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Tachinidae

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98-515: The Tachinidae are a large and variable family of true flies within the insect order Diptera , with more than 8,200 known species and many more to be discovered. Over 1,300 species have been described in North America alone. Insects in this family commonly are called tachinid flies or simply tachinids . As far as is known, they all are protelean parasitoids, or occasionally parasites, of arthropods , usually other insects. The family

196-603: 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 . The bacterial symbionts reproduce and release toxins, which then kill

294-573: 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

392-622: A diagnostically prominent postscutellum bulging beneath the scutellum (a segment of the mesonotum ). They are aristate flies, and the arista usually is bare, though sometimes plumose. The calypters (small flaps above the halteres ) are usually very large. Their fourth long vein bends away sharply. Adult flies feed on flowers and nectar from aphids and scale insects . As many species typically feed on pollen , they can be important pollinators of some plants, especially at higher elevations in mountains where bees are relatively few. The taxonomy of this family presents many difficulties. It

490-456: A length of up to 7 cm (2.8 in), is generally considered to be the largest fly in the world, while the smallest is Euryplatea nanaknihali , which at 0.4 mm (0.016 in) is smaller than a grain of salt. Brachycera are ecologically very diverse, with many being predatory at the larval stage and some being parasitic. Animals parasitised include molluscs , woodlice , millipedes , insects, mammals , and amphibians . Flies are

588-405: A meal. For visual course control, flies' optic flow field is analyzed by a set of motion-sensitive neurons. A subset of these neurons is thought to be involved in using the optic flow to estimate the parameters of self-motion, such as yaw, roll, and sideward translation. Other neurons are thought to be involved in analyzing the content of the visual scene itself, such as separating figures from

686-463: 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 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 lay their eggs on or in

784-401: A mobile head, with a pair of large compound eyes , and mouthparts designed for piercing and sucking (mosquitoes, black flies and robber flies), or for lapping and sucking in the other groups. Their wing arrangement gives them great manoeuvrability in flight, and claws and pads on their feet enable them to cling to smooth surfaces. Flies undergo complete metamorphosis ; the eggs are often laid on

882-805: 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 insects play an important part in limiting the densities of potential pests. Biological control agents such as these 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

980-732: 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. 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 . 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 weeds

1078-468: 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, Novius 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

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1176-487: 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 can control pests not affected by

1274-627: A 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. There are three basic biological pest control strategies: importation (classical biological control), augmentation and conservation. Importation or classical biological control involves

1372-602: 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 are more effective. The O. nubilalis integrated control releasing Tricogramma brassicae (egg parasitoid) and later Bacillus thuringiensis subs. kurstaki (larvicide effect) reduce pest damages more than insecticide treatments The population of Levuana iridescens ,

1470-438: A variety of hosts. The larvae of the shore flies (Ephydridae) and some Chironomidae survive in extreme environments including glaciers ( Diamesa sp., Chironomidae ), hot springs, geysers, saline pools, sulphur pools, septic tanks and even crude oil ( Helaeomyia petrolei ). Adult hoverflies (Syrphidae) are well known for their mimicry and the larvae adopt diverse lifestyles including being inquiline scavengers inside

1568-518: 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

1666-533: A wide spectrum of arthropods. Entomophaga is effective against pests such as the green peach aphid . Several members of Chytridiomycota and Blastocladiomycota have been explored as agents of biological control. From Chytridiomycota, Synchytrium solstitiale is being considered as a control agent of the yellow star thistle ( Centaurea solstitialis ) in the United States. Baculoviruses are specific to individual insect host species and have been shown to be useful in biological pest control. For example,

1764-485: Is Spilomyia longicornis , which is a fly but mimics a vespid wasp. Flies have a mobile head with a pair of large compound eyes on the sides of the head, and in most species, three small ocelli on the top. The compound eyes may be close together or widely separated, and in some instances are divided into a dorsal region and a ventral region, perhaps to assist in swarming behaviour. The antennae are well-developed but variable, being thread-like, feathery or comb-like in

1862-484: Is a method of controlling pests , whether pest animals such as insects and mites , weeds , or pathogens affecting animals or plants by using other organisms . 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 control: classical (importation), where

1960-461: Is a small parasitoid wasp attacking whiteflies , 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 : Mymaridae ) has been introduced to control

2058-447: 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 , 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

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2156-677: 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 . Pathogenic fungi may be controlled by other fungi, or bacteria or yeasts, such as: Gliocladium spp., mycoparasitic Pythium spp., binucleate types of Rhizoctonia spp., and Laetisaria spp. The fungi Cordyceps and Metacordyceps are deployed against

2254-542: 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

2352-480: 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 (the vedalia beetle). This success

2450-493: Is known from many habitats in all zoogeographical regions and is especially diverse in South America. Reproductive strategies vary greatly between tachinid species, largely, but not always clearly, according to their respective life cycles. Many species are generalists rather than specialists. Comparatively few are restricted to a single host species, so there is little tendency towards the close co-evolution one finds in

2548-589: Is largely based on morphological characters of the adult flies, but also on reproductive habits and on the immature stage. Some tachinid flies parasitize pest species. This has allowed them to be used as biological control agents by farmers. Some Tachinidae are generalists; for instance, Compsilura concinnata uses, at least, 200 different hosts, and thus are less safe to be used as biological controls because they will attack non-pest species, resulting in population decline. Others are more specialized and are safer; for instance, Istocheta aldrichi , which only attacks

2646-547: Is one of the major insect orders and of considerable ecological and human importance. Flies are major pollinators, second only to the bees and their Hymenopteran relatives. Flies may have been among the evolutionarily earliest pollinators responsible for early plant pollination . Fruit flies are used as model organisms in research, but less benignly, mosquitoes are vectors for malaria , dengue , West Nile fever , yellow fever , encephalitis , and other infectious diseases ; and houseflies , commensal with humans all over

2744-524: Is widely thought to be a member of Mecopterida , along with Lepidoptera (butterflies and moths), Trichoptera (caddisflies), Siphonaptera (fleas), Mecoptera (scorpionflies) and possibly Strepsiptera (twisted-wing flies). Diptera has been grouped with Siphonaptera and Mecoptera in the Antliophora, but this has not been confirmed by molecular studies. Diptera were traditionally broken down into two suborders, Nematocera and Brachycera , distinguished by

2842-532: 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. 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

2940-810: The Japanese beetle . This clade appears to have originated in the middle Eocene . The oldest known putatively tachinid fossil ( Lithexorista ) dates from the Eocene Green River Formation in Wyoming . Fly Housefly (Muscidae) (top left) Haematopota pluvialis (Tabanidae) (top right) Ctenophora pectinicornis (Tipulidae) (mid left) Ochlerotatus notoscriptus (Culicidae) (mid right) Milesia crabroniformis (Syrphidae) (bottom left) Holcocephala fusca (Asilidae) (bottom right) Nematocera ( paraphyletic ) (inc Eudiptera ) Brachycera Flies are insects of

3038-631: The Jurassic , some 180 million years ago. A third radiation took place among the Schizophora at the start of the Paleogene , 66 million years ago. The phylogenetic position of Diptera has been controversial. The monophyly of holometabolous insects has long been accepted, with the main orders being established as Lepidoptera, Coleoptera, Hymenoptera and Diptera, and it is the relationships between these groups which has caused difficulties. Diptera

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3136-526: The Lymantria dispar multicapsid nuclear polyhedrosis virus has been used to spray large areas of forest in North America where larvae of the spongy 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. A mammalian virus, the rabbit haemorrhagic disease virus

3234-1071: The Mecopterida , alongside the Mecoptera , Siphonaptera , Lepidoptera and Trichoptera . The possession of a single pair of wings distinguishes most true flies from other insects with "fly" in their names. However, some true flies such as Hippoboscidae (louse flies) have become secondarily wingless. The cladogram represents the current consensus view. Hymenoptera (sawflies, wasps, ants, bees) [REDACTED] Raphidioptera (snakeflies) [REDACTED] Megaloptera (alderflies and allies) [REDACTED] Neuroptera (Lacewings and allies) [REDACTED] Coleoptera (beetles) [REDACTED] Strepsiptera (twisted-wing parasites) [REDACTED] Trichoptera (caddisflies) [REDACTED] Lepidoptera (butterflies, moths) [REDACTED] Diptera [REDACTED] Mecoptera (scorpionflies) [REDACTED] Siphonaptera (fleas) [REDACTED] The first true dipterans known are from

3332-666: The Middle Triassic (around 240 million years ago), and they became widespread during the Middle and Late Triassic . Modern flowering plants did not appear until the Cretaceous (around 140 million years ago), so the original dipterans must have had a different source of nutrition other than nectar . Based on the attraction of many modern fly groups to shiny droplets, it has been suggested that they may have fed on honeydew produced by sap-sucking bugs which were abundant at

3430-651: The Strepsiptera . In contrast to the flies, the Strepsiptera bear their halteres on the mesothorax and their flight wings on the metathorax. Each of the fly's six legs has a typical insect structure of coxa, trochanter, femur, tibia and tarsus, with the tarsus in most instances being subdivided into five tarsomeres . At the tip of the limb is a pair of claws, and between these are cushion-like structures known as pulvilli which provide adhesion. Biological pest control Biological control or biocontrol

3528-596: The glassy-winged sharpshooter Homalodisca vitripennis (Hemiptera: Cicadellidae ) in French Polynesia and has successfully controlled ~95% of the pest density. 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. There are

3626-570: 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 mechanosensory organs known as halteres , which act as high-speed sensors of rotational movement and allow dipterans to perform advanced aerobatics. Diptera is a large order containing more than 150,000 species including horse-flies , crane flies , hoverflies , mosquitoes and others. Flies have

3724-466: The sericulture industry by attacking silkworm larvae. One particularly notorious silkworm pest is the Uzi fly ( Exorista bombycis ). Another reproductive strategy is to leave the eggs in the host's environment; for example, the female might lay on leaves, where the host is likely to ingest them. Some tachinids that are parasitoids of stem-boring caterpillars deposit eggs outside the host's burrow, letting

3822-548: The thorax , bears the wings and contains the flight muscles on the second segment, which is greatly enlarged; the first and third segments have been reduced to collar-like structures, and the third segment bears the halteres , which help to balance the insect during flight. The third tagma is the abdomen consisting of 11 segments, some of which may be fused, and with the 3 hindmost segments modified for reproduction. Some Dipterans are mimics and can only be distinguished from their models by very careful inspection. An example of this

3920-827: The Brachycera includes broader, more robust flies with short antennae. Many nematoceran larvae are aquatic. There are estimated to be a total of about 19,000 species of Diptera in Europe, 22,000 in the Nearctic region, 20,000 in the Afrotropical region, 23,000 in the Oriental region and 19,000 in the Australasian region. While most species have restricted distributions, a few like the housefly ( Musca domestica ) are cosmopolitan. Gauromydas heros ( Asiloidea ), with

4018-463: The Colombian cities of Bello , Medellín , and Itagüí . The project was executed by non-profit World Mosquito Program (WMP). Wolbachia prevents mosquitos from transmitting viruses such as dengue and zika . The insects pass the bacteria on to their offspring. The project covered a combined area of 135 square kilometres (52 sq mi), home to 3.3 million people. Most of the project area reached

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4116-509: The Levuana moth, a serious coconut pest in Fiji , was brought under control by a classical biological control program in the 1920s. 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

4214-1403: The Nematocera. The construction of a phylogenetic tree has been the subject of ongoing research. The following cladogram is based on the FLYTREE project. Ptychopteromorpha (phantom and primitive crane-flies) [REDACTED] Culicomorpha (mosquitoes, blackflies and midges) [REDACTED] Blephariceromorpha (net-winged midges, etc) [REDACTED] Bibionomorpha (gnats) [REDACTED] Psychodomorpha (drain flies, sand flies, etc) [REDACTED] Tipulomorpha (crane flies) [REDACTED] Stratiomyomorpha (soldier flies, etc) [REDACTED] Xylophagomorpha (stink flies, etc) [REDACTED] Tabanomorpha (horse flies, snipe flies, etc) [REDACTED] Nemestrinoidea [REDACTED] Asiloidea (robber flies, bee flies, etc) [REDACTED] Empidoidea (dance flies, etc) [REDACTED] Aschiza (in part) Phoroidea (flat-footed flies, etc) [REDACTED] Syrphoidea (hoverflies) [REDACTED] Hippoboscoidea (louse flies, etc) [REDACTED] Muscoidea (house flies, dung flies, etc) [REDACTED] Oestroidea (blow flies, flesh flies, etc) [REDACTED] Acalyptratae (marsh flies, etc) [REDACTED] Flies are often abundant and are found in almost all terrestrial habitats in

4312-677: The above mechanisms, especially when a species is introduced without a thorough understanding of the possible consequences. 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,

4410-467: The adaptations of many specialist species to their hosts, such as are typical of protelean parasitoids among the Hymenoptera . Larvae ( maggots ) of most members of this family are parasitoids (developing inside a living host, ultimately killing it). In contrast, a few are parasitic (not generally killing the host). Tachinid larvae feed on the host tissues, either after having been injected into

4508-462: 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

4606-519: The antennae and the maxillary palps bear the main olfactory receptors, while the gustatory receptors are in the labium, pharynx, feet, wing margins and female genitalia, enabling flies to taste their food by walking on it. The taste receptors in females at the tip of the abdomen receive information on the suitability of a site for ovipositing. Flies that feed on blood have special sensory structures that can detect infrared emissions, and use them to home in on their hosts, Many blood-sucking flies can detect

4704-562: 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

4802-671: 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 or wood wool . Green lacewings can be encouraged by using plastic bottles with an open bottom and

4900-509: 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; chalcidoid wasps , which parasitize eggs and larvae of many insect species; and tachinid flies , which parasitize

4998-750: 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

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5096-482: 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

5194-403: 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

5292-453: 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 , or beetle bank where beneficial insects such as parasitoidal wasps can live and reproduce, can help ensure

5390-595: The decades since, the same biocontrol methods that are routine on land have become common in the water. 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 , 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

5488-404: The differences in antennae. The Nematocera are identified by their elongated bodies and many-segmented, often feathery antennae as represented by mosquitoes and crane flies. The Brachycera have rounder bodies and much shorter antennae. Subsequent studies have identified the Nematocera as being non-monophyletic with modern phylogenies placing the Brachycera within grades of groups formerly placed in

5586-431: The different families. The mouthparts are adapted for piercing and sucking, as in the black flies, mosquitoes and robber flies, and for lapping and sucking as in many other groups. Female horse-flies use knife-like mandibles and maxillae to make a cross-shaped incision in the host's skin and then lap up the blood that flows. The gut includes large diverticulae , allowing the insect to store small quantities of liquid after

5684-431: 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 and other infectious diseases, biological control was attempted by a community action plan; copepods , baby turtles , and juvenile tilapia were added to

5782-467: The first instar larvae do the work of finding the host for themselves. In other species, the maggots use an ambush technique, waiting for the host to pass and then attacking it and burrowing into its body. Adult tachinids are not parasitic, but either do not feed at all or visit flowers, decaying matter, or similar sources of energy to sustain themselves until they have concluded their procreative activities. Their non-parasitic behaviour after eclosion from

5880-749: The flies, principally in the family Cecidomyiidae (gall midges). Many flies (most importantly in the family Agromyzidae) lay their eggs in the mesophyll tissue of leaves with larvae feeding between the surfaces forming blisters and mines. Some families are mycophagous or fungus feeding. These include the cave dwelling Mycetophilidae (fungus gnats) whose larvae are the only diptera with bioluminescence. The Sciaridae are also fungus feeders. Some plants are pollinated by fungus feeding flies that visit fungus infected male flowers. The larvae of Megaselia scalaris (Phoridae) are almost omnivorous and consume such substances as paint and shoe polish. The Exorista mella (Walker) fly are considered generalists and parasitoids of

5978-647: The genera Gymnosoma , Thrixion , Winthemia , and Eutachina as examples. In a closely related strategy some genera are effectively ovoviviparous (some authorities prefer the term ovolarviparous ) and deposit a hatching larva onto the host. For example, this occurs in Tachinidae species which parasitize the butterfly Danaus chrysippus in Ghana. The free larvae immediately bore into the host's body. Illustrative genera include Exorista and Voria . Many tachinid eggs hatch quickly, having partly developed inside

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6076-483: The genera Ocyptera , Alophora , and Compsilura are examples. In many species only one egg is laid on or in any individual host, and accordingly such an egg tends to be large, as is typical for eggs laid in small numbers. They are large enough to be clearly visible if stuck onto the outside of the host, and they generally are so firmly stuck that eggs cannot be removed from the skin of the host without killing them. Furthermore, scientists have observed in studies with

6174-527: The ground using motion parallax. The H1 neuron is responsible for detecting horizontal motion across the entire visual field of the fly, allowing the fly to generate and guide stabilizing motor corrections midflight with respect to yaw. The ocelli are concerned in the detection of changes in light intensity, enabling the fly to react swiftly to the approach of an object. Like other insects, flies have chemoreceptors that detect smell and taste, and mechanoreceptors that respond to touch. The third segments of

6272-400: 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

6370-788: The host cabbage looper that being glued to the host insect helps maggots burrow into the larva, where they remain until fully developed. Yet another strategy of oviposition among some Tachinidae is to lay large numbers of small, darkly coloured eggs on the food plants of the host species. Sturmia , Zenillia , and Gonia are such genera. Many tachinids are important natural enemies of major insect pests, and some species actually are used in biological pest control ; for example, some species of tachinid flies have been introduced into North America from their native lands as biocontrols to suppress populations of alien pests. Conversely, certain tachinid flies that prey on useful insects are themselves considered as pests; they can present troublesome problems in

6468-485: The host by the parent, or penetrating the host from outside. Various species have different modes of oviposition and of host invasion. Typically, tachinid larvae are endoparasites (internal parasites) of caterpillars of butterflies and moths, or the eruciform larvae of sawflies . For example, they have been found to lay eggs in African sugarcane borer larva, a species of moth common in sub-Saharan Africa, as well as

6566-403: The host insect. 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

6664-414: The interactions between each pest and control agent. 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 , while the predatory mite Phytoseiulus persimilis is used for control of the two-spotted spider mite. The egg parasite Trichogramma

6762-409: 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

6860-431: The larval food-source and the larvae, which lack true limbs, develop in a protected environment, often inside their food source. Other species are ovoviviparous , opportunistically depositing hatched or hatching larvae instead of eggs on carrion , dung, decaying material, or open wounds of mammals. The pupa is a tough capsule from which the adult emerges when ready to do so; flies mostly have short lives as adults. Diptera

6958-511: 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. 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

7056-413: The more northerly Arctic woolly bear moth . However, some species attack adult beetles and some attack beetle larvae. Others attack various types of true bugs , and others attack grasshoppers ; a few even attack centipedes . Also parasitised are bees, wasps and sawflies. Probably the majority of female tachinids lay white, ovoid eggs with flat undersides onto the skin of the host insect. Imms mentions

7154-563: 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 , cats 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

7252-403: The mother's uterus, which is long and often coiled for retaining developing eggs. However, it is suggested that the primitive state probably is to stick unembryonated eggs to the surface of the host. Many other species inject eggs into the host's body, using the extensible, penetrating part of their ovipositor , sometimes called the oviscapt, which roughly translates to "egg digger". Species in

7350-419: The nests of social insects. Some brachycerans are agricultural pests, some bite animals and humans and suck their blood, and some transmit diseases. Flies are adapted for aerial movement and typically have short and streamlined bodies. The first tagma of the fly, the head, bears the eyes, the antennae , and the mouthparts (the labrum, labium, mandible, and maxilla make up the mouthparts). The second tagma,

7448-406: The neural ganglia , and concentration of nerve tissue in the thorax, a feature that is most extreme in the highly derived Muscomorpha infraorder. Some flies such as the ectoparasitic Nycteribiidae and Streblidae are exceptional in having lost their wings and become flightless. The only other order of insects bearing a single pair of true, functional wings, in addition to any form of halteres, are

7546-530: 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. 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

7644-559: The population of weevils in the alfalfa area treated for alfalfa weevil in the Northeastern United States 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

7742-494: 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,

7840-773: 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 ). 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

7938-526: The predatory midge Feltiella acarisuga , and a ladybird Stethorus punctillum . 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,

8036-431: The pupa is what justifies the application of the term "protelean". Tachinid flies are extremely varied in appearance. Some adult flies may be brilliantly colored and resemble blow-flies (family Calliphoridae ). Most however are rather drab, some resembling house flies . However, tachinid flies commonly are more bristly and more robust. Also, they usually have a characteristic appearance. They have three-segmented antennae,

8134-405: The raised concentration of carbon dioxide that occurs near large animals. Some tachinid flies (Ormiinae) which are parasitoids of bush crickets , have sound receptors to help them locate their singing hosts. Diptera have one pair of fore wings on the mesothorax and a pair of halteres , or reduced hind wings, on the metathorax . A further adaptation for flight is the reduction in number of

8232-568: 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 ), in the Book of Tree Planting by Yu Zhen Mu ( Ming dynasty ), in

8330-568: The second largest group of pollinators after the Hymenoptera (bees, wasps and relatives). In wet and colder environments flies are significantly more important as pollinators. Compared to bees, they need less food as they do not need to provision their young. Many flowers that bear low nectar and those that have evolved trap pollination depend on flies. It is thought that some of the earliest pollinators of plants may have been flies. The greatest diversity of gall forming insects are found among

8428-469: 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 cucumeris ,

8526-696: 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. 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,

8624-466: 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

8722-579: The target of infecting 60% of local mosquitoes. The technique is not endorsed by WHO. Entomopathogenic fungi , which cause disease in insects, include at least 14 species that attack aphids . Beauveria bassiana is mass-produced and used to manage a wide variety of insect pests including whiteflies , thrips , aphids and weevils . 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

8820-606: The time, and dipteran mouthparts are well-adapted to softening and lapping up the crusted residues. The basal clades in the Diptera include the Deuterophlebiidae and the enigmatic Nymphomyiidae . Three episodes of evolutionary radiation are thought to have occurred based on the fossil record. Many new species of lower Diptera developed in the Triassic , about 220 million years ago. Many lower Brachycera appeared in

8918-578: 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., fluorescent Pseudomonads , and Streptomycetes are controls of various fungal pathogens. The largest-ever deployment of Wolbachia -infected A. aegypti mosquitoes reduced dengue incidence by 94–97% in

9016-491: 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 food sources ( nectar and to a lesser degree pollen ) 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

9114-473: The world apart from Antarctica. They include many familiar insects such as house flies, blow flies, mosquitoes, gnats, black flies, midges and fruit flies. More than 150,000 have been formally described and the actual species diversity is much greater, with the flies from many parts of the world yet to be studied intensively. The suborder Nematocera include generally small, slender insects with long antennae such as mosquitoes, gnats, midges and crane-flies, while

9212-607: The world, spread foodborne illnesses . Flies can be annoyances especially in some parts of the world where they can occur in large numbers, buzzing and settling on the skin or eyes to bite or seek fluids. Larger flies such as tsetse flies and screwworms cause significant economic harm to cattle. Blowfly larvae, known as gentles , and other dipteran larvae, known more generally as maggots , are used as fishing bait , as food for carnivorous animals, and in medicine in debridement , to clean wounds . Dipterans are holometabolans , insects that undergo radical metamorphosis. They belong to

9310-400: Was introduced to Australia to attempt to control the European rabbit populations there. 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. Introduction into New Zealand in the 1990s was similarly successful at first, but

9408-514: 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 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

9506-460: Was repeated in California using the beetle and a parasitoidal fly, Cryptochaetum 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

9604-445: 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

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