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Biological control or biocontrol is a method of controlling pests, such as insects, mites, weeds, and plant diseases, using other organisms. It relies on predation,
parasitism Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted structurally to this way of lif ...
, herbivory, or other natural mechanisms, but typically also involves an active human management role. It can be an important component of integrated pest management (IPM) programs. There are three basic strategies for biological pest control: classical (importation), where a natural enemy of a pest is introduced in the hope of achieving control; inductive (augmentation), in which a large population of natural enemies are administered for quick pest control; and inoculative (conservation), in which measures are taken to maintain natural enemies through regular reestablishment. Natural enemies of insect pests, also known as biological control agents, include predators,
parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
s, pathogens, and competitors. Biological control agents of plant diseases are most often referred to as antagonists. Biological control agents of weeds include seed predators,
herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
s, and plant pathogens. Biological control can have side-effects on biodiversity through attacks on non-target species by any of the above mechanisms, especially when a species is introduced without a thorough understanding of the possible consequences.


History

The term "biological control" was first used by Harry Scott Smith at the 1919 meeting of the Pacific Slope Branch of the American Association of Economic Entomologists, in Riverside, California. It was brought into more widespread use by the entomologist Paul H. DeBach (1914–1993) who worked on citrus crop pests throughout his life. However, the practice has previously been used for centuries. The first report of the use of an insect species to control an insect pest comes from " Nanfang Caomu Zhuang" (南方草木狀 ''Plants of the Southern Regions'') (), attributed to Western Jin dynasty botanist ''Ji Han'' (嵇含, 263–307), in which it is mentioned that "'' Jiaozhi people sell ants and their nests attached to twigs looking like thin cotton envelopes, the reddish-yellow ant being larger than normal. Without such ants, southern citrus fruits will be severely insect-damaged''". The ants used are known as ''huang gan'' (''huang'' = yellow, ''gan'' = citrus) ants ('' Oecophylla smaragdina''). The practice was later reported by Ling Biao Lu Yi (late Tang Dynasty or Early Five Dynasties), in ''Ji Le Pian'' by ''Zhuang Jisu'' (
Southern Song Dynasty The Song dynasty (; ; 960–1279) was an imperial dynasty of China that began in 960 and lasted until 1279. The dynasty was founded by Emperor Taizu of Song following his usurpation of the throne of the Later Zhou. The Song conquered the rest ...
), in the ''Book of Tree Planting'' by Yu Zhen Mu ( Ming Dynasty), in the book ''Guangdong Xing Yu'' (17th century), ''Lingnan'' by Wu Zhen Fang (Qing Dynasty), in ''Nanyue Miscellanies'' by Li Diao Yuan, and others. Biological control techniques as we know them today started to emerge in the 1870s. During this decade, in the US, the Missouri State Entomologist C. V. Riley and the Illinois State Entomologist W. LeBaron began within-state redistribution of parasitoids to control crop pests. The first international shipment of an insect as a biological control agent was made by Charles V. Riley in 1873, shipping to France the predatory mites ''Tyroglyphus phylloxera'' to help fight the grapevine phylloxera ( ''Daktulosphaira vitifoliae'') that was destroying grapevines in France. The United States Department of Agriculture (USDA) initiated research in classical biological control following the establishment of the Division of Entomology in 1881, with C. V. Riley as Chief. The first importation of a parasitoidal wasp into the United States was that of the braconid '' Cotesia glomerata'' in 1883–1884, imported from Europe to control the invasive cabbage white butterfly, '' Pieris rapae''. In 1888–1889 the vedalia beetle, ''
Rodolia cardinalis ''Novius cardinalis'' (common names vedalia beetle or cardinal ladybird) is a species of ladybird beetle native to Australia. It was formerly placed in the genus ''Rodolia'', but that genus was synonymized under the genus '' Novius'' in 2020.Pan ...
'', a lady beetle, was introduced from
Australia Australia, officially the Commonwealth of Australia, is a sovereign ''Sovereign'' is a title which can be applied to the highest leader in various categories. The word is borrowed from Old French , which is ultimately derived from the L ...
to California to control the cottony cushion scale, '' Icerya purchasi''. This had become a major problem for the newly developed citrus industry in California, but by the end of 1889, the cottony cushion scale population had already declined. This great success led to further introductions of beneficial insects into the US.Coulson, J. R.; Vail, P. V.; Dix M.E.; Nordlund, D.A.; Kauffman, W.C.; Eds. 2000. 110 years of biological control research and development in the United States Department of Agriculture: 1883–1993. U.S. Department of Agriculture, Agricultural Research Service. pages=3–11 In 1905 the USDA initiated its first large-scale biological control program, sending entomologists to Europe and Japan to look for natural enemies of the spongy moth, '' Lymantria dispar dispar'', and the brown-tail moth, ''Euproctis chrysorrhoea'', invasive pests of trees and shrubs. As a result, nine parasitoids (solitary wasps) of the spongy moth, seven of the brown-tail moth, and two predators of both moths became established in the US. Although the spongy moth was not fully controlled by these natural enemies, the frequency, duration, and severity of its outbreaks were reduced and the program was regarded as successful. This program also led to the development of many concepts, principles, and procedures for the implementation of biological control programs.
Prickly pear cacti ''Opuntia'', commonly called prickly pear or pear cactus, is a genus of flowering plants in the cactus family Cactaceae. Prickly pears are also known as ''tuna'' (fruit), ''sabra'', ''nopal'' (paddle, plural ''nopales'') from the Nahuatl word f ...
were introduced into Queensland, Australia as ornamental plants, starting in 1788. They quickly spread to cover over 25 million hectares of Australia by 1920, increasing by 1 million hectares per year. Digging, burning, and crushing all proved ineffective. Two control agents were introduced to help control the spread of the plant, the cactus moth ''
Cactoblastis cactorum ''Cactoblastis cactorum'', the cactus moth, South American cactus moth or nopal moth, is native to Argentina, Paraguay, Uruguay and southern Brazil. It is one of five species in the genus '' Cactoblastis'' that inhabit South America, where many pa ...
'', and the scale insect '' Dactylopius''. Between 1926 and 1931, tens of millions of cactus moth eggs were distributed around Queensland with great success, and by 1932, most areas of prickly pear had been destroyed. The first reported case of a classical biological control attempt in
Canada Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by tota ...
involves the parasitoidal wasp '' Trichogramma minutum''. Individuals were caught in New York State and released in Ontario gardens in 1882 by William Saunders, a trained chemist and first Director of the Dominion Experimental Farms, for controlling the invasive currantworm ''
Nematus ribesii ''Nematus ribesii'' is a species of sawfly in the family Tenthredinidae. English names include common gooseberry sawfly Importation or classical biological control involves the introduction of a pest's natural enemies to a new locale where they do not occur naturally. Early instances were often unofficial and not based on research, and some introduced species became serious pests themselves. To be most effective at controlling a pest, a biological control agent requires a colonizing ability which allows it to keep pace with changes to the habitat in space and time. Control is greatest if the agent has temporal persistence so that it can maintain its population even in the temporary absence of the target species, and if it is an opportunistic forager, enabling it to rapidly exploit a pest population. One of the earliest successes was in controlling '' Icerya purchasi'' (cottony cushion scale) in Australia, using a predatory insect ''
Rodolia cardinalis ''Novius cardinalis'' (common names vedalia beetle or cardinal ladybird) is a species of ladybird beetle native to Australia. It was formerly placed in the genus ''Rodolia'', but that genus was synonymized under the genus '' Novius'' in 2020.Pan ...
'' (the vedalia beetle). This success was repeated in California using the beetle and a parasitoidal fly, ''
Cryptochaetum ''Cryptochetum'' is a genus of scale parasite flies in the family Cryptochetidae. There are more than 30 described species in ''Cryptochetum''. Species These 35 species belong to the genus ''Cryptochetum'': *'' C. acuticornutum'' (Yang & Yang, ...
iceryae''. Other successful cases include the control of '' Antonina graminis'' in Texas by '' Neodusmetia sangwani'' in the 1960s. Damage from '' Hypera postica'', the alfalfa weevil, a serious introduced pest of forage, was substantially reduced by the introduction of natural enemies. 20 years after their introduction the population of weevils in the
alfalfa Alfalfa () (''Medicago sativa''), also called lucerne, is a perennial flowering plant in the legume family Fabaceae. It is cultivated as an important forage crop in many countries around the world. It is used for grazing, hay, and silage, as w ...
area treated for alfalfa weevil in the
Northeastern United States The Northeastern United States, also referred to as the Northeast, the East Coast, or the American Northeast, is a geographic region of the United States. It is located on the Atlantic coast of North America, with Canada to its north, the Southe ...
remained 75 percent down. Alligator weed was introduced to the United States from South America. It takes root in shallow water, interfering with navigation, irrigation, and flood control. The alligator weed flea beetle and two other biological controls were released in Florida, greatly reducing the amount of land covered by the plant. Another aquatic weed, the giant salvinia ('' Salvinia molesta'') is a serious pest, covering waterways, reducing water flow and harming native species. Control with the salvinia weevil ('' Cyrtobagous salviniae'') and the salvinia stem-borer moth ('' Samea multiplicalis)'' is effective in warm climates, and in Zimbabwe, a 99% control of the weed was obtained over a two-year period. Small commercially reared parasitoidal wasps, '' Trichogramma ostriniae'', provide limited and erratic control of the European corn borer (''Ostrinia nubilalis''), a serious pest. Careful formulations of the bacterium ''
Bacillus thuringiensis ''Bacillus thuringiensis'' (or Bt) is a gram-positive, soil-dwelling bacterium, the most commonly used biological pesticide worldwide. ''B. thuringiensis'' also occurs naturally in the gut of caterpillars of various types of moths and butterflie ...
'' are more effective. The O. nubilalis integrated control releasing Tricogramma brassicae (egg parasitoiod) and later Bacillus thuringiensis subs. kurstaki (larvicidae effect) reduce pest damages as better than insecticide treatments The population of ''
Levuana iridescens The levuana moth (''Levuana iridescens'') is an extinct species of moth in the family Zygaenidae. It is monotypic within the genus ''Levuana''. The levuana moth became a serious pest for coconut plants in 1877, in Viti Levu, Fiji. On the islan ...
'', the Levuana moth, a serious coconut pest in
Fiji Fiji ( , ,; fj, Viti, ; Fiji Hindi: फ़िजी, ''Fijī''), officially the Republic of Fiji, is an island country in Melanesia, part of Oceania in the South Pacific Ocean. It lies about north-northeast of New Zealand. Fiji consists ...
, was brought under control by a classical biological control program in the 1920s.


Augmentation

Augmentation involves the supplemental release of natural enemies that occur in a particular area, boosting the naturally occurring populations there. In inoculative release, small numbers of the control agents are released at intervals to allow them to reproduce, in the hope of setting up longer-term control and thus keeping the pest down to a low level, constituting prevention rather than cure. In inundative release, in contrast, large numbers are released in the hope of rapidly reducing a damaging pest population, correcting a problem that has already arisen. Augmentation can be effective, but is not guaranteed to work, and depends on the precise details of the interactions between each pest and control agent. An example of inoculative release occurs in the horticultural production of several crops in
greenhouse A greenhouse (also called a glasshouse, or, if with sufficient heating, a hothouse) is a structure with walls and roof made chiefly of Transparent ceramics, transparent material, such as glass, in which plants requiring regulated climatic condit ...
s. Periodic releases of the parasitoidal wasp, '' Encarsia formosa'', are used to control greenhouse whitefly, while the predatory mite '' Phytoseiulus persimilis'' is used for control of the two-spotted spider mite. The egg parasite '' Trichogramma'' is frequently released inundatively to control harmful moths. New way for inundative releases are now introduced i.e. use of drones. Egg parasitoids are able to find the eggs of the target host by means of several cues. Kairomones were found on moth scales. Similarly, ''Bacillus thuringiensis'' and other microbial insecticides are used in large enough quantities for a rapid effect. Recommended release rates for ''Trichogramma'' in vegetable or field crops range from 5,000 to 200,000 per acre (1 to 50 per square metre) per week according to the level of pest infestation. Similarly, nematodes that kill insects (that are entomopathogenic) are released at rates of millions and even billions per acre for control of certain soil-dwelling insect pests.


Conservation

The conservation of existing natural enemies in an environment is the third method of biological pest control. Natural enemies are already adapted to the habitat and to the target pest, and their conservation can be simple and cost-effective, as when nectar-producing crop plants are grown in the borders of rice fields. These provide nectar to support parasitoids and predators of planthopper pests and have been demonstrated to be so effective (reducing pest densities by 10- or even 100-fold) that farmers sprayed 70% less insecticides and enjoyed yields boosted by 5%. Predators of aphids were similarly found to be present in tussock grasses by field boundary hedges in England, but they spread too slowly to reach the centers of fields. Control was improved by planting a meter-wide strip of tussock grasses in field centers, enabling aphid predators to overwinter there. Cropping systems can be modified to favor natural enemies, a practice sometimes referred to as habitat manipulation. Providing a suitable habitat, such as a shelterbelt,
hedgerow A hedge or hedgerow is a line of closely spaced shrubs and sometimes trees, planted and trained to form a barrier or to mark the boundary of an area, such as between neighbouring properties. Hedges that are used to separate a road from adjoini ...
, or beetle bank where beneficial insects such as parasitoidal wasps can live and reproduce, can help ensure the survival of populations of natural enemies. Things as simple as leaving a layer of fallen leaves or mulch in place provides a suitable food source for worms and provides a shelter for insects, in turn being a food source for such beneficial mammals as hedgehogs and shrews. Compost piles and stacks of wood can provide shelter for invertebrates and small mammals. Long grass and ponds support amphibians. Not removing dead annuals and non-hardy plants in the autumn allow insects to make use of their hollow stems during winter. In California, prune trees are sometimes planted in grape vineyards to provide an improved overwintering habitat or refuge for a key grape pest parasitoid. The providing of artificial shelters in the form of wooden caskets, boxes or flowerpots is also sometimes undertaken, particularly in gardens, to make a cropped area more attractive to natural enemies. For example, earwigs are natural predators that can be encouraged in gardens by hanging upside-down flowerpots filled with
straw Straw is an agricultural byproduct consisting of the dry stalks of cereal plants after the grain and chaff have been removed. It makes up about half of the yield of cereal crops such as barley, oats, rice, rye and wheat. It has a number ...
or wood wool. Green lacewings can be encouraged by using plastic bottles with an open bottom and a roll of cardboard inside. Birdhouses enable insectivorous birds to nest; the most useful birds can be attracted by choosing an opening just large enough for the desired species. In cotton production, the replacement of broad-spectrum insecticides with selective control measures such as Bt cotton can create a more favorable environment for natural enemies of cotton pests due to reduced insecticide exposure risk. Such predators or
parasitoids In evolutionary ecology, a parasitoid is an organism that lives in close association with its host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionary strategies within parasitis ...
can control pests not affected by the Bt protein. Reduced prey quality and abundance associated with increased control from Bt cotton can also indirectly decrease natural enemy populations in some cases, but the percentage of pests eaten or parasitized in Bt and non-Bt cotton are often similar.


Biological control agents


Predators

Predators are mainly free-living species that directly consume a large number of prey during their whole lifetime. Given that many major crop pests are insects, many of the predators used in biological control are insectivorous species. Lady beetles, and in particular their larvae which are active between May and July in the northern hemisphere, are voracious predators of aphids, and also consume
mites Mites are small arachnids (eight-legged arthropods). Mites span two large orders of arachnids, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari, but genetic analysis does not show clear evid ...
,
scale insect Scale insects are small insects of the order Hemiptera, suborder Sternorrhyncha. Of dramatically variable appearance and extreme sexual dimorphism, they comprise the infraorder Coccomorpha which is considered a more convenient grouping than the ...
s and small
caterpillar Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder Sym ...
s. The spotted lady beetle (''
Coleomegilla maculata ''Coleomegilla maculata'', commonly known as the spotted lady beetle, pink spotted lady beetle or twelve-spotted lady beetle, is a large coccinellid beetle native to North America. The adults and larvae feed primarily on aphids and the species ...
'') is also able to feed on the eggs and larvae of the Colorado potato beetle (''Leptinotarsa decemlineata''). The larvae of many hoverfly species principally feed upon aphids, one larva devouring up to 400 in its lifetime. Their effectiveness in commercial crops has not been studied. The running crab spider '' Philodromus cespitum'' also prey heavily on aphids, and act as a biological control agent in European fruit orchards. Several species of entomopathogenic nematode are important predators of insect and other invertebrate pests. Entomopathogenic nematodes form a stress–resistant stage known as the infective juvenile. These spread in the soil and infect suitable insect hosts. Upon entering the insect they move to the hemolymph where they recover from their stagnated state of development and release their bacterial
symbionts Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasit ...
. The bacterial symbionts reproduce and release toxins, which then kill the host insect. '' Phasmarhabditis hermaphrodita'' is a microscopic
nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-Parasitism, parasitic nematodes also known as eelworms. They are a diverse animal phylum inhab ...
that kills slugs. Its complex life cycle includes a free-living, infective stage in the soil where it becomes associated with a pathogenic bacteria such as '' Moraxella osloensis''. The nematode enters the slug through the posterior mantle region, thereafter feeding and reproducing inside, but it is the bacteria that kill the slug. The nematode is available commercially in Europe and is applied by watering onto moist soil. Entomopathogenic nematodes have a limited shelf life because of their limited resistance to high temperature and dry conditions. The type of soil they are applied to may also limit their effectiveness. Species used to control spider mites include the predatory mites '' Phytoseiulus persimilis'', '' Neoseilus californicus,'' and ''
Amblyseius ''Amblyseius'' is a large genus of predatory mites belonging to the family Phytoseiidae.de Moraes, G. J. (2005)Phytoseiidae Species Listing Biology Catalog, Texas A&M University. Retrieved on August 19, 2010. Many members of this genus feed on ...
cucumeris'', the predatory midge '' Feltiella acarisuga'', and a ladybird ''
Stethorus punctillum ''Stethorus punctillum'', known generally as the lesser mite destroyer or spider mite destroyer, is a species of lady beetle in the family Coccinellidae Coccinellidae () is a widespread family of small beetles ranging in size from . They a ...
''. The bug '' Orius insidiosus'' has been successfully used against the two-spotted spider mite and the western flower thrips (''Frankliniella occidentalis''). Predators including ''Cactoblastis cactorum'' (mentioned above) can also be used to destroy invasive plant species. As another example, the poison hemlock moth (''Agonopterix alstroemeriana)'' can be used to control poison hemlock (''Conium maculatum''). During its larval stage, the moth strictly consumes its host plant, poison hemlock, and can exist at hundreds of larvae per individual host plant, destroying large swathes of the hemlock. For rodent pests,
cat The cat (''Felis catus'') is a domestic species of small carnivorous mammal. It is the only domesticated species in the family Felidae and is commonly referred to as the domestic cat or house cat to distinguish it from the wild members of ...
s are effective biological control when used in conjunction with reduction of "harborage"/hiding locations. While cats are effective at preventing rodent "population explosions", they are not effective for eliminating pre-existing severe infestations. Barn owls are also sometimes used as biological rodent control. Although there are no quantitative studies of the effectiveness of barn owls for this purpose, they are known rodent predators that can be used in addition to or instead of cats; they can be encouraged into an area with nest boxes. In Honduras, where the mosquito '' Aedes aegypti'' was transmitting
dengue fever Dengue fever is a mosquito-borne tropical disease caused by the dengue virus. Symptoms typically begin three to fourteen days after infection. These may include a high fever, headache, vomiting, muscle and joint pains, and a characterist ...
and other infectious diseases, biological control was attempted by a community action plan;
copepod Copepods (; meaning "oar-feet") are a group of small crustaceans found in nearly every freshwater and saltwater habitat (ecology), habitat. Some species are planktonic (inhabiting sea waters), some are benthos, benthic (living on the ocean floor) ...
s, baby turtles, and juvenile tilapia were added to the wells and tanks where the mosquito breeds and the mosquito larvae were eliminated. Even amongst arthropods usually thought of as obligate predators of animals (especially other arthropods),
floral A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechanism ...
food sources (
nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...
and to a lesser degree
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
) are often useful adjunct sources. It had been noticed in one study that adult '' Adalia bipunctata'' (predator and common biocontrol of '' Ephestia kuehniella'') could survive on flowers but never completed its life cycle, so a meta-analysis was done to find such an overall trend in previously published data, if it existed. In some cases floral resources are outright necessary. Overall, floral resources (and an imitation, i.e. sugar water) increase
longevity The word " longevity" is sometimes used as a synonym for "life expectancy" in demography. However, the term ''longevity'' is sometimes meant to refer only to especially long-lived members of a population, whereas ''life expectancy'' is always d ...
and fecundity, meaning even predatory population numbers can depend on non-prey food abundance. Thus biocontrol population maintenance - and success - may depend on nearby flowers.


Parasitoids

Parasitoids lay their eggs on or in the body of an insect host, which is then used as a food for developing larvae. The host is ultimately killed. Most insect
parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
s are wasps or flies, and many have a very narrow host range. The most important groups are the ichneumonid wasps, which mainly use
caterpillar Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder Sym ...
s as hosts; braconid wasps, which attack caterpillars and a wide range of other insects including aphids; chalcidoid wasps, which parasitize eggs and larvae of many insect species; and tachinid flies, which parasitize a wide range of insects including caterpillars, beetle adults and larvae, and true bugs. Parasitoids are most effective at reducing pest populations when their host organisms have limited refuges to hide from them. Parasitoids are among the most widely used biological control agents. Commercially, there are two types of rearing systems: short-term daily output with high production of parasitoids per day, and long-term, low daily output systems. In most instances, production will need to be matched with the appropriate release dates when susceptible host species at a suitable phase of development will be available. Larger production facilities produce on a yearlong basis, whereas some facilities produce only seasonally. Rearing facilities are usually a significant distance from where the agents are to be used in the field, and transporting the parasitoids from the point of production to the point of use can pose problems. Shipping conditions can be too hot, and even vibrations from planes or trucks can adversely affect parasitoids. '' Encarsia formosa'' is a small parasitoid wasp attacking
whiteflies Whiteflies are Hemipterans that typically feed on the undersides of plant leaves. They comprise the family Aleyrodidae, the only family in the superfamily Aleyrodoidea. More than 1550 species have been described. Description and taxonomy The ...
, sap-feeding insects which can cause wilting and black sooty moulds in glasshouse vegetable and ornamental crops. 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 pupate. '' Gonatocerus ashmeadi'' (
Hymenoptera Hymenoptera is a large order (biology), order of insects, comprising the sawfly, sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are Par ...
: Mymaridae) has been introduced to control the glassy-winged sharpshooter ''Homalodisca vitripennis'' (Hemiptera: Cicadellidae) in
French Polynesia )Territorial motto: ( en, "Great Tahiti of the Golden Haze") , anthem = , song_type = Regional anthem , song = " Ia Ora 'O Tahiti Nui" , image_map = French Polynesia on the globe (French Polynesia centered).svg , map_alt = Location of Frenc ...
and has successfully controlled ~95% of the pest density. The eastern spruce budworm is an example of a destructive insect in
fir Firs (''Abies'') are a genus of 48–56 species of evergreen coniferous trees in the family (biology), family Pinaceae. They are found on mountains throughout much of North America, North and Central America, Europe, Asia, and North Africa. The ...
and
spruce A spruce is a tree of the genus ''Picea'' (), a genus of about 35 species of coniferous evergreen trees in the family Pinaceae, found in the northern temperate and boreal (taiga) regions of the Earth. ''Picea'' is the sole genus in the subfami ...
forests. Birds are a natural form of biological control, but the ''Trichogramma minutum'', a species of parasitic wasp, has been investigated as an alternative to more controversial chemical controls. There are a number of recent studies pursuing sustainable methods for controlling urban cockroaches using parasitic wasps. Since most cockroaches remain in the sewer system and sheltered areas which are inaccessible to insecticides, employing active-hunter wasps is a strategy to try and reduce their populations.


Pathogens

Pathogenic micro-organisms include bacteria, fungi, and viruses. They kill or debilitate their host and are relatively host-specific. Various
microbial A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in olde ...
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. The use of pathogens against
aquatic weed Aquatic plant management involves the science and methodologies used to control invasive and non-invasive aquatic plant species in waterways. Methods used include spraying herbicide, biological controls, mechanical removal as well as habitat modifi ...
s was unknown until a groundbreaking 1972 proposal by Zettler and Freeman. Up to that point biocontrol of any kind had not been used against any water weeds. In their review of the possibilities, they noted the lack of interest and information thus far, and listed what was known of pests-of-pests - whether pathogens or not. They proposed that this should be relatively straightfoward to apply in the same way as other biocontrols. And indeed in the decades since, the same biocontrol methods that are routine on land have become common in the water.


Bacteria

Bacteria used for biological control infect insects via their digestive tracts, so they offer only limited options for controlling insects with sucking mouth parts such as aphids and scale insects. ''
Bacillus thuringiensis ''Bacillus thuringiensis'' (or Bt) is a gram-positive, soil-dwelling bacterium, the most commonly used biological pesticide worldwide. ''B. thuringiensis'' also occurs naturally in the gut of caterpillars of various types of moths and butterflie ...
'', a soil-dwelling bacterium, is the most widely applied species of bacteria used for biological control, with at least four sub-species used against
Lepidoptera Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...
n ( moth, butterfly),
Coleoptera Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
n (beetle) and
Diptera Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...
n (true fly) insect pests. The bacterium is available to organic farmers in sachets of dried spores which are mixed with water and sprayed onto vulnerable plants such as brassicas and fruit trees. Genes from ''B. thuringiensis'' have also been incorporated into
transgenic crops Genetically modified crops (GM crops) are plants used in agriculture, the DNA of which has been modified using genetic engineering methods. Plant genomes can be engineered by physical methods or by use of '' Agrobacterium'' for the delivery of ...
, making the plants express some of the bacterium's toxins, which are proteins. These confer resistance to insect pests and thus reduce the necessity for pesticide use. If pests develop resistance to the toxins in these crops, ''B. thuringiensis'' will become useless in organic farming also. The bacterium '' Paenibacillus popilliae'' which causes milky spore disease has been found useful in the control of Japanese beetle, killing the larvae. It is very specific to its host species and is harmless to vertebrates and other invertebrates. '' Bacillus'' spp.,p.94-5, II. Biocontrol Modes of Action fluorescent Pseudomonads, and
Streptomycete The ''Streptomycetaceae'' are a family of ''Actinomycetota'', making up the monotypic order ''Streptomycetales''. It includes the important genus ''Streptomyces''. This was the original source of many antibiotics, namely streptomycin, the first a ...
s are controls of various fungal pathogens.p.94


Fungi

Entomopathogenic fungi, which cause disease in insects, include at least 14 species that attack aphids. ''
Beauveria bassiana ''Beauveria bassiana'' is a fungus that grows naturally in soils throughout the world and acts as a parasite on various arthropod species, causing white muscardine disease; it thus belongs to the entomopathogenic fungi. It is used as a biological ...
'' is mass-produced and used to manage a wide variety of insect pests including
whiteflies Whiteflies are Hemipterans that typically feed on the undersides of plant leaves. They comprise the family Aleyrodidae, the only family in the superfamily Aleyrodoidea. More than 1550 species have been described. Description and taxonomy The ...
, thrips, aphids and weevils. '' Lecanicillium'' spp. are deployed against white flies, thrips and aphids. '' Metarhizium'' spp. are used against pests including beetles,
locusts Locusts (derived from the Vulgar Latin ''locusta'', meaning grasshopper) are various species of short-horned grasshoppers in the family Acrididae that have a swarming phase. These insects are usually solitary, but under certain circumst ...
and other grasshoppers,
Hemiptera Hemiptera (; ) is an order (biology), order of insects, commonly called true bugs, comprising over 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, Reduviidae, assassin bugs, Cimex, bed bugs, and shield bugs. ...
, and
spider mite Spider mites are members of the Tetranychidae family, which includes about 1,200 species. They are part of the subclass Acari (mites). Spider mites generally live on the undersides of leaves of plants, where they may spin protective silk webs, a ...
s. ''
Paecilomyces fumosoroseus ''Isaria fumosorosea'' is an entomopathogenic fungus, formerly known as ''Paecilomyces fumosoroseus''. It shows promise as a biological pesticide with an extensive host range. Life cycle When a conidium or blastospore of ''Isaria fumosorosea'' ...
'' is effective against white flies, thrips and aphids; '' Purpureocillium lilacinus'' is used against root-knot nematodes, and 89 ''
Trichoderma ''Trichoderma'' is a genus of fungi in the family Hypocreaceae that is present in all soils, where they are the most prevalent culturable fungi. Many species in this genus can be characterized as opportunistic avirulent plant symbionts. This ...
'' species against certain plant pathogens.p.93 '' Trichoderma viride'' has been used against Dutch elm disease, and has shown some effect in suppressing silver leaf, a disease of stone fruits caused by the pathogenic fungus '' Chondrostereum purpureum''. Pathogenic fungi may be controlled by other fungi, or bacteria or yeasts, such as: ''
Gliocladium ''Gliocladium''Corda (1840) ''Icon. fung. (Prague)'' 4: 30. is an asexual fungal genus in the Hypocreaceae. Certain other species including ''Gliocladium virens'' were recently transferred to the genus ''Trichoderma'' and ''G. roseum'' became ' ...
'' spp., mycoparasitic '' Pythium'' spp.,
binucleate Binucleated cells are cells that contain two nuclei. This type of cell is most commonly found in cancer cells and may arise from a variety of causes. Binucleation can be easily visualized through staining and microscopy. In general, binucleation ...
types of '' Rhizoctonia'' spp., and '' Laetisaria'' spp. The fungi ''
Cordyceps ''Cordyceps'' is a genus of ascomycete fungi (sac fungi) that includes about 600 species. Most ''Cordyceps'' species are endoparasitoids, parasitic mainly on insects and other arthropods (they are thus entomopathogenic fungi); a few are parasiti ...
'' and '' Metacordyceps'' are deployed against a wide spectrum of arthropods. ''