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14 December 2020

Bactrocera dorsalis (Oriental fruit fly)

Datasheet Types: Pest, Natural enemy, Invasive species

Abstract

This datasheet on Bactrocera dorsalis covers Identity, Overview, Distribution, Dispersal, Hosts/Species Affected, Diagnosis, Biology & Ecology, Environmental Requirements, Natural Enemies, Impacts, Prevention/Control, Further Information.

Identity

Preferred Scientific Name
Bactrocera dorsalis (Hendel, 1912)
Preferred Common Name
Oriental fruit fly
Other Scientific Names
Bactrocera (Bactrocera) dorsalis Drew & Hancock, 1994
Bactrocera (Bactrocera) invadens Drew et al., 2005
Bactrocera (Bactrocera) papayae Drew & Hancock, 1994
Bactrocera (Bactrocera) philippinensis Drew & Hancock, 1974
Bactrocera (Bactrocera) variabilis Lin & Wang
Bactrocera ferruginea Bezzi, 1913
Bactrocera invadens Drew, Tsuruta & White
Bactrocera papayae Drew & Hancock
Bactrocera philippinensis
Chaetodacus ferrugineus Bezzi, 1916
Chaetodacus ferrugineus dorsalis Bezzi, 1916
Chaetodacus ferrugineus var. dorsalis Hendel, 1915
Chaetodacus ferrugineus var. okinawanus Shiraki, 1933
Dacus (Bactrocera) dorsalis Hardy, 1977
Dacus (Bactrocera) semifemoralis Tseng et al., 1992
Dacus (Bactrocera) vilanensis Tseng et al., 1992
Dacus (Strumeta) dorsalis Hardy & Adachi, 1956
Dacus dorsalis Hendel, 1912
Dacus ferrugineus (Fabricius, 1805)
Musca ferruginea Fabricius, 1794, preocc.
Strumeta dorsalis Hering, 1956
Strumeta dorsalis okinawa Shiraki, 1968
Strumeta ferruginea Hering, 1956
International Common Names
Spanish
mosca oriental de la fruta
French
mouche de fruits asiatique
mouche orientale des arbres fruitiers
Portuguese
mosca oriental das frutas
Local Common Names
Germany
Orientalische Fruchtfliege
Japan
mikan-ko-mibae
Netherlands
mangga-vlieg
EPPO code
DACUDO (Bactrocera dorsalis)

Pictures

Bactrocera dorsalis (Oriental fruit fly); an aggregation of male flies on the spadix of an aroid, gathering eugenol to augment their pheromone. Wat Palad, Doi Suthep, Chiang Mai, Thailand. July 1994.
Aggregation of males
Bactrocera dorsalis (Oriental fruit fly); an aggregation of male flies on the spadix of an aroid, gathering eugenol to augment their pheromone. Wat Palad, Doi Suthep, Chiang Mai, Thailand. July 1994.
©R. Davies/Natural History Museum
Bactrocera dorsalis (Oriental fruit fly); adult.
Adult
Bactrocera dorsalis (Oriental fruit fly); adult.
©Clive Lau
Bactrocera dorsalis (Oriental fruit fly); adult.
Adult
Bactrocera dorsalis (Oriental fruit fly); adult.
©Clive Lau
Bactrocera dorsalis (Oriental fruit fly); adult.
Adult
Bactrocera dorsalis (Oriental fruit fly); adult.
©Clive Lau
Bactrocera dorsalis (Oriental fruit fly); adult. Museum set specimen.
Adult
Bactrocera dorsalis (Oriental fruit fly); adult. Museum set specimen.
©CABI BioScience
Bactrocera dorsalis; Adult specimen.
Museum set specimen
Bactrocera dorsalis; Adult specimen.
©CABI BioScience
Bactrocera dorsalis; Adult specimen.
Museum set specimen
Bactrocera dorsalis; Adult specimen.
©CABI BioScience
Bactrocera dorsalis; adult male fly at rest.
Adult fly
Bactrocera dorsalis; adult male fly at rest.
Marc De Meyer

Summary of Invasiveness

Bactrocera dorsalis is a highly invasive species. Native to Asia, Oriental fruit fly is now found in at least 65 countries, including parts of America and Oceania, and most of continental Africa (sub-Saharan countries). The potential risk of its introduction to a new area is facilitated by increasing international tourism and trade, and is influenced by changes in climate and land use. After introduction, it can easily disperse as it has a high reproductive potential, high biotic potential (short life cycle, up to 10 generations of offspring per year depending on temperature), a rapid dispersal ability and a broad host range. The economic impact would result primarily from the loss of the export markets and the costly requirement of quarantine restrictions and eradication measures. Furthermore, its establishment would have a serious impact on the environment, following the initiation of chemical and/or biological control programmes. Invasive B. dorsalis has been shown to be highly competitive with native fruit flies where it has established, quickly becoming the dominant fruit fly pest.
Oriental fruit fly is of quarantine significance to EPPO (European Plant Protection Organization), APPPC (Asia and Pacific Plant Protection Commission), COSAV (Comité de Sanidad Vegetal del Cono Sur), CPPC (Caribbean Plant Protection Commission), IAPSC (Inter-African Phytosanitary Council) and OIRSA (Organismo Internacional Regional de Sanidad Agropecuaria) countries.

Taxonomic Tree

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Notes on Taxonomy and Nomenclature

Bactrocera dorsalis is a member of the Oriental fruit fly (B. dorsalis) species complex. This species complex forms a group within the subgenus Bactrocera and the name may therefore be cited as Bactrocera (Bactrocera) dorsalis. B. dorsalis was originally treated as a single species, widespread over Asia, until it was split into several species, with the description of Bactrocera carambolae, B. papayae and B. philippinensis by Drew and Hancock (1994). Native range of true B. dorsalis became restricted primarily to continental Asian countries north of the Malay Peninsula. Bactrocera invadens was later described by Drew et al. (2005), when established populations were detected in East Africa (Lux et al., 2003) and in West Africa (Vayssières, 2004). Bactrocera philippinensis was designated a synonym of B. papayae by Drew and Romig (2013). Extensive research was carried out to delimitate species boundaries, based on morphological, molecular, cytogenetic, behavioural and chemoecological data by multidisciplinary teams, in great part coordinated under an FAO/IAEA Coordinated Research Project (CRP) on the ‘Resolution of cryptic species complexes of tephritid pests to overcome constraints to SIT application and international trade’. This resulted in the synonymization of B. invadens and B. papayae under B. dorsalis and leaving B. carambolae as a distinct species by Schutzeet al. (2015), who summarized the extensive research and evidence supporting the synonymization. Records of B. pedestris (Bezzi) from outside of the Philippines are mostly based on misidentifications of B. dorsalis.

Description

Eggs

The eggs of Bactrocera oleae were described in detail by Margaritis (1985) and those of other species are probably very similar. They are 0.8 mm long and 0.2 mm wide, with the micropyle protruding slightly at the anterior end, and white to yellow-white. The chorion is reticulate (requires scanning electron microscope examination).

Larvae

The following larval description was taken from White and Elson-Harris (1994):
B. dorsalis third-instar larva: medium-sized: 7.5-10.0 mm long and 1.5-2.0 mm wide;
Head: stomal sensory organ with three to four sensilla, surrounded by five large, unserrated preoral lobes; oral ridges with 11-14 rows of blunt edged, short teeth; accessory plates 12-15, shell-shaped with small, rounded teeth; mouth hooks moderately sclerotized, without pre-apical teeth.
Thoracic and abdominal segments: anterior portion of each thoracic segment with an encircling band of several discontinuous rows of small spinules; T1 with 9-11 rows of large, sharply pointed spinules; T2 spinules small, stout, sharply pointed with five to six rows dorsally, three to four rows laterally, five to seven rows ventrally; T3 spinules similar to those on T2, two to four rows dorsally, one to three rows laterally, three to five rows ventrally. Creeping welts with small, stout spinules, with one posterior row of spinules larger and stouter than remainder. A8 rounded with prominent intermediate areas and obvious sensilla.
Anterior spiracles: 8-12 tubules.
Posterior spiracles: spiracular slits thick walled, approximately 2.5-3.0 times as long as broad. Spiracular hairs just longer than a spiracular slit; dorsal and ventral bundles with 17-20 broad, flat hairs, branched apically; lateral bundles with 8-12 similarly shaped hairs.
Anal area: lobes protuberant, surrounded by three to five discontinuous rows of spinules. The inner rows of spinules stout, slightly curved, sharply pointed becoming larger just below the anal opening, outer rows with smaller spinules.

Puparium

Barrel-shaped with most larval features unrecognisable, the exception being the anterior and posterior spiracles, which are little changed by pupariation. White to yellow-brown. Usually approximately 60-80% the length of the larva.

Adults

Drew and Hancock (1994) distinguish the B. dorsalis species complex as follows: Bactrocera (Bactrocera) spp. with a clear wing membrane, except for a narrow costal band (not reaching R4+5); cells bc and c colourless (except in a few non-pests with a very pale tint) with microtrichia restricted to outer corner of cell c. Scutum generally black with lateral vittae present and medial vitta absent; yellow scutellum, except for basal band which is usually very narrow; abdomen with a medial dark stripe on T3-T5; dark laterally (but form of marking varies from species to species). B. dorsalis belongs to a subgroup that has yellow postpronotal lobes, parallel lateral vittae, and femora not extensively marked. Within this group it is distinguished by its short to long aculeus/aedeagus; tomentum with no gap; narrow costal band; generally narrow but sometimes extensive abdominal markings. It is noteworthy that colour of scutum varies in B. dorsalis from generally black to black with an extensive lanceolate red-brown pattern to almost entirely red-brown. Populations from the Indian subcontinent and Africa have extensive pale markings (Leblanc et al., 2013a), whereas specimens from Asia east of Myanmar mostly have dark scutum.

Distribution

The revision by Drew and Hancock (1994) split the original B. dorsalis into B. carambolae and three species, B. dorsalis, B. papayae and B. philippinensis, with mutually exclusive geographic ranges. B. dorsalis sensu stricto became restricted to mainland Asia (and Taiwan) and its adventive populations in Hawaii and French Polynesia, and newly described B. papayae ranged from southern Thailand and Peninsular Malaysia through East Malaysia and all Indonesian islands to New Guinea Island, and B. philippinensis was restricted to the Philippines and introduced to Palau. When B. dorsalis invaded continental Africa, around 2003, it was described as B. invadens, the origin and native range of which was believed to be Sri Lanka (Drew et al., 2005). With the exception of B. carambolae, all of these species are now treated together as B. dorsalis sensu lato, which is the most destructive, invasive and widespread of all Dacine fruit flies. The distribution and invasion history of B. dorsalis are summarized on a map published in Vargas et al. (2015). EPPO (2014) includes California, USA, in the distribution because the fly is repeatedly trapped there in small numbers. Whether or not B. dorsalis is actually established in continental America is a hotly debated topic (Papadopoulos et al., 2013; Suckling et al., 2014).
The distribution of B. dorsalis was mapped recently by IIE (1994a) and more recently by CABI/EPPO (2013). See also CABI/EPPO (1998, No. 24). Records of B. dorsalis in Guam, Northern Mariana Islands and New Caledonia published in IIE (1994a) and in previous versions of the Compendium were incorrect; B. dorsalis has been absent from Guam and the Northern Mariana Islands since 1965 (Leblanc, 1997) and was never present in New Caledonia. A distribution map of B. dorsalis across the world can be seen on the Global Biodiversity Information Facility (GBIF) website. A more detailed distribution of B. dorsalis in Africa can also be viewed on the website True Fruit flies (Diptera: Tephritidae) of the Afrotropical Region.

Distribution Map

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Distribution Table

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History of Introduction and Spread

Oriental fruit fly has been established since about 1945 and quickly became widespread in the Hawaiian Islands (Pemberton, 1946).
B. dorsalis (as B. papayae) is believed to have been introduced accidentally into the eastern Indonesian province of Irian Jaya [Papua Barat] prior to 1992, when it was first detected in the border areas of Papua New Guinea (Sar et al., 2001). By 2000 it had spread throughout much of mainland Papua New Guinea (Sar et al., 2001). In March 1993, it was detected on several northern islands in Torres Strait, Queensland (Fay et al., 1997). This represents the first detection of this known invasive species in Australian territory and it was quickly eradicated. In October 1995, it was detected in the Cairns region of northern Queensland (Fay et al., 1997; Hancock et al., 2000b). This was probably the result of accidental introduction from Papua New Guinea. The fly had spread throughout the Cairns-Mareeba-Mossman region and detections were made from Cooktown to Cardwell before it was eradicated during 1997-1998 (Hancock et al., 2000b; Cantrell et al., 2002). An isolated outbreak at Mount Isa in western Queensland was eradicated during 1997 (Hancock et al., 2000; Cantrell et al., 2002). Since then, there have been occasional incursions onto Torres Strait islands from Papua New Guinea. These have been eradicated whenever establishment occurred.
In 1991, the Ministry of Agriculture in Mauritius established a network of quarantine traps for exotic fruit flies. In June 1996, one Oriental fruit fly (B. dorsalis) was found in a trap near the airport in Mauritius (White, 2006). The quarantine trap grid was immediately extended in the area surrounding the airport, and fruit in the area was inspected for larval infestations. The larvae were reared from infested fruit found near the airport and it was clear that the oriental fruit fly had established in southern Mauritius. Morphological examination indicated that the flies had originated in southern India. An eradication programme for B. dorsalis infesting various tree crops was conducted from July 1996 to April 1998, in the southern region of Mauritius, using bait application technique (BAT), male annihilation technique (MAT), cover spray of trees with ripening fruits, soil drenching under trees with ripening and fallen fruits, and fruit clean-up and disposal. The introduction was probably accidental, as the first flies were detected in the airport neighbourhood.
In 2003, an unknown Bactrocera species was found in Kenya (Lux et al., 2003). Taxonomic expertise showed that it could not be a native species of Africa, but that it proved to be a member of the B. dorsalis complex. Identical specimens were collected earlier during a survey in Sri Lanka in 1993 and initially classified as aberrant forms of B. dorsalis. It was decided that both populations belonged to the same, hitherto undescribed species: B. invadens, which was formally described in 2005 (Drew et al., 2005) and designated a synonym of B. dorsalis a decade later (Schutze et al., 2015).
After its discovery in Kenya, it was recorded in a number of countries in eastern, central and western Africa in a relatively short time (Mwatawala et al., 2004; Drew et al., 2005; Meyer et al., 2007). This threat has also been reported in 2004 in Sudan (Luckman, 2004), Cameroon (Abanda et al., 2008), Senegal (Vayssières, 2004), Benin (Vayssières et al., 2005) and other West African countries. Recently, the species has also been reported from southern Africa (Meyer et al., 2007), southern India (Sithanantham et al., 2006) and Bhutan (Drew et al., 2007).
The exact invasion pathway into Africa is unknown. From 1999 to 2004, an intensive sampling programme was conducted in Kenya (Copeland et al., 2004). During that period, close to 4000 fruit samples (comprising approximately 980,000 pieces of fruit) belonging to 882 taxa and 116 plant families, were collected from coastal and western Kenya, as well as from the Central Highlands. Some trapping, including the use of methyl eugenol, was also carried out as a parallel study from 1999 to 2000 (IM White, UK [address available from CABI], personal communication, 2008). It was only in March 2003 that the first specimens were collected from the coastal region (Lux et al., 2003).
Vayssières and Kalabane (2000) and Vayssières et al. (2004) conducted intensive fruit fly sampling in commercial mango (Mangifera indica) orchards in different localities in western Africa, in Coastal Guinea and Mali, from 1992 to 1995 and 2000, respectively. None of these yielded any specimens of B. dorsalis. The first specimens in that part of the African mainland were detected in June 2004 (Drew et al., 2005; Vayssières, 2004). The presence of this species in those eastern or western African countries before the beginning of the twenty-first century is, therefore, very unlikely. Unfortunately, no similar studies were conducted at that time in other parts of the African continent where the fly is currently found. The fact that the first specimens were reported from the East African coast appears to indicate that the port of entry could have been an East African (coastal) locality, although there is no proof of this.

Introductions

Introduced toIntroduced fromYearReasonsIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
MauritiusIndia1996 YesNoEradicated in June 1998. The Ministry of Agriculture declared Mauritius free of oriental fruit fly

Risk of Introduction

The major risk is from the import of fruit containing larvae, either as part of cargo, or through the smuggling of fruit in airline passenger baggage or mail. For example, in New Zealand, Baker and Cowley (1991) recorded 7-33 interceptions of fruit flies per year in cargo and 10-28 per year in passenger baggage. Individuals who successfully smuggle fruit are likely to discard it when they discover that it is rotten. This method of introduction probably accounts for the discovery of at least one fly in a methyl eugenol trap in California, USA every year (Foote et al., 1993), although immediate implementation of eradication action plans has ensured that the fly has never been able to establish a proper breeding population, a view that has been challenged in recent years (Papadopoulos et al., 2013).

Means of Movement and Dispersal

B. dorsalis is known to have the potential to establish adventive populations in various tropical and subtropical areas. Using microsatellite markers, Qin et al. (2018) conducted a population genetic study of B. dorsalis across its entire geographical range. Based on this study, the authors concluded that the source of the pest within Asia was South Asia and populations of B. dorsalis introduced in Africa and Hawaii were most likely from South Asia (Qin et al., 2018).
Adult flight and the transport of infected fruit are the major means of movement and dispersal to previously uninfested areas.
[Erratum: In previous versions of this datasheet, it was stated that “many Bactrocera spp. can fly 50-100 km (Fletcher, 1989)” but a review of Fletcher (1989a) and Fletcher (1989b) by Hicks et al. (2019) found no evidence to support this statement and it has been removed. Fletcher (1989b) provides dispersal data for only 11 of 651 species of Bactrocera, many of the case studies lack the necessary numerical data, and the study did not discern between active flight and passive wind-assisted dispersal. There are differences among fruit fly species and further studies are required to determine dispersal distances for individual species. For further information on trapping Bactrocera species to monitor movement, see Weldon et al. (2014).] 

Pathway Causes

Pathway causeNotesLong distanceLocalReferences
Crop production (pathway cause) YesYes 
Food (pathway cause) YesYes 
Hitchhiker (pathway cause) YesYes 
Live food or feed trade (pathway cause) YesYes 
Smuggling (pathway cause) YesYes 

Pathway Vectors

Pathway vectorNotesLong distanceLocalReferences
Aircraft (pathway vector)Fruits infested with larvae and/or eggsYes  
Bulk freight or cargo (pathway vector)All life stages. In New Zealand, 7-33 interceptions of fruit flies per year in cargoYes  
Consumables (pathway vector)All life stagesYes  
Floating vegetation and debris (pathway vector)Fruits infested with larvae and/or eggsYesYes 
Land vehicles (pathway vector)Fruits infested with eggs, larvae and/or pupae Yes 
Luggage (pathway vector)Fruits infested with larvae and/or eggs. 10-28 interceptions per year in passenger baggage in NZYesYes
Mail (pathway vector)Fruits infested with eggs and/or larvaeYesYes 
Plants or parts of plants (pathway vector)Fruits infested with eggs and/or larvaeYesYes 
Soil, sand and gravel (pathway vector)PupaeYesYes 
Wind (pathway vector)Adults Yes 

Plant Trade

Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
Fruits (inc. pods)
arthropods/eggs
arthropods/larvae
Yes Pest or symptoms usually visible to the naked eye
Growing medium accompanying plants
arthropods/pupae
Yes Pest or symptoms usually visible to the naked eye
Plant parts not known to carry the pest in trade/transport
Bark
Bulbs/Tubers/Corms/Rhizomes
Flowers/Inflorescences/Cones/Calyx
Leaves
Roots
Seedlings/Micropropagated plants
Stems (above ground)/Shoots/Trunks/Branches
True seeds (inc. grain)
Wood

Wood Packaging

Wood packaging not known to carry the pest in trade/transportTimber typeUsed as packing
Loose wood packing material Yes
Non-wood Yes
Processed or treated wood Yes
Solid wood packing material with bark Yes
Solid wood packing material without bark Yes

Hosts/Species Affected

Bactrocera dorsalis has over 70 species of commercial/edible and wild hosts (White and Elson-Harris, 1994; Tsuruta et al., 1997; Allwood et al., 1999De Meyer and White, 2004; Clarke et al., 2005; Leblanc et al., 2012, 2013b). It is a serious pest of a wide range of fruit crops throughout its native range and wherever it has invaded. In order to summarise the common host range of B. dorsalis within and outside of its native range, records listed in the table below were taken from two regions and sources: South-East Asia (Clarke et al., 2005) and Africa (De Meyer and White, 2004). Records which only listed the genera as hosts for B. dorsalis in these reference sources were left out in order to comply with the definition of a fruit fly host for a plant species or cultivar as provided in ISPM 37 (FAO, 2016).
A few of the host records of B. dorsalis in surveys conducted in and outside of Africa were subsequently challenged in detailed host status studies following guidelines provided in the International Standards for Phytosanitary Measures - Determination of host status of fruit to fruit flies (ISPM No. 37) (FAO, 2016). Cultivated and export grade forms of a few fruit species found to be hosts for B. dorsalis in field surveys were later confirmed as non-hosts for the pest. The following sources provide further details of studies confirming non-host status of cultivated forms of some of the fruit species found to be hosts in field surveys: 

Carica papaya: fruit at particular maturity stages are non-host (Cugala et al., 2017)
Citrus limon: commercial export grade is non-host (Manrakhan et al., 2018)
Garcinia mangostana: commercial export grade is non-host (Unahawutti et al., 2014)
Musa acuminata: mature green (cv. Cavendish Dwarf) is non-host (Cugala et al., 2014)
Persea americana: some export grade cultivated varieties are non-host (Ware et al., 2016

Host Plants and Other Plants Affected

HostFamilyHost statusReferences
Abelmoschus esculentus (okra)MalvaceaeUnknown
Acca sellowianaLithomyrtusUnknown
Alangium lamarckiiAlangiaceaeUnknown
Anacardium occidentale (cashew nut)AnacardiaceaeOther
Annona cherimola (cherimoya)AnnonaceaeOther
Annona montanaAnnonaceaeOther
Annona muricata (soursop)AnnonaceaeOther
Annona reticulata (bullock's heart)AnnonaceaeOther
Annona senegalensis (wild custard apple)AnnonaceaeOther
Annona squamosa (sugar apple)AnnonaceaeOther
Artabotrys monteiroae Unknown
Artocarpus altilis (breadfruit)MoraceaeOther
Artocarpus dadahMoraceaeUnknown
Artocarpus heterophyllus (jackfruit)MoraceaeOther
Averrhoa carambola (carambola)OxalidaceaeOther
Baccaurea angulata Unknown
Benincasa hispida (wax gourd)CucurbitaceaeUnknown
Blighia sapida (akee apple)SapindaceaeOther
Bulbophyllum patens Unknown
Canarium album (Chinese white olive)StrombidaeUnknown
Capparis tomentosa (woolly caper bush)CapparaceaeUnknown
Capsicum annuum (bell pepper)SolanaceaeUnknown
Capsicum frutescens (chilli)SolanaceaeOther
Carica papaya (pawpaw)CaricaceaeOther
Choerospondias axillaris (Nepali hog plum)AnacardiaceaeUnknown
Chrysophyllum albidumSapotaceaeOther
Chrysophyllum cainito (caimito)SapotaceaeOther
Chrysophyllum roxburghiiSapotaceaeUnknown
Citrullus lanatus (watermelon)CucurbitaceaeOther
CitrusRutaceaeUnknown
Citrus aurantium (sour orange)RutaceaeOther
Citrus clementinaRutaceaeUnknown
Citrus limon (lemon)RutaceaeOther
Citrus madurensis (calamondin)RutaceaeUnknown
Citrus maxima (pummelo)RutaceaeUnknown
Citrus reticulata (mandarin)RutaceaeOther
Citrus reticulata x paradisi (tangelo)RutaceaeUnknown
Citrus sinensis (sweet orange)RutaceaeOther
Citrus x paradisi (grapefruit)RutaceaeOther
Coffea canephora (robusta coffee)RubiaceaeUnknown
Cordia dentata (white manjack)BoraginaceaeUnknown
Cordia myxa (sebesten)BoraginaceaeWild host
Rwomushana et al. (2008)
Cordyla africanaFabaceaeWild host
Crescentia cujete (calabash tree)BignoniaceaeUnknown
Cucumis ficifoliusCucurbitaceaeUnknown
Cucumis melo (melon)CucurbitaceaeOther
Cucumis sativus (cucumber)CucurbitaceaeOther
Cucurbita (pumpkin)CucurbitaceaeUnknown
Cucurbita argyrosperma (silver-seed gourd)CucurbitaceaeUnknown
Cucurbita moschata (pumpkin)CucurbitaceaeUnknown
Cucurbita pepo (marrow)CucurbitaceaeOther
Cydonia oblonga (quince)RosaceaeOther
Dacryodes edulis (African pear)BurseraceaeUnknown
Dimocarpus longan (longan tree)SapindaceaeOther
Diospyros dasyphylla Unknown
Diospyros decandra Unknown
Diospyros ebenaster (black sapote)EbenaceaeUnknown
Diospyros lotus (Date plum tree)EbenaceaeUnknown
Diospyros mespiliformis (ebony diospiros)EbenaceaeUnknown
Diospyros montanaEbenaceaeOther
Dracaena steudneriAgavaceaeWild host 
Drypetes roxburghiiEuphorbiaceaeUnknown
Eriobotrya japonica (loquat)RosaceaeOther
Eugenia palumbis Unknown
Eugenia uniflora (Surinam cherry)LithomyrtusOther
Fagraea berteroanaLoganiaceaeUnknown
Ficus carica (common fig)MoraceaeUnknown
Ficus lepicarpa Unknown
Ficus microcarpa (Indian laurel tree)MoraceaeWild host
Ficus ottoniifoliaMoraceaeWild host 
Ficus sycomorus (sycamore fig)MoraceaeUnknown
Flacourtia indica (governor's plum)FlacourtiaceaeUnknown
Fortunella japonica (round kumquat)RutaceaeUnknown
Fragaria vesca (wild strawberry)RosaceaeUnknown
Garcinia costata Unknown
Garcinia hombronianaClusiaceaeWild host
Garcinia indica (kokum butter tree)ClusiaceaeUnknown
Garcinia intermediaClusiaceaeUnknown
Garcinia mangostana (mangosteen)ClusiaceaeOther
Garcinia manniiClusiaceaeWild host
Garcinia venulosa Unknown
Gymnopetalum integrifoliumCucurbitaceaeUnknown
Haematostaphis barteri Unknown
Hedychium coronarium (white butterfly ginger lily)ZingiberaceaeUnknown
Hesperethusa crenulataRutaceaeUnknown
Hexalobus monopetalusAnnonaceaeOther 
Icacina senegalensisIcacinaceaeUnknown
Irvingia gabonensis (wild mango)IrvingiaceaeOther
Kedrostis hirtella Unknown
Lagenaria siceraria (bottle gourd)CucurbitaceaeUnknown
Lepisanthes alata Unknown
Litchi chinensis (lichi)SapindaceaeOther
Luffa acutangula (angled luffa)CucurbitaceaeUnknown
Luffa aegyptiaca (loofah)CucurbitaceaeUnknown
Malus domestica (apple)RosaceaeOther
Mangifera indica (mango)AnacardiaceaeOther
Manilkara zapota (sapodilla)SapotaceaeOther
Melothria wallichiiCucurbitaceaeUnknown
Momordica balsamina (common balsam apple)CucurbitaceaeUnknown
Momordica charantia (bitter gourd)CucurbitaceaeOther
Momordica cochinchinensisCucurbitaceaeUnknown
Mosiera longipesLithomyrtusUnknown
Musa acuminata (wild banana)MusaceaeMain
Musa paradisiaca Unknown
Musa x paradisiaca (plantain)MusaceaeOther
Myrica rubraMyricaceaeUnknown
Nauclea latifolia (pin cushion tree)RubiaceaeOther 
Nephelium lappaceum (rambutan)SapindaceaeOther
Ochrosia mariannensis Unknown
Opilia amentacea Unknown
Pandanus tectorius (screw pine)PandanaceaeUnknown
Parkia biglobosa (néré)FabaceaeUnknown
Passiflora edulis (passionfruit)PassifloraceaeOther
Persea americana (avocado)LauraceaeOther
Phaseolus vulgaris (common bean)FabaceaeMain
Phoenix dactylifera (date-palm)ArecaceaeUnknown
Phyllanthus acidus (star gooseberry)EuphorbiaceaeUnknown
Physalis minima (Sunberry)SolanaceaeUnknown
Physalis peruviana (Cape gooseberry)SolanaceaeUnknown
Pouteria campechiana (canistel)SapotaceaeMain
Pouteria duclitanSapotaceaeUnknown
Pouteria sapota (mammey sapote)SapotaceaeUnknown
Prunus armeniaca (apricot)RosaceaeOther
Prunus avium (sweet cherry)RosaceaeOther
Prunus bokhariensisRosaceaeUnknown
Prunus cerasoidesRosaceaeUnknown
Prunus domestica (plum)RosaceaeOther
Prunus persica (peach)RosaceaeOther
Prunus persica var. nucipersica (nectarine)RosaceaeUnknown
Psidium cattleianum (strawberry guava)LithomyrtusUnknown
Psidium guajava (guava)LithomyrtusOther
Pyrus pashiaRosaceaeUnknown
Quassia simarouba (mountain damson)SimaroubaceaeUnknown
Saba comorensis Unknown
Saba senegalensisApocynaceaeWild host
Sapium baccatumEuphorbiaceaeUnknown
Sclerocarya birrea (marula)AnacardiaceaeWild host
Securinega virosaEuphorbiaceaeUnknown
Sesbania grandiflora (sesbania)FabaceaeUnknown
Siphonodon celastrineusSalaciaUnknown
Solanum aculeatissimumSolanaceaeUnknown
Solanum aethiopicum (african scarlet eggplant)SolanaceaeOther
Solanum americanumSolanaceaeUnknown
Solanum anguiviSolanaceaeOther
Solanum erianthum (potato tree)SolanaceaeUnknown
Solanum ferox Unknown
Solanum granuloso-leprosum Unknown
Solanum incanum (grey bitter-apple)SolanaceaeWild host
Solanum linnaeanum (apple of Sodom)SolanaceaeUnknown
Solanum lycopersicum (tomato)SolanaceaeOther
Solanum mauritianum (tobacco tree)SolanaceaeUnknown
Solanum melongena (aubergine)SolanaceaeOther
Solanum seaforthianum (Brazilian nightshade)SolanaceaeUnknown
Solanum sodomeumSolanaceaeWild host 
Solanum verbascifolium Unknown
Sorindeia madagascariensisAnacardiaceaeWild host
Rwomushana et al. (2008)
Spondias bipinnataAnacardiaceaeUnknown
Spondias dulcis (otaheite apple)AnacardiaceaeOther
Spondias mombin (hog plum)AnacardiaceaeOther
Strychnos mellodoraLoganiaceaeWild host 
Strychnos nux-vomica (nux-vomica tree)LoganiaceaeUnknown
Syzygium cumini (black plum)LithomyrtusUnknown
Syzygium jambos (rose apple)LithomyrtusMain
Syzygium malaccense (Malay apple)LithomyrtusMain
Terminalia brasiliensisCombretaceaeUnknown
Terminalia catappa (Singapore almond)CombretaceaeWild host
Theobroma cacao (cocoa)MalvaceaeUnknown
Vaccinium reticulatumEricaceaeUnknown
Vangueria infausta (African medlar)RubiaceaeUnknown
Vitellaria paradoxa (shea tree)SapotaceaeOther
Ximenia americana (hog plum)OlacaceaeUnknown
Xylotheca kraussianaFlacourtiaceaeUnknown
Ziziphus mauritiana (jujube)RhamnaceaeMain
Ziziphus mucronataRhamnaceaeUnknown

Growth Stages

Fruiting stage
Post-harvest

Symptoms

Following oviposition there may be some necrosis around the puncture mark ('sting'). This is followed by decomposition of the fruit.

List of Symptoms/Signs

Symptom or signLife stagesSign or diagnosisDisease stage
Plants/Fruit/internal feeding   
Plants/Fruit/lesions: black or brown   
Plants/Fruit/premature drop   

Habitat List

CategorySub categoryHabitatPresenceStatus
TerrestrialTerrestrial – ManagedCultivated / agricultural landPrincipal habitatHarmful (pest or invasive)
TerrestrialTerrestrial – ManagedCultivated / agricultural landPrincipal habitatProductive/non-natural
TerrestrialTerrestrial – ManagedManaged forests, plantations and orchardsPrincipal habitatHarmful (pest or invasive)
TerrestrialTerrestrial – ManagedManaged forests, plantations and orchardsPrincipal habitatProductive/non-natural
TerrestrialTerrestrial ‑ Natural / Semi-naturalNatural forestsPresent, no further detailsNatural
TerrestrialTerrestrial ‑ Natural / Semi-naturalScrub / shrublandsPrincipal habitatHarmful (pest or invasive)
TerrestrialTerrestrial ‑ Natural / Semi-naturalScrub / shrublandsPrincipal habitatNatural

Biology and Ecology

The eggs of B. dorsalis are laid below the skin of the host fruit. These hatch within a day (although this can be delayed up to 5 days in cool conditions) and the larvae feed for another 6-17 days, depending on the temperature (Vargas et al., 1996). Pupariation is in the soil under the host plant for 12 days at 24°C and 60% RH, but may be delayed for up to 26 days under cool conditions (Vargas et al., 1996). The adults occur throughout the year and begin mating after 11 days (Arakaki et al., 1984), and may live for more than 3 months under laboratory conditions (Malod et al., 2020).
[Erratum: In previous versions of this datasheet, it was stated that “many Bactrocera spp. can fly 50-100 km (Fletcher, 1989)” but a review of Fletcher (1989a) and Fletcher (1989b) by Hicks et al. (2019) found no evidence to support this statement and it has been removed. Fletcher (1989b) provides dispersal data for only 11 of 651 species of Bactrocera, many of the case studies lack the necessary numerical data, and the study did not discern between active flight and passive wind-assisted dispersal. There are differences among fruit fly species and further studies are required to determine dispersal distances for individual species. For further information on trapping Bactrocera species to monitor movement, see Weldon et al. (2014).]

Climate

Climate typeDescriptionPreferred or toleratedRemarks
Af - Tropical rainforest climate> 60mm precipitation per monthTolerated 
Am - Tropical monsoon climateTropical monsoon climate ( < 60mm precipitation driest month but > (100 - [total annual precipitation(mm}/25]))Tolerated 
Aw - Tropical wet and dry savanna climate< 60mm precipitation driest month (in winter) and < (100 - [total annual precipitation{mm}/25])Preferred 
Cf - Warm temperate climate, wet all yearWarm average temp. > 10°C, Cold average temp. > 0°C, wet all yearPreferred 
Cw - Warm temperate climate with dry winterWarm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)Preferred 

Latitude/Altitude Ranges

Latitude North (°N)Latitude South (°S)Altitude lower (m)Altitude upper (m)
301500

Air Temperature

ParameterLower limit (°C)Upper limit (°C)
Absolute minimum temperature60
Mean annual temperature2332
Mean maximum temperature of hottest month3035
Mean minimum temperature of coldest month1626

Rainfall

ParameterLower limitUpper limitDescription
Dry season duration36number of consecutive months with <40 mm rainfall
Mean annual rainfall2502620mm; lower/upper limits

Rainfall Regime

Summer
Winter
Uniform

Natural enemy of

This content is currently unavailable.

Notes on Natural Enemies

Bactrocera spp. can be attacked as immature either by parasitoids and predators (invertebrates directly feeding or vertebrates eating fruit). Mortality due to vertebrate fruit consumption can be very high as can puparial mortality in the soil, either due to predation or environmental mortality (see White and Elson-Harris (1994) for brief review). Parasitoids appear to have little effect on the populations of most fruit flies and Fletcher (1987) noted that 0-30% levels of parasitism are typical. To date there are only a few records of partial biological control success for any Bactrocera or Dacus spp. (Wharton, 1989). Clausen (1978) noted that any benefit was almost entirely due to Fopius arisanus (then Opius oophilus) and gave the example of guava fruit (Psidium guajava) attack being reduced from 100 to 22% as a result of reduction in B. dorsalis populations through the effects of parasitism in Hawaii. More recently, F. arisanus introduction to French Polynesia has reduced infestations (larvae/kg) on guava, Tahitian chestnut and tropical almond by 70-95%, but reduction in percentage of infested fruits (by at least one larva) was not reduced as substantially (Leblanc et al., 2013c). A number of parasitoids were also released in Guam against B. dorsalis and this programme was reviewed by Waterhouse (1993). Due to difficulties in verifying the identifications of both parasitoids and (in some cases) the fruit fly hosts, no attempt has been made to catalogue all natural enemy records; see Clausen (1978), White and Elson-Harris (1994), Stibick (2004) and the website on parasitoids of fruit-infesting Tephritidae (http://paroffit.org/public/site/paroffit/home) for major sources of information. Bautista et al. (1998) described the phenology of F. arisanus. Van Mele et al. (2007) showed the positive effects of African weaver ants (Oecophylla longinoda) in controlling fruit flies in relation to the repellent cues of weaver ants (Adandonon et al., 2009), including B. invadens, in mango trees (Mangifera indica). 

Natural enemies

Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Aceratoneuromyia indicaParasite   Guam; Hawaii; Mariana Islandsvegetables
Aganaspis daciParasite   Hawaiivegetables
Austroopius fijiensisParasite   Hawaiivegetables
Fopius persulcatusParasite   Hawaii (presence uncertain) 
Fopius vandenboschiParasite
Arthropods|Larvae
  Guam; Hawaii; Saipanfruits; vegetables
Bracon fletcheriParasite   Hawaiivegetables
Diachasmimorpha albobalteataParasite   Hawaii (not established)vegetables
Diachasmimorpha kraussiiParasite
Larvae
  Hawaiivegetables
Diachasmimorpha tryoniParasite   USA; Hawaiiloquats; peaches
Dirhinus anthracinaParasite
Pupae
    
Dirhinus giffardiiParasite
Pupae
  Hawaii; Saipanfruits
Doryctobracon areolatusParasite   Hawaiivegetables
Oecophylla longinoda (weaver ant)Predator
Adults
Larvae
    
Psyttalia makiiParasite   Hawaiivegetables
Orius insidiosusPredator     
Pachycrepoideus vindemmiaeParasite
Pupae
  Hawaii, Benin 
Paratriphleps laevisculusPredator     
Psyttalia fijiensisParasite   Hawaii (not established)vegetables
Psyttalia incisiParasite
Larvae
  Guam; Hawaii; Mariana Islands; Saipanfruits
Psyttalia walkeriParasite     
Solenopsis geminata (tropical fire ant)Predator     
Spalangia endiusParasite
Pupae
    
Steinernema carpocapsaeParasite     
Tachinaephagus malayensisParasite     
Tetrastichus dacicidaParasite
Larvae
    
Tetrastichus giffardianusParasite
Larvae
  Hawaii 
Thyreocephalus albertisiPredator   Hawaiivegetables
Trybliographa daciParasite
Larvae
    
Tytthus mundulusPredator     
Utetes anastrephaeParasite   Hawaiivegetables
Utetes maniiParasite
Larvae
  Hawaii (not established) 
Fopius arisanusParasite   French Polynesia; Guam; Hawaii; Saipan ; Benin, Botswana, Cameroon, Comoros, Egypt, Kenya, Madagascar, Mauritius, Morocco, Mozambique, Namibia, Reunion, Senegal, Tanzania, Togo, Uganda, Zambia, Zimbabwefruits; loquats; peaches; vegetables
Diachasmimorpha longicaudata (fruit fly parasite, longtailed)Parasite
Larvae
  French Polynesia; Guam; Hawaii; Saipan; Kenya, Mauritius, Egypt, Cape Verde, Madagascar, Morocco, Mozambique, Reunion, Tanzania, Zambiafruits; loquats; peaches; vegetables
Psyttalia phaeostigmaParasite   Hawaiivegetables
Fopius skinneriParasite   Hawaii (not established)vegetables
Psyttalia fletcheriParasite     
Fopius deeralensisParasite   Hawaii (not established)vegetables
Fopius persulcatusParasite   Hawaii (presence uncertain) 

Impact Summary

CategoryImpact
Animal/plant collectionsNone
Animal/plant productsNegative
Biodiversity (generally)None
Crop productionNegative
Economic/livelihoodNegative
Environment (generally)None
Fisheries / aquacultureNone
Forestry productionNone
Human healthNone
Livestock productionNone
Native faunaNegative
Native floraNone
Rare/protected speciesNegative
TourismNone
Trade/international relationsNegative
Transport/travelNone

Impact

B. dorsalis is a very serious pest of a wide variety of fruits and vegetables throughout its range and damage levels can be anything up to 100% of unprotected fruit.

Impact: Economic

B. dorsalis is a very serious pest of a wide variety of fruits and vegetables throughout its range and damage levels can be anything up to 100% of unprotected fruit. As a result of its widespread distribution, pest status, invasive ability and potential impact on market access, B. dorsalis is considered to be a major threat to many countries, requiring costly quarantine restrictions and eradication measures. In Mauritius, the total cost of the eradication operation was approximately US$1 million (Seewooruthun et al., 2000). In Japan, eradication from the Ryukyu Islands has cost more than 200 million euros (Kiritani, 1998). In California, USA it has been estimated that the cost of not eradicating Oriental fruit fly would range from US$ 44 to 176 million in crop losses, additional pesticide use, and quarantine requirements. The cost for the eradication programme in northern Queensland (1995-1999) was AUS$ 33 million, but the estimated annual cost to control the pest, had it been left established, was estimated to be AUS$ 7-8 million (Cantrell et al., 2002). In Hawaii, annual losses in major fruit crops caused by B. dorsalis may exceed 13%, or US$ 3 million (Culliney, 2002).

Impact: Environmental

Due to the competition for food, oriental fruit flies would displace other less aggressive fruit fly species. Duyck et al. (2004) suggested that the r–K gradient could be used as a predictor of the potential invasive capacity of a species. Species with type K-demographic strategy traits, such as species of the genus Bactrocera, would be adapted for competition in saturated habitats. Duyck et al. (2004) reported that in all recorded cases, species further along the r–K gradient, such as B. dorsalis; have invaded over r-selected species, such as Ceratitis capitata, never the reverse.

Impact: Biodiversity

The environmental impact is rated high because the establishment of oriental fruit flies would likely trigger the initiation of chemical and/or biological control programmes. Chemical control would harm native insects and species of conservation significance. 

Impact: Social

Human health and tourism would be affected if plantations treated with insecticides are close to habitat and touristic resorts. However, the risk is very low because local protein bait application techniques (BAT) and male annihilation techniques (MAT) are the most common methods used for the area-wide management of the oriental fruit fly.

Risk and Impact Factors

Invasiveness

Invasive in its native range
Proved invasive outside its native range
Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
Highly mobile locally
Long lived
Fast growing
Has high reproductive potential
Has high genetic variability

Impact outcomes

Host damage
Negatively impacts agriculture
Threat to/ loss of native species

Impact mechanisms

Competition - monopolizing resources
Pest and disease transmission
Interaction with other invasive species

Likelihood of entry/control

Highly likely to be transported internationally accidentally
Highly likely to be transported internationally illegally
Difficult to identify/detect in the field
Difficult/costly to control

Detection and Inspection

Fruits (locally grown or samples of fruit imports) should be inspected for puncture marks and any associated necrosis. Suspect fruits should be cut open and checked for larvae. Larval identification is difficult, so if time allows, mature larvae should be transferred to saw dust (or similar dry medium) to allow pupariation. Upon emergence, adult flies must be fed with sugar and water for several days to allow hardening and full colour to develop, before they can be identified. Detection is described in the Prevention and Control section under Early Warning Systems.

Prevention and Control

Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

Regulatory Control

Systems approach and post-harvest treatments are currently being used in import protocols of fruit commodities regarded as susceptible to B. dorsalis from countries where the pest is present. Post-harvest treatments against B. dorsalis include heat treatments, cold treatments, ionizing radiation and fumigation (Dohino et al., 2017). In systems approach, at least two independent risk mitigation measures which can include pre- and/or post-harvest measures should be used. Pre-harvest control measures which would mitigate the risk of B. dorsalis are provided below. A combination of pre-harvest measures is recommended for effective control of B. dorsalis.

Cultural Control and Sanitary Methods

One of the most effective control techniques against fruit flies in general is to wrap fruit, either in newspaper, a paper bag, or in the case of long/thin fruits, a polythene sleeve. This is a simple physical barrier to oviposition, but it has to be applied well before the fruit is attacked. There is also some evidence that neem seed kernel extract can deter oviposition (Shivendra Singh and Singh, 1998). Early harvesting is also an effective control strategy for mango (Gajendra Singh et al., 1997). Little information is available on the attack time for most fruits but few Bactrocera spp. attack prior to ripening.
Other control and sanitary methods include the removal and destruction of fallen fruits because they may harbour larvae that could form a next generation. Destruction can either be by burning, deep burrowing (at least 0.5 m below the surface), feeding them to pigs, or putting the fruits in dark-coloured plastic bags and placing them in the sun (so that the inside temperature rises and kills the larvae).
Another method is raking or disturbing the soil below the fruit trees using other means. This will expose the puparia, leading to desiccation or predation by other organisms.

Biological control

Classical biological control using the hymenopteran parasitoid, Fopius arisanus, has been successful in Hawaii (Clausen et al., 1965). F. arisanus was also introduced in different African countries and Indian Ocean islands (Mohammed et al., 2016). In Benin, field parasitism rates of B. dorsalis by F. arisanus of up to 21% were recorded when sampling indigenous fruit (Gnanvossou et al., 2016).
Soil application of entomopathogenic fungi in combination with protein bait sprays reduced fruit infestation by B. dorsalis in Kenya (Ekesi et al., 2011).

Chemical Control

Although cover sprays of entire crops are sometimes used, the use of bait sprays is both more economical and more environmentally acceptable. A bait spray consists of a suitable insecticide (e.g. malathion, spinosad, fipronil) mixed with a protein bait. Both males and females of fruit flies are attracted to protein sources emanating ammonia, and so insecticides can be applied to just a few spots in an orchard and the flies will be attracted to these spots. The protein most widely used is hydrolysed protein, but some supplies of this are acid hydrolysed and so highly phytotoxic. Smith and Nannan (1988) have developed a system using autolysed protein. In Malaysia, this has been developed into a very effective commercial product derived from brewery waste.

Sterile Insect Technique

Sterile insect technique was used successfully to eradicate B. dorsalis from Okinawa and neighbouring islands in the Ryukyu Archipelago, Japan (FFEPO, 1987).
Male Suppression

The males of B. dorsalis are attracted to methyl eugenol (4-allyl-1,2-dimethoxybenzene), sometimes in very large numbers. On a small scale, many farmers use male suppression as a control technique; however, with flies attracted over a few hundred metres the traps may be responsible for increasing the fly level (at least of males) on a crop as much as for reducing it. However, the technique has been used as an eradication technique (male annihilation), in combination with bait (Bateman, 1982).

Early Warning Systems

Many countries that are free of Bactrocera spp., e.g. the USA (California and Florida) and New Zealand, maintain a grid of methyl eugenol and cue lure traps, at least in high-risk areas (ports and airports) if not around the entire climatically suitable area. The trap used will usually be modelled on the Steiner trap (White and Elson-Harris, 1994) or Jackson trap.

Field Monitoring

Monitoring is largely carried out by traps baited with methyl eugenol male lure (see Early Warning Systems) set in areas of infestation. However, there is evidence that some fruit flies have different host preferences in different parts of their range and host fruit surveys should also be considered as part of the monitoring process.

Gaps in Knowledge/Research Needs

Host plant surveys have not yet been carried out to show which hosts are of particular importance within the Asian range of true B. dorsalis.
Exploration of Fopius arisanus as a potential classical biological control of B. dorsalis outside Hawaii and French Polynesia is suggested.

Links to Websites

NameURLComment
Dacine Fruit Flies of the Asia-Pacifichttp://www.herbarium.hawaii.edu/fruitfly 
Featured Creatureshttp://entnemdept.ufl.edu/creatures/ 
Hawaii Area-Wide Fruit Fly IPMhttp://www.fruitfly.hawaii.edu/ 
Invasive Fruit Fly Pests in Africahttp://www.africamuseum.be/fruitfly/AfroAsia.htm 
Pacific Fruit Fly Webhttp://www.spc.int/pacifly/ 
Pest Fruit Flies of the Worldhttp://delta-intkey.com/ffa/www/_wintro.htm 
Tephritid Workers Databasehttp://www.tephritid.org/twd/srv/en/home 
True Fruit Flies (Diptera, Tephritidae) of the Afrotropical Regionhttp://projects.bebif.be/fruitfly/index.html 

Organizations

NameAddressCountryURL
IITA (Institut International d'Agriculture Tropicale)BP 08-0932 CotonouBeninhttp://www.iita.org/
CIRAD (Centre de Coopération Internationale en Recherche Agronomique pour le Développment)Head Office, 42, rue Scheffer
75116 Paris
Francehttp://www.cirad.fr
UHM (University of Hawaii at Manoa)College of Tropical Agriculture and Human Resources Department of Plant and Environmental Protecti
Honolulu, HI 96822
USAhttp://www.uhm.hawaii.edu/
USDA-ARSTropical Plant Pests Research Unit,
64 Nowelo Street, Hilo, HI 96720
USAhttp://www.ars.usda.gov/
ICMPFF (International Centre for the Management of Pest Fruit Flies)Griffith UniversityNathan Campus
Queensland 4111
Australiahttp://www.griffith.edu.au/centre/icmpff/

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