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Toxotrypana curvicauda Gerstaecker
Papaya fruit fly

Lateral female habitus and dorsal male habitus (from Knab & Yothers 1914).

Recognition
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This species, like other Toxotrypana species, is wasplike in appearance. The body is elongate, yellow with dark brown markings, the abdomen is petiolate, and the wing pattern includes only a broad costal band extending the length of the wing and a faint streak on the cubital cells. Toxotrypana species differ from other Tephritidae in having a medial, longitudinal depression on the scutum, vein R₂₊₃ with 3 sharp bends, often with spur veins arising from them, and most head and thoracic setae reduced in size. T. curvicauda is the only species of Toxotrypana known from the United States and the West Indies (it also occurs in Mexico, Central and northern South America). It is one of the species of this genus in which the female oviscape is extremely long (at least as long as the thorax and abdomen combined) and strongly arched.

Toxotrypana curvicauda differs from the other species of the genus with such long terminalia in the pattern of its dark brown thoracic markings. In particular, on the scutum the posterior brown mark is broader than long and separate from the submedial and sublateral stripes, and the latter are strongly laterally curved posteriorly. The anatergite at most has a dorsal and a ventrolateral spot (as opposed to a complete stripe or being mostly brown), and the scutellum at most has the base and lateral third of the apical margin brown.

The larvae are difficult to distinguish from those of other fruit flies. They are opaque whitish or yellowish, cylindrical, tapering anteriorly, with 13-15 oral ridges, 22-28 tubules on the anterior spiracle, relatively short posterior spiracular hairs, and with inconspicuous tubercles on the caudal segment (Heppner 1986).

Classification and Evolutionary Relationships
Order: Diptera. Family: Tephritidae. Subfamily: Trypetinae. Genus: Toxotrypana. Species: curvicauda. Author: Gerstaecker.
Relationships among the species of Toxotrypana have not been analyzed other than the recognition of two species groups by Norrbom et al. (1999). The phylogenetic relationships of Toxotrypana to other fruit flies were discussed by Norrbom et al. (1999) and McPheron et al. (1999). Click here for a more extensive discussion of Toxotrypana phylogeny.

Names Used for this Species
Toxotrypana curvicauda Gerstaecker 1860: 194.
Toxotrypana fairbatesi Munro 1984: 160; synonymy (Steyskal 1986: 114).
Mikimyia furcifera Bigot 1884: xxix.; synonymy (Mik 1890: 251, Snow 1895: 119).
Toxotrypana curvicaude: Munro 1984: 160 [misspelling].
Papaya fruit fly
Click here to link to fly names database

Type data
T. curvicauda: Syntype(s) - female, number unstated (ZMHU), "St. Jean" [St. John, Virgin Islands], Moritz.
M. furcifera: Holotype - male (UMO), Brazil [probably erroneous].
T. fairbatesi: Holotype - male (SANC), CUBA: Soledad, Sta. Clara, 1 Aug.1932 [not VII.1932 as stated by Munro], Bates & Fairchild.

Distribution
USA (s. Texas) south to Colombia & Venezuela, West Indies; introduced USA (Florida). This species has been reported from the following countries: Bahamas, Belize, Colombia, Costa Rica, Cuba, Dominican Republic, El Salvador, Guatemala, Honduras, Mexico (Morelos, Oaxaca, Tamaulipas, Veracruz), Nicaragua, Panama, Puerto Rico, Trinidad, U.S.A. (Florida (introduced), Texas), Venezuela, Virgin Islands. It does not occur in Brazil, Peru or South Carolina, USA; records from those areas were based on errors or misidentifications by Bigot (1884), Röder (1891: 31), and Hendel (1914).
Click here for specimen data

Toxotrypana curvicauda was introduced to Florida sometime prior to December, 1905 (Knab & Yothers 1914).  It has been found in all parts of the state where papaya is grown, including the following counties: Brevard, Broward, Charlotte, Dade, Highlands, Hillsborough, Indian River, Lee, Manatee, Martin, Monroe, Orange, Palm Beach, Pinellas, St. Lucie, Sarasota, and Volusia (Weems 1969, Foote et al. 1993).
Click here for map of US distribution

Biology
This species is a common pest of papaya (Carica papaya L.) and is commonly known as the papaya fruit fly, although other hosts are known. Like other species of Toxotrypana, T. curvicauda appears to breed in the fruits of a variety of Asclepiadaceae, Caricaceae, and perhaps Apocynaceae.  All of these plants produce copious, milky white latex and have fruits with very thick skins.  The larvae feed on developing seeds and associated tissues within the central cavity of the fruit. Landolt (1999) extensively reviewed the behavior and other aspects of the biology of this species.  The adults are vespid wasp mimics.   Males stake out territories on host plants and produce a pheromone to call females.  The female lays its eggs in immature fruits, using the long ovipositor to penetrate the skin and thick pulp, and deposit the eggs in the seed cavity. There are three larval stages.  The mature larva tunnels out of the fruit and pupates in the soil.  See Selman (1998, http://www.ifas.ufl.edu/~insect/fruit/tropical/papaya_fruit_fly.htm) for additional description of the life cyle.

Natural host plants, besides papaya, include: Carica cauliflora Jacq. (Caricaceae), commonly known as "papayito" in Venezuela (Guagliumi 1966); a Carica sp., commonly known as papayuelo in Colombia (Figueroa 1977); Jacaratia mexicana (Caricaceae), commonly known as "cuaguayote" or "bonete" in Morelos, Mexico (Castrejón-Ayala 1987); an Apocynaceae sp., probably Tabernaemontana sp. in Cuba (Bates 1933); and at least three species of Gonolobus (Asclepiadaceae). The latter include: G. barbatus Kunth, commonly known as "pancololote" in Morelos, Mexico (Castrejón-Ayala & Camino-Lavín 1991, as G. sorodius); a wild species of Asclepiadaceae called "talayote" or "talayotillo" in northeastern Mexico (Santa Engracia and Cañon de Rosario) reported by Baker et al. (1944), and suggested by Castrejón-Ayala (1987) to be a species of Gonolobus, and by Landolt (1994) as possibly G. erianthus D.C.; and G. salvinii Hemsl., commonly called "champeron", in Guatemala (Norrbom & Muñiz, unpubl. data). In Florida, curvicauda has been reported to attack two exotic plants, Morrenia odorata (Asclepiadaceae) (Landolt 1994) and mango, Mangifera indica L. (Anacardiaceae), although the latter record appears to have been based on isolated or very rare occurrences (Butcher 1952), and mango is not considered a normal host plant (Landolt 1999).
Click here to link to host plant database

Economic Significance
As indicated by its common name, papaya fruit fly, T. curvicauda is a pest of papaya (Carica papaya L.).  According to Weems (1969) and Heppner (1986), it is one of the most important pests of this fruit; according to Wolfenbarger & Walker (1974) and Lamberts & Crane (1990), it is one of the three most important insect pest of papaya in Florida. The larvae feed on the developing seeds and other tissues within the central cavity of the fruit, and the holes they burrow through the pulp to exit the fruit destroy the edible part of the papaya.

Management and control of this pest are discussed on the following sites:
http://www.ifas.ufl.edu/~insect/fruit/tropical/papaya_fruit_fly.htm
http://edis.ifas.ufl.edu/scripts/htmlgen.exe?DOCUMENT_IG074

References
Click here to access fruit fly literature database
Aldrich, J. M. 1905. A catalog of North American Diptera (or two-winged flies). Smithson. Misc. Collect. 46(2): 680 p. [p. 600, in catalog]
Baker, A. C., W. E. Stone, C. C. Plummer & M. McPhail. 1944. A review of studies on the Mexican fruitfly and related Mexican species. U.S. Dep. Agric. Misc. Publ. 531: 155 p. [p. 142, Mexico, hosts]
Bates, M. 1933. Notes on West Indian Trypetidae (Diptera). Bull. Brooklyn Entomol. Soc. 28: 160-172. [p. 160, Cuba, host]
Benjamin, F. H. 1934. Descriptions of some native trypetid flies with notes on their habits. U.S. Dep. Agric. Tech. Bull. 401: 95 p. [p. 11, taxonomy, immature stages]
Bigot, J. M. F. 1884. Description d'un nouveau genre et d'une nouvelle espece de Dipteres de la famille des Ortalidae. Ann. Soc. Entomol. Fr. (6) 4 (Bull.): xxix-xxx. [taxonomy]
Butcher, F. G. 1952. The occurrence of papaya fruit fly in mango. Proc. Fla. State Hortic. Soc. 65: 196. [host plant]
Castrejón-Ayala, F. 1987. Aspectos de la biologia y habitos de Toxotrypana curvicauda Gerst. (Diptera: Tephritidae) en condiciones de laboratorio y su distribucion en una plantacion de Carica papaya L. en Yautepec, Mor. Thesis, Escuela Nacional de Ciencias Biológicas, Mexico, D.F., iv + 88 p. [host plants]
Castrejón-Ayala, F. & M. Camino-Lavín. 1991. New host plant record for Toxotrypana curvicauda (Diptera: Tephritidae). Fla. Entomol. 74: 466. [host plants]
Enderlein, G. 1911. Trypetiden-Studien. Zool. Jahrb. Abt. Syst. Geogr. Biol. Tiere 31: 407-460. [p. 407, Costa Rica]
Figueroa Potes, A. 1977. Insectos y acarinos de Colombia. Facultad de Ciencias Agropecuarias, Universidad Nacional, Palmira, 685 p. [host plants, Colombia]
Foote, R. H. 1965. Family Tephritidae, p. 658-678. In A. Stone, C. W. Sabrosky, W. W. Wirth, R. H. Foote & J. R. Coulson, eds., A catalog of the Diptera of America north of Mexico. U.S. Dep. Agric. Agric. Handb. 276: 1696 p. [p. 659, in catalog]
Foote, R. H. 1967. Family Tephritidae (Trypetidae, Trupaneidae), 91 p. In N. Papavero, ed., A catalogue of the Diptera of the Americas south of the United States. 57. Departamento de Zoologia, Secretaria da Agricultura, São Paulo. [p. 48, in catalog]
Foote, R. H., F. L. Blanc & A. L. Norrbom. 1993. Handbook of the fruit flies (Diptera: Tephritidae) of America north of Mexico. Comstock Publishing Associates, Ithaca. xii + 571 p. [p. 415, USA, references]
Gerstaecker, A. 1860. Beschreibung einiger ausgezeichneten neuen Dipteren aus der Familie Muscariae. Stettin. Entomol. Ztg. 21: 163-202. [p. 194, original description]
Greene, C. T. 1929. Characters of the larvae and pupae of certain fruit flies. J. Agric. Res. 38: 489-504. [p. 491, immature stages]
Guagliumi, P. 1966. Insetti e aracnidi delle piante comuni del Venezuela segnalati nel periodo 1938-1963. Relaz. Monogr. Agr. Subtrop. Trop. (N.S.) 86: 392 p. [host plants, Venezuela]
Hendel 1914. Die Bohrfliegen Südamerikas. Übersicht und Katalog der bisher aus der neotropischen Region beschriebenen Tephritinen. Abh. Ber. K. Zool. Anthrop. Ethnogr. Mus. (1912) 14 (3): 1-84. [p. 10, review]
Heppner, J. B. 1986. Larvae of fruit flies. III. Toxotrypana curvicauda (Papaya fruit fly) (Diptera: Tephritidae). Fla. Dep. Agric. Consum. Serv. Div. Plant Ind. Entomol. Circ. 282: 2 p. [larva]
Kitto, G. B. 1983. An immunological approach to the phylogeny of the Tephritidae, p. 203-211. In R. Cavalloro, ed., 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. [p. 205, biochemical taxonomy]
Knab, F. & W. W. Yothers. 1914. Papaya fruit fly. J. Agric. Res. 2: 447-453. [host, biology, immature stages, Florida]
Lamberts, M. and J.H. Crane. 1990. Tropical fruits, p. 337-355. In J. Janick and J.E. Simon, eds., Advances in new crops. Timber Press, Portland, OR. [pest status]
Landolt, P. J. 1994. Fruit of Morrenia odorata (Asclepiadaceae) as a host for the papaya fruit fly, Toxotrypana curvicauda (Diptera: Tephritidae). Fla. Entomol. 77: 287-288. [host plants]
Landolt, P. J. 1999. Behavior of flies in the genus Toxotrypana (Trypetinae: Toxotrypanini), p. 363-373. In M. Aluja & A. L. Norrbom, eds., Fruit flies (Tephritidae): Phylogeny and evolution of behavior. CRC Press, Boca Raton. [16] + 944 p. [behavior, biology].
Mason, A. C. 1922. Biology of the papaya fruit fly, Toxotrypana curvicauda, in Florida. U.S. Dep. Agric. Bull. 1081: 10 p. [biology, control]
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. [phylogenetic relationships]
Mik, J. 1890. Ueber Toxotrypana curvicauda Gerst. und Mikimyia furcifera Big. (Ein dipterologischer Beitrag.). Wien. Entomol. Ztg. 9: 251-254.[classification]
Munro, H. K. 1984. A taxonomic treatise on the Dacidae (Tephritoidea, Diptera) of Africa. Entomol. Mem. S. Afr. Dep. Agric. 61: [i] + ii-ix + 313 p. [p. 159, taxonomy]
Norrbom, A. L., L. E. Carroll, F. C. Thompson, I. M. White & A. Freidberg. 1999. Systematic database of names, pp. 65-251. In F. C. Thompson (ed.), Fruit Fly Expert Identification System and Systematic Information Database. Myia 9, vii + 524 pp. & Diptera Data Dissemination Disk (CD-ROM) (1998) 1. [type data, distribution, synonymy]
Norrbom, A. L., R. A. Zucchi & V. Hernández-Ortiz. 1999. Phylogeny of the genera Anastrepha and Toxotrypana (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. [phylogenetic relationships]
Osten Sacken, C. R. 1878. Catalogue of the described Diptera of North America. [Ed. 2]. Smithson. Misc. Collect. 16(2): xlvi + [2] + 276 p. [p. 181, in catalog]
Röder, V. von. 1891. Bemerkungen zu dem dipterologischen Beitrage von Prof. Mik in der Wiener Entomologischen Zeitung, Jahrg. 1890, pag. 251, über Toxotrypana curvicauda Gerst. Wien. Entomol. Ztg. 10: 31-32. [misidentification]
Sarma, R., G. B. Kitto, S. H. Berlocher & G. L. Bush. 1987. Biochemical and immunological studies on an alpha-glycerophosphate dehydrogenase from the tephritid fly, Anastrepha suspensa. Arch. Insect Biochem. Physiol. 4: 271-286. [p. 283, biochemical taxonomy]
Snow, W. A. 1895. On Toxotrypana of Gerstaecker. Kans. Univ. Q. 4: 117-119. [classification, Mexico]
Steyskal, G. C. 1986. A taxonomic treatise on the Dacidae (Tephritoidea, Diptera) of Africa, H. K. Munro [Book review]. Bull. Entomol. Soc. Am. 32: 114. [taxonomy]
Wulp, F. M. van der. 1899. Fam. Muscidae, p. 393-408. In F. D. Godman & O. Salvin, eds., Biologia Centrali-Americana. Zoologia. Class Insecta. Diptera. Or, contributions to the knowledge of the fauna and flora of Mexico and Central America. Vol. 2. Taylor & Francis, London. 489 p. [p. 379, Mexico]
Wasbauer, M. S. 1972. An annotated host catalog of the fruit flies of America north of Mexico (Diptera: Tephritidae). Occas. Pap. Calif. Dep. Agric. Bur. Entomol. 19: [i] + 172 p. [p. 136, hosts]
Weems, H. V. 1969. Papaya fruit fly (Toxotrypana curvicauda Gerstaecker), 1860 (Diptera: Tephritidae). Fla. Dep. Agric. Div. Plant Industry, Entomol. Circ. 86: 2 p. [pest circular, Florida]
White, I. M. & M. M. Elson-Harris. 1992. Fruit flies of economic significance: Their identification and bionomics. C A B International, Wallingford. xii + 601 p. [p. 390, review]
Wolfenbarger, D. O. & S. D. Walker. 1974. Two major pest problems of papayas. Proc. Fla. State Hortic. Soc. 1974: 384-385. [pest status, control]

Contact Information
Identification:
The Systematic Entomology Laboratory provides identifications for most agriculturally important groups of insects and mites on a per request basis. See http://www.sel.barc.usda.gov/selhome/requests.htm.
In the United States, identifications are often provided by or through county extension agents.