BIOLOGICAL CONTROL OF TEPHRITIDAE
----CLICK on desired underlined categories [ to search for Subject Matter, depress Ctrl/F ]:
The biological control efforts against fruit flies of the genus Tephritidae have been extensive over the past half century, a thorough review being given in Clausen (1987). However, as it becomes increasingly apparent that the Mediterranean fruit fly, Ceratitis capitata (Wiedemann), and Mexican fruit fly, Anastrepha ludens (Loew) pose a continued threat to California's agriculture through periodic invasions of our borders, there is an urgent need to consider the application of alternative methods to chemicals in eradication and control programs. The implementation of effective biological controls at the sources of invasion as well as within the state boundaries where breeding may occur, offers an environmentally sound, non-polluting alternative. There is an urgent need to (1) search for, procure and initially evaluate natural enemies of Mediterranean and Mexican fruit flies fruit fly in natural ranges in central Africa and southern Mexico (parasites, predators and pathogens); (2) introduce and study foreign natural enemies in the adult stage, and evaluate their respective effectiveness under field conditions in Hawaii, southern Mexico, and if applicable, California; (3) attempt development of a mass production scheme of resident California fruit flies (e.g., walnut husk fly) to serve as acceptable hosts for Mediterranean fruitfly natural enemies for use in laboratory study and periodic colonization efforts in infested areas of California, and (4) to test the feasibility of building a culture bank of Medfly and Mexican fruit fly natural enemies on resident California fruit flies for use in conjunction with other eradication and control methods (e.g., sterile-male releases, adult fly baiting) during periodic invasions of these pests and in anticipation of their possible permanent establishment in the State of California.
The fruit flies of the family Tephritidae constitute a group of agricultural pests of worldwide importance, as they attack a wide range of fruits and vegetables. The most important are the several species of Dacus and Ceratitis, which occur in many countries of warm temperate and subtropical climates; Anastrepha, an American genus occurring from Mexico and the West Indies through South America; and Rhagoletis, with a more restricted host range, occurring in the north temperate region (Legner & Goeden (1987). The Mediterranean fruit fly, although eradicated periodically from the state of Florida where it had "peninsular" distribution, and recently from California where it repeatedly reappears, is presently firmly established in southern Mexico. There it has been temporarily contained by a massive sterile-male and parasite release effort by the U. S. Department of Agriculture. The appearance of Anastrepha in southern Baja California during the past two years suggests that it may eventually move north and pose a continuous threat along the Mexican border. Another chronic threat has been the permanently established population in the Hawaiian Islands, from which periodic accidental invasions of California are thought to occur. Recently, Carey & Dowell (1989), Greathead & Waage (1983), Gilstrap et al (1987), Wharton (1989) and Wong & Ramadan (1990) have noted that further biological control efforts are definitely justified against fruit flies.
Several studies have investigated the potential economic of C. capitata in California and elsewhere. Details on this and various abatement tactics may be found in UC/AID (1977) Galt & Albertson (1981), Carey (1982, 1984), Gilmore (1983), Dowell (1983), Schreibner (1983), Spitler & Couey (1983), Williamson (1983), Krainaker et al. (1987) and Carter (1990).
The need for investigation into the biological control of fruit flies in Hawaii, Mexico and California is ever more important as it becomes recognized that insecticides, although offering expedient and predictable results under certain conditions, are often inadequate and at least perceived as dangerous, if not physically dangerous to wildlife and humans alike. As problems involving insecticidal residues and insect resistance to chemicals continue to increase, many programs directed at the control of fruit flies must ultimately be modified with increased dosages and costs to such an extent that they invariably arouse the concern and ire of naturalist and conservationist organizations. A case in point is the fire ant eradication program. By 1959 extensive damage to wildlife and domestic animals had positively been attributed to the effects of several insecticides used in the program (Clawson 1959). Fire ant control was finally declared unsuccessful in 1960, and in some states, fire ant numbers were actually reported to have increased since the eradication program began (Byrd 1960, Cottam 1959). Presently, a new effort to control fire ant is being attempted with natural enemies imported from Brazil and Argentina.
Some investigators believe that the Mediterranean and Mexican fruit flies are already permanently established in California and that unless the current eradication effort is greatly increased, it is just a matter of time before at least one species, Medfly, will spread throughout the state (Barinaga 1990). The malathion baits currently in use against them may not be potent enough for fast eradication, as it is recognized that Medflies will not eat the bait unless that is the only substance placed in their cages (Citrograph 1990). Under outdoor conditions they may prefer to seek out clean ripening fruit.
Mediterranean Fruit Fly.--The Mediterranean fruit fly is a major pest throughout the Mediterranean region, portions of Africa, the Middle East, Central and South America, Mexico, and Hawaii, and has become established in Australia. In France, it is able to persist from year to year only in areas bordering the Mediterranean, yet survival is reported in Austria, where severe winters, with continuous frosts, can cause up to 90% mortality of the pupae (Clausen 1978). Although parasitic insects have been imported against it, 95% of the species were obtained from areas outside the fly's accepted native range in central Africa and Madagascar. However, some reductions in infestations are attributable to natural enemy activity in the invaded areas, especially when parasitoids are mass released as biotic insecticides (Wong & Ramadan 1990, Wong et al. 1990).
The medfly was first described fin 1824 and was first noted as a pest in citrus in 1829 from shipments of oranges to England from the Azores. The fly spread throughout the world over the next 100 years and was continually noted as a destructive pest wherever it was found. The first program for the biological control of medfly was undertaken by the government of Western Australia in 1902 with the engagement of George Compere to search for natural enemies and to determine the aboriginal home of the medfly. Unfortunately Compere was never able to acertain the aboriginal home nor did he establish the parasitoids he collected from India, Sri Lanka and Brazil in Western Australia.
The medfly invaded Hawaii in 1910 and soon thereafter the Board of Commissioners hired Filipi Silvestri to again search for natural enemies of this fly. It was determined by experts of the day that collections should concentrate in Western Africa. Therefore, Silvestri traveled for eight months through West and East Africa and South Africa. He found only six specimens of the medfly on the entire journey, but reared many parasitoids from other fruit-infesting tephritids collected along the way. He managed to establish four species in Hawaii: Opius concolor Szepligeti, Biosteres tryoni (Cameron), Coptera silvestrii Kieffer and Dirhinus anthracina Walker. Two more missions over the next 30 years were sent out in hopes of obtaining parasitoids, but only Tetrastichus giffardianus Silvestri and Biosteres fullaway (Silvestri) were established.
Other biological control programs were undertaken in several countries where the medfly was firmly established, but these programs have not been well documented, and the extent of control of any of the parasitoid species is virtually unknown, the notable exception being Hawaii. Even in Hawaii control was never noteworthy and the medfly proglem was finally overshadowed by the introduction of Dacus dorsalis Hendel. For North America the answer to the medfly invasions starting in 1929 was complete eradication by means of fruit stripping and poisoned bait sprays.
The success of these early and subsequent biological control programs against medfly has been variable (Gilstrap & Hart 1987, Wharton & Gilstrap 1983). In Hawaii, a cooperative biological control program initiated in 1948 involved the release of 32 entomophagous species to compat both medfly and the oriental fruit fly. Three parasitoids, Biosteres longicaudatus (Ashmead), B. vandenboschi (Fullaway), and B. oophilus (Fullaway) became widespread and abundant (Bess et al. 1961). During 1966-1968, parasitization of the medfly and the oriental fruit fly was high (ca. 70%); it was mainly due to the egg-pupal parasitoid, B. oophilus (Haramoto & Bess 1970). During 1978-1981, Biosteres oophilus was still the predominant parasitoid as it accounted for ca. 80% of the total parasitization. Occasionally the larval-pupal parasitoids, Biosteres longicaudatus and B. tryoni (Cameron) achieved a parasitization of 32 and 8%, respectively (Wong et al. 1984). Extensive fruit collections done between 1949-1985 showed that Jerusalem cherry, coffee and peach were among the most important hosts of the medfly. These fruits yielded more than 100 larvae/Kg of infested fruits (Liquido et al. 1990; Nishida et al. 1985). The fruits that yielded a high number of medfly larvae were elliptical to spherical and yellowish to reddish. They had a diameter of 1-7 cm and a weight of 1-30 grams. Most of these hosts belonged to five plant families: Myrtaceae, Rutaceae, Rosaceae, Sapotaceae and Solanaceae (Liquido et al. 1990).
In Costa Rica a classical biological control program was initiated in 1955. During 1979-1980 parasitoids were collected from <10% of C. capitata populations. These were two introduced braconids, B. longicaudatus and B. oophilus, and two indigenous cynipids, Ganaspis carvalhoi (Dettmer) and Odontosema anastrephae (Borgmeier) (Wharton et al. 1981). An exploration for medfly parasitoids conducted in West-Central Africa during 1980-1982 showed that C. capitata occurred in low frequency in coffee plantations in Cameroon. Parasitization of tephritids in coffee by braconids ranged from 10-56% (Steck et al. 1986). In Guatemala infestation of C. capitata was serious in coffee and tangerine. The rest of the fruits were mainly infested by Anastrepha spp. (Eskafi 1988, 1990). Parasitization rate of C. capitata and Anastrepha spp. was low, ranging from 0.04 to 7.95%. The most common parasitoids recovered from both flies were B. longicaudatus and Doryctobracon crawfordi (Viereck) (Eskafi 1990).
The behavior of the ectoparasitoid Muscidifurax raptor (Girault & Sanders) in searching for the potential host C. capitata pupae was analyzed under laboratory conditions. The searching efficiency of M. raptor females decreased with increasing density. The proportion of avoidance of superparasitism was 0.615. The response to a high parasitoid density was to increase the proportion of males in the progeny, as males searching for mates interfered and decreased the searching activity of the females (Podoler & Menzel 1977, 1979). The medfly was susceptible to the Mexican strain of the nematode Steinernema feltiae. Emerging adults and pupae were not susceptible to the nematode, but the third instars (prior to pupating in the soil) suffered high mortalities (50-90%) when exposed to high nematode concentrations (150,000 - 500,000 nematodes/cup) (Lindegren & Vail 1986). Field exposure of mature larvae to a dose of 500 nematodes/cm2 yielded high mortality of C. capitata (Lindegren et al. 1990). In addition to the hymenopterous parasitoids and insect pathogenic nematodes, arthropod predators such as ants could play an important role in reducing fruit fly populations. Under laboratory conditions, the argentine ant, Iridomyrmex humilis (Mayr) caused 50% mortality of medfly pupae after a 10 min. attack. However, ant predation could be important only in localized areas; it is not adequate to regulate medfly populations (Wong et al. 1984).
Typically, the most effective natural enemies of an insect occur in regions where the pest evolved. The natural range of Mediterranean fruit fly is the sub-Saharan central African region, including the Island of Madagascar. Although no information is available from Madagascar, a number of promising natural enemies have been discovered in Central Africa (Bianchi & Krauss 1936, 1937; Gilstrap & Hart 1987, Greathead 1976, Silvestri 1914, Steck et al. 1986, van Zwaluwenburg 1936, 1937, Wharton, 1989, Wharton & Gilstrap 1983). However, because of technological difficulties associated with transportation and culture, only two species attacking Ceratitis capitata have been successfully translocated out of central Africa. A concentrated effort to locate natural enemies there might yield the kind of species capable of regulating this pest at low densities, as it has been known to be rare in that general region since the early 1900's (Silvestri 1913). We believe that parasitic Hymenoptera are the most effective natural enemies of Mediterranean fruit flies. At least six species of fruit flies in the genus Ceratitis are known from central Africa, and numerous parasitic Hymenoptera have been reported active on them at very low host densities (Table 1, Silvestri 1913, Clausen 1978, F. Gilstrap, pers. comm.). Entomologists in California have not tested these because the Mediterranean fruit fly has been quarantined. Therefore, promising species of natural enemies for Medfly might be found among these related species. Also, there has been no concentrated effort to locate disease organisms, such as viruses, bacteria and fungi, which might prove invaluable in eradication campaigns.
Mexican fruit fly.--Some of the natural enemies of oriental and Mediterranean fruit flies have shown activity on Anastrepha spp. in southern Mexico, and may be influential in partial biological control of that species (Aluja et al. 1990). However, there have been no formal attempts to obtain natural enemies from other areas where different species of Anastrepha occur, such as South America.
Two species of parasitic insects are already proven and available as biotic insecticides (augmentive releases) against Medfly. These are Diachasmimorpha longicaudata and D. tryoni, which have been used with some success in Mexico and Hawaii (USDA 1988, Wong et al. 1990a,b). The use of these parasites in lieu of malathion during the establishment phase of specific natural enemies from central Africa, would greatly aid their survival and while providing some economic control of Medfly.
Table 1. Known parasitic species attacking fruitflies of the genus Ceratitis in their natural range in Central Africa.
Parasite species Host stage attacked
Biosteres caudatus Szepligeti larva
Diachasma fullawayi Silvestri larva
Diachasmimorpha longicaudata (Ashmead) larva
Diachasmimorpha tryoni (Cameron) larva
Dirhinus ehrhorni Silvestri pupa
Dirhinus giffardii Silvestri pupa
Ganaspis sp. larva?
Halticoptera sp. larva?
Hedylus giffardii Silvestri larva?
Hedylus sp. larva?
Galesus silvestrii Kieffer pupa
Microbracon celer (Szepligeti) larva?
Opius humilis Silvestri larva
Opius inconsuetus Silvestri larva
Opius perproximus Silvestri larva
Opius n. sp. larva
Spalangia afra Silvestri pupa
Tetrastichus dacicida Silvestri larva
Tetrastichus giffardii Silvestri larva
Tetrastichus oxyurus Silvestri larva
Tetrastichus n. sp. larva
Misc. Fruitflies.--Several species of Rhagoletis are very important pests of cultivated cherries in North America and Europe, with some species having been considered as subjects for biological control, despite the low economic threshold. Infestation rates of less than 0.2% are currently required for commercial marketing of cherries in the United States. Four species of parasitoids associated with the Oriental fruit fly, Dacus dorsalis Hendel, were introduced against such fruit flies. These included Opius longicaudatus compensans (Silv.), Opius longicaudatus farmosanus (Full.), Opius oophilus Full., and Opius longicaudatus novacaledonicus Full. These parasitoids were introduced from Hawaii and released against Rhagoletis indifferens Cueran and Rhagoletis fausta Osten Sak in Oregon and Washington in the 1950's (Clausen 1956b). However, none became established probably because they all originated in tropical regions. A parasitoid of R. cerasi, the European cherry fruit fly, was imported against the eastern cherry fruit fly, R. cingulata Loew during 1959-64 in New Jersey, without successful establishment. Other species including Biosteres sublaevis Wharton, Coptera occidentalis and Phygadeuon wiesmanni are under investigation in California and Oregon (Croft & AliNiazee 1996, Legner & Goeden (1987).).
Abasa, R. O. 1973. Observations on the seasonal emergence of fruit flies on a Kenya coffee estate and studies of the pest status of Ceratitis capitata (Wied.) in coffee [Diptera: Tephritidae]. E. Afr. Agric. For. J. 39: 144-148.
Aluja, M. 1985. Manejo integrado de las moscas de la fruit [Diptera: Tephritidae]. Programa Mosca Med. DGSV-SARH, Mexico. 241 pp.
Aluja, M. & P. Liedo. 1986. Future perspectives on integrated management of fruit flies in Mexico. In: Pest Control: operations and systems analysis in fruit fly management (M. Mangel, ed.). Springer, New York, 12-48.
Aluja, M., M. Cabrera, E. Rios, J. Guillen, H. Celedonio, J. Hendrichs & P. Ledo. 1987a. A survey of the economically important fruit flies [Diptera: Tephritidae] present in Chapas and a few other fruit growing regions in Mexico. Fl. Entomol. 70: 320-329.
Aluja, M., J. Guillen, G. de la Rosa, M. Cabrera, H. Celedonio, P. Liedo & J. Hendrichs. 1987b. Natural host plant survey of the economically important fruit flies [Diptera: Tephritidae] of Chiapas, Mexico. Fl. Entomol. 70: 329-338.
Aluja, M., J. Guillen, P. Liedo, M. Cabrera, E. Rios, G. de la Rosa, H. Celedonio & D. Mota. 1990. Fruit-infesting Tephritidae [Dipt.: Tephritidae] and associated parasitoids in Chiapas, Mexico. Entomophaga 35: 39-48.
Anonymous. 1951. Distribution maps of insect pests. Map No. 1, Ceratitis capitata. Commonw. Inst. Entomol., Ser. A. 1p.
Anonymous. 1956. Distribution maps of insect pests. Map No. 64, Dacus curcubitae. Commonw. Inst. Entomol.Ser. A. 1 p.
Anonymous. 1960. Distribution maps of insect pests. Map No. 109, Dacus dorsalis. Commonw. Inst. Entomol., Ser. A. 1 p.
Anonymous. 1978. Medfly on the move. Citrograph 63: 99, 100, 114.
Arietta, M. D., L. Terrazas, & J. E. Jimenez. 1961. La mosca Mediterráneo (Ceratitis capitata Wied.) en Centro America. Situación actual en Nicaragua y posibilidades de erradicación. Fitofilo 14: 29-38.
Autter, S. H. 1977. Current fruit fly situation in Chile. FAO Plant Protection Bull. 24: 118-9.
Baas, J. 1959. The Mediterranean fruit fly Ceratitis capitata Wied. in central Europe (Part 2). Hofchen-Briefe 12: 113-40.
Back, E. A. 1914. The Mediterranean fruit fly in Bermuda. U. S. Dept. Agr. Bull. No. 161. 8 pp.
Back, E. A. 1917. Danger of introducing fruit flies into the United States. U. S. Dept. Agric. Yearbk. 1917, p. 185-96.
Back, E. A. & C. E. Pemberton. 1914. Life history of the melon fly. J. Agr. Res. 3: 269-74.
Back, E. A. & C. E. Pemberton. 1915. Parasitism among the larvae of the Mediterranean fruit fly (C. capitata) in Hawaii during 1914. Report to the Board of Commissioners for Agriculture and Forestry, Territory of Hawaii. 1914: 153-161.
Back, E. A. & C. E. Pemberton. 1916. Parasitism among the larvae of the Mediterranean fruit fly (Ceratitis capitata) in Hawaii in 1915. J. Econ. Entomol. 9: 306-11.
Back, E. A. & C. E. Pemberton. 1917. The melon fly in Hawaii. U. S. Dept. Agr. Bull. No. 491. 64 pp.
Back, E. A. & C. E. Pemberton. 1918. The Mediterranean fruit fly in Hawaii. U. S. Dept. Agric. Bull. 536: 1-119.
Baker, A. C., W. E. Stone, C. C. Plummer & M. McPhail. 1944. A review of studies on the Mexican fruitfly and related Mexican species. USDA Misc. Publ. No. 531, Wash., D.C.
Barinaga, M. 1990. Entomologists in the medfly maelstrom. Science 247: 1168-1169.
Baranowski, R. M. 1974. Release of Opius longicaudatus against Anastrepha suspensa in Florida. Fruit Fly News 3: 17.
Baranowski, R. M. & R. W. Swanson. 1971. The utilization of Parachasma cereum (Hymenoptera: Braconidae) as a means of suppression Anastrepha suspensa (Diptera: Tephritidae) populations. Proc. Tall Timbers Conf. on Ecol. Anim. Contr. by Habitat Management 1971: 249-252.
Bartlett, K. A. 1941. The introductions into Puerto Rico of beneficial insects parasitic on West Indian fruit flies. J. Agric. Univ. of Puerto Rico. 25: 25-31.
Bateman, M. A. 1973. The eradication of Queensland fruit fly from Easter Island. FAO Plant Protec. Bull. 21: 114.
Bellows, T. S., Jr. & T. W. Fisher, (eds) 1999. Handbook of Biological Control: Principles and Applications. Academic Press, San Diego, CA. 1046 p.
Benjamin, F. H. 1934. Description of some native trypetid flies with notes on their habits. U. S. Dept. Agr. Tech. Bull. No. 401. 95 pp.
Bennett, F. D., D. Rosen, P. Cochereau & B. J. Wood. 1976. Biological control of pests of tropical fruits and nuts. In: C. B. Huffaker & P. S. Messenger (eds.), Theory and Practice of Biological Control. Academic Press, New York, San Francisco, London. pp. 359-395.
Bennett, F. D., M. Yaseen, M. N. Beg & M. J. Sommeijer. 1977. Anastrepha spp.--Investigations on their natural enemies and establishment of Biosteres longicaudatus in Trinidad, West Indies. Commonw. Inst. Biol. Control Tech. Bull. 18: 1-12.
Berlese, A. 1911. Diffusione in Italia di un Opius australiano contro il Dacus oleae. Redia 7: 470.
Bess, H. A. 1953. Status of Ceratitis capitata in Hawaii following the introduction of Dacus dorsalis and its parasites. Proc. Hawaiian Entomol. Soc. 15: 221-34.
Bess, H. A., R. van den Bosch & F. Haramoto. 1950. Progress and status of two recently introduced parasites of the Oriental fruit fly, Daucs dorsalis Hendel, in Hawaii. Proc. Hawaiian Entomol. Soc. 14: 29-33.
Bess, H. A., R. van den Bosch & F. H. Haramoto. 1961. Fruit fly parasites and their activities in Hawaii. Proc. Hawaiian Entomol. Soc. 17: 367-378.
Bianchi, F. A. & N. Krauss. 1936. Report on the USDA East-African fruit fly expedition of 1935-36. U. S. Dept. Agr. Bur. Entomol. Plant Quarantine Rept. 33 pp.
Bianchi, F. A. & N. Krauss. 1937. Fruit fly investigations in East Africa. Hawaiian Planters' Record 41: 299-306.
Bou ek, Z. & T. C. Narendran. 1981. Indian chalcid wasps (Hymenoptera) of the genus Dirhinus parasitic on synanthropic and other Diptera. Systematic Entomol. 6: 229-251.
Bravo-Mojica, H. 1959. Parasitismo en larvas de diferentes edades y mejor relación parásito huésped de Syntomosphyrum indicum Silv., un enemigo natural de moscas de la fruta. Unpub. B.S. thesis, Escuela Nacional de Agricultura. Chapingo, México. 29 pp.
Breme, F. de. 1842. Note sur le genre Ceratitis de Macleay. Ann. Soc. Entomol. de France 11: 183-90.
Bridwell, J. C. 1914. Report for the period from October 1 to December 31 (1913). Hawaii Bd. Commrs. Agr. & Forest., Div. Entomol. Bull. No. 3: 154-65.
Bridwell, J. C. 1916. Breeding fruit-fly parasites in the Hawaiian Islands. J. Econ. Entomol. 9: 412-6.
Bush, G. L. 1966. The taxonomy, cytology and evolution of the genus Rhagoletis in North America (Diptera: Tephritidae). Harvard Univ. Mus. Compar. Zool. Bull. 134: 431-562.
Byrd, I. B. 1960. What are the side effects of the imported fire ant control program? Biological Problems in Water Pollution Trans., 1959 seminar. U. S. Public Health Service Tech. Rept. W60-3, 46-50.
California Department of Food and Agriculture. 1990. Medfly information kit. Tech. Publ. CDFA, April 30, 1990.
Carey, J. R. 1982. Demograph and population dynamics for the Mediterranean fruit fly. Ecol. Modeling 16: 125-50.
Carey, J. R. 1984. Host specific demographic studies of the Mediterranean fruit fly (Ceratitis capitata). Ecol. Ent. 9: 261-70.
Carey, J. R. & R. V. Dowell. 1989. Exotic fruit fly pests and California agriculture. Calif. Agric. 43(3): 38-40.
Carter, H. O. 1990. Testimony on the economic implications of the Medfly infestation: assembly hearings. University of California Agricultural Issues Center, March 6, 1990.
Carter, W. 1950. The oriental fruit fly: Progress on research. J. Econ. Entomol. 43: 677-83.
Christensen, L. D. & W. E. Stone. 1956. Mediterranean fruit fly invades Costa Rica. Calif. Citrogr. Feb 1956. 4 pp.
Citrograph. 1980. Eradication of Medfly by sterile-male release: a case study. January 1980, p. 63.
Citrograph. 1990. The Medfly: A man with a super plan. pp. 163-164.
Clancy, D. W., P. E. Marucci & R. Dresner. 1952. Importation of natural enemies to control the Oriental fruit fly in Hawaii. J. Econ. Entomol. 45: 85-90.
Clausen, C. P. 1956. Biological control of fruit flies. J. Econ. Entomol. 49: 176-178.
Clausen, C. P. 1978. Tephritidae (Trypetidae, Trupaneidae). In: Introduced parasites and predators of arthropod pests and weeds: A world review; pp. 320-24. Agric. Handbk. No. 480. ARS, USDA, U. S. Govt. Printing Off., Wash., D.C. 546 pp.
Clausen, C. P., D. W. Clancy & Q. C. Chock. 1965. Biological control of the Oriental fruit fly (Dacus dorsalis Hendel) and other fruit flies in Hawaii. U. S. Dept. Agric. Tech. Bull. 1322: 1-102.
Clawson, S. G. 1959. Fire ant eradication--the quail. Alabama Conservation 30(4): 14.
Cochereau, P. 1970. Les mouches des fruits et leurs parsites dans la zone Indo-Australo-Pacifique et particulierment en Nouvelle Caledonie. Cahiers d'Office Recherche de Science et Technologie d'Outre-Mer (Biologie) 12: 15-50.
Compere, G. 1910. Fruit flies. 38th Fruit Growers' Convention of California, 1910. pp. 106-9.
Compere, G. 1912. A few facts concerning the fruit flies of the World. Calif. Dept. Agric. Mon. Bull. 1: 707-370, 842-845, 907-911, 929-932.
Coquillett, D. W. 1899. A new trypetid from Hawaii. Entomol. News 10: 129-30.
Corbett, G. H. 1937. Division of Entomology Annual Report for the Year 1936. Dept. Agr., South Seas and Federated Malay States, Gen. Ser. Dept. Agr. Straits Settl. 26: 29-48.
Cottam, C. 1959. The uncontrolled use of pesticides in the southeast. Address to Southeastern Assn. Fish, Game and Conservation Commissioners. October 1959.
Coulson, J. R., A. Carrell & D. L. Vincent. 1988. Releases of beneficial organisms in the United States and Territories--1981. U. S. Dept. Agr., Misc. Publ. No. 1464. 324 pp.
Croft, B. A. & M. T. AliNiazee. 1996. Biological control in deciduous tree fruit crops. Principles and Application of Biological Control. University of California Press, Berkeley, CA. (in press).
Cunningham, R. T., W. Routhier, E. J. Harris, G. Cunningham, L. Johnson, W. Edwards, R. Rosander & W. G. Vettle. 1980. A case study: Eradication of Medfly by sterile release. Citrograph (Jan.): 63-7.
DeBach, P. 1974. Biological Control by Natural Enemies. Cambridge Univ. Press. 323 pp.
Delucchi, V. 1977. La lutte biologique contre les revageurs d l'olivier. In: J. Humanes Guillen & J. M. Philippe (eds.), Manuel d'oleiculture. United Nations Food & Agric. Org., Rome. pp. 207-227.
Dresner, E. 1954. Observations on the biology and habits of pupal parasites of the Oriental fruit fly. Proc. Hawaiian Entomol. Soc. 15: 299-310.
Etia, P. M. 1979. Climate. In: Atlas de la Republique Unie du Cameroun (G. Laclavere ed.). Editions Jeune Afrique, Paris. 16-19.
Eskafi, F. M. 1987. Host plants of fruit flies (Diptera: Tephritidae) of economic importance in Guatemala. The Florida Entomol. 70: 116-123.
Eskafi, F. M. 1988. Infestation of citrus by Anastrepha spp. and Ceratitis capitata (Diptera: Tephritidae) in high coastal plains of Guatemala. Environ. Entomol. 17: 52-58.
Eskafi, F. M. 1990. Parasitism of fruit flies Ceratitis capitata and Anastrepha spp. (Diptera: Tephritidae) in Guatemala. Entomophaga 35: 355-62.
Fischer, M. 1973. Hymenoptera Braconidae [Opiinae I.]. Das Tierreich 91: 620 pp.
Fonseca, J. Pinto da & M. Autuori. 1940. Processos de criacao da "Vespinha africana" parasita de "mosca de Mediterraneo." O. Biologico 6: 354-351.
Foote, R. H. 1967. A catalogue of the Diptera of the Americas south of the United States. No. 57: Family Tephritidae. Dept. de Zool., Sec. da Agr., Sao Paulo. 91 pp.
Foote, R. H. 1980. Fruit fly genera south of the United States. U. S. Dept. Agr. Tech. Bull. No. 1600. 79 pp.
Froggatt, W. W. 1899. Notes on fruit-maggot flies with descriptions of new species. Agr. Gaz. New South Wales 10: 497-504.
Froggatt, W. W. 1910. Notes on fruit flies with description of new species. Proc. Linn. Soc. N. S. Wales 35: 862-72.
Fullaway, D. T. 1914. Report for the period from May 16 to Sept. 30 Territory of Hawaii, Board of Commissioners for Agriculture and Forestry, Division of Entomology Bulletin 3: 148-153.
Fullaway, D. T. 1915. Report of the work of the insectary. Report to the Board of Commissioners for Agriculture and Forestry, Territory of Hawaii. 1914: 143-151.
Fullaway, D. T. 1920. The melonfly: its control in Hawaii by a parasite introduced from India. Hawaiian Forester & Agriculturist 17: 101-105.
Fullaway, D. T. 1936. Fruit flies and their parasites in Brazil, 1935-36. United States Dept. of Agric., Bureau of Entomol. & Plant Quarantine Rept. 6pp.
Fuller, C. 1897. The fruit fly. J. Bur. Agr. W. Australia, March: 1185-6.
Galt, D. & B. Albertson. 1981. Potential economic impact of the Mediterranean fruit fly, Ceratitis capitata (Wied.), upon establishment in California. Univ. of Calif. Cooperative Extension and Giannini Foundation of Agricultural Economics, December 1981.
Gilmore, J. E. 1983. Introduction to the symposium, "The Medfly in California: The Threat, Defense Strategies and Public Policy." HortScience 18(1): Feb. 1983.
Gilstrap, F. E. & W. G. Hart. 1987. Biological control of the Mediterranean fruit fly in the United States and Central America. U. S. Dept. of Agric. ARS-56: 1-64.
Gingrich, R. E. 1987. Demonstration of Bacillus thuringiensis as a potential control agent for the adult Mediterranean fruit fly, Ceratitis capitata (Wied.). J. Appl. Entomol. 104: 378-385.
Girard, A. 1906. The spread of the fruit fly in the neighborhood of Paris. Compt. Rend. Acad. Sci., Paris 143: 353-4.
Goeden, R. D. 1985. Host-plant relations of Trupanea spp. (Diptera: Tephritidae) in southern California. Proc. Entomol. Soc. Wash. 87: 564-571.
Goeden, R. D. 1987a. Host-plant relations of native Urophora spp. (Diptera: Tephritidae) in southern California. Proc. Entomol. Soc. Wash. 89: 269-274.
Goeden, R. D. 1987b. Life history of Trupanea conjuncta (Adams) on Trixis californica Kellogg in southern California (Diptera: Tephritidae). Pan-Pac. Entomol. 63: 284-291.
Goeden, R. D. 1988. Gall formation by the capitulum-infesting fruit fly, Tephritis stigmatica (Diptera: Tephritidae). proc. Entomol. Soc. Wash. 90: 37-43.
Goeden, R. D. 1989. Host-plants of Neaspilota in California (Diptera: Tephritidae). Proc. Entomol. Soc. Wash. 91: 164-168.
Goeden, R. D. & F. L. Blanc. 1986. New synonomy, host-preference, and California records in the genera Dioxyna and Paroxyna (Diptera: Tephritidae). Pan-Pac. Entomol. 62: 88-90.
Goeden, R. D. & D. W. Ricker. 1971. Biology of Zonosemata vittigera relative to silverleaf nightshade (Diptera: Tephridae). J. Econ. Entomol. 64: 417-21.
Gonzales-A., T. 1963. Informe actividades sección mosca de las frutas. Anexo 1 (pp. 1-19). Proc. OIRSA 11 Reunión del Comité Inrternacional Regional de Sanidad Agropecuaria. 100 pp.
Gonzales-A., T. 1966. Informe del Programa ampliado de control biológico de la mosca de Mediterráneo. Documento III (pp. 35-57). Proc. OIRSA 14 Reunión del Comité Internacional Regional de Sanidad Agropecuaria. 105 pp.
Gonzales-A., T. 1972. Programa de control biológico de las moscas de las frutas. Documento III (pp. 12-30). Proc. OIRSA 20 Reunión del Comité Internacional Regional de Sanidad Agropecuaria. 95 pp.
Gonzales-A., T. 1981. Programa cooperativo Costa Rica - OIRSA de investigaciones biológicas. Parte VI (pp. 99-118). Proc. 29 Reunión del Comité Internacional Regional de Sanidad Agropecuaria. 182 pp.
Gonzalez-hernandez, A. & L. O. Tejada. 1979. Fluctuación de la población de Anastrepha ludens (Loew) y de sus enemigos naturales en Sargentia greggii S. Watts. Filia Entomol. Mexicana 41: 49-60.
Greathead, D. J. 1971. A review of biological control in the Ethiopian Region. Commonw. Inst. Biol. Contr. Tech. Comm. 5: 1-162.
Greathead, D. J. 1972. Notes on coffee fruit flies and their parasites at Kawanda, Uganda. Commonw. Ins. Biol. Control, Tech. Bull. 15: 11-18.
Greathead, D. J. 1976. Mediterranean fruit fly, olive fly. In: D. J. Greathead (ed.). A review of biological control in western and southern Europe. Commonw. Inst. Biol. Contr. Tech. Comm. 7: 37-43.
Greathead, D. J. & J. K. Waage. 1983. Opportunities for biological control of agricultural pests in developing countries. World Bank Tech. Paper 11: 1-44.
Gurney, W. B. 1936. In search of fruit-fly parasites in India. A summary of investigations made in 1935. Agricultural Gazette. New South Wales 47: 374-378.
Gutierrez-S., J. 1976. La mosca del Mediterráneo, Ceratitis capitata (Wiedemann), y los factores ecológicos que favorecerían su establecimiento y propagación en México. Secretaría de Agricultura y Ganadería, Dirección General de Sanidad Vegetal, México. 233 pp.
Hadden, F. C. 1936. Fruit flies and their parasites in Malaya, Ceylon and India. U. S. Dept. Agr. Bur. Entomol. Plant Quarantine Rept. 7 pp.
Hagen, K. S., W. W. Allen & R. L. Tassan. 1981. Mediterranean fruit fly: The worst may be yet to come. Calif. Agric. 35(3-4): 5-7.
Hancock, D. L. 1984. Ceratitinae [Diptera: Tephritidae] from the Malagasy subregion. J. Entomol. Soc. Sth. Afr. 47: 277-301.
Haramoto, F. H. & H. A. Bess. 1970. Recent studies on the abundance of the Oriental and Mediterranean fruit flies and the status of their parasites. Proc. Hawaiian Entomol. Soc. 20: 551-566.
Headrick, D. and R. D. Goeden. 1989. Life history of Pteromalus coloradensis (Ashmead) (Hymenoptera: Pteromalidae) a parasite of Paracantha gentilis Hering (Diptera: Tephritidae) in Cirsium thistle capitula. Proc. Entomol. Soc. Wash. 91: 594-603.
Herrera, A. L. 1905. El gusano de la naranja. Biol. Com. Parasitol. Agric. (México), Tomo II: 307-415.
Hinckley, A. D. 1965. Fruit fly infestation and parasitization in Fiji. Proc. Hawaiian Entomol. Soc. 19: 91-95.
Houston, W. W. K. 1981. Fluctuations in numbers and the significance of the sex ratio of the Mexican fruit fly, Anastrepha ludens caught in McPhail traps. Entomol. Exp. & Appl. 30: 140-150.
Howard, L. O. 1917. The practical use of the insect enemies of injurious insects. U. S. Dept. Agric. Yearbook 1916: 282.
Hunt, E. G. & A. I. Bischoff. 1960. Inimical effects on wildlife of periodic DDD applications to Clear Lake. Calif. Fish & Game 49(1): 91-106.
Ingram, W. R. 1965. An evaluation of several insecticides against berry borer and fruit fly in Uganda robusta coffee. E. Afr. Agric. For. J. 30: 259-262.
Jimenez-Jimenez, E. 1956. Las moscas de la fruta y sus enemigos naturales. Fitófilo (México) 9: 4-11.
Jimenez-Jimenez, E. 1961. Resumen de los trabajos de control biológico que se efectuan en México para el combate de plagas agrícolas. Fitófilo (México) 14: 9-15.
Knipling, E. F. 1986. The estimated relative effects of Biosteres oophilus and B. longicaudatus releases on rates of parasitism and on the dynamics of Dacus dorsalis populations in Hawaii. USDA Working Paper, USDA-ARS.
Krainaker, D. A., J. R. Carey and R. I. Vargas. 1987. Effect of larval host on life history traits of the Mediterranean fruit fly, Ceratitis capitata. Oecologia (Berlin) 73: 583-90.
Laing, J. E. & J. Hamai. 1976. Biological control of insect pests and weeds by imported parasites, predators, and pathogens. In: C. B. Huffaker & P. S. Messenger (eds.). Theory and Practice of Biological Control. Academic Press, New York, San Francisco, London. pp. 685-743.
Landeros, J. F. 1978. Evaluación de algunos factores reguladores de la población de Anastrepha ludens (Loew) en su hospedera silvestre Sargentia greggii Watts, en la región de Rincón de la Sierra, Guadalupe, Nuevo León, México. Unpubl. B.S. thesis, ITESM. 53 pp.
Le Pelley, R. H. 1968. Pests of coffee. Longmans, London. 590 pp.
Le Pelley, R. H. 1973. Coffee insects. Ann. Rev. Entomol. 18: 121-142.
1986 Legner, E. F. 1986. Importation of exotic natural enemies. In: pp. 19-30, "Biological Control of Plant Pests and of Vectors of Human and Animal Diseases." Fortschritte der Zool. Bd. 32: 341 pp.
1987 Legner, E. F. 1987. The importance of single species in determining the average density of plants and animals. Proc. Calif. Mosq. & Vector Contr. Assoc., Inc. 55: 121-123.
1987 Legner, E. F. & R. D. Goeden. 1987. Larval parasitism of Rhagoletis completa (Diptera: Tephritidae) on Juglans microcarpa (Juglandaceae) in western Texas and southeastern New Mexico. Proc. Entomol. Soc. Wash. 89(4): 739-743.
Lindergren, J. E. & P. V. Vail. 1986. Susceptibility of Mediterranean fruit fly, melon fly and oriental fruit fly [Diptera: Tephritidae] to the entomogenous nematode Steinernema feltiae in laboratory tests. Environ. Entomol. 15: 465-468.
Lindegren, J. E., T. T. Wong & D. O. McInnis. 1990. Response of the Mediterranean fruit fly (Diptera: Tephritidae) to the entomogenous nematode Steinernema feltiae in field tests in Hawaii. Environ. Ent. 19: 383-86.
Liquido, N. J., R. T. Cunningham & S. Nakagawa. 1990. Host plants of Mediterranean fruit fly (Diptera: Tephritidae) on the island of Hawaii (1949-1985 survey). J. Econ. Ent. 83: 1863-78.
Macleay, W. S. 1829. Notice of Ceratitis citriperda, an insect very destructive to orange. Zool. J. 4: 475-82.
McBride, O. C. & Y. Tanada. 1949. A revised list of host plants of the melon fly in Hawaii. Proc. Hawaiian Entomol. Soc. 13: 411-21.
Messenger, P. S., & N. E. Flitters. 1954. Bioclimatic studies of three species of fruit flies in Hawaii. J. Econ. Entomol. 47: 756-65.
Mitchell, W. C., C. O. Andrew, K. S. Hagen, R. A. Hamilton, E. J. Harris, K. L. Maehler & R. A. Rhode. 1977. The Mediterranean fruit fly and its economic impact in Central American countries and Panama. University of California/Agency for International Development Pest Management and Related Environmental Protection Project. 189 pp.
Monastero, S. 1931. Un nuovo parassita endofago della mosca delle olive trovato in Altavilla Milicia (Sicilia) (Fam. Braconidae Gen. Opius). Atti della Reale Accademia Scienze, Palermo 16: 195-200.
Morales, M. E. 1963. The Mediterranean fruit fly, Ceratitis capitata (Wied.), problem in Costa Rica. Unpubl. M.S. Thesis, University of Florida. 215 pp.
Moutia, A. 1934. Report of the Department of Agriculture, Mauritius. Entomological Division 1933: 25-29.
Munro, H. K. 1964. Some fruit flies of economic importance in South Africa. Dept. Agr. Tech. Serv. Rep. S. Afr. Circ. 18 pp.
Narayanan, E. S. & S. S. Chawla. 1962. Parasites of fruit fly pests of the World with brief notes on their bionomics, habits and distribution. Beitrage zur Entomologie 12: 437-476.
Newell, I. M. & F. H. Haramoto. 1968. Biotic factors influencing populations of Dacus dorsalis in Hawaii. Proc. Hawaiian Entomol. Soc. 20: 81-139.
Newell, I. M., W. C. Mitchell & F. L. Rathburn. 1952. Infestation norms for Dacus cucurbitae in Momordica balsamina, and seasonal differences in activity of the parasite, Opius fletcheri. Proc. Hawaiian Entomol. Soc. 14: 497-508.
Newell, W. 1930. The Mediterranean fruit fly situation. J. Econ. Ent. 23: 512-23.
Nishida, T., & F. Haramoto. 1953. Immunity of Dacus curcubitae to attack by certain parasites of Dacus dorsalis. J. Econ. Entomol. 46: 61-4.
Nishida, T. 1955. Natural enemies of the melon fly, Dacus cucurbitae Coq. in Hawaii. Ann. Entomol. Soc. Amer. 48: 171-178.
Nishida, T., E. J. Harris, R. I. Vargas & T. T. Y. Wong. 1985. Distributional loci and host fruit utilization patterns of the Mediterranean fruit fly, Ceratitis capitata (Diptera: Tephritidae), in Hawaii. Environ. Ent. 14: 602-606.
Noble, N. S. 1942. Melittobia (Syntomosphyrum) indicum (Silv.) (Hymenoptera, Chalcidoidea), a parasite of the Queensland fruit fly, Strumeta tryoni (Frogg.). Linn. Soc. N. S. Wales. Proc. 67: 269-76.
O'Connor, B. A. 1960. A decade of biological control work in Fiji. Agricultural Journal, Fiji Dept. of Agric. 30: 44-54.
Orian, A. J. E. & L. A. Moutia. 1960. Fruit flies (Trypetidae) of economic importance in Mauritius. Revue Agricole et Sucriere de l'Ile Maurice 39: 142-150.
Ortiz, H. J. J. 1958. Observaciones sobre la biología de la mosca de la fruta Anastrepha ludens y sus enemigos naturales en la zona cítrica de Río Ramos, Nuevo León, México y sus alrededores. Unpubl. B. S. thesis, Univ. de Nuevo León, México.
Pemberton, C. E. & H. F. Willard. 1918a. Fruitfly parasitism in Hawaii during 1916. J. Agr. Res. 12: 103-8.
Pemberton, C. E. & H. F. Willard. 19l8b. Interrelations of fruit-fly parasites in Hawaii. J. Agric. Res. 12: 285-295.
Pemberton, C. E. & H. F. Willard. 1918c. Work on parasitism of the Mediterranean fruit fly in Hawaii in 1917. J. Agr. Res. 14: 605-10.
Pemberton, C. E. & H. F. Willard. 1918c. A contribution to the biology of fruit fly parasites in Hawaii. J. Agr. Res. 15: 419-65.
Podoler, H. & Z. Menzel. 1977. Analysis of solitariness in a parasite-host system (Muscidifurax raptor, Hymenoptera: Pteromalidae--Ceratitis capitata, Diptera: Tephritidae). Ecol. Ent. 2: 153-60.
Podoler, H. & Z. Menzel. 1979. Analysis of a host-parasite (Ceratitis-Muscidifurax) relationship under laboratory conditions. Ecol. Ent. 4: 45-59.
Quaintance, A. L. 1912. The Mediterranean fruit-fly. U. S. Dept. Agr. Bur. Entomol. Cir. No. 160. 25 pp.
Quayle, H. J. 1938. Insects of Citrus and Other Subtropical Fruits. Ithaca, N. Y., Comstock Publ. Co., Inc. 538 pp.
Rao, V. P., M. A. Ghani, T. Sankaran & K. C. Mathur. 1971. A review of the biological control of insects and other pests in Southeast Asia and the Pacific Region. Commonwealth Inst. Biol. Contr. Tech. Commun. 6: 43-45, 81-83.
Rhode, R. H. 1976. The history and current status of the Mediterranean fruit fly in North America. J. Rio Grande Valley Hort. Soc. 30: 11-17.
Roder, V. 1885. Ceratitis capitata. Berl. Ent. Zeitschr. 29: 132.
Rondani, C. 1870. Ortalidinae italicae, collectae, distinctae et en ordinem dispositae. Bull. Soc. Entomol. Ital. 2: 29.
Ruiz-Cancino, E. 1979. Parasitismo natural en Anastrepha ludens (Loew.) en hospederas silvestres y cultivadas en zona centro de Tamaulipas, México. Unpubl. Seminario de Investigación II, Univ. Autónoma de Tamaulipas. Nov. 1979. 20 pp.
Russo, G. 1959. Esperimenti di lotta biologica contro la mosca delle olive (Dacus oleae) e la mosca della frutta (Ceratitis capitata) in Italia. Bollettino del Laboratorio di Entomologia Agraria "Filippo Silvestri" Portici 17: 131-142.
San Juana, S. 1974. Control biológico de las moscas de la fruta en el CaÁon del Tomellín, Oaxaca, México. Dir. Gen. Sanidad Vegetal, SARH.
Scribner, J. 1983. The Medfly in California: Organization of the eradication program and public policy. HortScience 18(1): Feb. 1983.
Silverman, J. & R. D. Goeden. 1980. Life history of a fruit fly, Procecidochares sp. on the ragweed, Ambrosia dumosa (Gray) Payne, in southern California (Diptera: Tephritidae). Pan-Pacific Entomol. 56: 283-288.
Silvestri, F. 1913. Viaggio in Africa per cercare parassiti mosche dei frutti. Bull. Lab. di Zollogia generale e agraria, Portici 8: 1-164.
Silvestri, F. 1914. Report on an expedition to Africa in search of the natural enemies of fruit flies (Trypaneidae) with descriptions, observations and biological notes. Territory of Hawaii, Board of Agriculture and Forestry, Division of Entomology Bull. 3: 1-176.
Silvestri, F. 1915. Descrizione di nuovi imenotteri calcididi africani. Boll. Lab. Zool. gen. agr. Portici 9: 337-77.
Silvestri, F. 1939. La lotta biologica contro le mosche dei frutti della famiglia Trypetidae. Proc. VII Intern. Congress of Entomology, Berlin 4: 2396-2418.
Smith, H. S. 1929. Multiple parasitism: its relation to the biological control of insect pests. Bull. Entomol. Res. 20: 141-9.
Snowball, G. J. 1966. Status of introduced parasites of Queensland fruit fly (Strumeta tryoni). 1962-1965. Australian J. Agric. Res. 17: 719-739.
Spitler, G. H. & H. M. Couey. 1983. Methyl bromide fumigation treatments of fruits infested by the Mediterranean fruit fly (Diptera: Tephritidae). J. Econ. Ent. 73(3): June 1983.
Steck, G. J., F. E. Gilstrap, R. A. Wharton & W. G. Hart. 1986. Braconid parasitoids of Tephritidae (Diptera) infesting coffee and other fruits in West-Central Africa. Entomophaga 31: 59-67.
Stone, A. 1942. The fruit flies of the genus Anastrepha. U. S. Dept. Agr. Misc. Publ. No. 439. 112 pp.
Stone, A., C. W. Sabrosky, W. W. Wirth, R. H. Foote & J. R. Coulson. 1965. A catalog of the Diptera of America north of Mexico. U. S. Dept. Agr. Handb. No. 279: 1696 pp.
Turica, A. 1968. Lucha biologica como medio de control de las moscas de los frutos. Idia. 241: 29-38.
UC/AID Pest Management & Environmental Protection Project. 1977. The Mediterranean fruit fly and its economic impact on Central American countries and Panama. 1977 Report.
U. S. Department of Agriculture Annual Report. 1988. Proyecto piloto de control biológico de moscas de la fruta, mediante liberaciones masivas de parasitoides en mazapa de madero, Chiapas, México. Tapachula, Chiapas, Mexico. U.S.D.A. Contract a53-6395-7-50. 16 pp.
van den Bosch, R. & F. H. Haramoto. 1951. Opius oophilus Fullaway, an egg-larval parasite of the Oriental fruit fly discovered in Hawaii. Proc. Hawaiian Entomol. Soc. 14: 251-5.
van den Bosch, R. & F. H. Haramoto. 1953. Competition among parasites of the Oriental fruit fly. Proc. Hawaiian Entomol. Soc. 15: 201-6.
van den Bosch, R., H. A. Bess & F. H. Haramoto. 1951. Status of Oriental fruit fly parasites in Hawaii. J. Econ. Entomol. 44: 753-9.
van Zwaluwenburg, R. H. 1936. Report on an expedition to west Africa in search of fruit fly parasites (1935-36). U. S. Dept. Agr. Bur. Entomol. Plant Quar. Rept. 20 pp.
van Zwaluwenburg, R. H. 1937. West African Notes. The Hawaiian Planters' Record 41: 57-83.
Weems, H. V. 1981. Mediterranean fruit fly, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae). Fla. Dept. Agr. & Consumer Serv., Div. Plant Industry, Entomol. Cir. No. 230. 8 pp.
Wharton, R. A. 1989. Chapter 9.1 Classical Biological Control of Fruit-infesting Tephritidae. pp. 303-313. In: A. S. Robinson & G. Harper (eds.). World Crop Pests. Fruit Flies , Their Biology, Natural Enemies and Control. Vol. 3B. Elsevier.
Wharton, R. A. & F. E. Gilstrap. 1983. Key to and status of opiine braconid (Hymenoptera) parasitoids used in biological control of Ceratitis and Dacus s. l. (Diptera: Tephritidae). Ann. Entomol. Soc. Amer. 76: 721-742.
Wharton, R. A., F. E. Gilstrap, R. H. Rhode, M. Fischel-M & W. G. Hart. 1981. Hymenopterous egg-pupal and larval-pupal parasitoids of Ceratitis capitata and Anastrepha spp. [Dip.: Tephritidae] in Costa Rica. Entomophaga 26: 285-290.
Wiedemann, C. R. W. 1824. Munus rectoris in Acadamia Christiana Albertina aditurus Analecta entomologica ex Museo Regio Havniesi maxime congesta profert inconibusque illustrat. 60 p. Kiel.
Willard, H. F. 1920a. Opius fletcheri as a parasite of the melon fly in Hawaii. J. Agr. Res. 20: 423-38.
Willard, H. F. 1920b. Work and parasitism of the Mediterranean fruit fly in Hawaii during 1918. J. Agr. Res. 18: 441-6.
Willard, H. F. 1923. Work on parasitism of the Mediterranean fruit fly in Hawaii during 1919 and 1920. J. Agr. Res. 25: 1-7.
Willard, H. F. & T. L. Bissell. 1926. Work and parasitism of the Mediterranean fruit fly in Hawaii in 1921. J. Agr. Res. 33: 9-15.
Willard, H. F. & T. L. Bissell. 1930. Parasitism of the Mediterranean fruit fly in Hawaii, 1922-24. U. S. Dept. Agr. Cir. 109: 12 pp.
Willard, H. F. & A. C. Mason. 1937. Parasitization of the Mediterranean fruit fly in Hawaii. 1914-33. U. S. Dept. Agric. Circ. 439: 1-18.
Williamson, D. L. 1983. The Medfly in California: methods of attack. HortScience 18(1): Feb. 1983.
Wilson, F. 1960. A review of the biological control of insects and weeds in Australia and Australian New Guinea. Commonw. Inst. Biol. Control. Tech. Comm. 1: 24-26, 50.
Wong, T. T. Y., J. I. Nishimoto & H. M. Couey. 1983. Mediterranean fruit fly (Diptera: Tephritidae): further studies on selective mating response of wild and of unirradiated and irradiated, laboratory-reared flies in field cages. Ann. Entomol. Soc. Amer. 76: 51-55.
Wong, T. T. Y., J. I. Nishimoto & N. Mochizuki. 1983. Infestation patterns of Mediterranean fruit fly and the oriental fruit fly (Diptera: Tephritidae) in the Kula area of Maui, Hawaii. Environ. Entomol. 12: 1031-1039.
Wong, T. T. Y., N. Mochizuki & J. I. Nishimoto. 1984. Seasonal abundance of parasitoids of the Mediterranean and Oriental fruit flies (Diptera: Tephritidae) in the Kula area of Maui, Hawaii. Environ. Entomol. 13: 140-145.
Wong, T. T. Y., R. M. Kobayashi, L. C. Whitehand, D. C. Henry, D. A. Zadig & C. L. Denny. 1984. Mediterranean fruit fly (Diptera: Tephritidae): mating choices of irradiated laboratory-reared and untreated wild flies of California in laboratory cages. J. Econ. Entomol. 77: 58-62.
Wong, T. T. Y., D. O. McInnis, J. I. Nishimoto, A. K. Ota & V. C. S. Chang. 1984. Predation of the Mediterranean fruit fly [Diptera: Tephritidae] by the Argentine ant [Hymenoptera: Formicidae] in Hawaii. J. Econ. Entomol. 77: 1454-1458.
Wong, T. T. Y., D. O. McInnis, R. M. Kobayashi & J. I. Nishimoto. 1985. Distribution and seasonal abundance of adult male Mediterranean fruit flies (Diptera: Tephritidae) in Kula, Maui, Hawaii. J. Econ. Entomol. 78: 552-557.
Wong, T. T. Y., R. M. Kobayashi & D. O. McInnis. 1986. Mediterranean fruit fly (Diptera: Tephritidae): methods of assessing the effectiveness of sterile insect releases. J. Econ. Entomol. 79: 1501-1506.
Wong, T. T. Y., M. M. Ramadan, D. O. McInnis & N. Mochizuki. 1990a. Influence of cohort age and host age on oviposition activity and offspring sex-ratio of Biosteres tryoni (Hymenoptera: Braconidae), a larval parasitoid of Ceratitis capitata (Diptera: Tephritidae). J. Econ. Entomol. 83: (in press).
Wong, T. T. Y. & M. H. Ramadan. 1990b. Mass-rearing biology of larval parasitoids (Hymenoptera: Braconidae: Opiinae) of tephritid flies (Diptera: Tephritidae) in Hawaii. In: Advances in Insect Rearing For "Research and Pest Management," T. E. Anderson & N. C. Leppla (eds.)
Wong, T. T. Y., M. M. Ramadan, D. O. McInnis, N. Mochizuki, J. I. Nishimoto & J. C. Herr. (in press 1990c). Augmentative releases of Diachasmimorpha tryoni (Hymenoptera: Braconidae) to suppress a Mediterranean fruit fly (Diptera: Tephritidae) population in Kula, Maui, Hawaii. Environ. Entomol. (in press).
Yamada, T., S. Nakagawa & H. Kamasaki. 1962. Identification of three species of reared Hawaiian fruit fly pupae. Proc. Hawaiian Entomol. Soc. 18: 319-21.
Young, L. A. & H. P. Nicholson. 1951. Stream pollution resulting from the use of organic insecticides. Progressive Fish Culturist 13(4): 193-98.