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ters and granddaughters 12 59 60 aphid giving birth to live young populations are often entirely female this process repeats itself throughout the summer producing multiple generations that typically live 20 to 40 days for example some species of cabbage aphids like brevicoryne brassicae can produce up to 41 generations of females in a season thus one female hatched in spring can theoretically produce billions of descendants were they all to survive 61 in autumn aphids reproduce sexually and lay eggs environmental factors such as a change in photoperiod and temperature or perhaps a lower food quantity or quality causes females to parthenogenetically produce sexual females and males 58 the males are genetically identical to their mothers except that with the aphids x0 sex determination system they have one fewer sex chromosome 58 these sexual aphids may lack wings or even mouthparts 21 sexual females and males mate and females lay eggs that develop outside the mother the eggs survive the winter and hatch into winged alate or wingless females the following spring this occurs in for example the life cycle of the rose aphid macrosiphum rosae which may be considered typical of the family however in warm environments such as in the tropics or a greenhouse aphids may go on reproducing asexually for many years 29 the life stages of the green apple aphid aphis pomi drawing by robert evans snodgrass 1930 aphids reproducing asexually by parthenogenesis can have genetically identical winged and non winged female progeny control is complex some aphids alternate during their life cycles between genetic control polymorphism and environmental control polyphenism of production of winged or wingless forms 62 winged progeny tend to be produced more abundantly under unfavorable or stressful conditions some species produce winged progeny in response to low food quality or quantity e g when a host plant is starting to senesce 63 the winged females migrate to start new colonies on a new host plant for example the apple aphid aphis pomi after producing many generations of wingless females gives rise to winged forms that fly to other branches or trees of its typical food plant 64 aphids that are attacked by ladybugs lacewings parasitoid wasps or other predators can change the dynamics of their progeny production when aphids are attacked by these predators alarm pheromones in particular beta farnesene are released from the cornicles these alarm pheromones cause several behavioral modifications that depending on the aphid species can include walking away and dropping off the host plant additionally alarm pheromone perception can induce the aphids to produce winged progeny that can leave the host plant in search of a safer feeding site 65 viral infections which can be extremely harmful to aphids can also lead to the production of winged offspring 66 for example densovirus infection has a negative impact on dysaphis plantaginea rosy apple aphid reproduction but contributes to the development of aphids with wings which can transmit the virus more easily to new host plants 67 additionally symbiotic bacteria that live inside of the aphids can also alter aphid reproductive strategies based on the exposure to environmental stressors 68 in the autumn host alternating heteroecious aphid species produce a special winged generation that flies to different host plants for the sexual part of the life cycle flightless female and male sexual forms are produced and lay eggs 69 some species such as aphis fabae black bean aphid metopolophium dirhodum rose grain aphid myzus persicae peach potato aphid and rhopalosiphum padi bird cherry oat aphid are serious pests they overwinter on primary hosts on trees or bushes in summer they migrate to their secondary host on a herbaceous plant often a crop then the gynoparae return to the tree in autumn another example is the soybean aphid aphis glycines as fall approaches the soybean plants begin to senesce from the bottom upwards the aphids are forced upwards and start to produce winged forms first females and later males which fly off to the primary host buckthorn here they mate and overwinter as eggs 52 ecology edit ant mutualism edit some species of ants farm aphids protecting them on the plants where they are feeding and consuming the honeydew the aphids release from the terminations of their alimentary canals this is a mutualistic relationship with these dairying ants milking the aphids by stroking them with their antennae c 70 although symbiotic the feeding behaviour of aphids is altered by ant attendance aphids attended by ants tend to increase the production of honeydew in smaller drops with a greater concentration of amino acids 71 macrosiphoniella yomogicola has green and red morphs in japan where lasius japonicus ants prefer the green morph which provides more honeydew and select for them their honeydew contains dopamine and makes the attendant ants act more aggressively against intruders the aphids are thought to be selfishly manipulating the ants to provide them better protection from predators and parasites 72 some farming ant species gather and store the aphid eggs in their nests over the winter in the spring the ants carry the newly hatched aphids back to the plants some species of dairying ants such as the european yellow meadow ant lasius flavus 73 manage large herds of aphids that feed on roots of plants in the ant colony queens leaving to 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 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