as a model organism in research because of the ease with which it can be bred and reared in the laboratory, its small genome, and the fact that many of its genes have counterparts in higher eukaryotes. A large number of genetic studies have been undertaken based on this species; these have had a profound impact on the study of gene expression, gene regulatory mechanisms and mutation. Other studies have investigated physiology, microbial pathogenesis and development among other research topics.[38]

Blowflies feeding on the fresh corpse of a porcupine, Hystrix africaeaustralis

Maggots found on corpses are useful to forensic entomologists. Maggot species can be identified by their anatomical features and by matching their DNA. Maggots of different species of flies visit corpses and carcases at fairly well-defined times after the death of the victim, and so do their predators, such as beetles in the family Histeridae. Thus, the presence or absence of particular species provides evidence for the time since death, and sometimes other details such as the place of death, when species are confined to particular habitats such as woodland.[39]

Maggots used as animal feed at London Zoo

Some species of maggots such as blowfly larvae (gentles) and bluebottle larvae (casters) are bred commercially; they are sold as bait in angling, and as food for carnivorous animals (kept as pets, in zoos, or for research) such as some mammals,[40] fishes, reptiles, and birds[41] including poultry.[42] It has been suggested that fly larvae could be used at a large scale as food for farmed chickens, pigs, and fish. However, consumers are opposed to the inclusion of insects in their food, and the use of insects in animal feed remains illegal in areas such as the European Union.[43][44]

Casu marzu is a traditional Sardinian sheep milk cheese that contains larvae of the cheese fly, Piophila casei.

Fly larvae can be used as a biomedical tool for wound care and treatment. Maggot debridement therapy (MDT) is the use of blow fly larvae to remove the dead tissue from wounds, most commonly being amputations. Historically, this has been used for centuries, both intentional and unintentional, on battlefields and in early hospital settings.[45] Removing the dead tissue promotes cell growth and healthy wound healing. The larvae also have biochemical properties such as antibacterial activity found in their secretions as they feed.[46] These medicinal maggots are a safe and effective treatment for chronic wounds.[47]

The Sardinian cheese casu marzu is exposed to flies known as cheese skippers such as Piophila casei, members of the family Piophilidae. The digestive activities of the fly larvae soften the cheese and modify the aroma as part of the process of maturation. At one time European Union authorities banned sale of the cheese and it was becoming hard to find,[48] but the ban has been lifted on the grounds that the cheese is a traditional local product made by traditional methods.[49]

Notes[edit]

  1. ^ Some authors draw a distinction in writing the common names of insects. True flies are in their view best written as two words, such as crane fly, robber fly, bee fly, moth fly, and fruit fly. In contrast, common names of non-dipteran insects that have "fly" in their names are written as one word, e.g. butterfly, stonefly, dragonfly, scorpionfly, sawfly, caddisfly, whitefly.[1] In practice, however, this is a comparatively new convention; especially in older books, names like "saw fly" and "caddis fly", or hyphenated forms such as house-fly and dragon-fly are widely used.[2] In any case, non-entomologists cannot, in general, be expected to tell dipterans, "true flies", from other insects, so it would be unrealistic to expect rigour in the use of common names. Also, exceptions to this rule occur, such as the hoverfly, which is a true fly, and the Spanish fly, a type of blister beetle.

References[edit]

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  2. ^ Comstock, J. H. An Introduction to Entomology, Comstock Publishing. 1949. May be downloaded from: [1]
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  27. ^ Gibb, Timothy J.; Oseto, Christian (2010). Arthropod Collection and Identification: Laboratory and Field Techniques. Academic Press. p. 189. ISBN 978-0-08-091925-6. 
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  35. ^ Leland Clifton Wyman (1983). "Navajo Ceremonial System". Handbook of North American Indians (PDF). Humboldt State University. p. 539. Nearly every element in the universe may be thus personalized, and even the least of these such as tiny Chipmunk and those little insect helpers and mentors of deity and man in the myths, Big Fly (Dǫ’ soh) and Ripener (Corn Beetle) Girl (’Anilt’ ánii ’At’ ééd) (Wyman and Bailey 1964:29–30, 51, 137–144), are as necessary for the harmonious balance of the universe as is the great Sun. 
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Further reading[edit]

  • Blagoderov, V.A., Lukashevich, E.D. & Mostovski, M.B. 2002. Order Diptera. In: Rasnitsyn, A.P. and Quicke, D.L.J. The History of Insects, Kluwer pp.–227–240.
  • Colless, D.H. & McAlpine, D.K. 1991 Diptera (flies), pp. 717–786. In: The Division of Entomology. Commonwealth Scientific and Industrial Research Organisation, Canberra (spons.), The insects of Australia. Melbourne University Press.
  • Hennig, Willi Diptera (Zweifluger). Handb. Zool. Berl. 4 (2) (31):1–337. General introduction with key to World Families. In German.
  • Oldroyd, Harold The Natural History of Flies. W. W. Norton. 1965.
  • Séguy, Eugène Diptera: recueil d'etudes biologiques et systematiques sur les Dipteres du Globe (Collection of biological and systematic studies on Diptera of the World). 11 vols. Part of Encyclopedie Entomologique, Serie B II: Diptera. 1924–1953.
  • Séguy, Eugène La Biologie des Dipteres 1950.
  • Thompson, F. Christian. "Sources for the Biosystematic Database of World Diptera (Flies)" (PDF). United States Department of Agriculture, Systematic Entomology Laboratory. 

External links[edit]

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