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Fruit Flies (Diptera: Tephritoidea): Biology, Host Plants, Natural Enemies, and the Implications to Their Natural Control

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Fruit Flies (Diptera: Tephritoidea): Biology, Host Plants, Natural Enemies, and the Implications to Their Natural Control 270 Integrated Pest Management and Pest Control – Current and Future Tactics Smith, H.S., 1919. On some phases of insect control by the biological method. J. Econ. Entomol. 12: 288–292. 12 Solomon, M. 1949. The natural control of animal populations. Jour. Anim. Ecol. 18:1–35. Sun, C. N., G. P. Georghiou and K. Weiss. 1980. Toxicity of Bacillus thuringiensis subsp. israelensis to mosquito larvae variously resistant to conventional insecticides. Mosq. News 40:614-618. Fruit Flies (Diptera: Tephritoidea): Steffan, W. A. and N. L. Evenhuis. 1981. Biology of Toxorhycnhites. Annu. Rev. Entomol. 26: Biology, Host Plants, Natural Enemies, 159-181. Trpis, M. 1972. Development and predatory behavior of Toxorhynchites brevipalpis (Diptera, and the Implications to Their Natural Control Culicidae) in relation to temperature. Environ. Entomol. 1: 537-546. Trpis, M. 1973. Interaction between the predator Toxorhynchites brevipalpis and its prey Aedes M. A. Uchoa aegypti. Bull. Wld. Hlth. Org. 49: 359-365. Laboratório de Insetos Frugívoros, Tully, T. P. Cassey and R. Ferriere. 2005. Functional response: rigorous estimation and Universidade Federal da Grande Dourados sensitivity to genetic variation in prey. Oikos 111: 479-487. Brazil Van Dam, A. R., and W. E. Walton. 2008. The effect of predatory fish exudates on the ovipositional behavior of three mosquito species: Cules quinquefasciatus, Aedes aegypti and Culex tarsalis. Medical and Veterinary Entomology 22: 399-404. 1. Introduction Vanlerberghe, V., M. E. Toledo, M. Rodriguez, D. Gomez, A. Baly, J. R. Benitez, and P. Van der Stuyft. 2009. Community involvement in dengue vector control: cluster Brazil is the third world largest producer of fruits, surpassed only by China (94.4 millions of randomised trial. British Medical Journal 338: b1959. tons) and India (51.14 million tons) (Vitti, 2009). The fruit growing area in Brazil currently Vanlerberghe, V., E. Villegas, M. Oviedo, A. Baly, A. Lenhart, P. J. McCall, and P. Van der takes up 2.3 millions of hectares, with an annual production superior to 36.8 millions of Stuyft. 2011. Evaluation of the effectiveness of insecticide treated materials for tones. The horticulture generates six millions of direct jobs, totalizing about 27% of total household level dengue vector control. PLoS Negl Trop Dis 5: e994. labor force employed in agriculture in the Country, and makes a gross domestic product Various. 1995. Biological Control Benefits and Risks. Cambridge University press. (GDP) of about US$ 11 billion. In the farms of fruit growing, in general, there are a demand Vezzani, D. and A. P. Albicocco. 2009. The effect of shade on the container index and pupal for intensive and qualified labor, creating jobs and ensuring a rural Well-being of the productivity of the mosquitoes Aedes aegypti and Culex pipiens breeding in artificial farmers and their employees, both on small farms as on large farms. However, Brazil containers. Medical and Veterinary Entomology, 23: 78–84. occupies the 17th position among world exporters of fruits (Ibraf, 2009; Vitti, 2009). Wandscheer, C. B., J. E. Duque, M. A. N. da Silva, Y. Fukuyama, J. L. Wohlke, J. Adelmann, Part of Brazilian fruit production is lost in the field due the attack by larvae of different and J. D. Fontana. 2004. Larvicidal action of ethanolic extracts from fruit endocarps species of fruit flies (Diptera: Tephritoidea). Herein, fruit flies are referred as the guild of all of Melia azedarach and Azadirachta indica against the dengue mosquito Aedes aegypti. specialized species with frugivorous larvae, that in South America, especially in Toxicon 44: 829-835. Brazil, belong to two families: Tephritidae and Lonchaeidae (Diptera: Tephritoidea) Wang, C. H., N. T. Chang, H. H. Wu, and C. M. Ho. 2000. Integrated control of the dengue (Uchoa & Nicácio, 2010). On the other hand, the fruit flies are interesting animals of the vector Aedes aegypti in Lui-Chui village, Ping-Tung country, Taiwan. Journal of the scientific point of view, because they have polytene chromosomes like those found in American Mosquito Control Association 16: 93-99. species of Drosophila (Drosophilidae), which are very important for genetics studies. Fruit Williams, F. M. and S. A. Juliano. 1985. Further difficulties in the analysis of functional Flies also can be easily reared in the laboratory to serve as experimental animals for response experiments and a resolution. Can. Entomol. 117: 631-640. research in several areas of the biological and environmental sciences (Uchoa et al., 2004). Watts, R. B. and S. M. Smith. 1978. Oogenesis in Toxorhynchites rutilus (Diptera: Culicidae). Can. J. Zool. 56: 136-139. The fruit flies belong to two families: Tephritidae and Lonchaeidae (Tephritoidea). They World Health Organisation. 2002. Malaria entomology and vector control. Learner's guide. have great economic importance because they are considered the key pests that most WHO/CDS/CPE/SMT/2002.18. adversely affect the production and marketing of fruits and vegetables around the world. World Health Organisation. 2004. Global strategic framework for integrated vector The tephritids are able of inserting the ovipositor to drop their eggs into the living tissues of management. WHO/CDS/CPE/PVC/2004.10. host plants, such as green fruit, fruit in process of maturation or ripe fruits. If females of World Health Organisation. 2009. Dengue guidelines for diagnosis, treatment, prevention Lonchaeidae lay their eggs inside or over the fruits, flowers, or inside terminal shoots of and control WHO/HTM/NTD/DEN/2009.1. Euphorbiaceae is still unknown. According Lourenção et al. (1996), Neosilba perezi (Romero Wright, J. W., R. F. Fritz, and J. Haworth. 1972. Changing concepts of vector control in & Ruppel) is a key pest in shoots of cassava clones. Both families of fruit flies cause direct malaria eradication. Ann. Rev. Entomol. 17: 75-102. and indirect damages. The direct ones are because their eggs hatch and the larvae eat the underlying flesh of the fruits. The indirect damage is due to depreciation of the fruits in the Fruit Flies (Diptera: Tephritoidea): 272 Integrated Pest Management and Pest Control – Current and Future Tactics Biology, Host Plants, Natural Enemies, and the Implications to Their Natural Control 273 market retailers; opening holes through which can penetrate pathogenic microorganisms or probably by airplane flights (aircraft) between Indonesia and Suriname (Oliveira et al., 2006). decomposers, or yet, causing the early fall of fruits attacked in the field. Some species of B. carambolae is a species in process of eradication from the Region North of Brazil. fruit flies are also the major bottleneck in the exports of fresh fruits and vegetables between The genus Ceratitis has 89 described species worldwide, occurring mainly in tropical Africa. nations. This is because the importing countries generally impose stringent quarantine In Brazil occurs only Ceratitis capitata which is distributed in almost all tropical and warm barriers to the producing and exporting Countries where fruit flies do occur, fearing the temperate areas in the world (Virgilio et al., 2008). C. capitata is originally from Africa, with entry exotic species inside the imported products in their territories (Uchoa & Nicácio, 2010; abundant populations in the Mediterranean region which borders with Europe. It has been White & Elson-Harris, 1992). found in Brazil for the first time in 1901, in the state of São Paulo (Uchôa & Zucchi, 1999). Tephritidae is the most species rich family of fruit flies, with around 5,000 described species, The genus Rhagoletis, with 70 described species occurs mainly in the Holarctic and in six subfamilies (Tachiniscinae, Blepharoneurinae, Phytalmyiinae, Trypetinae, Dacinae, Neotropical regions, being reported 21 species in the last one. Rhagoletis species infest mostly and Tephritinae); about 500 genera, and probably many undescribed species worldwide. fruits of Juglandaceae, Rosaceae, Rutaceae, and Solanaceae. In the Brazilian territory are Tephritids are peculiars because they are among the few groups of dipterans strictly reported three species (Ragoletis adusta Foote, from the state of São Paulo, R. ferruginea phytophagous, except the Tachiniscinae, which are thought be parasitoids of Lepidoptera, R macquarti and at least, some species of Phytalmyiinae that feed on live or dead bamboos (Poaceae) or Hendel, in Bahia, Paraná, and Santa Catarina, and . (Loew), in Goiás, and Minas on trees recently fallen of other plant families. Blepharoneurinae feed in flowers, fruits, and Gerais (Foote, 1981; Ramírez et al., 2008), but the species of Rhagoletis have not been make galls in Cucurbitaceae; Trypetinae and Dacinae feed in fruits or in seeds of a wide considered as key pests in Brazil. On the other hand, some species in this genus are pest of range of plant families, and Tephritinae eat in flowers, make gall, or are leaf-miners in a fruits in Peru and Chile (Salazar et al., 2002). wide array of plant taxa: Aquifoliaceae, Scrophulariaceae, Verbenaceae, but mainly in Lonchaeidae is the second family of fruit flies with economic importance in South America, flowerheads of Asteraceae (Norrbom, 2010; Uchoa & Nicácio, 2010). where some species of the genera Dasiops and Neosilba are primary pests in crop fruits. The The Lonchaeidae fruit flies have about 500 described species worldwide, in two subfamilies, species of Dasiops attack cultivated or wild passion fruit species: green or ripe fruits, or floral and nine genera. Dasiopinae is represented only by Dasiops Rondani, and the Lonchaeinae, buds (Passifloraceae), depending on the Dasiops species (Norrbom & Mcalpine, 1997; Uchôa et al., 2002; Uchôa & Nicácio, 2010). The Neosilba species are generally polyphagous, with the other eight remaining genera, being Neosilba the most studied and economically attacking many species of fruit, native or exotic, cultivated or wild ones. The Neosilba species important genus in Neotropics, with 20 described species, from which 16 are reported in most commonly involved in the infestation of fruits and vegetables are: N. zadolicha Steyskal Brazil. The genus Dasiops, with about 120 described species worldwide, have few species & McAlpine, N. pendula (Bezzi), N.
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