Fly Temporal range: Middle Triassic - Recent 245–0 Ma PreЄ Є O S D C P T J K Pg N
A poster with sixteen different species of flies
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Superorder:	Panorpida
(unranked):	Antliophora
Order:	Diptera Linnaeus, 1758
Suborders
Nematocera (includes Eudiptera) Brachycera Data related to Diptera at Wikispecies

True flies are insects of the order Diptera (from the Greek di = two, and ptera = wings). They possess a pair of wings on the mesothorax and a pair of halteres, derived from the hind wings, on the metathorax. Apart from secondarily flightless insects (including some flies), the only other order of insects with any form of halteres are the Strepsiptera, and theirs are on the mesothorax, with the flight wings on the metathorax.

Order Diptera[link]

The presence of a single pair of patent, metathoracic flight wings distinguishes true flies from other insects with "fly" in their names, such as mayflies, dragonflies, damselflies, stoneflies, whiteflies, fireflies, alderflies, dobsonflies, snakeflies, sawflies, caddisflies, butterflies or scorpionflies. Some true flies have become secondarily wingless, especially in the superfamily Hippoboscoidea, or among those that are inquilines in social insect colonies.

Some authors draw a distinction in writing the common names of insects. True flies are written as two words, such as crane fly, robber fly, bee fly, moth fly, and fruit fly. In contrast, common names of nondipteran 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, and, especially in older books, one commonly might see the likes of "saw fly" and "caddis fly". In any case, non-entomologists cannot in general be expected to tell dipterans, "true flies", from other insects.

The Diptera comprise a large order, containing an estimated 240,000 species of mosquitoes, gnats, midges and others, although under half of these (about 120,000 species) have been described.^[2] It is one of the major insect orders both in terms of ecological and human (medical and economic) importance. The Diptera, in particular the mosquitoes (Culicidae), are of great importance as disease transmitters, acting as vectors for malaria, dengue, West Nile virus, yellow fever, encephalitis and other infectious diseases.

Anatomy and biology[link]

See Morphology of Diptera and Biology of Diptera

Portrait of a flesh fly (Sarcophagidae)

An image of a house fly eye surface by using scanning electron microscope at 450× magnification

Flies are adapted for aerial movement and typically have short and streamlined bodies. The first tagma of the fly, the head, consists of ocelli, antennae, compound eyes, 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. The third segment of the thorax bears the halteres, which help to balance the insect during flight. 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.^[3]

A scan of a house fly taken at 40 magnifications under a scanning electron microscope

Flies have a mobile head with eyes and in most cases have large compound eyes on the sides of the head, with three small ocelli on the top. The antennae take a variety of forms, but are often short, which reduces drag while flying.

Because no species of fly has teeth or any other organ or limb that allows them to eat solid foods, flies consume only liquid food or finely granular foods, such as pollen, and their mouthparts and digestive tracts show various modifications for such diets. Female Tabanidae use knife-like mandibles and maxillae to make a cross-shaped incision in the hosts' skin and then lap up the blood. The gut includes large diverticulae, allowing the insect to store small quantities of liquid after a meal.^[3]

For detailed anatomy explore [1] which uses examples from the 4 major fly groups (Lower Diptera, Lower Brachycera, Acalyptrate, Calyptrate) representing different anatomical expressions.

Reproduction and development[link]

Mating anthomyiid flies

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 located below the genitals, or, in the case of 360° torsion, to the sperm duct being wrapped around the gut, despite 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 round 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 great populations due to their ability to mate effectively and in a short period of time during the mating season.^[3]

The female lays her eggs as close to the food source as possible, and development is rapid, allowing the larvae to consume as much food as possible in a short period of time before transforming into adults. The eggs hatch immediately after being laid, or the flies are ovoviviparous, with the larvae hatching inside the mother.^[3]

Mosquito larva. A typical Nematoceran. Note the head capsule and clearly defined three segments of the thorax

Syrphid larva. Maggot-like anatomy, typical of a Brachycerid fly, lacking both a conspicuously demarcated thorax and head capsule

Larval flies have no true legs. Some Dipteran larvae, such as species of Simuliidae, Tabanidae, and Vermileonidae, have prolegs adapted to such functions as holding onto a substrate in flowing water, holding onto host tissues, or holding prey.^[4] Roughly speaking, there is some anatomical distinction between the larvae of the Nematocera and the Brachycera (see Classification section, below); especially in the Brachycera, there is little demarcation between the thorax and abdomen, though the demarcation may be very visible in many Nematocera, such as mosquitoes (see image, both here and in the mosquitoes article); in the Brachycera, the head of the larva is not clearly distinguishable from the rest of the body, and there are few, if any, sclerites. Informally, such Brachyceran larvae are called maggots,^[5] but the term is nontechnical 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.^[3] Nematoceran larvae generally have visible eyes and antennae, though usually small and of limited function.

The pupae take various forms, and in some cases develop inside a silk cocoon. After emerging from the pupa, the adult fly rarely lives more than a few days, and serves mainly to reproduce and to disperse in search of new food sources.

Classification[link]

Cleaning

A fly

The Nematocera are recognized by their elongated bodies and feathery antennae as represented by mosquitoes and crane flies. The Brachycera have a more roundly proportioned body and much shorter antennae. In 1964, B.B. Rohdendorf proposed a classification in which the Nematocera is split into two suborders, the Archidiptera and the Eudiptera.^[6]

1. Suborder Nematocera (77 families, 35 of them extinct) – long antennae, pronotum distinct from mesonotum, in Nematocera, larvae are either eucephalic or hemicephalic and often aquatic.
2. Suborder Brachycera (141 families, 8 of them extinct) – short antennae, the pupa is inside a puparium formed from the last larval skin, they are generally robust flies with larvae having reduced mouthparts.
   1. Infraorders Tabanomorpha and Asilomorpha – these comprise the majority of what was the Orthorrhapha under older classification schemes. The antennae are short, but differ in structure from those of the Muscomorpha.
   2. Infraorder Muscomorpha – (largely the Cyclorrhapha of older schemes). Muscomorpha have three-segmented, aristate (with a bristle) antennae and larvae with three acephalic instars (maggots).

Most of the Muscomorpha are further subdivided into the Acalyptratae and Calyptratae based on whether or not they have a calypter (a wing flap that extends over the halteres).

Beyond that, considerable revision in the taxonomy of the flies has taken place since the introduction of modern cladistic techniques, and much remains uncertain. The secondary ranks between the suborders and the families are out of practical or historical considerations than out of strict respect for phylogenetic classifications (modern cladists spurn the use of Linnaean rank names). All classifications in use now, including this article, contain some paraphyletic groupings; this is emphasized where the numerous alternative systems are most greatly at odds. See list of families of Diptera.

Dipterans belong to the taxon Mecopterida, that also contains Mecoptera, Siphonaptera, Lepidoptera (butterflies and moths) and Trichoptera. Inside it, they are classified closely together with Mecoptera and Siphonaptera in the superorder Antliophora.^[7]

Evolution[link]

Diptera derive from Mecoptera or a strictly related group. The first true dipterans known are from the Middle Triassic, and they became widespread during the Middle and Late Triassic .^[8]

The basal clades in the Diptera are the Deuterophlebiidae and Nymphomyiidae.^[9] The Bibionomorpha are a sister clade to Brachycera. The branching order of the remaining clades of the lower Diptera - infraorders Culicomorpha, Psychodomorpha and Tipulomorpha - has yet to be resolved.

Within the Brachycera, several progressively nested groups exist: Eremoneura (three larval instars), Cyclorrhapha (pupation occurs within a puparium), Schizophora (flies that escape from their puparium using the ptilinal sac, an evertable frontal pouch) and Calyptratae (larger flies with wings that have the calypter, an enlarged basal lobe).

The Schizophora include most of the family-level diversity in Diptera (∼85 families) and more than 50,000 species. The Calyptratae form a monophyletic superfamily. Other monophyletic superfamilies include the Ephydroidea, Lauxanioidea, Nerioidea, Sciomyzoidea and Tephritoidea. The relationships between the remaining families have yet to be clarified.

Sister groups to the Drosophilidae consist of two families, Braulidae and Cryptochetidae.

There were three episodes of rapid evolution in the lower Diptera (~220 million years ago), lower Brachycera (180 million years ago) and Schizophora (65 million years ago).^[9]

Maggots[link]

Maggots being used to treat a wound

Maggots found on corpses are useful to forensic scientists; specifically forensic entomology. By their stage of development, these maggots indicate the time elapsed since death, as well as the place of death.

Maggot species can be identified using their DNA. The housefly maggot measures 10–20 mm (⅜–¾ in) in size. At the height of the summer season, a generation of flies (egg to adult) may be produced in 12 to 14 days. Other insect families, such as Histeridae, feed on maggots. Thus the lack of maggots would increase the estimated time of death.

Maggots are bred commercially, as a popular bait in angling, and as a food for carnivorous pets such as reptiles or birds.

Maggots are used in medicine to clean out necrotic wounds and in food production, particularly of cheeses such as casu marzu designed to rot as part of their aging process.

Flies are reared in large numbers in Japan to serve as pollinators of sunflowers in greenhouses, especially the maggots.

Flies in culture[link]

Flies have been depicted in mythology and literature. In the Biblical fourth plague of Egypt, flies represent death and decay. Myiagros was a god in Greek mythology who chased away flies during the sacrifices to Zeus and Athena, and Zeus sent a fly to bite Pegasus, causing Bellerophon to fall back to Earth when he attempted to ride the winged steed to Mount Olympus. In the traditional Navajo religion, Big Fly is an important spirit being.

In the 15th-century trompe l'oeil painting Portrait of a Carthusian (1446) by Petrus Christus, a fly sits on a fake frame.^[10]

Emily Dickinson's 1955 poem "I Heard a Fly Buzz When I Died" also makes reference to flies in the context of death. In fact, flies such as the genus Hydrotaea are used in forensic cases to determine time of death for corpses. In William Golding's 1954 novel Lord of the Flies, the fly is a symbol of the children involved.

In the 1958 science fiction film The Fly (and its 1986 remake), a scientist accidentally exchanges parts of his body with those of a fly.

Musical works that mention flies: Yoko Ono's 1971 album Fly, U2's 1991 song "The Fly", Wire's 1978 song "I Am The Fly", Dave Matthews's 2007 song "The Fly" and Béla Bartók's 1920s piano work "From the Diary of a Fly".

Damien Hirst's provocative 1990 artwork, titled A Thousand Years, featured a severed cow's head contained in a box with thousands of flies and a bug zapper, creating an entire life cycle within a glass box. In 2001 Dr. Garnet Hertz implanted a complete web server into a dead fly.^[11]

The ability of flies to cling to surfaces has also inspired the title of "Human Fly" for real and fictional stunt performers whose feats involve climbing buildings.

See also[link]

• Female sperm storage

Notes[link]

References[link]

Biology[link]

• Harold Oldroyd The Natural History of Flies. New York: W. W. Norton. 1965.
• Eugène Séguy 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. Text figs. Part of Encyclopedie Entomologique, Serie B II: Diptera. 1924–1953.
• Eugène Seguy. La Biologie des Dipteres 1950. pp. 609. 7 col + 3 b/w plates, 225 text figs.

Classification[link]

• Brown, B.V., Borkent, A., Cumming, J.M., Wood, D.M., Woodley, N.E., and Zumbado, M. (Editors) 2009 Manual of Central American Diptera. Volume 1 NRC Research Press, Ottawa ISBN 978-0-660-19833-0
• 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 Univ. Press, Melbourne.
• Griffiths, G.C.D. The phylogenetic classification of Diptera Cyclorrhapha, withspecial reference to the structure of the male postabdomen. Ser. Ent. 8, 340 pp. [Dr. W. Junk, N. V., The Hague] (1972).
• Willi Hennig Die Larvenformen der Dipteren. 3. Teil. Akad.-Verlag, Berlin. 185 pp., 3 pls. 1948
• Willi Hennig (1954) Flugelgeader und System der Dipteren unter Berucksichtigung der aus dem Mesozoikum beschriebenen Fossilien. Beitr. Ent. 4: 245-388 (1954).
• F. Christian Thompson. "Sources for the Biosystematic Database of World Diptera (Flies)" (PDF). United States Department of Agriculture, Systematic Entomology Laboratory. http://www.sel.barc.usda.gov/diptera/names/BDWDsour.pdf. 
• Willi Hennig: Diptera (Zweifluger). Handb. Zool. Berl. 4 (2 ) (31):1–337. General introduction with key to World Families. In German.

Evolution[link]

• 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 Publ., Dordrecht, Boston, London, pp. 227–240.

External links[link]

Wikiquote has a collection of quotations related to: Flies

Wikimedia Commons has media related to: Diptera

Wikispecies has information related to: Diptera

Wikisource has the text of the 1911 Encyclopædia Britannica article Fly.

General

• The Diptera Site
• The Dipterists Forum - The Society for the study of flies
• The Bishop Museum Catalog of Fossil Diptera
• The Tree of Life Project
• Diptera at the Open Directory Project
• Manual of Afrotropical Diptera

Identification help

• The Diptera.info Portal Expert Diptera site and Diptera information portal.
• BugGuide

Anatomy

• Fly anatomy
• Drawing Wing venation

Describers

• Authors of fly names

v t e Orders of insects Extant Archaeognatha Blattodea (Cockroaches) Coleoptera (Beetles) Dermaptera (Earwigs) Diptera (Flies) Embioptera (Webspinners) Ephemeroptera (Mayflies) Hemiptera Hymenoptera Isoptera (Termites) Lepidoptera Mantodea (Mantises) Mecoptera Megaloptera Neuroptera Notoptera Odonata Orthoptera Phasmatodea (Sticks) Phthiraptera (Lice) Plecoptera Psocoptera Raphidioptera (Snakeflies) Siphonaptera (Fleas) Strepsiptera Thysanoptera (Thrips) Trichoptera (Caddisflies) Zoraptera Zygentoma (Thysanura) Extinct Coxoplectoptera Diaphanopterodea Glosselytrodea Meganisoptera Megasecoptera Palaeodictyoptera Wikispecies

Jools Holland
Holland at the British Academy Television Awards 2009
Background information
Birth name	Julian Miles Holland
Born	(1958-01-24) 24 January 1958 (age 54) Blackheath, London, England
Genres	Boogie-woogie, jazz, blues, R&B
Occupations	Musician, composer, television presenter, bandleader
Instruments	Piano, keyboard, guitar
Years active	1974–present
Associated acts	Squeeze Rhythm & Blues Orchestra
Website	Official site

Julian Miles "Jools" Holland OBE, DL (born 24 January 1958) is an English pianist, bandleader, singer, composer, and television presenter. He was a founder of the band Squeeze (1974-1980 & 1985-1990) and his work has involved him with many artists including Sting, Eric Clapton, George Harrison, David Gilmour, Magazine and Bono.

Holland is a published author and appears on television shows besides his own and contributes to radio shows. In 2004, he collaborated with Tom Jones on an album of traditional R&B music. He currently hosts Later... with Jools Holland, a music-based show aired on BBC2, on which his annual show Hootenanny, is based.^[1]

Life and career[link]

His great grandfather came from Ireland.^[2]

Holland played as a session musician before finding fame, and his first studio session was with Wayne County & the Electric Chairs in 1976 on their track "F*ck Off."^[3]

Holland was a founding member of the British pop band Squeeze, formed in March 1974, in which he played keyboards until 1981 and helped the band to achieve millions of record sales, before pursuing his solo career.^[3]

Holland began issuing solo records in 1978, his first EP being Boogie Woogie '78. He continued his solo career through the early 1980s, releasing an album and several singles between 1981 and 1984. He branched out into TV, co-presenting the Newcastle-based TV music show The Tube with Paula Yates. Holland achieved notoriety by inadvertently using the phrase "groovy fuckers" in a live, early evening TV trailer for the show, causing it to be suspended for six weeks.^[4] He referred to this in his sitcom "The Groovy Fellers" with Rowland Rivron.

Holland at the Tsunami Relief concert in Cardiff's Millennium Stadium, 22 January 2005

In 1983 Holland played an extended piano solo on The The's re-recording of "Uncertain Smile" for the album Soul Mining. In 1985, Squeeze (which had continued in Holland's absence through to 1982) unexpectedly regrouped including Jools Holland as their keyboard player. Holland remained in the band until 1990, at which point, he again departed Squeeze on amicable terms to resume his solo career as a musician and a TV host.

In 1987, Holland formed The Jools Holland Big Band which consisted of himself and Gilson Lavis from Squeeze. This gradually became his 18-piece Rhythm & Blues Orchestra.^[3]

Between 1988 and 1990 he performed and co-hosted along with David Sanborn during the two seasons of the music performance program Sunday Night on NBC late-night television.^[5] Since 1992 he has presented the music program Later... with Jools Holland, plus an annual New Year's Eve Hootenanny.

In 1996, Holland signed a recording contract with Warner Bros. Records,^[3] and his records are now marketed through Rhino Records.

Holland has a touring band, The Rhythm And Blues Orchestra, which often includes singers Sam Brown and Ruby Turner. In January 2005 Holland and his band performed with Eric Clapton as the headline act of the Tsunami Relief Cardiff.

Personal[link]

Holland lives in the Westcombe Park area of Blackheath in South East London, where he had his studio, Helicon Mountain, built to his design, inspired by Portmeirion, the setting for the 1960s TV series The Prisoner.^[6] Holland owns costumes and props from the series and occasionally appears wearing the trademark brown-with-white-pipe blazer featured in the series. In 1987, Holland demonstrated his love of the series and starred in a spoof documentary, The Laughing Prisoner, with Stephen Fry, Terence Alexander and Hugh Laurie.^[6] Much of it was shot on location in Portmeirion, with archive footage of Patrick McGoohan, and featuring musical numbers from Siouxsie and the Banshees, Magnum and XTC. Holland performed a number towards the end of the program. His younger brother, Christopher, also plays organ in the Rhythm & Blues Orchestra, as well as being a singer-songwriter.

Holland was an interviewer for The Beatles Anthology TV project, and appeared in the 1997 film Spiceworld as a musical director.

He received an OBE in 2003 in the Queen's Birthday Honours list, for services to the British music industry as a television presenter and musician. In September 2006 Holland was appointed a Deputy Lieutenant for Kent.^[7] He is also known for his charity work: in June 2006 he performed in Southend for HIV/AIDS charity Mildmay,^[8] and in early 2007 he performed at Wells and Rochester Cathedrals to raise money for maintaining cathedral buildings.^[9] He is also patron of the Drake Music Project^[10] and has raised many thousands of pounds for the charity.

Holland was appointed an Honorary Fellow of Canterbury Christ Church University at a ceremony held at Canterbury Cathedral on 30 January 2009.^[11]

On 29 August 2005 Holland married Christabel McEwen, his girlfriend of 15 years (between 1983 and 1995 she had been married to Edward Lambton, 7th Earl of Durham, before they divorced). The wedding, at St James's Church, Cooling near Rochester, was attended by many celebrities, including Ringo Starr, Robbie Coltrane, Stephen Fry, Lenny Henry, Noel Gallagher, Dawn French and Jennifer Saunders.^[12]

Holland is also a patron for The Milton Rooms, an Arts centre in Malton, North Yorkshire, along with Bill Nighy, Imelda Staunton and Kathy Burke.^[13]

Writing[link]

His 2007 biography, Barefaced Lies and Boogie Woogie Boasts was BBC Radio 4 "Book of the Week" in the week beginning 8 October 2007 and was read by Holland. He is the author or joint author of four other books.

Discography[link]

Albums that have charted and gained Certifications[link]

Releases[link]

• 1978 "Boogie Woogie '78" (EP)
• 1981 Jools Holland and His Millionaires
• 1984 Jools Holland Meets Rock 'A' Boogie Billy (U.S. release only)
• 1990 World Of His Own
• 1991 The Full Complement
• 1992 "Together Again" (single with Sam Brown)
• 1992 The A-Z Geographer's Guide To The Piano
• 1994 Solo Piano
• 1994 Live Performance
• 1996 " Jazz & Rock & Roll
• 1997 Lift The Lid
• 1998 Best Of
• 1999 Sunset Over London
• 2000 Hop The Wag
• 2001 Small World Big Band
• 2002 SWBB Volume Two: More Friends
• 2003 Jack O The Green (SWBB Friends 3)
• 2004 Tom Jones & Jools Holland
• 2005 Beatroute
• 2005 Swinging the Blues, Dancing the Ska
• 2006 Moving Out To The Country
• 2007 Best of Friends
• 2008 The Collection
• 2008 The Informer (With Ruby Turner)
• 2008 "The Informer" (single with Ruby Turner)
• 2009 "I Went By" (single with Louise Marshall)
• 2010 Rockinghorse

Film and television[link]

• 1981 Urgh! A Music War
• 1982 Police: Around the World
• 1982 - 1987 The Tube (Host for 121 editions)
• 1983 Rebellious Jukebox: Compere
• 1984 The Young Ones: Punk (Episode entitled "Summer Holiday")
• 1985 Walking To New Orleans (Jools Holland in New Orleans)
• 1987 Eat the Rich: Sun Reporter
• 1987 Filthy Rich & Catflap: Strip Show Pianist (Episode #1.3)
• 1987 The Laughing Prisoner: No. 7
• 1987 French and Saunders (Episode 1.5)
• 1988 Sunday Night: Host (unknown episodes)
• 1989 Juke Box Jury: Host (unknown episodes)
• 1989 The Groovy Fellers Himself, 6 episodes
• 1991 Mr Roadrunner (Jools Holland in Memphis)
• 1994 There's No Business: Pianist
• 1994 - 1995 Don't Forget Your Toothbrush
• 1995 The Beatles Anthology
• 1997 Spice World: Musical Director
• 1997 Name That Tune: Host and Pianist
• 2003 Jool's History of the Piano : Presenter
• 2006 Top Gear: Star in a reasonably priced car.
• 2007 Fairport@Forty: Interview
• 2009 Chop Shop Rover Concept: The Jet 1 Car : Customer

Current television programmes[link]

• 1992–present Later... with Jools Holland
• 1993–present Hootenanny

Books[link]

• "Rolling Stones": A Life on the Road, (with Dora Loewenstein), Viking/Allen Lane (1998), (ISBN 0-670-88051-5)
• Beat Route: Journeys Through Six Counties, Weidenfeld & Nicholson (1998), (ISBN 0-575-06700-4)
• Ray Charles: Man and Music, (with Michael Lydon), Payback Press (1999), (ISBN 0-86241-929-8)
• Hand That Changed Its Mind , International Music Publications (2007), (ISBN 1-84328-645-9)
• Barefaced Lies and Boogie-woogie Boasts , Michael Joseph Ltd (2007), (ISBN 0-7181-4915-7)

References[link]

External links[link]

• Official website
• Jools Holland at the Internet Movie Database
• Jools Holland's Radio 2 show at BBC Programmes
• Later...with Jools Holland at BBC Online