Invasive Species Compendium

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Datasheet

Anastrepha ludens
(Mexican fruit fly)

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Datasheet

Anastrepha ludens (Mexican fruit fly)

Summary

  • Last modified
  • 10 December 2020
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Natural Enemy
  • Preferred Scientific Name
  • Anastrepha ludens
  • Preferred Common Name
  • Mexican fruit fly
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Uniramia
  •         Class: Insecta
  • 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 i...

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Pictures

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PictureTitleCaptionCopyright
Adult females ovipositing on citrus fruit skin.
TitleAdults
CaptionAdult females ovipositing on citrus fruit skin.
CopyrightUSDA-ARS
Adult females ovipositing on citrus fruit skin.
AdultsAdult females ovipositing on citrus fruit skin.USDA-ARS
TitleLine artwork of adult female
Caption
CopyrightCAB International
Line artwork of adult femaleCAB International

Identity

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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

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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

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

Notes on Taxonomy and Nomenclature

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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

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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

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Baker et al. (1944) considered this species to be native only to northeastern Mexico, although Jirón et al. (1988) suggested it also may have been native to Nicaragua and Costa Rica and that further evaluation of the status of the original southern populations was needed.

A. ludens was rare in Costa Rica and was not a pest of citrus prior to the mid-1990s, when it suddenly became common in the central highlands and was found attacking orange (Citrus spp.) and grapefruit (Citrus paradisi), including at a research station where Jirón et al. (1988) had worked extensively and had not found it. It has subsequently been detected attacking citrus at high elevations in western Panama. This information suggests that a new population was introduced to Costa Rica from further north and has spread into Panama.

The record of this species from Colombia (Núñez Bueno, 1981) was based on misidentification of Anastrepha manizaliensis (Norrbom et al., 2005). There are no valid reports of A. ludens from Colombia.

See also CABI/EPPO (2001, Map no. 89).


Distribution Table

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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.

Last updated: 17 Feb 2021
Continent/Country/Region Distribution Last Reported Origin First Reported Invasive Reference Notes

Europe

SloveniaAbsent

North America

BelizePresentNative
Costa RicaPresent, LocalizedHigher elevations. More common and widely distributed since the 1990s
El SalvadorPresentNative
GuatemalaPresentNative
HondurasPresentNative
MexicoPresent, Widespread
NicaraguaPresentNative
PanamaPresent, LocalizedIntroducedInvasiveHigher elevations, western Panama; First reported: <1996
United StatesPresent, Transient under eradication
-ArizonaAbsent, Formerly present
-CaliforniaPresent, Transient under eradication
-FloridaAbsent, Intercepted onlyRarely trapped, not established
-TexasPresent, Transient under eradication

Oceania

New ZealandAbsent, Confirmed absent by survey

South America

ArgentinaAbsent, Invalid presence record(s)
ColombiaAbsent, Invalid presence record(s)

Risk of Introduction

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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

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

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial ManagedCultivated / agricultural land Present, no further details Harmful (pest or invasive)
Terrestrial ManagedManaged forests, plantations and orchards Present, no further details Harmful (pest or invasive)
Terrestrial ManagedDisturbed areas Present, no further details
Terrestrial ManagedUrban / peri-urban areas Present, no further details Harmful (pest or invasive)
Terrestrial Natural / Semi-naturalNatural forests Present, no further details Natural

Hosts/Species Affected

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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 and Other Plants Affected

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Plant nameFamilyContextReferences
Anacardium occidentale (cashew nut)AnacardiaceaeOther
AnnonaAnnonaceaeOther
Annona cherimola (cherimoya)AnnonaceaeOther
    Annona liebmanniana (hardshell custard-apple)AnnonaceaeOther
    Annona muricata (soursop)AnnonaceaeOther
      Annona reticulata (bullock's heart)AnnonaceaeOther
      Annona squamosa (sugar apple)AnnonaceaeOther
      Capsicum pubescens (rocoto)SolanaceaeUnknown
      Carica papaya (pawpaw)CaricaceaeOther
      Casimiroa edulis (white sapote)RutaceaeMain
      Casimiroa greggiiUnknown
      Casimiroa pubescensRutaceaeOther
      CitrusRutaceaeMain
      Citrus aurantiifolia (lime)RutaceaeOther
      Citrus aurantium (sour orange)RutaceaeMain
      Citrus limetta (sweet lemon tree)RutaceaeOther
      Citrus maxima (pummelo)RutaceaeMain
      Citrus medica (citron)RutaceaeOther
      Citrus reshni (Cleopatra mandarin)RutaceaeUnknown
      Citrus reticulata (mandarin)RutaceaeOther
      Citrus reticulata x paradisi (tangelo)RutaceaeOther
        Citrus sinensis (navel orange)RutaceaeMain
        Citrus x paradisi (grapefruit)RutaceaeMain
        Coffea arabica (arabica coffee)RubiaceaeOther
        Diospyros ebenaster (black sapote)EbenaceaeUnknown
        Diospyros kaki (persimmon)EbenaceaeOther
        Ficus carica (common fig)MoraceaeUnknown
        fruitsOther
          Heteromeles salicifolia (toyon)RosaceaeUnknown
          IngaFabaceaeOther
            Inga michelianaUnknown
            Malus domestica (apple)RosaceaeOther
            MammeaClusiaceaeUnknown
            Mammea americana (mamey apple)ClusiaceaeOther
            Mangifera indica (mango)AnacardiaceaeMain
            Morus (mulberrytree)MoraceaeUnknown
            Mosiera longipesMyrtaceaeUnknown
            Passiflora edulis (passionfruit)PassifloraceaeOther
            Persea americana (avocado)LauraceaeOther
              Pouteria campechiana (canistel)SapotaceaeUnknown
              Prunus persica (peach)RosaceaeMain
              Psidium cattleianum (strawberry guava)MyrtaceaeOther
                Psidium guajava (guava)MyrtaceaeOther
                Punica granatum (pomegranate)PunicaceaeOther
                Pyrus communis (European pear)RosaceaeOther
                Sargentia greggiiRutaceaeUnknown
                Sideroxylon capiri subsp. tempisqueUnknown
                Spondias purpurea (red mombin)AnacardiaceaeOther
                Syzygium jambos (rose apple)MyrtaceaeOther
                Talisia olivaeformisSapindaceaeOther
                  Terminalia catappa (Singapore almond)CombretaceaeUnknown

                  Growth Stages

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                  Fruiting stage, Post-harvest

                  Symptoms

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                  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.

                  List of Symptoms/Signs

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                  SignLife StagesType
                  Fruit / internal feeding

                  Biology and Ecology

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                  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

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                  ClimateStatusDescriptionRemark
                  A - Tropical/Megathermal climate Preferred Average temp. of coolest month > 18°C, > 1500mm precipitation annually
                  Af - Tropical rainforest climate Preferred > 60mm precipitation per month
                  Am - Tropical monsoon climate Preferred Tropical monsoon climate ( < 60mm precipitation driest month but > (100 - [total annual precipitation(mm}/25]))
                  As - Tropical savanna climate with dry summer Preferred < 60mm precipitation driest month (in summer) and < (100 - [total annual precipitation{mm}/25])
                  Aw - Tropical wet and dry savanna climate Preferred < 60mm precipitation driest month (in winter) and < (100 - [total annual precipitation{mm}/25])
                  Cs - Warm temperate climate with dry summer Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers
                  Cw - Warm temperate climate with dry winter Preferred Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)

                  Latitude/Altitude Ranges

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                  Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
                  27 8

                  Natural enemies

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                  Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
                  Aceratoneuromyia indica Parasite Larvae Belize; Mexico Citrus; grapefruits; oranges
                  Biosteres arisanus Parasite Mexico Citrus
                  Biosteres longicaudatus Parasite Larvae Belize; Mexico Citrus; grapefruits; oranges
                  Biosteres vandenboschi Parasite Mexico Citrus
                  Diachasmimorpha tryoni Parasite Mexico Citrus
                  Dirhinus giffardii Parasite Mexico Citrus
                  Doryctobracon crawfordi Parasite Larvae
                  Lyssomanes pescadero Predator
                  Opius incisi Parasite Mexico Citrus
                  Pachycrepoideus vindemmiae Parasite Pupae Belize Citrus; grapefruits; oranges
                  Peromyscus boylii Predator
                  Peromyscus leucopus Predator
                  Solenopsis geminata Predator
                  Trybliographa daci Parasite Mexico Citrus

                  Notes on Natural Enemies

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                  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

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                  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

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                  CauseNotesLong DistanceLocalReferences
                  Crop production Yes Yes

                  Pathway Vectors

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                  VectorNotesLong DistanceLocalReferences
                  AircraftImmatures in fruit Yes Yes
                  Containers and packaging - woodOf fruit cargo. Yes
                  LuggageImmatures in fruit Yes Yes
                  MailFruit in post. Yes
                  Plants or parts of plantsImmatures in fruit Yes Yes
                  Soil, sand and gravelRisk of puparia in soil. Yes

                  Plant Trade

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                  Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
                  Fruits (inc. pods) eggs; larvae; pupae Yes Pest or symptoms usually visible to the naked eye
                  Growing medium accompanying plants larvae; pupae Yes 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

                  Impact Summary

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                  CategoryImpact
                  Economic/livelihood Negative

                  Impact

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                  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

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                  Invasiveness
                  • Proved invasive outside its native range
                  • Has a broad native range
                  • Abundant in its native range
                  • Highly adaptable to different environments
                  • Is a habitat generalist
                  • Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
                  • Capable of securing and ingesting a wide range of food
                  • Has high reproductive potential
                  • Has high genetic variability
                  Impact outcomes
                  • Host damage
                  • Negatively impacts agriculture
                  • Negatively impacts livelihoods
                  Likelihood of entry/control
                  • Highly likely to be transported internationally accidentally
                  • Highly likely to be transported internationally illegally
                  • Difficult to identify/detect as a commodity contaminant
                  • Difficult to identify/detect in the field
                  • Difficult/costly to control

                  Diagnosis

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                  Non-morphological methods for fruit fly diagnosis are being investigated, but few species have so far been compared. A. ludens was included in a comparative study of cuticular hydrocarbons of larvae and adults of six species by Carlson and Yocom (1986).

                  Detection and Inspection

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                  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

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                  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

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                  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.

                  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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                  Aluja M, Guillen J, Liedo P, Cabrera M, Rios E, Rosa G de la, Celedonio H, Mota D, 1990. Fruit infesting tephritids (Dipt.: Tephritidae) and associated parasitoids in Chiapas, Mexico. Entomophaga, 35(1):39-48

                  Aluja, M., Arredondo, J., Díaz-Fleischer, F., Birke, A., Rull, J., Niogret, J., Epsky, N., 2014. Susceptibility of 15 mango (Sapindales: Anacardiaceae) cultivars to the attack by Anastrepha ludens and Anastrepha obliqua (Diptera: Tephritidae) and the role of underdeveloped fruit as pest reservoirs: management implications. Journal of Economic Entomology, 107(1), 375-388. doi: 10.1603/EC13045

                  Aluja, M., Díaz-Fleischer, F., Arredondo, J., Valle-Mora, J., Rull, J., 2010. Effect of cold storage on larval and adult Anastrepha ludens (Diptera: Tephritidae) viability in commercially ripe, artificially infested Persea americana 'Hass'. Journal of Economic Entomology, 103(6), 2000-2008. doi: 10.1603/EC09425

                  Aluja, M., Guillén, J., Rosa, G. de la, Cabrera, M., Celedonio, H., Liedo, P., Hendrichs, J., 1987. Natural host plant survey of the economically important fruit flies (Diptera: Tephritidae) of Chiapas, Mexico. Florida Entomologist, 70(3), 329-330. doi: 10.2307/3495066

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                  CABI, Undated. Compendium record. Wallingford, UK: CABI

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                  Carroll L E, Wharton R A, 1989. Morphology of the immature stages of Anastrepha ludens (Diptera; Tephritidae). Annals of the Entomological Society of America. 82 (2), 201-214. DOI:10.1093/aesa/82.2.201

                  Celedonio-Hurtado H, Liedo P, Aluja M, Guillen J, Berrigan D, Carey J, 1988. Demography of Anastrepha ludens, A. obliqua and A. serpentina (Diptera: Tephritidae) in Mexico. Florida Entomologist. 71 (2), 111-120. DOI:10.2307/3495357

                  Darby H H , Kapp E H, 1934. Technical Bulletin. United States Department of Agriculture, Washington, D.C. 20 pp.

                  Déctor N, Malo E A, Rojas J C, Liedo P, 2016. Comparative responses of Anastrepha ludens and Anastrepha obliqua (Diptera: Tephritidae) to the synthetic attractant BioLure. Journal of Economic Entomology. 109 (5), 2054-2060. DOI:10.1093/jee/tow169

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                  EPPO, 2020. EPPO Global database. In: EPPO Global database, Paris, France: EPPO. https://gd.eppo.int/

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                  García-Ramírez A de J, Medina H R E, López-Martínez V, Vázquez L M, Duarte U I E, Delfín-González H, 2010. Talisia olivaeformis (Sapindaceae) and Zuelania guidonia (Flacourtiaceae): new host records for Anastrepha spp. (Diptera: Tephritidae) in México. Florida Entomologist. 93 (4), 633-634. http://www.fcla.edu/FlaEnt/ DOI:10.1653/024.093.0421

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                  Houston W W K, 1981. Fluctuations in numbers and the significance of the sex ratio of the Mexican fruit fly, Anastrepha ludens caught in McPhail traps. Entomologia Experimentalis et Applicata. 30 (2), 140-150. DOI:10.1007/BF00300879

                  IPPC, 2020. Anastrepha ludens (Mexican Fruit Fly): APHIS Establishes a Quarantine in Cameron County, Texas. In: IPPC Official Pest Report, Rome, Italy: FAO. https://www.ippc.int/

                  Jirón L F, Soto-Manitiu J, Norrbom A L, 1988. A preliminary list of the fruit flies of the genus Anastrepha (Diptera: Tephritidae) in Costa Rica. Florida Entomologist. 71 (2), 130-137. DOI:10.2307/3495360

                  Lasa R, Toxtega Y, Herrera F, Cruz A, Navarrete M A, Antonio S, 2014. Inexpensive traps for use in mass trapping Anastrepha ludens (Diptera: Tephritidae). Florida Entomologist. 97 (3), 1123-1130. DOI:10.1653/024.097.0319

                  Loew H, 1873. Monographs of the Diptera of North America, pt. III. 351 pp.

                  Mangan R L, 2003. Adult diet and male-female contact effects on female reproductive potential in Mexican fruit fly (Anastrepha ludens Loew) (Diptera Tephritidae). Journal of Economic Entomology. 96 (2), 341-347. DOI:10.1603/0022-0493-96.2.341

                  Martinez A J, Salinas E J, Rendon P, 2007. Capture of Anastrepha species (Diptera: Tephritidae) with multilure traps and biolure attractants in Guatemala. Florida Entomologist. 90 (1), 258-263. DOI:10.1653/0015-4040(2007)90[258:COASDT]2.0.CO;2

                  NAPPO, 2007. Anastrepha ludens (Mexican fruit fly)-Quarantined Area in San Diego County, California - United States., https://www.pestalerts.org/official-pest-report/anastrepha-ludens-mexican-fruit-fly-quarantined-area-san-diego-county

                  NAPPO, 2007a. Anastrepha ludens (Mexican Fruit Fly) - Quarantined Area in Webb County, Texas - United States., https://www.pestalerts.org/official-pest-reports?keyword=Anastrepha%20ludens&oprID=262

                  NAPPO, 2009. Phytosanitary Alert System: Anastrepha ludens (Mexican fruit fly) - Removal of Quarantine Area in Los Angeles County, California - United States., NAPPO. http://www.pestalert.org/oprDetail.cfm?oprID=385&keyword=anastrepha%20ludens

                  NAPPO, 2012. Phytosanitary Alert System: Anastrepha ludens (Mexican fruit fly) eradicated in the United States., NAPPO. http://www.pestalert.org/oprDetail.cfm?oprID=511&keyword=anastrepha%20ludens

                  NAPPO, 2013. Mexican fruit fly, Anastrepha ludens, eradicated in Texas, US., https://www.pestalerts.org/official-pest-reports?oprID=565

                  NAPPO, 2014. Anastrepha ludens (Mexican fruit fly) - Regulated area established in Texas., https://www.pestalerts.org/official-pest-reports?oprID=580

                  NAPPO, 2014a. Mexican Fruit Fly (Anastrepha ludens) - Removal of Quarantine Area in Weslaco, Hidalgo County, Texas., https://www.pestalerts.org/official-pest-report/mexican-fruit-fly-anastrepha-ludens-removal-quarantine-area-weslaco-hidalgo

                  NAPPO, 2015. Anastrepha ludens (Mexican Fruit Fly) - Establishment of Quarantine Areas in the Rangerville, Cameron County and McAllen, Mission, and La Villa areas of Hidalgo County, Texas., USA: North American Plant Protection Organization. https://www.pestalerts.org/official-pest-report/anastrepha-ludens-mexican-fruit-fly-establishment-quarantine-areas-rangerville

                  NAPPO, 2016. Anastrepha ludens (Mexican Fruit Fly): APHIS Establishes a Quarantine in the San Ygnacio Area of Zapata County, Texas., USA: North American Plant Protection Organization. https://www.pestalerts.org/official-pest-report/anastrepha-ludens-mexican-fruit-fly-aphis-establishes-quarantine-san-ygnacio

                  NAPPO, 2016a. Anastrepha ludens (Mexican Fruit Fly) - APHIS Establishes Quarantine Area in the Encantada Area of Cameron County, Texas., USA: North American Plant Protection Organization. https://www.pestalerts.org/official-pest-report/anastrepha-ludens-mexican-fruit-fly-aphis-establishes-quarantine-area

                  NAPPO, 2016b. Anastrepha ludens (Mexican Fruit Fly) - APHIS Establishes Quarantine Area in the Granjeno Area of Hidalgo County, Texas., USA: North American Plant Protection Organization. https://www.pestalerts.org/official-pest-report/anastrepha-ludens-mexican-fruit-fly-aphis-establishes-quarantine-area-granjeno

                  NAPPO, 2016c. Anastrepha ludens (Mexican Fruit Fly) - APHIS Establishes Quarantine Area in the Hargill Area of Willacy County, Texas., USA: North American Plant Protection Organization. https://www.pestalerts.org/official-pest-report/anastrepha-ludens-mexican-fruit-fly-aphis-establishes-quarantine-area-hargill

                  NAPPO, 2016d. Anastrepha ludens (Mexican Fruit Fly) - APHIS Establishes Quarantine Area in the Rio Hondo Area of Cameron and Willacy Counties, Texas., USA: North American Plant Protection Organization. https://www.pestalerts.org/official-pest-report/anastrepha-ludens-mexican-fruit-fly-aphis-establishes-quarantine-area-rio

                  NAPPO, 2016e. Anastrepha ludens (Mexican fruit fly) - APHIS Establishes Quarantine Area in the Zapata Area of Zapata County, Texas., USA: North American Plant Protection Organization. https://www.pestalerts.org/official-pest-report/anastrepha-ludens-mexican-fruit-fly-aphis-establishes-quarantine-area-zapata

                  NAPPO, 2016f. Anastrepha ludens (Mexican Fruit Fly) - APHIS Establishes Quarantine in the Bayview Area of Cameron County, Texas., USA: North American Plant Protection Organization. https://www.pestalerts.org/official-pest-report/anastrepha-ludens-mexican-fruit-fly-aphis-establishes-quarantine-bayview-area

                  NAPPO, 2016g. Anastrepha ludens (Mexican Fruit Fly) - APHIS Establishes Quarantine in the McCook Area of Hidalgo County, Texas., USA: North American Plant Protection Organization. https://www.pestalerts.org/official-pest-report/anastrepha-ludens-mexican-fruit-fly-aphis-establishes-quarantine-mccook-area

                  NAPPO, 2016h. Anastrepha ludens (Mexican Fruit Fly) - APHIS removes a quarantine in the Bayview area of Cameron County, Texas., USA: North American Plant Protection Organization. https://www.pestalerts.org/official-pest-report/anastrepha-ludens-mexican-fruit-fly-aphis-removes-quarantine-bayview-area

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                  NAPPO, 2016j. Anastrepha ludens (Mexican Fruit Fly) - APHIS removes quarantine in the Rio Hondo area of Cameron and Willacy Counties, Texas., USA: North American Plant Protection Organization. https://www.pestalerts.org/official-pest-report/anastrepha-ludens-mexican-fruit-fly-aphis-removes-quarantine-rio-hondo-area

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                  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

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