Mosquito

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This article is about the insect. For other uses, see Mosquito (disambiguation).

Mosquito
Temporal range: 99–0 
Ma
 Late Cretaceous (Cenomanian) – Recent
Aedes aegypti, vector of yellow fever
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Diptera
Superfamily: Culicoidea
Family: Culicidae
Meigen, 1818[1]
Subfamilies
Diversity
112 genera

Mosquitoes, the Culicidae, are a

micropredators, small animals that parasitise larger ones by drinking their blood without immediately killing them. Medical parasitologists view mosquitoes instead as vectors of disease, carrying protozoan parasites or bacterial or viral pathogens from one host
to another.

The mosquito life cycle consists of four stages:

organic material. These larvae are important food sources for many freshwater animals, such as dragonfly nymphs, many fish, and some birds. Adult females of many species have mouthparts adapted to pierce the skin of a host and feed on blood of a wide range of vertebrate hosts, and some invertebrates, primarily other arthropods
. Some species only produce eggs after a blood meal.

The mosquito's

arboviral diseases such as yellow fever and dengue 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, and adult. The first three stages—egg, larva, and pupa—are largely aquatic,[4] the eggs usually being laid in stagnant water.[5] They hatch to become larvae, which feed, grow, and molt until they change into pupae. 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 in diapause.[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 similar

driven indirectly by muscles which vibrate the thorax.[9][10] Mosquitoes are mainly small flies; the largest are in the genus Toxorhynchites, at up to 18 mm (0.71 in) in length and 24 mm (0.94 in) in wingspan.[11] Those in the genus Aedes 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.

swarms, 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
    Anatomy of an adult female mosquito
  • Adult yellow fever mosquito Aedes aegypti, typical of subfamily Culicinae. Male (left) has bushy antennae and longer palps than female (right)
    Adult yellow fever mosquito Aedes aegypti, typical of subfamily Culicinae. Male (left) has bushy antennae and longer palps 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 a bromeliad. Some lay near the water's edge while others attach their eggs to aquatic plants. A few, like Opifex fuscus, can breed in salt-marshes.[5] Wyeomyia smithii breeds in the pitchers of pitcher 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 or dapping 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 as Culex, Culiseta, and Uranotaenia 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, but Aedes eggs in diapause hatch irregularly over an extended period.[19]

Larva

The mosquito larva's head has prominent mouth brushes used for feeding, a large

thorax with no legs, and a segmented abdomen. It breathes air through a siphon on its abdomen, so must come to the surface frequently. It spends most of its time feeding on algae, 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, or instars, molting each time, after which it metamorphoses into a pupa.[13] Aedes larvae, except when very young, can withstand drying; they go into diapause for several months if their pond dries out.[19]

Pupa

The head and thorax of the pupa are merged into a cephalothorax, 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.
    Mosquito pupae, shortly before the adults emerged. The head and thorax are fused into the cephalothorax.

Feeding by adults

Diet

Further information: Anautogeny
Female Aedes notoscriptus feeding on blood from a human arm.

Both male and female mosquitoes feed on

anautogenous, requiring blood to lay eggs. Many Culex 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 favor

Eastern equine encephalitis virus in North America.[27] Loss of blood from many bites can add up to a large volume, occasionally causing the death of livestock as large as cattle and horses.[28] Malaria-transmitting mosquitoes seek out caterpillars and feed on their haemolymph,[29] impeding their development.[30]

  • Feeding on a snake
    Feeding on a snake
  • Feeding on a frog
    Feeding on a frog
  • Feeding on a bird
    Feeding on a bird

Finding hosts

odorants
.

Most mosquito species are

heritable, 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 more carboxylic 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 of volatile 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 of Plasmodium-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 aldehydes heptanal, octanal, and nonanal. 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 mosquito Aedes aegypti showed that the evolution of preference for human odour is linked to increases in the expression of the olfactory receptor AaegOr4. This recognises a compound present at high levels in human odour called sulcatone. However, the malaria mosquito Anopheles 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]

Mouthparts

Further information: Insect mouthparts

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 the

maxillae, the hypopharynx, and the labrum. 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, with grasshopper mouthparts (shown both in situ and separately) representing a more primitive condition. All the mouthparts except the labium are stylets, formed into a fascicle or bundle.
    Evolution of mosquito mouthparts, with grasshopper mouthparts (shown both in 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 flexible labium sheath supporting the piercing and sucking tube which penetrates the host's skin
    Mouthparts of a female mosquito while feeding on blood, showing the flexible
    labium
    sheath supporting the piercing and sucking tube which penetrates the host's skin

Saliva

Mosquito saliva contains

antimicrobial agents that control bacterial growth in the sugar meal.[50]

For a mosquito to obtain a blood meal, it must circumvent its

IFN-γ production;[54][55] suppress T cell populations;[56][57][58] decrease expression of interferon−α/β, making virus infections more severe;[59][60] increase natural killer T cells in the blood; and decrease cytokine production.[61]

Egg development and blood digestion

An Anopheles 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 her proboscis, and as the gut fills up, the stomach lining secretes a peritrophic membrane that surrounds the blood. This keeps the blood separate from anything else in the stomach. Like many Hemiptera 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 synthesizes protease enzymes, primarily trypsin assisted by aminopeptidase, that hydrolyze the blood proteins into free amino acids. These are used in the synthesis of vitellogenin, which in turn is made into egg yolk protein.[64]

Distribution

Cosmopolitan

Mosquitoes have a

subpolar climates, such as Iceland, 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 of

caribou herd.[71]

Effect of climate change

For a mosquito to transmit disease, there must be favorable seasonal conditions,

El Niño affects the location and number of outbreaks in East Africa, Latin America, Southeast Asia and India. 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, and India. 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 of Plasmodium vivax malaria by Anopheles 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 mosquito Aedes albopictus, has by 2023 established across southern Europe and as far north as much of northern France, Belgium, Holland, and both Kent and West London in England.[78]

Ecology

Predators and parasites

Mosquito larvae are among the commonest animals in ponds, and they form an important food source for freshwater

giant tree frog, freshwater turtles such as the red-eared slider, and birds such as ducks.[79]

Emerging adults are consumed at the pond surface by predatory flies including Empididae and Dolichopodidae, and by spiders. Flying adults are captured by dragonflies and damselflies, by birds such as swifts and swallows, and by vertebrates including bats.[80]

Mosquitoes are parasitised by hydrachnid mites, ciliates such as Glaucoma, microsporidians such as Thelania, and fungi including species of Saprolegniaceae and Entomophthoraceae.[80]

Pollination

A mosquito visiting a marigold flower for nectar

Several flowers including members of the

pollinated by mosquitoes, which visit to obtain sugar-rich nectar. They are attracted to flowers by a range of semiochemicals such as alcohols, aldehydes, ketones, and terpenes. Mosquitoes have visited and pollinated flowers since the Cretaceous period. It is possible that plant-sucking exapted mosquitoes to blood-sucking.[17]

Parasitism

Further information: Parasitism

Ecologically, blood-feeding mosquitoes are

ectoparasites, feeding on blood from the outside of their hosts, using their piercing mouthparts, rather than entering their bodies. Unlike some other ectoparasites such as fleas and lice, mosquitoes do not remain constantly on the body of the host, but visit only to feed.[83]

Evolution

Fossil record

Fossilized mosquito encased in amber
Culex malariager mosquito infected with the malarial parasite Plasmodium dominicana, in Dominican amber of Miocene age, 15–20 million years ago[84]

A 2023 study suggested that

Anophelinae, indicating that the split between this subfamily and the Culicinae took place over 99 million years ago.[88] Molecular estimates suggest that this split occurred 197.5 million years ago, during the Early Jurassic, but that major diversification did not take place until the Cretaceous.[90]

Taxonomy

Further information: List of mosquito genera

Over 3,600 species of mosquitoes in 112

described. They are traditionally divided into two subfamilies, the Anophelinae and the Culicinae, which carry different diseases. Roughly speaking, protozoal diseases like malaria are transmitted by anophelines, while viral diseases such as yellow fever and dengue fever are transmitted by culicines.[91]

The name Culicidae was introduced by the German entomologist

British Museum (Natural History) from around the world, on the 1898 instruction of the Secretary 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]

Phylogeny

External

Mosquitoes are members of a family of the true flies (order Diptera): the Culicidae (from the Latin culex, genitive culicis, meaning "midge" or "gnat").[95] They are members of the infraorder Culicomorpha and superfamily Culicoidea. The phylogenetic tree is based on the FLYTREE project.[96][97]

Diptera

Ptychopteromorpha (phantom and primitive crane-flies)

Culicomorpha

Chironomidae (non-biting midges)

Simulioidea (blackflies and biting midges)

Culicoidea

Dixidae (meniscus midges)

Corethrellidae (frog-biting midges)

Chaoboridae (phantom midges)

Culicidae

other midges and gnats

all other flies, inc. Brachycera

(true flies)

Internal

The two subfamilies of mosquitoes are

Anophelinae, containing three genera and approximately 430 species, and Culicinae, which contains 11 tribes, 108 genera and 3,046 species. Kyanne Reidenbach and colleagues analysed mosquito phylogenetics 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]

Culicidae

basal spp.

Anophelinae

Culicinae

other spp.

Aedini

other spp.

Sabethini

Interactions with humans

Anopheles albimanus feeding on a human arm. As mosquitoes are the only vectors of malaria, controlling them reduces its incidence.

Vectors of disease

Main article: Mosquito-borne disease

Mosquitoes are

elephantiasis, is spread by a wide variety of mosquitoes.[102] A bacterial disease spread by Culex and Culiseta mosquitoes is tularemia.[103]

Control

Main article: Mosquito control
Mosquito nets can prevent people being bitten while they sleep.

Many measures have been tried for

copepods,[110] dragonfly nymphs and adults, and some species of lizard and gecko.[111] Another approach is to introduce large numbers of sterile males.[112] Genetic modification methods including cytoplasmic incompatibility, chromosomal translocations, sex distortion and gene replacement, solutions seen as inexpensive and not subject to vector resistance, have been explored.[113] Control of disease-carrying mosquitoes using gene drives has been proposed.[114][115]

Repellents

Main article: Insect repellent
Mosquito repellents (including a mosquito coil) in a Finnish store

Insect repellents are applied on skin and give short-term protection against mosquito bites. The chemical

Electronic insect repellent devices that produce ultrasounds intended to keep away insects (and mosquitoes) are marketed. No EPA or university study has shown that these devices prevent humans from being bitten by a mosquito.[119]

Bites

Further information:
Mosquito bite allergies

Mosquito bites lead to a variety of skin reactions and more seriously to

Aqueous ammonia (3.6%) also provides relief.[127] Both topical heat[128] and cold may be useful as treatments.[129]

In human culture

Greek mythology

The Bull and the Mosquito
", 1912

The Bull and the Mosquito", with the general moral that the large beast does not even notice the small one, derive ultimately from Mesopotamia.[130]

Origin myths

The peoples of

Ostiak 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 the Yakuts, Goldes (Nanai people), and Samoyed 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. The Tatars of the Altai 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 of Jiki-ketsu-gaki, or "blood-drinking pretas".[132]

Modern era

How a Mosquito Operates (1912)

Winsor McCay's 1912 film How 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 the Royal Navy have borne the name HMS Mosquito or the archaic form of the name, HMS Musquito.[135]

The de Havilland Mosquito was a high-speed aircraft manufactured between 1940 and 1950, and used in many roles.[136]

The Russian city of Berezniki 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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Further reading

  • Winegard, Timothy Charles (2019). The mosquito: a human history of our deadliest predator. Penguin Random House.
    OCLC 1111638283
    .
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