The mosquito life cycle consists of four stages:egg,larva,pupa, andadult. Eggs are laid on the water surface; they hatch intomotile larvae that feed on aquaticalgae andorganic material. These larvae are important food sources for many freshwater animals, such asdragonfly nymphs, many fish, and some birds. Adult females of many species have mouthparts adapted to pierce the skin of ahost andfeed on blood of a wide range ofvertebrate hosts, and someinvertebrates, primarily otherarthropods. Some species only produce eggs after a blood meal.
The mosquito'ssaliva is transferred to the host during the bite, and can cause an itchyrash. In addition, blood-feeding species can ingest pathogens while biting, and transmit them to other hosts. Those species include vectors ofparasitic diseases such asmalaria andfilariasis, andarboviral diseases such asyellow fever anddengue fever. By transmitting diseases, mosquitoes cause the deaths of over 725,000 people each year.
Description and life cycle
Like all flies, mosquitoes go through four stages in their life cycles:egg,larva,pupa, andadult. The first three stages—egg, larva, and pupa—are largely aquatic,[4] the eggs usually being laid in stagnant water.[5] They hatch to becomelarvae, which feed, grow, and molt until they change intopupae. The adult mosquito emerges from the mature pupa as it floats at the water surface. Mosquitoes have adult lifespans ranging from as short as a week to around a month. Some species overwinter as adults indiapause.[6]
Adult
Mosquitoes have one pair of wings, with distinct scales on the surface. Their wings are long and narrow, while the legs are long and thin. The body, usually grey or black, is slender, and typically 3–6 mm long. When at rest, mosquitoes hold their first pair of legs outwards, whereas the somewhat similarChironomid midges hold these legs forwards.[7]Anopheles mosquitoes can fly for up to four hours continuously at 1 to 2 km/h (0.62 to 1.24 mph),[8] traveling up to 12 km (7.5 mi) in a night. Males beat their wings between 450 and 600 times per second,driven indirectly by muscles which vibrate the thorax.[9][10] Mosquitoes are mainly small flies; the largest are in the genusToxorhynchites, at up to 18 mm (0.71 in) in length and 24 mm (0.94 in) in wingspan.[11] Those in the genusAedes are much smaller, with a wingspan of 2.8 to 4.4 mm (0.11 to 0.17 in).[12]
Mosquitoes can develop from egg to adult in hot weather in as few as five days, but it may take up to a month.[13] At dawn or dusk, within days of pupating, males assemble inswarms, mating when females fly in.[14] The female mates only once in her lifetime, attracted by the pheromones emitted by the male.[15][16] As a species that need blood for the eggs to develop, the female finds a host and drinks a full meal of blood. She then rests for two or three days to digest the meal and allow her eggs to develop. She is then ready to lay the eggs and repeat the cycle of feeding and laying.[14] Females can live for up to three weeks in the wild, depending on temperature, humidity, their ability to obtain a blood meal, and avoiding being killed by their vertebrate hosts.[14][17]
Anatomy of an adult female mosquito
Adult yellow fever mosquitoAedes aegypti, typical of subfamilyCulicinae. Male (left) has bushy antennae and longerpalps than female (right)
Eggs
The eggs of most mosquitoes are laid in stagnant water, which may be a pond, a marsh, a temporary puddle, a water-filled hole in a tree, or the water-trapping leaf axils of abromeliad. Some lay near the water's edge while others attach their eggs to aquatic plants. A few, likeOpifex fuscus, can breed in salt-marshes.[5]Wyeomyia smithii breeds in the pitchers ofpitcher plants, its larvae feeding on decaying insects that have drowned there.[18]
Oviposition, egg-laying, varies between species.Anopheles females fly over the water, touching down ordapping to place eggs on the surface one at a time; their eggs are roughly cigar-shaped and have floats down their sides. A female can lay 100–200 eggs in her lifetime.[14]Aedes females drop their eggs singly, on damp mud or other surfaces near water; their eggs hatch only when they are flooded.[19] Females in genera such asCulex,Culiseta, andUranotaenia lay their eggs in floating rafts.[20][21]Mansonia females in contrast lay their eggs in arrays, attached usually to the under-surfaces of waterlily pads.[22]
Clutches of eggs of most mosquito species hatch simultaneously, butAedes eggs in diapause hatch irregularly over an extended period.[19]
The mosquito larva's head has prominent mouth brushes used for feeding, a largethorax with no legs, and a segmentedabdomen. It breathes air through a siphon on its abdomen, so must come to the surface frequently. It spends most of its time feeding onalgae, bacteria, and other microbes in the water's surface layer. It dives below the surface when disturbed. It swims either by propelling itself with its mouth brushes, or by jerkily wriggling its body. It develops through several stages, orinstars, molting each time, after which itmetamorphoses into apupa.[13]Aedes larvae, except when very young, can withstand drying; they go intodiapause for several months if their pond dries out.[19]
The head and thorax of thepupa are merged into acephalothorax, with the abdomen curving around beneath it. The pupa or "tumbler" can swim actively by flipping its abdomen. Like the larva, the pupa of most species must come to the surface frequently to breathe, which they do through a pair of respiratory trumpets on their cephalothoraxes. They do not feed; they pass much of their time hanging from the surface of the water by their respiratory trumpets. If alarmed, they swim downwards by flipping their abdomens in much the same way as the larvae. If undisturbed, they soon float up again. The adult emerges from the pupa at the surface of the water and flies off.[13]
Mosquito pupae, shortly before the adults emerged. The head and thorax are fused into the cephalothorax.
Both male and female mosquitoes feed onnectar, aphid honeydew, and plant juices,[17] but in many species the females are alsoblood-suckingectoparasites. In some of those species, a blood meal is essential for egg production; in others, it just enables the female to lay more eggs.[23] Both plant materials and blood are useful sources of energy in the form of sugars. Blood supplies more concentrated nutrients, such aslipids, but the main function of blood meals is to obtain proteins for egg production.[24][25] Mosquitoes likeToxorhynchites reproduce autogenously, not needing blood meals. Disease vector mosquitoes likeAnopheles andAedes areanautogenous, requiring blood to lay eggs. ManyCulex species are partially anautogenous, needing blood only for their second and subsequent clutches of eggs.[26]
Host animals
Blood-sucking mosquitoes favour particular host species, though they are less selective when food is short. Different mosquito species favoramphibians,reptiles includingsnakes,birds, andmammals. For example,Culiseta melanura sucks the blood ofpasserine birds, but as mosquito numbers rise they attack mammals including horses and humans, causing epidemics ofEastern equine encephalitis virus in North America.[27] Loss of blood from many bites can add up to a large volume, occasionally causing the death oflivestock as large ascattle andhorses.[28]Malaria-transmitting mosquitoes seek outcaterpillars and feed on their haemolymph,[29] impeding their development.[30]
Feeding on a snake
Feeding on a frog
Feeding on a bird
Finding hosts
Blood-feeding female mosquitoes find their hosts using multiple cues, including exhaledcarbon dioxide, heat, and many differentodorants.
Most mosquito species arecrepuscular, feeding at dawn or dusk, and resting in a cool place through the heat of the day.[31] Some species, such as theAsian tiger mosquito, are known to fly and feed during daytime.[32] Female mosquitoes hunt for hosts by smelling substances such ascarbon dioxide (CO2) and1-octen-3-ol (mushroom alcohol, found in exhaled breath) produced from the host, and through visual recognition.[33] Thesemiochemical that most strongly attractsCulex quinquefasciatus isnonanal.[34] Another attractant issulcatone.[35] A large part of the mosquito's sense of smell, or olfactory system, is devoted to sniffing out blood sources. Of 72 types of odor receptors on its antennae, at least 27 are tuned to detect chemicals found in perspiration.[36] InAedes, the search for a host takes place in two phases. First, the mosquito flies about until it detects a host's odorants; then it flies towards them, using the concentration of odorants as its guide.[37] Mosquitoes prefer to feed on people withtype O blood, an abundance of skin bacteria, high body heat, and pregnant women.[38][39] Individuals' attractiveness to mosquitoes has aheritable, genetically controlled component.[40]
The multitude of characteristics in a host observed by the mosquito allows it to select a host to feed on. This occurs when a mosquito notes the presence of CO2, as it then activates odour and visual search behaviours that it otherwise would not use. In terms of a mosquito’s olfactory system, chemical analysis has revealed that people who are highly attractive to mosquitoes produce significantly morecarboxylic acids.[41] A human's unique body odour indicates that the target is actually a human host rather than some other living warm-blooded animal (as the presence of CO2 shows). Body odour, composed ofvolatile organic compounds emitted from the skin of humans, is the most important cue used by mosquitoes.[42] Variation in skin odour is caused by body weight, hormones, genetic factors, and metabolic or genetic disorders. Infections such as malaria can influence an individual’s body odour. People infected by malaria produce relatively large amounts ofPlasmodium-induced aldehydes in the skin, creating large cues for mosquitoes as it increases the attractiveness of an odour blend, imitating a "healthy" human odour. Infected individuals produce larger amounts of aldehydesheptanal,octanal, andnonanal. These compounds are detected by mosquito antennae. Thus, people infected with malaria are more prone to mosquito biting.[43]
Contributing to a mosquito's ability to activate search behaviours, a mosquito's visual search system includes sensitivity to wavelengths from different colours. Mosquitoes are attracted to longer wavelengths, correlated to the colours of red and orange as seen by humans, and range through the spectrum of human skin tones. In addition, they have a strong attraction to dark, high-contrast objects, because of how longer wavelengths are perceived against a lighter-coloured background.[44]
Scanning electron microscope image of the Labium tip of Culex mosquito
Different species of mosquitoes have evolved different methods of identifying target hosts. Study of a domestic form and an animal-biting form of the mosquitoAedes aegypti showed that the evolution of preference for human odour is linked to increases in the expression of theolfactory receptor AaegOr4. This recognises a compound present at high levels in human odour calledsulcatone. However, the malaria mosquitoAnopheles gambiae also has OR4 genes strongly activated by sulcatone, yet none of them are closely related to AaegOr4, suggesting that the two species have evolved to specialise in biting humans independently.[44]
Female mosquito mouthparts are highly adapted to piercing skin and sucking blood. Males only drink sugary fluids, and have less specialized mouthparts.[45]
Externally, the most obvious feeding structure of the mosquito is the proboscis, composed of thelabium, U-shaped in section like arain gutter, which sheaths a bundle (fascicle) of six piercing mouthparts or stylets. These are twomandibles, twomaxillae, thehypopharynx, and thelabrum. The labium bends back into a bow when the mosquito begins to bite, staying in contact with the skin and guiding the stylets downwards. The extremely sharp tips of the labrum and maxillae are moved backwards and forwards to saw their way into the skin, with just one thousandth of the force that would be needed to penetrate the skin with a needle, resulting in a painless insertion.[46][47][48]
Evolution of mosquito mouthparts, withgrasshopper mouthparts (shown bothin situ and separately) representing a more primitive condition. All the mouthparts except the labium are stylets, formed into a fascicle or bundle.
Mouthparts of a female mosquito while feeding on blood, showing the flexiblelabium sheath supporting the piercing and sucking tube which penetrates the host's skin
AnAnopheles stephensi female is engorged with blood and beginning to pass unwanted liquid fractions to make room in its gut for more of the solid nutrients.
Females of many blood-feeding species need a blood meal to begin the process of egg development. A sufficiently large blood meal triggers a hormonal cascade that leads to egg development.[62] Upon completion of feeding, the mosquito withdraws herproboscis, and as the gut fills up, the stomach lining secretes aperitrophic membrane that surrounds the blood. This keeps the blood separate from anything else in the stomach. Like manyHemiptera that survive on dilute liquid diets, many adult mosquitoes excrete surplus liquid even when feeding. This permits females to accumulate a full meal of nutrient solids. The blood meal is digested over a period of several days.[63] Once blood is in the stomach, the midgut synthesizesprotease enzymes, primarilytrypsin assisted byaminopeptidase, that hydrolyze the bloodproteins into freeamino acids. These are used in the synthesis ofvitellogenin, which in turn is made into egg yolk protein.[64]
Distribution
Cosmopolitan
Mosquitoes have acosmopolitan distribution, occurring in every land region except Antarctica and a few islands with polar orsubpolar climates, such asIceland, which is essentially free of mosquitoes.[65] This absence is probably caused by Iceland's climate. Its weather is unpredictable, freezing but often warming suddenly in mid-winter, making mosquitoes emerge from pupae in diapause, and then freezing again before they can complete their life cycle.[66][67]
Eggs oftemperate zone mosquitoes are more tolerant of cold than the eggs of species indigenous to warmer regions.[68][69] Many can tolerate subzero temperatures, while adults of some species can survive winter by sheltering in microhabitats such as buildings or hollow trees.[70] In warm and humid tropical regions, some mosquito species are active for the entire year, but in temperate and cold regions they hibernate or enterdiapause.Arctic or subarctic mosquitoes, like some other arctic midges in families such asSimuliidae andCeratopogonidae may be active for only a few weeks annually as melt-water pools form on the permafrost. During that time, though, they emerge in huge numbers in some regions; a swarm may take up to 300 ml of blood per day from each animal in acaribou herd.[71]
Effect of climate change
For a mosquito to transmit disease, there must be favorable seasonal conditions,[72] primarily humidity, temperature, and precipitation.[73]El Niño affects the location and number of outbreaks in East Africa, Latin America, Southeast Asia andIndia.Climate change impacts the seasonal factors and in turn the dispersal of mosquitoes.[74] Climate models can use historic data to recreate past outbreaks and to predict the risk of vector-borne disease, based on an area's forecasted climate.[75]Mosquito-borne diseases have long been most prevalent in East Africa, Latin America, Southeast Asia, andIndia. An emergence in Europe was observed early in the 21st century. It is predicted that by 2030, the climate of southern Great Britain will be suitable for transmission ofPlasmodium vivax malaria byAnopheles mosquitoes for two months of the year, and that by 2080, the same will be true for southern Scotland.[76][77]Dengue fever, too, is spreading northwards with climate change. The vector, the Asian tiger mosquitoAedes albopictus, has by 2023 established across southern Europe and as far north as much of northern France, Belgium, Holland, and bothKent and West London in England.[78]
Emerging adults are consumed at the pond surface by predatory flies includingEmpididae andDolichopodidae, and byspiders. Flying adults are captured by dragonflies and damselflies, by birds such asswifts andswallows, and by vertebrates includingbats.[80]
Several flowers including members of theAsteraceae,Rosaceae andOrchidaceae arepollinated by mosquitoes, which visit to obtain sugar-richnectar. They are attracted to flowers by a range of semiochemicals such as alcohols, aldehydes, ketones, and terpenes. Mosquitoes have visited and pollinated flowers since theCretaceous period. It is possible that plant-suckingexapted mosquitoes to blood-sucking.[17]
Ecologically, blood-feeding mosquitoes aremicropredators, small animals that feed on larger animals without immediately killing them. Evolutionary biologists see this as a form ofparasitism; inEdward O. Wilson's phrase "Parasites ... are predators that eat prey in units of less than one."[81] Micropredation is one of six majorevolutionarily stable strategies within parasitism. It is distinguished by leaving the host still able to reproduce, unlike the activity ofparasitic castrators orparasitoids; and having multiple hosts, unlike conventional parasites.[82][83] From this perspective, mosquitoes areectoparasites, feeding on blood from the outside of their hosts, using their piercing mouthparts, rather than entering their bodies. Unlike some other ectoparasites such asfleas andlice, mosquitoes do not remain constantly on the body of the host, but visit only to feed.[83]
A 2023 study suggested thatLibanoculex intermedius found inLebanese amber, dating to theBarremian age of the Early Cretaceous, around 125 million years ago was the oldest known mosquito.[85] However its identification as a mosquito is disputed, with other authors considering it to be achaoborid fly instead.[86] Three other unambiguous species ofCretaceous mosquito are known.Burmaculex antiquus andPriscoculex burmanicus are known fromBurmese amber from Myanmar, which dates to the earliest part of theCenomanian age of the Late Cretaceous, around 99 million years ago.[87][88]Paleoculicis minutus, is known fromCanadian amber from Alberta, Canada, which dates to theCampanian age of the Late Cretaceous, around 79 million years ago.[89]P. burmanicus has been assigned to theAnophelinae, indicating that the split between this subfamily and theCulicinae took place over 99 million years ago.[88] Molecular estimates suggest that this split occurred 197.5 million years ago, during the EarlyJurassic, but that major diversification did not take place until the Cretaceous.[90]
Over 3,600 species of mosquitoes in 112genera have beendescribed. They are traditionally divided into two subfamilies, theAnophelinae and theCulicinae, which carry different diseases. Roughly speaking, protozoal diseases like malaria are transmitted by anophelines, while viral diseases such asyellow fever anddengue fever are transmitted by culicines.[91]
The name Culicidae was introduced by the German entomologistJohann Wilhelm Meigen in his seven-volume classification published in 1818–1838.[92] Mosquito taxonomy was advanced in 1901 when the English entomologistFrederick Vincent Theobald published his 5-volume monograph on the Culicidae.[93] He had been provided with mosquito specimens sent in to theBritish Museum (Natural History) from around the world, on the 1898 instruction of theSecretary of State for the Colonies,Joseph Chamberlain, who had written that "in view of the possible connection of Malaria with mosquitoes, it is desirable to obtain exact knowledge of the different species of mosquitoes and allied insects in the various tropical colonies. I will therefore ask you ... to have collections made of the winged insects in the Colony which bite men or animals."[94]
The two subfamilies of mosquitoes areAnophelinae, containing three genera and approximately 430 species, andCulicinae, which contains 11 tribes, 108 genera and 3,046 species. Kyanne Reidenbach and colleagues analysed mosquitophylogenetics in 2009, using both nuclear DNA and morphology of 26 species. They note that Anophelinae is confirmed to be rather basal, but that the deeper parts of the tree are not well resolved.[98]
Insect repellents are applied on skin and give short-term protection against mosquito bites. The chemicalDEET repels some mosquitoes and other insects.[116] SomeCDC-recommended repellents arepicaridin,eucalyptus oil (PMD), andethyl butylacetylaminopropionate (IR3535).[117]Pyrethrum (fromChrysanthemum species, particularlyC. cinerariifolium andC. coccineum) is an effective plant-based repellent.[118]Electronic insect repellent devices that produceultrasounds intended to keep away insects (and mosquitoes) are marketed. NoEPA or university study has shown that these devices prevent humans from being bitten by a mosquito.[119]
Ancient Greek beast fables including "The Elephant and the Mosquito" and "The Bull and the Mosquito", with the general moral that the large beast does not even notice the small one, derive ultimately fromMesopotamia.[130]
Origin myths
The peoples ofSiberia haveorigin myths surrounding the mosquito. OneOstiak myth tells of a man-eating giant,Punegusse, who is killed by a hero but will not stay dead. The hero eventually burns the giant, but the ashes of the fire become mosquitoes that continue to plague mankind. Other myths from theYakuts, Goldes (Nanai people), andSamoyed have the insect arising from the ashes or fragments of some giant creature or demon. Similar tales found in Native North American myth, with the mosquito arising from the ashes of a man-eater, suggest a common origin. TheTatars of theAltai had a variant of the same myth, involving the fragments of the dead giant,Andalma-Muus, becoming mosquitoes and other insects.[131]
Lafcadio Hearn tells that in Japan, mosquitoes are seen as reincarnations of the dead, condemned by the errors of their former lives to the condition ofJiki-ketsu-gaki, or "blood-drinkingpretas".[132]
Modern era
How a Mosquito Operates (1912)
Winsor McCay's 1912 filmHow a Mosquito Operates was one of the earliest works of animation. It has been described as far ahead of its time in technical quality.[133] It depicts a giant mosquito tormenting a sleeping man.[134]
Twelve ships of theRoyal Navy have borne the name HMSMosquito or the archaic form of the name, HMSMusquito.[135]
Thede Havilland Mosquito was a high-speed aircraft manufactured between 1940 and 1950, and used in many roles.[136]
The Russian city ofBerezniki annually celebrates its mosquitoes from the 17th of July to the 20th in a "most delicious girl" competition. In the competition, the girls stand for 20 minutes in their shorts and vests, and the one who receives the most bites wins.[137]
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