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14-September-2008 18:38:39 - Aphid This article is about the insect. For other uses, see Aphid disambiguation. Aphids Scientific classification Kingdom: Animalia Phylum: Arthropoda Class: Insecta Order: Hemiptera Suborder: Sternorrhyncha Superfamily: Aphidoidea Families There are 10 families: Anoeciidae Aphididae Drepanosiphidae Greenideidae Hormaphididae Lachnidae Mindaridae Pemphigidae Phloeomyzidae Thelaxidae Aphids, also known as plant lice,1 are small plant-eating insects, and members of the superfamily Aphidoidea.2 Aphids are the among the most destructive insect pests on Earth.3 About 4,400 species of 10 families are known. Historically, many fewer families were recognized, as most species were included in the family Aphididae. Around 250 species are serious pests for agriculture and forestry as well as an annoyance for gardeners. They vary in length from one to ten millimetres. Natural enemies include predatory lady beetles Coleoptera: Coccinellidae, hoverfly larvae Diptera: Syrphidae, parasitic wasps, aphid midge larvae, aphid lions, crab spiders4 lacewings Neuroptera: Chrysopidae, and entomopathogenic fungi like Lecanicillium lecanii and the Entomophthorales. Aphids are distributed world-wide, but are most common in temperate zones. Also, in contrast to many taxa, species diversity is much lower in the tropics than in the temperate zones. They can migrate great distances, mainly through passive dispersal by riding on winds. For example, the currant lettuce aphid Nasonovia ribisnigri Mosley is believed to have spread from New Zealand to Tasmania in this way.5 Aphids have also been spread by human transportation of infested plant materials. Contents 1 Taxonomy 1.1 Relation to phylloxera and adelgids 2 Anatomy 3 Diet 4 Symbioses 5 Reproduction 6 Evolution 7 Threats 8 Defenses 9 Effects on plants 10 In popular culture 11 Gallery 12 See also 13 References 14 External links Taxonomy Aphids are in the superfamily Aphidoidea in the homopterous division of the order Hemiptera. Recent classification within Hemiptera has reduced the old taxon Homoptera to two suborders: Sternorrhyncha e.g., aphids, whiteflies, scales, psyllids, etc. and Auchenorrhyncha e.g., cicadas, leafhoppers, treehoppers, planthoppers, etc. with the suborder Heteroptera containing a large group of insects known as the true bugs. More recent reclassifications have substantially rearranged the families within Aphidoidea: some old families were reduced to subfamily rank e.g., Eriosomatidae, and many old subfamilies elevated to family rank. Furthermore, woolly conifer aphids like the pine aphid, the spruce aphid and the balsam woolly aphid are not aphids, but adelgids. Relation to phylloxera and adelgids Aphids, adelgids, and phylloxerids are very closely related, and are either placed in the insect super family Aphidoidea Blackman and Eastop, 1994 or into two super families Phylloxeroidea and Aphidoidea within the order Homoptera, the plant-sucking bugs.6 Like aphids, phylloxera feed on the roots, leaves and shoots of grape plants, but unlike aphids do not produce honeydew or cornicle secretions.7 Phylloxera Daktulosphaira vitifoliae, are insects which caused the Great French Wine Blight and a plague that devastated European viticulture in the 19th century. Similarly, adelgids also feed on plant phloem. Adelgids are sometimes described as aphids, but more properly as classified as aphid-like insects, because they have no cauda or cornicles.3 Anatomy The life stages of the green apple aphid Aphis pomi The life stages of the green apple aphid Aphis pomi Most aphids have soft, green bodies, but other colors are common, such as black, brown, and pink. Aphids have antennas with as many as six segments.3 Aphids feed themselves through sucking mouthparts called stylets, enclosed in a sheath called a rostrum, which is formed from modifications of the mandible and maxilla.8 They have long, thin legs and two-jointed, two-clawed tarsi. Most aphids have a pair of cornicles or siphunculi, abdominal tubes through which they exude droplets of a quick-hardening defensive fluid8 containing triacylglycerols, called cornicle wax. Other defensive compounds can also be produced by some types of aphids.3 Aphids have a tail-like protrustion called a cauda above their rectal apertures. They have two compound eyes, and an ocular tubercle behind and above each eye, made up of three lenses called triommatidia.910 When host plant quality becomes poor or conditions become crowded, some aphid species produce winged offspring, alates, that can disperse to other food sources. The mouthparts or eyes are smaller or missing in some species and forms.3 Diet Many aphid species are monophagous that is, they feed on only one plant species. Others, like the green peach aphid Myzus persicae, feed on hundreds of plant species across many families. Aphids passively feed on sap of phloem vessels in plants, as do many of their fellow members of Hemiptera such as scale insects and cicadas. Once a phloem vessel is punctured, the sap, which is under high pressure, is forced into the aphid's food canal. As they feed, aphids often transmit plant viruses to the plants, such as to potatoes, cereals, sugarbeets and citrus plants.8 These viruses can sometimes kill the plants. Plants contain low densities of the nitrogen compounds needed for building proteins. This requires aphids to consume an excess of sap to satisfy their nutritional requirements. The excess is expelled as honeydew, out of the recta of aphids, in such large volumes that in sometimes it can fall like rain.3 Aphid honeydew is rich in carbohydrates, like the phloem it derives from. Symbioses Ant tending aphids Ant tending aphids Some species of ants farm aphids, protecting them on the plants they eat, and eating the honeydew that the aphids release from the terminations of their alimentary canals. This is a mutualistic relationship. These dairying ants milk the aphids by stroking them with their antennae.1112 Therefore, sometimes aphids are called ant cows. 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 flavus13 manage large herds of aphids that feed on roots of plants in the ant colony. Queens that are 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.12 An interesting variation in ant-aphid relationships involves Lycaenid butterflies such as the Sievers blue butterfly and the Japanese copper butterfly and the 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, but carry the caterpillars to their nest. In the nest, the ants feed the caterpillars, which produce honeydew for the ants. When the caterpillars reach full size, they crawl to the colony entrance and form cocoons. After two weeks, butterflies emerge and take flight.14 Some bees in coniferous forests also collect aphid honeydew to make forest honey.8. Many aphids are host to endosymbiont bacteria, Buchnera, inside specialized cells called bacteriocytes. These bacteria synthesize some essential amino acids that are absent from the phloem that the aphids eat.15 Reproduction Aphid giving birth to live young Aphid giving birth to live young Some aphid species have unusual and complex reproductive adaptations, while others have fairly simple reproduction. Adaptations include having both sexual and asexual reproduction, creation of eggs or live nymphs and switches between woody and herbaceous types of host plant at different times of the year.316 Many aphids undergo cyclical parthenogenesis. In the spring and summer, mostly or only females are present in the population. The overwintering eggs that hatch in the spring result in females, called fundatrices. Reproduction is typically parthenogenetic and viviparous. Females undergo a modified meiosis that results in eggs that are genetically identical to their mother parthenogenetic. The embryos develop within the mothers' ovarioles, which then give live birth to first instar female nymphs viviparous. The offspring resemble their parent in every way except size, and are called virginoparae. This process iterates throughout the summer, producing multiple generations that typically live 20 to 40 days. Thus one female hatched in spring may produce many billions of descendants. For example, some species of cabbage aphids like Brevicoryne brassicae can produce up to 41 generations of females, or more than 1.5 x 1027 offspring if they all live. In autumn, aphids undergo sexual, oviparous reproduction. A change in photoperiod and temperature, or perhaps a lower food quantity or quality, causes females to parthenogenetically produce sexual females and males. The males are genetically identical to their mothers except that they have one less sex chromosome. These sexual aphids may lack wings or even mouthparts.3 Sexual females and males mate, and females lay eggs that develop outside the mother. The eggs endure the winter and emerge as winged or wingless females the following spring. This is, for example, the life cycle of the rose aphid Aphis rosae, which may be considered typical of the family. However in warm environments, such as a in the tropics or in a glasshouse or greenhouse, aphids may go on reproducing asexually for many years.8 Some species produce winged females in the summer, sometimes in response to low food quality or quantity. The winged females migrate to start new colonies on a new plant, often of quite a different kind. For example, the apple aphid Aphis mali, after producing many generations of apterous females on its typical food-plant, gives rise to winged forms which fly away and settle on grass or corn-stalks. Some aphids have telescoping generations. That is, the parthenogenetic, viviparous female has a daughter within her, who is already parthenogenetically producing her own daughter. Thus a female's diet can effect the body size and birth rate of more than one generation daughters and granddaughters.1718 Evolution Aphids probably appeared around 280 million years ago, in the early Permian period. They probably fed on plants like Cordaitales or Cycadophyta. The oldest known aphid fossil is of the species Triassoaphis cubitus Evans from the Triassic.19 The number of species was small, but increased considerably with the appearance of angiosperms 160 million years ago. Angiosperms allowed aphids to specialize. Organs like the cornicles did not appear until the Cretaceous period. Threats Aphids are soft-bodied, and have a wide variety of insect predators. Aphids also are often infected by bacteria, viruses and fungi. Aphids are affected by the weather, such as precipitation,20 temperature21 and wind.22 Lady beetle larva consuming an aphid Lady beetle larva consuming an aphid Insects that attack aphids include predatory lady beetles Coleoptera: Coccinellidae, hoverfly larvae Diptera: Syrphidae, parasitic wasps, aphid midge larvae, aphid lions, crab spiders23 and lacewings Neuroptera: Chrysopidae. Fungi that attack aphids include Neozygites fresenii, Entomophthora, Beauveria bassiana, Metarhizium anisopliae and entomopathogenic fungi like Lecanicillium lecanii. Aphids brush against the spores, which are typically about 1/4 inch long. These spores stick to the aphid, germinate and penetrate the aphid's skin. The fungus grows in the aphid hemolymph i.e., the counterpart of blood for aphids. After about 3 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 type of fungus that killed the aphid, but a secondary fungus.20 Aphids can be easily killed by unfavorable weather, such as late spring freezes.24 Excessive heat kills the symbiotic bacteria that some aphids depend on, which makes the aphids infertile.25 Rain prevents winged aphids from dispersing, and knocks aphids off plants and thus kills them from the impact or by starvation.202627 However, Ken Ostlie, an entomologist with the University of Minnesota Extension Service, suggests that rain should not be relied on for aphid control.28 Defenses A P-14 lady beetle consuming an aphid A P-14 lady beetle consuming an aphid29 Aphid excreting defensive fluid from the cornicles. Aphid excreting defensive fluid from the cornicles. Aphids are soft-bodied, and have little protection from predators and diseases. Some species of aphid 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.308 For example, Alexander's horned aphids are a type of soldier aphid that has a hard exoskeleton and pincer-like mouthparts.31 Infestation of a variety of Chinese trees by Chinese sumac aphids Melaphis chinensis Bell can create a Chinese gall which is valued as a commercial product. As Galla Chinensis, Chinese galls are used as a medication used in Chinese medicine to treat coughs, diarrhea, night sweats, dysentry and to stop intestinal and uterine bleeding. Chinese galls are also an important source of tannins.8 Some species of aphid, known as woolly aphids Eriosomatinae, excrete a fluffy wax coating for protection.8 The cabbage aphid Brevicoryne brassicae stores and releases chemicals that produce a violent chemical reaction and strong mustard oil smell to repel predators. 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 Dictionary32 and the 2008 ion of the World Book Encyclopedia.33. In fact, honeydew secretions are produced from the anus of the aphid34, while cornicles mostly produce defensive chemicals such as waxes. There also is evidence of cornicle wax attracting aphid predators in some cases.35 Aphids are also known to defend themselves from attack by parasitoid wasps by kicking. Effects on plants 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 vigour in the plant, and aphid saliva is toxic to plants. Aphids frequently transmit disease-causing organisms like plant viruses to their hosts. The green peach aphid Myzus persicae is a vector for more than 110 plant viruses. Cotton aphids Aphis gossypii often infect sugarcane, papaya and groundnuts with viruses.3 Aphids contributed to the spread of late blight Phytophthora infestans among potatoes in the Great Irish Potato Famine of the 1840s.36 The coating of plants with honeydew can contribute to the spread of fungi which can damage plants.373839 Honeydew produced by aphids has been observed to reduce the effectiveness of fungicides as well.40 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 a noninfested plant. However, this does not appear to be supported by the observational evidence.41 The damage of plants, and in particular commercial crops, has resulted in large amounts of resources and efforts being spent attempting to control the activities of aphids.3 In popular culture In the movie Antz, Weaver says to Zee Don't you want your aphid beer? Zee replies I can't help it. I have a thing about drinking from the anus of another creature. Call me crazy. In the movie A Bug's Life, the Ant Queen has a pet aphid named Aphie.42 In the book A Scanner Darkly, Jerry Fabin is tormented by hallucinations of aphids crawling all over his body. In the film Freck is the one tormented by the aphids. Gallery Aphid infestation on broccoli plant Lupine stalk infested with aphids Two aphids fighting Two aphids Cluster of aphids Aphids under attack on a thistle Aphid colors Aphids Aphid infestation on a sunflower stalk Aphids on rose foliage Aphids feeding on a rose bud. A lady beetle can be seen climbing the stalk. Ladybird larva eating wooly apple aphids See also Aeroplankton Pineapple gall Economic entomology References This article incorporates text from the Encyclopædia Britannica Eleventh ion, a publication now in the public domain. Monograph of the British Aphides, Volumes I-IV, George Bowdler Buckton, Ray Society, 1876-1883. ^ Not to be confused with jumping plant lice ^ Aphids are also sometimes known as blackflies and greenflies page 86 of Bugs of the World, George C. McGavin, Facts on File, 1993 ^ a b c d e f g h i j Bugs of the World, George C. McGavin, Facts on File, 1993, ISBN 0816027374 ^ Photo of crab spider eating Aphis asclepiadis aphids on common milkweed, Anurag Agrawal, Phytophagy Laboratory, Department of Ecology and Evolutionary Biology and Department of Entomology at Cornell University. ^ Scientist battles lettuce aphid, Pip Courtney, Landline, Australian Broadcasting Corporation, October 30, 2005, Retrieved 1 January 2007 ^ Aphid Ecology - An optimization approach, Second ion, A.F.G. Dixon, Springer; 2nd ed. ion 1997, ISBN 0412741806 ^ Biology and Management of Grape Phylloxera, Jeffrey Granett, M. Andrew Walker,-Laszlo Kocsis, and Amir D. Omer, Annual Review of Entomology, Vol. 46: 387-412, January 2001, doi 10.1146/annurev.ento.46.1.387 ^ a b c d e f g h Aphid, Henry G. Stroyan, McGraw-Hill Encyclopedia of Science and Technology, 8th ion, 1997, ISBN 0-07-911504-7 ^ Molecular Studies of the Salivary Glands of the Pea Aphid, Acyrthosiphon Pisum Harris, Navdeep S. Mutti, PhD Thesis, Kansas State University, 2006. ^ Aphid Ecology, A. F. G. Dixon, Chapman and Hall, 1998, ISBN 0412741806 ^ There are also dairying ants that milk mealybugs and other insects. ^ a b Ant, Linda M. Hooper-Bui, World Book Encyclopedia, 2008, ISBN 978-0-7166-0108-1 ^ Insects of the World, Anthony Wootton, Blandford, Cassell Plc, 1984, reprinted 1999, ISBN 0713723661 ^ pages 78 and 79 of Insects and Spiders, Time-Life Books, ISBN 0809496879 ^ Douglas, A E 1998. Nutritional interactions in insect-microbial symbioses: Aphids and their symbiotic bacteria Buchnera. Annual Review of Entomology 43: 178. doi:10.1146/annurev.ento.43.1.17. ISSN 00664170. Retrieved on 2007-05-16. ^ About 10% of aphid species produce generations that alternate between woody and herbaceous plants page 87 of Bugs of the World, George C. McGavin, Facts on File, 1993. ^ Effect of nitrogen fertilization on Aphis gossypii Homoptera: Aphididae: variation in size, color, and reproduction, E. Nevo and M. Coll, J. Econ. Entomol. 94: 27-32, 2001. ^ Effect of nitrogen fertilizer on the intrinsic rate of increase of the rusty plum aphid, Hysteroneura setariae Thomas Homoptera: Aphididae on rice Oryza sativa L., G. C. Jahn, L. P. Almazan, and J. Pacia, Environmental Entomology 34 4: 938-943, 2005. ^ Acropyga and Azteca Ants Hymenoptera: Formicidae with Scale Insects Sternorrhyncha: Coccoidea: 20 Million Years of Intimate Symbiosis, Christine Johnson, Dant Agosti, Jacques H. Delabie, Klaus Dumpert, D.J. Williams, Michael von Tschirnhaus and Ulrich Maschwitz, American Museum Novitates, June 22, 2001. ^ a b c Early Season Aphid and Thrips Populations, Gerald E. Brust, University of Maryland College of Agriculture and Natural Resources News Article, June 22, 2006 ^ Some Effects of Fluctuating Temperatures on Metabolism, Development, and Rate of Population Growth in the Cabbage Aphid, Brevicoryne Brassicae, K. P. Lamb, Ecology, Vol. 42, No. 4 Oct., 1961, pp. 740-745 ^ Abundance of Aphids on Cereals from Before 1973 to 1977, Margaret G. Jones, The Journal of Applied Ecology, Vol. 16, No. 1 Apr., 1979, pp. 1-22 ^ Photo of crab spider eating Aphis asclepiadis aphids on common milkweed, Anurag Agrawal, Phytophagy Laboratory, Department of Ecology and Evolutionary Biology and Department of Entomology at Cornell University. ^ Soybean Aphid, A New Beginning for 2007, Christian Krupke, John Obermeyer, and Robert Oeil, Pest and Crop, May 11, 2007 - Issue 7, Purdue Extension Service. ^ Why Some Aphids Can't Stand The Heat, Science Daily, April 23, 2007. ^ Population Dynamics of the Cabbage Aphid, Brevicoryne brassicae L., R. D. Hughes, The Journal of Animal Ecology, Vol. 32, No. 3 Oct., 1963, pp. 393-424 ^ Stable Age Distributions of Lucerne Aphid Populations in SE-Tasmania, S. Suwanbutr, page 38-43, Thammasat International Journal of Science and Technology, Vol 1, No. 5, 1996 ^ Spider Mites, Aphids and Rain Complicating Spray Decisions in Soybean, Ken Ostlie, Minnesota Crop eNews, University of Minnesota Extension Service, August 3, 2006 ^ The fourteen-spotted lady beetles are also known as P-14 lady beetles, or propylea quatuordecimpunctata. ^ Aoki, S. 1977 Colophina clematis Homoptera, Pemphigidae, an aphid species with soldiers. Kontyu 45, 27682 ^ page 144 of Insects and Spiders, Time-Life Books, ISBN 0809496879 ^ Defence by Smear: Supercooling in the Cornicle Wax of Aphids, John S. Edwards, Letters to Nature, Nature, 211, 73 - 74, 02 July 1966; doi:10.1038/211073a0 ^ Aphid, Candace Martinson, World Book Encyclopedia, 2008, ISBN 978-0-7166-0108-1 ^ Mutualism between ants and honeydew producing homoptera. MJ Way. Annual Review of Entomology ^ Kairomonal effect of an aphid cornicle secretion onLysiphlebus testaceipes Cresson Hymenoptera: Aphidiidae, Tessa R. Grasswitz and Timothy D. Paine, Journal of Insect Behavior, Springer Netherlands, ISSN 0892-7553 Print 1572-8889 Online, Issue Volume 5, Number 4 / July, 1992, DOI 10.1007/BF01058190 ^ page 61 of The Most Extreme Bugs, Catherine Nichols, Forward by Kevin Mohs and Ian McGee, Jossey-Bass, John Wiley and Sons, 2007, ISBN 9780787986636 ^ Sooty mold fungus growing on honeydew deposited on lower sugarcane leaves by yellow sugarcane aphids, Sipha flava Forbes, University of Florida ^ Sooty mold, Daniel H. Gillman, University of Massachusetts Extension, Fall 2005 ^ Scorias spongiosa, the beech aphid poop-eater, Hannah T. Reynolds and Tom Volk, Tom Volk's Fungus of the Month, University of Wisconsin-La Crosse, September 2007 ^ Interaction between phyllosphere yeasts, aphid honeydew and fungicide effectiveness in wheat under field conditions, J. Dika and J. A. Van Pelt, Plant pathology, vol. 41, no6, pp. 661-675 1 p., 1992, ISSN 0032-0862 CODEN PLPAAD ^ Aphid Honeydew: A re-appraisal of the hypothesis of Owen and Wiegert, Dhurpad Choudhury, Oikos, Vol. 45, No. 2 Oct., 1985, pp. 287-290 ^ Wikia article on Aphie External links on the UF / IFAS Featured Creatures Web site Aphis gossypii, melon or aphid Aphis nerii, oleander aphid Hyadaphis coriandri, corianderaphid Longistigma caryae, giant bark aphid Myzus persicae, green peach aphid Sarucallis kahawaluokalani, crapemyrtle aphid Shivaphis celti, an Asian woolly hackberry aphid Toxoptera citricida, brown citrus aphid Retrieved from http://en..org/wiki/Aphid Categories: Hemiptera | Pest insects | Insect vectors of plant pathogens Views Article Discussion this page History Personal tools Log in / create account Navigation Main page Contents Featured content Current events Random article Search Go Search Interaction Community portal Recent changes Contact Donate to Help Toolbox What links here Related changes Upload file Special pages Printable version Permanent link Cite this page Languages العربية ÄŒesky Dansk Deutsch Eesti Español Esperanto Ù?ارسی Français í•œêµì–´ Ido Italiano עברית Lietuvių Magyar Nederlands 日本語 ‪Norsk bokmÃ¥l‬ ‪Norsk nynorsk‬ Polski Português Runa Simi РуÑ?Ñ?кий SlovenÅ¡Ä?ina Suomi Svenska УкраїнÑ?ька This page was last modified on 12 September 2008, at 08:00
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