Biological pest control

Biological control of pests in agriculture is a method of controlling pests (including insects, mites, weeds and plant diseases) that relies on predation, parasitism, herbivory, or other natural mechanisms. It can be an important component of integrated pest management (IPM) programs.

Overview

Biological Control is defined as the reduction of pest populations by natural enemies and typically involves an active human role. Natural enemies of insect pests, also known as biological control agents, include predators, parasitoids, and pathogens. Biological control agents of plant diseases are most often referred to as antagonists. Biological control agents of weeds include herbivores and plant pathogens. Predators, such as lady beetles and lacewings, are mainly free-living species that consume a large number of prey during their lifetime. Parasitoids are species whose immature stage develops on or within a single insect host, ultimately killing the host. Most have a very narrow host range. Many species of wasps and some flies are parasitoids. Pathogens are disease-causing organisms including bacteria, fungi, and viruses. They kill or debilitate their host and are relatively specific to certain insect groups. There are three basic types of biological control strategies; conservation, classical biological control, and augmentation. These are discussed in more detail below.

Conservation

The conservation of natural enemies is probably the most important and readily available biological control practice available to homeowners and gardeners. Natural enemies occur in all areas, from the backyard garden to the commercial field. They are adapted to the local environment and to the target pest, and their conservation is generally simple and cost-effective. Lacewings, lady beetles, hover fly larvae, and parasitized aphid mummies are almost always present in aphid colonies. Fungus-infected adult flies are often common following periods of high humidity. These naturally occurring biological controls are often susceptible to the same pesticides used to target their hosts. Preventing the accidental eradication of natural enemies is termed simple conservation."'

Effects of biological control on biodiversity

Effects on native biodiversity

Biological control can potentially have positive and negative effects on biodiversity. Most of the time a biological control is introduced to an area to protect a native species from an invasive or exotic species that has moved into its area. The control is introduced to lessen the competition among native and invasive species. However, the introduced control does not always target only the intended species. It can also target native species.Fact|date=December 2007

When introducing a biological control to a new area, a primary concern is the host- or prey-specificity of the control agent. Generalist feeders (control agents that are not restricted to a single species or a small range of species) often make poor biological control agents, and may become invasive species themselves. For this reason, potential biological control agents should be subject to extensive testing and quarantine before release into any new environment. If a species is introduced and attacks a native species, the biodiversity in that area can decrease dramatically. When one native species is removed from an area, it may have filled an essential niche, When this niche is absent it will directly affect the entire ecosystem. Fact|date=December 2007 Because they tend to be generalist feeders, vertebrate animals seldom make good biological control agents, and many of the classic cases of "biocontrol gone awry" involve vertebrates. For example, the cane toad ("Bufo marinus") was introduced as a biological control and had significant negative impact on biodiversity. The cane toad was intentionally introduced to Australia to control the cane beetle. When introduced, the cane toad thrived very well and did not only feed on cane beetles but other insects as well. The cane toad soon spread very rapidly, thus taking over native habitat. The introduction of the cane toad also brought foreign disease to native reptiles. This drastically reduced the population of native toads and frogs. “The cane toad also exudes and can squirt poison from the parotid glands on their shoulders when threatened or handled. This toxin contains a cocktail of chemicals that can kill animals that eat it. Freshwater crocodiles, goannas, tiger snakes, dingos and northern quolls have all died after eating cane toads, as have pet dogs (Cane toad,2003) ”. This goes to show a small but deadly organism can alter the native biodiversity in an ecosystem in a very expedient manner. A pyramid effect can take place if native species are reduced or eradicated. The domino effect keeps on going and can potentially exude on other bordering ecosystems until an equilibrium is reached.

A second example of a biological control agent that subsequently crossed over to native species is the "Rhinocyllus conicus". The seed feeding weevil was introduced to North America to control exotic thistles (Musk and Canadian). However, the weevil did not target only the exotic thistles, it also targeted native thistles that are essential to various native insects. The native insects rely solely on native thistles and do not adapt to other plant species. Therefore, they cannot survive. Biological controls do not always have negative impacts on biodiversity (Corry 2000).Successful biological control reduces the density of the target species over several years, thus providing the potential for native species to re-establish. In addition, regeneration and reestablishment programs can aid to the recovery of native species. Native species can be affected in a positive way as well. To develop or find a biological control that exerts control only on the targeted species is a very lengthy process of research and experiments. In the late 1800’s, the citrus industry was in great fear when the cottony cushion scale was discovered. This organism could cause a great deal of economic loss to the industry. However, a biological control was introduced. The vedalia beetle and a parasitoid fly were introduced to control the pest. Within a few years time, the cottony cushion scale was controlled by the natural enemies and the citrus industry suffered little financial loss. Many exotic or invasive species can suppress the development of native species. The introduction of an effective biological control that reduces the population of the invasive species allows the rejuvenation of the native species. Biological controls can reduce competition for biotic and abiotic factors which can result in the re-establishment of the once over ran native species.Fact|date=December 2007

Effects on invasive species

Invasive species are closely associated with biological controls because the environment in which they are invasive most likely does not contain their natural enemies. If invasive species are not controlled, biodiversity may be at great threat in the affected area. An example of an invasive species is the alligator weed. [cite web
url=http://www.ars-grin.gov/cgi-bin/npgs/html/taxon.pl?101502
title=Alternanthera philoxeroides information from NPGS/GRIN
publisher=www.ars-grin.gov
accessdate=2008-08-09
last=
first=] This plant was introduced to the United States from South America. This aquatic weed spreads very rapidly and causes many problems in lakes and rivers. The weed takes root in shallow water causing major problems such as navigation, irrigation, and flood control. The alligator weed flea beetle and two other biological controls were released in Florida. Because of their success, Florida banned the use of herbicides to control alligator weed three years after the controls were introduced (Cofrancesco 2007). Biological controls for invasive species also can have a negative impact on biodiversity.

The cane toad, as mentioned previously, is a great example of trying to control an invasive species. The cane toad was introduced to eradicate an invasive species. However, it became invasive, thus altering the biodiversity. The introduction of the cane toad could have potentially caused more of a disturbance in biodiversity than the targeted species did.

Effects on future

With further research and more scientific experiments, biological control could potentially play a huge role in the future of pest prevention. Biological control is being used among society today; however, it could someday reduce the use of many pesticides and herbicides. Since biological control could potentially have a large economic value, if found to be successful, research and job fields would increase continually. By increasing awareness of biological controls among more people, new successful biological controls could be discovered in the future. This could eliminate the overuse of chemicals. Biodiversity would increase because untargeted species that are exterminated with chemicals would no longer occur.Fact|date=December 2007

Economic effects

Therefore, biological control is heavily analyzed by the amount of economic gain that directly comes from biological control. Many of the known economics of biological control are related directly to agriculture practices. Since agriculture has a huge impact on biodiversity this could potentially increase the biodiversity among agricultural practices. In order for agriculture to keep up with the growing population, many inputs are increased resulting in the loss of un-harmful species. Biological control use has been very minimal in agriculture. Less than 1% of global pest control sales of $30 billion involve biological methods (Griffiths 2007:in press). Very few case studies on the cost-benefit analysis of biological control have been done however a few have taken place. A Critical evaluation of augmentative biological control has found four case studies. In one case, “the releases of a parasitoid "Gryon pennsylvanicum" Ashmead to control the true bug "Anasa tristis" DeGeer on pumpkins produced lower net benefit (in dollars) than applications of esfenvalerate (pesticide); 18% lower in one year and 120% lower in the next. In 1 year of the study, a combination of augmentative releases and use of a resistant pumpkin variety produced greater net benefit than pesticide alone, but not pesticide combined with the resistant variety (Olson et al. 1996) ”. Another case study found that “calculated that releases of "T. nubilale" were considerably less cost-effective than pesticide applications used to control ECB on feed corn and fresh-market sweet corn. Pesticide applications produced 87% and 45% more net benefit (in dollars) than augmentation for feed corn and fresh market corn, respectively. In seed corn, however, "Trichogramma" releases produced essentially equivalent net benefits to pesticide treatments. In a third cost-benefit analysis of augmentation, Lundgren et al. (2002) showed that "Trichogramma brassicae" Bezdenko releases produced considerably less net benefit (94%; measured in cabbage head production) than methomyl treatments (Andow 1997). In two other studies, “biological control releases were about two times the cost of pesticide applications; this was true for releases of a parasitoid, "Choetospila elegans" Westwood, used to control a stored product pest, "Rhyzopertha dominica" (F.) (Flinn et al.,1996) and releases of green lacewings, "Chrysoperla carnea" Stephens to control leafhoppers in grapes (Daane et al., 1996). Finally Prokrym et al. (1992) suggested that "Trichogramma" releases were about six times as expensive as pesticide treatments for "O. nubilalis" in sweet corn,” (Collier 2003). These case studies offer us some idea of how economical biological control can be. These show that biological control is less cost effective than chemical applications and in result raises a flag that more research needs to be done. With progression in research, we can use more controls at a cheaper cost and increase the amount of biodiversity in areas because of the minimal use of chemicals that cannot target a specific species of pest.

Classical biological control

Classical biological control is the introduction of natural enemies to a new locale where they did not originate or do not occur naturally. This is usually done by government authorities. In many instances the complex of natural enemies associated with an insect pest may be inadequate. This is especially evident when an insect pest is accidentally introduced into a new geographic area without its associated natural enemies. These introduced pests are referred to as exotic pests and comprise about 40% of the insect pests in the United States. Examples of introduced vegetable pests include the European corn borer ("Ostrinia nubilalis"), one of the most destructive insects in North America. To obtain the needed natural enemies, scientists turned to classical biological control. This is the practice of importing, and releasing for establishment, natural enemies to control an introduced (exotic) pest, although it is also practiced against native insect pests. The first step in the process is to determine the origin of the introduced pest and then collect appropriate natural enemies associated with the pest or closely related species. The natural enemy is then passed through a rigorous quarantine process, to ensure that no unwanted organisms (such as hyperparasitoids) are introduced, then they are mass produced, and released. Follow-up studies are conducted to determine if the natural enemy becomes successfully established at the site of release, and to assess the long-term benefit of its presence. There are many examples of successful classical biological control programs. One of the earliest successes was in controlling "Icerya purchasi", the cottony cushion scale, a pest that was devastating the California citrus industry in the late 1800s. A predatory insect "Rodolia cardinalis" (the Vedalia Beetle), and a parasitoid fly were introduced from Australia. Within a few years the cottony cushion scale was completely controlled by these introduced natural enemies.

Damage from "Hypera postica" Gyllenhal, the alfalfa weevil, a serious introduced pest of forage, was substantially reduced by the introduction of several natural enemies. About 20 years after their introduction, the population of weevils, in the alfalfa area treated for alfalfa weevil in the Northeastern United States, was reduced by 75 percent. A small wasp, "Trichogramma ostriniae", was introduced from China to help control the European corn borer making it a recent example of a long history of classical biological control efforts for this major pest. Many classical biological control programs for insect pests and weeds are under way across the United States and Canada. The population of "Levuana irridescens" (the Levuana moth), a serious coconut pest in Fiji was brought under control by a classical biological control program in the 1920s.

Classical biological control is long lasting and inexpensive. Other than the initial costs of collection, importation, and rearing, little expense is incurred. When a natural enemy is successfully established it rarely requires additional input and it continues to kill the pest with no direct help from humans and at no cost. Unfortunately, classical biological control does not always work. It is usually most effective against exotic pests and less so against native insect pests. The reasons for failure are not often known, but may include the release of too few individuals, poor adaptation of the natural enemy to environmental conditions at the release location, and lack of synchrony between the life cycle of the natural enemy and host pest.

Augmentation

This third type of biological control involves the supplemental release of natural enemies. Relatively few natural enemies may be released at a critical time of the season (inoculative release) or literally millions may be released (inundative release). Additionally, the cropping system may be modified to favor or augment the natural enemies. This latter practice is frequently referred to as habitat manipulation.

An example of inoculative release occurs in greenhouse production of several crops. Periodic releases of the parasitoid, "Encarsia formosa", are used to control greenhouse whitefly, and the predaceous mite, "Phytoseiulus persimilis", is used for control of the two-spotted spider mite.

Lady beetles, lacewings, or parasitoids such as those from the genus "Trichogramma" are frequently released in large numbers (inundative release). Recommended release rates for Trichogramma in vegetable or field crops range from 5,000 to 200,000 per acre per week depending on level of pest infestation. Similarly, entomopathogenic nematodes are released at rates of millions and even billions per acre for control of certain soil-dwelling insect pests.

Habitat or environmental manipulation is another form of augmentation. This tactic involves altering the cropping system to augment or enhance the effectiveness of a natural enemy. Many adult parasitoids and predators benefit from sources of nectar and the protection provided by refuges such as hedgerows, cover crops, and weedy borders. Also, the provisioning of natural shelters in the form of wooden caskets, boxes or (turnaround) flowerpots is a form of this. For example, the stimulation of the natural predator "Dermaptera" is done in gardens by hanging up turnaround flowerpots with straw or wood wool. Mixed plantings and the provision of flowering borders can increase the diversity of habitats and provide shelter and alternative food sources. They are easily incorporated into home gardens and even small-scale commercial plantings, but are more difficult to accommodate in large-scale crop production. There may also be some conflict with pest control for the large producer because of the difficulty of targeting the pest species and the use of refuges by the pest insects as well as natural enemies.

Examples of habitat manipulation include growing flowering plants (pollen and nectar sources) near crops to attract and maintain populations of natural enemies. For example, hover fly adults can be attracted to umbelliferous plants in bloom.

Biological control experts in California have demonstrated that planting prune trees in grape vineyards provides an improved overwintering habitat or refuge for a key grape pest parasitoid. The prune trees harbor an alternate host for the parasitoid, which could previously overwinter only at great distances from most vineyards. Caution should be used with this tactic because some plants attractive to natural enemies may also be hosts for certain plant diseases, especially plant viruses that could be vectored by insect pests to the crop. Although the tactic appears to hold much promise, only a few examples have been adequately researched and developed.

Examples of predators

Ladybugs, and in particular their larvae which are active between May and July in the northern hemisphere, are voracious predators of aphids such as greenfly and blackfly, and will also consume mites, scale insects and small caterpillars. The ladybug is a very familiar beetle with various colored markings, whilst its larvae are initially small and spidery, growing up to 17 mm long. The larvae have a tapering segmented grey/black body with orange/yellow markings and ferocious mouthparts. They can be encouraged by cultivating a patch of nettles in the garden and by leaving hollow stems and some plant debris over winter so that they can hibernate.

Hoverflies resemble slightly darker bees or wasps and they have characteristic hovering, darting flight patterns. There are over 100 species of hoverfly whose larvae principally feed upon greenfly, one larva devouring up to fifty a day, or 1000 in its lifetime. They also eat fruit tree spider mites and small caterpillars. Adults feed on nectar and pollen, which they require for egg production. Eggs are minute (1 mm), pale yellow white and laid singly near greenfly colonies. Larvae are 8-17 mm long, disguised to resemble bird droppings, they are legless and have no distinct head. Semi-transparent in a range of colours from green, white, brown and black.

Hoverflies can be encouraged by growing attractant flowers such as the poached egg plant "(Limnanthes douglasii)", marigolds or phacelia throughout the growing season.

Dragonflies are important predators of mosquitoes, both in the water, where the dragonfly naiads eat mosquito larvae, and in the air, where adult dragonflies capture and eat adult mosquitoes. Community-wide mosquito control programs that spray adult mosquitoes also kill dragonflies, thus removing an important biocontrol agent, and can actually increase mosquito populations in the long term.

Other useful garden predators include lacewings, pirate bugs, rove and ground beetles, aphid midge, centipedes, predatory mites, as well as larger fauna such as frogs, toads, lizards, hedgehogs,
slow-worms and birds. Cats and rat terriers kill field mice, rats, June bugs, and birds. Dogs chase away many types of pest animals. Dachshunds are bred specifically to fit inside tunnels underground to kill badgers.

* "Phytoseiulus persimilis" (against spider mites)
* "Amblyseius californicus" (against spider mites)
* "Amblyseius cucumeris" (against spider mites) [ [http://207.5.17.151/biobest/en/films/dvd.htm Spider mites and their natural enemies] ]
* "Typhlodromips swirskii" (against spider mites, thrips, and white flies)
* "Feltiella acarisuga" (against spider mites)
* "Stethorus punctillum" (against spider mites)
* "Macrolophus caluginosus" (against spider mites)
* "Encarsia formosa" (against white flies)
* "Eretmocerus" spp. (against white flies) [ [http://207.5.17.151/biobest/en/films/dvd.htm White flies and their natural enemies] ]

Parasitoid insects

Most insect parasitoids are wasps or flies. Parasitiods comprise a diverse range of insects that lay their egg on or in the body of an insect host, which is then used as a food for developing larvae. Parasitic wasps take much longer than predators to consume their victims, for if the larvae were to eat too fast they would run out of food before they became adults. Such parasites are very useful in the organic garden, for they are very efficient hunters, always at work searching for pest invaders. As adults they require high energy fuel as they fly from place to place, and feed upon nectar, pollen and sap, therefore planting plenty of flowering plants, particularly buckwheat, umbellifers, and composites will encourage their presence.

Four of the most important groups are:

*Ichneumonid wasps: (5-10 mm). Prey mainly on caterpillars of butterflies and moths.
*Braconid wasps: Tiny wasps (up to 5 mm) attack caterpillars and a wide range of other insects including greenfly. A common parasite of the cabbage white caterpillar- seen as clusters of sulphur yellow cocoons bursting from collapsed caterpillar skin.
*Chalcid wasps: Among the smallest of insects (<3 mm). Parasitize eggs/larvae of greenfly, whitefly, cabbage caterpillars, scale insects and Strawberry Tortrix Moth ("Acleris comariana").
*Tachinid flies: Parasitize a wide range of insects including caterpillars, adult and larval beetles, true bugs, and others.

Biological control with micro-organisms

Various microbial insect diseases occur naturally, but may also be used as biological pesticides. When naturally occurring, these outbreaks are density dependent in that they generally only occur as insect populations become denser.

Bacteria and biological control

Bacteria used for biological control infect insects via their digestive tracts, so insects with sucking mouth parts like aphids and scale insects are difficult to control with bacterial biological control. [ L.A. Swan. 1964. Beneficial Insects. 1st ed. page 249.] "Bacillus thuringiensis" is the most widely applied species of bacteria used for biological control, with at least four sub-species used to control Lepidopteran (moth, butterfly), Coleopteran (beetle) and Dipteran (true flies) insect pests.

Fungi and biological control

Fungi that cause disease in insects are known as entomopathogenic fungi, including at least fourteen species of entomophthoraceous fungi attack aphids. [I.M. Hall & P.H. Dunn, Entomophthorous Fungi Parasitic on the Spotted Alfalfa Aphid, Hilgardia, Sept 1957.] Species in the genus "Trichoderma" are used to manage some soilborne plant pathogens.

Plants to regulate insect pests

Choosing a diverse range of plants for the garden can help to regulate pests in a variety of ways, including;

*Masking the crop plants from pests, depending on the proximity of the companion or intercrop.
*Producing olfactory inhibitors, odors that confuse and deter pests.
*Acting as trap plants by providing an alluring food that entices pests away from crops.
*Serving as nursery plants, providing breeding grounds for beneficial insects.
*Providing an alternative habitat, usually in a form of a shelterbelt, hedgerow, or beetle bank where beneficial insects can live and reproduce. Nectar-rich plants that bloom for long periods are especially good, as many beneficials are nectivorous during the adult stage, but parasitic or predatory as larvae. A good example of this is the soldier beetle which is frequently found on flowers as an adult, but whose larvae eat aphids, caterpillars, grasshopper eggs, and other beetles.

Plants to regulate plants

The legume vine "Mucuna pruriens" is used in the countries of Benin and Vietnam as a biological control for problematic "Imperata cylindrica" grass.cite web
url=http://www.tropicalforages.info/key/Forages/Media/Html/Mucuna_pruriens.htm
title=Factsheet - Mucuna pruriens
publisher=www.tropicalforages.info
accessdate=2008-05-21
last=
first=] "Mucuna pruriens" is said not to be invasive outside its cultivated area.

Directly introducing biological controls

Most of the biological controls listed above depend on providing incentives in order to 'naturally' attract beneficial insects to the garden. However there are occasions when biological controls can be directly introduced. Common biocontrol agents include parasitoids, predators, pathogens or weed feeders. This is particularly appropriate in situations such as the greenhouse, a largely artificial environment, and are usually purchased by mail order.

Some biocontrol agents that can be introduced include;
*"Encarsia formosa". This is a small predatory chalcid wasp which is parasitical on whitefly, a sap-feeding insect which can cause wilting and black sooty moulds. It is most effective when dealing with low level infestations, giving protection over a long period of time. The wasp lays its eggs in young whitefly 'scales', turning them black as the parasite larvae pupates. It should be introduced as soon as possible after the first adult whitefly are seen. Should be used in conjunction with insecticidal soap.

*Red spider mite, another pest found in the greenhouse, can be controlled with the predatory mite "Phytoseilus persimilis". This is slightly larger than its prey and has an orange body. It develops from egg to adult twice as fast as the red spider mite and once established quickly overcomes infestation.

*A fairly recent development in the control of slugs is the introduction of 'Nemaslug', a microscopic nematode ("Phasmarhabditis hermaphrodita") which will seek out and parasitize slugs, reproducing inside them and killing them. The nematode is applied by watering onto moist soil, and gives protection for up to six weeks in optimum conditions, though is mainly effective with small and young slugs under the soil surface.

*A bacterial biological control which can be introduced in order to control butterfly caterpillars is "Bacillus thuringiensis". This available in sachets of dried spores which are mixed with water and sprayed onto vulnerable plants such as brassicas and fruit trees. The bacterial disease will kill the caterpillars, but leave other insects unharmed. There are strains of "Bt" that are effective against other insect larvae. "Bt israelensis" is effective against mosquito larvae and some midges.

*A viral biological control which can be introduced in order to control the overpopulation of European rabbit in Australia is the rabbit haemorrhagic disease virus that causes the rabbit haemorrhagic disease.

*A biological control being developed for use in the treatment of plant disease is the fungus "Trichoderma viride". This has been used against Dutch Elm disease, and to treat the spread of fungal and bacterial growth on tree wounds. It may also have potential as a means of combating silver leaf disease.

*The parasitoid "Gonatocerus ashmeadi" (Hymenoptera: Mymaridae) has been introduced to control the glassy-winged sharpshooter "Homalodisca vitripennis" (Hemipterae: Cicadellidae) in French Polynesia and has successfully controlled ~95% of the pest density [cite journal
author=Hoddle M.S., Grandgirard J., Petit J., Roderick G.K., Davies N.
date=2006
title=Glassy-winged sharpshooter Ko'ed - First round - in French Polynesia
journal=Biocontrol News and Information
volume=27
issue=3
pages=47N–62N]

Economics of biological pest control

Biological control proves to be very successful economically, and even when the method has been less successful, it still produces a benefit-to-cost ratio of 11:1. One study has estimated that a successful biocontrol program returns £32 in benefits for each £1 invested in developing and implementing the program, i.e., a 32:1 benefit-to-cost ratio. The same study had shown that an average chemical pesticide program only returned profits in the ratio of 13:1.Fact|date=February 2007

Negative results of biological pest control

In some cases, biological pest control can have unforeseen negative results that could outweigh all benefits. For example, when the mongoose was introduced to Hawaii in order to control the rat population, it preyed on the endemic birds of Hawaii, especially their eggs, more often than it ate the rats.

Cane toads ("Bufo marinus") were introduced to Australia in the 1930s in a failed attempt to control the cane beetle, a pest of sugar cane crops. 102 toads were obtained from Hawaii and bred in captivity to increase their numbers until they were released into the sugar cane fields of the tropic north in 1935. It was later discovered that the toads can't jump very high so they did not eat the cane beetles which stayed up on the upper stalks of the cane plants. The toads soon became very numerous and out-competed native species and became very harmful to the Australian environment, including being very toxic to would-be predators such as native snakes. [http://www.fdrproject.org/pages/toads.htm]

References

Biological control

*Wiedenmann, R. 2000. Introduction to Biological Control. Midwest Institute for Biological Control. Illinois. Available from http://www.inhs.uiuc.edu/research/biocontrol

Building organic pest-free gardens
* [http://www.readersdigest.co.uk/timesaving-garden-p-691.html?cPath=41_50 The Time Saving Garden] by David and Charles PLC/Reader's Digest, ISBN 13: 9780276442452

Effects on native biodiversity
*Pereira, M.J. et al. (1998) Conservation of natural vegetation in Azores Islands. Bol. Mus. Munic. Funchal 5, 299–305

*Weeden, C.R., A. M. Shelton, and M. P. Hoffman. Biological Control: A Guide to Natural Enemies in North America. Available from [http://www.nysaes.cornell.edu/ent/biocontrol/predators/rodolia_cardinalis.html] (accessed December 2007)

*Cane toad: a case study. 2003. Available from [http://www.biotechnologyonline.gov.au/enviro/canetoad.cfm] (accessed December 2007)

*Humphrey, J. and Hyatt. 2004. CSIRO Australian Animal Health Laboratory. Biological Control of the Cane Toad Bufo marinus in Australia

*Cory, J. and Myers, J. 2000. Direct and indirect ecological effects of biological control. Trends in Ecology & Evolution. 15, 4, 137-139.

*Johnson, M. 2000. Nature and Scope of Biological Control. Biological Control of Pest. 675

Effects on invasive species

*Cofrancesco, A. 2007. U.S. National Management Plan: An Action Plant for the Nation- Control and Management. Army Corps of Engineers. Available from [http://www.invasivespeciesinfo.gov/index.shtml]

*Lass, D. and Miller, R. 1995. BioScience. 45, 10. 680.

Effects on the future

*Cooksey D. 2002. [http://nature.berkeley.edu/bicon/what%20is%20Biological%20Control.htm Biological Control in Pest Management Systems of Plants.] Western Regional Committee, Riverside, CA.

Economic effects

*Griffiths, G.J.K. 2007. Efficacy and economics of shelter habitats for conservation. Biological Control: in press. doi:10.1016/j.biocontrol.2007.09.002

*Collier T. and Steenwyka, R. 2003. A critical evaluation of augmentative biological control. Economics of augmentation: 31, 245-256.

ee also

* Insectary plants
* Integrated Pest Management
* Japanese beetle (article includes information on biological control methods)
* Organic farming
* Biological pesticide
* Sterile insect technique
* Mating disruption

External links and references

* [http://www.nysaes.cornell.edu/ent/biocontrol/ Biological Control: A Guide to Natural Enemies in North America]
* [http://www.beyondpesticides.org Beyond Pesticides] - Provides information on pesticides and alternatives to their use.
* [http://grow.ars-informatica.ca/pest_control_index.php Grow'Em Organic Pest Control] - organic pest control solutions available to small- and large-scale gardeners, described in detail
* [http://www.ovam.be/jahia/Jahia/pid/467 Construction of a ecological garden and implementation of natural pest control]
* [http://www.provant.be/leefomgeving/natuur_en_bos/natuurvriendelijke_t/ 2 other pdf-files about the construction of a ecological garden and implementation of natural pest control]
* [http://207.5.17.151/biobest/en/films/dvd.htm Education movies about biological pest control of white flies, aphids and spider mites in greenhouses]
* [http://www.anbp.org Association of Natural Biocontrol Producers (ANBP)] Trade association of biological pest control industry.
* [http://gwss.mooreascience.org/index-en.htm Successful biocontrol of the GWSS in French Polynesia]
* cite journal
author=R. James Cook
journal=Annual Review of Phytopathology
date=September 1993
volume=31
pages=53–80
url=http://arjournals.annualreviews.org/doi/abs/10.1146/annurev.py.31.090193.000413
title=Making Greater Use of Introduced Microorganisms for Biological Control of Plant Pathogens
doi=10.1146/annurev.py.31.090193.000413
* cite paper
author=U.S. Congress, Office of Technology Assessment
publisher=US Government Printing Office, Washington, DC
date=1995
title=Biologically based technologies for pest control
work=OTA-ENV-636
url=http://www.princeton.edu/~ota/disk1/1995/9506/9506.PDF
* cite paper
author=Felix Wäckers, Paul van Rijn and Jan Bruin
publisher=Cambridge University Press, UK, 2005
date=2005
title=Plant-Provided Food for Carnivorous Insects - a protective mutualism and its applications.
ISBN 139780521819411
* [http://molecrickets.ifas.ufl.edu/mcri0001.htm Alternative Methods of Mole Cricket Control] at the University of Florida