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The Tephritidae are one of two
fly Flies are insects of 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 ...
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 Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
''
Drosophila ''Drosophila'' () is a genus of flies, belonging to the family Drosophilidae, whose members are often called "small fruit flies" or (less frequently) pomace flies, vinegar flies, or wine flies, a reference to the characteristic of many speci ...
'' (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 the Tephritidae. Description, recategorization, and
genetic analyses Genetic analysis is the overall process of studying and researching in fields of science that involve genetics and molecular biology. There are a number of applications that are developed from this research, and these are also considered parts of ...
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 realm A biogeographic realm or ecozone is the broadest biogeographic division of Earth's land surface, based on distributional patterns of terrestrial organisms. They are subdivided into bioregions, which are further subdivided into ecoregions. ...
s.


Description

For terms see Morphology of Diptera an
Tephritidae glossary
Tephritids are small to medium-sized (2.5–10 mm) flies that are often colourful, and usually with pictured wings, 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, 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 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 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 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.


Ecology

The larvae of almost all Tephritidae are phytophagous. Females deposit eggs in living, healthy plant tissue using their telescopic
ovipositor The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
s. Here, the
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...
e 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 Behavioral ecology, also spelled behavioural ecology, is the study of the evolutionary basis for ethology, animal behavior due to ecology, ecological pressures. Behavioral ecology emerged from ethology after Niko Tinbergen outlined Tinbergen's f ...
of tephritid fruit flies 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 Batesian mimicry is a form of mimicry where a harmless species has evolved to imitate the warning signals of a harmful species directed at a predator of them both. It is named after the English naturalist Henry Walter Bates, after his work on ...
, 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 the host plant and feeding on pollen, nectar, rotting plant debris, or honeydew. Natural enemies include
parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
wasps of the genera
Diapriidae The Diapriidae are a family of parasitoid wasps. These tiny insects have an average length of 2–4 mm and never exceed 8 mm. They typically attack larvae and pupae of a wide range of insects, especially flies. The about 2,300 described ...
and
Braconidae The Braconidae are a family of parasitoid wasps. After the closely related Ichneumonidae, braconids make up the second-largest family in the order Hymenoptera, with about 17,000 recognized species and many thousands more undescribed. One analysis ...
.


Economic importance

Tephritid fruit flies are of major economic importance in
agriculture Agriculture or farming is the practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that enabled people ...
. Some have negative effects, some positive. Various species of fruit flies cause damage to fruit and other plant crops. The genus ''
Bactrocera ''Bactrocera'' is a large genus of tephritid fruit flies, with close to 500 species currently described and accepted. Name The genus name is derived from Ancient Greek ''bakter'' "rod" and ''kera'' "horn". Systematics Prior to the 1990s, almo ...
'' is of worldwide notoriety for its destructive impact on agriculture. The
olive fruit fly The olive fruit fly (''Bactrocera oleae'') is a species of fruit fly which belongs to the subfamily Dacinae. It is a phytophagous species whose larvae feed on the fruit of olive trees, hence the common name. It is considered a serious pest in ...
(''B. oleae''), for example, feeds on only one plant: the wild or commercially cultivated
olive The olive, botanical name ''Olea europaea'', meaning 'European olive' in Latin, is a species of small tree or shrub in the family Oleaceae, found traditionally in the Mediterranean Basin. When in shrub form, it is known as ''Olea europaea'' ' ...
, ''Olea europaea''. It has the capacity to ruin 100% of an olive crop by damaging the fruit. ''
Bactrocera dorsalis ''Bactrocera dorsalis'', previously known as ''Dacus dorsalis'' and commonly referred to as the oriental fruit fly, is a species of tephritid fruit fly that is endemic to Southeast Asia. It is one of the major pest species in the genus ''Bact ...
'' is another highly invasive pest species that damages tropical fruit, vegetable, and nut crops.''
Euleia heraclei ''Euleia heraclei'', known as the celery fly or the hogweed picture-wing fly is a species of tephritid or fruit flies in the genus '' Euleia'' of the family Tephritidae. Distribution This species is widespread in most of Europe (Austria, Belgi ...
'' is a pest of celery and parsnips. The genus ''
Anastrepha ''Anastrepha'' is the most diverse genus in the American tropics and subtropics. Currently, it comprises more than 300 described species, including nine major pest species, such as the Mexican fruit fly (''A. ludens''), the South American fruit ...
'' includes several important pests, notably '' A. grandis'', '' A. ludens'' (Mexican fruit fly), '' A. obliqua'', and '' A. suspensa''. Other pests are ''
Strauzia longipennis ''Strauzia longipennis'' is a large species of tephritid fruit fly known by the common name sunflower maggot. It is a minor pest whose larvae mine stems of sunflowers (Helianthus spp.). Damage from larval feeding on spongy tissue is usually lig ...
'', a pest of sunflowers and ''
Rhagoletis mendax ''Rhagoletis mendax'' is a species of tephritid fruit fly known by the common name blueberry maggot. The blueberry maggot is closely related to the apple maggot (''R. pomonella''), a larger fruit fly in the same genus. It is a major pest of plan ...
'', 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 The Queensland fruit fly (''Bactrocera tryoni'') is a species of fly in the family Tephritidae in the insect order Diptera. ''B. tryoni'' is native to subtropical coastal Queensland and northern New South Wales. They are active during the day ...
'') is responsible for more than $28.5 million in damage to Australian fruit crops 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 ''Urophora'' is a genus of tephritid or fruit flies in the family Tephritidae. Species *'' Urophora acompsa'' ( Hendel, 1914) *'' Urophora aerea'' ( Hering, 1942) *'' Urophora affinis'' (Frauenfeld, 1857) *'' Urophora agnata'' ( Hering, 1942) ...
'' are used as control agents against rangeland-destroying noxious weeds such as starthistles and knapweeds, but their effectiveness is questionable. ''
Urophora sirunaseva ''Urophora sirunaseva'' is a species of tephritid or fruit flies in the genus '' Urophora'' of the family Tephritidae. Distribution Greece, Turkey, Moldova, Ukraine, Israel; introduced to North America North America is a continent ...
'' produces larvae that pupate within a woody
gall Galls (from the Latin , 'oak-apple') or ''cecidia'' (from the Greek , anything gushing out) are a kind of swelling growth on the external tissues of plants, fungi, or animals. Plant galls are abnormal outgrowths of plant tissues, similar to be ...
within the flower and disrupt seed production. ''
Chaetorellia acrolophi ''Chaetorellia acrolophi'' is a species of tephritid fruit fly known as the knapweed peacock fly. It is used as an agent of biological pest control against noxious knapweeds, especially spotted knapweed (''Centaurea maculosa''). The adult fl ...
'' is an effective biocontrol agent against knapweeds ''
Chaetorellia australis ''Chaetorellia australis'' is a species of tephritid fruit fly known as the yellow starthistle peacock fly. It is used as an agent of biological pest control against the noxious weed yellow starthistle (''Centaurea solstitialis''). The adult f ...
'' and ''
Chaetorellia succinea ''Chaetorellia succinea'' is a species of tephritid fruit fly that was accidentally released in 1991 into the United States and had since become one of the major biological pest controls against the noxious weed yellow starthistle (''Centaurea ...
'', deposit eggs into the starthistle seedheads, where their larvae consume the seeds and flower ovaries.Turner, C.E., G.L. Piper and E.M. Coombs. 1996. ''Chaetorellia australis'' (Diptera: Tephritidae) for biological control of yellow starthistle, ''Centaurea solstitialis'' (Compositae), in the western USA: establishment and seed destruction. Bull. Entomol. Res. 86: 1 77-182. 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 the use of cover sprays with conventional
pesticides Pesticides are substances that are meant to control pests. This includes herbicide, insecticide, nematicide, molluscicide, piscicide, avicide, rodenticide, bactericide, insect repellent, animal repellent, microbicide, fungicide, and la ...
, 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 Integrated pest management (IPM), also known as integrated pest control (IPC) is a broad-based approach that integrates both chemical and non-chemical practices for economic control of pests. IPM aims to suppress pest populations below the eco ...
.


Systematics

Tephritidae is divided into several subfamilies: * Blepharoneurinae (5 genera, 34 species) * Dacinae (41 genera, 1066 species) * Phytalmiinae (95 genera, 331 species) * Tachiniscinae (8 genera, 18 species) * Tephritinae (211 genera, 1859 species) * Trypetinae (118 genera, 1012 species) The genera ''Oxyphora'', '' Pseudorellia'', and ''Stylia'' comprise 32 species, and are not included in any subfamily ('' incertae sedis'').


Identification

*Richard H. Foote, P. L. Blanc, Allen L. Norrbom, 1993 ''Handbook of the Fruit Flies (Diptera: Tephritidae) of America North of Mexico'' Cornell University Press (Comstock Publishing). *Merz, B. 1994. Diptera Tephritidae. ''Insecta Helvetica Fauna'' 10: 1-198. *White, I.M. 1988. Tephritid flies. Diptera: Tephritidae. '' Handbooks for the Identification of British Insects'' 10(5a): 1-134. *White I.M. & Elson-Harris M.M. 1994 ''Fruit Flies of Economic Significance: their Identification and Bionomics''. 2nd ed. International Institute of Entomology, London. *R.A.I. Drew and Meredith C Romig ''Tropical Fruit Flies of South-East Asia'' (Tephritidae: Dacinae) CABI * Hendel1914. Die Gattungen der Bohrfliegen. ''Wein. Entomol. Ztg.'' 33: 73–98. Keys to World genera Out of date but still the only world
monograph A monograph is a specialist work of writing (in contrast to reference works) or exhibition on a single subject or an aspect of a subject, often by a single author or artist, and usually on a scholarly subject. In library cataloging, ''monogra ...
. *Hendel, F., 1927. Trypetidae.In: Lindner, E. (Ed.). ''Die Fliegen der Paläarktischen Region'' 5, 49, 1-221. Keys to Palaearctic species but now needs revision (in German). * Séguy, E. (1934) Diptères: Brachycères. II. Muscidae acalypterae, Scatophagidae. Paris: Éditions Faune de France 28
virtuelle numérique
*Rikhter, V.A. Family Conopidae in Bei-Bienko, G. Ya, 1988 ''Keys to the insects of the European Part of the USSR'' Volume 5 (Diptera) Part 2 English edition. Keys to Palaearctic species but now needs revision.


Species lists


West Palaearctic including RussiaJapanWorld list


Gallery

Image:Urophora.cardui.male.jpg, '' Urophora cardui'', male Image:Strauzia longipennis.jpg, ''
Strauzia longipennis ''Strauzia longipennis'' is a large species of tephritid fruit fly known by the common name sunflower maggot. It is a minor pest whose larvae mine stems of sunflowers (Helianthus spp.). Damage from larval feeding on spongy tissue is usually lig ...
'' Image:Fruit fly Euaresta.jpg, '' Euaresta aequalis'' Image:Procecidochares atra.jpg, ''
Procecidochares atra ''Procecidochares atra'' is a species of tephritid or fruit flies in the genus '' Procecidochares'' of the family Tephritidae. Its galls are most common on ''Solidago altissima, S. gigantea, S. rugosa''.http://bugguide.net/node/view/68596 Bugg ...
'' Image:Euleia heraclei01.jpg, ''
Euleia heraclei ''Euleia heraclei'', known as the celery fly or the hogweed picture-wing fly is a species of tephritid or fruit flies in the genus '' Euleia'' of the family Tephritidae. Distribution This species is widespread in most of Europe (Austria, Belgi ...
'' Image:Chaetostomella_cylindrica.jpg, '' Cheatostomella cylindrica'' Image:Xyphosia.miliaria.male.jpg, ''
Xyphosia miliaria ''Xyphosia miliaria'' is a species of tephritid or fruit flies in the family Tephritidae. Distribution This species is present in most of Europe, in the eastern Palearctic realm, and in the Near East. Habitat This species occurs in wet meadows ...
'' Image:GoldenrodGallFlyLarva.jpg, ''Eurosta solidaginis'', gall maker on goldenrod File:Procecidochares atra galls.jpg, Gall of ''Procecidochares atra'' on ''Solidago'' File:Rhagoletis suavis.jpg, '' Rhagoletis suavis'' File:Eurosta solidaginis female.jpg, ''Eurosta solidaginis'', female


References


Further reading

*Christenson, L. D. and Foote, R.H. 1960. Biology of fruit flies, ''Annu. Rev. Entomol.'', vol. 5, pp. 171–192. *Bruce A. McPheron, Gary J. Steck (Editors), 1996 ''Fruit fly pests : a world assessment of their biology and management'' International Symposium on Fruit Flies of Economic Importance (4th : 1994 : Sand Key, Florida, USA) Delray Beach, Fla. : St Lucie Press *Foote R.H., Steyskal G.C. 1981 Tephritidae. in: McAlpine J.F. (Ed.), ''Manual of Nearctic Diptera''. Agriculture Canada, Ottawa, pp. 817–831
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External links

;General
Natural Enemies of True Fruit Flies (Tephritidae)
USDA The United States Department of Agriculture (USDA) is the federal executive department responsible for developing and executing federal laws related to farming, forestry, rural economic development, and food. It aims to meet the needs of comme ...

Tephritidae Information from the Diptera Site
* Tephritid Workers Database ;Identification
The Diptera site
Comprehensive guide to identification literature with a worldwide perspective. ;Galleries
Diptera.info imagesImages at BugGuideFamily Tephritidae at EOL
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