Anastrepha ludens (Mexican fruit fly)

Pictures

Picture	Title	Caption	Copyright
Title Adults Caption Adult females ovipositing on citrus fruit skin. Copyright USDA-ARS	Adults	Adult females ovipositing on citrus fruit skin.	USDA-ARS
Title Line artwork of adult female Caption Copyright CAB International	Line artwork of adult female		CAB International

Identity

Preferred Scientific Name

• Anastrepha ludens (Loew)

Preferred Common Name

• Mexican fruit fly

Other Scientific Names

• Acrotoxa ludens (Loew)
• Anastrepha lathana Stone
• Trypeta ludens Loew

International Common Names

• English: fruitfly, Mexican
• Spanish: gusano de la fruta; gusano de la naranja; mosca mexicana de la fruta
• French: mouche mexicaine des fruits

Local Common Names

• Germany: Fruchtfliege, Mexikanische
• Italy: anastrefa; mosca de la fruta

EPPO code

• ANSTLU (Anastrepha ludens)

Summary of Invasiveness

A. ludens has a broad host range and is a major pest, especially of citrus and mango (Mangifera indica) in most parts of its range. This species and Anastrepha obliqua are the most important pest species of Anastrepha in Central America and Mexico. It occurs in subtropical areas as far north as southern Texas, thus it may be more of a threat of introduction to other subtropical areas of the world than other species of Anastrepha. It is invasive at least in Panama and has been trapped in California, USA. 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 ludens

Notes on Taxonomy and Nomenclature

This species was first described in 1873 by Loew as Trypeta ludens. The current combination was proposed by Wulp (1900). The name Anastrepha lathana is recognized as a synonym. A record for presence in Colombia was based on misidentification of Anastrepha manizaliensis (Norrbom et al., 2005).

Description

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

Adult

As in most other Anastrepha spp., the adults of A. ludens 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 that runs from near the wing base to about half-way along the wing length.

Identification to species is more difficult. In particular, it is essential to dissect the aculeus (ovipositor piercer) of a female specimen for positive identification. The adults of A. ludens are unlikely to be confused with those of any of the other species of Anastrepha occurring within its range, except perhaps Anastrepha distincta, which has considerably shorter male and female terminalia. Several species from South America, such as Anastrepha manizaliensis and Anastrepha schultzi are more difficult to distinguish from A. ludens (Norrbom et al., 2005). For a positive identification, the females should be dissected to check the aculeus dimensions and shape carefully.

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 usually with irregular dark-brown spot. Subscutellum dark-brown laterally; brown mark often extending onto lateral part of mediotergite. Scutum entirely microtrichose or at most with small presutural, medial bare area.

Wing: 7-9 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; distal section of band moderately broad, well-separated from apex of vein M. V-band with distal arm usually complete and connected to proximal arm, although often weaker anteriorly; proximal arm extended to vein R₄₊₅, not 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 5.2-6.1 mm long; ratio to mesonotum length 1.51-1.84. Glans with basolateral membranous lobe, mostly membranous medially, with isolated, T-shaped apical sclerite.

Female terminalia: oviscape straight, 3.4-6.3 mm long; ratio to mesonotum length 1.10-1.55. Dorsobasal scales of eversible membrane numerous, hook-like, in triangular pattern. Aculeus length 3.35-5.75 mm; tip 0.32-0.40 mm long, 0.12-0.14 mm wide, gradually tapering, but with slight medial constriction, distal half or less serrate. Three spermathecae ovoid.

Larva: the larvae of Anastrepha are extremely difficult to identify, but the third-instar larvae of A. ludens can usually be distinguished from those of other species of the genus using the key by Steck et al. (1990) or the interactive key by Carroll et al. (2004). Carroll and Wharton (1989) provided a detailed description of the egg, larva (all three instars) and puparium. Berg (1979), Heppner (1984), and White and Elson-Harris (1992) also provided descriptions of the third-instar.

The following diagnostic description of the third-instar is based on Carroll and Wharton (1989) and White and Elson-Harris (1992).

Larvae: medium-large, 5.8-11.1 mm long and 1.2-2.5 mm wide.

Head: stomal sensory organ large, rounded, with five small sensilla; 11-17 oral ridges with margins entire or slightly undulant; accessory plates small; mandible moderately sclerotised, with a large slender curved apical tooth.

Thoracic and abdominal segments: T1-T3 with spinulose areas on anterior margins; mid-dorsally with four to six, three to five and one to two rows of spinules, respectively. Dorsal spinules absent from A1-A8. Creeping welt on A1 with seven to nine rows of spinules, those on A2-A8 with 9-17 rows. A8 with intermediate lobes moderately developed; tubercles and sensilla small, but obvious.

Anterior spiracles: with 12-21 tubules.

Posterior spiracles: spiracular slits approximately 3.5 times as long as broad, with moderately sclerotised rimae. Spiracular hairs short (about one-third to one-fifth the length of the spiracular slit), often branched in the apical third; dorsal and ventral bundles of 6-13 hairs, lateral bundles of four to seven hairs.

Anal area: lobes large, protuberant, usually distinctly bifid; surrounded by three to four discontinuous rows of small spinules.

Egg: 1.37-1.60 mm long, greatest width 0.18-0.21 mm. White, spindle-shaped, broad anteriorly, tapering posteriorly; micropyle slightly to one side of apex of anterior pole; faint reticulation near micropyle consisting primarily of irregular pentagons and hexagons, these becoming very faint and elongated in the posterior portion of the egg; distinct openings into chorion at vertices of polygons in anterior end (from Carroll and Wharton, 1989).

Distribution

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.

Risk of Introduction

Consignments of fruits of Citrus spp., Malus spp., mango (Mangifera indica) and guava (Psidium guajava) from countries where the pest occurs should be inspected for symptoms of infestation and 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. ludens does not occur or from a place of production found free from the pest by regular inspection for 3 months before harvest.

Fruits may also be treated in transit by cold treatment (for example, 18, 20 or 22 days at 0.5, 1.0 or 1.5°C, respectively) or, for certain types of fruits, by vapour heat (for example maintaining at 43°C for 4-6 h) (USDA, 1994), or forced hot-air treatment (Mangan and Ingle, 1994). Shellie et al. (1997) found that heating in a controlled atmosphere was more effective. Hot-water immersion has also been tested and found to be inadequate (Thomas and Mangan, 1995).

Ethylene dibromide was previously widely used as a fumigant, but is now generally withdrawn because of its carcinogenicity. The use of fruit coating has also been investigated as a means of killing the larvae (Hallman, 1997).

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

Habitat

A. ludens may be found in fruit-growing areas with suitable hosts and in natural forests.

Habitat List

Hosts/Species Affected

Baker et al. (1944) considered Casimiroa greggii (Rutaceae) to be the only native wild host of A. ludens, although Casimiroa edulis may also have been an original native host (Jirón et al., 1988). Citrus spp. and mango (Mangifera indica) are the most important introduced hosts (Hernandez-Ortiz, 1992). Myrtaceae (e.g. guavas, Psidium guajava), Rosaceae (e.g. peaches, Prunus persica) and a variety of other fruits are occasional hosts (Norrbom, 2004a).  

Host Plants/Plants Affected

Growth Stages

Symptoms

Symptoms List

Biology and Ecology

As in many Anastrepha spp., generally, the eggs are laid below the skin of the host fruit in clutches of 1-23 eggs. They hatch within 6-12 days and the larvae feed for another 15-32 days at 25°C. 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).

Climate

Latitude/Altitude

Natural Enemies

Notes on Natural Enemies

Aluja et al. (1990) found that Biosteres longicaudatus was the major parasitoid with up to 29% parasitism of Anastrepha spp., including A. ludens. In another area of Mexico, Gonzalez-Hernandez and Tejada (1979) found that Doryctobracon crawfordi was the most abundant parasitoid from 1954-1959. Thomas (1993) found that Peromyscus spp. could destroy up to 34% of puparia. Other predators were also reported by Thomas (1995), for example, ants and staphylinid beetles, which attacked larvae (up to 5%) on the ground while they were hunting for pupariation sites.

Spiders may be important natural regulators of numbers and Lyssomanes pescadero has been regarded as a potential biocontrol agent of A. ludens (Jimenez and Tejas, 1996).

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 fruit containing live larvae. The most important fruits liable to carry A. ludens are Citrus spp. and mango (Mangifera indica), and to a lesser extent peaches (Prunus persica) and guava (Psidium guajava). 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 (CABI/EPPO, 1998). A. ludens is mainly important on Citrus spp. and mangoes [Mangifera indica]. It is the most abundant fruit fly in some areas of Guatemala (Eskafi, 1988) and Mexico (Malo et al., 1987).

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
• Highly adaptable to different environments
• 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

Diagnosis

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.

Trap shape and design is important. Epsky et al. (1995) and Heath et al. (1995) described dry traps for use with synthetic lures. Robacker (1992) tested spheres and rectangles (vertical and horizontal) and found that the most efficient trap shapes and colours varied between seasons. Blanco-Montero and Sanchez-Salas (1990) showed that the traditional McPhail trap was more effective than yellow circular or rectangular traps.

Similarities to Other Species/Conditions

The adults of A. ludens are unlikely to be confused with those of any other species of Anastrepha occurring within its range, except perhaps Anastrepha distincta, which has considerably shorter male and female terminalia. Several species from South America, such as Anastrepha manizaliensis and Anastrepha schultzi are more difficult to distinguish from A. ludens (Norrbom et al., 2005). For a positive identification, the females should be dissected to carefully check the aculeus dimensions and shape.

The larvae of Anastrepha are extremely difficult to identify and specialist help should be sought to confirm critical identifications. The third stage larva of A. ludens can usually be distinguished using the key by Steck et al. (1990) or the interactive key by Carroll et al. (2004).

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. 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) and Silva-Contreras (1978) give an example specific to A. ludens.

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. Many different attractants have been developed including fermented corn extract (Lee et al., 1997), host-fruit volatiles (Robacker and Heath, 1996), pheromones (Landolt and Heath, 1996), Staphylococcus aureus odour (Robacker and Flath, 1995) and corn hydrolysate (Heath et al., 1994).

Biological Control

Control of A. ludens using Bacillus thuringiensis has been tested in the laboratory (Martinez et al., 1997) and found to cause up to 90% adult mortality.

Biological control has been tried against A. ludens, but introduced parasitoids have had little impact (Wharton, 1989). Sterile insect release has been tested against A. ludens (Gilmore, 1989) and although no major eradication programme has been carried out, sterile flies are used as part of a programme to keep a fly free zone in southern Texas, USA (Mangan, 1996).

Parasitoids of the mediterranean and oriental fruit flies were imported from Hawaii, USA, in 1954-59, but only Biosteres longicaudatus and Aceratoneuromyia indica became established. It was claimed that A. indica accounted for up to 80% parasitism (Clausen, 1978).

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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