Anopheles 542071 225014283 2008-07-11T13:41:34Z Mark-mitchell-aldershot 3401544 {{Taxobox | name = ''Anopheles'' | image = Anopheles stephensi.jpeg | image_width = 300px | image_caption = ''[[Anopheles stephensi]]'' | regnum = [[Animal]]ia | phylum = [[Arthropod]]a | subphylum = [[Hexapoda]] | classis = [[Insect]]a | subclassis = [[Pterygota]] | infraclassis = [[Neoptera]] | superordo = [[Endopterygota]]| | ordo = [[Fly|Diptera]] | subordo = [[Nematocera]] | infraordo = [[Culicomorpha]] | superfamilia = [[Culicoidea]] | familia = [[Culicidae]] | genus = '''''Anopheles''''' | genus_authority = [[Johann Wilhelm Meigen|Meigen]], 1818 | range_map = Anopheles-range-map.png | range_map_width = 300px | range_map_caption = ''Anopheles'' range map | subdivision_ranks = Some Species | subdivision = *''[[Anopheles acanthotorynus]]'' *''[[Anopheles albimanus]]'' *''[[Anopheles albitarsis]]'' *''[[Anopheles amictus]]'' *''[[Anopheles annulipalpis]]'' *''[[Anopheles anomalophyllus]]'' *''[[Anopheles antunesi]]'' *''[[Anopheles apicimacula]]'' *''[[Anopheles aquasalis]]'' *''[[Anopheles argyritarsis]]'' *''[[Anopheles atroparvus]]'' *''[[Anopheles aztecus]]'' *''[[Anopheles bambusicolus]]'' *''[[Anopheles barberi]]'' *''[[Anopheles beklemishevi]]'' *''[[Anopheles bellator]]'' *''[[Anopheles benarrochi]]'' *''[[Anopheles boliviensis]]'' *''[[Anopheles bonnei]]'' *''[[Anopheles bradleyi]]'' *''[[Anopheles braziliensis]]'' *''[[Anopheles bustamentei]]'' *''[[Anopheles canorii]]'' *''[[Anopheles coustani]]'' *''[[Anopheles crucians]]'' *''[[Anopheles cruzii]]'' *''[[Anopheles crypticus]]'' *''[[Anopheles culicifacies]]'' *''[[Anopheles darlingi]]'' *''[[Anopheles eiseni]]'' *''[[Anopheles earlei]]'' *''[[Anopheles evandroi]]'' *''[[Anopheles evansae]]'' *''[[Anopheles farauti]]'' *''[[Anopheles fausti]]'' *''[[Anopheles fluminensis]]'' *''[[Anopheles fluviatilis]]'' *''[[Anopheles forattinii]]'' *''[[Anopheles franciscanus]]'' *''[[Anopheles freeborni]]'' *''[[Anopheles funestus]]'' *''[[Anopheles gabaldoni]]'' *''[[Anopheles galvaoi]]'' *''[[Anopheles gambiae]]'' *''[[Anopheles georgianus]]'' *''[[Anopheles gilesi]]'' *''[[Anopheles gomezdelatorrei]]'' *''[[Anopheles grabhamii]]'' *''[[Anopheles guarao]]'' *''[[Anopheles hailarensis]]'' *''[[Anopheles halophylus]]'' *''[[Anopheles hectoris]]'' *''[[Anopheles homunculus]]'' *''[[Anopheles hyrcanus]]'' *''[[Anopheles ininii]]'' *''[[Anopheles intermedius]]'' *''[[Anopheles introlatus]]'' *''[[Anopheles judithae]]'' *''[[Anopheles kompi]]'' *''[[Anopheles kosiensis]]'' *''[[Anopheles lanei]]'' *''[[Anopheles latens]]'' *''[[Anopheles lutzii]]'' *''[[Anopheles maculipennis]]'' *''[[Anopheles mattogrossensis]]'' *''[[Anopheles mediopunctatus]]'' *''[[Anopheles melas]]'' *''[[Anopheles merus]]'' *''[[Anopheles minimus]]'' *''[[Anopheles minor]]'' *''[[Anopheles moucheti]]'' *''[[Anopheles neivai]]'' *''[[Anopheles neomaculipalpus]]'' *''[[Anopheles nigritarsis]]'' *''[[Anopheles nili]]'' *''[[Anopheles nimbus]]'' *''[[Anopheles noroestensis]]'' *''[[Anopheles nuneztovari]]'' *''[[Anopheles occidentalis]]'' *''[[Anopheles oiketorakras]]'' *''[[Anopheles oswaldoi]]'' *''[[Anopheles ovengensis]]'' *''[[Anopheles pampanae]]'' *''[[Anopheles parapunctipennis]]'' *''[[Anopheles parvus]]'' *''[[Anopheles perplexens]]'' *''[[Anopheles peryassui]]'' *''[[Anopheles peytoni]]'' *''[[Anopheles pictipennis]]'' *''[[Anopheles pseudomaculipes]]'' *''[[Anopheles pseudopunctipennis]]'' *''[[Anopheles pseudotibiamaculatus]]'' *''[[Anopheles punctimacula]]'' *''[[Anopheles punctulatus]]'' *''[[Anopheles punctipennis]]'' *''[[Anopheles quadrimaculatus]]'' *''[[Anopheles rachoui]]'' *''[[Anopheles rangeli]]'' *''[[Anopheles rennellensis]]'' *''[[Anopheles rivulorum]]'' *''[[Anopheles rondoni]]'' *''[[Anopheles sanctielii]]'' *''[[Anopheles shannoni]]'' *''[[Anopheles squamifemur]]'' *''[[Anopheles stephensi]]'' *''[[Anopheles thomasi]]'' *''[[Anopheles tibiamaculatus]]'' *''[[Anopheles triannulatus]]'' *''[[Anopheles vargasi]]'' *''[[Anopheles vestitipennis]]'' *''[[Anopheles walkeri]]'' *''[[Anopheles xelajuensis]]'' }} '''''Anopheles''''' is a [[genus]] of [[mosquito]] (''Culicidae''). There are approximately 400 ''Anopheles'' species, of which 30-40 transmit five different species of parasites of the genus ''[[Plasmodium]]'' that cause [[malaria]] which affects humans in [[endemic (ecology)|endemic]] areas. ''[[Anopheles gambiae]]'' is one of the best known, because of its predominant role in the transmission of the most dangerous ''[[Plasmodium falciparum]]''. Some species of ''Anopheles'' also can serve as the vectors for canine [[heartworm]] ''[[Dirofilaria immitis]]'', the [[Filariidae]] ''[[Wuchereria bancrofti]]'' and ''[[Brugia malayi]]'', and [[virus]]es like the one that is the cause of [[O'nyong'nyong]] fever. Mosquitoes in other genera ([[Aedes]], [[Culex]]) can also serve as vectors of disease agents. == Life stages == Like all mosquitoes, anophelines go through four stages in their life cycle: [[egg (biology)|egg]], [[larva]], [[pupa]], and [[imago]]. The first three stages are aquatic and last 5-14 days, depending on the species and the ambient temperature. The adult stage is when the female ''Anopheles'' mosquito acts as [[malaria]] [[vector (biology)|vector]]. The adult females can live up to a month (or more in captivity) but most probably do not live more than 1-2 weeks in nature. === Eggs === Adult females lay 50-200 eggs per [[oviposition]]. Eggs are laid singly directly on water and are unique in having floats on either side. Eggs are not resistant to drying and hatch within 2-3 days, although hatching may take up to 2-3 weeks in colder climates. === Larvae === [[Image:Anopheles larve.jpg|left|thumb|Anopheles larva from southern Germany, about 8 mm long.]] Mosquito larvae have a well-developed head with mouth brushes used for feeding, a large [[thorax]] and a segmented [[abdomen]]. They don't have legs. In contrast to other mosquitoes, ''Anopheles'' larvae lack a respiratory siphon and for this reason position themselves so that their body is parallel to the surface of the water. Larvae breathe through [[spiracle]]s located on the 8th abdominal segment and therefore must come to the surface frequently. The larvae spend most of their time feeding on [[algae]], [[bacteria]], and other microorganisms in the surface microlayer. They dive below the surface only when disturbed. Larvae swim either by jerky movements of the entire body or through [[Marine propulsion|propulsion]] with the mouth brushes. Larvae develop through 4 stages, or [[instar]]s, after which they [[metamorphosis|metamorphose]] into [[pupae]]. At the end of each instar, the larvae molt, shedding their exoskeleton, or skin, to allow for further growth. The larvae occur in a wide range of habitats but most species prefer clean, unpolluted water. Larvae of ''Anopheles'' mosquitoes have been found in fresh- or salt-water marshes, mangrove swamps, rice fields, grassy ditches, the edges of streams and rivers, and small, temporary rain pools. Many species prefer habitats with vegetation. Others prefer habitats that have none. Some breed in open, sun-lit pools while others are found only in shaded breeding sites in forests. A few species breed in tree holes or the leaf axils of some plants. === Pupae === The pupa is comma-shaped when viewed from the side. The head and [[thorax]] are merged into a [[cephalothorax]] with the abdomen curving around underneath. As with the larvae, pupae must come to the surface frequently to breathe, which they do through a pair of respiratory trumpets on the cephalothorax. After a few days as a pupa, the dorsal surface of the cephalothorax splits and the adult mosquito emerges. === Adults === The duration from egg to adult varies considerably among species and is strongly influenced by ambient temperature. Mosquitoes can develop from egg to adult in as little as 5 days but usually take 10-14 days in tropical conditions. Like all mosquitoes, adult ''Anopheles'' have slender bodies with 3 sections: head, thorax and abdomen. The head is specialized for acquiring sensory information and for feeding. The head contains the eyes and a pair of long, many-segmented [[Antenna (biology)|antennae]]. The antennae are important for detecting host odors as well as odors of breeding sites where females lay eggs. The head also has an elongate, forward-projecting [[proboscis]] used for feeding, and two sensory palps. The [[thorax]] is specialized for locomotion. Three pairs of legs and a pair of wings are attached to the thorax. The abdomen is specialized for food digestion and egg development. This segmented body part expands considerably when a female takes a blood meal. The blood is digested over time serving as a source of [[protein]] for the production of eggs, which gradually fill the [[abdomen]]. ''Anopheles'' mosquitoes can be distinguished from other mosquitoes by the [[palp]]s, which are as long as the proboscis, and by the presence of discrete blocks of black and white scales on the wings. Adult ''Anopheles'' can also be identified by their typical resting position: males and females rest with their abdomens sticking up in the air rather than parallel to the surface on which they are resting. Adult mosquitoes usually mate within a few days after emerging from the pupal stage. In most species, the males form large [[swarm]]s, usually around dusk, and the females fly into the swarms to mate. Males live for about a week, feeding on [[nectar]] and other sources of [[sugar]]. Females will also feed on sugar sources for energy but usually require a blood meal for the development of eggs. After obtaining a full blood meal, the female will rest for a few days while the blood is digested and eggs are developed. This process depends on the temperature but usually takes 2-3 days in tropical conditions. Once the eggs are fully developed, the female lays them and resumes host seeking. The cycle repeats itself until the female dies. While females can live longer than a month in captivity, most do not live longer than 1-2 weeks in nature. Their lifespan depends on temperature, humidity, and also their ability to successfully obtain a blood meal while avoiding host defenses. == Habitat == Although malaria is nowadays limited to tropical areas, most notoriously regions of sub-Saharan Africa, many ''Anopheles'' species live in colder latitudes (see [http://www.cdc.gov/Malaria/biology/mosquito/map.htm this map] from the CDC). Indeed, malaria outbreaks have, in the past, occurred in colder climates, for example during the construction of the [[Rideau Canal]] in Canada during the 1820s. Since then, the ''Plasmodium'' parasite (not the ''Anopheles'' mosquito) has been eradicated from first world countries. The [http://www.cdc.gov/Malaria/biology/mosquito/ CDC] warns, however, that "''Anopheles'' that can transmit malaria are found not only in malaria-endemic areas, but also in areas where malaria has been eliminated. The latter areas are thus constantly at risk of re-introduction of the disease." == Susceptibility to become a vector of disease == Some species are poor vectors of [[malaria]], as the [[parasite]]s do not develop well (or at all) within them. There is also variation within species. In the laboratory, it has been possible to select for strains of ''A. gambiae'' that are refractory to infection by [[malaria]] parasites. These refractory strains have an [[immune]] response that encapsulates and kills the parasites after they have invaded the mosquito's [[stomach]] wall. Scientists are studying the [[genetics|genetic]] mechanism for this response. It is hoped that some day, genetically modified mosquitoes that are refractory to malaria can replace wild mosquitoes, thereby limiting or eliminating malaria transmission. === Malaria transmission and control === Understanding the biology and behavior of ''Anopheles'' mosquitoes can help understand how [[malaria]] is transmitted and can aid in designing appropriate control strategies. Factors that affect a mosquito's ability to transmit malaria include its innate susceptibility to [[Plasmodium]], its host choice and its longevity. Factors that should be taken into consideration when designing a control program include the susceptibility of malaria vectors to [[insecticide]]s and the preferred feeding and resting location of adult mosquitoes. On [[December 21]], [[2007]], a study published in [[PLoS Pathogens]] found that the hemolytic C-type [[lectin]] CEL-III from ''[[Dendrochirotida|Cucumaria echinata]]'', a [[sea cucumber]] found in the [[Bay of Bengal]], impaired the development of the malaria parasite when produced by transgenic ''A.&nbsp;stephensi''.<ref>{{cite journal |author=Yoshida S, Shimada Y, Kondoh D, ''et al'' |title=Hemolytic C-type lectin CEL-III from sea cucumber expressed in transgenic mosquitoes impairs malaria parasite development |journal=PLoS Pathog. |volume=3 |issue=12 |pages=e192 |year=2007 |pmid=18159942 |doi=10.1371/journal.ppat.0030192 |url=http://www.plospathogens.org/article/info:doi/10.1371/journal.ppat.0030192}}</ref> This could potentially be used one day to control malaria by spreading genetically modified mosquitoes refractory to the parasites, although there are numerous scientific and ethical issues to be overcome before such a control strategy could be implemented. === Preferred sources for blood meals === One important behavioral factor is the degree to which an ''Anopheles'' species prefers to feed on humans ([[anthropophily]]) or animals such as cattle ([[zoophilic|zoophily]]). Anthropophilic ''Anopheles'' are more likely to transmit the malaria parasites from one person to another. Most ''Anopheles'' mosquitoes are not exclusively anthropophilic or zoophilic. However, the primary malaria vectors in [[Africa]], ''A. gambiae'' and ''A. funestus'', are strongly anthropophilic and, consequently, are two of the most efficient malaria vectors in the world. Once ingested by a mosquito, malaria parasites must undergo development within the mosquito before they are infectious to humans. The time required for development in the mosquito (the extrinsic [[incubation period]]) ranges from 10-21 days, depending on the [[parasite]] species and the temperature. If a mosquito does not survive longer than the extrinsic incubation period, then she will not be able to transmit any malaria parasites. It is not possible to measure directly the life span of mosquitoes in nature. But indirect estimates of daily survivorship have been made for several ''Anopheles'' species. Estimates of daily survivorship of ''A. gambiae'' in [[Tanzania]] ranged from 0.77 to 0.84 meaning that at the end of one day between 77% and 84% will have survived.<ref>(Charlwood et al., 1997, Survival And Infection Probabilities of Anthropophagic Anophelines From An Area of High Prevalence of Plasmodium falciparum in Humans, Bulletin of Entomological Research, 87, 445-453)</ref> Assuming this survivorship is constant through the adult life of a mosquito, less than 10% of female ''A. gambiae'' would survive longer than a 14-day extrinsic incubation period. If daily survivorship increased to 0.9, over 20% of mosquitoes would survive longer than a 14-day extrinsic incubation period. Control measures that rely on [[insecticide]]s (e.g. indoor residual spraying) may actually impact malaria [[Transmission (medicine)|transmission]] more through their effect on adult longevity than through their effect on the population of adult mosquitoes. === Patterns of feeding and resting === Most ''Anopheles'' [[mosquito]]es are [[crepuscular]] (active at dusk or dawn) or [[nocturnal]] (active at night). Some ''Anopheles'' mosquitoes feed indoors (endophagic) while others feed outdoors (exophagic). After feeding on some blood mosquitoes prefer to rest indoors (endophilic) while others prefer to rest outdoors (exophilic), though this can differ regionally based on local vector ecotype, and vector chromosomal makeup, as well as housing type and local microclimatic conditions. Biting by nocturnal, endophagic ''Anopheles'' mosquitoes can be markedly reduced through the use of [[insecticide]]-treated bed nets (ITNs) or through improved housing construction to prevent mosquito entry (e.g. window screens). Endophilic mosquitoes are readily controlled by indoor spraying of residual insecticides. In contrast, exophagic/exophilic vectors are best controlled through source reduction (destruction of the breeding sites). == Insecticide resistance == [[Insecticide]]-based control measures (e.g. indoor spraying with insecticides, ITNs) are the principal way to kill mosquitoes that bite indoors. However, after prolonged exposure to an insecticide over several [[generation]]s, mosquitoes, like other [[insect]]s, may develop resistance, a capacity to survive contact with an insecticide. Since mosquitoes can have many generations per year, high levels of resistance can arise very quickly. Resistance of mosquitoes to some insecticides has been documented with just within a few years after the insecticides were introduced. There are over 125 mosquito [[species]] with documented resistance to one or more insecticides. The development of resistance to insecticides used for indoor residual spraying was a major impediment during the Global Malaria Eradication Campaign. Judicious use of insecticides for mosquito control can limit the development and spread of resistance. However, use of insecticides in agriculture has often been implicated as contributing to resistance in mosquito populations. It is possible to detect developing resistance in mosquitoes and control programs are well advised to conduct surveillance for this potential problem. == See also == *[[Tropical disease]] *[[O'nyong'nyong virus]] === Source === * ''Original version from http://www.cdc.gov/malaria/biology/mosquito/'' * ''http://animaldiversity.ummz.umich.edu/site/accounts/classification/Anopheles.html'' == References == <references/> === External links === * [http://www.anobase.org/ ''Anopheles'' Database] * [http://www.cdc.gov/Malaria/biology/mosquito/ CDC - National Center for Infectious Diseases, Division of Parasitic Diseases; ''Anopheles'' Mosquitoes] * [http://www.cdc.gov/Malaria/biology/mosquito/map.htm World map showing distribution of various ''Anopheles'' species] * [http://www.wrbu.org/ Walter Reed Biosystematics Unit] * [http://www.map.ox.ac.uk Malaria Atlas Project] [[Category:Culicidae]] [[Category:Insects of Africa]] [[Category:Insect vectors of human pathogens]] [[Category:Malaria]] [[Category:Flies]] [[de:Anopheles]] [[es:Anopheles]] [[fr:Anophèle]] [[id:Anopheles]] [[it:Anopheles]] [[he:אנופלס]] [[ka:მალარიის კოღო]] [[hu:Maláriaszúnyogok]] [[ms:Anopheles]] [[nl:Anopheles]] [[no:Malariamygg]] [[pt:Anopheles]] [[ru:Малярийные комары]] [[sl:Komar mrzličar]] [[sv:Malariamyggor]] [[zh:瘧蚊]]