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Anastrepha Schiner (Diptera: Tephritidae)

Anastrepha pulchella female, dorsal habitus; Annastrepha striata, wing.

Recognition
Most species of Anastrepha can be easily distinguished from other Tephritidae by the strongly anteriorly curved apex of vein M, which in most species meets the costa without a distinct angle.  In a few species, the curve is not as strong, althougth the distal half of the last section of M is always curved anteriorly.   There is overlap in this character with Toxotrypana, which differs in having many of the thoracic setae (postpronotal, presutural supra-alar, dorsocentral, acrostichal and scutellar setae) reduced or absent, the scutum with a median longitudinal furrow, and the abdomen petiolate. Additional useful diagnostic characters for Anastrepha include: Ocellar seta short and weak (except in A. tripunctata); dorsocentral seta much closer to level of postalar seta than to level of postsutural supra-alar seta; wing usually with pattern including C-, S-, and V-bands (see description), although some species have parts of these bands reduced or fused, and others have a wasp mimic pattern, with only a complete costal band and a streak in the cubital cells; lateral surstylus short (less than height of epandrium), usually flattened; glans with basal membranous lobe with minute spicules (except in spp. of dentata and daciformis groups, which lack glans); oviscape tube-shaped, often elongate, basally with flangelike lateral lobe; eversible membrane enlarged basally, this area dorsally with enlarged, toothlike scales; aculeus long and slender, well sclerotized; 3 spermathecae.
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Classification and Evolutionary Relationships
Order: Diptera. Family: Tephritidae. Subfamily: Trypetinae. Tribe: Toxotrypanini. Genus: Anastrepha. Author: Schiner.
The phylogenetic relationships of Anastrepha to other fruit flies were discussed by Norrbom et al. (1999) and McPheron et al. (1999) (see Phylogeny of Anastrepha and Toxotrypana). The status of Anastrepha as a natural (i.e., monophyletic) group is uncertain.  It and the genus Toxotrypana Gerstaecker clearly form a natural group, but there is some question as to whether some species now included in Anastrepha may be more closely related to Toxotrypana. Synapomorphies (shared derived characters) indicating the relationship of these two genera include: 1) the eversible membrane of the female is enlarged basally and bears a group of enlarged dorsal teeth (the teeth are secondarily reduced in some species of the A. dentata group, but still larger than in this area in other tephritids, and the basal area is still enlarged); 2) the base of the oviscape of the female has lateral, flange-like lobes (smaller in Toxotrypana); 3) the male glans is weakly sclerotized medially and has a T-shaped, somewhat hook-like apical sclerite; 4) the surstyli are relatively short, the lateral surstylus without anterior or posterior lobes; and 5) vein M is anteriorly curved in the distal half of its last segment (distal to DM-Cu). Other characters of uncertain polarity, such as the strongly sclerotized secondary connection of the subepandrial sclerite, the number, size and arrangement of the sensilla of the aculeus tip (3 pairs, none extended beyond lateral margin), and the posterior location of the dorsocentral seta, are similar in Toxotrypana and Anastrepha. These similarities do not contradict the hypothesis that Anastrepha and Toxotrypana form a monophyletic group, and some or all may be additional synapomorphies. Analysis of mitochondrial DNA sequences further supported the close relationship of these two genera (McPheron et al. 1999).

The phylogenetic relationships within Anastrepha were also analyzed by Norrbom et al. (1999) and McPheron et al. (1999), who discussed the morphological and molecular (mitochondrial DNA) character data, respectively, supporting each of the 18 species groups recognized by the former authors, as well as the relationships among them.  Some of these groups are well supported as natural (i.e., monophyletic) groups, whereas the status of others remains uncertain.
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Diversity & Distribution
Anastrepha includes 198 currently recognized species as well as numerous other undescribed species. Their combined range includes the southern Nearctic Region (north to southern Texas and Florida) and all of the Neotropical Region, except Chile, southern Argentina, and several of the Lesser Antilles.
Click here for information on species identification.
Click here for list of included species and links to species pages.
Click here for character matrix for species.

Names Used for this Genus
Anastrepha Schiner 1868: 263, type species Dacus serpentinus Wiedemann (OD).
Acrotoxa Loew 1873: 227, type species Dacus fraterculus Wiedemann, Bezzi 1909: 280 (SD).
Pseudodacus Hendel 1914: 66, type species Anastrepha daciformis Bezzi (OD). Proposed as a subgenus.
Phobema Aldrich 1925: 7, type species atrox Aldrich (OD).
Lucumaphila Stone 1939: 340, type species sagittata Stone (OD).
Instrypetas Herrera, Rangel & Barreda 1900: 5, Nomen nudum.
Anastrepa Cresson 1908: 98, missp. Anastrepha Schiner.
Instrypeta Foote 1967: 11, missp. Instrypetas Herrera, Rangel & Barreda.
Anastraphe Hardy 1968: 145, missp. Anastrepha Schiner.
Click here to link to fly names database for full nomenclatural data.

Biology
There is an extensive literature on the biology of Anastrepha species, although much of the information is based on a handful of pest species.  Considerable information can be found in general references such as Aluja (1994), Aluja & Liedo (1993), Aluja & Norrbom (1999), Cavalloro (1983, 1989), Economopoulos (1987), Malavasi & Zucchi (2000), McPheron & Steck (1996), Robinson & Hooper (1989) and references cited within. 

Anastrepha species breed in fruit, either in the pulp and/or the seeds, except for one species (A. manihoti) which attacks buds and stems.  The host range for the entire genus is very broad, with host plants reported from 143 genera in 54 families (see Norrbom, in press, for a comprehensive database of host plant records), although the range is narrower if exotic host plants are excluded.  Although there are generalist species, especially in the fraterculus species group, which includes the majority of the pest species, most species individually attack a fairly narrow range of plants.  Except for single species in the serpentina, striata, leptozona and pseudoparallela groups, feeding on more than a few related hosts or 1-2 families by a single Anastrepha species is rare outside of the fraterculus group. Many species, including the majority of those in the primitive clades, breed in fruits of latex-bearing plants, especially Sapotaceae. Certain species groups are associated mainly with other plant families, such as the spatulata group on Euphorbiaceae and Olacaceae, the pseudoparallela group on Passifloraceae, the grandis group (or at least the one species of the group whose hosts are known) on Cucurbitaceae, the striata group on Myrtaceae, and some species of the mucronota group on Bombacaceae.

In most published host plant records for Anastrepha, the part of the fruit attacked has not been mentioned. At least in the case of A. sagittata (Stone) and A. hamata (Loew) (dentata group) and A. katiyari Norrbom and A. pallens Coquillett (daciformis group), the larva feeds exclusively in the seed (McPhail & Berry 1936, Baker et al. 1944, Norrbom et al. 1999, Aluja et al. 1999). Larvae of A. pseudoparallela (pseudoparallela group), and A. montei Lima and A. pickeli Lima (spatulata group) feed on developing seeds and associated tissues (Morgante et al. 1996, Stefani & Morgante 1996) similar to species of Toxotrypana. Larvae of A. cordata Aldrich (cryptostrepha group) feed mainly on the seeds and later on the pulp of the fruit, apparently because immature fruits have large quantities of latex (Hernández-Ortiz & Pérez-Alonso 1993). Larvae of A. crebra Stone (mucronota group) have been found feeding in the pulp and seeds at same time (Hernández-Ortiz & Pérez-Alonso 1993), and those of A. steyskali Korytkowski (leptozona group), A. anomala Stone and A. serpentina (Wiedemann) (serpentina group), and A. ludens (Loew) (fraterculus group) can feed on both seeds and the fleshy mesocarp (Stone 1942, Korytkowski 1974, Aluja et al. 1999). Conversely, in most of the generalist species, such as A. obliqua (Macquart), fraterculus (Wiedemann), striata Schiner and distincta Greene, feeding is primarily or exclusively on the mesocarp (Aluja 1994).

Economic Significance
Anastrepha is the most economically important genus in the American tropics and subtropics. White & Elson-Harris (1992) listed 15 species as significant pests and 28 others that have been reported to attack economically important plants. The worst pest species are the Mexican fruit fly (A. ludens (Loew)), the West Indian fruit fly (A. obliqua (Macquart)), the South American fruit fly (A. fraterculus (Wiedemann) complex), the Caribbean fruit fly (A. suspensa (Loew)), A. sororcula Zucchi, A. serpentina (Wiedemann), A. striata Schiner, the South American cucurbit fruit fly (A. grandis (Macquart)), and the Inga fruit fly (A. distincta Greene).

References
Click here to access fruit fly literature database
Aluja, M. 1994. Bionomics and management of Anastrepha. Annu. Rev. Entomol. 39: 155-178.
Aluja S., M. & P. Liedo F., eds. 1993. Fruit flies: Biology and management. Proceedings of the International Symposium on Fruit Flies of Economic Importance, 1990, Antigua Guatemala, Guatemala, October 14-20, 1990. Springer-Verlag, New York. xxxiii + 492 p.
Aluja, M. & A. L. Norrbom, eds. 1999. Fruit Flies (Tephritidae): Phylogeny and Evolution of Behavior. CRC Press, Boca Raton.
Baker, A. C., Stone, W. E., Plummer, C. C. & McPhail, M. 1944. A review of studies on the Mexican fruitfly and other related Mexican species. U. S. Dept. Agric. Misc. Publ. No. 531. 155 pp.
Blanchard, E. E. 1961. Especies argentinas del género Anastrepha Schiner (sens. lat.) (Diptera, Trypetidae). Rev. Invest. Agric. 15 (2): 281-342. [monograph of 38 spp. in Argentina]
Bush, G. L. 1962. The cytotaxonomy of the larvae of some Mexican fruit flies in the genus Anastrepha (Tephritidae, Diptera). Psyche (Camb.) 68: 87-101. [cytotaxonomy of 9 spp.]
Caraballo, J. 1981. Las moscas de frutas del genero Anastrepha Schiner, 1868 (Diptera: Tephritidae) de Venezuela. Dissertation, Universidad Central de Venezuela, Maracay. xi + 210 p. [monograph of 37 spp. in Venezuela]
Caraballo, J. 1985. Nuevas especies del género Anastrepha Schiner, 1868 (Diptera: Tephritidae) de Venezuela. Bol. Entomol. Venez. 4: 25-32.
Cavalloro, R., ed. 1983. Fruit flies of economic importance. Proceedings of the CEC/IOBC International Symposium, Athens, Greece, November 16-19, 1982. A. A. Balkema, Rotterdam. xii + 642 p.
Cavalloro, R., ed. 1989. Fruit flies of economic importance 87. Proceedings of the CEC/IOBC International Symposium, Rome, 7-10 April 1987. A. A. Balkema, Rotterdam. 626 p.
Economopoulos, A. P., ed. 1987. Fruit flies: Proceedings of the Second International Symposium, 16-21 September 1986, Colymbari, Crete, Greece. Elsevier Science Publishers (distributors), Amsterdam & New York. 590 p.
Foote, R. H., Blanc, F. L. & Norrbom, A. L. 1993. Handbook of the fruit flies (Diptera: Tephritidae) of America north of Mexico. Comstock Publishing Associates, Ithaca. xii + 571 p. [key to 19 spp. in USA]
Hendel, F. 1914. Die Bohrfliegen Südamericas. Abhandl. Bericht. Königl. Zool. Anthrop. Ethnograph. Mus. Dresden (1912) 14: 1-84.
Hernández-Ortiz, V. 1992. El genero Anastrepha Schiner en Mexico (Diptera: Tephritidae). Taxonomia, distribucion y sus plantas huespedes. Instituto de Ecología and Sociedad Mexicana de Entomología, Xalapa, Veracruz, Mexico. 162 p. [monograph of 28 spp. in Mexico]
Lima, A. da Costa. 1934. Moscas de frutas do genero Anastrepha Schiner, 1868 (Diptera: Trypetidae). Mem. Instit. Oswaldo Cruz 28: 487-575. [revision of 62 spp.]
Loew, H. 1873. Monographs of the Diptera of North America. Part III. Smithsonian Misc. Coll. 11 (Publ. 256): 1-351.
Malavasi, A. & R. A. Zucchi, eds. 2000. Moscas-das-frutas de importância econômica no Brasil. Conhecimento básico e aplicado. Holos, Riberão Preto. 327 p. [Brazil]
McPheron, B. A. & G. J. Steck, eds. 1996. Fruit fly pests: A world assessment of their biology and management. St. Lucie Press, Delray Beach. 586 p.
McPheron, B. A., H.-Y. Han, J. G. Silva & A. L. Norrbom. 1999. Phylogeny of the genera Anastrepha and Toxotrypana (Trypetinae: Toxotrypanini) based upon 16S rRNA mitochondrial DNA sequences, p. 343-361. In M. Aluja & A. L. Norrbom, eds., Fruit flies (Tephritidae): Phylogeny and evolution of behavior. CRC Press, Boca Raton. [16] + 944 p. [phylogeny]
Norrbom, A. L. 1991. The species of Anastrepha (Diptera: Tephritidae) with a grandis-type wing pattern. Proc. Entomol. Soc. Wash. 93: 101-124.
Norrbom, A. L. 1997. Revision of the Anastrepha benjamini species group and the A. pallidipennis complex (Diptera: Tephritidae). Insecta Mundi 11: 141-157.
Norrbom, A. L. 1998. A revision of the Anastrepha daciformis species group (Diptera: Tephritidae). Proc. Entomol. Soc. Wash. 100: 160-192.
Norrbom, A.L. Host plant database for Anastrepha and Toxotrypana (Diptera: Tephritidae: Toxotrypanini). Diptera Data Dissemination Disk 2 (in press).
Norrbom, A. L. & Kim, K. C. 1988. Revision of the schausi group of Anastrepha Schiner (Diptera: Tephritidae), with a discussion of the terminology of the female terminalia in the Tephritoidea. Ann. Entomol. Soc. Am. 81: 164-173.
Norrbom, A. L. & Kim, K. C. 1988. A list of the reported host plants of the species of Anastrepha (Diptera: Tephritidae). U.S. Dep. Agric. Animal Plant Health Insp. Serv. APHIS 81-52: 114 p.
Norrbom, A.L., R.A. Zucchi & V. Hernández-Ortiz. 1999. Phylogeny of the genera Anastrepha and Toxtrypana (Trypetinae: Toxotrypanini) based on morphology, p. 299-342. In M. Aluja & A. L. Norrbom, eds., Fruit flies (Tephritidae): Phylogeny and evolution of behavior. CRC Press, Boca Raton. [16] + 944 p. [phylogeny]
Robinson, A. S. & G. Hooper, eds. 1989. Fruit flies. Their biology, natural enemies, and control, xii + 372 + xv + 447 p. W. Helle, ed., World crop pests, Vol. 3(A) & 3(B). Elsevier, Amsterdam.
Steck, G. J., Carroll, L. E., Celedonio-Hurtado, H. & Guillen-Aguilar, J. 1990. Methods for identification of Anastrepha larvae (Diptera: Tephritidae), and key to 13 species. Proc. Entomol. Soc. Wash. 92: 333-346. [key to larvae of 13 spp.]
Steyskal, G. C. 1977. Pictorial key to species of the genus Anastrepha (Diptera: Tephritidae). Entomological Society of Washington, Washington, D.C. 35 pp. [key to 141 spp.]
Stone, A. 1939. A revision of the genus Pseudodacus Hendel (Dipt. Trypetidae). Rev. Entomol. (Rio J.) 10: 282-289. [revision of 4 spp.]
Stone, A. 1939. A new genus of Trypetidae near Anastrepha (Diptera). J. Wash. Acad. Sci. 29: 340-350. [revision of 10 spp.]
Stone, A. 1942. The fruitflies of the genus Anastrepha. U. S. Dept. Agric. Misc. Publ. No. 439. 112 pp. [revision of 126 spp.]
Stone, A. 1942. New species of Anastrepha and notes on others. J. Wash. Acad. Sci. 32: 298-304. [revision of 7 spp.]
Tigrero, J. O. 1998. Revisión de especies de moscas de la fruta presentes en el Ecuador. Published by the author, Sangolquí, Ecuador. 55 p.
Zucchi, R. A. 1978. Taxonomia das espécies de Anastrepha Schiner, 1868 (Diptera, Tephritidae) assinaladas no Brasil. Dissertation, Universidade de São Paulo, Piracicaba. vi + 105 p. [monograph of 77 spp. in Brazil]
Zucchi, R. A. 1979. Novas especies de Anastrepha Schiner, 1868 (Diptera, Tephritidae). Rev. Bras. Entomol. 23: 35-41.

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