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54-710: Muscidae are a family of flies found in the superfamily Muscoidea . Muscidae, some of which are commonly known as house flies or stable flies due to their synanthropy , are worldwide in distribution and contain almost 4,000 described species in over 100 genera. Most species are not synanthropic. Adults can be predatory(like humans), hematophagous , saprophagous , or feed on a number of types of plant and animal exudates . They can be attracted to various substances including sugar, sweat, tears [1] and blood. Larvae occur in various habitats including decaying vegetation, dry and wet soil, nests of insects and birds, fresh water, and carrion. The housefly, Musca domestica ,

108-401: A larval stage undergo metamorphosis, and during metamorphosis the organism loses larval characteristics. The word metamorphosis derives from Ancient Greek μεταμόρφωσις , "transformation, transforming", from μετα- ( meta- ), "after" and μορφή ( morphe ), "form". In insects, growth and metamorphosis are controlled by hormones synthesized by endocrine glands near

162-500: A yolk sac ), then to motile larvae (often known as fingerlings due to them roughly reaching the length of a human finger ) that have to forage for themselves after the yolk sac resorbs, and then to the juvenile stage where the fish progressively start to resemble adult morphology and behaviors until finally reaching sexual maturity . In typical amphibian development, eggs are laid in water and larvae are adapted to an aquatic lifestyle. Frogs , toads , and newts all hatch from

216-399: A butterfly 2 – The pupa is now spewing the thread to form chrysalis 3 – The chrysalis is fully formed 4 – Adult butterfly coming out of the chrysalis In cephalochordata , metamorphosis is iodothyronine -induced and it could be an ancestral feature of all chordates . Some fish, both bony fish (Osteichthyes) and jawless fish (Agnatha) , undergo metamorphosis. Fish metamorphosis

270-648: A complete metamorphosis, including a pupal or resting stage between the larval and adult forms. A number of hypotheses have been proposed to explain the evolution of holometaboly from hemimetaboly, mostly centering on whether or not the intermediate stages of hemimetabolous forms are homologous in origin to the pupal stage of holometabolous forms. According to a 2009 study, temperature plays an important role in insect development as individual species are found to have specific thermal windows that allow them to progress through their developmental stages. These windows are not significantly affected by ecological traits, rather,

324-413: A considerable role during physiological processes of multicellular organisms, particularly during embryogenesis , and metamorphosis. Additional research in 2019 found that both autophagy and apoptosis , the two ways programmed cell death occur, are processes undergone during insect metamorphosis. Below is the sequence of steps in the metamorphosis of the butterfly (illustrated): 1 – The larva of

378-456: 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

432-454: A long stage of growth followed by a more gradual metamorphosis to the migrating phase. In the pre-adult freshwater stage, the eel also has phenotypic plasticity because fish-eating eels develop very wide mandibles, making the head look blunt. Leptocephali are common, occurring in all Elopomorpha ( tarpon - and eel -like fish). Most other bony fish undergo metamorphosis initially from egg to immotile larvae known as sac fry ( fry with

486-405: 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

540-401: 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

594-420: A relatively long, spiral‐shaped gut to digest that diet. Recent studies suggest tadpoles do not have a balanced homeostatic feedback control system until the beginning stages of metamorphosis. At this point, their long gut shortens and begins favoring the diet of insects. Rapid changes in the body can then be observed as the lifestyle of the frog changes completely. The spiral‐shaped mouth with horny tooth ridges

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648-438: 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

702-485: 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

756-555: Is iodothyronine-induced and an ancestral feature of all chordates . All three categories of metamorphosis can be found in the diversity of insects, including no metamorphosis ("ametaboly"), incomplete or partial metamorphosis ("hemimetaboly"), and complete metamorphosis ("holometaboly"). While ametabolous insects show very little difference between larval and adult forms (also known as " direct development "), both hemimetabolous and holometabolous insects have significant morphological and behavioral differences between larval and adult forms,

810-547: 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

864-421: Is outside the parental body, development is subject to many adaptations due to specific ecological circumstances. For this reason tadpoles can have horny ridges for teeth, whiskers, and fins. They also make use of the lateral line organ. After metamorphosis, these organs become redundant and will be resorbed by controlled cell death, called apoptosis . The amount of adaptation to specific ecological circumstances

918-769: Is present. The R5 cell is either parallel-sided or narrowed distally. Vein 2A is short and does not reach the wing margin. The Fanniidae , which used to be a subfamily (Fanniinae) of the Muscidae, share these characters, but may be separated from them by the absence of the identifying characteristics for the family Fanniidae. Larvae mainly develop in decaying plant material or manure. Adults of many species are passive vectors of pathogens for diseases such as typhoid fever , dysentery , anthrax , and African sleeping sickness . Larvae of some Atherigona species are important pests in cultivation of cereals, like rice and maize. Seven species in six described genera have been recorded from

972-549: Is reabsorbed, due to the higher thyroxin concentrations required for tail resorption. Salamander development is highly diverse; some species go through a dramatic reorganization when transitioning from aquatic larvae to terrestrial adults, while others, such as the axolotl , display pedomorphosis and never develop into terrestrial adults. Within the genus Ambystoma , species have evolved to be pedomorphic several times, and pedomorphosis and complete development can both occur in some species. In newts, metamorphosis occurs due to

1026-402: Is remarkable, with many discoveries still being made. With frogs and toads, the external gills of the newly hatched tadpole are covered with a gill sac after a few days, and lungs are quickly formed. Front legs are formed under the gill sac, and hindlegs are visible a few days later. Following that there is usually a longer stage during which the tadpole lives off a vegetarian diet. Tadpoles use

1080-401: Is resorbed together with the spiral gut. The animal develops a big jaw, and its gills disappear along with its gill sac. Eyes and legs grow quickly, a tongue is formed, and all this is accompanied by associated changes in the neural networks (development of stereoscopic vision, loss of the lateral line system, etc.) All this can happen in about a day. It is not until a few days later that the tail

1134-476: Is the best known and most important species. Some, from the genera Hydrotaea and Muscina , are involved in forensic case studies. The antennae are three-segmented and aristate; vein Rs is two-branched, a frontal suture is present, and the calypters are well developed. The arista is often plumose for the entire length. The hypopleuron is usually without bristles; generally, more than one sternopleural bristle

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1188-403: Is typically under strong control by the thyroid hormone. Examples among the non-bony fish include the lamprey . Among the bony fish, mechanisms are varied. The salmon is diadromous , meaning that it changes from a freshwater to a saltwater lifestyle. Many species of flatfish begin their life bilaterally symmetrical , with an eye on either side of the body; but one eye moves to join

1242-518: 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

1296-615: 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

1350-1060: 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

1404-649: 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

1458-618: 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. Metamorphosis Generally organisms with

1512-566: 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

1566-548: 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

1620-795: 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

1674-1392: 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

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1728-519: 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

1782-411: The change in habitat, not a change in diet, because newt larvae already feed as predators and continue doing so as adults. Newts' gills are never covered by a gill sac and will be resorbed only just before the animal leaves the water. Adults can move faster on land than in water. Newts often have an aquatic phase in spring and summer, and a land phase in winter. For adaptation to a water phase, prolactin

1836-462: The development of adult characteristics during ecdysis . In holometabolous insects, molts between larval instars have a high level of juvenile hormone, the moult to the pupal stage has a low level of juvenile hormone, and the final, or imaginal , molt has no juvenile hormone present at all. Experiments on firebugs have shown how juvenile hormone can affect the number of nymph instar stages in hemimetabolous insects. In chordates, metamorphosis

1890-404: 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

1944-431: 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

1998-538: The eggs as larvae with external gills but it will take some time for the amphibians to interact outside with pulmonary respiration. Afterwards, newt larvae start a predatory lifestyle, while tadpoles mostly scrape food off surfaces with their horny tooth ridges. Metamorphosis in amphibians is regulated by thyroxin concentration in the blood, which stimulates metamorphosis, and prolactin , which counteracts its effect. Specific events are dependent on threshold values for different tissues. Because most embryonic development

2052-650: The evolution of metamorphosis in insects is thought to have fuelled their dramatic radiation (1,2). Some early ametabolous "true insects" are still present today, such as bristletails and silverfish . Hemimetabolous insects include cockroaches , grasshoppers , dragonflies , and true bugs . Phylogenetically, all insects in the Pterygota undergo a marked change in form, texture and physical appearance from immature stage to adult. These insects either have hemimetabolous development, and undergo an incomplete or partial metamorphosis, or holometabolous development, which undergo

2106-749: 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

2160-805: The fossil record. Lambrecht (1980: 369) estimated that the family Muscidae originated as long ago as the Permian , although no fossil record exists for the family any older than the Eocene . List of genera according to the Catalogue of Life : 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

2214-411: The front of the body ( anterior ). Neurosecretory cells in an insect's brain secrete a hormone, the prothoracicotropic hormone (PTTH) that activates prothoracic glands, which secrete a second hormone, usually ecdysone (an ecdysteroid ), that induces ecdysis (shedding of the exoskeleton). PTTH also stimulates the corpora allata , a retrocerebral organ, to produce juvenile hormone , which prevents

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2268-527: 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

2322-431: 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

2376-566: The most significant being the inclusion, in holometabolous organisms, of a pupal or resting stage between the larval and adult forms. In hemimetabolous insects , immature stages are called nymphs . Development proceeds in repeated stages of growth and ecdysis (moulting); these stages are called instars . The juvenile forms closely resemble adults, but are smaller and lack adult features such as wings and genitalia. The size and morphological differences between nymphs in different instars are small, often just differences in body proportions and

2430-419: 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,

2484-406: 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

2538-488: The number of segments; in later instars, external wing buds form. The period from one molt to the next is called a stadium. In holometabolous insects, immature stages are called larvae and differ markedly from adults. Insects which undergo holometabolism pass through a larval stage, then enter an inactive state called pupa (called a "chrysalis " in butterfly species), and finally emerge as adults. The earliest insect forms showed direct development ( ametabolism ), and

2592-464: The other side of the fish – which becomes the upper side – in the adult form. The European eel has a number of metamorphoses, from the larval stage to the leptocephalus stage, then a quick metamorphosis to glass eel at the edge of the continental shelf (eight days for the Japanese eel ), two months at the border of fresh and salt water where the glass eel undergoes a quick metamorphosis into elver, then

2646-405: 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

2700-562: 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

2754-588: 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

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2808-529: The windows are phylogenetically adapted to the ecological circumstances insects are living in. According to research from 2008, adult Manduca sexta is able to retain behavior learned as a caterpillar . Another caterpillar, the ornate moth caterpillar , is able to carry toxins that it acquires from its diet through metamorphosis and into adulthood, where the toxins still serve for protection against predators. Many observations published in 2002, and supported in 2013 indicate that programmed cell death plays

2862-473: 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

2916-607: 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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