Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Anastrepha fraterculus
(South American fruit fly)

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Datasheet

Anastrepha fraterculus (South American fruit fly)

Summary

  • Last modified
  • 16 November 2021
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Natural Enemy
  • Preferred Scientific Name
  • Anastrepha fraterculus
  • Preferred Common Name
  • South American fruit fly
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Uniramia
  •         Class: Insecta
  • Summary of Invasiveness
  • A. fraterculus has a broad host range, particularly in the family Myrtaceae, but it is also a pest of citrus and apples [Malus domestica] in some areas. It is the most important pest species of Anastrepha in subtropical areas...

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Pictures

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PictureTitleCaptionCopyright
Anastrepha fraterculus, wing markings and venation.
TitleWing
CaptionAnastrepha fraterculus, wing markings and venation.
CopyrightUSDA-ARS
Anastrepha fraterculus, wing markings and venation.
WingAnastrepha fraterculus, wing markings and venation.USDA-ARS
Anastrepha fraterculus, genitalia.
TitleGenitalia
CaptionAnastrepha fraterculus, genitalia.
CopyrightUSDA-ARS
Anastrepha fraterculus, genitalia.
GenitaliaAnastrepha fraterculus, genitalia.USDA-ARS
Anastrepha fraterculus, genitalia.
TitleGenitalia.
CaptionAnastrepha fraterculus, genitalia.
CopyrightUSDA-ARS
Anastrepha fraterculus, genitalia.
Genitalia.Anastrepha fraterculus, genitalia.USDA-ARS
Anastrepha fraterculus, genitalia.
TitleGenitalia.
CaptionAnastrepha fraterculus, genitalia.
CopyrightUSDA-ARS
Anastrepha fraterculus, genitalia.
Genitalia.Anastrepha fraterculus, genitalia.USDA-ARS
Anastrepha fraterculus, adult female.
TitleAdult
CaptionAnastrepha fraterculus, adult female.
CopyrightUSDA-ARS
Anastrepha fraterculus, adult female.
AdultAnastrepha fraterculus, adult female.USDA-ARS

Identity

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Preferred Scientific Name

  • Anastrepha fraterculus (Wiedemann)

Preferred Common Name

  • South American fruit fly

Other Scientific Names

  • Acrotoxa fraterculus (Wiedemann)
  • Anastrepha braziliensis Greene
  • Anastrepha costarukmanii Capoor
  • Anastrepha fraterculus var. soluta Bezzi
  • Anastrepha lambayecae Korytkowski & Ojeda
  • Anastrepha peruviana Townsend
  • Anastrepha pseudofraterculus Capoor
  • Anastrepha scholae Capoor
  • Anthomyia frutalis Weyenburgh
  • Dacus fraterculus Wiedemann
  • Tephritis mellea Walker
  • Trypeta fraterculus (Wiedemann)
  • Trypeta unicolor Loew

International Common Names

  • English: fruit fly, South American
  • Spanish: mosca de la ciruela; mosca de la fruta suramericana; mosca sudamericana de la fruta
  • French: mouche des fruits sud-américaine
  • Portuguese: mosca das frutas sul-americana

Local Common Names

  • Germany: Fruchtfliege, Suedamerikanische

EPPO code

  • ANSTFR (Anastrepha fraterculus)

Summary of Invasiveness

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A. fraterculus has a broad host range, particularly in the family Myrtaceae, but it is also a pest of citrus and apples [Malus domestica] in some areas. It is the most important pest species of Anastrepha in subtropical areas of South America, thus it and Anastrepha ludens may be more of a threat of introduction to other subtropical areas of the world than other species of Anastrepha. It is invasive in the Galapagos Islands. As it is probably a complex of cryptic species whose ranges and delimitation remain unresolved, there is also the threat of introduction of particular cryptic species to other areas within the range of the complex. 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 fraterculus

Notes on Taxonomy and Nomenclature

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This is probably a complex of cryptic species, which has not yet been studied in sufficient detail to permit clear delimitation of the individual species. Data of various types, including isozymes, karyotypes, morphometric analysis of morphology, mating incompatability and pest status (e.g. Steck, 1991, 1999; Alberti et al., 2002; Hernández-Ortiz et al., 2004; Selivon et al., 2005; Vera et al., 2006), indicate that certain populations (e.g. Andean and lowland populations in Venezuela; sympatric populations in southern Brazil) are likely to be distinct species, but comprehensive analysis is needed to resolve the status of other populations from throughout the range of the complex.

The oldest name pertaining to the complex is Dacus fraterculus Wiedemann, 1830. The current combination was proposed by Wulp (1899). Numerous names are currently recognized as synonyms of A. fraterculus, but remain available and may become valid if the complex is split into multiple species.

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

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

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

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

Thorax: mostly yellow to orange-brown, with the following areas yellow to white and often contrasting: postpronotal lobe; single medial and paired sublateral vittae on scutum, the slender medial vitta extended nearly 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 dark-brown spot. Subscutellum and mediotergite dark-brown laterally. Scutum entirely microtrichose or at most with small presutural, medial bare area.

Wing: vein M strongly curved apically. Vein R2+3 nearly straight. Pattern mostly orange-brown and moderate brown. C-band and S-band usually connected along vein R4+5, but sometimes separated; marginal hyaline spot (or end of band) present in cell r1 at apex of vein R4+5. S-band with middle section between costa and vein Cu1 largely yellow to orange with narrow brown margins, darkening distally; distal section of band relatively narrow, well-separated from apex of vein M. V-band with distal arm usually complete and connected to proximal arm; 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 2.7-3.2 mm long; ratio to mesonotum length 0.90-1.10. Glans with basolateral membranous lobe, mostly membranous medially, with isolated, T-shaped apical sclerite.

Female terminalia: oviscape straight, 1.40-2.15 mm long; ratio to mesonotum length 0.59-0.75. Dorsobasal scales of eversible membrane numerous, hook-like, in triangular pattern. Aculeus length 1.50-1.95 mm; tip 0.20-0.30 mm long, 0.12-0.15 mm wide, gradually tapering, but with slight constriction proximal to serrate part, distal 0.50-0.67 serrate. Three spermathecae ovoid.

Immature Stages

Larva: it is very difficult, and in some cases impossible, to identify larvae of Anastrepha species from morphological characteristics. The key by Steck et al. (1990) and the interactive key by Carroll et al. (2004) are the best tools for larval identification. Descriptions of A. fraterculus larvae are provided by Weems (1980), Steck et al. (1990) and White and Elson-Harris (1994). White and Elson-Harris (1994) described the third-instar larva as follows:

Larvae: medium-sized; 8.0-9.5 mm long; 1.4-1.8 mm wide.

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

Thoracic and abdominal segments: anterior margin of T1 with a broad, encircling band of 4-11 discontinuous rows of small, sharply pointed spinules; T2 and T3 with three to seven rows of smaller spinules encircling each segment. Dorsal spinules occasionally on A1-A3, but absent from A4-A8. Creeping welts with 7-12 rows of small spinules. A8 with dorsal and sensilla well-developed; intermediate areas obvious, with large sensilla; ventral sensilla present.

Anterior spiracles: with 14-18 tubules.

Posterior spiracles: spiracular slits about three times as long as broad, with heavily sclerotised, dark-brown rimae. Spiracular hair bundles large; dorsal and ventral bundles of 12-16 long hairs, many branched in apical third; lateral bundles of six to nine hairs similarly branched.

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

Distribution

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A record of A. fraterculus in Texas, USA (Stone, 1942; EPPO, 2014) published in previous versions of the Compendium is unreliable as the original source of the record (Stone, 1942) is old and was published before the fraterculus group and complex was distinguished (Norrbom et al., 1999). A. fraterculus is considered a quarantine pest in the USA and is regulated at ports of entry (PestID, 2016). USDA-APHIS has an ongoing Mexican fruit fly trapping network in southern Texas, which includes traps capable of attracting A. fraterculus, ensuring that it will be detected if it enters the area (USDA-APHIS-PPQ, 2010, 2015) and triggering response plans, including eradication (USDA-APHIS-PPQ, 2016). There have been no detections of A. fraterculus in the USA in six years of surveys from 2011 to 2016 (NAPIS, 2017).

See also CABI/EPPO (1997).

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 Last Reported Origin First Reported Invasive Reference Notes

Europe

SloveniaAbsent

North America

BelizePresent, WidespreadNative
Costa RicaPresent, WidespreadNative
El SalvadorPresent, WidespreadNative
GuatemalaPresent, WidespreadNative
HondurasPresent, WidespreadNative
MexicoPresent, LocalizedNativeLowlands north to Nuevo Leon, absent from northwest
NicaraguaPresent, WidespreadNative
PanamaPresent, WidespreadNative
Trinidad and TobagoPresent, WidespreadNative
United StatesPresent, Localized
-TexasPresent, Localized

Oceania

New ZealandAbsent, Confirmed absent by survey

South America

ArgentinaPresent, LocalizedNativeOccurs south to San Luis
BoliviaPresent
BrazilPresent, WidespreadNativeAbsent from Amazonia
-AlagoasPresentNative
-AmapaPresent
-AmazonasPresent
-BahiaPresentNativeOriginal citation: Malavasi Zucchi (2000)
-CearaPresentNativeOriginal citation: Malavasi Zucchi (2000)
-Espirito SantoPresentNativeOriginal citation: Malavasi Zucchi (2000)
-GoiasPresentNativeOriginal citation: Malavasi Zucchi (2000)
-MaranhaoPresent
-Mato GrossoPresent
-Mato Grosso do SulPresentNativeOriginal citation: Malavasi Zucchi (2000)
-Minas GeraisPresentNativeOriginal citation: Malavasi Zucchi (2000)
-ParaPresent
-ParaibaPresentNativeOriginal citation: Malavasi Zucchi (2000)
-ParanaPresentNative
-PernambucoPresentNativeOriginal citation: Malavasi Zucchi (2000)
-PiauiPresentNativeOriginal citation: Malavasi Zucchi (2000)
-Rio de JaneiroPresentNativeOriginal citation: Malavasi Zucchi (2000)
-Rio Grande do NortePresentNativeOriginal citation: Malavasi Zucchi (2000)
-Rio Grande do SulPresentNativeOriginal citation: Malavasi Zucchi (2000)
-RoraimaPresent
-Santa CatarinaPresentNativeOriginal citation: Malavasi Zucchi (2000)
-Sao PauloPresentNativeOriginal citation: Malavasi Zucchi (2000)
-SergipePresentNative
-TocantinsPresentNative
ChileAbsent, EradicatedEradicated 1964
ColombiaPresent, WidespreadIntroducedInvasive
EcuadorPresent, WidespreadNative and not invasive on the mainland, exotic and invasive on the Galapagos Islands, where it was first recorded in 1979
-Galapagos IslandsPresent
French GuianaPresent
GuyanaPresentNative
ParaguayPresent, WidespreadNative
PeruPresent, WidespreadNative
SurinamePresent
UruguayPresent, WidespreadNative
VenezuelaPresent, WidespreadNative

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
Chile No No Enkerlin et al. (1989) Eradicated after establishment
Galapagos Islands <1979 Yes No Foote (1982); Harper et al. (1989) Probably introduced from mainland Ecuador

Risk of Introduction

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EPPO lists A. fraterculus as an A1 quarantine pest (OEPP/EPPO, 1983) within the broad category 'non European Trypetidae'; it is also of quarantine significance to APPPC, CPPC and NAPPO. A. fraterculus, like the other Anastrepha spp., derives from tropical wet forest habitats and therefore represents a high risk to similar areas.

Consignments of fruits of Annona, Citrus, Fortunella, Malus, mango [Mangifera indica], peach [Prunus persica] and guava [Psidium guajava] from countries where A. fraterculus 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 that such fruits should come from an area where A. fraterculus 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 (e.g. 13, 15 or 17 days at 0.5, 1 or 1.5°C, respectively) or, for certain types of fruits, by vapour heat (for example, keeping at 43°C for 4-6 h) (USDA, 1994), or by hot water immersion (Nascimento et al., 1992). Ethylene dibromide was previously widely used as a fumigant, but is now generally withdrawn because of its carcinogenicity. Carneiro and Salles (1994) showed that an entomopathogenic fungus (Paecilomyces fumosoroseus isolate CG 260) could be used to treat larvae entering soil, which then die upon pupariation.

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

Habitat

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A. fraterculus 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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Pest status varies among populations of the fraterculus complex. The populations in Mexico and Central America are less significant pests (e.g. of guava [Psidium guajava], rose apple [Syzygium jambos] and occasionally peach [Prunus persica] and mango [Mangifera indica], but not citrus or apple [Malus domestica]) than some populations in the Andes, southern Brazil, and Argentina that also attack apples, citrus and a variety of other crops (even blackberries [Rubus fruticosus]). Some populations range into subtropical areas in southern Brazil and Argentina and temperate elevations in the Andes, and are pests in those zones.

The complex as a whole is broadly polyphagous (Norrbom, 2004a), but the preferred hosts are Myrtaceae, particularly the native American guava (P. guajava). Other cultivated mytaceous hosts include other Psidium spp., Campomanesia spp., rose apple, Horn of plenty [Feijoa sellowiana], Surinam cherry [Eugenia uniflora] and other Eugenia spp. Prunus spp., especially peach, and loquat [Eriobotrya japonica] are commonly reported hosts. The most frequent introduced hosts in Mexico are S. jambos and Terminalia catappa [Indian-almond] (Hernandez-Ortiz, 1992). Apple, pear, kumquat [Fortunella spp.], peach, loquat and various Citrus spp. are among the cultivated crops attacked in southern Brazil, although most of the primary hosts are Myrtaceae (Salles, 1995b). Guava, Surinam cherry, grapefruit [Citrus paradisi], cherimoya [Annona cherimola], apricot [Prunus armeniaca], plum [Prunus domestica] and peach are significant hosts in Argentina (Ovruski et al., 2003; Segura et al., 2006).

In common with other polyphagous and difficult to identify species, many host records cannot be substantiated and only records confirmed by Norrbom (2004a) or subsequent reliable sources have been accepted here.

Host Plants and Other Plants Affected

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Plant nameFamilyContextReferences
Acca sellowianaLithomyrtusUnknown
Actinidia chinensis (Chinese gooseberry)ActinidiaceaeOther
Actinidia deliciosa (kiwifruit)ActinidiaceaeOther
Alchornea latifoliaEuphorbiaceaeUnknown
Ampelocera hottleiUlmaceaeOther
Anacardium occidentale (cashew nut)AnacardiaceaeUnknown
Annona cherimola (cherimoya)AnnonaceaeOther
Ovruski et al. (2003); Prezotto et al. (2019); Vera et al. (2006)
Annona muricata (soursop)AnnonaceaeOther
Annona rugulosaUnknown
Annona squamosa (sugar apple)AnnonaceaeOther
Averrhoa carambola (carambola)OxalidaceaeOther
Butia eriospathaArecaceaeOther
Byrsonima crassifolia (wild cherry)MalpighiaceaeUnknown
Calycolpus moritzianusUnknown
CampomanesiaLithomyrtusUnknown
Campomanesia adamantiumLithomyrtusUnknown
Campomanesia espiritosantensisLithomyrtusUnknown
Campomanesia guazumifoliaUnknown
Campomanesia lineatifoliaLithomyrtusUnknown
Campomanesia pubescensUnknown
Campomanesia xanthocarpaLithomyrtusMain
Carica papaya (pawpaw)CaricaceaeOther
Celtis iguanaeaUnknown
Chrysophyllum gonocarpumSapotaceaeMain
CitrusRutaceaeUnknown
Araujo et al. (2019); Prezotto et al. (2019)
Citrus aurantium (sour orange)RutaceaeOther
Citrus limetta (sweet lemon tree)RutaceaeOther
Citrus maxima (pummelo)RutaceaeOther
Citrus reticulata (mandarin)RutaceaeOther
Citrus sinensis (sweet orange)RutaceaeOther
Citrus x paradisi (grapefruit)RutaceaeOther
Coffea arabica (arabica coffee)RubiaceaeOther
Coffea liberica (Liberian coffee tree)RubiaceaeOther
Crataegus (hawthorns)RosaceaeUnknown
Cryptocarya aschersonianaLauraceaeUnknown
Cydonia oblonga (quince)RosaceaeOther
Diospyros kaki (persimmon)EbenaceaeOther
Diospyros malabaricaEbenaceaeOther
Eriobotrya japonica (loquat)RosaceaeMain
EugeniaLithomyrtusUnknown
Eugenia brasiliensis (brazil cherry)LithomyrtusMain
Eugenia dysentericaLithomyrtusUnknown
Araujo and Zucchi (2006); Silva et al. (2010)
Eugenia involucrataLithomyrtusMain
Eugenia lambertianaUnknown
Eugenia myrcianthesUnknown
Eugenia platyphyllaUnknown
Eugenia platysemaUnknown
Eugenia pyriformisLithomyrtusMain
Eugenia stipitataLithomyrtusOther
Eugenia uniflora (Surinam cherry)LithomyrtusMain
Feijoa sellowiana (Horn of plenty)LithomyrtusMain
Ficus carica (common fig)MoraceaeOther
Fortunella japonica (round kumquat)RutaceaeMain
Fragaria ananassa (strawberry)RosaceaeUnknown
Fragaria vesca (wild strawberry)RosaceaeOther
Garcinia brasiliensisClusiaceaeUnknown
Hexachlamys edulisLithomyrtusUnknown
IngaFabaceaeUnknown
Inga edulis (ice-cream bean)FabaceaeOther
Inga marginataFabaceaeOther
Inga michelianaUnknown
Inga paternoFabaceaeUnknown
Inga sellowianaUnknown
Juglans australisJuglandaceaeOther
Gómez-Cendra et al. (2016); Ovruski et al. (2003)
Juglans neotropica (andean walnut)JuglandaceaeOther
Juglans regia (walnut)JuglandaceaeOther
MalpighiaMalpighiaceaeUnknown
Malpighia emarginataMalpighiaceaeUnknown
Malpighia glabra (acerola)MalpighiaceaeUnknown
Araujo et al. (2019); Leal et al. (2009)
Malus (ornamental species apple)RosaceaeOther
Malus domestica (apple)RosaceaeOther
Mangifera indica (mango)AnacardiaceaeOther
Manilkara zapota (sapodilla)SapotaceaeOther
Araujo et al. (2019); Raga et al. (2011)
Mosiera longipesLithomyrtusUnknown
Mouriri acutifloraUnknown
Mouriri glaziovianaUnknown
Myrcianthes pungensLithomyrtusUnknown
Myrciaria cauliflora (jaboticaba)LithomyrtusUnknown
Myrciaria floribundaLithomyrtusUnknown
Myrciaria glaziovianaUnknown
Araujo et al. (2019)
Myrciaria glomerataUnknown
Myrciaria strigipesUnknown
Olea europaea subsp. europaea (European olive)OleaceaeOther
Passiflora alataPassifloraceaeOther
Passiflora caerulea (blue passionflower)PassifloraceaeUnknown
Passiflora edulis (passionfruit)PassifloraceaeUnknown
Araujo et al. (2019)
Passiflora tripartita var. mollissima (banana passionfruit)PassifloraceaeUnknown
Vera et al. (2006)
Peritassa campestrisSalaciaUnknown
Araujo and Zucchi (2006)
Persea americana (avocado)LauraceaeOther
Araujo et al. (2019); Putruele (1996)
PicramniaUnknown
Plinia caulifloraUnknown
Plinia edulisLithomyrtusUnknown
Pouteria caimitoSapotaceaeOther
Araujo et al. (2019); Melo et al. (2012); Raga et al. (2011)
Pouteria gardnerianaSapotaceaeUnknown
Araujo and Zucchi (2006)
Pouteria obovataSapotaceaeOther
Pouteria sapota (mammey sapote)SapotaceaeUnknown
Prunus (stone fruit)RosaceaeMain
Araujo et al. (2019); Malavasi et al. (1980)
Prunus armeniaca (apricot)RosaceaeOther
Ovruski et al. (2003); Putruele (1996)
Prunus avium (sweet cherry)RosaceaeUnknown
Prunus domestica (plum)RosaceaeOther
Prunus dulcis (almond)RosaceaeOther
Prunus mume (Japanese apricot tree)RosaceaeUnknown
Prunus persica (peach)RosaceaeMain
Prunus serotina (black cherry)RosaceaeUnknown
Psidium (guava)LithomyrtusUnknown
Araujo and Zucchi (2006)
Psidium acutangulumLithomyrtusUnknown
Psidium cattleianum (strawberry guava)LithomyrtusMain
Psidium guajava (guava)LithomyrtusMain
Psidium guineense (Guinea guava)LithomyrtusMain
Psidium kennedyanumUnknown
Psidium myrtoidesUnknown
Psidium sartorianumLithomyrtusUnknown
Punica granatum (pomegranate)PunicaceaeOther
Pyrus communis (European pear)RosaceaeOther
Quassia guianensisSimaroubaceaeOther
Rollinia laurifoliaUnknown
Rollinia sericeaAnnonaceaeUnknown
Rubus (blackberry, raspberry)RosaceaeUnknown
Araujo et al. (2019); Manni et al. (2015)
Rubus fruticosus (blackberry)RosaceaeUnknown
Rubus glaucusRosaceaeOther
Rubus idaeus (raspberry)RosaceaeUnknown
Rubus ulmifoliusRosaceaeUnknown
Sideroxylon capiri subsp. tempisqueUnknown
Solanum (nightshade)SolanaceaeUnknown
Solanum quitoense (naranjilla)SolanaceaeOther
Spondias (purple mombin)AnacardiaceaeUnknown
Spondias dulcis (otaheite apple)AnacardiaceaeOther
Araujo et al. (2019)
Spondias mombin (hog plum)AnacardiaceaeOther
Spondias purpurea (red mombin)AnacardiaceaeOther
Spondias tuberosaAnacardiaceaeMain
Syagrus romanzoffiana (queen palm)ArecaceaeUnknown
Araujo et al. (2019)
Syzygium jambos (rose apple)LithomyrtusMain
Syzygium malaccense (Malay apple)LithomyrtusOther
Syzygium uniflorumUnknown
Talisia olivaeformisSapindaceaeOther
Terminalia catappa (Singapore almond)CombretaceaeOther
Theobroma cacao (cocoa)MalvaceaeOther
Vitis vinifera (grapevine)VitaceaeOther
Ziziphus joazeiroRhamnaceaeMain
Zuelania guidoniaFlacourtiaceaeOther

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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A. fraterculus eggs are laid below the skin of the host fruit. Many Anastrepha spp. lay their eggs deeper inside the fruit or in the seeds. The life cycle includes: the egg, three larval stages, pupa and adult. Salles (2000) presented a table showing the length of development at temperatures from 15-30°C. At 25°C the eggs hatch in 2.6-3.2 days and the larvae feed for another 11-14 days (up to 34.5 days at 15°C). Pupariation is in the soil under the host plant and the adults emerge after 10-15 days (43.2 days at 15°C) and may live up to 161 days in laboratory conditions. Salles (2000) reported that females can produce 278-437 eggs. The adults occur throughout the year (Christenson and Foote, 1960). They have no winter diapause or quiescence in more temperate areas such as southern Brazil (Salles, 1993). Reproductive behaviour in the laboratory and field has been studied by Lima et al. (1994).

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)
26 35

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Aganaspis pelleranoi Parasite
Coptera haywardi Parasite
Doryctobracon areolatus Parasite Arthropods|Larvae; Arthropods|Pupae
Doryctobracon brasiliensis Parasite Arthropods|Larvae
Opius bellus Parasite Arthropods|Larvae
Pachycrepoideus vindemmiae Parasite
Trichopria anastrephae Parasite Cruz et al. (2011)
Utetes anastrephae Parasite Arthropods|Larvae

Notes on Natural Enemies

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Average parasitism ranged from 0.44 to 29.3% (Aluja et al., 1990). For further information, see Loiacono (1981) and Salles (1996).

Means of Movement and Dispersal

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There is evidence that adults of Anastrepha spp. can fly up to 135 km (Fletcher, 1989) and therefore natural movement can be 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. For most regions, the most important fruits liable to carry A. fraterculus are guavas [Psidium guajava] and other Myrtaceae; Citrus, Malus and Prunus are attacked in some areas. 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
Clothing, footwear and possessionsFruit in case or handbag. Yes
Containers and packaging - woodOf fruit cargo. Yes
Land vehiclesAeroplanes and boats, with 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) arthropods/eggs; arthropods/larvae; arthropods/pupae Yes Pest or symptoms not visible to the naked eye but usually visible under light microscope
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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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 introduced Ceratitis capitata (EPPO/CABI, 1996). A. fraterculus is an important pest of guavas [Psidium guajava] (and locally significant Myrtaceae) and mangoes [Mangifera indica], and also to some extent of Citrus and Prunus spp. (Hernandez Ortiz, 1992; White and Elson Harris, 1994). Some populations in South America, particularly in southern Brazil to northern Argentina, and in the Andean countries, are considered more significant pests than those in Mexico and Central America, and these probably represent different cryptic species.

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

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. McPhail traps are usually used for the capture of Anastrepha spp. (White and Elson-Harris, 1994) 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 approximately 13% of the released flies.

Some studies have shown that egg morphology can be used to separate closely related species found in host fruits (Souza et al., 1983; Murillo and Jiron, 1994). The larvae of some species may also be differentiated using cuticular hydrocarbons (Sutton and Carlson, 1993). Neither method has yet been generalized for application outside of very specific circumstances.

Similarities to Other Species/Conditions

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Identification of the fraterculus complex and closely related species is very difficult and is based mainly on measurements and subtle differences in the shape of the aculeus and its tip (Araujo et al., 1996; Araujo and Zucchi, 2006). The species most likely to be confused with A. fraterculus are Anastrepha obliqua, Anastrepha suspensa, and especially Anastrepha sororcula, Anastrepha zenildae and Anastrepha turpiniae.

A. obliqua differs in having the subscutellum entirely orange, not dark-brown laterally. It also lacks the medial spot on the scuto-scutellar suture that is usually present in the fraterculus complex. The aculeus tip is more serrate and less tapered basal to the serrate part. A. suspensa has the apical part of the S-band (the band on the anterior apical margin of the wing) much broader and touching or almost touching the apex of vein M. The other three species are distinguished from the fraterculus complex by the length of the aculeus and its tip, although there is slight overlap in these characters among these species. A. sororcula has a shorter aculeus, with a shorter, stouter tip, whereas A. zenildae and A. turpiniae have longer aculeus tips.

The larvae of Anastrepha are extremely difficult to identify and specialist help should be sought to confirm critical identifications. The third-stage larva is very similar to those of A. obliqua and A. suspensa, and these species usually cannot be distinguished (Steck et al., 1990). The larvae of A. sororcula, A. zenildae and A. turpiniae have not been described.

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). 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 in that they can be applied as a spot treatment so that the flies are attracted to the insecticide and there is minimal impact on most natural enemies. However, Nasca et al. (1983) warned that bait sprays could have a substantial effect on Chrysopidae (Neuroptera), which are natural enemies of many pests.

The toxicity to A. fraterculus of different insecticides used in baits was recently compared by Salles (1995) and Lerenzato et al. (1984). They advocated control measures be applied when 0.5-1.0 flies per day per trap were found. Like many fruit flies, shape and colour play a role in host seeking behaviour and Cytrynowicz et al. (1982) found a preference for yellow spheres. The sterile insect technique was tried in Peru (Gonzalez et al., 1971), but has not been applied on a realistic scale.

References

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Caetano F H, Solferini V N, Britto F B de, Lins D S, Aluani T, Brito V G de, Zara F J, 2006. Ultra morphology of the digestive system of Anastrepha fraterculus and Ceratitis capitata (Diptera Tephritidae). Brazilian Journal of Morphological Sciences. 23 (3/4), 455-462.

Calvo M V, Delgado S, Scatoni I, Mello Garcia F R, 2017. First report of Neosilba pradoi and Dasiops frieseni (Diptera: Lonchaeidae) in cultivated and wild hosts in Uruguay. Florida Entomologist. 100 (4), 831-832. DOI:10.1653/024.100.0427

Caraballo J, 1981. Fruit flies of the genus Anastrepha Schiner, 1868 (Diptera: Tephritidae) in Venezuela. (Las moscas de frutas del genero Anastrepha Schiner, 1868 (Diptera: Tephritidae) de Venezuela.). Maracay, Venezuela: Universidad Central de Venezuela. xi + 210 pp.

Cardoso V V, Ferreira M P, Montagner J M, Fernandez C G, Moreira J C, Oliveira A K, 2002. The effects of constant and alternating temperatures on the reproductive potential, life span, and life expectancy of Anastrepha fraterculus (Wiedemann) (Diptera: Tephritidae). Brazilian Journal of Biology. 62 (4B), 775-786. DOI:10.1590/S1519-69842002000500006

Cardoso V V, Prestes P R, Casali E A, Moreira J C F, Oliveira A K, 2004. Ornithine decarboxylase activity during the development of Anastrepha fraterculus (Diptera, Tephritidae). Archives of Insect Biochemistry and Physiology. 57 (4), 151-159. DOI:10.1002/arch.20022

Destéfano R H R, Bechara I J, Messias C L, Piedrabuena A E, 2005. Effectiveness of Metarhizium anisopliae against immature stages of Anastrepha fraterculus fruitfly (Diptera: Tephritidae). Brazilian Journal of Microbiology. 36 (1), 94-99. DOI:10.1590/S1517-83822005000100018

Enkerlin D, Garcia R L, Lopez F, 1989. Mexico, Central and South America. In: Fruit flies: their biology, natural enemies and control. [ed. by Robinson A S, Hooper G]. Amsterdam, Netherlands: Elsevier Science Publishers B. V. 83-90.

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

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

Feitosa S S, Silva P R R, Pádua L E de M, Sousa M P da S, Passos E P de, Soares A A R A, 2007. First register of fruit flies (Diptera: Tephritidae) in star fruit in Teresina, Altos and Parnaiba, state of Piaui, Brazil. (Primeiro registro de moscas-das-frutas (Diptera: Tephritidae) em carambola nos municípios de Teresina, Altos e Parnaíba no estado do Piauí.). Semina: Ciencias Agrárias (Londrina). 28 (4), 629-634. http://www.uel.br/proppg/semina/pdf/semina_28_4_19_10.pdf

Foote R H, 1982. The Tephritidae (Diptera) of the Galapagos Archipelago. Memoirs of the Entomological Society of Washington. 48-55.

Garcia F R M, Campos J V, Corseuil E, 2003. Population fluctuation of Anastrepha fraterculus (Wiedemann, 1830) (Diptera, Tephritidae) in the Western Region of Santa Catarina State, Brazil. (Flutuação populacional de Anastrepha fraterculus (Wiedemann, 1830) (Diptera, Tephritidae) na Região Oeste de Santa Catarina, Brasil.). Revista Brasileira de Entomologia. 47 (3), 415-420. DOI:10.1590/S0085-56262003000300009

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

Goncalves G B, dos Santos J C G, da Silva C E, dos Santos E S, do Nascimento R R, Santana A E G, Zucchi RA, 2006. Occurrence Of Fruit Flies (Diptera: Tephritidae) In The State Of Alagoas, Brazil. Florida Entomologist. 89 (1), 93-94. DOI:10.1653/0015-4040(2006)89[93:OOFFDT]2.0.CO;2

Gonçalves G B, Santos J C G dos, Silva C E da, Santos E S dos, Nascimento R R do, Sant'ana A E G, Zucchi R A, 2006. Occurrence of fruit flies (Diptera: Tephritidae) in the state of Alagoas, Brazil. Florida Entomologist. 89 (1), 93-94. DOI:10.1653/0015-4040(2006)89[93:OOFFDT]2.0.CO;2

Harper J D, Escobar J S, Cereceda G, 1989. Collection of Anastrepha fraterculus on Santa Cruz Island, Galápagos Province, Ecuador. Florida Entomologist. 72 (1), 205-206. DOI:10.2307/3494984

Hernández-Ortiz V, 1992. El género Anastrepha en México. Taxonomia, distribución y sus plantas huéspedes. Xalapa, Mexico: Instituto de Ecologia.

Hernández-Ortiz V, Canal N A, Salas J O T, Ruíz-Hurtado F M, Dzul-Cauich J F, 2015. Taxonomy and phenotypic relationships of the Anastrepha fraterculus complex in the Mesoamerican and Pacific Neotropical dominions (Diptera, Tephritidae). ZooKeys. 95-124. http://zookeys.pensoft.net/articles.php?id=6027

Hernández-Ortiz V, Gómez-Anaya J A, Sánchez A, McPheron B A, Aluja M, 2004. Morphometric analysis of Mexican and South American populations of the Anastrepha fraterculus complex (Diptera: Tephritidae) and recognition of a distinct Mexican morphotype. Bulletin of Entomological Research. 94 (6), 487-499. http://www.ingenta.com/journals/browse/cabi/ber DOI:10.1079/BER2004325

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

Katiyar K P, Camacho Molina J, Matheus R, 2000. Fruit flies (Diptera: Tephritidae) infesting fruits of the genus Psidium (Myrtaceae) and their altitudinal distribution in western Venezuela. Florida Entomologist. 83 (4), 480-486. DOI:10.2307/3496724

Klesener D F, Menezes Júnior A de O, Santos R S S dos, 2016. Occurrence and population fluctuation of insects-pest in apple orchards in north Paraná. (Ocorrência e flutuação populacional de insetos-praga em pomares de macieira da região norte do Paraná.). Revista de la Facultad de Agronomía (La Plata). 115 (2), 201-208. http://www.agro.unlp.edu.ar/revista/index.php/revagro/article/view/408/417

Korytkowski C A, 2001. Status of the genus Anastrepha Schiner, 1868 (Diptera: Tephritidae) in Peru. (Situación actual del género Anastrepha Schiner, 1868 (Diptera: Tephritidae) en el Perú.). Revista Peruana de Entomología. 97-158.

Leal M R, Souza S A da S, Aguiar-Menezes E de L, Lima Filho M, Menezes E B, 2009. Fruit fly diversity, their host plants and their parasitoids in the northern and northwestern regions of Rio de Janeiro State, Brazil. (Diversidade de moscas-das-frutas, suas plantas hospedeiras e seus parasitóides nas regiões Norte e Noroeste do Estado do rio de Janeiro, Brasil.). Ciência Rural. 39 (3), 627-634. DOI:10.1590/S0103-84782009005000002

Lemos L do N, Deus E da G de, Nascimento D B do, Jesus-Barros C R de, Costa Neto S V da, Adaime R, 2017. Species of Anastrepha (Diptera: Tephritidae), their host plants, and parasitoids in small fruit production areas in the state of Amapá, Brazil. Florida Entomologist. 100 (2), 403-410. DOI:10.1653/024.100.0201

Lopes G N, Arias O R, Cônsoli F L, Zucchi R A, 2013. The identity of specimens of the Anastrepha fraterculus complex (Diptera, Tephritidae) with atypical aculeus tip. Neotropical Entomology. 42 (6), 618-627. DOI:10.1007/s13744-013-0162-0

Maciel A A S, Lemos R N S de, Araújo A A R, Machado K K G, Silva E A da, Araújo J R G, Mesquita M L R, 2017. Diversity and infestation indices of fruit flies (Diptera: Tephritidae) in guava (Psidium guajava L.). African Journal of Agricultural Research. 12 (24), 2087-2092. DOI:10.5897/ajar2016.11973

Manni M, Lima K M, Guglielmino C R, Lanzavecchia S B, Juri M, Vera T, Cladera J, Scolari F, Gomulski L, Bonnizoni M, Gasperi G, Silva J G, Malacrida A R, 2015. Relevant genetic differentiation among Brazilian populations of Anastrepha fraterculus (Diptera, Tephritidae). ZooKeys. 157-173. http://zookeys.pensoft.net/articles.php?id=6713&display_type=element&element_type=7&element_id=0&element_name=Anastrepha

Manuel G, Manuel L-M, Smeltekop H, Nicanor C, Carlos A J, Marin R, 2012. Adult population dynamics of the bolivian fruit flies Anastrepha sp. (Diptera: Tephritidae) at Municipality Coroico, Department of The La Paz, Bolivia. (Dinámica poblacional de adultos de la mosca boliviana de la fruta Anastrepha sp. (Díptera: Tephritidae) en el Municipio de Coroico, Departamento de La Paz, Bolivia). Journal of the Selva Andina Research Society. 2 (2), 2-12. https://www.ingentaconnect.com/content/doaj/20729294/2011/00000002/00000002/art00002

Molineros J, Tigrero J O, Sandoval D, 1992. Status of the fruit fly problem in Ecuador. (Diagnostico de la situacion actual del problema de las moscas de la fruta en el Ecuador.). In: Diagnostico de la situacion actual del problema de las moscas de la fruta en el Ecuador. Quito, Ecuador: Comision Ecuatoriana de Energia Atomica, Direccion de Investigaciones. 53 pp.

Norrbom A L, 2004. Host plant database for Anastrepha and Toxotrypana (Diptera: Tephritidae: Toxotrypanini). In: Diptera Data Dissemination Disk,

Núñez Bueno L, 1981. Contribution to a survey of the fruit flies (Diptera: Tephriridae [sic]) of Colombia. (Contribucion al reconocimiento de las moscas de las frutas (Diptera: Tephriridae [sic]) en Colombia.). Revista do Instituto Colombiano Agropecuario, Bogotá. 173-179.

Oroño L E, Ovruski S M, Norrbom A L, Schliserman P, Colin C, Martin C B, 2005. Two new native host plant records for Anastrepha fraterculus (Diptera: Tephritidae) in Argentina. Florida Entomologist. 88 (2), 228-232. DOI:10.1653/0015-4040(2005)088[0228:TNNHPR]2.0.CO;2

Ovruski S M, Oroño L E, Schliserman P, Nuñez-Campero S, 2007. The effect of four fruit species on the parasitization rate of Anastrepha fraterculus (Diptera: Tephritidae, Trypetinae) by Diachasmimorpha longicaudata (Hymenoptera: Braconidae, Opiinae) under laboratory rearing conditions. Biocontrol Science and Technology. 17 (9/10), 1079-1085. DOI:10.1080/09583150701661620

Ovruski S M, Schliserman P, Nieuwenhove G A van, Bezdjian L P, Núñez-Campero S, Albornoz-Medina P, 2010. Occurrence of Ceratitis capitata and Anastrepha fraterculus (Diptera: Tephritidae) on cultivated, exotic fruit species in the highland valleys of Tucuman in Northwest Argentina. Florida Entomologist. 93 (2), 277-282. DOI:10.1653/024.093.0219

Ovruski S, Schliserman P, Aluja M, 2003. Native and introduced host plants of Anastrepha fraterculus and Ceratitis capitata (Diptera: Tephritidae) in Northwestern Argentina. Journal of Economic Entomology. 96 (4), 1108-1118. DOI:10.1603/0022-0493-96.4.1108

Savaris M, Lampert S, Marsaro-Júnior A L, Adaime R, Souza Filho M F de, 2013. First record of Anastrepha fraterculus and Ceratitis capitata (Diptera, Tephritidae) on Arecaceae in Brazil. Florida Entomologist. 96 (4), 1597-1599. http://www.fcla.edu/FlaEnt/ DOI:10.1653/024.096.0445

Scally M, Into F, Thomas D B, Ruiz-Arce R, Barr N B, Schuenzel E L, 2016. Resolution of inter and intra-species relationships of the West Indian fruit fly Anastrepha obliqua. Molecular Phylogenetics and Evolution. 286-293. DOI:10.1016/j.ympev.2016.04.020

Schliserman P, Ovruski S M, Coll O R de, Wharton R, 2010. Diversity and abundance of hymenopterous parasitoids associated with Anastrepha fraterculus (Diptera: Tephritidae) in native and exotic host plants in Misiones, Northeastern Argentina. Florida Entomologist. 93 (2), 175-182. DOI:10.1653/024.093.0205

Segura D F, Vera M T, Cagnotti C L, Vaccaro N, Coll O de, Ovruski S M, Cladera J L, 2006. Relative abundance of Ceratitis capitata and Anastrepha fraterculus (Diptera: Tephritidae) in diverse host species and localities of Argentina. Annals of the Entomological Society of America. 99 (1), 70-83. http://docserver.esa.catchword.org/deliver/cw/pdf/esa/freepdfs/00138746/v99n1s9.pdf DOI:10.1603/0013-8746(2006)099[0070:RAOCCA]2.0.CO;2

Silva J G, Dutra V S, Santos M S, Silva N M O, Vidal D B, Nink R A, Guimarães J A, Araujo E L, 2010. Diversity of Anastrepha spp. (Diptera: Tephritidae) and associated braconid parasitoids from native and exotic hosts in Southeastern Bahia, Brazil. Environmental Entomology. 39 (5), 1457-1465. DOI:10.1603/EN10079

Silva W R da, Silva R A da, 2007. Survey of fruit flies and its parasitoids in Ferreira Gomes, Amapá State. (Levantamento de moscas-das-frutas e de seus parasitóides no município de Ferreira Gomes, Estado do Amapá.). Ciência Rural. 37 (1), 265-268. DOI:10.1590/S0103-84782007000100043

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Stone A, 1942. New species of Anastrepha and notes on others. Journal of the Washington Academy of Sciences. 298-304.

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Uramoto K, Martins D S, Zucchi R A, 2008. Fruit flies (Diptera, Tephritidae) and their associations with native host plants in a remnant area of the highly endangered Atlantic Rain Forest in the State of Espírito Santo, Brazil. Bulletin of Entomological Research. 98 (5), 457-466. DOI:10.1017/S0007485308005774

Vera MT, Caceres C, Wornoayporn V, Islam A, Robinson AS, De La Vega MH, Hendrichs J, Cayol JP, 2006. Mating Incompatibility Among Populations of the South American Fruit Fly Anastrepha fraterculus (Diptera: Tephritidae). Annals of the Entomological Society of America. 99 (2), 387-397. DOI:10.1603/0013-8746(2006)099[0387:MIAPOT]2.0.CO;2

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Zucchi R A, 1978. Taxonomy of Anastrepha Schiner species, 1868 (Diptera, Tephritidae) present in Brazil. (Taxonomia das espécies de Anastrepha Schiner, 1868 (Diptera, Tephritidae) assinaladas no Brasil.). Piracicaba, Brazil: Universidade de São Paulo. vi + 105 pp.

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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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