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53-452: The Tephritidae are one of two fly families referred to as fruit flies , the other family being the Drosophilidae . The family Tephritidae does not include the biological model organisms of the genus Drosophila (in the family Drosophilidae), which is often called the "common fruit fly". Nearly 5,000 described species of tephritid fruit fly are categorized in almost 500 genera of

106-399: A dorsal preapical bristle. The female has an oviscape. The larva is amphipneustic (having only the anterior and posterior pairs of spiracle). The body varies from white to yellowish or brown. The posterior end of pale-coloured species is sometimes black. The body tapers at the anterior. The two mandibles sometimes have teeth along the ventral margin. The antennomaxillary lobes at each side of

159-432: A few insects leave them in the wounds. For example, of the 20,000 species of bees worldwide, only the half-dozen species of honeybees ( Apis ) are reported to have a barbed stinger that cannot be withdrawn; of wasps, nearly all are reported to have smooth stingers with the exception of two species, Polybia rejecta and Synoeca surinama . A few non-insect arthropods, such as scorpions , also sting. Among arthropods ,

212-454: A humeral and a subcostal break. The apical part of the subcostal is usually indistinct or even transparent and at about a right angle with respect to the basal part. Crossvein BM-Cu is present; the cell cup (posterior cubital cell or anal cell) is closed and nearly always narrowing to an acute angle. It is closed by a geniculated vein (CuA2). The CuA2 vein is rarely straight or convex. The tibiae lack

265-456: A length of up to 7 cm (2.8 in), is generally considered to be the largest fly in the world, while the smallest is Euryplatea nanaknihali , which at 0.4 mm (0.016 in) is smaller than a grain of salt. 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 , and amphibians . Flies are

318-405: A meal. For visual course control, flies' optic flow field is analyzed by a set of motion-sensitive neurons. 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. Other neurons are thought to be involved in analyzing the content of the visual scene itself, such as separating figures from

371-401: 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 often laid on

424-403: A pain-producing agent and the other in the distribution and penetration of the paralyzing components to vulnerable sites in the offender. This helps in the rapid immobilization of the animal or of the body parts receiving the venom. Spiders only bite, although some tarantulas have barbed bristles called urticating hairs . Certain caterpillars also have urticating hairs. Centipedes also possess

477-582: A pest of sunflowers and Rhagoletis mendax , a pest of blueberries. Another notorious agricultural pest is the Mediterranean fruit fly or Medfly, Ceratitis capitata , which is responsible for millions of dollars' worth in expenses by countries for control and eradication efforts, in addition to costs of damage to fruit crops. Similarly, the Queensland fruit fly ( Bactrocera tryoni ) is responsible for more than $ 28.5 million in damage to Australian fruit crops

530-404: A sting or stinger is a sharp organ, often connected with a venom gland and adapted to inflict a wound by piercing, as with the caudal sting of a scorpion . Stings are usually located at the rear of the animal. Animals with stings include bees, wasps (including hornets ), some ants like fire ants , and scorpions , as well as a single beetle species ( Onychocerus albitarsis ) that can deliver

583-438: A variety of hosts. The larvae of the shore flies (Ephydridae) and some Chironomidae survive in extreme environments including glaciers ( Diamesa sp., Chironomidae ), hot springs, geysers, saline pools, sulphur pools, septic tanks and even crude oil ( Helaeomyia petrolei ). Adult hoverflies (Syrphidae) are well known for their mimicry and the larvae adopt diverse lifestyles including being inquiline scavengers inside

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636-431: A venomous sting from its antennae, whose terminal segments have evolved to resemble a scorpion's tail. In all stinging Hymenoptera the sting is a modified ovipositor . Unlike most other stings, honey bee workers' stings are strongly barbed and lodge in the flesh of mammals upon use, tearing free from the honey bee's body, killing the bee within minutes. The sting has its own ganglion , and it continues to saw into

689-399: A year. This species lays eggs in a wide variety of unripe fruit hosts, causing them to rot prior to ripening. Some fruit flies are used as agents of biological control , thereby reducing the populations of pest species. Several species of the genus Urophora are used as control agents against rangeland-destroying noxious weeds such as starthistles and knapweeds , but their effectiveness

742-485: Is Spilomyia longicornis , which is a fly but mimics a vespid wasp. 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

795-404: Is complicated by its introduction of venom , although not all stings are venomous. Bites, which can introduce saliva as well as additional pathogens and diseases, are often confused with stings, and vice versa. Specific components of venom are believed to give rise to an allergic reaction , which in turn produces skin lesions that may vary from a small itching weal, or slightly elevated area of

848-587: Is divided into several subfamilies: The genera Oxyphora , Pseudorellia , and Stylia comprise 32 species, and are not included in any subfamily ( incertae sedis ). Fly Housefly (Muscidae) (top left) Haematopota pluvialis (Tabanidae) (top right) Ctenophora pectinicornis (Tipulidae) (mid left) Ochlerotatus notoscriptus (Culicidae) (mid right) Milesia crabroniformis (Syrphidae) (bottom left) Holcocephala fusca (Asilidae) (bottom right) Nematocera ( paraphyletic ) (inc Eudiptera ) Brachycera Flies are insects of

901-621: Is not barbed and can be used to sting repeatedly. The description of barbed or unbarbed is not precise: there are barbs on the stings of yellowjacket wasps and the Mexican honey wasp , but the barbs are so small that the wasp can sometimes withdraw its sting apparatus from victim's skin. The stings of some wasps, such as those of the Polistes versicolor , contain relatively large amounts of 5-hydroxytryptamine (5-HT) in its venoms. The 5-HT in these venoms has been found to play at least two roles: one as

954-419: Is of great interest to biologists. Some fruit flies have extensive mating rituals or territorial displays. Many are brightly colored and visually showy. Some fruit flies show Batesian mimicry , bearing the colors and markings of dangerous arthropods such as wasps or jumping spiders because it helps the fruit flies avoid predation, though the flies lack stingers . Adult tephritid fruit flies are often found on

1007-547: Is one of the major insect orders and of considerable ecological and human importance. Flies are major pollinators, second only to the bees and their Hymenopteran relatives. Flies may have been among the evolutionarily earliest pollinators responsible for early plant pollination . 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 , commensal with humans all over

1060-683: Is questionable. Urophora sirunaseva produces larvae that pupate within a woody gall within the flower and disrupt seed production. Chaetorellia acrolophi is an effective biocontrol agent against knapweeds Chaetorellia australis and Chaetorellia succinea , deposit eggs into the starthistle seedheads, where their larvae consume the seeds and flower ovaries. Since economically important tephritid fruit flies exist worldwide, vast networks of researchers, several international symposia, and intensive activities on various subjects extend from ecology to molecular biology ( Tephritid Workers Database ). Pest management techniques applied to tephritid include

1113-518: Is widely thought to be a member of Mecopterida , along with Lepidoptera (butterflies and moths), Trichoptera (caddisflies), Siphonaptera (fleas), Mecoptera (scorpionflies) and possibly Strepsiptera (twisted-wing flies). Diptera has been grouped with Siphonaptera and Mecoptera in the Antliophora, but this has not been confirmed by molecular studies. Diptera were traditionally broken down into two suborders, Nematocera and Brachycera , distinguished by

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1166-615: 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. The phylogenetic position of Diptera has been controversial. The monophyly of holometabolous insects has long been accepted, with the main orders being established as Lepidoptera, Coleoptera, Hymenoptera and Diptera, and it is the relationships between these groups which has caused difficulties. Diptera

1219-1060: The Mecopterida , alongside the Mecoptera , Siphonaptera , Lepidoptera and Trichoptera . The possession 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. The cladogram represents the current consensus view. Hymenoptera (sawflies, wasps, ants, bees) [REDACTED] Raphidioptera (snakeflies) [REDACTED] Megaloptera (alderflies and allies) [REDACTED] Neuroptera (Lacewings and allies) [REDACTED] Coleoptera (beetles) [REDACTED] Strepsiptera (twisted-wing parasites) [REDACTED] Trichoptera (caddisflies) [REDACTED] Lepidoptera (butterflies, moths) [REDACTED] Diptera [REDACTED] Mecoptera (scorpionflies) [REDACTED] Siphonaptera (fleas) [REDACTED] The first true dipterans known are from

1272-649: The Middle Triassic (around 240 million years ago), and they became widespread during the Middle and Late Triassic . Modern flowering plants did not appear until the Cretaceous (around 140 million years ago), so the original dipterans must have had a different source of nutrition other than nectar . Based on the attraction of many modern fly groups to shiny droplets, it has been suggested that they may have fed on honeydew produced by sap-sucking bugs which were abundant at

1325-605: The Strepsiptera . In contrast to the flies, the Strepsiptera bear their halteres on the mesothorax and their flight wings on the metathorax. 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 . At the tip of the limb is a pair of claws, and between these are cushion-like structures known as pulvilli which provide adhesion. Stinger An insect sting

1378-596: The larvae find their food upon emerging. The larvae develop in leaves , stems, flowers, seeds, fruits, and roots of the host plant, depending on the species. Some species are gall-forming . One exception to the phytophagous lifestyle is Euphranta toxoneura (Loew) whose larvae develop in galls formed by sawflies . The adults sometimes have a very short lifespan. Some live for less than a week. Some species are monophagous (feeding on only one plant species) others are polyphagous (feeding on several, usually related plant species). The behavioral ecology of tephritid fruit flies

1431-566: 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 mechanosensory organs known as halteres , which act as high-speed sensors of rotational movement and allow dipterans to perform advanced aerobatics. Diptera is a large order containing more than 150,000 species including horse-flies , crane flies , hoverflies , mosquitoes and others. Flies have

1484-548: 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 hindmost segments modified for reproduction. Some Dipterans are mimics and can only be distinguished from their models by very careful inspection. An example of this

1537-795: The Brachycera includes broader, more robust flies with short antennae. Many nematoceran larvae are aquatic. 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. While most species have restricted distributions, a few like the housefly ( Musca domestica ) are cosmopolitan. Gauromydas heros ( Asiloidea ), with

1590-1392: The Nematocera. The construction of a phylogenetic tree has been the subject of ongoing research. The following cladogram is based on the FLYTREE project. Ptychopteromorpha (phantom and primitive crane-flies) [REDACTED] Culicomorpha (mosquitoes, blackflies and midges) [REDACTED] Blephariceromorpha (net-winged midges, etc) [REDACTED] Bibionomorpha (gnats) [REDACTED] Psychodomorpha (drain flies, sand flies, etc) [REDACTED] Tipulomorpha (crane flies) [REDACTED] Stratiomyomorpha (soldier flies, etc) [REDACTED] Xylophagomorpha (stink flies, etc) [REDACTED] Tabanomorpha (horse flies, snipe flies, etc) [REDACTED] Nemestrinoidea [REDACTED] Asiloidea (robber flies, bee flies, etc) [REDACTED] Empidoidea (dance flies, etc) [REDACTED] Aschiza (in part) Phoroidea (flat-footed flies, etc) [REDACTED] Syrphoidea (hoverflies) [REDACTED] Hippoboscoidea (louse flies, etc) [REDACTED] Muscoidea (house flies, dung flies, etc) [REDACTED] Oestroidea (blow flies, flesh flies, etc) [REDACTED] Acalyptratae (marsh flies, etc) [REDACTED] Flies are often abundant and are found in almost all terrestrial habitats in

1643-745: The Tephritidae. Description , recategorization , and genetic analyses are constantly changing the taxonomy of this family. To distinguish them from the Drosophilidae, the Tephritidae are sometimes called peacock flies , in reference to their elaborate and colorful markings. The name comes from the Greek τεφρος, tephros , meaning "ash grey". They are found in all the biogeographic realms . For terms see Morphology of Diptera and Tephritidae glossary Tephritids are small to medium-sized (2.5–10 mm) flies that are often colourful, and usually with pictured wings,

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1696-519: The antennae and the maxillary palps bear the main olfactory receptors, while the gustatory receptors are in the labium, pharynx, feet, wing margins and female genitalia, enabling flies to taste their food by walking on it. The taste receptors in females at the tip of the abdomen receive information on the suitability of a site for ovipositing. Flies that feed on blood have special sensory structures that can detect infrared emissions, and use them to home in on their hosts, Many blood-sucking flies can detect

1749-404: The differences in antennae. The Nematocera are identified 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. Subsequent studies have identified the Nematocera as being non-monophyletic with modern phylogenies placing the Brachycera within grades of groups formerly placed in

1802-431: 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. 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

1855-749: The flies, principally in the family Cecidomyiidae (gall midges). Many flies (most importantly in the family Agromyzidae) lay their eggs in the mesophyll tissue of leaves with larvae feeding between the surfaces forming blisters and mines. Some families are mycophagous or fungus feeding. These include the cave dwelling Mycetophilidae (fungus gnats) whose larvae are the only diptera with bioluminescence. The Sciaridae are also fungus feeders. Some plants are pollinated by fungus feeding flies that visit fungus infected male flowers. The larvae of Megaselia scalaris (Phoridae) are almost omnivorous and consume such substances as paint and shoe polish. The Exorista mella (Walker) fly are considered generalists and parasitoids of

1908-405: The frontal bristles are inserted on a raised tubercle. Interfrontal setulae are usually absent or represented by one or two tiny setulae near the lunula. True vibrissae are absent, but several genera have strong bristles near the vibrissal angle. The wings usually have yellow, brown, or black markings or are dark-coloured with lighter markings. In a few species, the wings are clear. The costa has both

1961-527: The ground using motion parallax. 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. The ocelli are concerned in the detection of changes in light intensity, enabling the fly to react swiftly to the approach of an object. Like other insects, flies have chemoreceptors that detect smell and taste, and mechanoreceptors that respond to touch. The third segments of

2014-532: The host plant and feeding on pollen, nectar, rotting plant debris, or honeydew. Natural enemies include parasitoid wasps of the genera Diapriidae and Braconidae . Tephritid fruit flies are of major economic importance in agriculture . Some have negative effects, some positive. Various species of fruit flies cause damage to fruit and other plant crops. The genus Bactrocera is of worldwide notoriety for its destructive impact on agriculture. The olive fruit fly ( B. oleae ), for example, feeds on only one plant:

2067-431: The larval food-source and the larvae, which lack true limbs, develop in a protected environment, often inside their food source. Other species are ovoviviparous , opportunistically depositing hatched or hatching larvae instead of eggs on carrion , dung, decaying material, or open wounds of mammals. The pupa is a tough capsule from which the adult emerges when ready to do so; flies mostly have short lives as adults. Diptera

2120-675: The mandibles have several transverse oral ridges or short laminae directed posteriorly. The anterior spiracles (prothoracic spiracles) end bluntly and are not elongated. Each has at least three openings or up to 50 arranged transversely in one to three groups or irregularly. Each posterior spiracle (anal spiracle) lacks a clearly defined peritreme and each has three spiracular openings (in mature larvae). These are usually more or less horizontal, parallel and usually bear branched spiracular hairs in four tufts. The larvae of almost all Tephritidae are phytophagous . Females deposit eggs in living, healthy plant tissue using their telescopic ovipositors . Here,

2173-419: The nests of social insects. Some brachycerans are agricultural pests, some bite animals and humans and suck their blood, and some transmit diseases. 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,

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2226-406: The neural ganglia , and concentration of nerve tissue in the thorax, a feature that is most extreme in the highly derived Muscomorpha infraorder. Some flies such as the ectoparasitic Nycteribiidae and Streblidae are exceptional in having lost their wings and become flightless. The only other order of insects bearing a single pair of true, functional wings, in addition to any form of halteres, are

2279-406: The other secreting an alkaline neurotoxin ; acting independently, each toxin is rather mild, but when they combine through the sting, the combination has strong irritating properties. In a small number of cases, the second occasion of a bee or wasp sting causes a severe allergic reaction known as anaphylaxis . While the overwhelming majority of insects withdraw their stingers from their victims,

2332-405: The raised concentration of carbon dioxide that occurs near large animals. Some tachinid flies (Ormiinae) which are parasitoids of bush crickets , have sound receptors to help them locate their singing hosts. 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

2385-562: The second largest group of pollinators after the Hymenoptera (bees, wasps and relatives). In wet and colder environments flies are significantly more important as pollinators. Compared to bees, they need less food as they do not need to provision their young. Many flowers that bear low nectar and those that have evolved trap pollination depend on flies. It is thought that some of the earliest pollinators of plants may have been flies. The greatest diversity of gall forming insects are found among

2438-430: The skin, to large areas of inflamed skin covered by vesicles and crusted lesions . Stinging insects produce a painful swelling of the skin, the severity of the lesion varying according to the location of the sting, the identity of the insect and the sensitivity of the subject. Many species of bees and wasps have two poison glands, one gland secreting a toxin in which formic acid is one recognized constituent, and

2491-432: The subcostal vein curving forward at a right angle. The head is hemispherical and usually short. The face is vertical or retreating and the frons is broad. Ocelli and cellar bristles are present. The postvertical bristles are parallel to divergent. Two to eight pairs of frontal bristles are seen (at least one but usually several lower pairs curving inwards and at least one of the upper pairs curving backwards). In some species,

2544-548: The target's flesh and release venom for several minutes. This trait is of obvious disadvantage to the individual but protects the hive from attacks by large animals; aside from the effects of the venom, the remnant also marks the stung animal with honey bee alarm pheromone. The barbs of a honey bee's attack are only suicidal if the skin is elastic, as is characteristic of vertebrates such as birds and mammals; honey bees can sting other insects repeatedly without dying. The sting of nearly all other bees and other sting-bearing organisms

2597-588: The time, and dipteran mouthparts are well-adapted to softening and lapping up the crusted residues. The basal clades in the Diptera include the Deuterophlebiidae and the enigmatic Nymphomyiidae . Three episodes of evolutionary radiation are thought to have occurred based on the fossil record. Many new species of lower Diptera developed in the Triassic , about 220 million years ago. Many lower Brachycera appeared in

2650-420: The use of cover sprays with conventional pesticides , however, due to deleterious impact of these pesticides, new, less impactful and more targeted pest control techniques have been used, such as toxic food baits , male annihilation technique using specific male attractant parapheromones in toxic baits or mass trapping , or even sterile insect technique as part of integrated pest management . Tephritidae

2703-504: The wild or commercially cultivated olive , Olea europaea . It has the capacity to ruin 100% of an olive crop by damaging the fruit. Bactrocera dorsalis is another highly invasive pest species that damages tropical fruit, vegetable, and nut crops. Euleia heraclei is a pest of celery and parsnips. The genus Anastrepha includes several important pests, notably A. grandis , A. ludens (Mexican fruit fly), A. obliqua , and A. suspensa . Other pests are Strauzia longipennis ,

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2756-473: The world apart from Antarctica. They include many familiar insects such as house flies, blow flies, mosquitoes, gnats, black flies, midges and fruit flies. More than 150,000 have been formally described and the actual species diversity is much greater, with the flies from many parts of the world yet to be studied intensively. The suborder Nematocera include generally small, slender insects with long antennae such as mosquitoes, gnats, midges and crane-flies, while

2809-607: The world, spread foodborne illnesses . Flies can be annoyances especially in some parts of the world where they can occur in large numbers, buzzing and settling on the skin or eyes to bite or seek fluids. Larger flies such as tsetse flies and screwworms 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 , as food for carnivorous animals, and in medicine in debridement , to clean wounds . Dipterans are holometabolans , insects that undergo radical metamorphosis. They belong to

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