Biological pest control

Biological control or biocontrol is a method of controlling pests such as insects, mites, weeds and plant diseases using other organisms.[1] It relies on predation, parasitism, 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.[2]

Natural enemies of insect pests, also known as biological control agents, include predators, parasitoids, pathogens, and competitors. Biological control agents of plant diseases are most often referred to as antagonists. Biological control agents of weeds include seed predators, herbivores and plant pathogens.

Biological control can have side-effects on biodiversity through attacks on non-target species by any of the same mechanisms, especially when a species is introduced without thorough understanding of the possible consequences.

Syrphid.maggot3554.5.13.08cw
Syrphus hoverfly larva (below) feed on aphids (above), making them natural biological control agents.
Cotesia9061.8.15.07.c
A parasitoid wasp (Cotesia congregata) adult with pupal cocoons on its host, a tobacco hornworm (Manduca sexta, green background), an example of a hymenopteran biological control agent

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.[3] It was brought into more widespread use by the entomologist Paul H. DeBach (1914–1993) who worked on citrus crop pests throughout his life.[4][5] 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) (c. 304 AD), 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".[6] 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), 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.[6]

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 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, a lady beetle, was introduced from Australia 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.[7][8]

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 gypsy moth, Lymantria dispar dispar, and brown-tail moth, Euproctis chrysorrhoea, invasive pests of trees and shrubs. As a result, nine parasitoids (solitary wasps) of gypsy moth, seven of brown-tail moth, and two predators of both moths became established in the US. Although the gypsy 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.[7][8][9]

Larvaefeedingoncacti
Cactoblastis cactorum larvae feeding on Opuntia prickly pear cacti

Prickly pear cacti 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, 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.[10]

The first reported case of a classical biological control attempt in Canada involves the parasitoidal wasp Trichogramma minutum. Individuals were caught in New York State and released in Ontario gardens in 1882 by William Saunders, trained chemist and first Director of the Dominion Experimental Farms, for controlling the invasive currantworm Nematus ribesii. Between 1884 and 1908, the first Dominion Entomologist, James Fletcher, continued introductions of other parasitoids and pathogens for the control of pests in Canada.[11]

Types of biological pest control

There are three basic biological pest control strategies: importation (classical biological control), augmentation and conservation.[12]

Importation

Vedalia Beetle (15959056801)
Rodolia cardinalis, the vedalia beetle, was imported from Australia to California in the 19th century, successfully controlling cottony cushion scale.

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.[13]

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.[14]

Joseph Needham noted a Chinese text dating from 304 AD, Records of the Plants and Trees of the Southern Regions, by Hsi Han, which describes mandarin oranges protected by large reddish-yellow citrus ants which attack and kill insect pests of the orange trees. The citrus ant (Oecophylla smaragdina)[15] was rediscovered in the 20th century, and since 1958 has been used in China to protect orange groves.[16]

One of the earliest successes in the west was in controlling Icerya purchasi (cottony cushion scale) in Australia, using a predatory insect Rodolia cardinalis (the vedalia beetle). This success was repeated in California using the beetle and a parasitoidal fly, Cryptochaetum iceryae.[17] Other successful cases include the control of Antonina graminis in Texas by Neodusmetia sangwani in the 1960s.[18]

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 area treated for alfalfa weevil in the Northeastern United States remained 75 percent down.[19]

Alternanthera philoxeroides NRCS-1
The invasive species Alternanthera philoxeroides (alligator weed) was controlled in Florida (U.S.) by introducing alligator weed flea beetle.

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.[20] 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,[21][22] and in Zimbabwe, a 99% control of the weed was obtained over a two-year period.[23]

Small commercially reared parasitoidal wasps,[12] Trichogramma ostriniae, provide limited and erratic control of the European corn borer (Ostrinia nubilalis), a serious pest. Careful formulations of the bacterium Bacillus thuringiensis are more effective.[24]

The population of Levuana iridescens, the Levuana moth, a serious coconut pest in Fiji, was brought under control by a classical biological control program in the 1920s.[25]

Augmentation

Lady bugs are a beneficial insect commonly sold for biological control of aphids.
Hippodamia convergens, the convergent lady beetle, is commonly sold for biological control of aphids.

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.[26]

An example of inoculative release occurs in the horticultural production of several crops in greenhouses. Periodic releases of the parasitoidal wasp, Encarsia formosa, are used to control greenhouse whitefly,[27] while the predatory mite Phytoseiulus persimilis is used for control of the two-spotted spider mite.[28]

The egg parasite Trichogramma is frequently released inundatively to control harmful moths. Similarly, Bacillus thuringiensis and other microbial insecticides are used in large enough quantities for a rapid effect.[26] 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.[29] 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.[30]

Conservation

The conservation of existing natural enemies in an environment is the third method of biological pest control.[31] 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%.[32] 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 centres of fields. Control was improved by planting a metre-wide strip of tussock grasses in field centres, enabling aphid predators to overwinter there.[31]

Dermaptera flowerpot
An inverted flowerpot filled with straw to attract earwigs

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, 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 allows insects to make use of their hollow stems during winter.[33] In California, prune trees are sometimes planted in grape vineyards to provide an improved overwintering habitat or refuge for a key grape pest parasitoid.[34] 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 which can be encouraged in gardens by hanging upside-down flowerpots filled with straw 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.[33]

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 can control pests not affected by the Bt protein. Reduced prey quality and abundance associated 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.[35]

Biological control agents

Predators

Chrysopidae 3035
Predatory lacewings are available from biocontrol dealers.

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, scale insects and small caterpillars. The spotted lady beetle (Coleomegilla maculata) is also able to feed on the eggs and larvae of the Colorado potato beetle (Leptinotarsa decemlineata).[36]

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.[37]

Organic-agriculture biocontrol-cotton polistes-wasp3
Predatory Polistes wasp searching for bollworms or other caterpillars on a cotton plant

Several species of entomopathogenic nematode are important predators of insect and other invertebrate pests.[38] Phasmarhabditis hermaphrodita is a microscopic nematode 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.[39]

Species used to control spider mites include the predatory mites Phytoseiulus persimilis,[40] Neoseilus californicus,[41] and Amblyseius cucumeris, the predatory midge Feltiella acarisuga,[41] and a ladybird Stethorus punctillum.[41] The bug Orius insidiosus has been successfully used against the two-spotted spider mite and the western flower thrips (Frankliniella occidentalis).[42]

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.[43]

Aleiodes indiscretus wasp parasitizing gypsy moth caterpillar
The parasitoid wasp Aleiodes indiscretus parasitizing a gypsy moth caterpillar, a serious pest of forestry[44]

For rodent pests, cats are effective biological control when used in conjunction with reduction of "harborage"/hiding locations.[45][46][47] While cats are effective at preventing rodent "population explosions", they are not effective for eliminating pre-existing severe infestations.[47] Barn owls are also sometimes used as biological rodent control.[48] Although there are no quantitative studies of the effectiveness of barn owls for this purpose,[49] they are known rodent predators that can be used in addition to or instead of cats;[50][51] they can be encouraged into an area with nest boxes.[52][53]

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 parasitoids are wasps or flies, and many have a very narrow host range. The most important groups are the ichneumonid wasps, which mainly use caterpillars as hosts; braconid wasps, which attack caterpillars and a wide range of other insects including aphids; chalcid 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.[54] Parasitoids are most effective at reducing pest populations when their host organisms have limited refuges to hide from them.[55]

Encarsia formosa, an endoparasitic wasp, is used for whitefly control
Encarsia formosa, widely used in greenhouse horticulture, was one of the first biological control agents developed.
Waspcycle
Life cycles of greenhouse whitefly and its parasitoid wasp Encarsia formosa

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.[56] 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.[57] 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.[58] Shipping conditions can be too hot, and even vibrations from planes or trucks can adversely affect parasitoids.[56]

Encarsia formosa is a small predatory chalcid wasp which is a parasitoid of whitefly, a sap-feeding insect 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.[27] Gonatocerus ashmeadi (Hymenoptera: Mymaridae) has been introduced to control the glassy-winged sharpshooter Homalodisca vitripennis (Hemiptera: Cicadellidae) in French Polynesia and has successfully controlled ~95% of the pest density.[59]

The eastern spruce budworm is an example of a destructive insect in fir and spruce 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.[60]

There are a number of recent studies pursuing sustainable methods for controlling urban cockroaches using parasitic wasps.[61][62] 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 insect diseases occur naturally, but may also be used as biological pesticides.[63] When naturally occurring, these outbreaks are density-dependent in that they generally only occur as insect populations become denser.[64]

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.[65] Bacillus thuringiensis, a soil-dwelling bacterium, is the most widely applied species of bacteria used for biological control, with at least four sub-species used against Lepidopteran (moth, butterfly), Coleopteran (beetle) and Dipteran (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.[66][67] Genes from B. thuringiensis have also been incorporated into transgenic crops, 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.[68] If pests develop resistance to the toxins in these crops, B. thuringiensis will become useless in organic farming also.[69][67] 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.[70]

Fungi

Pandora neoaphidis
Green peach aphid, a pest in its own right and a vector of plant viruses, killed by the fungus Pandora neoaphidis (Zygomycota: Entomophthorales) Scale bar = 0.3 mm.

Entomopathogenic fungi, which cause disease in insects, include at least 14 species that attack aphids.[71] Beauveria bassiana is mass-produced and used to manage a wide variety of insect pests including whiteflies, thrips, aphids and weevils.[72] Lecanicillium spp. are deployed against white flies, thrips and aphids. Metarhizium spp. are used against pests including beetles, locusts and other grasshoppers, Hemiptera, and spider mites. Paecilomyces fumosoroseus is effective against white flies, thrips and aphids; Purpureocillium lilacinus is used against root-knot nematodes, and 89 Trichoderma species against certain plant pathogens. 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.[73]

The fungi Cordyceps and Metacordyceps are deployed against a wide spectrum of arthropods.[74] Entomophaga is effective against pests such as the green peach aphid.[75]

Several members of Chytridiomycota and Blastocladiomycota have been explored as agents of biological control.[76][77] From Chytridiomycota, Synchytrium solstitiale is being considered as a control agent of the yellow star thistle (Centaurea solstitialis) in the United States.[78]

Viruses

Baculoviruses are specific to individual insect host species and have been shown to be useful in biological pest control. For example, the Lymantria dispar multicapsid nuclear polyhedrosis virus has been used to spray large areas of forest in North America where larvae of the gypsy moth are causing serious defoliation. The moth larvae are killed by the virus they have eaten and die, the disintegrating cadavers leaving virus particles on the foliage to infect other larvae.[79]

A mammalian virus, the rabbit haemorrhagic disease virus was introduced to Australia to attempt to control the European rabbit populations there.[80] It escaped from quarantine and spread across the country, killing large numbers of rabbits. Very young animals survived, passing immunity to their offspring in due course and eventually producing a virus-resistant population.[81] Introduction into New Zealand in the 1990s was similarly successful at first, but a decade later, immunity had developed and populations had returned to pre-RHD levels.[82]

Oomycota

Lagenidium giganteum is a water-borne mold that parasitizes the larval stage of mosquitoes. When applied to water, the motile spores avoid unsuitable host species and search out suitable mosquito larval hosts. This mold has the advantages of a dormant phase, resistant to desiccation, with slow-release characteristics over several years. Unfortunately, it is susceptible to many chemicals used in mosquito abatement programmes.[83]

Competitors

The legume vine Mucuna pruriens is used in the countries of Benin and Vietnam as a biological control for problematic Imperata cylindrica grass: the vine is extremely vigorous and suppresses neighbouring plants by out-competing them for space and light. Mucuna pruriens is said not to be invasive outside its cultivated area.[84] Desmodium uncinatum can be used in push-pull farming to stop the parasitic plant, witchweed (Striga).[85]

The Australian bush fly, Musca vetustissima, is a major nuisance pest in Australia, but native decomposers found in Australia are not adapted to feeding on cow dung, which is where bush flies breed. Therefore, the Australian Dung Beetle Project (1965–1985), led by George Bornemissza of the Commonwealth Scientific and Industrial Research Organisation, released forty-nine species of dung beetle, to reduce the amount of dung and therefore also the potential breeding sites of the fly.[86]

Combined use of parasitoids and pathogens

In cases of massive and severe infection of invasive pests, techniques of pest control are often used in combination. An example is the emerald ash borer, Agrilus planipennis, an invasive beetle from China, which has destroyed tens of millions of ash trees in its introduced range in North America. As part of the campaign against it, from 2003 American scientists and the Chinese Academy of Forestry searched for its natural enemies in the wild, leading to the discovery of several parasitoid wasps, namely Tetrastichus planipennisi, a gregarious larval endoparasitoid, Oobius agrili, a solitary, parthenogenic egg parasitoid, and Spathius agrili, a gregarious larval ectoparasitoid. These have been introduced and released into the United States of America as a possible biological control of the emerald ash borer. Initial results for Tetrastichus planipennisi have shown promise, and it is now being released along with Beauveria bassiana, a fungal pathogen with known insecticidal properties.[87][88][89]

Difficulties

Many of the most important pests are exotic, invasive species that severely impact agriculture, horticulture, forestry and urban environments. They tend to arrive without their co-evolved parasites, pathogens and predators, and by escaping from these, populations may soar. Importing the natural enemies of these pests may seem a logical move but this may have unintended consequences; regulations may be ineffective and there may be unanticipated effects on biodiversity, and the adoption of the techniques may prove challenging because of a lack of knowledge among farmers and growers.[90]

Side effects

Biological control can affect biodiversity[14] through predation, parasitism, pathogenicity, competition, or other attacks on non-target species.[91] An introduced control does not always target only the intended pest species; it can also target native species.[92] In Hawaii during the 1940s parasitic wasps were introduced to control a lepidopteran pest and the wasps are still found there today. This may have a negative impact on the native ecosystem; however, host range and impacts need to be studied before declaring their impact on the environment.[93]

Cane toad distribution stills
Cane toad (introduced into Australia 1935) spread from 1940 to 1980: it was ineffective as a control agent. Its distribution has continued to widen since 1980.

Vertebrate animals tend to be generalist feeders, and seldom make good biological control agents; many of the classic cases of "biocontrol gone awry" involve vertebrates. For example, the cane toad (Rhinella marina) was intentionally introduced to Australia to control the greyback cane beetle (Dermolepida albohirtum),[94] and other pests of sugar cane. 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 could not jump very high and so were unable to eat the cane beetles which stayed on the upper stalks of the cane plants. However, the toad thrived by feeding on other insects and soon spread very rapidly; it took over native amphibian habitat and brought foreign disease to native toads and frogs, dramatically reducing their populations. Also, when it is threatened or handled, the cane toad releases poison from parotoid glands on its shoulders; native Australian species such as goannas, tiger snakes, dingos and northern quolls that attempted to eat the toad were harmed or killed. However, there has been some recent evidence that native predators are adapting, both physiologically and through changing their behaviour, so in the long run, their populations may recover.[95]

Rhinocyllus conicus, a seed-feeding weevil, was introduced to North America to control exotic musk thistle (Carduus nutans) and Canadian thistle (Cirsium arvense). However, the weevil also attacks native thistles, harming such species as the endemic Platte thistle (Cirsium neomexicanum) by selecting larger plants (which reduced the gene pool), reducing seed production and ultimately threatening the species' survival.[96] Similarly, the weevil Larinus planus was also used to try to control the Canadian thistle, but it damaged other thistles as well.[97][98] This included one species classified as threatened.[99]

The small Asian mongoose (Herpestus javanicus) was introduced to Hawaii in order to control the rat population. However, the mongoose was diurnal, and the rats emerged at night; the mongoose therefore preyed on the endemic birds of Hawaii, especially their eggs, more often than it ate the rats, and now both rats and mongooses threaten the birds. This introduction was undertaken without understanding the consequences of such an action. No regulations existed at the time, and more careful evaluation should prevent such releases now.[100]

The sturdy and prolific eastern mosquitofish (Gambusia holbrooki) is a native of the southeastern United States and was introduced around the world in the 1930s and '40s to feed on mosquito larvae and thus combat malaria. However, it has thrived at the expense of local species, causing a decline of endemic fish and frogs through competition for food resources, as well as through eating their eggs and larvae.[101] In Australia, control of the mosquitofish is the subject of discussion; in 1989 researchers A. H. Arthington and L. L. Lloyd stated that "biological population control is well beyond present capabilities".[102]

Grower education

A potential obstacle to the adoption of biological pest control measures is that growers may prefer to stay with the familiar use of pesticides. However, pesticides have undesired effects, including the development of resistance among pests, and the destruction of natural enemies; these may in turn enable outbreaks of pests of other species than the ones originally targeted, and on crops at a distance from those treated with pesticides.[103] One method of increasing grower adoption of biocontrol methods involves letting them learn by doing, for example showing them simple field experiments, enabling them to observe the live predation of pests, or demonstrations of parasitised pests. In the Philippines, early season sprays against leaf folder caterpillars were common practice, but growers were asked to follow a 'rule of thumb' of not spraying against leaf folders for the first 30 days after transplanting; participation in this resulted in a reduction of insecticide use by 1/3 and a change in grower perception of insecticide use.[104]

See also

References

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Further reading

General
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.; Shelton, A. M.; Hoffman, M. P. Biological Control: A Guide to Natural Enemies in North America.
  • Cane toad: a case study. 2003.
  • Humphrey, J. and Hyatt. 2004. CSIRO Australian Animal Health Laboratory. Biological Control of the Cane Toad Bufo marinus in Australia
  • Cory, J.; Myers, J. (2000). "Direct and indirect ecological effects of biological control". Trends in Ecology & Evolution. 15 (4): 137–139. doi:10.1016/s0169-5347(99)01807-8.
  • Johnson, M. 2000. Nature and Scope of Biological Control. Biological Control of Pests.
Effects on invasive species
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.; Steenwyka, R. 2003. A critical evaluation of augmentative biological control. Economics of augmentation: 31, 245–256.

External links

Alastor (wasp)

Alastor is a Palearctic, Indomalayan and Afrotropical genus of potter wasps.

Aphelinidae

The Aphelinidae are a moderate-sized family of tiny parasitic wasps, with about 1160 described species in some 35 genera. These minute insects are challenging to study, as they deteriorate rapidly after death unless extreme care is taken (e.g., preservation in ethanol), making identification of most museum specimens difficult. The larvae of the majority are primary parasitoids on Hemiptera, though other hosts are attacked, and details of the life history can be variable (e.g., some attack eggs, some attack pupae, and others are hyperparasites). Males and females may have different hosts and different life histories [1].

They are found throughout the world in virtually all habitats, and are extremely important as biological control agents.

They are difficult to separate from other Chalcidoidea except by subtle features of the wing venation and other difficult characters, and the family appears to be paraphyletic, so is likely to be split up in the future (e.g., the Azotinae and Calesinae may become separate families).

Biocide

A biocide is defined in the European legislation as a chemical substance or microorganism intended to destroy, deter, render harmless, or exert a controlling effect on any harmful organism. The US Environmental Protection Agency (EPA) uses a slightly different definition for biocides as "a diverse group of poisonous substances including preservatives, insecticides, disinfectants, and pesticides used for the control of organisms that are harmful to human or animal health or that cause damage to natural or manufactured products". When compared, the two definitions roughly imply the same, although the US EPA definition includes plant protection products and some veterinary medicines.

The terms "biocides" and "pesticides" are regularly interchanged, and often confused with "plant protection products". To clarify this, pesticides include both biocides and plant protection products, where the former refers to substances for non-food and feed purposes and the latter refers to substances for food and feed purposes.

When discussing biocides a distinction should be made between the biocidal active substance and the biocidal product. The biocidal active substances are mostly chemical compounds, but can also be microorganisms (e.g. bacteria). Biocidal products contain one or more biocidal active substances and may contain other non-active co-formulants that ensure the effectiveness as well as the desired pH, viscosity, colour, odour, etc. of the final product. Biocidal products are available on the market for use by professional and/or non-professional consumers.

Although most of the biocidal active substances have a relative high toxicity, there are also examples of active substances with low toxicity, such as CO2, which exhibit their biocidal activity only under certain specific conditions such as in closed systems. In such cases, the biocidal product is the combination of the active substance and the device that ensures the intended biocidal activity, i.e. suffocation of rodents by CO2 in a closed system trap. Another example of biocidal products available to consumers are products impregnated with biocides (also called treated articles), such as clothes and wristbands impregnated with insecticides, socks impregnated with antibacterial substances etc.

Biocides are commonly used in medicine, agriculture, forestry, and industry. Biocidal substances and products are also employed as anti-fouling agents or disinfectants under other circumstances: chlorine, for example, is used as a short-life biocide in industrial water treatment but as a disinfectant in swimming pools. Many biocides are synthetic, but there are naturally occurring biocides classified as natural biocides, derived from, e.g., bacteria and plants.A biocide can be:

A pesticide: this includes fungicides, herbicides, insecticides, algicides, molluscicides, miticides, rodenticides, and slimicides.

An antimicrobial: this includes germicides, antibiotics, antibacterials, antivirals, antifungals, antiprotozoals, and antiparasites. See also spermicide.

Braconidae

The Braconidae are a family of parasitoid wasps. After the closely related Ichneumonidae, braconids make up the second-largest family in the order Hymenoptera, with about 17,000 recognized species and many thousands more undescribed. One analysis estimated a total between 30,000 and 50,000, and another provided a narrower estimate between 42,000 and 43,000 species.

Chalcid wasp

Chalcid wasps (, from Greek khalkos, meaning 'copper', for their metallic colour) are insects within the superfamily Chalcidoidea, part of the order Hymenoptera. The superfamily contains some 22,500 known species, and an estimated total diversity of more than 500,000 species, meaning the vast majority have yet to be discovered and described. The name "chalcid" is often confused with the name "chalcidid", though the latter refers strictly to one constituent family, the Chalcididae, rather than the superfamily as a whole; accordingly, most recent publications (e.g.,) use the name "chalcidoid" when referring to members of the superfamily.

Most of the species are parasitoids of other insects, attacking the egg or larval stage of their host, though many other life cycles are known. These hosts are to be found in at least 12 different insect orders including Lepidoptera (butterflies and moths), Diptera (true flies), Coleoptera (beetles), Hemiptera (true bugs), and other Hymenoptera, as well as two orders of Arachnida, and even one family of nematodes. For example, the chalcid fly is responsible for a small percentage of egg death in the wood white butterfly (L. sinapis). When the host is itself a parasitoid, they are referred to as hyperparasitoids. A small percentage are phytophagous and the larvae feed inside seeds, stems, and galls, including some that act as pollinators (e.g. fig wasps). Generally beneficial to humans as a group, chalcidoids help keep various crop pests under control, and many species have been imported as biocontrol agents. Moth parasitoid Copidosoma floridanum is one such species, whose genome is being sequenced by the Human Genome Sequencing Center as part of the i5K project, which aims to sequence the genomes of 5,000 arthropods.

Chalcidoids are tiny, dark-coloured wasps, typically black or brown, but often metallic blue or green, with complex sculpturing on the body. They are also recognized by the characteristic reduced wing venation, similar to that seen in other superfamilies of parasitoid wasps.

Companion planting

Companion planting in gardening and agriculture is the planting of different crops in proximity for any of a number of different reasons, including pest control, pollination, providing habitat for beneficial insects, maximizing use of space, and to otherwise increase crop productivity. Companion planting is a form of polyculture.

Companion planting is used by farmers and gardeners in both industrialized and developing countries for many reasons. Many of the modern principles of companion planting were present many centuries ago in cottage gardens in England and forest gardens in Asia, and thousands of years ago in Mesoamerica.

Encarsia formosa

Encarsia formosa is a species of chalcidoid wasp and a well known parasitoid of greenhouse whitefly, one of the first to be used commercially for biological pest control, from the 1920s. They can use at least 15 species of whitefly as a host, including Bemisia tabaci and Aleyrodes proletella.The tiny females (about 0.6 mm long) are black with a yellow abdomen and opalescent wings. This species reproduces asexually via thelytoky induced by Wolbachia infection. Males are produced only rarely. They are slightly larger than females and are completely black in coloration.

Flufenoxuron

Flufenoxuron is an insecticide that belongs to the benzoylurea group, which also includes diflubenzuron, triflumuron, and lufenuron.Flufenoxuron is a white crystalline powder. It is insoluble in water, is not flammable, and is not an oxidizer.

Ichneumonoidea

The superfamily Ichneumonoidea contains the two largest families within Hymenoptera: Ichneumonidae and Braconidae. The group is thought to contain as many as 100,000 species, many of which have not yet been described. Like other parasitoid wasps, they were long placed in the "Parasitica", variously considered as an infraorder or an unranked clade, now known to be paraphyletic.

Isaria fumosorosea

Isaria fumosorosea is an entomopathogenic fungus, formerly known as Paecilomyces fumosoroseus. It shows promise as a biological pesticide with an extensive host range.

Metarhizium rileyi

Metarhizium rileyi is a species of entomopathogenic fungus in the family Clavicipitaceae; there is extensive literature under its synonym Nomuraea rileyi.

Orange oil

Orange oil is an essential oil produced by cells within the rind of an orange fruit (Citrus sinensis fruit). In contrast to most essential oils, it is extracted as a by-product of orange juice production by centrifugation, producing a cold-pressed oil. It is composed of mostly (greater than 90%) d-limonene, and is often used in place of pure d-limonene. D-limonene can be extracted from the oil by distillation.

Paper wasp

Paper wasps are vespid wasps that gather fibers from dead wood and plant stems, which they mix with saliva, and use to construct water-resistant nests made of gray or brown papery material. Some types of paper wasps are also sometimes called umbrella wasps, due to the distinctive design of their nests.

Parasitoid

A parasitoid is an organism that lives in close association with its host and at the host's expense, and which sooner or later kills it. Parasitoidism is one of six major evolutionary strategies within parasitism, distinguished by the fatal prognosis for the host, which makes the strategy close to predation.

Among parasitoids, strategies range from living inside the host, allowing it to go on growing until the parasitoid emerges as an adult, to paralysing the host and living outside it. Hosts include other parasitoids, resulting in hyperparasitism; in the case of oak galls, up to five levels of parasitism are possible. Some parasitoids influence their host's behaviour in ways that favour the propagation of the parasitoid.

Parasitoids are found in a variety of taxa across the endopterygote insects, whose complete metamorphosis may have pre-adapted them for a split lifestyle, with parasitoid larvae and freeliving adults. Most are in the Hymenoptera, where the ichneumons and many other parasitoid wasps are highly specialised for a parasitoidal way of life. Other parasitoids are in the Diptera, Coleoptera and other orders of endopterygote insects. Some of these, usually but not only wasps, are used in biological pest control.

The biology of parasitoidism has inspired science fiction authors and scriptwriters to create numerous parasitoidal aliens that kill their human hosts, such as the alien species in Ridley Scott's 1979 film Alien.

Potter wasp

Potter wasps (or mason wasps), the Eumeninae, are a cosmopolitan wasp group presently treated as a subfamily of Vespidae, but sometimes recognized in the past as a separate family, Eumenidae.

Pyrethrum

Pyrethrum was a genus of several Old World plants now classified as Chrysanthemum or Tanacetum (e.g., C. coccineum) which are cultivated as ornamentals for their showy flower heads. Pyrethrum continues to be used as a common name for plants formerly included in the genus Pyrethrum. Pyrethrum is also the name of a natural insecticide made from the dried flower heads of Chrysanthemum cinerariifolium and Chrysanthemum coccineum. Its active ingredient are pyrethrins.

Scelionidae

The hymenopteran family Scelionidae is a very large cosmopolitan group (over 3000 described species in some 160 genera) of exclusively parasitoid wasps, mostly small (0.5–10 mm), often black, often highly sculptured, with (typically) elbowed antennae that have a 9- or 10-segmented flagellum. Nowadays, it is considered to be a subfamily of the Platygastridae.

They are generally idiobionts, attacking the eggs of many different types of insects, spiders, butterflies (the hackberry emperor, for example) and many are important in biological control. Several genera are wingless, and a few attack aquatic insect eggs underwater.

Soldier beetle

The soldier beetles (Cantharidae) are relatively soft-bodied, straight-sided beetles. They are cosmopolitan in distribution. One of the first described species has a color pattern reminiscent of the red coats of early British soldiers, hence the common name. They are also known commonly as leatherwings because of their soft elytra.Historically, these beetles were placed in a superfamily "Cantharoidea", which has been subsumed by the superfamily Elateroidea; the name is still sometimes used as a rankless grouping, including the families Cantharidae, Drilidae, Lampyridae, Lycidae, Omalisidae, Omethidae, Phengodidae (which includes Telegeusidae), and Rhagophthalmidae.

Torymidae

The Torymidae are a family of wasps that consists of attractive metallic species with enlarged hind legs, and generally with long ovipositors in the females. Many are parasitoids on gall-forming insects, and some are phytophagous (plant-eating) species, sometimes usurping the galls formed by other insects. Over 960 species in about 70 genera are found worldwide. They are best recognized in that they are one of the few groups of Chalcidoidea in which the cerci are visible.

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