Anastrepha ludens (Mexican fruit fly)

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Adults

Adult females ovipositing on citrus fruit skin.

USDA-ARS

Line artwork of adult female

CAB 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: 12 May 2022

Continent/Country/Region	Distribution	Origin	Invasive	Reference	Notes
Europe
Slovenia	Absent			EPPO (2022)
North America
Belize	Present	Native		UK, CAB International (1958) Van Whervin (1966) Houston (1981) Ruiz-Arce et al. (2015) EPPO (2022)
Costa Rica	Present, Localized			Stone (1942) UK, CAB International (1958) Jirón et al. (1988) Epsky (2003) Ruiz-Arce et al. (2015) Scally et al. (2016) EPPO (2022) CABI (Undated)	Higher elevations. More common and widely distributed since the 1990s
El Salvador	Present	Native		UK, CAB International (1958) Delmi et al. (1996) EPPO (2022)
Guatemala	Present	Native		Stone (1942) UK, CAB International (1958) Eskafi and Cunningham (1987) Martinez et al. (2007) Ruiz-Arce et al. (2015) Scally et al. (2016) EPPO (2022)
Honduras	Present	Native		UK, CAB International (1958) Epsky (2003) Ruiz-Arce et al. (2015) EPPO (2022)
Mexico	Present, Widespread			EPPO (2022) Loew (1873) Darby and Kapp (1934) Stone (1942) UK, CAB International (1958) Celedonio-Hurtado et al. (1988) Carroll and Wharton (1989) Hernandez-Ortiz (1992) Epsky (2003) Mangan (2003) Thomas (2003) Rull et al. (2007) García-Ramírez et al. (2010) Ruiz-Montiel et al. (2013) Aluja et al. (2014) Lasa et al. (2014) Ruiz-Arce et al. (2015) Vanoye-Eligio et al. (2015) Déctor et al. (2016) Scally et al. (2016) Vanoye-Eligio et al. (2017)
Nicaragua	Present	Native		UK, CAB International (1958) Borge and Basedow (1997) EPPO (2022) CABI (Undated)
Panama	Present, Localized	Introduced	Invasive	Norrbom et al. (2005) Ruiz-Arce et al. (2015) EPPO (2022)	Higher elevations, western Panama; First reported: <1996
United States	Present, Transient under eradication			Stone (1942) UK, CAB International (1958) Foote et al. (1993) Steck (2003) NAPPO (2007) NAPPO (2007a) NAPPO (2012) NAPPO (2013) Aluja et al. (2014) NAPPO (2014) NAPPO (2014a) NAPPO (2015) NAPPO (2016) NAPPO (2016a) NAPPO (2016b) NAPPO (2016c) NAPPO (2016d) NAPPO (2016e) NAPPO (2016f) NAPPO (2016g) NAPPO (2016h) NAPPO (2016i) NAPPO (2016j) NAPPO (2016k) NAPPO (2017) NAPPO (2017a) NAPPO (2018) EPPO (2022)
-Arizona	Absent, Formerly present			EPPO (2022) UK, CAB International (1958) Foote et al. (1993)
-California	Present, Transient under eradication			EPPO (2022) UK, CAB International (1958) Foote et al. (1993) NAPPO (2007) NAPPO (2009)
-Florida	Absent, Intercepted only			UK, CAB International (1958) Aluja et al. (2014) EPPO (2022)	Rarely trapped, not established
-Texas	Present, Transient under eradication			Stone (1942) UK, CAB International (1958) Foote et al. (1993) NAPPO (2012) NAPPO (2017) NAPPO (2017a) NAPPO (2018) IPPC (2020) EPPO (2022) CABI (Undated)
Oceania
New Zealand	Absent, Confirmed absent by survey			EPPO (2022)
South America
Argentina	Absent, Invalid presence record(s)			EPPO (2022)
Colombia	Absent, Invalid presence record(s)			EPPO (2022) Núñez Bueno (1981) Norrbom et al. (2005)

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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Category	Sub-Category	Habitat	Presence	Status
Terrestrial	Managed	Cultivated / agricultural land	Present, no further details	Harmful (pest or invasive)
Terrestrial	Managed	Managed forests, plantations and orchards	Present, no further details	Harmful (pest or invasive)
Terrestrial	Managed	Disturbed areas	Present, no further details
Terrestrial	Managed	Urban / peri-urban areas	Present, no further details	Harmful (pest or invasive)
Terrestrial	Natural / Semi-natural	Natural 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 name	Family	Context	References
Anacardium occidentale (cashew nut)	Anacardiaceae	Other	Eskafi and Cunningham (1987)
Annona	Annonaceae	Other	Ruiz-Montiel et al. (2013)
Annona cherimola (cherimoya)	Annonaceae	Other
Annona liebmanniana (hardshell custard-apple)	Annonaceae	Other	Ruiz-Montiel et al. (2013)
Annona muricata (soursop)	Annonaceae	Other
Annona reticulata (bullock's heart)	Annonaceae	Other	Ruiz-Montiel et al. (2013)
Annona squamosa (sugar apple)	Annonaceae	Other	Aluja et al. (1987)
Capsicum pubescens (rocoto)	Solanaceae	Unknown	Steck (2003); Thomas (2004)
Carica papaya (pawpaw)	Caricaceae	Other	Eskafi and Cunningham (1987)
Casimiroa edulis (white sapote)	Rutaceae	Main	Aluja et al. (1987); Aluja et al. (2000); Birke and Aluja (2011); Bush (1962); Eskafi and Cunningham (1987); Jimenez et al. (1992); Orozco-Dávila et al. (2007)
Casimiroa greggii		Unknown	Aluja et al. (2009); Pecina et al. (2009)
Casimiroa pubescens	Rutaceae	Other	Vanoye-Eligio et al. (2015)
Citrus	Rutaceae	Main	Aluja et al. (1987); Hernández-Ortiz et al. (2006); Jimenez et al. (1992)
Citrus aurantiifolia (lime)	Rutaceae	Other	Eskafi and Cunningham (1987)
Citrus aurantium (sour orange)	Rutaceae	Main	Déctor et al. (2016); Mangan (2003); Eskafi and Cunningham (1987); Aluja et al. (1987); Aluja et al. (2000); Birke and Aluja (2011); Bush (1962); Dor et al. (2014); Hernandez et al. (2013); Hernández-Ortiz et al. (2006); Jimenez et al. (1992); Molina-Nery et al. (2014); Orozco-Dávila et al. (2007)
Citrus limetta (sweet lemon tree)	Rutaceae	Other	Aluja et al. (1987); Eskafi and Cunningham (1987); Jimenez et al. (1992)
Citrus maxima (pummelo)	Rutaceae	Main	Bush (1962); Aluja et al. (1987); Aluja et al. (2000); Jimenez et al. (1992)
Citrus medica (citron)	Rutaceae	Other	Aluja et al. (1987)
Citrus reshni (Cleopatra mandarin)	Rutaceae	Unknown	Eskafi and Cunningham (1987)
Citrus reticulata (mandarin)	Rutaceae	Other	Aluja et al. (1987); Eskafi and Cunningham (1987); Aluja et al. (2000)
Citrus reticulata x paradisi (tangelo)	Rutaceae	Other
Citrus sinensis (sweet orange)	Rutaceae	Main	Aluja et al. (1987); Aluja et al. (2000); Aluja et al. (2009); Bush (1962); Eskafi and Cunningham (1987); Hernández-Ortiz et al. (2006); Jimenez et al. (1992); Lasa et al. (2014); López et al. (1999); Mangan et al. (2011); Molina-Nery et al. (2014); Pecina et al. (2009)
Citrus x paradisi (grapefruit)	Rutaceae	Main	Eskafi and Cunningham (1987); Aluja et al. (2014); Aluja et al. (1987); Aluja et al. (2009); Aluja et al. (2000); Aluja et al. (2009); Birke and Aluja (2011); Díaz-Fleischer and Aluja (2003); Dor et al. (2014); Foote and Blanc (1963); Hernández-Ortiz et al. (2006); Jimenez et al. (1992); Mangan et al. (2011); Molina-Nery et al. (2014); Rull et al. (2007)
Coffea arabica (arabica coffee)	Rubiaceae	Other	Eskafi and Cunningham (1987)
Diospyros ebenaster (black sapote)	Ebenaceae	Unknown	Orozco-Dávila et al. (2007)
Diospyros kaki (persimmon)	Ebenaceae	Other	Eskafi and Cunningham (1987)
Ficus carica (common fig)	Moraceae	Unknown	Foote and Blanc (1963)
fruits		Other
Heteromeles salicifolia (toyon)	Rosaceae	Unknown	Foote and Blanc (1963)
Inga	Fabaceae	Other
Inga micheliana		Unknown	Eskafi and Cunningham (1987)
Malus domestica (apple)	Rosaceae	Other	Eskafi and Cunningham (1987)
Mammea	Clusiaceae	Unknown	Da (1934)
Mammea americana (mamey apple)	Clusiaceae	Other	Bush (1962)
Mangifera indica (mango)	Anacardiaceae	Main	Aluja et al. (2010); Aluja et al. (1987); Aluja et al. (2000); Berrigan et al. (1988); Carroll and Wharton (1989); Celedonio-Hurtado et al. (1988); Darby and Kapp (1934); Eskafi and Cunningham (1987); Hernández et al. (2018); Hernández-Ortiz et al. (2006); Jimenez et al. (1992); López et al. (1999); Malo and Zapien (1994)
Morus		Unknown	Foote and Blanc (1963)
Mosiera longipes	Lithomyrtus	Unknown	Eskafi and Cunningham (1987)
Passiflora edulis (passionfruit)	Passifloraceae	Other	Eskafi and Cunningham (1987)
Persea americana (avocado)	Lauraceae	Other
Pouteria campechiana (canistel)	Sapotaceae	Unknown	Orozco-Dávila et al. (2007)
Prunus persica (peach)	Rosaceae	Main	Aluja et al. (2000); Eskafi and Cunningham (1987)
Psidium cattleianum (strawberry guava)	Lithomyrtus	Other
Psidium guajava (guava)	Lithomyrtus	Other	Darby and Kapp (1934); Eskafi and Cunningham (1987); Jimenez et al. (1992); López et al. (1999)
Punica granatum (pomegranate)	Punicaceae	Other	Foote and Blanc (1963); Jimenez et al. (1992)
Pyrus communis (European pear)	Rosaceae	Other	Eskafi and Cunningham (1987)
Sargentia greggii	Rutaceae	Unknown	Bush (1962); Gonzalez-Hernandez and Tejada (1979); Mangan (2003); Orozco-Dávila et al. (2007); Plummer and Mcphail (1941)
Sideroxylon capiri subsp. tempisque		Unknown	Aluja et al. (1987)
Spondias purpurea (red mombin)	Anacardiaceae	Other	Eskafi and Cunningham (1987)
Syzygium jambos (rose apple)	Lithomyrtus	Other	Eskafi and Cunningham (1987)
Talisia olivaeformis	Sapindaceae	Other
Terminalia catappa (Singapore almond)	Combretaceae	Unknown	Jimenez et al. (1992)

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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Sign	Life Stages	Type
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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Climate	Status	Description
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 enemy	Type	Life stages	Biological control in	Biological control on
Aceratoneuromyia indica	Parasite	Arthropods\|Larvae	Belize; Mexico	Citrus; grapefruits; oranges
Biosteres arisanus	Parasite		Mexico	Citrus
Biosteres longicaudatus	Parasite	Arthropods\|Larvae	Belize; Mexico	Citrus; grapefruits; oranges
Biosteres vandenboschi	Parasite		Mexico	Citrus
Diachasmimorpha tryoni	Parasite		Mexico	Citrus
Dirhinus giffardii	Parasite		Mexico	Citrus
Doryctobracon crawfordi	Parasite	Arthropods\|Larvae
Lyssomanes pescadero	Predator
Opius incisi	Parasite		Mexico	Citrus
Pachycrepoideus vindemmiae	Parasite	Arthropods\|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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Cause	Notes	Long Distance	Local	References
Crop production		Yes	Yes

Pathway Vectors

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Vector	Notes	Long Distance	Local
Aircraft	Immatures in fruit	Yes	Yes
Containers and packaging - wood	Of fruit cargo.	Yes
Luggage	Immatures in fruit	Yes	Yes
Mail	Fruit in post.	Yes
Plants or parts of plants	Immatures in fruit	Yes	Yes
Soil, sand and gravel	Risk of puparia in soil.	Yes

Plant Trade

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Plant parts liable to carry the pest in trade/transport	Pest stages	Borne internally	Borne externally	Visibility of pest or symptoms
Fruits (inc. pods)	arthropods/eggs; arthropods/larvae; arthropods/pupae	Yes		Pest or symptoms usually visible to the naked eye
Growing medium accompanying plants	arthropods/larvae; arthropods/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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Category	Impact
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

Aluja, M., Ordano, M., Teal, P. E. A., Sivinski, J., García-Medel, D., Anzures-Dadda, A., 2009. Larval feeding substrate and species significantly influence the effect of a juvenile hormone analog on sexual development/performance in four tropical tephritid flies. Journal of Insect Physiology, 55(3), 231-242. doi: 10.1016/j.jinsphys.2008.11.013

Aluja, M., Piñero, J., López, M., Ruíz, C., Zúñiga, A., Piedra, E., Díaz-Fleischer, F., Sivinski, J., 2000. New host plant and distribution records in Mexico for Anastrepha spp., Toxotrypana curvicauda Gerstacker, Rhagoletis zoqui Bush, Rhagoletis sp., and Hexachaeta sp. (Diptera: Tephritidae). Proceedings of the Entomological Society of Washington, 102(4), 802-815.

Aluja, M., Rull, J., Pérez-Staples, D., Díaz-Fleischer, F., Sivinski, J., 2009. Random mating among Anastrepha ludens (Diptera: Tephritidae) adults of geographically distant and ecologically distinct populations in Mexico. Bulletin of Entomological Research, 99(2), 207-214. doi: 10.1017/S0007485308006299

Aluja, M.ín, Sivinski, J., Ovruski, S., Guillén, L., López, M., Cancino, J., Torres-Anaya, A., Gallegos-Chan, G., Ruíz, Lía, 2009. Colonization and domestication of seven species of native New World hymenopterous larval-prepupal and pupal fruit fly (Diptera: Tephritidae) parasitoids. Biocontrol Science and Technology, 19, 49-79. doi: 10.1080/09583150802377373

BAKER AC, STONE WE, PLUMMER CC, MCPAHAIL M, 1944. A Review of Studies on the Mexican Fruitfly and related Mexican Species. Miscellaneous Publications. United States Department of Agriculture. Washington, D.C., 155 pp.

Bateman MA, 1982. III. Chemical methods for suppression or eradication of fruit fly populations, In: Drew RAI, Hooper GHS, Bateman MA eds. Economic Fruit Flies of the South Pacific Region. 2nd edn. Brisbane, Australia: Queensland Department of Primary Industries, 115-128.

Bateman MA, 1982. III. Chemical methods for suppression or eradication of fruit fly populations. In: Economic Fruit Flies of the South Pacific Region [ed. by Drew RAI, Hooper GHS, Bateman MA] Brisbane, Australia: Queensland Department of Primary Industries, 115-128.

Berg GH, 1979. Pictorial key to fruit fly larvae of the family Tephritidae. Pictorial key to fruit fly larvae of the family Tephritidae. Organismo Internacional Regional de Sanidad Agropecuaria. San Salvador El Salvador, 36 pp.

Berrigan, D. A., Carey, J. R., Guillen, J., Celedonio, H., 1988. Age and host effects on clutch size in the Mexican fruit fly, Anastrepha ludens. Entomologia Experimentalis et Applicata, 47(1), 73-80. doi: 10.1007/BF00186718

Birke, A., Aluja, M., 2011. Anastrepha ludens and Anastrepha serpentina (Diptera: Tephritidae) do not infest Psidium guajava (Myrtaceae), but Anastrepha obliqua occasionally shares this resource with Anastrepha striata in nature. Journal of Economic Entomology, 104(4), 1204-1211. doi: 10.1603/EC11042

Blanco Montero CA, Sanchez Salas JA, 1990. Evaluation of six different traps for detecting the Mexican fruit fly. Southwestern Entomologist, 15(3):327-331

Bush, G. L., 1962. The Cytotaxonomy of the Larvae of Some Mexican Fruit Flies in the Genus Anastrepha (Tephritidae, Diptera). Psyche, 69(3), 87-101. doi: 10.1155/1962/20495

CABI/EPPO, 1998. Distribution maps of quarantine pests for Europe (edited by Smith IM, Charles LMF). Wallingford, UK: CAB International, xviii + 768 pp.

CABI/EPPO, 2001. Anastrepha ludens. Distribution Maps of Plant Pests, Map No. 89. Wallingford, UK: CAB International.

Calkins CO, Schroeder WJ, Chambers DL, 1984. Probability of detecting Caribbean fruit fly, Anastrepha suspensa (Loew) (Diptera: Tephritidae), populations with McPhail traps. Journal of Economic Entomology, 77(1):198-201.

Carlson DA, Yocom SR, 1986. Cuticular hydrocarbons from six species of tephritid fruit flies. Archives of Insect Biochemistry and Physiology, 3(4):397-412

Carroll LE, Norrbom AL, Dallwitz MJ, Thompson FC, 2004. Pest fruit flies of the world - larvae. http://delta-intkey

Carroll LE, Wharton RA, 1989. Morphology of the immature stages of Anastrepha ludens (Diptera; Tephritidae). Annals of the Entomological Society of America, 82(2):201-214

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

Christenson LD, Foote RH, 1960. Biology of fruit flies. Annual Review of Entomology, 5:171-192.

Clausen CP, 1978. Tephritidae (Trypetidae, Trupaneidae), In: Clausen CP, ed. Introduced Parasites and Predators of Arthropod Pests and Weeds: A World Review. Agricultural Handbook, United States Department of Agriculture, 480:320-335.

Da Costa Lima, A. , 1934. Fruit-flies of the Genus Anastrepha. (Moscas de frutas do genero Anastrepha Schiner, 1868). Memorias do Instituto Oswaldo Cruz, 28(4), 487-575 pp.

Darby, H. H. , Kapp, E. H. , 1934. Technical Bulletin. United States Department of Agriculture, Washington, D.C (444), 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

Díaz-Fleischer, F., Aluja, M., 2003. Clutch size in frugivorous insects as a function of host firmness: the case of the tephritid fly Anastrepha ludens. Ecological Entomology, 28(3), 268-277. doi: 10.1046/j.1365-2311.2003.00517.x

Dor, A., Valle-More, J., Rodriguez-Rodriguez, S. E., Liedo, P., 2014. Predation of Anastrepha ludens (Diptera: Tephritidae) by Norops serranoi (Reptilia: Polychrotidae): Functional Response and Evasion Ability. Environmental Entomology, 43(3), 706-715. doi: 10.1603/EN13281

Drew RAI, 1982. Fruit fly collecting. In: Drew RAI, Hooper GHS, Bateman MA, eds. Economic Fruit Flies of the South Pacific Region, 2nd edition. Brisbane, Australia: Queensland Department of Primary Industries, 129-139.

EPPO, 1983. Data sheets on quarantine organisms. Set 6. EPPO Bulletin, 13(1). unnumbered.

EPPO, 1990. Specific quarantine requirements. EPPO Technical Documents, No. 1008. Paris, France: European and Mediterranean Plant Protection Organization.

EPPO, 2014. PQR database. Paris, France: European and Mediterranean Plant Protection Organization. http://www.eppo.int/DATABASES/pqr/pqr.htm

Epsky ND, Heath RR, Guzman A, Meyer WL, 1995. Visual cue and chemical cue interactions in a dry trap with food-based synthetic attractant for Ceratitis capitata and Anastrepha ludens (Diptera: Tephritidae). Environmental Entomology, 24(6):1387-1395; 16 ref.

Eskafi FM, 1988. Infestation of citrus by Anastrepha spp. and Ceratitis capitata (Diptera: Tephritidae) in high coastal plains of Guatemala. Environmental Entomology, 17(1):52-58

Eskafi, F. M., Cunningham, R. T., 1987. Host plants of fruit flies (Diptera: Tephritidae) of economic importance in Guatemala. Florida Entomologist, 70(1), 116-123. doi: 10.2307/3495098

FAO/IAEA, 2003. Trapping Guidelines for area-wide fruit fly programmes. Vienna, Austria: International Atomic Energy Agency, 47 pp.

Fletcher BS, 1989. Ecology; movements of tephritid fruit flies. In: Robinson AS, Hooper G, eds. Fruit Flies; Their Biology, Natural Enemies and Control. World Crop Pests, 3(B). Amsterdam, Netherlands: Elsevier, 209-219.

Foote RH, Blanc FL, Norrbom AL, 1993. Handbook of the Fruit Flies (Diptera: Tephritidae) of America North of Mexico. Ithaca, USA: Comstock.

Foote, R. H., Blanc, F.L., 1963. THE FRUIT FLIES OR TEPHRITIDAE OF CALIFORNIA. In: BULLETIN OF THE CALIFORNIA INSECT SURVEY,7, USA: University of California Press. 1-117. https://essig.berkeley.edu/documents/cis/cis07.pdf

García-Ramírez Ade J, Medina H RE, López-Martínez V, Vázquez L M, Duarte U IE, 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/

Gilmore JE, 1989. Control; sterile insect technique (SIT); overview. In: Robinson AS, Hooper G, eds. Fruit Flies; Their Biology, Natural Enemies and Control. World Crop Pests 3(B). Amsterdam, Netherlands: Elsevier, 353-363.

Gonzalez-Hernandez A, Tejada LO, 1979. Population fluctuations of Anastrepha ludens (Loew) and of its natural enemies on Sargentia greggii S. Watts. Folia Entomologica Mexicana, No. 41:49-60

Gonzalez-Hernandez A, Tejada LO, 1980. Especies de Anastrepha (Diptera: Tephritidae) en el Estado de Nuevo Leon, Mexico. Folia Entomologica Mexicana, 44:121-128.

Hallman GJ, 1997. Mortality of Mexican fruit fly (Diptera: Tephritidae) immatures in coated grapefruits. Florida Entomologist, 80(3):324-328; 11 ref.

Heath RR, Epsky ND, Bloem S, Bloem K, Acajabon F, Guzman A, Chambers D, 1994. pH Effect on the attractiveness of a corn hydrolysate to the Mediterranean fruit fly and several Anastrepha species (Diptera: Tephritidae). Journal of Economic Entomology, 87(4):1008-1013

Heath RR, Epsky ND, Guzman A, Dueben B, Manukian A, Meyer WL, 1995. Development of a dry plastic insect trap with food-based synthetic attractant for the Mediterranean and Mexican fruit flies (Diptera: Tephritidae). Journal of Economic Entomology, 88(5):1307-1315; 26 ref.

Hedstrom I, Jimenez J, 1988. Field evaluation of attractants in the capture of Anastrepha spp. (Diptera, Tephritidae), pests of fruit trees in tropical America. II. Ammonium acetate and torula with sodium borate. Revista Brasileira de Entomologia, 32(2):319-322.

Hedström I, Jirón LF, 1985. Field evaluation of attractants in the trapping of Anastrepha spp. (Diptera, Tephritidae), pests of fruit trees in tropical America. I. Molasses and torula yeast. Revista Brasileira de Entomologia, 29(3/4):515-520.

Heppner JB, 1984. larvae of fruit flies. I. Anastrepha ludens (Mexican fruit fly) and Anastrepha suspensa (Caribbean fruit fly) (Diptera: Tephritidae). Entomology Circular, Division of Plant Industry, Florida Department of Agricultural and Consumer Services, No. 260, 4 pp.

Hernandez Fuentes, L. M., Urías López, M. A., Castañeda Vildózola, A., Hernandez Ortiz, V., Orozco Santos, M., 2013. No-host status of fruit flies Anastrepha ludens and A. obliqua (Diptera: Tephritidae) in soursop fruit, Annona muricata (Magnoliales: Annonaceae). Southwestern Entomologist, 38(1), 85-97. doi: 10.3958/059.038.0109

Hernández, E., Aceituno-Medina, M., Toledo, J., Gómez-Simuta, Y., Villarreal-Fuentes, J. M., Carrasco, M., Liedo, P., Hallman, G. J., Montoya, P., 2018. Generic irradiation and hot water phytosanitary treatments for mango fruits cv. 'Ataulfo' niño infested by Anastrepha ludens and Anastrepha obliqua (Diptera: Tephritidae). Journal of Economic Entomology, 111(5), 2110-2119. doi: 10.1093/jee/toy198

Hernandez-Ortiz V, 1992. El género Anastrepha en México. Taxonomfa, distribución y sus plantas huéspedes. Xalapa, Mexico: Instituto de Ecologfa.

Hernandez-Ortiz V, Perez-Alonso R, 1993. The natural host plants of Anastrepha (Diptera: Tephritidae) in a tropical rain forest of Mexico. Florida Entomologist, 76(3):447-460

Hernández-Ortiz, V., Delfín-González, H., Escalante-Tio, A., Manrique-Saide, P., 2006. Hymenopteran parasitoids of Anastrepha fruit flies (Diptera: Tephritidae) reared from different hosts in Yucatan, Mexico. Florida Entomologist, 89(4), 508-515. doi: 10.1653/0015-4040(2006)89[508:HPOAFF]2.0.CO;2

Jimenez ML, Tejas A, 1996. Variacion temporal de la araneofauna en frutales de la region del Cabo, Baja California Sur Mexico. Southwestern Entomologist, 21:331-336.

Jimenez, M. L., Tejas, A., Servin, R., 1992. Natural hosts of Anastrepha ludens (Loew) (Diptera: Tephritidae) in the La Paz area, B.C.S. Mexico. (Hospederos naturales de Anastrepha ludens (Loew) (Diptera: Tephritidae) en el municipio de La Paz, B.C.S. Mexico). Folia Entomológica Mexicana, (No. 84), 157-158.

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

Kisliuk M, Cooley DE, 1933. Fruit-fly survey in the West Indies, Brazil, Uruguay, Chile, and Peru. Department of Agriculture, Plant Quarantine Service Regulatory Announcements, 227-243.

Landolt PJ, Heath RR, 1996. Development of pheromone-based trapping systems for monitoring and controlling tephritid fruit flies in Florida. In: Rosen D, Bennett FD, Capinera JL, eds. Pest Management in the Subtropics: Integrated Pest Management a Florida Perspective. Andover, UK: Intercept Limited, 197-207.

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

Lee ChangJoo, DeMilo AL, Moreno DS, Mangan RL, 1997. Identification of the volatile components of E802 Mazoferm steepwater, a condensed fermented corn extractive highly attractive to the Mexican fruit fly (Diptera: Tephritidae). Journal of Agricultural and Food Chemistry, 45(6):2327-2331; 19 ref.

Loew H, 1873. Monographs of the Diptera of North America, pt. III. Smithsn. Misc. Collect. 11, (no. 256), 1-351.

López, M., Aluja, M., Sivinski, J., 1999. Hymenopterous larval-pupal and pupal parasitoids of Anastrepha flies (Diptera: Tephritidae) in Mexico. Biological Control, 15(2), 119-129. doi: 10.1006/bcon.1999.0711

Malo E, Baker PS, Valenzuela J, 1987. The abundance of species of Anastrepha (Diptera; Tephritidae) in the coffee producing area of coastal Chiapas, southern Mexico. Folia Entomologica Mexicana, No. 73:125-140

Malo, E. A., Zapien, G. I., 1994. McPhail trap captures of Anastrepha obliqua and Anastrepha ludens (Diptera: Tephritidae) in relation to time of day. Florida Entomologist, 77(2), 290-294. doi: 10.2307/3495518

Mangan RL, 1996. Effects of adult chill treatments on mate attracting potential for irradiated Mexican fruit fly males. Entomologia Experimentalis et Applicata, 79(2):153-159; 20 ref.

Mangan RL, Ingle SJ, 1994. Forced hot-air quarantine treatment for grapefruit infested with Mexican fruit fly (Diptera: Tephritidae). Journal of Economic Entomology, 87(6):1574-1579

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

Mangan, R. L., Thomas, D. B., Moreno, A. M. T., 2011. Host status of grapefruit and valencia oranges for Anastrepha serpentina and Anastrepha ludens (Diptera: Tephritidae). Journal of Economic Entomology, 104(2), 388-397. doi: 10.1603/EC10147

Martinez AJ, Robacker DC, Garcia JA, 1997. Toxicity of an isolate of Bacillus thuringiensis subspecies darmstadiensis to adults of the Mexican fruit fly (Diptera: Tephritidae) in the laboratory. Journal of Economic Entomology, 90(1):130-134; 20 ref.

McAlister Jr LC, 1936, April. Observations on the West Indian Fruit Fly at Key West in 1932-33. Journal of Economic Entomology, 29(2):440-445 pp.

Molina-Nery, M. C., Ruiz-Montoya, L., Zepeda-Cisneros, C. S., Liedo, P., 2014. Genetic structure of populations of Anastrepha ludens (Diptera: Tephritidae) in Mexico. Florida Entomologist, 97(4), 1648-1661. doi: 10.1653/024.097.0439

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. Phytosanitary Alert System: Mexican fruit fly, Anastrepha ludens - regulated area established in Willacy County, Texas. Phytosanitary Alert System: Mexican fruit fly, Anastrepha ludens - regulated area established in Willacy County, Texas. NAPPO. http://www.pestalert.org/oprDetail.cfm?oprID=548

NAPPO, 2013. Phytosanitary Alert System: Mexican fruit fly, Anastrepha ludens, eradicated in Texas, US. Phytosanitary Alert System: Mexican fruit fly, Anastrepha ludens, eradicated in Texas, US. NAPPO. http://www.pestalert.org/oprDetail.cfm?oprID=565

Norrbom AL, 2004. Host plant database for Anastrepha and Toxotrypana (Diptera: Tephritidae: Toxotrypanini). Diptera Data Dissemination Disk (CD-ROM) 2.

Norrbom AL, Foote RH, 1989. Taxonomy and zoogeography; the taxonomy and zoogeography of the genus Anastrepha (Diptera: Tephritidae). In: Robinson AS, Hooper G, eds. Fruit flies; Their Biology, Natural Enemies and Control. World Crop Pests, 3(A). Amsterdam, Netherlands: Elsevier, 15-26.

Norrbom AL, Kim KC, 1988. A list of the reported host plants of the species of Anastrepha (Diptera: Tephritidae). Hyattsville, MD, USA: US Dept. Agric., Animal and Plant Health Inspection Service, Plant Protection and Quarantine.

Norrbom AL, Korytkowski CA, Gonzalez F, Orduz B, 2005. A new species of Anastrepha from Colombia related to Mexican fruit fly (Diptera: Tephritidae). Revista Colombiana de Entomologia, 31(1):67-70.

Núñez Bueno L, 1981. [English title not available]. (Contribucion al reconocimiento de las moscas de las frutas (Diptera: Tephriridae [sic]) en Colombia) Revista do Instituto Colombiano Agropecuario, 16:173-179.

OEPP/EPPO, 1990. Specific quarantine requirements. EPPO Technical Documents, No. 1008. Paris, France: EPPO.

Orozco-Dávila, D., Hernández, R., Meza, S., Domínguez, J., 2007. Sexual competitiveness and compatibility between mass-reared sterile flies and wild populations of Anastrepha ludens (Diptera: Tephritidae) from different regions in Mexico. Florida Entomologist, 90(1), 19-26. doi: 10.1653/0015-4040(2007)90[19:SCACBM]2.0.CO;2

Pecina Quintero, V., López Arroyo, J. I., Loera Gallardo, J., Rull, J., Rosales Robles, E., Cortez Mondaca, E., Hernández Delgado, S., Mayek Perez, N., Aluja Schuneman, M., 2009. Genetic differences between Anastrepha ludens (Loew) populations stemming from a native and an exotic host in NE Mexico. Agricultura Técnica en México, 35(3), 323-331.

Plummer, C. C., Mcphail, M., 1941. The yellow chapóte (Sargentia greggii) a native host of the Mexican fruitfly (Anastrepha ludens). Technical Bulletin. United States Department of Agriculture, 775, 12 pp.

Robacker DC, 1992. Effects of shape and size of colored traps on attractiveness to irradiated, laboratory-strain Mexican fruit flies (Diptera: Tephritidae). Florida Entomologist, 75(2):230-241

Robacker DC, Flath RA, 1995. Attractants from Staphylococcus aureus cultures for Mexican fruit fly, Anastrepha ludens. Journal of Chemical Ecology, 21(11):1861-1874; 24 ref.

Robacker DC, Heath RR, 1996. Attraction of Mexican fruit flies (Diptera: Tephritidae) to lures emitting host-fruit volatiles in a citrus orchard. Florida Entomologist, 79(4):600-602; 4 ref.

Roessler Y, 1989. Control; insecticides; insecticidal bait and cover sprays. In: Robinson AS, Hooper G, eds. Fruit Flies. Their Biology, Natural Enemies and Control. World Crop Pests 3(B). Amsterdam, Netherlands: Elsevier, 329-336.

Ruiz-Montiel C, Flores-Peredo R, Hernández-Librado V, Illescas-Riquelme CP, Domínguez-Espinosa PI, Piñero JC, 2013. Annona liebmanniana and A. cherimola × A. reticulata (Magnoliales: Annonaceae): two new host plant species of Anastrepha ludens (Diptera: Tephritidae) in Mexico. Florida Entomologist, 96(1):232-234. http://www.fcla.edu/FlaEnt/

Rull, J., Diaz-Fleischer, F., Arredondo, J., 2007. Irradiation of Anastrepha ludens (Diptera: Tephritidae) revisited: optimizing sterility induction. Journal of Economic Entomology, 100(4), 1153-1159. doi: 10.1603/0022-0493(2007)100[1153:IOALDT]2.0.CO;2

Sharp JL, 1987. Laboratory and field experiments to improve enzymatic casein hydrolysate as an arrestant and attractant for Caribbean fruit fly, Anastrepha suspensa (Diptera: Tephritidae). Florida Entomologist, 70(2):225-233.

Shellie KC, Mangan RL, Ingle SJ, 1997. Tolerance of grapefruit and Mexican fruit fly larvae to heated controlled atmospheres. Postharvest Biology and Technology, 10(2):179-186; 18 ref.

Silva-Contreras JdeJ, 1978. Efectividad de dos metodos de control quimico de la mosca mexicana de la fruta, Anastrepha ludens Loew, en el Istmo de Tehuantepec, Oaxaca. Instituto Nacional de Investigaciones Agricolas, Mexico: Technical Report 1978. Informe Tecnico de la Coordinacion Nacional del Apoyo Entomologico, 3:60-64.

Smith IM McNamara DG Scott PR Holderness M, 1997. Ceratitis capitata. In: Quarantine Pests for Europe. Wallingford, UK: CABI, 146-152.

Smith IM, McNamara DG, Scott PR, Holderness M, 1997. Quarantine pests for Europe. Second Edition. Data sheets on quarantine pests for the European Union and for the European and Mediterranean Plant Protection Organization. Quarantine pests for Europe. Second Edition. Data sheets on quarantine pests for the European Union and for the European and Mediterranean Plant Protection Organization., Ed. 2:vii + 1425 pp.; many ref.

Steck GJ, 2001. Concerning the occurrence of Anastrepha obliqua (Diptera: Tephritidae) in Florida. Florida Entomologist, 84(2):320-321.

Steck GJ, Carroll LE, Celedonio-Hurtado H, Guillen-Aguilar J, 1990. Methods for identification of Anastrepha larvae (Diptera: Tephritidae), and key to 13 species. Proceedings of the Entomological Society of Washington, 92(2):333-346

Steck, G. J., 2003. Pest Alert: Mexican fruit fly Anastrepha ludens (Loew) (Tephritidae). USA: Florida Department of Agriculture and Consumer Services, Division of Plant Industry .https://www.fdacs.gov/content/download/66386/file/Pest%20Alert%20-%20Anastrepha%20ludens%20-%20Mexican%20Fruit%20Fly.pdf

Stone A, 1942. The fruitflies of the genus Anastrepha. Miscellaneous Publications of the United States Department of Agriculture, 439:1-112.

Thomas DB, 1993. Survivorship of the pupal stages of the Mexican fruit fly Anastrepha ludens (Loew) (Diptera: Tephritidae) in an agricultural and a nonagricultural situation. Journal of Entomological Science, 28(4):350-362

Thomas DB, 1995. Predation on the soil inhabiting stages of the Mexican fruit fly. Southwestern Entomologist, 20(1):61-71

Thomas DB, Mangan RL, 1995. Morbidity of the pupal stage of the Mexican and West Indian fruit flies (Diptera: Tephritidae) induced by hot-water immersion in the larval stage. Florida Entomologist, 78(2):235-246

Thomas, D. B., 2004. Hot peppers as a host for the Mexican fruit fly Anastrepha ludens (Diptera: Tephritidae). Florida Entomologist, 87(4), 603-608. doi: 10.1653/0015-4040(2004)087[0603:HPAAHF]2.0.CO;2

UK CAB International, 2001. Anastrepha ludens. [Distribution map]. Distribution Maps of Plant Pests. Wallingford, UK: CAB International, Map 89.

USDA, 1994. Treatment manual. Frederick, USA: USDA/APHIS.

Vanoye-Eligio, V., Barrientos-Lozano, L., Gaona-García, G., Lara-Villalón, M., 2015. New wild host of Anastrepha ludens in Northeastern Mexico. Southwestern Entomologist, 40(2), 435-437. http://www.bioone.org/loi/swen

Wharton RH, 1989. Control; classical biological control of fruit-infesting Tephritidae, In: Robinson AS, Hooper G, eds. Fruit Flies; their Biology, Natural Enemies and Control. World Crop Pests 3(B). Amsterdam, Netherlands: Elsevier, 303-313.

White IM, Elson-Harris MM, 1992. Fruit flies of economic significance: their identification and bionomics. Wallingford, UK: CAB International, 601 pp.

Wulp FMder, 1900. Family Muscidae. (Biologia Centrali-Americana. Zoologia. Class Insecta. Diptera) In: Contributions to the knowledge of the fauna and flora of Mexico and Central America. Volume 2 [ed. by Godman FD, Salvin O] London, UK: Taylor & Francis, 417-428.

Distribution References

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

Borge M N R, Basedow T, 1997. A survey on the occurrence and flight periods of fruit fly species (Diptera: Tephritidae) in a fruit growing area in southwest Nicaragua, 1994/95. Bulletin of Entomological Research. 87 (4), 405-412. DOI:10.1017/S000748530003741X

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

CABI, Undated a. CABI Compendium: Status as determined by CABI editor. Wallingford, UK: CABI

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

Delmi M, Morán S, Núñez F, Granados G, 1996. The efficiency of bait to attract fruit flies in El Salvador. (Eficiencia de cebos como atrayentes de moscas de la fruta en El Salvador.). Agronomia Mesoamericana. 7 (2), 13-22.

EPPO, 2022. EPPO Global database. In: EPPO Global database, Paris, France: EPPO. 1 pp. https://gd.eppo.int/

Epsky N D, 2003. Development of Lures for detection and Delimitation of Invasive Anastrepha Fruit Flies. [Proceedings of the Caribbean Food Crops Society], 39 (1) Puerto Rico: Caribbean Food Crop Society. 84-89. http://ageconsearch.umn.edu/record/256637/files/1-12.pdf

Eskafi F M, Cunningham R T, 1987. Host plants of fruit flies (Diptera: Tephritidae) of economic importance in Guatemala. Florida Entomologist. 70 (1), 116-123. DOI:10.2307/3495098

Foote R H, Blanc F L, Norrbom A L, 1993. Handbook of the fruit flies (Diptera: Tephritidae) of America north of Mexico. Ithaca, USA: Comstock Publishing Associates. xii + 571 pp.

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. DOI:10.1653/024.093.0421

Hernandez-Ortiz V, 1992. (El género Anastrepha en México). In: Taxonomfa, distribución y sus plantas huéspedes, Xalapa, Mexico: Instituto de Ecologfa.

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