Anastrepha obliqua (West Indian fruit fly)

Pictures

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Title Line artwork of adult female Caption Copyright CAB International	Line artwork of adult female		CAB International

Identity

Preferred Scientific Name

• Anastrepha obliqua (Macquart)

Preferred Common Name

• West Indian fruit fly

Other Scientific Names

• Acrotoxa obliqua (Macquart)
• Anastrepha acidusa Greene
• Anastrepha ethalea Greene
• Anastrepha fraterculus var. ligata Costa Lima
• Anastrepha fraterculus var. mombinpraeoptans Seín
• Anastrepha mombinpraeoptans Seín
• Anastrepha trinidadensis Greene
• Tephritis obliqua Macquart
• Trypeta obliqua (Macquart)

International Common Names

• English: Antillean fruit fly; fruit fly, West Indian
• Spanish: mosca de la fruta; mosca de la fruta de las Indias Occidentales
• French: mouche antillaise des fruits

Local Common Names

• Germany: Fruchtfliege, Westindische

EPPO code

• ANSTOB (Anastrepha obliqua)

Summary of Invasiveness

A. obliqua is the most important fruit fly pest of mango (Mangifera indica) in the Neotropics and attacks a broad range of other fruits. It is widespread in Mexico, Central and South America and the West Indies. It is invasive in the Lesser Antilles and was temporarily established in Key West, Florida, USA. It should be considered a serious threat to other tropical parts of the world, particularly mango-producing regions. It is considered an A1 quarantine pest by EPPO.

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Metazoa
•         Phylum: Arthropoda
•             Subphylum: Uniramia
•                 Class: Insecta
•                     Order: Diptera
•                         Family: Tephritidae
•                             Genus: Anastrepha
•                                 Species: Anastrepha obliqua

Notes on Taxonomy and Nomenclature

This species was first described by Macquart (1835) as Tephritis obliqua, although that name was confused and not recognized as pertaining to this species for many years. The species was also described as Anastrepha fraterculus var. mombinpraeoptans Seín, Anastrepha fraterculus var. ligata Lima, and Anastrepha trinidadensis Greene. Most records of Anastrepha acidusa (Walker) are misidentifications of this species. The current combination was proposed by Schiner in 1868.

Description

For a general description of the genus, see the datasheet on Anastrepha.

Adult

As in most other Anastrepha spp., the adults of A. obliqua are easily separated from those of other tephritid genera by a simple wing venation character; vein M, the vein that reaches the wing margin just behind the wing apex, curves forwards before joining the wing margin. Furthermore, most Anastrepha spp. have a very characteristic wing pattern; the apical half of the wing has two inverted 'V'-shaped markings, one fitting within the other; and a stripe along the forward edge of the wing, which runs from near the wing base to about half-way along the wing length.

Identification to species is more difficult. In particular, for positive identification it is essential to dissect the aculeus (the distal, piercing part of the ovipositor that is normally retracted into the oviscape) of a female specimen. A. obliqua adults are difficult to separate from those of Anastrepha fraterculus, Anastrepha sororcula, Anastrepha zenildae, Anastrepha turpiniae, Anastrepha suspensa and several other species of the fraterculus group; if necessary, specimens should be referred to a specialist.

The body is predominantly yellow to orange-brown, and the setae are red-brown to dark-brown.

Head: yellow except ocellar tubercle brown. Facial carina, in profile, concave. Frons with three or more frontal setae, two orbital setae. Antenna not extended to ventral facial margin.

Thorax: mostly yellow to orange-brown, with the following areas yellow to white and often contrasting: postpronotal lobe; single medial and paired sublateral vittae on scutum, the slender medial vitta extended nearly the full length of the scutum, slightly broadened posteriorly, ovoid; sublateral vitta extended from transverse suture almost to posterior margin, including intra-alar seta; scutellum; propleuron; dorsal margin of anepisternum; dorsal margin of katepisternum; katepimeron; and most of anatergite and katatergite. Area bordering scutoscutellar suture medially without dark-brown spot. Subscutellum without dark markings; mediotergite usually dark-brown laterally. Scutum entirely microtrichose or at most with small presutural, medial bare area.

Wing: 5.7-7.5 mm long. Vein M strongly curved apically. Vein R₂₊₃ nearly straight. Pattern mostly orange-brown and moderate-brown. C-band and S-band usually connected along vein R₄₊₅, but sometimes separated; marginal hyaline spot (or end of band) present in cell r₁ at apex of vein R₄₊₅. S-band with middle section between costa and vein Cu₁ largely yellow to orange with narrow brown margins, darkening distally; distal section of band moderately broad, well-separated from apex of vein M. V-band with distal arm complete and connected to proximal arm; proximal arm extended to vein R₄₊₅, sometimes connected to S-band.

Abdomen: tergites yellow to orange-brown, without dark-brown markings.

Male terminalia: lateral surstylus moderately long, in posterior view slightly tapered, somewhat truncate apically. Phallus 2.3–2.7 mm long; ratio to mesonotum length 0.8-0.9. Glans with basolateral membranous lobe, mostly membranous medially, with isolated, T-shaped apical sclerite.

Female terminalia: oviscape straight, 1.6-1.9 mm long; ratio to mesonotum length 0.53-0.61 mm. Dorsobasal scales of eversible membrane numerous, hook-like, in triangular pattern. Aculeus length 1.30-1.65 mm; tip 0.16-0.21 mm long, 0.08-0.12 mm wide, gradually tapering, but with slight constriction proximal to serrate part, distal 0.67-0.82 mm serrate. Three spermathecae ovoid.

Immature Stages

Larva: it is very difficult, and in some cases impossible to identify larvae of Anastrepha species from morphological characteristics. The key by Steck et al. (1990) and the interactive key by Carroll et al. (2004) are the best tools for larval identification. Descriptions of A. obliqua larvae are provided by Berg (1979), Steck et al. (1990) and White and Elson-Harris (1994). White and Elson-Harris (1994) described the third-instar larva as follows:

Larvae: medium-sized; 7.5-9.0 mm long; 1.4-1.8 mm wide.

Head: stomal sensory organ rounded, only slightly protuberant, with two to three very small sensilla; 7-10 oral ridges; accessory plates small; mandible moderately to heavily sclerotised, with a large slender curved apical tooth.

Thoracic and abdominal segments: T1 with a broad anterior band of 5-10 discontinuous rows of small, sharply pointed spinules; T2 and T3 with two to five rows of spinules. Dorsal spinules absent from A1-A8. Creeping welts on A1-A8 with 7-11 rows of stout spinules. A8 with large dorsal tubercles and stout sensilla; intermediate areas well-developed with obvious sensilla; ventral sensilla small, but well defined.

Anterior spiracles: with 12-16 tubules.

Posterior spiracles: spiracular slits about three times as long as broad, with heavily sclerotised, dark-brown rimae. Spiracular hairs in dorsal and ventral bundles of 10-16 stout hairs branched in apical third; lateral bundles of three to six hairs similarly branched.

Anal area: lobes very large, protuberant, not grooved; surrounded by two to five discontinuous rows of small, sharp spinules.

Egg: the eggs of A. obliqua bear a conspicuous lobe on the anterior (micropyle) end, which projects outside the fruit peel and is believed to aid in respiration (Murillo and Jiron, 1994). This lobe is lacking in related species, thus eggs inside the abdomens of gravid females can provide a useful diagnostic character for this species.

Distribution

See also CABI/EPPO (2011) and CABI/EPPO (1988, No. 90).

Distribution Table

The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

History of Introduction and Spread

A. obliqua is known from Mexico south to northern Argentina, and from most of the West Indies. Its original native range is obscure. It is the only species of Anastrepha widespread in both the West Indies and the mainland. Although commonly known as the West Indian fruit fly, the original range of A. obliqua in the Antilles is uncertain, and in at least some of the Lesser Antilles it is an invasive species. It was originally described from Cuba and at least has long been present in the Greater Antilles. Its current distribution on the mainland and the Greater Antilles is presumed to be natural, but could have been affected by human activities.

Within the Lesser Antilles, Kisliuk and Cooley (1933) reported it (as Anastrepha acidusa) from Dominica, Martinique, Nevis, St. Kitts, St. Lucia, Trinidad and the Virgin Islands. They reported Anastrepha larvae (presumably of A. obliqua), but did not rear adults from Guadeloupe. They did not find it in Antigua, the Bahamas, Barbados, and St. Vincent. A survey from 1988-1990 confirmed its presence in Dominica and St. Lucia (EC Ambrose, Inter-American Institute for Cooperation on Agriculture [IICA], St. Lucia, unpublished data). It has been present in Barbados since 2001 (I Gibbs, Ministry of Agriculture and Rural Development, Barbados, personal communication, 2004), in Grenada since 2002 (T Peters, Ministry of Agriculture, Botanic Gardens, St. George's, Grenada, personal communication, 2004), and in St. Vincent since 2002 (M Delpeche, Ministry of Agriculture, Richmond Hill, Kingstown, St. Vincent, West Indies, personal communication, 2004). White and Elson-Harris (1992) reported it from the Bahamas, but the basis for that record is unclear and it is probably erroneous. A. obliqua has not been collected in extensive trapping in the last decade (S Miller, Department of Agriculture, Nassau, Bahamas, personal communication, 2004). Reports of its introduction to Bermuda were erroneous (Woodley and Hilburn, 1994). It was established in southern Florida, USA (Key West) from 1930-1936 (McAlister, 1936), but there is no evidence of a breeding population being present since then (Steck, 2001). It has been trapped in California and Texas, USA, but is not established there.

Introductions

Risk of Introduction

EPPO lists A. obliqua as an A1 quarantine pest (OEPP/EPPO, 1983) within the broad category 'non-European Trypetidae'; it is also of quarantine significance to APPPC, CPPC and NAPPO.

Consignments of fruits of Citrus spp., mango [Mangifera indica] and guava [Psidium guajava] from countries where A. obliqua occurs should be inspected for symptoms of infestation. Those suspected should be cut open in order to look for larvae. For example, EPPO recommends (OEPP/EPPO, 1990) that such fruits should come from an area where A. obliqua does not occur or from a place of production found free from the pest by regular inspection for 3 months prior to harvest. Fruits may also be treated in transit by cold treatment (for example, 13, 15 or 17 days at 0.5, 1 or 1.5°C, respectively) or, for certain types of fruits, by vapour heat (for example, keeping at 43°C for 4-6 h) (USDA, 1994), or by hot water immersion (Nascimento et al., 1992; Thomas and Mangan, 1995) or forced hot air quarantine treatment (Mangan and Ingle, 1992).

Plants of host species transported with roots from countries where A. obliqua occurs should be free from soil, or the soil should be treated against puparia, and should not carry fruits. Importation of such plants may be prohibited.

Habitat List

Hosts/Species Affected

The main native hosts are Spondias spp. (Anacardiaceae), but these are only of local interest. Mangoes [Mangifera indica], also Anacardiaceae, are the economically important host, on which the species has extended its range (Hernandez-Ortiz, 1992). Citrus spp. and guavas [Psidium guajava] are only occasional hosts. Like other Anastrepha spp., A. obliqua has been recorded incidentally on a wider range of fruits, both tropical and temperate, but these records are incidental occurrences, of no economic significance.

In common with other polyphagous and difficult to identify species, many host records cannot be substantiated and only records confirmed by Norrbom and Kim (1988) or subsequent reliable sources have been accepted here. Post 1988 records include Eugenia stipitata (Couturier et al., 1996).

Host Plants/Plants Affected

Growth Stages

Symptoms

Attacked fruit can show signs of oviposition punctures, but these, or any other symptoms of damage, are often difficult to detect in the early stages of infestation. Much damage may occur inside the fruit before external symptoms are seen, often as networks of tunnels accompanied by rotting. Very sweet fruits may produce a sugary exudate.

Symptoms List

Biology and Ecology

The eggs are laid singly, below the skin of the host fruit. The larvae hatch within 3-12 days and feed for another 15-32 days. Pupariation is in the soil under the host plant and the adults emerge after 15-19 days (longer in cool conditions); the adults occur throughout the year (Christenson and Foote, 1960), with little seasonal variation (Hedstrom, 1993).

Climate

Latitude/Altitude

Natural Enemies

Notes on Natural Enemies

Several braconid larval parasitoids are recorded but their impact appears low, for example, Biosteres longicaudatus in 2.7% of larvae (Borge and Basedow, 1997). For further information, see Ohashi et al. (1997).

Means of Movement and Dispersal

There is evidence that the adults of Anastrepha spp. can fly as far as 135 km (Fletcher, 1989) and therefore natural movement is an important means of spread.

In international trade, the major means of dispersal to previously uninfested areas is the transport of fruits containing live larvae. For the EPPO region, the most important fruits liable to carry A. obliqua are mango (Mangifera indica), and to a lesser extent Citrus spp. and guava (Psidium guajava). A. obliqua has been intercepted in France on mangoes from Mexico. The various tropical fruit hosts that may be locally important in America are infrequently traded to Europe. There is also a risk from the transport of puparia in soil or packaging with plants that have already fruited.

Pathway Causes

Pathway Vectors

Plant Trade

Impact Summary

Impact

Anastrepha spp. are the most serious fruit fly pests in the tropical Americas (Norrbom and Foote, 1989), with the possible exception of the introduced Ceratitis capitata (EPPO/CABI, 1996). A. obliqua is recorded from Citrus spp., but they are not important hosts (Enkerlin et al., 1989). It mainly attacks mangoes (Mangifera indica) and other Anacardiaceae (Whervin, 1974).

Risk and Impact Factors

Impact outcomes

• Host damage
• Negatively impacts agriculture
• Negatively impacts livelihoods

Invasiveness

• Abundant in its native range
• Capable of securing and ingesting a wide range of food
• Has a broad native range
• Has high genetic variability
• Has high reproductive potential
• Is a habitat generalist
• Proved invasive outside its native range
• Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc

Likelihood of entry/control

• Difficult to identify/detect as a commodity contaminant
• Difficult to identify/detect in the field
• Difficult/costly to control
• Highly likely to be transported internationally accidentally
• Highly likely to be transported internationally illegally

Detection and Inspection

No male lures have yet been identified for Anastrepha spp. However, they are captured by traps emitting ammonia and it is likely that traps already set for Rhagoletis cerasi in the cherry-growing areas of the EPPO region may attract Anastrepha spp. if they should ever occur in those areas. McPhail traps are usually used for the capture of Anastrepha spp. (Drew, 1982) and possible baits are ammonium acetate (Hedstrom and Jimenez, 1988), casein hydrolysate (Sharp, 1987) and torula yeast (Hedstrom and Jiron, 1985). The number of traps required per unit area is high; in a release and recapture test, Calkins et al. (1984) placed 18 traps per 0.4 ha and only recovered about 13% of the released flies.

Some studies have shown that egg morphology can be used to separate closely related species found in host fruits (Murillo and Jiron, 1994). The larvae of some species may also be differentiated using cuticular hydrocarbons (Sutton and Carlson, 1993). Neither method has yet been generalized for application outside of very specific circumstances.

Similarities to Other Species/Conditions

The adults of A. obliqua are difficult to separate from those of various other species of the fraterculus group, such as the A. fraterculus complex, Anastrepha sororcula, Anastrepha zenildae, Anastrepha turpiniae and Anastrepha suspensa. The females can be distinguished by the dimensions and shape of the aculeus, particularly its tip, which is two-thirds to three-fourths serrate. The adults lack the dark-brown spot on the scutoscutellar suture and lateral dark-brown mark on the subscutellum that are usually present in the above species.

The larvae of Anastrepha are extremely difficult to identify and specialist help should be sought to confirm critical identifications. The third-instar larvae are very similar to those of A. suspensa and the A. fraterculus complex, and these species usually cannot be distinguished (Steck et al., 1990).

Prevention and Control

Control can be considerably aided by good cultural practices, for example, by gathering all fallen and infected host fruits and destroying them, and by the selection of suitable varieties (Jiron, 1996). Insecticidal protection is possible by using a cover spray or a bait spray. Malathion is the usual choice of insecticide for fruit fly control and this is usually combined with protein hydrolysate to form a bait spray (Roessler, 1989). Practical details are given by Bateman (1982). Bait sprays work on the principle that both male and female tephritids are strongly attracted to a protein source from which ammonia emanates. Bait sprays have the advantage over cover sprays that they can be applied as a spot treatment so that the flies are attracted to the insecticide and there is minimal impact on natural enemies. The use of natural plant substances, such as a leaf infusion of Piper auritum combined with gathering fallen fruit resulted in local eradication (Perales-Segovia et al., 1996).

Many baits have been evaluated including those based on yeasts (Fragenas et al., 1996), borated hydrolysed protein and yeast (Jiron and Soto-Manitiu, 1989), human urine (Hedstrom, 1988) and molasses (Hedstrom and Jiron, 1985). The longevity of some baits were compared by Malo (1992).

Phytosanitary Measures

Consignments of fruits of Citrus spp., mango [Mangifera indica] and guava [Psidium guajava] from countries where A. obliqua occurs should be inspected for symptoms of infestation. Those suspected should be cut open in order to look for larvae. For example, EPPO recommends (OEPP/EPPO, 1990) that such fruits should come from an area where A. obliqua does not occur or from a place of production found free from the pest by regular inspection for 3 months prior to harvest. Fruits may also be treated in transit by cold treatment (for example, 13, 15 or 17 days at 0.5, 1 or 1.5°C, respectively) or, for certain types of fruits, by vapour heat (for example, keeping at 43°C for 4-6 h) (USDA, 1994), or by hot water immersion (Nascimento et al., 1992; Thomas and Mangan, 1995) or forced hot air quarantine treatment (Mangan and Ingle, 1992).

Ethylene dibromide was previously widely used as a fumigant, but is now generally withdrawn because of its carcinogenicity.

Plants of host species transported with roots from countries where A. obliqua occurs should be free from soil, or the soil should be treated against puparia, and should not carry fruits. Importation of such plants may be prohibited.

References

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Links to Websites

Contributors

27/02/2008 Updated by:

Allen Norrbom, Systematic Entomology Laboratory, USDA, c/o National Museum of Natural History, MRC 168, PO Box 37012, Washington, DC 20013-7012, USA

Distribution Maps

• = Present, no further details
• = Evidence of pathogen
• = Widespread
• = Last reported
• = Localised
• = Presence unconfirmed
• = Confined and subject to quarantine
• = See regional map for distribution within the country
• = Occasional or few reports

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