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Bio-Control Cases Biological resistance among
arthropods to chlorinated hydrocarbon, organophosphate and carbamate
insecticides in the late 1950's resulted in an expected turn toward suitable
alternatives and especially to biological control. Attention was directed to the control of Diptera of medical and
veterinary importance at a 1960 symposium in Washington, D.C. (Anonymous
1960), where biological control possibilities were emphasized. Jenkins (1964) reviewed the literature
listing the known natural enemies of all arthropods of medical and veterinary
importance, noting over 1,500 parasites, pathogens and predators. Renewed research emphasis on natural
enemies followed the Washington symposium and Jenkins' review, and by 1999
there has been a substantial increase in research treating of the existence
and biologies of natural enemies, as well as further reviews of the subject
(Laird 1971a,b,c; Bay 1974; Brown 1973, Chapman 1974, Bay et al. 1976, Legner et al. 1974,
Federici 1981, Murdoch 1982, Service 1983, Legner and Sjogren 1984,
Laird 1986, Garcia and Legner 1999). The American Mosquito Control Association has maintained a
quarterly accounting of publications pertaining to mosquito biological
control agents since the Jenkins (1964) review, and the World Health
Organization began issuing a series of reports in 1979 which described the
characteristics of specific proven biological control organisms. Interest in biological
control of aquatic Diptera actually began in the late 1800's (Lamborn
1890). At that time the possible use
of dragonflies as natural enemies for the control of mosquitoes was clearly
recognized. However, to the present
day the difficulties associated with the colonization and management of these
insects has discouraged their practical use in mosquito control. In the early 1900's the small mosquitofish,
Gambusia affinis* (Baird and Girard) (Microcyprini: Cyprinodontidae), was
stressed for biological control, and being much easier manipulate than
dragonflies, it was quickly utilized and transported throughout the world
during the early decades of the 1900's in attempts to control mosquitoes
(Legner and Sjogren 1984). The mosquitofish, and a few
other natural enemies were employed with some enthusiasm until the
mid-1940's, when all biological control measures were curtailed sharply with
the introduction of synthetic organophosphorus insecticides after World War
II. Their rapid killing power was so
dramatic for flies and mosquitoes, that other control tactics were temporarily
dismissed to a minor role. Interest
in biological control resumed when the succession of insecticides developed
during the 1940s and 1950s began to fail, due to the development of
biological resistance in vector and pest populations and in the 1990's when
environmental contamination became an increasing public concern. Progress in the biological control of
Diptera has been uninterrupted since its revival, even with problems of
establishing pest tolerance levels, and the temporary unstable habitats exploited
by Diptera (Legner and Sjogren 1984). Bay et
al. (1976) noted that dipterous pests are
usually in the adult stage, which is of some advantage for control because it
allows the control action to be taken against the immature stages, thus
eliminating the adult before it can cause problems. However, it is difficult to establish tolerance levels for such
pests. For example, an individual
mosquito can be extremely annoying, which may lead to a reaction for control;
and low population levels of a vector may still transmit a disease. However, reductions of any kind are
desirable in the absence of more effective strategies, even though such
partial controls may seem unacceptable (e.g.,
Service 1983). Setting tolerance
levels for veterinary pests is comparatively more practical than for
humans. The frequently temporary
habitats utilized by aquatic Diptera poses a problem for biological control
in that natural enemies cannot always coexist with pests to thus regulate
their populations. Also, the habitat
exploited by the pests is often only an undesirable extension of human
activity, such as in the cultivation of rice, where the production of
mosquitoes is usually of little concern to the rice producer. As studies on biological
control agents progressed, it became evident that their practical application
for control would not be simple. The
classical biological control approach involving the introduction of exotic
natural enemies followed by substantial and sustained declines in host
population densities have been reported in only a few cases. Often significant decreases in the pest
population density were still not acceptable to the general public or health
authority that desired an even lower population threshold, or investigations
were terminated early before long-term benefits could be recorded. Problems of mass production, packaging and
distribution of biological control agents have burdened commercial
involvement. However, not until the
1990's did the desire for expedient and thorough effectiveness of commercial
insecticides begin to give way to the slower and usually less potent
biological controls. The present review includes
pertinent literature of major dipterous taxonomic groups where some success
has been achieved or where work is currently being conducted on species
breeding in aquatic habitats (mosquitoes, chironomids, blackflies and
tabanids). Emphasis is on biological
control agents that can be manipulated, that have been used successfully,
that are being researched and which show at least some promise for successful
deployment. While progress in the
development of biological control agents has been substantial and current
work is expanding, a present overall evaluation is that biological control
will continue to be implemented only gradually for Diptera of medical and
veterinary importance. The majority
of research is still driven by economic forces in the search for marketable
products, especially evidenced by the disproportionate attention given to
fungal and bacterial pathogens.
However, the importance of maintaining maximum impact of resident
natural enemies is almost universally accepted, and with continued effort,
biolological control should become a major component in the overall strategy
for the control of these important pests (Legner and Sjogren 1984,
Garcia and Legner 1999). Mosquitoes The successful widespread use
of biological control agents against mosquitoes requires a precise
understanding of the ecology of predator/prey and pathogen/host
relationships. The opportunistic
characteristics of many species, including their ability to take advantage of
temporary habitats, coupled with their short generation time, high natural
mortality, great dispersal potential, and other R-strategist characteristics,
pose difficult problems for any biological control agent (Garcia and Legner 1999). Mosquitoes typically exploit many aquatic
habitats. Often a good biological
control agent will have a much narrower range of environmental activity than
the target species. Thus, in many
situations a number of different biological control agents and/or appropriate
methods are necessary to control even one species of mosquito across its
range of exploitable breeding sources. Insectivorous Fish.--Various species of fishes are used for
the biological control of mosquitoes, which together constitute the major
successes in biological control.
However, their usefulness is limited to relatively permanent bodies of
water, where their impact on the target species is usually only partially
successful. Bay et
al. (1976) remarked that many kinds of fish
consume mosquito larvae, but only a few species have been manipulated to
manage mosquito populations. The mosquitofish, G. affinis
<PHOTO> , is the best known
biological mosquito control. Native
to the southeastern United States, eastern Mexico and the Caribbean area, it
was first used as an introduced agent for mosquito control when transported
from North Carolina to New Jersey in 1905 (Lloyd 1987). Later it was introduced to the Hawaiian
Islands to control mosquitoes, and during the next 70 years to over 50
countries. The mosquitofish ranks as
the most widely disseminated biological control agent (Bay 1969, Lloyd 1987). Many of these introductions were to
control Anopheles species that were
transmitting malaria. Hackett (1937)
described its usefulness in malaria control programs in Europe, noting that
the fish had a definite impact on the suppression of the disease. Tabibzadeh et al. (1970) reported an expansive release program in Iran and
concluded that the fish was an important component in malaria
eradication. Nevertheless, Sasa and
Kurihara (1981) and Service (1983) judged that the fish had little impact on
the disease and that most evidence was circumstantial. Gambusia*
spp. no longer are recommended by the World Health Organization for
malaria control programs, primarily because of their harmful interference
with indigenous species of fish (Service 1983, Lloyd 1987). The biological attributes of G. affinis
are a high reproductive capacity, high survivorship, small size, omnivorous
foraging in shallow water, relatively high tolerance to variations in
temperature, salinity and organic waste, which make this species an excellent
biological control agent (Bay et al.
1976,
Moyle 1976, Moyle et al.
1982). Whether this fish leads to
effective mosquito control at practical costs in many situations is still
debated, however. Probably an
accurate assessment is revealed in a statement by Kligler (1930) that
"... their usefulness as larvae-destroyers under local conditions where
vegetation is abundant and micro fauna rich enough to supply their needs
without great trouble, is limited. In
moderately clear canals, on the other hand, or in pools having a limited food
supply, they yielded excellent results ...." In California this fish had
been used extensively for control of mosquitoes in various habitats (Bay et al. 1976). Many mosquito abatement districts in
California have developed technology for culturing, harvesting and winter
storage of the mosquito fish in order to facilitate stocking early in the
spring (Coykendall 1982, 1986). This
is particularly important in the northern rice producing areas of California
where early stocking appears to be of critical importance for build-up of
fish populations to control mosquitoes during late summer. Some results of the use of G. affinis*
these rice fields illustrate the mixed successes achieved in the field. Rice cultivation in California
continuously poses one of the most difficult control problems for Anopheles spp. and Culex species. Hoy and Reed (1971) showed that good
control of Culex tarsalis* Coquillett (Culicidae) could
be achieved at stocking rates of about 480 or more females per ha., and Stewart and Miura (1985) reported
excellent control with similar stocking rates against this mosquito in the
San Joaquin Valley. Although Cx. tarsalis appears to
be controlled effectively by G. affinis*, the control of Anopheles freeborni* Aitken
(Culicidae) in northern California rice fields is less apparent. Hoy et al. (1972)
showed a reduction of An. freeborni* populations at various stocking
rates of about 120 to 720 fish per ha., but the reduction was not nearly as
striking as for Cx. tarsalis. It was suggested that improved control could be achieved by
earlier season stocking, involving
multiple release points in fields and a reliable source of healthy
fish for stocking. Despite an ample
research effort in mass culture, management and storage for G. affinis*
by the State of California (Hoy and Reed 1971), a mass production procedure
has never provided adequate numbers (Downs et al. 1986, Cech and
Linden 1987). Studies of G. affinis*
for control of mosquitoes in wild rice show that relatively high stocking
rates can effectively reduce An. freeborni* and Cx. tarsalis
populations within a three-month period (Kramer et al. 1987). Wild rice
is a more vigorous and taller plant than white rice, requiring only 90
instead of 150 days to mature (Garcia and Legner 1999). Commercial production has been increasing
in the 1980's in California (Kramer et
al. 1988). Kramer et al. (1987) stocked at rates of
1.7 kg./ha. (ca. 2400 fish/kg.)
released in 1/10 ha. wild rice plots, but failed to show a significant
difference in reduction of mosquitoes from plots with no fish. A decrease in numbers of larvae was noted
just prior to harvest which suggested that the fish were beginning to have an
impact on mosquito numbers (Kramer et
al. 1987). The abundance of fish
in these experimental plots, based on recovery after drainage, reached about
100,000 individuals per ha. (ca. 32 kg./ha) or a density of about 10 fish per
square meter, which did not produce significant control. This study was repeated a
year later at the rates of 1.7 and 3.4 kg./ha. of fish. Results showed an average suppression of larvae
(primarily An. freeborni*) of <1 and 0.5 per dip for the low and high rate
respectively, compared to control plots which averaged >4.5 per dip. Fish densities in the second study
surpassed those of the first by about two fold at the 1.7 kg./ha. rate and
three fold at the 3.4 kg./ha. rate, and these greater numbers accounted for
the control differences observed in the second year, although mosquitoes were
not eliminated. Differences between
test plots and control plots were first observed eight weeks after the fish
had been planted and mosquitoes remained under control until the fields were
drained (Kramer et al. 1988). Davey et al. (1974) and Davey and Meisch (1977) showed that at
inundative release rates of 4,800 fish per ha., G. affinis* was effective for control of Psorophora columbiae*
(Dyar and Knab) in Arkansas rice fields.
Fish released at the water flow inlets scattered quickly throughout
the fields. This is an important
attribute for controlling Psorophora
spp. and Aedes spp., whose hatch
and larval development are completed within a few days. A combination of 1,200 G. affinis*
and about 300 green sunfish [Lepomis
cyanellus* Rafinesque (Perciformes:
Centrarchidae)] gave better control than either four times the amount of G. affinis*
or L. cyanellus* used separately.
This synergistic effect reduces logistic problems associated with
having enough fish available at the times fields are inundated. Blaustein (1986) found enhanced control of
An. freeborni* by mosquitofish in California rice fields after the
addition of green sunfish. Addition
of the latter forced mosquitofish to remain longer in protected areas where
mosquitoes were more abundant in order to elude the green sunfish. The lack of available large numbers of fish for stocking fields
either by inundation, such as in Arkansas or for control later in the season
as practiced in California, is the main reason why fish have not been used
more extensively in rice fields (Garcia and Legner 1999). An unusual use of the
mosquito fish by inundative release was reported by Farley and Caton
(1982). The fish were released in
subterranean urban storm drains to control Culex quinquefasciatus*
Say (Culicidae) breeding in entrapped water at low points in the system. Fish releases were made following the last
major rains to avoid having them flushed out of the system. Fish survived for more than three months
during the summer and were found throughout the system. Gravid females produced progeny, but no subsequent
mating occurred, and after the initial increase in numbers fish populations
declined as summer progressed.
Reductions of mosquitoes from 75 to 94% were observed for three months
compared to untreated areas (Mulligan et
al. 1983). This control practice
is now conducted on a routine basis by the Fresno Mosquito Abatement District
(Garcia and Legner 1999). Although G. affinis* has been
useful for control of mosquitoes in a number of situations, there are definitely
some environmental drawbacks to its use.
This fish probably never would have been intentionally introduced into
foreign areas if today's environmental concerns existed in the early 1900's
(Pelzman 1975, Lloyd 1987). A major
objection to mosquitofish has been their direct impact on native fishes
through predation, or their indirect impact through competition (Bay et al. 1976, Schoenherr 1981, Lloyd
1987). More than 30 species of native
fish have been adversely affected by the introduction of Gambusia* spp. (Schoenherr 1981, Lloyd 1987). Introductions of Gambusia* spp. have also reduced numbers of other aquatic
organisms coinhabiting the same waters (Hoy et al. 1972, Farley and Younce 1977a,b; Rees 1979, Walters and
Legner 1980,
Hurlbert and Mulla 1981). However,
there are no reports of this species, through its feeding on zooplankton
(Hurlbert and Mulla 1981, Hurlbert et
al. 1972) causing algal blooms outside of the experimental aquarium
environment (Walters and Legner 1980). Another widely used fish for
mosquito control is the common guppy, Poecilia
reticulata* (Peters)
(Microcyprini: Cyprinodontidae),
which has been deployed successfully in Asia for the control of waste water
mosquitoes, especially Cx. quinquefasciatus. Like their poeciliid relatives, Gambusia* spp., they are native to
tropical South America. But, rather
than being intentionally introduced to control mosquitoes, this fish was
spread to other parts of the world through the tropical fish trade. Sasa et al. (1965)
observed feral populations of this fish breeding in drains in Bangkok and
concluded from their observations that it was controlling mosquitoes common to
that habitat. The practical use of
guppies is primarily restricted to subtropical climates because they do not
tolerate low temperate-zone water temperatures (Sasa and Kurihara 1981). However, their most important attribute is
a tolerance to relatively high levels of organic pollutants, which makes them
ideal for urban water sources that are rich in organic wastes (Sjogren
1972). In Sri Lanka, wild populations
have been harvested and used for the control of mosquitoes in abandoned
wells, coconut husks and other sources rich in organic rubbish (Sasa and
Kurihara 1981, Sabatinelli 1990).
This fish also now occurs in
India, Indonesia and China and has been intentionally introduced for
filariasis control into Burma (Sasa and Kurihara 1981). Mian et al. (1986)
evaluated its use for control of mosquitoes in sewage treatment facilities in
southern California and concluded that guppies showed great potential for
mosquito control in these situations. Imported fish have also been
used to clear aquatic vegetation
from waterways which concurrently produced excellent mosquito control. In the irrigation canals and drains of
southeastern California, which extend to over 8,000 km., three species of
subtropical cichlids <PHOTO>,
Tilapia zillii (Gervais) (Percomorphi:
Cichlidae), Oreochromis mossambica* (Peters)
(Percomorphi: Cichlidae) and Oreochromis
hornorum* (Trewazas)
(Percomorphi: Cichlidae) were
introduced and became established over some 2,000 ha. of Cx. tarsalis breeding
habitat (Legner and Sjogren 1984). In this situation, mosquito populations are under control by a
combination of direct predation and the consumption of aquatic plants by these
omnivorous fishes (Legner and Medved 1973, Legner 11978a, 1978b, 1983;
Legner and Fisher 1980, Legner and Murray 1981 , Legner and Pelsue 1980,
1983). This is a unique example of persistent
biological control and probably only apropos for relatively sophisticated
irrigation systems where a permanent water supply is assured, and water
conditions are suitable to support the fish (Legner et al. 1980). Advantages in the use of these fish are the clearing of
vegetation to keep waterways open, mosquito control, and the fish are large
enough to be captured for human consumption.
Some sophistication is necessary when stocking these cichlids for
aquatic weed control, which is often not understood by irrigation management
personnel (Hauser et al. 1976 , 1977;
Legner 1979b). Otherwise competitive displacement may
eliminate T. zillii, the most efficient weed eating species (Legner 1986). The numerous crater nests of these
cichlids found in irrigation drains attests to their firm establishment and
aquatic weed cleansing action <PHOTO>. Storage of water in open
containers has frequently been the cause for outbreaks of human disease
transmitted by Aedes aegypti (Linnaeus) (Culicidae) in less
developed parts of the world. While conducting
Ae. aegypti surveys in Malaysia during the mid 1960s, Dr. Richard
Garcia of the University of California,
Berkeley (pers. commun.)
observed P. reticulata* being utilized by town residents for the control of
mosquitoes in bath and drinking water storage containers. The origin of this control technique was
not clear but it appeared to be a custom brought to the area by Chinese
immigrants. Not all residents used
fish, but those that did had no breeding of Ae. aegypti in their
vicinity. Neng (1986) reported that
catfish, Claris* sp., controlled Ae. aegypti in water storage tanks in coastal villages of southern
China. This indigenous, edible fish
consumed large numbers of mosquito larvae, had a tolerance for a wide range
of environmental extremes, and could be acquired in the local markets. One fish was placed in each water source
with survey teams monitoring for its presence about every 10-15 days. If fish were not found on inspection the
occupant was persuaded to replace the fish.
The study was conducted from 1981 to 1985, during which
mosquito-breeding surveys showed a great initial reduction in Ae. aegypti followed by a sustained control of mosquitoes over the
four-year study period. Outbreaks of
dengue were observed in neighboring provinces during this period, but not in
the fishing villages under observation.
The cost of the program was estimated to be about 1/15th that of
indoor house spraying (Neng 1986). Alio et
al. (1985) described the use of a local
species of fish for the control of a malaria vector similar to that reported
by Kligler (1930). Oreochromis sp., a tilapine, was
introduced into human-made water catchment basins called "barkits"
in the semi arid region of northern Somalia.
These small-scattered impoundments were the only sources of water
during the dry season for the large pastoral human population. Anopheles
arabiensis Patton, a local vector
of malaria, was essentially restricted to these sites, and introduction of
fish into the "barkits" dramatically reduced both the vector and
nonvector populations of mosquitoes.
Treatment of the human population with antimalarial drugs during the
initial phase of this two-year study, combined with the lower vector
population reduced the transmission rate of malaria to insignificance over a
21 month period whereas the control villages remained above 10 percent. Alio et al. (1985)
suggested that the added benefits of reduced vegetation and insects in the
water sources was also recognized by the local population, resulting in
community cooperation. This was
expected to further benefit the control strategy by providing assistance in
fish distribution and maintenance as the program expanded to other
areas. The last two examples involve
the use of indigenous rather than imported fish in vector control
programs. There are other examples
where native fishes have been used in specialized circumstances (Kligler
1930, Legner et al. 1974,
Menon and Rajagopalan 1977, 1978, Walters and Legner 1980, Ataur-Rahim 1981 and
Luh 1980, 1981). Lloyd (1987)
reasoned that only indigenous fish should be employed for mosquito control
because of the environmental disruption affected by imports such as G. affinis*. However, he urged careful examinations for
prey selectivity, reproductive potential and competence in suppression of
mosquitoes before attempting their use.
Lloyd (1987) also encouraged a multidisciplinary approach involving
entomologists and fisheries biologists when utilizing indigenous fish for
mosquito control. Paradoxically, in
California where native pup fishes in the genus Cyprinodon* may afford a greater potential for mosquito control
under a wider range of environmental extremes than Gambusia* spp. (Walters and Legner 1980), the California
Department of Fish and Game discourages their use on the basis that unknown
harmful effects might occur to other indigenous fishes, and that certain rare
species of Cyprinodon <PHOTO>
might be lost through hybridization. An effective tactic was used
in China where native fish serve both for mosquito control and as a protein source (Petr 1987, Garcia and
Legner 1999). However, this approach for mosquito
control is not novel, as Kligler (1930) used a tilapine fish to control Anopheles spp. in citrus irrigation
systems in Palestine, where farmers cared for the fish, consuming the larger
ones. According to Luh (1980, 1981),
rearing of edible fish for the purpose of mosquito control and human food has
been widely encouraged in China. The
common carp, Cyprinus carpio Linnaeus (Cypriniformes:
Cyprinidae), and the grass carp, Ctenopharygodon
idella* Valenciennes
(Cypriniformes: Cyprinidae), are generally used. Fish are liberated as fry when rice seedlings are planted. Fields are specially prepared with a
central "fish pit" and radiating ditches for refuge when water
levels are low. Pisciculture in rice fields
give benefits of a significant reduction in culicine larvae, a lesser extent
anopheline larvae, the fish are harvested as food, and rice yields are
increased probably by a reduction of aquatic weeds and by fertilization of
the plants through fish excreta (Luh 1981). Annual or "instant"
fishes, (Cyprinodontidae), native to South America and Africa, have been
considered as possible biological control agents for mosquitoes (Vanderplank
1941, 1967; Hildemann and Wolford 1963, Bay 1976, 1972; Markofsky and
Matias 1979). The desiccation
resistant eggs of these cyprinodontids enable them to persist in temporary
water habitats. They may also impact
mosquito populations in native areas (Vanderplank 1941, Hildemann and Wolford
1963, Markofsky and Matias 1979). In
California the South American Cynolebias
nigripinnis* Regan (Cyprinoformes:
Cyprinodontidae) and Cynolebias bellottii * (Steindachner)
(Cyprinoformes: Cyprinodontidae), survived one summer in rice fields, but no
reproduction was observed over a three-year period (Coykendall 1980). It was speculated that further research
may enable their establishment in temporary pools and possibly rice fields. Cynolebias
bellottii <PHOTO>,
reproduced repeatedly and persisted
in small intermittently dried ponds in Riverside, California for 11
consecutive years, 1968-1979 (Legner and Walters unpubl.). Four drying/flooding operations over two
months were required to eliminate this species from ponds that were being
used for native fish studies (Walters and Legner 1980). Because they survive an annual dry period,
these fish might be successfully integrated into mosquito control programs,
especially in newly created sources in geographic areas where they naturally
occur (Vaz-Ferreira et al. 1963,
Anon 1981, and Geberich and Laird 1985). The practical use of fish species
other than Gambusia* spp. in
mosquito control often has been restricted by inadequate supplies, as the
cost of tropical and semitropical species obtainable from commercial sources
has been prohibitive for stocking large mosquito habitats. Low water temperatures during spring
months are unfavorable for tropical species and frequently predispose them to
fungal pathogens or predation by cold water fish species (Legner 1979b,
1983). Predacious Arthropods.--Numerous species of predatory arthropods
have been observed preying on mosquitoes, and in some cases are considered
important in control (James 1967, Service 1977, Collins and Washino 1985,
McDonald and Buchanan 1981). However,
among the several hundred predatory species observed, only a few have been
deployed to control mosquitoes.
Dragonflies, or "mosquito hawks", were one of the first
arthropods to be examined; but difficulties in colonization, production and
handling have limited their use to only a few areas (Urabe et al. 1986, Sebastian et al. 1990). Thus, they probably never will be used
extensively other than in a conservation sense. Aquatic Coleoptera have been
extensively studied in the field, with research facilitated by their habits
of consuming solid prey. Although their
value in effective mosquito predation has been minimized (Kühlhorn 1961),
techniques in serology and radioactive labeling have established the
importance of several species in mosquito predation (Baldwin et al. 1955, Bay 1974). The Dytiscidae appeared valuable to a
number of workers, with common dytiscid genera including Dytiscus, Laccophilus, Agabus, and Rhantus. Laccophilus terminalus* Sharp
(Coleoptera: Dytiscidae) was
extensively studied (Borland 1971), but Washino (1969) and Kühlhorn (1961) found
this predator to be of limited value in California and Germany, respectively. Sometimes difficulties
associated with the manipulative use of arthropods may be partially
overcome. For example, the mosquito
genus Toxorhynchites, whose larvae
are predators of other mosquitoes, was liberated on several Pacific Islands
in an effort to control natural and artificial container breeding mosquitoes
such as Ae. aegypti and Aedes albopictus (Skuse) (Culicidae) (Paine
1934, Bonnet and Hu 1951, Peterson 1956).
The introductions were not considered successful, even though
predatory mosquitoes did establish in some areas (Steffan 1975). Follow-up studies showed low egg
production, lack of synchrony between predator and prey life cycles, and
selection of only a relatively small number of prey breeding sites (Muspratt
1951, Nakagawa 1963, Trpiš 1973, Bay 1974, Rivière and Pichon 1978, Rivière
1985). There is still considerable
interest in the use of various Toxorhynchites
spp. for inundative liberations (Gerberg and Visser 1978, Chadee et al. 1987, Lane 1992). Trpiš (1981) studying Toxorhynchites brevipalpis*
(Theobald) showed a high daily
consumption rate and long survival of larvae without prey, making this
species a prime candidate for biological control. Observations on adult females showed a 50% survivorship over a
10-week period with a relatively high oviposition rate per female. The above attributes suggest that this
species would be useful for inundative liberations against container breeding
mosquitoes. Studies by Focks et al. (1979, 1980, 1982, 1983) with Toxorhynchites rutilis rutilis*
Coquillett in Florida, showed that this species had a high success rate in
artificial breeding containers. In a
12.6 ha. residential area, about 70% of the available oviposition sites were
located over a 14-day period by two inoculations of 175 females. Mass culturing techniques have been
developed for this species and Toxorhynchites
amboinensis* (Doleschall) (Focks
and Boston 1979, Rivière et al.
1987b). Focks et al. (1986) reported that inoculations of 100 T. amboinensis*
females per block for several weeks, combined with ultra low volume
application of malathion, reduced Ae.
aegypti populations by about 96% in
a residential area of New Orleans.
The T. amboinensis* and not the insecticide treatment apparently
accounted for most of the reduction.
Reducing both the number of predators and malathion applications
without lowering efficacy could further refine the procedure. Mosquitoes such as Ae. aegypti and Ae. albopictus, which breed in and whose eggs are dispersed by means
of artificial containers, pose major health hazards as vectors of human
pathogens throughout the warmer latitudes.
Containerized products and rubber tires, which are discarded or
stockpiled, give these mosquito species a considerable ecological
advantage. The incapacity of
governments to control disposal of these containers and difficulties in
location once they are discarded makes inundative liberations of Toxorhynchites spp, either alone or in
combination with other controls, a logical approach (Focks et al. 1986, Rivière et al. 1987b). Other mosquito genera that
are predatory on mosquitoes breeding in temporary restricted habitats, such
as containers include species of Megarhinus,
Anopheles, Lutzia, Armigeres, Eretmapodites and Psorophora. Other Diptera
that are predacious on mosquito larvae include Chaoboridae, Dolichopodidae
and Empidae. However, manipulation of
species in these genera and families has not been attempted directly, although
their importance in natural predation of pestiferous mosquitoes is
recognized. Among the Hemiptera, the
Notonectidae are voracious predators of mosquito larvae under experimental
conditions and in waterfowl refuges in California's Central Valley (Garcia
and Legner 1999, Legner and Sjogren, unpub.
data). Notonecta undulata*
Say (Hemiptera: Notonectidae) and Notonecta unifasciata* Guerin (Hemiptera: Notonectidae) have been colonized
in the laboratory. In addition,
collection of large numbers of eggs, nymphs and adults is feasible from such
breeding sites as sewage oxidation ponds (Garcia and Legner 1999and
Sjogren and Legner 1974). Studies on storage of eggs at low temperatures show a rapid
decrease of viability with time (Sjogren and Legner 1989). The most workable use of these predators
appears to be the recovery of eggs from wild populations on artificial
oviposition materials and their redistribution to mosquito breeding
sites. Such investigations were
carried out in central California rice fields by Miura (1986). Floating vegetation such as algal mats and
duck weed (Lemna spp.) form protective
refugia for mosquito larvae, and consequently populations of mosquitoes can
be high in the presence of notonectids (Garcia et al. 1974). High costs
of colonization and mass production, coupled with the logistics of
distribution, handling and timing of release at the appropriate breeding
site, thwart the use of notonectids in mosquito control. Other hemipterous genera that
have been given some attention as useful mosquito predators are Belostoma, Abedus (Washino 1969) and species of Corixidae (Sailer and Lienk
1954). Immature dragonflies also are
predatory on mosquitoes, but they do not possess the searching ability
demonstrated by certain Hemiptera and Coleoptera. Spiders (Araneae) also have been shown to be effective
predators of adult mosquitoes (Dabrowska-Prot et al. 1968, Garcia and Schlinger 1972, Service 1973). Parasitic aquatic mites
frequently occur on mosquitoes but their biological control importance has
not been evaluated (Mullen 1975). Predacious Crustaceans.--In addition to insect predators,
several crustaceans feed on mosquito larvae, among which are the tadpole
shrimp, Triops longicaudatus (LeConte) (Notostraca: Triopsidae)., and several
copepods. Scott and Grigarick (1979)
and Mulla et al. (1986),
investigating the tadpole shrimp, showed that it was an effective predator
under laboratory conditions and considered that it may play an important role
in the field against flood water Aedes
spp. and Psorophora spp in
southern California. Drought
resistance in predator eggs is an appealing attribute for egg production,
storage and manipulation in field situations against these mosquitoes (Fry
and Mulla 1992). However, synchrony
in hatch and development between the predator and the prey is crucial if this
is to be a successful biological control agent for the rapidly developing Aedes spp. and Psorophora spp. Tadpole
shrimp are considered important pests in commercial rice fields. Miura and Takahashi (1985)
reported that Cyclops vernalis* Fisher (Copepoda) was an
effective predator on early instar Cx.
tarsalis larvae in the
laboratory. It was speculated that
copepods could have an important role in suppressing mosquito populations in
rice fields because of their feeding behavior and abundance. Another crustacean that may be
suited for more extensive application is the cyclopoid predator, Mesocyclops aspericornis* Daday (Copepoda)(Rivière et al. 1987a,b). Studies
have shown >90% reductions of Ae.
aegypti and Aedes polynesiensis*
Marks (Culicidae) after inoculation into artificial containers, wells,
treeholes and land crab burrows.
Although not able to survive desiccation, the small cyclopod predator
has persisted almost 2.5 years in crabholes and up to five years in wells,
tires and treeholes under subtropical conditions. It can be mass-produced, but its occurrence in large numbers in
local water sources allows for the inexpensive and widespread application to
mosquito breeding sites in Polynesia (Rivière et al. 1987a,b). The
species is also very tolerant of salinities greater than 50 parts per
thousand. The benthic feeding
behavior of M. aspericornis* makes it an effective predator of the benthos
foraging Aedes spp., but limits
effectiveness against surface foraging mosquitoes. Rivière
et al. (1987a,b)
reported that the effectiveness against Aedes
spp. was due to a combination of predation and competition for food. Perhaps the greatest value of this Mesocyclops is in the control of
crabhole breeding species, such as Ae.
polynesiensis* in the South
Pacific. Other Invertebrate Predators.--The
most important nonarthropod invertebrates to receive attention for mosquito
control are the turbellarian flatworms and a coelenterate. Several flatworm species have been shown
to be excellent predators of mosquito larvae in a variety of aquatic habitats
(Yu and Legner 1976a,b; Collins and Washino 1978,
Case and Washino 1979, Legner 1979a, Meyer and Learned 1981,
Ali and Mulla 1983, George 1983, George et
al. 1983, Perich et al. 1990,
and Legner 1991 ). Several biological and ecological
attributes of flatworms make them ideal candidates for manipulative use. Among them are ease of mass production, an
overwintering embryo, effective predatory behavior in shallow waters with
emergent vegetation, on site exponential reproduction following inoculation
(Legner and Tsai 1977 ,1978,
Legner 1977,
1979a;
Darby et al. 1988) and tolerance to
environmental contaminants (Levy and Miller 1978, Nelson 1979). Collins and Washino (1978)
and Case and Washino (1979) suggested that flatworms, particularly Mesostoma spp.* (Microturbellaria),
may play an important role in the natural regulation of mosquitoes in some
California rice fields because of their densities and their predatory attack
on mosquito larvae in sentinel cages.
An analysis using extensive sampling showed a significant negative
correlations between the presence of flatworms and population levels of Cx. tarsalis and An. freeborni* (Case and Washino
1979). However, these workers
cautioned that an alternative hypothesis related to the ecology of these
species may have accounted for the correlations. Subsequent investigations by Palchick and Washino (1984),
employing more restrictive sampling, were not able to confirm the
correlations between Mesostoma spp.*
and mosquito populations. However,
problems associated with sampling in California rice fields, coupled with the
complexity of the prey and predator interactions (Palchik and Washino 1986),
indicate that further studies are necessary before the role of this group of
flatworms in rice fields can be clearly established. Considering all the
attributes for manipulative use of flatworms, it is surprising that they have
not been developed further for use in mosquito control. Undoubtedly the contemporary development
of Bacillus thuringiensis var. israelensis DeBarjac (H-14), a highly
selective easily applied and "marketable" microbial insecticide,
has been partially responsible for slowing further work and development of
these predators. Their mass culture
must be continuous and demands skilled technical assistants (Legner and Tsai 1978). Their persistence in field habitats may
also depend on the presence of other organisms, such as ostracods, which can
be utilized for food during low mosquito abundance (Legner et al. 1976 ). The coelenterates, like the
flatworms, showed great promise for further development and use in selected
breeding habitats (Qureshi and Bay 1969).
Chlorohydra viridissima (Pallas) (Hydrazoa) is
efficient in suppressing culicine larvae in ponds with dense vegetation and
this species also can be mass-produced (Lenhoff and Brown 1970, Yu et al. 1974). However, like the flatworms, work on these
predators has declined, probably for similar reasons as speculated for the
flatworms. Microbial pesticides can
be employed over an extensive range of different mosquito breeding
habitats. Yet the relative seasonal
permanence of control achieved with the flatworms and hydra should restore
their importance as resistance to and costs of microbial pesticides
accelerates. Pathogenic Fungi.--Species of fungi such as Beauveria bassiana (Bolsano), Metarrhizium
anisopliae (Metsch.), Entomophthora spp., Coelomomyces spp. and Lagenidium spp. have been used to
control mosquitoes (Garcia and Legner 1999); but the most promising fungal
pathogen is a highly selective and environmentally safe oomycete, Lagenidium giganteum* Couch (Oomycetes: Lagenidiales) which it is applied by aircraft to rice
fields (Kerwin and Washino 1987). Lagenidium giganteum* develops
asexually and sexually in mosquito larvae, and recycles in standing bodies of
water. This creates the potential for
prolonged infection in overlapping generations of mosquitoes. Lagenidium
giganteum* may also remain
dormant after the water source has dried up and then become active again when
water returns. The sexually produced
oospore offers the most promising stage for commercial production because of
its resistance to desiccation and long-term stability. Nevertheless, problems with production and
activation of the oospores remain (Garcia and Legner 1999). Field trials with the sexual oospore and
the asexual zoospore indicate that this mosquito pathogen is near the goal of
practical utilization. Kerwin et
al. (1986) reported that the asynchronous
germination of the oospore is of particular advantage in breeding sources
where larval populations of mosquitoes are relatively low, but recruitment of
mosquitoes is continuous due to successive and overlapping generations, as in
California rice fields. The
germination of oospores over several months provides long-term control for
these continuous low level populations.
In addition, the asexual zoospores arising from the oospore infected
mosquito is available every two to three days to respond in a density dependent
manner to suppress any resurging mosquito population. This stage survives about 48 hours after
emerging from the infected host. Kerwin et
al. (1986) indicated that laboratory
fermentation production of the asexual stage of Lagenidium for controlling mosquitoes in the field may approach
the development requirements and costs for the production of Bacillus thuringiensis israelensis. A distinct advantage of this pathogen over
the Bacillus is its ability to
recycle through successive host generations.
There are disadvantages in that the asexual stage is relatively
fragile, cannot be dried and has a maximum storage life of only eight weeks,
thus, the focus of attention for commercial production is on the oospore,
which is resistant to desiccation and can be easily stored. Axtell and Guzman (1987) succeeded to
encapsulate both the sexual and asexual stages in calcium alginate and
reported activity against mosquito larvae after storage for up to 35 and 75
days, respectively. Limitations on
the use of this pathogen include intolerance to polluted water, salinity and
other environmental factors (Garcia and Legner 1999). However, there are numerous
mosquito-breeding sources where these limitations do not exist and, therefore,
this selective and persistent pathogen may become available for routine
mosquito control. The fungus Culicinomyces clavosporus Couch, Romney and Rao, first isolated from laboratory
mosquito colonies and later from field habitats, has been studied for
biological control (Sweeney 1987).
The fungus is active against a wide range of mosquito species and also
causes infections in other aquatic Diptera.
The relatively inexpensive media in fermentation tanks facilitates
production. However, problems in
storage must be overcome if this fungus is to be widely used (Sweeney
1987). Although the fungus has shown
high infection rates in field trials with high dosage rates, appreciable
persistence at the site has not been demonstrated (Sweeney 1987). Various species of Coelomomyces have been studied for use
in mosquito control, with epizootic infection rates in excess of 90% being
recorded. Although these fungi
persist in certain habitats for long periods, the factors responsible for
triggering outbreaks are not well understood (Chapman 1974). Field-testing that has been done shows
great variability (Federici 1981).
Difficulties associated with the complex life cycle of these fungi
have encumbered research. Federici
(1981) and Lacey and Undeen (1986) reviewed the potential of these fungi for
mosquito control. Nevertheless,
infections of up to 100% have been reported on some populations of Anopheles gambiae* Giles (Culicidae) in Zambia (Muspratt 1963), but lower
rates of 24-48% were reported in Anopheles
quadrimaculatus Say (Culicidae) and Ae.
crucians* Wiedemann (Culicidae) in
the southeastern U.S. (Umphlett 1970, Chapman et al. 1972). Higher
infections exceeding 95% were reported from Culiseta inornata
(Williston) and Psorophora howardii Coquillett by Coelomomyces psorophorae* Couch and in Aedes
triteriatus* (Say) (Culicidae) by Coelomomyces
macleayae Laird and 85% in Culex
peccator* Dyar et Knab (Culicidae)
by Coelomomyces pentangulatus* Couch (Bay et al. 1976). Although Coelomomyces species have been difficult to mass produce, new
introductions of these fungi were made by Laird (1967) on a tiny Pacific
Island against Ae. polynesiensis* Mark, a vector of
filariasis. This represents one of
the few attempts to establish new mosquito pathogens in an area where they
did not exist. Further application of
Coelomomyces spp. as a direct
mosquito control is dependent on the development of easily cultured
inoculum. Reports of research with B. bassiana
on Culex tarsalis and Aedes
nigromaculis* (Ludlow) (Culicidae) (Legner et al. 1974) substantiates that of Clark et al. (1968): Aedes
nigromaculis* was more susceptible
than Cx.. tarsalis with the third host
passage resulting in 100% infection under laboratory conditions. Parasitic Nematodes.--Among the various mermithid and
rhabditoid nematodes pathogenic for mosquitoes, Romanomermis culicivorax*
Ross and Smith (Mermithidae: Nematoda), has received the most attention
(Poinar 1979, Platzer 1990, Kaya and Gaugler 1993). This mermithid is active against a wide range of mosquito
species, and has been mass-produced and deployed in a number of field trials. The nematode was commercially produced and
sold as Skeeter Doom TMR, but the eggs showed reduced viability in
transport and the product currently is no longer sold (Service 1983). However, the nematode's ability to recycle
through multigenerations of mosquitoes and overwinter in various habitats,
including drained, harvested, stubble-burned, cultivated and replanted rice
fields, favors further research and development for biological control
(Petersen and Willis 1975, Brown-Westerdahl et al. 1982). Several
field applications showing good results have included both the preparasitic
stage and post parasitic stages with the former more applicable to a
"rapid kill" and the latter for more long-term continuous control
such as in rice fields (Levy et al.
1979, Brown-Westerdahl et al.
1982). Obstacles to its widespread
use include intolerance to low levels of salinity, polluted water and low
oxygen levels, predation by aquatic organisms and the potential for
development of resistance by the host (Brown-Westerdahl 1982). Although such environmental problems are not
as important for anopheline control, the cost of in vivo mass production
is a disadvantage for use of this pathogen.
However, it may be adapted for use in specialized habitats integrated
with other controls (Brown-Westerdahl et
al. 1982). Neoaplectana carpocapsae*
Weiser (Mermithidae: Nematoda) and
other nematodes have shown a high level of infection in nature (Platzer
1990). Pathogenic Bacteria.--Bacteria are not commonly associated
with mosquitoes in nature, but one spore forming bicrystalliferous strain of Bacillus thuringiensis var. israelensis
(H-14), was isolated by Goldberg and Margalit (1977) and the toxin it
produces has been shown by numerous studies to be an effective and
environmentally sound microbial insecticide against mosquitoes and blackflies. A high degree of specificity and toxicity,
coupled with the relative ease of production, have made it the most widely
used microbial product to date for mosquito and blackfly control. Several formulations have been available
commercially throughout the world.
Nevertheless, its efficacy varies under different environmental
conditions and there are some problems associated with its use (Garcia 1987,
Lacey and Undeen 1986, Garcia and Legner 1999). The bacterium as applied commercially
cannot multiply in the environment, thus it acts essentially as a synthesized
insecticide. Evolution of the
bacterium to counteract developing resistance in the host is thus precluded,
and there are limitations on the development of new strains in the laboratory
(Smits 1987). Another spore forming
bacterium, Bacillus sphaericus* Neide, is larvicidal
against certain mosquito species (Mulla 1986, Mulla et al. 1991, Singer 1990, Weiser 1984). Several strains of this pathogen show a high degree of toxic
variability among species of mosquitoes.
Culex spp. appear to be
highly susceptible, whereas other species such as Ae. aegypti respond
poorly to treatment. Unlike the
transitory larvicidal activity of Bt.
toxin (Cry IV), some strains of B. sphaericus persist and apparently
recycle in certain aquatic habitats (DesRochers and Garcia 1984, Lacey 1990,
Yap 1990, Yousten et al.
1992). Although evolution to
counteract resistance in the insect is thus possible, real resistance has developed
nonetheless (Rodcharoen and Mulla 1993). Parasitic Protozoa.--Many species of protozoa have been
isolated from mosquitoes and other medically important Diptera (Roberts et al. 1983, Lacey and Undeen
1986). These include flagellates (Blastocrithidia spp. and Crithidia spp.), eugregarines (Lankesteria spp.), ciliates (Vorticella spp. and Tetrahymena spp.), and
schizogregarines (Caulleryella spp.)
and microsporidians. Due to their
complex life cycle and the in vivo production methods necessary for
maintaining them, research on their practical utility has been limited. However, if more information is developed
on their life cycle, it may be found that they could play a role in
suppressing mosquitoes through inoculative and augmentive releases in certain
habitats (Lacey and Undeen 1986).
Infection of mosquitoes by most Microsporida
is transovum and field transmission has yet to be shown. Only a few species including Nosema, Stegomyiae and Stempellia
sp. possess the ability to infect their hosts per os (Chapman 1974). Among other promising
protozoa is the endoparasitic ciliate, Lambornella
clarki Corliss and Coats
(Ciliophora: Tetrahymenidae), a natural pathogen of the treehole mosquito, Aedes sierrensis* Ludlow (Culicidae), which has received considerable
attention as a potential biological control agent for container breeding
mosquitoes (Egerter and Anderson 1985, Egerter et al. 1986, Washburn and Anderson 1990). Cysts resistant to desiccation allow
persistence of the ciliate from one year to the next. In
vitro production methods have been
sought and field trials initiated to determine its efficacy for biological
control (Anderson and Washburn 1990). Viruses.--A number of pathogenic
viruses have been isolated from mosquitoes and blackflies (Granados and
Federici 1986). A natural population
of Aedes sollicitans* Walker (Culicidae) in Louisiana sustained an
epizootic by a cytoplasmic and a nuclear polyhedrosis virus where more than
71% infection occurred (Clark and Fukuda 1971). Bay et al. (1974) also reported that
H. C. Chapman observed a similar epizootic infecting over 65% of the larvae
of Ae. sollicitans*, but reflooding after drying of these habitats
greatly reduced infection. Mosquito
iridescent viruses have been reported from various mosquito species in Europe
and the United States (Clark et al.
1965, Weiser 1965), but natural infection levels rarely exceed 1%. Therefore, viruses do not appear
practical for use in control (Lacey and Undeen 1986). Larvicidal Plants.--Certain plants and plant products are
lethal to developing mosquitoes (Azmi et
al.1998, Joshi et al 1998, Su
and Mulla 1998a, 1998b, Sukumar et al.
1991). However, practical deployment
has not been demonstrated, and in the absence of insect population
interaction with the substance, insect resistance should rapidly
develop. Nevertheless, the
possibility of some plant extracts such as Neem, Azadirachta indica A. Juss, being innocuous to nontarget
organisms (e.g., mosquito predators) makes such substances highly desirable
for integrated control (Su and Mulla 1998, 1998b). Particularly interesting
is the activity of ethanol extracts of fresh Neem showing antimalarial
activity against chloroquine resistant Plasmodium
falciparum strain K1. (Joshi
et al. 1998). Chironomidae Chironomid midges pose
nuisances in metropolitan areas such as southwestern California wherever
there is a great proximity of urban development to paved flood control river
channels, sewage oxidation ponds and recreational lakes. Infestations in paved river channels
characteristically become especially severe following winters with above
average rainfall. Rapid
recolonization of the scoured habitat occurs due to fertile urban runoff
water which stimulates algal growth. Fish have been used for
chironomid midge abatement in lentic habitats as an adjunct to chemical
pesticides. Such species as the
common carp, Cyprinus carpio L. and goldfish, Carassius auratus (L.) and pupfish, Cyprinodon
macularius Baird and Girard, have
been effective in shallow California ponds (Anderson and Ingram 1960, Bay and
Anderson 1965, Legner et al. 1975,
Walters and Legner 1980, Legner and Warkentin 1990). However, other cichlid species in the
genera Tilapia and Oreochromis are useful for the lotic
situation in the paved storm drain habitats (Legner 1983). The addition of three species of tilapine
fishes to drainages in the Los Angeles area in the 1970's resulted in
widespread establishment of an apparent hybrid of Oreochromis mossambica* (Peters) (Cichlidae) and
Oreochromis hornorum* Trewazas
(Legner 1983). Densities of Chironomidae, principally Chironomus attenuatus* Johannsen larvae, declined significantly in the
drainages and resulted in complete adult midge control. The foraging on Chironomidae in certain detritus
substrates by very dense populations of the fish influenced the ability of
such substrates to produce chironomids.
The chironomid-sustained fish biomass in autumn may exceed 4 X 105
kg.. over a distance of 18 km. of one studied paved river channel. By 1990 the tilapine fish were regularly
ranging in the neritic zone along the southwestern California coast, and
their contribution to predatory marine fish biomass was considered
significant (Legner and Pelsue 1980, Legner et al. 1980). The Planaria
and Hydra
noted previously in mosquito control also significantly reduced chironomid
population densities in experiments (Yu and Legner 1976a, Garcia and Legner 1999). However, they were never deployed
specifically for chironomid control.
Hilsenoff (1964), Hilsenoff and Lovett (1966) reported on leeches and
a microsporidian as significant natural enemies of chironomids. Tabanidae
Tabanidae, or horseflies and
deerflies, although widespread and on occasion serious pests and vectors of
disease to livestock, have not received much attention. Only one successful inundative release of
the egg parasitoid, Phanurus emersoni* Girault (Hymenoptera:
Scelionidae), has been recorded (Parman 1928). Apparently, this effort was precipitated by a severe outbreak
of anthrax at the time and since this disease diminished and other control
tactics are available, interest in their biological control has not been
continued. Other references to
natural enemies of tabanids include James (1963) and Magnarelli and Anderson
(1980). Simuliidae
The genera Simulium and Eusimulium are of special importance because adults emerge in
great numbers to inflict vicious bites on humans. Moreover, some species are vectors of onchocerciasis. Attempts were made in 1931 to establish
certain dragonflies and a predacious chironomid, Cardiocladius sp., in New Zealand on Simulium sp., but results were not positive (Clausen et al. 1978). This group apparently does not lend itself
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