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BIOLOGICAL
CONTROL OF FRUIT FLIES (e.legner@ucr.edu) 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.
The Mediterranean fruit fly, although eradicated periodically from the state
of Florida where it had "peninsular" distribution, is presently
firmly established in southern Mexico where it is temporarily contained by a
massive sterile-male and parasite release effort by the U. S. Department of
Agriculture. Eventually this species may move north and pose a continuous
threat along the Mexico-United States 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. Principal
Fruit Flies in California The Mediterranean fruit fly in
particular has received a lot of attention in recent times. It 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). 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). Although parasitic insects have
been imported against it, all except one species were obtained from areas
outside the fly\'s native range in central Africa. However, some reductions
in infestations are attributable to natural enemy activity in the invaded
areas, especially when parasitic insects are mass released as biotic
insecticides (Wong & Ramadan 1990, Wong et al. 1990).
Some investigators believe that the Medfly is already permanently established
in California and that unless the current eradication effort is greatly
increased, it is just a matter of time before it will spread throughout the
state (Barinaga 1990). The Malathion and other baits currently in use against
Medflies 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.
As it becomes increasingly apparent that fruit flies pose continuous threats
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 an invasion as well as within
the state boundaries where breeding may occur, offers an environmentally
sound, non-polluting alternative. There is a need to (1) develop and improve
techniques for the search, procurement and evaluation of natural enemies of
fruit flies in their natural ranges (parasites, predators and pathogens); (2)
introduce and study foreign natural enemies and evaluate their respective
effectiveness under field conditions in Hawaii, southern Mexico, and
California; (3) develop a mass production scheme for periodic releases of introduced
natural enemies in infested areas of California, and to observe their
effectiveness under natural conditions; and (4) build a culture bank of
natural enemies for use in conjunction with other eradication and control
methods (e.g., sterile-male releases) during periodic invasions of this pest
and in anticipation of its possible permanent establishment in the State of
California.
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 damages 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 ants is being attempted with natural
enemies imported from Brazil and Argentina. Biological
Control Efforts Against Fruit Flies
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 an 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 in their 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 Medfly
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. Specific
Examples Medfly.
Cerititis capitata (Wiedemann)-- The Mediterranean fruit fly
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 the 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 ascertain the aboriginal home nor did he establish the
parasites 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 parasitic insects 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 parasitic insects, but only Tetrastichus giffardianus
Silvestri and Biosteres fullawayi (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 parasitic species is
virtually unknown, the notable exception being Hawaii. Even in Hawaii control
was never noteworthy and the medfly problem 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
the 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 combat both
medfly and the oriental fruit fly. Three parasitic species, 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 parasite, B. oophilus
(Haramoto & Bess 1970). During 1978-1981, Biosteres oophilus
was still the predominant parasite as it accounted for ca. 80% of the total
parasitization. Occasionally the larval-pupal parasite, 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 or spherical and
yellowish or 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 parasitic insects 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 natural enemies of the medfly, 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 parasitic species recovered from both flies were B. longicaudatus
and Doryctobracon crawfordi (Viereck) (Eskafi 1990).
The behavior of the ectoparasite 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 parasite
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 parasites 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 originated. The natural range of the 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 (Table 1;
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). The parasitic Hymenoptera are believed to be the
most effective natural enemies of 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.). These have not been tested by entomologists in California 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. Table 1. Known parasitic species
attacking fruit flies of the genus Ceratitis in their natural Central African range ( Parasite species & Host
stage attacked) _________________________________________________________________________________________________________ Biosteres caudatus Szepligeti larva Hedylus sp. larva,
Diachasma fullawayi Silvestri larva, Galesus silvestrii
Kieffer pupa, Diachasmimorpha longicaudata (Ashmead) larva , Microbracon celer (Szepligeti) larva?, Diachasmimorpha tryoni
(Cameron) larva, Opius humilis Silvestri larva, Dirhinus
ehrhorni Silvestri pupa, Opius inconsuetus Silvestri
larva Dirhinus giffardii Silvestri pupa, Hedylus giffardii
Silvestri larva?, Ganaspis sp. larva? Opius n. sp. larva, Halticoptera
sp. larva? , Spalangia afra Silvestri pupa Opius perproximus Silvestri larva, Tetrastichus dacicida
Silvestri larva, Tetrastichus giffardii Silvestri larva,
Tetrastichus oxyurus Silvestri larva, Tetrastichus n.
sp. larva _____________________________________________________________________________________ Mexican fruit fly.
Anastrepha ludens (Loew)--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. Misc. fruit flies--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 parasitic insects
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 parasites 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 parasite 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
1992). Pertinent References: 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.:
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