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o start a new colony take an aphid egg to found a new herd of underground aphids in the new colony these farming ants protect the aphids by fighting off aphid predators 70 some bees in coniferous forests collect aphid honeydew to make forest honey 29 an interesting variation in ant aphid relationships involves lycaenid butterflies and myrmica ants for example niphanda fusca butterflies lay eggs on plants where ants tend herds of aphids the eggs hatch as caterpillars which feed on the aphids the ants do not defend the aphids from the caterpillars since the caterpillars produce a pheromone which deceives the ants into treating them like ants and carrying the caterpillars into their nest once there the ants feed the caterpillars which in return produce honeydew for the ants when the caterpillars reach full size they crawl to the colony entrance and form cocoons after two weeks the adult butterflies emerge and take flight at this point the ants attack the butterflies but the butterflies have a sticky wool like substance on their wings that disables the ants jaws allowing the butterflies to fly away without being harmed 74 another ant mimicking gall aphid paracletus cimiciformis eriosomatinae has evolved a complex double strategy involving two morphs of the same clone and tetramorium ants aphids of the round morph cause the ants to farm them as with many other aphids the flat morph aphids are aggressive mimics with a wolf in sheep s clothing strategy they have hydrocarbons in their cuticle that mimic those of the ants and the ants carry them into the brood chamber of the ants nest and raise them like ant larvae once there the flat morph aphids behave like predators drinking the body fluids of ant larvae 75 an ant guards its aphids ants tending aphids ant extracting honeydew from an aphid ants tending aphids and collecting honeydew secreted a wrinkled solder beetle flies in and eats an aphid before being chased away by the ants bacterial endosymbiosis edit endosymbiosis with micro organisms is common in insects with more than 10 of insect species relying on intracellular bacteria for their development and survival 76 aphids harbour a vertically transmitted from parent to its offspring obligate symbiosis with buchnera aphidicola the primary symbiont inside specialized cells the bacteriocytes 77 five of the bacteria genes have been transferred to the aphid nucleus 78 the original association is estimated to have occurred in a common ancestor 280 to 160 million years ago and enabled aphids to exploit a new ecological niche feeding on phloem sap of vascular plants b aphidicola provides its host with essential amino acids which are present in low concentrations in plant sap 79 the metabolites from endosymbionts are also excreted in honeydew 80 the stable intracellular conditions as well as the bottleneck effect experienced during the transmission of a few bacteria from the mother to each nymph increase the probability of transmission of mutations and gene deletions 81 82 as a result the size of the b aphidicola genome is greatly reduced compared to its putative ancestor 83 despite the apparent loss of transcription factors in the reduced genome gene expression is highly regulated as shown by the ten fold variation in expression levels between different genes under normal conditions 84 buchnera aphidicola gene transcription although not well understood is thought to be regulated by a small number of global transcriptional regulators and or through nutrient supplies from the aphid host 85 some aphid colonies also harbour secondary or facultative optional extra bacterial symbionts these are vertically transmitted and sometimes also horizontally from one lineage to another and possibly from one species to another 86 87 so far the role of only some of the secondary symbionts has been described regiella insecticola plays a role in defining the host plant range 88 89 hamiltonella defensa provides resistance to parasitoids but only when it is in turn infected by the bacteriophage apse 90 91 and serratia symbiotica prevents the deleterious effects of heat 92 predators edit a western polished lady beetle and her eggs near a milkweed aphid colony aphids are eaten by many bird and insect predators in a study on a farm in north carolina six species of passerine bird consumed nearly a million aphids per day between them the top predators being the american goldfinch with aphids forming 83 of its diet and the vesper sparrow 93 insects that attack aphids include the adults and larvae of predatory ladybirds hoverfly larvae parasitic wasps aphid midge larvae aphid lions the larvae of green lacewings and arachnids such as spiders among ladybirds myzia oblongoguttata is a dietary specialist which only feeds on conifer aphids whereas adalia bipunctata and coccinella septempunctata are generalists feeding on large numbers of species the eggs are laid in batches each female laying several hundred female hoverflies lay several thousand eggs the adults feed on pollen and nectar but the larvae feed voraciously on aphids eupeodes corollae adjusts the number of eggs laid to the size of the aphid colony 94 predators eating aphids ladybird larva hoverfly larva the ladybird beetle propylea quatuordecimpunctata lady bug on whorled milkweed aphids are often infected by bacteria viruses and fungi they are affected by the weather such as precipitation 95 temperature 96 and wind 97 fungi that attack aphids include neozygites fresenii entomophthora beauveria bassiana metarhizium anisopliae and entomopathogenic fungi such as lecanicillium lecanii aphids brush against the microscopic spores these stick to the aphid germinate and penetrate the aphid s skin the fungus grows in the aphid s hemolymph after about three days the aphid dies and the fungus releases more spores into the air infected aphids are covered with a woolly mass that progressively grows thicker until the aphid is obscured often the visible fungus is not the one that killed the aphid but a secondary infection 95 aphids can be easily killed by unfavourable weather such as late spring freezes 98 excessive heat kills the symbiotic bacteria that some aphids depend on which makes the aphids infertile 99 rain prevents winged aphids from dispersing and knocks aphids off plants and thus kills them from the impact or by starvation 95 100 101 but cannot be relied on for aphid control 102 anti predator defences edit aphid secreting defensive fluid from the cornicles a milkweed aphid on narrow leaf milkweed is attacked by a hoverfly larvae it thrashs and release pheromones and sticky wax from its cornicles nearby aphids flee video played at 4x speed most aphids have little protection from predators some species interact with plant tissues forming a gall an abnormal swelling of plant tissue aphids can live inside the gall which provides protection from predators and the elements a number of galling aphid species are known to produce specialised soldier forms sterile nymphs with defensive features which defend the gall from invasion 29 103 104 for example alexander s horned aphids are a type of soldier aphid that has a hard exoskeleton and pincer like mouthparts 74 144 a woolly aphid colophina clematis has first instar soldier nymphs that protect the aphid colony killing larvae of ladybirds hoverflies and the flower bug anthocoris nemoralis by climbing on them and inserting their stylets 105 although aphids cannot fly for most of their life cycle they can escape predators and accidental ingestion by herbivores by dropping off the plant onto the ground 106 others species use the soil as a permanent protection feeding on the vascular systems of roots and remaining underground all their lives they are often attended by ants for the honeydew they produce and are carried from plant to plant by the ants through their tunnels 93 some species of aphid known as woolly aphids eriosomatinae excrete a fluffy wax coating for protection 29 the cabbage aphid brevicoryne brassicae sequesters secondary metabolites from its host stores them and releases chemicals that produce a violent chemical reaction and strong mustard oil smell to repel predators 107 peptides produced by aphids thaumatins are thought to provide them with resistance to some fungi 108 it was common at one time to suggest that the cornicles were the source of the honeydew and this was even included in the shorter oxford english dictionary 109 and the 2008 edition of the world book encyclopedia 110 in fact honeydew secretions are produced from the anus of the aphid 111 while cornicles mostly produce defensive chemicals such as waxes there also is evidence of cornicle wax attracting aphid predators in some cases 112 some clones of aphis craccivora are sufficiently toxic to the invasive and dominant predatory ladybird harmonia axyridis to suppress it locally favouring other ladybird species the toxicity is in this case narrowly specific to the dominant predator species 113 parasitoids edit further information parasitoid a milkweed aphid pushes the sticky wax drop from its cornicle against an attacking parasitic wasp extruding another drop two scenes at one tenth speed a different aphid has captured a wasp aphids are abundant and widespread and serve as hosts to a large number of parasitoids many of them being very small c 0 1 inches 2 5 mm long parasitoid wasps 114 one species aphis ruborum for example is host to at least 12 species of parasitoid wasps 115 parasitoids have been investigated intensively as biological control agents and many are used commercially for this purpose 116 plant aphid interactions edit aphids on plant host plants mount local and systemic defenses against aphid attack young leaves in some plants contain chemicals that discourage attack while the older leaves have lost this resistance while in other plant species resistance is acquired by older tissues and the young shoots are vulnerable volatile products from interplanted onions have been shown to prevent aphid attack on adjacent potato plants by encouraging the production of terpenoids a benefit exploited in the traditional practice of companion planting while plants neighboring infested plants showed increased root growth at the expense of the extension of aerial parts 32 the wild potato solanum berthaultii produces an aphid alarm pheromone e β farnesene as an allomone a pheromone to ward off attack it effectively repels the aphid myzus persicae at a range of up to 3 millimetres 117 s berthaultii and other wild potato species have a further anti aphid defence in the form of glandular hairs which when broken by aphids discharge a sticky liquid that can immobilise some 30 of the aphids infesting a plant 118 plants exhibiting aphid damage can have a variety of symptoms such as decreased growth rates mottled leaves yellowing stunted growth curled leaves browning wilting low yields and death the removal of sap creates a lack of vigor in the plant and aphid saliva is toxic to plants aphids frequently transmit plant viruses to their hosts such as to potatoes cereals sugarbeets and citrus plants 29 there are two types of virus transmission between plant aphid interactions non circulative transmission and circulative transmission in non circulative transmission the virus attaches itself to the aphids mouthparts and is released when the aphids feed on a different plant these non circulatory transmitted viruses promotes rapid dispersion of the vector or aphids in circulative transmission the virus is ingested and passes through the gut lining to enter the hemolymph where it is then circulated throughout the entire body after reaching the salivary glands the virus is then released into the saliva upon transmission sites in plants circulatory transmitted viruses allows for long term feeding by the aphids and increases the chances of being infected with the virus 119 the green peach aphid myzus persicae is a vector for more than 110 plant viruses cotton aphids aphis gossypii often infect sugarcane papaya and peanuts with viruses 21 in plants which produce the phytoestrogen coumestrol such as alfalfa damage by aphids is linked with higher concentrations of coumestrol 120 aphid with honeydew from the anus not the cornicles the coating of plants with honeydew can contribute to the spread of fungi which can damage plants 121 122 honeydew produced by aphids has been observed to reduce the effectiveness of fungicides as well 123 a hypothesis that insect feeding may improve plant fitness was floated in the mid 1970s by owen and wiegert it was felt that the excess honeydew would nourish soil micro organisms including nitrogen fixers in a nitrogen poor environment this could provide an advantage to an infested plant over an uninfested plant however this does not appear to be supported by observational evidence 124 sociality edit some aphids show some of the traits of eusociality joining insects such as ants bees and termites however there are differences between these sexual social insects and the clonal aphids which are all descended from a single female parthenogenetically and share an identical genome about fifty species of aphid scattered among the closely related host alternating lineages eriosomatinae and hormaphidinae have some type of defensive morph these are gall creating species with the colony living and feeding inside a gall that they form in the host s tissues among the clonal population of these aphids there may be several distinct morphs and this lays the foundation for a possible specialization of function in this case a defensive caste the soldier morphs are mostly first and second instars with the third instar being involved in eriosoma moriokense and only in smythurodes betae are adult soldiers known the hind legs of soldiers are clawed heavily sclerotized and the stylets are robust making it possible to rupture and crush small predators 125 the larval soldiers are altruistic individuals unable to advance to breeding adults but acting permanently in the interests of the colony another requirement for the development of sociality is provided by the gall a colonial home to be defended by the soldiers 126 the soldiers of gall forming aphids also carry out the job of cleaning the gall the honeydew secreted by the aphids is coated in a powdery wax to form liquid marbles 127 that the soldiers roll out of the gall through small orifices 104 aphids that form closed galls use the plant s vascular system for their plumbing the inner surfaces of the galls are highly absorbent and wastes are absorbed and carried away by the plant 104 interactions with humans edit pest status edit about 5000 species of aphid have been described and of these some 450 species have colonized food and fiber crops as direct feeders on plant sap they damage crops and reduce yields but they have a greater impact by being vectors of plant viruses the transmission of these viruses depends on the movements of aphids between dif...
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