View source for Mosquito

{{good article}}
{{Short description|Family of flies}}
{{About|the insect|}}
{{pp-semi-indef|small=yes}}
{{use British English|date=February 2024}}
{{use dmy dates|date=February 2024}}
{{Automatic taxobox
| fossil_range = {{fossilrange |99|0|earliest=125}} <small>[[Late Cretaceous]] ([[Cenomanian]]) – Recent</small>
| image = Aedes aegypti.jpg
| image_caption = ''[[Aedes aegypti]]'', [[Disease vector|vector]] of [[yellow fever]]
| taxon = Culicidae
| authority = [[Johann Wilhelm Meigen|Meigen]], 1818<ref name=mosqInv>{{cite web |url=https://mosquito-taxonomic-inventory.myspecies.info/family-culicidae-meigen-1818 |title=Family Culicidae Meigen, 1818 |date=November 2, 2008 |author=Harbach, Ralph |work=Mosquito Taxonomic Inventory |access-date=March 15, 2022 |archive-date=October 3, 2022 |archive-url=https://web.archive.org/web/20221003061320/https://mosquito-taxonomic-inventory.myspecies.info/family-culicidae-meigen-1818 |url-status=live }}, see also [https://mosquito-taxonomic-inventory.myspecies.info/valid-species-list Valid Species List] {{Webarchive |url=https://web.archive.org/web/20220315103505/https://mosquito-taxonomic-inventory.myspecies.info/valid-species-list |date=2022-03-15 }}</ref>
| subdivision_ranks = Subfamilies
| subdivision = *[[List of mosquito genera#Subfamily Anophelinae|Anophelinae]]
*[[Culicinae]]
| diversity = 112 genera
| diversity_link = List of mosquito genera <!--Culicidae#Systematics-->
}}
'''Mosquitoes''', the '''Culicidae''', are a [[Family (biology)|family]] of small [[Diptera|flies]] consisting of 3,600 [[species]]. The word ''mosquito'' (formed by ''[[Musca (fly)|mosca]]'' and [[diminutive]] ''-ito'')<ref>{{cite web |url=http://dle.rae.es/?id=PuiMZGa |title=mosquito |publisher=[[Real Academia Española]] |access-date=24 July 2016 |archive-date=24 July 2016 |archive-url=https://web.archive.org/web/20160724225319/http://dle.rae.es/?id=PuiMZGa |url-status=live }}</ref> is Spanish and Portuguese for ''little fly''.<ref name="isbn0-19-861271-0">{{cite book |last=Brown |first=Lesley |title=The New Shorter Oxford English Dictionary on Historical Principles |publisher=[[Clarendon Press]] |location=Oxford, England |year=1993 |isbn=978-0-19-861271-1 |url=https://archive.org/details/newshorteroxford00lesl }}</ref> Mosquitoes have a slender [[segmented body]], one pair of wings, three pairs of long hair-like legs, and specialized, highly elongated, [[piercing-sucking mouthpart]]s. All mosquitoes drink [[nectar]] from [[flower]]s; females of some species have in addition adapted to drink blood. The group diversified during the [[Cretaceous]] period. [[Evolutionary biology|Evolutionary biologists]] view mosquitoes as [[micropredator]]s, small animals that [[Parasitism|parasitise]] larger ones by drinking their blood without immediately killing them. [[Parasitology|Medical parasitologists]] view mosquitoes instead as [[Disease vector|vectors of disease]], carrying [[protozoa]]n parasites or [[bacteria]]l or [[virus|viral]] [[pathogen]]s from one [[Host (biology)|host]] to another.
The mosquito life cycle consists of four stages: [[egg]], [[larva]], [[pupa]], and [[adult]]. Eggs are laid on the water surface; they hatch into [[Motility|motile]] larvae that feed on aquatic [[algae]] and [[organic material]]. These larvae are important food sources for many freshwater animals, such as [[Dragonfly#Life cycle|dragonfly nymphs]], many fish, and some birds. Adult females of many species have mouthparts adapted to pierce the skin of a [[Host (biology)|host]] and [[Hematophagy|feed on blood]] of a wide range of [[vertebrate]] hosts, and some [[invertebrate]]s, primarily other [[arthropod]]s. Some species only produce eggs after a blood meal.
The mosquito's [[saliva]] is transferred to the host during the bite, and can cause an itchy [[rash]]. In addition, blood-feeding species can ingest pathogens while biting, and transmit them to other hosts. Those species include vectors of [[parasitic disease]]s such as [[malaria]] and [[filariasis]]<!--NOT A LIST-->, and [[arboviruses|arboviral]] diseases such as [[yellow fever]] and [[dengue fever]]<!--NOT A LIST-->. By transmitting diseases, mosquitoes cause the deaths of over 725,000 people each year.
== Description and life cycle ==
[[File:Mosquito Life-Cycle.svg|frameless|right|upright=1.2]]
Like all flies, mosquitoes go through four stages in their life cycles: [[egg (biology)|egg]], [[larva]], [[pupa]], and [[imago|adult]]. The first three stages—egg, larva, and pupa—are largely aquatic,<ref>{{cite web |title=FAQs |publisher=[[American Mosquito Control Association|AMCA]] |url=https://www.mosquito.org/page/faq |url-status=live |archive-url=https://web.archive.org/web/20190716212718/https://www.mosquito.org/page/FAQ |archive-date=2019-07-16 }}</ref> the eggs usually being laid in stagnant water.<ref name="Wigglesworth 1933">{{cite journal |last=Wigglesworth |first=Vincent B. |author-link=Vincent Wigglesworth |year=1933 |journal=Journal of Experimental Biology |volume=10 |pages=27–36 |title=The Adaptation of Mosquito Larvae to Salt Water |url=http://jeb.biologists.org/content/10/1/27 |issue=1 |doi=10.1242/jeb.10.1.27 |doi-access=free |access-date=2013-04-01 |archive-date=2014-06-24 |archive-url=https://web.archive.org/web/20140624025943/http://jeb.biologists.org/content/10/1/27 |url-status=live}}</ref> They hatch to become [[larva]]e, which feed, grow, and molt until they change into [[pupa]]e. 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]].<ref>Kosova, Jonida (2003) [http://digitalcommons.unf.edu/ojii_volumes/94 "Longevity Studies of Sindbis Virus Infected ''Aedes Albopictus''"] {{Webarchive |url=https://web.archive.org/web/20120425162236/http://digitalcommons.unf.edu/ojii_volumes/94/ |date=2012-04-25 }}. All Volumes (2001–2008). Paper 94.</ref>
=== 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&nbsp;mm long. When at rest, mosquitoes hold their first pair of legs outwards, whereas the somewhat similar [[Chironomidae|Chironomid]] midges hold these legs forwards.<ref>{{Cite web |url=https://nature.mdc.mo.gov/discover-nature/field-guide/midges |title=Midges |website=MDC Discover Nature |access-date=2019-11-19 |archive-date=2019-10-26 |archive-url=https://web.archive.org/web/20191026055541/https://nature.mdc.mo.gov/discover-nature/field-guide/midges |url-status=live}}</ref> ''[[Anopheles]]'' mosquitoes can fly for up to four hours continuously at {{cvt|1|to|2|km/h}},<ref>{{cite journal |last1=Kaufmann |first1=C. |last2=Briegel |first2=H. |title=Flight performance of the malaria vectors Anopheles gambiae and Anopheles atroparvus |journal=Journal of Vector Ecology |volume=29 |issue=1 |pages=140–153 |date=June 2004 |pmid=15266751 |url=http://www.sove.org/Journal%20PDF/June%202004/Kaufmann.pdf |archive-url=https://web.archive.org/web/20110728043927/http://www.sove.org/Journal%20PDF/June%202004/Kaufmann.pdf |archive-date=2011-07-28 }}</ref> traveling up to {{cvt|12|km}} in a night. Males beat their wings between 450 and 600 times per second, [[Indirect flight|driven indirectly]] by muscles which vibrate the thorax.<ref>{{cite web |first=Diana |last=Leung |url=https://hypertextbook.com/facts/2000/DianaLeung.shtml |title=Frequency of mosquito wings |year=2000 |website=The Physics Factbook |editor-last=Elert |editor-first=Glenn |accessdate=2022-01-24 |archive-date=2022-01-25 |archive-url=https://web.archive.org/web/20220125000341/https://hypertextbook.com/facts/2000/DianaLeung.shtml |url-status=live}}</ref><ref>{{cite journal |last=Smith |first=David S. |title=Flight muscles of insects |journal=[[Scientific American]] |volume=212 |pages=76–88 |year=1965 |issue=6 |doi=10.1038/scientificamerican0665-76 |pmid=14327957 |bibcode=1965SciAm.212f..76S }}</ref> Mosquitoes are mainly small flies; the largest are in the genus ''[[Toxorhynchites]]'', at up to {{cvt|18|mm}} in length and {{cvt|24|mm}} in wingspan.<ref name="Cook 2009">{{cite book |author1=Cook, G.C. |author2=Zumla, A |year=2009 |title=Manson's Tropical Diseases |publisher=Saunders Elsevier |page=1735 |edition=22 |isbn=978-1-4160-4470-3}}</ref> Those in the genus ''[[Aedes]]'' are much smaller, with a wingspan of {{cvt|2.8|to|4.4|mm}}.<ref name="UF/IFAS A. gambiae">{{cite web |title=African malaria mosquito |url=https://entnemdept.ufl.edu/creatures/aquatic/Anopheles_gambiae.htm |publisher=[[University of Florida]] |access-date=11 February 2024}}</ref>
Mosquitoes can develop from egg to adult in hot weather in as few as five days, but it may take up to a month.<ref name="EPA life cycle"/> At dawn or dusk, within days of pupating, males assemble in [[swarm]]s, mating when females fly in.<ref name="CDC Anopheles"/><!-- Males live for about 5–7 days. --> The female mates only once in her lifetime, attracted by the pheromones emitted by the male.<ref>{{cite web |url=https://www.timeshighereducation.com/hub/university-witwatersrand/p/male-mosquito-odours-reveal-how-mozzies-mate |title=Male mosquito odours reveal how mozzies mate |publisher=[[University of the Witwatersrand]] |date=5 August 2020 |access-date=25 January 2024 |archive-date=25 January 2024 |archive-url=https://web.archive.org/web/20240125141528/https://www.timeshighereducation.com/hub/university-witwatersrand/p/male-mosquito-odours-reveal-how-mozzies-mate |url-status=live }}</ref><ref>{{cite journal |last1=Mozūraitis |first1=R. |last2=Hajkazemian |first2=M. |last3=Zawada |first3=J.W. |display-authors=etal |title=Male swarming aggregation pheromones increase female attraction and mating success among multiple African malaria vector mosquito species |journal=Nature Ecology & Evolution |volume=4 |issue=10 |pages=1395–1401 |date=3 August 2020 |doi=10.1038/s41559-020-1264-9 |pmid=32747772|bibcode=2020NatEE...4.1395M |s2cid=220948478 |url=https://ora.ox.ac.uk/objects/uuid:e28af752-35bf-4ed0-b0c6-7d612b3e44e3 }}</ref> 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.<ref name="CDC Anopheles"/> 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.<ref name="CDC Anopheles"/><ref name="Peach-Gries-2020"/>
<gallery class=center mode="nolines" heights="450" widths="300">
File:Culex pipiens diagram en.svg|Anatomy of an adult female mosquito
File:Aedes aegypti E-A-Goeldi 1905.jpg|Adult yellow fever mosquito ''[[Aedes aegypti]]'', typical of subfamily [[Culicinae]]. Male (left) has bushy antennae and longer [[Insect mouthparts|palps]] than female (right)
</gallery>
=== 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 [[Bromeliaceae|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.<ref name="Wigglesworth 1933"/> ''[[Wyeomyia smithii]]'' breeds in the pitchers of [[pitcher plant]]s, its larvae feeding on decaying insects that have drowned there.<ref>Crans, Wayne J.; [http://www.rci.rutgers.edu/~insects/sp27.htm ''Wyeomyia smithii'' (Coquillett)] {{webarchive |url=https://web.archive.org/web/20130605173233/http://www.rci.rutgers.edu/~insects/sp27.htm |date=2013-06-05 }}. Rutgers University, Center for Vector Biology.</ref>
[[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.<ref name="CDC Anopheles">{{cite web |title=Anopheles Mosquitoes |publisher=[[Centers for Disease Control and Prevention]] |url=https://www.cdc.gov/malaria/about/biology/mosquitoes/ |access-date=December 13, 2023 |date=July 16, 2020 |archive-date=18 May 2012 |archive-url=https://web.archive.org/web/20120518210447/http://www.cdc.gov/malaria/about/biology/mosquitoes/ |url-status=live }}</ref> ''Aedes'' females drop their eggs singly, on damp mud or other surfaces near water; their eggs hatch only when they are flooded.<ref name="Huang Walker Vulule Miller 2006"/> Females in genera such as ''Culex'', ''[[Culiseta]]'', and ''[[Uranotaenia]]'' lay their eggs in floating rafts.<ref>{{cite book |last1=Gullan |first1=P. J. |last2=Cranston |first2=P. S. |title=The Insects: An Outline of Entomology |publisher=[[Wiley-Blackwell]] |location=Oxford |year=2014 |edition=5th |isbn=978-1-118-84616-2 |url=https://books.google.com/books?id=lF5hBAAAQBAJ |page=280 |access-date=14 December 2023 |archive-date=11 December 2023 |archive-url=https://web.archive.org/web/20231211165406/https://books.google.com/books?id=lF5hBAAAQBAJ |url-status=live }}</ref><ref name="Spielman, D'Antonio">{{cite book |last1=Spielman |first1=Andrew |last2=D'Antonio |first2=M. |chapter=Part One: Magnificent Enemy |title=Mosquito: a natural history of our most persistent and deadly foe |publisher=[[Hyperion (publisher)]] |location=New York |year=2001 |isbn=978-0-7868-6781-3 |url=https://archive.org/details/mosquitonaturalh00spie }}</ref> ''[[Mansonia (fly)|Mansonia]]'' females in contrast lay their eggs in arrays, attached usually to the under-surfaces of waterlily pads.<ref name="Amorim Sa Rojas Santos Neto 2022 pp. 631–637">{{cite journal |last1=Amorim |first1=J. A. |last2=Sa |first2=I. L. R. |last3=Rojas |first3=M. V. R. |last4=Santos Neto |first4=N. F. |last5=Galardo |first5=A. K. R. |last6=Carvalho |first6=D. P. |last7=Ribeiro |first7=K. A. N. |last8=Sallum |first8=M. A. M. |display-authors=5 |title=Aquatic Macrophytes Hosting Immature Mansonia (Mansonia) Blanchard, 1901 (Diptera, Culicidae) in Porto Velho, Rondonia State, Brazil |journal=Journal of Medical Entomology |volume=59 |issue=2 |date=2022-03-16 |doi=10.1093/jme/tjab223 |pages=631–637|pmid=35043213 }}</ref>
Clutches of eggs of most mosquito species hatch simultaneously, but ''Aedes'' eggs in diapause hatch irregularly over an extended period.<ref name="Huang Walker Vulule Miller 2006">{{cite journal |last1=Huang |first1=Juan |last2=Walker |first2=Edward D |last3=Vulule |first3=John |last4=Miller |first4=James R. |title=Daily temperature profiles in and around Western Kenyan larval habitats of Anopheles gambiae as related to egg mortality |journal=Malaria Journal |volume=5 |issue=1 |date=2006 |page=87 |pmid=17038186 |pmc=1617108 |doi=10.1186/1475-2875-5-87 |doi-access=free }}</ref>
<gallery mode="packed">
File:Anopheles_egg_2_(cropped).jpg|''[[Anopheles]]'' eggs with side floats
File:Mosquito egg SEM.jpg|[[Scanning electron microscope|Electron micrograph]] of a culicine egg
File:Gelege1 (cropped).jpg|''Culex'' egg raft
</gallery>
=== Larva ===
The mosquito larva's head has prominent mouth brushes used for feeding, a large [[Thorax (insect anatomy)|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 [[instar]]s, molting each time, after which it [[metamorphosis|metamorphoses]] into a [[pupa]].<ref name="EPA life cycle">{{cite web |title=Mosquito Life Cycle |date=21 February 2013 |url=https://www.epa.gov/mosquitocontrol/mosquito-life-cycle |publisher=[[Environmental Protection Agency]] |access-date=12 December 2023 |archive-date=19 December 2020 |archive-url=https://web.archive.org/web/20201219184319/https://www.epa.gov/mosquitocontrol/mosquito-life-cycle |url-status=live }}</ref> ''Aedes'' larvae, except when very young, can withstand drying; they go into [[diapause]] for several months if their pond dries out.<ref name="Huang Walker Vulule Miller 2006"/>
<gallery class="center" mode="nolines" heights=200 widths=200>
File:AnophelesLarvaPhoto CDCHarryWeinburgh publicdomain.jpg|''[[Anopheles]]'' larva
File:Culex restuans larva diagram en.svg|Anatomy of a ''[[Culex]]'' larva
File:Culex sp larvae.png|''Culex'' larvae plus one pupa
</gallery>
=== 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.<ref name="EPA life cycle"/>
<gallery class="center" mode="nolines" heights=200 widths=400>
File:Underwater view of mosquito pupae in standing water.jpg|Mosquito pupae, shortly before the adults emerged. The head and thorax are fused into the cephalothorax.
</gallery>
== Feeding by adults ==
=== Diet ===
{{further|Anautogeny}}
[[File:Mosquito Tasmania crop.jpg|thumb|Female ''[[Ochlerotatus notoscriptus]]'' feeding on blood from a human arm. ]]
Both male and female mosquitoes feed on [[nectar source|nectar]], aphid honeydew, and plant juices,<ref name="Peach-Gries-2020">{{cite journal |last1=Peach |first1=Daniel A. H. |last2=Gries |first2=Gerhard |title=Mosquito phytophagy – sources exploited, ecological function, and evolutionary transition to haematophagy |journal=Entomologia Experimentalis et Applicata|issue=2 |doi=10.1111/eea.12852 |year=2019 |volume=168 |doi-access=free |pages=120–136 }}</ref> but in many species the females are also [[Hematophagy|blood-sucking]] [[Parasitism|ectoparasites]]. In some of those species, a blood meal is essential for egg production; in others, it just enables the female to lay more eggs.<ref name="pmid30846726">{{cite journal |title=Multimodal floral cues guide mosquitoes to tansy inflorescences |journal=Scientific Reports |date=March 2019 |volume=9 |issue=1 |page=3908 |pmid=30846726 |last1=Peach |first1=Daniel A. H. |last2=Gries |first2=R. |last3=Zhai |first3=H. |last4=Young |first4=N. |last5=Gries |first5=G. |doi=10.1038/s41598-019-39748-4 |pmc=6405845 |bibcode=2019NatSR...9.3908P }}</ref> Both plant materials and blood are useful sources of energy in the form of sugars. Blood supplies more concentrated nutrients, such as [[lipid]]s, but the main function of blood meals is to obtain proteins for egg production.<ref name="Tyagi 2004">{{cite book |last=Tyagi |first=B.K. |title=The Invincible Deadly Mosquitoes |publisher=Scientific Publishers |year=2004 |isbn=978-93-87741-30-0 |url=https://books.google.com/books?id=4g9fDwAAQBAJ&pg=PA79 |access-date=2021-04-06 |page=79 |quote=Only female mosquitoes require a blood meal (protein)...The number of egg formation and development in ovary of the female is directly dependent on quantum and nature supply of blood meal. |archive-date=2022-01-29 |archive-url=https://web.archive.org/web/20220129181746/https://books.google.com/books?id=4g9fDwAAQBAJ&pg=PA79 |url-status=live }}</ref><ref name="AMCA">{{cite web |title=Biology |url=https://www.mosquito.org/page/biology |website=mosquito.org |publisher=[[American Mosquito Control Association]] |access-date=6 April 2021 |quote=Acquiring a blood meal (protein) is essential for egg production, but mostly both male and female mosquitoes are nectar feeders for their nutrition. |archive-date=29 March 2021 |archive-url=https://web.archive.org/web/20210329163726/https://www.mosquito.org/page/biology |url-status=live }}</ref> Mosquitoes like ''[[Toxorhynchites]]'' reproduce autogenously, not needing blood meals. Disease vector mosquitoes like ''Anopheles'' and ''Aedes'' are [[anautogenous]], requiring blood to lay eggs. Many ''Culex'' species are partially anautogenous, needing blood only for their second and subsequent clutches of eggs.<ref>{{cite journal |last1=Sawabe |first1=K. |last2=Moribayashi |first2=A. |title=Lipid utilization for ovarian development in an autogenous mosquito, Culex pipiens molestus (Diptera: Culicidae) |journal=Journal of Medical Entomology |volume=37 |issue=5 |pages=726–731 |date=September 2000 |pmid=11004785 |doi=10.1603/0022-2585-37.5.726 |doi-access=free }}</ref>
=== Host animals ===
Blood-sucking mosquitoes favour particular host species, though they are less selective when food is short. Different mosquito species favor [[amphibian]]s, [[reptile]]s including [[snake]]s, [[bird]]s, and [[mammal]]s. For example, ''[[Culiseta melanura]]'' sucks the blood of [[passerine]] birds, but as mosquito numbers rise they attack mammals including horses and humans, causing epidemics of [[Eastern equine encephalitis virus]] in North America.<ref name="Lehane2005">{{cite book |last=Lehane |first=M. J. |title=The Biology of Blood-Sucking in Insects |url=https://books.google.com/books?id=ZV5EKCEGHIoC&pg=PA151 |date=9 June 2005 |publisher=[[Cambridge University Press]] |isbn=978-0-521-83608-1 |page=151 |access-date=18 February 2016 |archive-date=28 May 2016 |archive-url=https://web.archive.org/web/20160528091322/https://books.google.com/books?id=ZV5EKCEGHIoC&pg=PA151 |url-status=live}}</ref> Loss of blood from many bites can add up to a large volume, occasionally causing the death of [[livestock]] as large as [[cattle]] and [[horse]]s.<ref name="LSUAgCenter">{{cite web |title=Hurricane Laura exacerbates mosquito problems with livestock |website=[[LSU AgCenter]] |date=2020-09-09 |url=http://www.lsuagcenter.com/profiles/rbogren/articles/page1599664054105 |access-date=2022-02-26 |archive-date=2022-02-26 |archive-url=https://web.archive.org/web/20220226191729/https://www.lsuagcenter.com/profiles/rbogren/articles/page1599664054105 |url-status=live }}</ref> [[Malaria]]-transmitting mosquitoes seek out [[caterpillar]]s and feed on their haemolymph,<ref name="George Blanford Thomas Baker 2014">{{cite journal |last1=George |first1=Justin |last2=Blanford |first2=Simon |last3=Thomas |first3=Matthew B. |last4=Baker |first4=Thomas C. |title=Malaria Mosquitoes Host-Locate and Feed upon Caterpillars |journal=PLOS ONE |volume=9 |issue=11 |date=2014-11-05 |pmid=25372720 |pmc=4220911 |doi=10.1371/journal.pone.0108894 |page=e108894 |bibcode=2014PLoSO...9j8894G |doi-access=free}}</ref> impeding their development.<ref>{{cite journal |last1=Martel |first1=Véronique |last2=Schlyter |first2=Fredrik |last3=Ignell |first3=Rickard |last4=Hansson |first4=Bill S. |last5=Anderson |first5=Peter |title=Mosquito feeding affects larval behaviour and development in a moth |journal=PLOS ONE |volume=6 |issue=10 |pages=e25658 |year=2011 |pmid=21991329 |pmc=3185006 |doi=10.1371/journal.pone.0025658 |bibcode=2011PLoSO...625658M |doi-access=free }}</ref>
<gallery mode=packed heights=150>
File:Chironius scurrulus (Yasuni) (cropped) with mosquitoes.jpg|Feeding on a snake
File:Mosquitoes vs. Frog (14555480700) (cropped).jpg|Feeding on a frog
File:JJeffreyApapaneMosquito.jpg|Feeding on a bird
</gallery>
=== Finding hosts ===
[[File:How to attract mosquitoes.svg|thumb|upright=1.5|Blood-feeding female mosquitoes find their hosts using multiple cues, including exhaled [[carbon dioxide]], heat, and many different [[odorant]]s.]]
Most mosquito species are [[crepuscular]], feeding at dawn or dusk, and resting in a cool place through the heat of the day.<ref>{{cite conference |conference=Proceedings of the Eighty-Second Annual Meeting of the New Jersey Mosquito Control Association |year=1989 |pages=53–57 |last=Crans |first=Wayne J. |url=http://www.rci.rutgers.edu/~insects/restbox.htm |title=Resting boxes as mosquito surveillance tools |url-status=dead |archive-url=https://web.archive.org/web/20060720195034/http://www.rci.rutgers.edu/~insects/restbox.htm |archive-date=2006-07-20 }}</ref> Some species, such as the [[Asian tiger mosquito]], are known to fly and feed during daytime.<ref>{{cite web |last1=Maruniak |first1=James E. |title=Asian tiger mosquito |url=http://entnemdept.ufl.edu/creatures/aquatic/asian_tiger.htm |website=Featured Creatures |publisher=[[University of Florida]] |access-date=October 2, 2014 |location=[[Gainesville, Florida]] |date=July 2014 |archive-date=September 7, 2014 |archive-url=https://web.archive.org/web/20140907181247/http://entnemdept.ufl.edu/creatures/aquatic/asian_tiger.htm |url-status=live }}</ref> Female mosquitoes hunt for hosts by smelling substances such as [[carbon dioxide]] (CO<sub>2</sub>) and [[1-Octen-3-ol|1-octen-3-ol]] (mushroom alcohol, found in exhaled breath) produced from the host, and through visual recognition.<ref>{{cite journal |last1=Hallem |first1=Elissa A. |last2=Nicole Fox |first2=A. |last3=Zwiebel |first3=Laurence J. |last4=Carlson |first4=John R. |title=Olfaction: mosquito receptor for human-sweat odorant |journal=Nature |volume=427 |issue=6971 |pages=212–213 |date=January 2004 |pmid=14724626 |doi=10.1038/427212a |bibcode=2004Natur.427..212H |s2cid=4419658 }}</ref> The [[semiochemical]] that most strongly attracts ''[[Culex]] quinquefasciatus'' is [[nonanal]].<ref>{{cite magazine |url=https://www.usnews.com/science/articles/2009/10/28/scientists-identify-key-smell-that-attracts-mosquitoes-to-humans.html |title=Scientists identify key smell that attracts mosquitoes to humans |magazine=[[US News]] |date=October 28, 2009 |access-date=5 September 2017 |archive-date=10 September 2024 |archive-url=https://web.archive.org/web/20240910225722/https://www.usnews.com/science/articles/2009/10/28/scientists-identify-key-smell-that-attracts-mosquitoes-to-humans |url-status=live }}</ref> Another attractant is [[sulcatone]].<ref>{{cite news |url=https://richarddawkins.net/2014/11/scientists-have-identified-the-gene-that-makes-mosquitoes-crave-human-blood/ |title=Scientists have identified the gene that makes mosquitoes crave human blood |publisher=[[Richard Dawkins Foundation]] |date=November 21, 2014 |access-date=November 21, 2014 |archive-date=November 25, 2014 |archive-url=https://web.archive.org/web/20141125134812/https://richarddawkins.net/2014/11/scientists-have-identified-the-gene-that-makes-mosquitoes-crave-human-blood/ |url-status=live }}</ref> 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.<ref>{{cite news |url=http://www.timesonline.co.uk/tol/news/science/medicine/article7014046.ece |newspaper=The Times |location=London |title=Sweat and blood why mosquitoes pick and choose between humans |first=Hannah |last=Devlin |date=February 4, 2010 |access-date=May 13, 2010 |archive-date=October 3, 2022 |archive-url=https://web.archive.org/web/20221003061323/https://www.thetimes.co.uk/ |url-status=dead }}</ref> In ''Aedes'', 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.<ref name="Estrada">{{cite book |last1=Estrada-Franco |first1=R. G. |last2=Craig |first2=G. B. |year=1995 |title=Biology, disease relationship and control of ''Aedes albopictus'' |publisher=[[Pan American Health Organization]] |location=Washington, D.C. |series=Technical Paper No. 42}}</ref> Mosquitoes prefer to feed on people with [[type O blood]], an abundance of skin bacteria, high body heat, and pregnant women.<ref>{{cite journal |last1=Shirai |first1=Yoshikazu |last2=Funada |first2=Hisashi |last3=Takizawa |first3=Hisao |last4=Seki |first4=Taisuke |last5=Morohashi |first5=Masaaki |last6=Kamimura |first6=Kiyoshi |title=Landing preference of ''Aedes albopictus'' (Diptera: Culicidae) on human skin among ABO blood groups, secretors or nonsecretors, and ABH antigens |journal=Journal of Medical Entomology |volume=41 |issue=4 |pages=796–799 |date=July 2004 |pmid=15311477 |doi=10.1603/0022-2585-41.4.796 |doi-access=free }}</ref><ref name="Chappell, NPR 2013.07.12: Mosquitos' perceptions of humans">{{cite news |last1=Chappell |first1=Bill |title=5 Stars: A Mosquito's Idea Of A Delicious Human |url=https://www.npr.org/blogs/thetwo-way/2013/07/12/201521175/5-stars-a-mosquitos-idea-of-a-delicious-human |access-date=23 July 2021 |publisher=[[NPR]] |date=12 July 2013 |archive-date=14 October 2014 |archive-url=https://web.archive.org/web/20141014202549/http://www.npr.org/blogs/thetwo-way/2013/07/12/201521175/5-stars-a-mosquitos-idea-of-a-delicious-human |url-status=live }}</ref> Individuals' attractiveness to mosquitoes has a [[heritable]], genetically controlled component.<ref>{{cite journal  |last1=Fernández-Grandon |first1=G. Mandela |last2=Gezan |first2=Salvador A. |last3=Armour |first3=John A. L. |last4=Pickett |first4=John A. |last5=Logan |first5=James G. |title=Heritability of attractiveness to mosquitoes |journal=PLOS ONE |volume=10 |issue=4 |pages=e0122716 |date=22 April 2015 |pmid=25901606 |pmc=4406498 |doi=10.1371/journal.pone.0122716 |bibcode=2015PLoSO..1022716F |doi-access=free }}</ref>
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 CO<sub>2</sub>, 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 acid|carboxylic acids]].<ref>{{Cite journal |last1=De Obaldia |first1=Maria Elena |last2=Morita |first2=Takeshi |last3=Dedmon |first3=Laura C. |last4=Boehmler |first4=Daniel J. |last5=Jiang |first5=Caroline S. |last6=Zeledon |first6=Emely V. |last7=Cross |first7=Justin R. |last8=Vosshall |first8=Leslie B. |display-authors=3 |date=2022-10-27 |title=Differential mosquito attraction to humans is associated with skin-derived carboxylic acid levels |journal=Cell |volume=185 |issue=22 |pages=4099–4116.e13 |doi=10.1016/j.cell.2022.09.034 |pmid=36261039|pmc=10069481 }}</ref> 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 CO<sub>2</sub> shows). Body odour, composed of [[volatile organic compound]]s emitted from the skin of humans, is the most important cue used by mosquitoes.<ref>{{Cite journal |last=McBride |first=Carolyn |date=12 November 2014 |title=Evolution of mosquito preference for humans linked to an odorant receptor |journal=Nature|volume=515 |issue=7526 |pages=222–227 |doi=10.1038/nature13964 |pmid=25391959 |pmc=4286346 |bibcode=2014Natur.515..222M }}</ref> 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.<ref>{{Cite journal |last1=Robinson |first1=Ailie |last2=Busula |first2=Annette O. |last3=Voets |first3=Mirjam A. |last4=Beshir |first4=Khalid B. |last5=Caulfield |first5=John C. |last6=Powers |first6=Stephen J. |last7=Verhulst |first7=Niels O. |last8=Winskill |first8=Peter |last9=Muwanguzi |first9=Julian |last10=Birkett |first10=Michael A. |last11=Smallegange |first11=Renate C. |last12=Masiga |first12=Daniel K. |last13=Mukabana |first13=W. Richard |last14=Sauerwein |first14=Robert W. |last15=Sutherland |first15=Colin J. |display-authors=3 |date=May 2018 |title=Plasmodium -associated changes in human odor attract mosquitoes |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=115 |issue=18 |pages=E4209–E4218 |doi=10.1073/pnas.1721610115 |doi-access=free |pmc=5939094 |pmid=29666273 |bibcode=2018PNAS..115E4209R }}</ref>
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.<ref>{{Cite journal |last1=Alonso San Alberto |first1=Diego |last2=Rusch |first2=Claire |last3=Zhan |first3=Yinpeng |last4=Straw |first4=Andrew D. |last5=Montell |first5=Craig |last6=Riffell |first6=Jeffrey A. |date=2022-02-04 |title=The olfactory gating of visual preferences to human skin and visible spectra in mosquitoes |journal=[[Nature Communications]] |volume=13 |issue=1 |pages=555 |doi=10.1038/s41467-022-28195-x |pmid=35121739 |pmc=8816903 |bibcode=2022NatCo..13..555A}}</ref>
[[File:Culex mosquito proboscis.jpg|thumb|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.<ref>{{Cite journal |last1=Alonso San Alberto |first1=Diego |last2=Rusch |first2=Claire |last3=Zhan |first3=Yinpeng |last4=Straw |first4=Andrew D. |last5=Montell |first5=Craig |last6=Riffell |first6=Jeffrey A. |date=2022-02-04 |title=The olfactory gating of visual preferences to human skin and visible spectra in mosquitoes |journal=[[Nature Communications]] |volume=13 |issue=1 |pages=555 |doi=10.1038/s41467-022-28195-x |pmid=35121739 |pmc=8816903 |bibcode=2022NatCo..13..555A}}</ref>
=== Mouthparts ===
{{further|Insect mouthparts}}
Female mosquito mouthparts are highly adapted to piercing skin and sucking blood. Males only drink sugary fluids, and have less specialized mouthparts.<ref name="Wahid Sunahara Mogi 2003">{{cite journal |last1=Wahid |first1=Isra |last2=Sunahara |first2=Toshihiko |last3=Mogi |first3=Motoyoshi |title=Maxillae and Mandibles of Male Mosquitoes and Female Autogenous Mosquitoes (Diptera: Culicidae) |journal=Journal of Medical Entomology |volume=40 |issue=2 |date=2003-03-01 |doi=10.1603/0022-2585-40.2.150 |pages=150–158 |pmid=12693842 |s2cid=41524028 }}</ref>
Externally, the most obvious feeding structure of the mosquito is the proboscis, composed of the [[Labium (insect)|labium]], U-shaped in section like a [[rain gutter]], which sheaths a bundle (fascicle) of six piercing mouthparts or stylets. These are two [[Mandible (insect mouthpart)|mandibles]], two [[Galea (insects)|maxillae]], the [[Insect mouthparts#Hypopharynx|hypopharynx]], and the [[Labrum (arthropod mouthpart)|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.<ref name="NPR mouthparts">{{cite web |last1=Quirós |first1=Gabriela |title=WATCH: Mosquitoes Use 6 Needles To Suck Your Blood |url=https://www.npr.org/sections/health-shots/2016/06/07/480653821/watch-mosquitoes-use-6-needles-to-suck-your-blood |publisher=NPR |access-date=13 December 2023 |date=June 7, 2016 |archive-date=3 January 2024 |archive-url=https://web.archive.org/web/20240103011402/https://www.npr.org/sections/health-shots/2016/06/07/480653821/watch-mosquitoes-use-6-needles-to-suck-your-blood |url-status=live }}</ref><ref name="Choo Buss Tan Leal 2015">{{cite journal |last1=Choo |first1=Young-Moo |last2=Buss |first2=Garrison K. |last3=Tan |first3=Kaiming |last4=Leal |first4=Walter S. |title=Multitasking roles of mosquito labrum in oviposition and blood feeding |journal=Frontiers in Physiology |volume=6 |date=2015-10-29 |page=306 |pmid=26578978 |pmc=4625056 |doi=10.3389/fphys.2015.00306 |doi-access=free }}</ref><ref name="Zahran Sawires Hamza 2022">{{cite journal |last1=Zahran |first1=Nagwan |last2=Sawires |first2=Sameh |last3=Hamza |first3=Ali |title=Piercing and sucking mouth parts sensilla of irradiated mosquito, Culex pipiens (Diptera: Culicidae) with gamma radiation |journal=Scientific Reports |volume=12 |issue=1 |date=2022-10-25 |page=17833 |pmid=36284127 |pmc=9596698 |doi=10.1038/s41598-022-22348-0 |bibcode=2022NatSR..1217833Z }}</ref>
<gallery class=center mode=nolines heights=275 widths=350>
File:Evolution of mosquito mouthparts.svg|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.
File:Feeding mosquito, mouthparts labelled.svg|Mouthparts of a female mosquito while feeding on blood, showing the flexible  [[Labium (insect)|labium]] sheath supporting the piercing and sucking tube which penetrates the host's skin
</gallery>
=== Saliva ===
Mosquito saliva contains [[enzyme]]s that aid in sugar feeding,<ref>{{cite journal |last1=Grossman |first1=G. L. |last2=James |first2=A. A. |title=The salivary glands of the vector mosquito, ''Aedes aegypti'', express a novel member of the amylase gene family |journal=Insect Molecular Biology |volume=1 |issue=4 |pages=223–232 |year=1993 |pmid=7505701 |doi=10.1111/j.1365-2583.1993.tb00095.x |s2cid=13019630 }}</ref> and [[antimicrobial agents]] that control bacterial growth in the sugar meal.<ref>{{cite journal |last1=Rossignol |first1=P.A. |last2=Lueders |first2=A.M. |title=Bacteriolytic factor in the salivary glands of Aedes aegypti |journal=Comparative Biochemistry and Physiology. B, Comparative Biochemistry |volume=83 |issue=4 |pages=819–822 |year=1986 |pmid=3519067 |doi=10.1016/0305-0491(86)90153-7 }}</ref>
For a mosquito to obtain a blood meal, it must circumvent its [[vertebrate]] host's physiological responses.  Mosquito saliva blocks the host's [[hemostasis]] system, with proteins that reduce [[vascular constriction]], [[blood clotting]], and [[platelet]] aggregation, to ensure the blood keeps flowing.<ref name="Valenzuela Pham Garfield 2002"/> It modulates the host's [[immune response]] via a mixture of proteins which lower [[angiogenesis]] and [[immunity (medical)|immunity]]; create [[inflammation]];<ref name="Valenzuela Pham Garfield 2002">{{cite journal |last1=Valenzuela |first1=J.G. |last2=Pham |first2=V.M. |last3=Garfield |first3=M.K. |last4=Francischetti |first4=I.M.B. |last5=Ribeiro |first5=J.M.C. |title=Toward a description of the sialome of the adult female mosquito Aedes aegypti |journal=Insect Biochemistry and Molecular Biology |volume=32 |issue=9 |pages=1101–1122 |date=September 2002 |pmid=12213246 |doi=10.1016/S0965-1748(02)00047-4 |bibcode=2002IBMB...32.1101V }}</ref><ref name="autogenerated4">{{cite journal |last1=Ribeiro |first1=J.M. |last2=Francischetti |first2=I.M. |title=Role of arthropod saliva in blood feeding: sialome and post-sialome perspectives |journal=Annual Review of Entomology |volume=48 |pages=73–88 |year=2003 |pmid=12194906 |doi=10.1146/annurev.ento.48.060402.102812 |url=https://zenodo.org/record/1234963 |access-date=2019-06-29 |archive-date=2020-08-04 |archive-url=https://web.archive.org/web/20200804040649/https://zenodo.org/record/1234963 |url-status=live }}</ref> suppress [[tumor necrosis factor]] release from activated [[mast cells]];<ref>{{cite journal |last1=Bissonnette |first1=Elyse Y. |last2=Rossignol |first2=Philippe A. |last3=Befus |first3=A. Dean|title=Extracts of mosquito salivary gland inhibit tumour necrosis factor alpha release from mast cells |journal=Parasite Immunology |volume=15 |issue=1 |pages=27–33 |date=January 1993 |pmid=7679483 |doi=10.1111/j.1365-3024.1993.tb00569.x }}</ref> suppress [[interleukin]] (IL)-2 and [[IFN-γ]] production;<ref name="Cross Cupp Enriquez 1994">{{cite journal |last1=Cross |first1=Martin L. |last2=Cupp |first2=Eddie W. |last3=Enriquez |first3=F. Javier |title=Differential modulation of murine cellular immune responses by salivary gland extract of Aedes aegypti |journal=The American Journal of Tropical Medicine and Hygiene |volume=51 |issue=5 |pages=690–696 |date=November 1994 |pmid=7985763 |doi=10.4269/ajtmh.1994.51.690 }}</ref><ref name="Zeidner Higgs Happ 1999">{{cite journal |last1=Zeidner |first1=Nordin S. |last2=Higgs |first2=Stephen |last3=Happ |first3=Christine M. |last4=Beaty |first4=Barry J. |last5=Miller |first5=Barry R. |title=Mosquito feeding modulates Th1 and Th2 cytokines in flavivirus susceptible mice: an effect mimicked by injection of sialokinins, but not demonstrated in flavivirus resistant mice |journal=Parasite Immunology |volume=21 |issue=1 |pages=35–44 |date=January 1999 |pmid=10081770 |doi=10.1046/j.1365-3024.1999.00199.x |s2cid=26774722 |url=https://zenodo.org/record/1236522 |access-date=2020-09-25 |archive-date=2022-04-10 |archive-url=https://web.archive.org/web/20220410004355/https://zenodo.org/record/1236522 |url-status=live }}</ref> suppress [[T cell]] populations;<ref>{{cite journal |last1=Wanasen |first1=N. |last2=Nussenzveig |first2=R. H. |last3=Champagne |first3=D. E. |last4=Soong |first4=L. |last5=Higgs |first5=S.|title=Differential modulation of murine host immune response by salivary gland extracts from the mosquitoes ''Aedes aegypti'' and ''Culex quinquefasciatus'' |journal=Medical and Veterinary Entomology |volume=18 |issue=2 |pages=191–199 |date=June 2004 |pmid=15189245 |doi=10.1111/j.1365-2915.2004.00498.x |s2cid=42458052 }}</ref><ref>{{cite journal |last1=Wasserman |first1=H. A. |last2=Singh |first2=S. |last3=Champagne |first3=D. E. |title=Saliva of the Yellow Fever mosquito, ''Aedes aegypti'', modulates murine lymphocyte function |journal=Parasite Immunology |volume=26 |issue=6–7 |pages=295–306 |year=2004 |pmid=15541033 |doi=10.1111/j.0141-9838.2004.00712.x |s2cid=32742815 }}</ref><ref>{{cite journal |last1=Depinay |first1=Nadya |last2=Hacini |first2=Fériel |last3=Beghdadi |first3=Walid |last4=Peronet |first4=Roger |last5=Mécheri |first5=Salaheddine |title=Mast cell-dependent down-regulation of antigen-specific immune responses by mosquito bites |journal=Journal of Immunology |volume=176 |issue=7 |pages=4141–4146 |date=April 2006 |pmid=16547250 |doi=10.4049/jimmunol.176.7.4141 |doi-access=free }}</ref> decrease expression of [[interferon]]−α/β, making virus infections more severe;<ref>{{cite journal |last1=Schneider |first1=Bradley S. |last2=Soong |first2=Lynn |last3=Zeidner |first3=Nordin S. |last4=Higgs |first4=Stephen |title=Aedes aegypti salivary gland extracts modulate anti-viral and TH1/TH2 cytokine responses to sindbis virus infection |journal=Viral Immunology |volume=17 |issue=4 |pages=565–573 |year=2004 |pmid=15671753 |doi=10.1089/vim.2004.17.565 }}</ref><ref>{{cite journal |last1=Taylor |first1=J. L. |last2=Schoenherr |first2=C. |last3=Grossberg |first3=S. E.|title=Protection against Japanese encephalitis virus in mice and hamsters by treatment with carboxymethylacridanone, a potent interferon inducer |journal=The Journal of Infectious Diseases |volume=142 |issue=3 |pages=394–399 |date=September 1980 |pmid=6255036 |doi=10.1093/infdis/142.3.394 }}</ref> increase [[natural killer T cell]]s in the blood; and decrease cytokine production.<ref>{{cite journal |last1=Vogt |first1=Megan B. |last2=Lahon |first2=Anismrita |last3=Arya |first3=Ravi P. |last4=Kneubehl |first4=Alexander R. |last5=Spencer Clinton |first5=Jennifer L. |last6=Paust |first6=Silke |last7=Rico-Hesse |first7=Rebecca |title=Mosquito saliva alone has profound effects on the human immune system |journal=PLOS Neglected Tropical Diseases |volume=12 |issue=5 |pages=e0006439 |date=May 2018 |pmid=29771921 |pmc=5957326 |doi=10.1371/journal.pntd.0006439 |doi-access=free}}</ref>
=== Egg development and blood digestion ===
[[File:Anopheles stephensi.jpeg|thumb|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.<!--<ref name="Mullen2009">{{cite book |first1=Gary |last1=Mullen |first2=Lance |last2=Durden |year=2009 |title=Medical and Veterinary Entomology |publisher=Academic Press |location=London |pages= }}</ref>--><ref>{{cite journal |last1=Zhu |first1=J. |last2=Miura |first2=K. |last3=Dittmer |first3=N.T. |last4=Raikhel |first4=A.S. |title=AaSvp, a mosquito homolog of COUP-TF is involved in termination of vitellogenesis by repressing the 20-hydroecdysone response |journal=Journal of Insect Science |date=2002 |volume=2 |issue=17 |page=17 |pmid=15455051|pmc=405832 }}</ref> Upon completion of feeding, the mosquito withdraws her [[proboscis]], and as the gut fills up, the stomach lining secretes a [[Peritrophic matrix|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.<ref name="Curic Hercog Vrselja Wagner 2014">{{cite journal |last1=Curic |first1=Goran |last2=Hercog |first2=Rajna |last3=Vrselja |first3=Zvonimir |last4=Wagner |first4=Jasenka |title=Identification of person and quantification of human DNA recovered from mosquitoes (Culicidae) |journal=Forensic Science International: Genetics |volume=8 |issue=1 |date=2014 |doi=10.1016/j.fsigen.2013.07.011 |pages=109–112|pmid=24315597 }}</ref> Once blood is in the stomach, the midgut synthesizes [[protease]] enzymes, primarily [[trypsin]] assisted by [[aminopeptidase]], that hydrolyze the blood [[protein]]s into free [[amino acid]]s. These are used in the synthesis of [[vitellogenin]], which in turn is made into egg yolk protein.<ref name="Billingsley">{{cite journal |last1=Billingsley |first1=P.F. |last2=Hecker |first2=H. |title=Blood digestion in the mosquito, Anopheles stephensi Liston (Diptera: Culicidae): activity and distribution of trypsin, aminopeptidase, and alpha-glucosidase in the midgut |journal=Journal of Medical Entomology |volume=28 |issue=6 |pages=865–871 |date=November 1991 |pmid=1770523 |doi=10.1093/jmedent/28.6.865 }}</ref>
== Distribution ==
=== Cosmopolitan ===
Mosquitoes have a [[cosmopolitan distribution]], occurring in every land region except Antarctica and a few islands with polar or [[subpolar climate]]s, such as [[Iceland]], which is essentially free of mosquitoes.<ref name="Visindavefur.hi.is">{{cite web |url=http://visindavefur.hi.is/svar.php?id=2166 |title=Vísindavefurinn: Af hverju lifa ekki moskítóflugur á Íslandi, fyrst þær geta lifað báðum megin á Grænlandi? |language=is |publisher=Visindavefur.hi.is |access-date=2013-10-15 |archive-url=https://archive.today/20130802162121/http://visindavefur.hi.is/svar.php?id=2166 |archive-date=2013-08-02 |url-status=dead }}</ref> 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.<ref>{{cite journal |last=Peterson |first=B.V. |year=1977 |journal=The Canadian Entomologist |volume=109 |issue=3 |pages=449–472 |doi=10.4039/Ent109449-3 |title=The Black Flies of Iceland (Diptera: Simuliidae) |s2cid=86752961 }}</ref><ref>{{cite journal |last1=Gislason |first1=G.M. |author2=Gardarsson A. |year=1988 |title=Long term studies on ''Simulium vittatum'' Zett. (Diptera: Simuliidae) in the River Laxá, North Iceland, with particular reference to different methods used in assessing population changes |journal=Verb. Int. Ver. Limnol |volume=23 |issue=4 |pages=2179–2188 |doi=10.1080/03680770.1987.11899871 |bibcode=1988SILP...23.2179G }}</ref>
Eggs of [[Temperate climate|temperate zone]] mosquitoes are more tolerant of cold than the eggs of species indigenous to warmer regions.<ref>{{cite journal |last1=Hawley |first1=W. A. |last2=Pumpuni |first2=C. B. |last3=Brady |first3=R. H. |last4=Craig |first4=G. B. |title=Overwintering survival of ''Aedes albopictus'' (Diptera: Culicidae) eggs in Indiana |journal=Journal of Medical Entomology |volume=26 |issue=2 |pages=122–129 |date=March 1989 |pmid=2709388 |doi=10.1093/jmedent/26.2.122 }}</ref><ref>{{cite journal |last1=Hanson |first1=S.M. |last2=Craig |first2=G.B. |title=''Aedes albopictus'' (Diptera: Culicidae) eggs: field survivorship during northern Indiana winters |journal=Journal of Medical Entomology |volume=32 |issue=5 |pages=599–604 |date=September 1995 |pmid=7473614 |doi=10.1093/jmedent/32.5.599 }}</ref> Many can tolerate subzero temperatures, while adults of some species can survive winter by sheltering in microhabitats such as buildings or hollow trees.<ref>{{cite journal |last1=Romi |first1=Roberto |last2=Severini |first2=Francesco |last3=Toma |first3=Luciano |title=Cold acclimation and overwintering of female ''Aedes albopictus'' in Roma |journal=Journal of the American Mosquito Control Association |volume=22 |issue=1 |pages=149–151 |date=March 2006 |pmid=16646341 |doi=10.2987/8756-971X(2006)22[149:CAAOOF]2.0.CO;2 |s2cid=41129725 }}</ref> In warm and humid tropical regions, some mosquito species are active for the entire year, but in temperate and cold regions they hibernate or enter [[diapause]]. [[Arctic]] or subarctic mosquitoes, like some other arctic midges in families such as [[Simuliidae]] and [[Ceratopogonidae]] 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 a [[caribou]] herd.<ref name=n1>{{cite journal |last=Fang |first=J. |title=Ecology: A world without mosquitoes |journal=Nature |volume=466 |issue=7305 |pages=432–434 |date=July 2010 |pmid=20651669 |doi=10.1038/466432a |doi-access=free }}</ref>
=== Effect of climate change ===
For a mosquito to transmit disease, there must be favorable seasonal conditions,<ref name=":0">{{Cite journal |last=Reiter |first=Paul |date=2001 |title=Climate Change and Mosquito-Borne Disease |journal=Environmental Health Perspectives |volume=109 |issue=Suppl 1 |pages=142–158 |via=EHP |doi=10.1289/ehp.01109s1141 |pmid=11250812 |pmc=1240549}}</ref> primarily humidity, temperature, and precipitation.<ref name="Bai Morton Liu 2013">{{cite journal |last1=Bai |first1=Li |last2=Morton |first2=Lindsay Carol |last3=Liu |first3=Qiyong |title=Climate change and mosquito-borne diseases in China: a review |journal=Globalization and Health |volume=9 |page=10 |date=March 2013 |pmid=23497420 |pmc=3605364 |doi=10.1186/1744-8603-9-10 |doi-access=free }}</ref> [[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.<ref name="pmid30120891">{{cite journal |last1=Caminade |first1=Cyril |last2=McIntyre |first2=K. Marie |last3=Jones |first3=Anne E. |title=Impact of recent and future climate change on vector-borne diseases |journal=Annals of the New York Academy of Sciences |volume=1436 |issue=1 |pages=157–173 |date=January 2019 |pmid=30120891 |pmc=6378404 |doi=10.1111/nyas.13950 |bibcode=2019NYASA1436..157C }}</ref> 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.<ref name="pmid29229233">{{cite journal |last1=Tjaden |first1=Nils Benjamin |last2=Caminade |first2=Cyril |last3=Beierkuhnlein |first3=Carl |last4=Thomas |first4=Stephanie Margarete |title=Mosquito-Borne Diseases: Advances in Modelling Climate-Change Impacts |journal=Trends in Parasitology |volume=34 |issue=3 |pages=227–245 |date=March 2018 |pmid=29229233 |doi=10.1016/j.pt.2017.11.006 }}</ref>
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.<ref name=":2">{{Cite journal |last=Baylis |first=Matthew |date=5 December 2017 |title=Potential impact of climate change on emerging vector-borne and other infections in the UK |journal=Environmental Health |volume=16 |issue=Suppl 1 |page=112 |pmc=5773876 |pmid=29219091 |doi=10.1186/s12940-017-0326-1 |bibcode=2017EnvHe..16S.112B |doi-access=free }}</ref><ref name="pmid29219091">{{cite journal |last=Baylis |first=M. |title=Potential impact of climate change on emerging vector-borne and other infections in the UK |journal=Environmental Health |volume=16 |issue=Suppl 1 |page=112 |date=December 2017 |pmid=29219091 |pmc=5773876 |doi=10.1186/s12940-017-0326-1 |bibcode=2017EnvHe..16S.112B |doi-access=free }}</ref>
[[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.<ref name="Horton 2024">{{cite news |last1=Horton |first1=Helena |title=Mosquito-borne diseases spreading in Europe due to climate crisis, says expert |url=https://www.theguardian.com/environment/2024/apr/25/mosquito-borne-diseases-spreading-in-europe-due-to-climate-crisis-says-expert |access-date=25 April 2024 |work=[[The Guardian]] |date=25 April 2024 |archive-date=10 September 2024 |archive-url=https://web.archive.org/web/20240910225716/https://www.theguardian.com/environment/2024/apr/25/mosquito-borne-diseases-spreading-in-europe-due-to-climate-crisis-says-expert |url-status=live }}</ref>
== Ecology ==
=== Predators and parasites ===
Mosquito larvae are among the commonest animals in ponds, and they form an important food source for freshwater [[Predation|predator]]s. Among the many aquatic insects that catch mosquito larvae are [[dragonfly]] and [[damselfly]] nymphs, [[whirligig beetle]]s, and [[water strider]]s. Vertebrate predators include fish such as catfish and the [[mosquitofish]], amphibians including the [[Scaphiopodidae|spadefoot toad]] and the [[giant tree frog]], freshwater turtles such as the [[red-eared slider]], and birds such as ducks.<ref>{{Cite web |last=Beck |first=Kevin |title=What Eats Mosquitoes? |date=November 22, 2019 |url=https://sciencing.com/eats-mosquitoes-8568515.html |url-status=live |access-date=31 May 2021 |website=Sciencing |archive-date=2 June 2021 |archive-url=https://web.archive.org/web/20210602215704/https://sciencing.com/eats-mosquitoes-8568515.html}}</ref>
Emerging adults are consumed at the pond surface by predatory flies including [[Empididae]] and [[Dolichopodidae]], and by [[spider]]s. Flying adults are captured by dragonflies and damselflies, by birds such as [[Apus (bird)|swifts]] and [[swallow]]s, and by vertebrates including [[bat]]s.<ref name="Medlock Snow 2008"/>
Mosquitoes are parasitised by [[Hydrachnidia|hydrachnid]] mites, [[ciliate]]s such as ''[[Glaucoma (ciliate)|Glaucoma]]'', [[microsporidia]]ns such as ''Thelania'', and fungi including species of [[Saprolegniaceae]] and [[Entomophthoraceae]].<ref name="Medlock Snow 2008">{{cite journal |last1=Medlock |first1=J. M. |last2=Snow |first2=K. R. |title=Natural predators and parasites of British mosquitoes–a review |journal=European Mosquito Bulletin |volume=25 |issue=1 |year=2008 |pages=1–11 |url=https://e-m-b.myspecies.info/sites/e-m-b.org/files/European_Mosquito_Bulletin_Publications811/EMB25/EMB25_1.pdf |access-date=15 December 2023 |archive-date=15 December 2023 |archive-url=https://web.archive.org/web/20231215100621/https://e-m-b.myspecies.info/sites/e-m-b.org/files/European_Mosquito_Bulletin_Publications811/EMB25/EMB25_1.pdf |url-status=live }}</ref>
=== Pollination ===
[[File:Mosquito on marigold flower.JPG|thumb|A mosquito visiting a [[Calendula|marigold]] flower for [[nectar]] ]]
Several flowers including members of the [[Asteraceae]], [[Rosaceae]] and [[Orchidaceae]] are [[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 [[Exaptation|exapted]] mosquitoes to blood-sucking.<ref name="Peach-Gries-2020"/>
=== Parasitism ===
{{further |Parasitism}}
Ecologically, blood-feeding mosquitoes are [[micropredator]]s, small animals that feed on larger animals without immediately killing them. Evolutionary biologists see this as a form of [[parasitism]]; in [[E. O. Wilson|Edward O. Wilson]]'s phrase "Parasites ... are predators that eat prey in units of less than one."<ref name="Wilson 2014">{{cite book |last=Wilson |first=Edward O. |author-link=E. O. Wilson |title=The Meaning of Human Existence  |year=2014 |publisher=W. W. Norton & Company |isbn=978-0-87140-480-0 |page=112 |quote=Parasites, in a phrase, are predators that eat prey in units of less than one. Tolerable parasites are those that have evolved to ensure their own survival and reproduction but at the same time with minimum pain and cost to the host.}}</ref> Micropredation is one of six major [[Behavioral ecology#Evolutionarily stable strategy|evolutionarily stable strategies]] within parasitism. It is distinguished by leaving the host still able to reproduce, unlike the activity of [[Parasitic castration|parasitic castrator]]s or [[parasitoid]]s; and having multiple hosts, unlike conventional parasites.<ref name="Poulin Randhawa 2011">{{cite journal |last=Poulin |first=Robert |author-link=Robert Poulin (zoologist) |editor1=Rollinson, D. |editor2=Hay, S. I. |title=The Many Roads to Parasitism: A Tale of Convergence |journal=Advances in Parasitology |url=https://books.google.com/books?id=9y4AlXka7t0C&pg=PA28 |year=2011 |volume=74 |publisher=Academic Press |isbn=978-0-12-385897-9 |pages=27–28 |doi=10.1016/B978-0-12-385897-9.00001-X |pmid=21295676 }}</ref><ref name="Poulin Randhawa 2015">{{cite journal |last1=Poulin |first1=Robert |author1-link=Robert Poulin (zoologist) |last2=Randhawa |first2=Haseeb S. |title=Evolution of parasitism along convergent lines: from ecology to genomics |journal=Parasitology |date=February 2015 |volume=142 |issue=Suppl 1 |pages=S6–S15 |doi=10.1017/S0031182013001674 |pmc=4413784 |pmid=24229807}} {{open access}}</ref> From this perspective, mosquitoes are [[ectoparasite]]s, feeding on blood from the outside of their hosts, using their piercing mouthparts, rather than entering their bodies. Unlike some other ectoparasites such as [[flea]]s and [[Louse|lice]], mosquitoes do not remain constantly on the body of the host, but visit only to feed.<ref name="Poulin Randhawa 2015"/>
== Evolution ==
=== Fossil record ===
[[File:Culex malariager in Dominican amber.jpg|thumb|upright|alt=Fossilized mosquito encased in amber |''Culex malariager'' mosquito infected with the [[malaria]]l parasite ''Plasmodium dominicana'', in [[Dominican Republic|Dominican]] amber of [[Miocene]] age, 15–20 million years ago<ref name="Poinar 2014">{{cite journal |last=Poinar |first=George |title=Evolutionary History of Terrestrial Pathogens and Endoparasites as Revealed in Fossils and Subfossils |journal=Advances in Biology |volume=2014 |date=2014-06-12 |doi=10.1155/2014/181353 |pages=1–29 |doi-access=free }}</ref>]]
A 2023 study suggested that ''[[Libanoculex |Libanoculex intermedius]]'' found in [[Lebanese amber]], dating to the [[Barremian]] age of the Early Cretaceous, around 125 million years ago was the oldest known mosquito.<ref>{{Cite journal |last1=Azar |first1=Dany |last2=Nel |first2=André |last3=Huang |first3=Diying |last4=Engel |first4=Michael S. |date=December 2023 |title=The Earliest Fossil Mosquito |journal=Current Biology |volume=33 |issue=23 |pages=5240–5246.e2 |doi=10.1016/j.cub.2023.10.047 |pmid=38052162 |s2cid=265612144 |doi-access=free |bibcode=2023CBio...33E5240A }}</ref> However its identification as a mosquito is disputed, with other authors considering it to be a [[Chaoboridae|chaoborid]] fly instead.<ref>{{Cite journal |last=Harbach |first=Ralph E. |date=2024-03-12 |title=Libanoculex&nbsp;intermedius is not a mosquito (Diptera: Culicidae): It is a chaoborid (Chaoboridae) |url=https://mapress.com/zt/article/view/zootaxa.5424.1.9 |journal=Zootaxa |language=en |volume=5424 |issue=1 |pages=139–144 |doi=10.11646/zootaxa.5424.1.9 |issn=1175-5334 |access-date=9 August 2024 |archive-date=9 August 2024 |archive-url=https://web.archive.org/web/20240809084423/https://mapress.com/zt/article/view/zootaxa.5424.1.9 |url-status=live }}</ref> Three other unambiguous species of [[Cretaceous]] mosquito are known. ''[[Burmaculex antiquus]]'' and ''[[Priscoculex burmanicus]]'' are known from [[Burmese amber]] from Myanmar, which dates to the earliest part of the [[Cenomanian]] age of the Late Cretaceous, around 99 million years ago.<ref>{{cite journal |last1=Borkent |first1=A. |last2=Grimaldi |first2=D.A. |year=2004 |title=The earliest fossil mosquito (Diptera: Culicidae), in Mid-Cretaceous Burmese amber |journal=[[Annals of the Entomological Society of America]] |volume=97 |issue=5 |pages=882–888 |doi=10.1603/0013-8746(2004)097[0882:TEFMDC]2.0.CO;2 |doi-access=free}}</ref><ref name=":3">{{Cite journal |last1=Poinar |first1=George |last2=Zavortink |first2=Thomas J. |last3=Brown |first3=Alex |date=2019-01-30 |title=''Priscoculex burmanicus'' n. gen. et sp. (Diptera: Culicidae: Anophelinae) from mid-Cretaceous Myanmar amber |journal=Historical Biology |volume=32 |issue=9 |pages=1157–1162 |doi=10.1080/08912963.2019.1570185 |s2cid=92836430}}</ref> ''[[Paleoculicis minutus]]'', is known from [[Canadian amber]] from Alberta, Canada, which dates to the [[Campanian]] age of the Late Cretaceous, around 79 million years ago.<ref>{{cite journal |author=Poinar, G. O. |display-authors=etal |year=2000 |title=''Paleoculicis minutus'' (Diptera: Culicidae) n. gen., n. sp., from Cretaceous Canadian amber with a summary of described fossil mosquitoes |url=http://www.geologica-acta.com/pdf/aghv3501a12.pdf |url-status=dead |journal=[[Geologica Acta|Acta Geológica Hispánica]] |volume=35 |pages=119–128 |archive-url=https://web.archive.org/web/20131029193407/http://www.geologica-acta.com/pdf/aghv3501a12.pdf |archive-date=2013-10-29 |access-date=2009-12-10}}</ref> ''P. burmanicus'' has been assigned to the [[Anophelinae]], indicating that the split between this subfamily and the [[Culicinae]] took place over 99 million years ago.<ref name=":3" /> 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.<ref>{{Cite journal |last1=Lorenz |first1=Camila |last2=Alves |first2=João M.P. |last3=Foster |first3=Peter G. |last4=Suesdek |first4=Lincoln |last5=Sallum |first5=Maria Anice M. |date=2021-05-10 |title=Phylogeny and temporal diversification of mosquitoes (Diptera: Culicidae) with an emphasis on the Neotropical fauna |journal=Systematic Entomology |volume=46 |issue=4 |pages=798–811 |doi=10.1111/syen.12489 |bibcode=2021SysEn..46..798L |s2cid=236612378}}</ref>
=== Taxonomy ===
{{further|List of mosquito genera}}
Over 3,600 species of mosquitoes in 112 [[genera]] have been [[Alpha taxonomy|described]]. They are traditionally divided into two subfamilies, the [[List of mosquito genera#Subfamily Anophelinae|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.<ref name="Molina-Cruz Lehmann Knöckel 2013 pp. 530–537">{{cite journal |last1=Molina-Cruz |first1=Alvaro |last2=Lehmann |first2=Tovi |last3=Knöckel |first3=Julia |title=Could culicine mosquitoes transmit human malaria? |journal=Trends in Parasitology |volume=29 |issue=11 |date=2013 |doi=10.1016/j.pt.2013.09.003 |pages=530–537 |pmid=24140295 |pmc=10987011 }}</ref>
The name Culicidae was introduced by the German entomologist [[Johann Wilhelm Meigen]] in his seven-volume classification published in 1818–1838.<ref name="Meigen 1818">{{cite book |last=Meigen |first=Johann Wilhelm |author-link=Johann Wilhelm Meigen |date=1818–1838 |title=Systematische Beschreibung der bekannten Europäischen zweiflügeligen Insekten |language=de |trans-title=Systematic description of the known European two-winged insects |location=Aachen |publisher=Friedrich Wilhelm Forstmann |volume=1–7 |url=https://www.biodiversitylibrary.org/item/45833#page/9/mode/1up |access-date=16 December 2023 |archive-date=26 February 2023 |archive-url=https://web.archive.org/web/20230226175413/https://www.biodiversitylibrary.org/item/45833#page/9/mode/1up |url-status=live }}</ref> Mosquito taxonomy was advanced in 1901 when the English entomologist [[Frederick Vincent Theobald]] published his 5-volume monograph on the Culicidae.<ref name=Theobald1901>{{cite book |last=Theobald |first=Frederick Vincent |author-link=Frederick Vincent Theobald |date=1901 |title=A Monograph of the Culicidae, or Mosquitoes |location=London |publisher=[[British Museum (Natural History)]] |volume=1 |isbn=978-1178519037 |url=https://archive.org/details/amonographculic03zoolgoog/page/n3/mode/2up |page=4<!--ie whole book-->}}</ref> He had been provided with mosquito specimens sent in to the [[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."<ref name="Harbach Kitching 2016">{{Cite journal |last1=Harbach |first1=R. E. |last2=Kitching |first2=I. |date=January 2016 |title=The phylogeny of Anophelinae revisited: inferences about the origin and classification of Anopheles (Diptera: Culicidae) |journal=[[Zoologica Scripta]] |volume=45 |pages=34–47 |url=https://nhm.openrepository.com/handle/10141/612216 |doi=10.1111/zsc.12137 |hdl=10141/612216 |s2cid=46364692 |hdl-access=free |access-date=16 December 2023 |archive-date=10 September 2024 |archive-url=https://web.archive.org/web/20240910225816/https://nhm.openrepository.com/handle/10141/612216 |url-status=live }}</ref>
=== Phylogeny ===
==== External ====
Mosquitoes are members of a [[family (biology)|family]] of the [[Fly|true flies (order Diptera)]]: the Culicidae (from the [[Latin]] {{lang|la|culex}}, [[Genitive case|genitive]] {{lang|la|culicis}}, meaning "midge" or "gnat").<ref name="isbn0-398-06179-3">{{cite book |last=Jaeger |first=Edmund C. |author-link=Edmund Jaeger |title=A Source-Book of Biological Names and Terms |publisher=Thomas |location=Springfield, Ill |year=1959 |isbn=978-0-398-06179-1 }}</ref> They are members of the infraorder [[Culicomorpha]] and superfamily [[Culicoidea]]. The phylogenetic tree is based on the FLYTREE project.<ref>{{cite web |last1=Yeates |first1=David K. |last2=Meier |first2=Rudolf |last3=Wiegmann |first3=Brian |title=Phylogeny of True Flies (Diptera): A 250 Million Year Old Success Story in Terrestrial Diversification |url=http://www.inhs.illinois.edu/research/flytree/flyphylogeny/ |website=Flytree |publisher=Illinois Natural History Survey |archive-url=https://web.archive.org/web/20151228210535/http://inhs.illinois.edu/research/flytree/flyphylogeny/ |url-status=dead |archive-date=28 December 2015 |access-date=24 May 2016}}</ref><ref>{{cite journal |last1=Yeates |first1=David K. |last2=Weigmann |first2=Brian M. |last3=Courtney |first3=Greg W. |last4=Meier |first4=Rudolf |last5=Lambkins |first5=Christine |last6=Pape |first6=Thomas |title=Phylogeny and systematics of Diptera: Two decades of progress and prospects |year=2007 |journal=Zootaxa |volume=1668 |pages=565–590 |doi=10.11646/zootaxa.1668.1.27 }}</ref>
{{clade|style=font-size:100%;line-height:110%;
|label1=[[Diptera]]
|sublabel1= (true flies) &nbsp;
|1={{clade
|1={{clade
|1=[[Ptychopteromorpha]] (phantom and primitive crane-flies) [[File:Ptychoptera contaminata male Walker 1856 plate-XXVIII.png|70px]]
|label2=[[Culicomorpha]]
|2={{clade
|1=[[Chironomidae]] (non-biting midges) [[File:Chironomidae indet. (Chironomidae) (Unidentified Non-biting Midge) - (male imago), Arnhem, the Netherlands - 2.jpg|70px]]
|2={{clade
|1=Simulioidea (blackflies and biting midges) [[File:CSIRO ScienceImage 11052 Biting midge on human skin (cropped).jpg|60px]]
|label2=[[Culicoidea]]
|2={{clade
|1=[[Dixidae]] (meniscus midges) [[File:Dixa nebulosa adult John Curtis British Entomology 409.png|70px]]
|2={{clade
|1=[[Corethrellidae]] (frog-biting midges)
|2={{clade
|1=[[Chaoboridae]] (phantom midges) [[File:Chaoborus crystallinus male as Corethra plumicornis (Walker Insecta Britanica Vol 3 page 397 plate XXV).png|60px]]
|2='''Culicidae''' [[File:Stegomyia fasciata.jpg|100px]]
}}
}}
}}
}}
}}
}}
|2={{clade
|1=other midges and gnats [[File:Isoneuromyia annandalei.jpg|70px]]
|2=all other flies, inc. [[Brachycera]] [[File:Syrphidae icon.jpg|70px]]
}}
}}
}}
==== 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.<ref name="Reidenbach Cook Bertone Harbach 2009">{{cite journal |last1=Reidenbach |first1=Kyanne R. |last2=Cook |first2=Shelley |last3=Bertone |first3=Matthew A. |last4=Harbach |first4=Ralph E. |last5=Wiegmann |first5=Brian M. |last6=Besansky |first6=Nora J. |title=Phylogenetic analysis and temporal diversification of mosquitoes (Diptera: Culicidae) based on nuclear genes and morphology |journal=[[BMC Evolutionary Biology]] |volume=9 |issue=1 |date=2009 |page=298|pmid=20028549 |pmc=2805638 |doi=10.1186/1471-2148-9-298 |bibcode=2009BMCEE...9..298R |doi-access=free }}</ref>
{{clade|style=font-size:100%;line-height:110%;
|label1='''Culicidae'''
|1={{clade
|1=basal spp.
|2={{clade
|label1=[[Anophelinae]]
|1=[[File:Anopheles gambiae mosquito feeding 1354.p lores.jpg|125px]]
|label2=[[Culicinae]]
|2={{clade
|1=other spp.
|2={{clade
|1=[[Aedini]] [[File:Aedes aegypti feeding (cropped).jpg|125px]]
|2={{clade
|1=other spp.
|2=[[Sabethini]] [[File:Sabethes albiprivus (cropped).jpg|125px]]
}}
}}
}}
}}
}}
}}
== Interactions with humans ==
[[File:Anopheles albimanus mosquito.jpg|thumb|''[[Anopheles albimanus]]'' feeding on a human arm. As mosquitoes are the only vectors of [[malaria]], controlling them reduces its incidence.]]
=== Vectors of disease ===
{{Main|Mosquito-borne disease}}
Mosquitoes are [[Vector (epidemiology)|vectors]] for many disease-causing [[microorganism]]s including [[bacteria]], [[virus]]es, and [[protozoa]]n parasites. Nearly 700 million people acquire a [[mosquito-borne illness]] each year, resulting in over 725,000 deaths.<ref name="EBMED2014">{{cite web |title=Mosquito as Deadly Menace |url=https://www.pfizer.com/news/articles/mosquito_as_deadly_menace#.Yweo2_DM88I.link |publisher=[[Pfizer]] |access-date=10 December 2023 |archive-date=25 August 2022 |archive-url=https://web.archive.org/web/20220825170028/https://www.pfizer.com/news/articles/mosquito_as_deadly_menace#.Yweo2_DM88I.link |url-status=live }}</ref> Common mosquito-borne viral diseases include [[yellow fever]]<ref name="WHO2 014">{{cite web |title=Yellow fever Fact sheet N°100 |url=https://www.who.int/mediacentre/factsheets/fs100/en/ |publisher=[[World Health Organization]] |access-date=23 February 2014 |date=May 2013 |url-status=live |archive-url=https://web.archive.org/web/20140219011908/http://www.who.int/mediacentre/factsheets/fs100/en/ |archive-date=19 February 2014}}</ref> and [[dengue fever]] transmitted mostly by ''[[Aedes aegypti]]''.<ref name="WHO 2009">{{cite book |url=http://whqlibdoc.who.int/publications/2009/9789241547871_eng.pdf |title=Dengue Guidelines for Diagnosis, Treatment, Prevention and Control |publisher=[[World Health Organization]] |year=2009 |isbn=978-92-4-154787-1 |access-date=2013-08-13 |archive-date=2012-10-17 |archive-url=https://web.archive.org/web/20121017230435/http://whqlibdoc.who.int/publications/2009/9789241547871_eng.pdf |url-status=live}}</ref> Parasitic diseases transmitted by mosquitoes include [[malaria]] and [[lymphatic filariasis]]. The ''[[Plasmodium]]'' parasites that cause malaria are carried by female ''[[Anopheles]]'' mosquitoes. Lymphatic filariasis, the main cause of [[elephantiasis tropica|elephantiasis]], is spread by a wide variety of mosquitoes.<ref>{{cite web |title=Lymphatic Filariasis |url=https://www.who.int/mediacentre/factsheets/fs102/en/ |publisher=[[World Health Organization|World Health Organisation]] |access-date=24 August 2011 |archive-date=5 May 2016 |archive-url=https://web.archive.org/web/20160505163529/http://www.who.int/mediacentre/factsheets/fs102/en/|url-status=live}}</ref> A bacterial disease spread by ''Culex'' and ''Culiseta'' mosquitoes is [[tularemia]].<ref name="pmid21776596">{{cite journal |last1=Muslu |first1=H. |last2=Kurt |first2=O. |last3=Özbilgin |first3=A. |title=[Evaluation of mosquito species (Diptera: Culicidae) identified in Manisa province according to their breeding sites and seasonal differences] |language=tr |journal=Turkiye Parazitolojii Dergisi |volume=35 |issue=2 |pages=100–104 |year=2011 |pmid=21776596 |doi=10.5152/tpd.2011.25 |doi-access=free }}</ref>
=== Control ===
{{main|Mosquito control}}
[[File:Mosquito Netting.jpg|thumb|Mosquito nets can prevent people being bitten while they sleep.]]
Many measures have been tried for [[mosquito control]], including the elimination of breeding places, exclusion via [[window screen]]s and [[mosquito net]]s, [[biological control]] with parasites such as fungi<ref>{{cite news |url=https://www.nytimes.com/2005/06/10/science/10mosquito.html |title=Fungus Fatal to Mosquito May Aid Global War on Malaria |date=10 June 2005 |work=The New York Times |access-date=19 February 2017 |archive-date=9 May 2015 |archive-url=https://web.archive.org/web/20150509040906/http://www.nytimes.com/2005/06/10/science/10mosquito.html |url-status=live}}</ref><ref>{{cite journal |last=Kramer |first=J.P. |title=''Entomophthora culicis'' (Zygomycetes, Entomophthorales) as a pathogen of ''adultaedes aegypti'' (diptera, culicidae) |journal=Aquatic Insects |volume=4 |issue=2 |pages=73–79 |year=1982 |doi=10.1080/01650428209361085 |bibcode=1982AqIns...4...73K }}</ref> and nematodes,<ref>{{cite journal |last1=Shamseldean |first1=M.M. |last2=Platzer |first2=E.G. |title=''Romanomermis culicivorax'': penetration of larval mosquitoes |journal=Journal of Invertebrate Pathology |volume=54 |issue=2 |pages=191–199 |date=September 1989 |pmid=2570111 |doi=10.1016/0022-2011(89)90028-1 |bibcode=1989JInvP..54..191S }}</ref> or predators such as fish,<ref>{{cite journal |last=Krumholz |first=Louis A. |jstor=1948627 |title=Reproduction in the Western Mosquitofish, ''Gambusia affinis affinis'' (Baird & Girard), and Its Use in Mosquito Control |journal=Ecological Monographs |volume=18 |issue=1 |year=1948 |pages=1–43 |doi=10.2307/1948627 |bibcode=1948EcoM...18....1K }}</ref><ref name="Jianguo-Dashu-1995">{{cite book |chapter-url=http://www.idrc.ca/en/ev-27870-201-1-DO_TOPIC.html |chapter=Part III: Interactions - 31. A Comparative Study of the Ability of Fish to Catch Mosquito Larva |title=Rice-fish culture in China |isbn=978-1-55250-313-3 |year=1995 |publisher=International Development Research Centre |url-status=dead |archive-url=https://web.archive.org/web/20110609010556/https://www.idrc.ca/en/ev-27870-201-1-DO_TOPIC.html |archive-date=2011-06-09 |first1=Wang |last1=Jianguo |first2=Ni |last2=Dashu |editor1-first=Kenneth T. |editor1-last=MacKay}}</ref><ref>{{cite journal |last=Fradin |first=M.S. |title=Mosquitoes and mosquito repellents: a clinician's guide |journal=[[Annals of Internal Medicine]] |volume=128 |issue=11 |pages=931–940 |date=June 1998 |pmid=9634433 |doi=10.7326/0003-4819-128-11-199806010-00013 |citeseerx=10.1.1.691.2193 |s2cid=35046348 }}</ref> [[copepods]],<ref>{{cite journal |last1=Marten |first1=G.G. |last2=Reid |first2=J.W. |title=Cyclopoid copepods |journal=Journal of the American Mosquito Control Association |volume=23 |issue=2 Suppl |pages=65–92 |year=2007 |pmid=17853599 |doi=10.2987/8756-971X(2007)23[65:CC]2.0.CO;2 |s2cid=7645668 }}</ref> [[dragonfly]] nymphs and adults, and some species of lizard and [[gecko]].<ref>{{cite journal |last1=Canyon |first1=D.V. |last2=Hii |first2=J.L. |title=The gecko: an environmentally friendly biological agent for mosquito control |journal=Medical and Veterinary Entomology |volume=11 |issue=4 |pages=319–323 |date=October 1997 |pmid=9430109 |doi=10.1111/j.1365-2915.1997.tb00416.x |s2cid=26987818 }}</ref> Another approach is to [[sterile insect technique|introduce large numbers of sterile males]].<ref name=twsE44>{{cite news |last=Carpenter |first=Jennifer |title=Spermless mosquitoes hold promise to stop malaria |publisher=[[BBC]] |quote=Scientists have created spermless mosquitoes in an effort to curb the spread of malaria. |date=8 August 2011 |url=https://www.bbc.co.uk/news/science-environment-14422440 |access-date=5 August 2011 |archive-date=9 August 2011 |archive-url=https://web.archive.org/web/20110809002749/http://www.bbc.co.uk/news/science-environment-14422440 |url-status=live }}</ref> 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.<ref>Webb, Jonathan (10 June 2014) [https://www.bbc.co.uk/news/science-environment-27765974 GM lab mosquitoes may aid malaria fight] {{Webarchive |url=https://web.archive.org/web/20220816191930/https://www.bbc.com/news/science-environment-27765974 |date=2022-08-16 }}. ''[[BBC]]''.</ref> Control of disease-carrying mosquitoes using [[gene drive]]s has been proposed.<ref>{{cite journal |last1=Kyrou |first1=Kyros Kyrou |display-authors=etal |title=A CRISPR–Cas9 gene drive targeting ''doublesex'' causes complete population suppression in caged ''Anopheles gambiae'' mosquitoes |journal=Nature Biotechnology |volume=36 |issue=11 |pages=1062–1066 |date=Sep 24, 2018 |doi=10.1038/nbt.4245 |pmid=30247490 |pmc=6871539 |url=http://spiral.imperial.ac.uk/bitstream/10044/1/64952/11/nbt.4245.pdf |access-date=September 23, 2019 |archive-date=April 29, 2019 |archive-url=https://web.archive.org/web/20190429023635/http://spiral.imperial.ac.uk/bitstream/10044/1/64952/11/nbt.4245.pdf |url-status=live}}</ref><ref>{{cite journal |last1=Michael Le Page |title=Gene tool could halt malaria spread |journal=[[New Scientist]] |date=Sep 29, 2018 |url=https://www.newscientist.com/article/mg23931974-000-evolution-defying-dna-makes-mosquitoes-infertile-by-changing-their-sex |access-date=November 2, 2018 |archive-date=November 12, 2018 |archive-url=https://web.archive.org/web/20181112061002/https://www.newscientist.com/article/mg23931974-000-evolution-defying-dna-makes-mosquitoes-infertile-by-changing-their-sex/ |url-status=live }}</ref>
=== Repellents ===
{{main|Insect repellent}}
[[File:Insect repellents in Finnish store.jpg |thumb |[[Insect repellent|Mosquito repellents]] (including a [[mosquito coil]]) in a [[Finland|Finnish]] store]]
Insect repellents are applied on skin and give short-term protection against mosquito bites. The chemical [[DEET]] repels some mosquitoes and other insects.<ref>{{cite journal |last1=Syed |first1=Z. |last2=Leal |first2=W.S. |title=Mosquitoes smell and avoid the insect repellent DEET |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=105 |issue=36 |pages=13598–13603 |date=September 2008 |pmid=18711137 |pmc=2518096 |doi=10.1073/pnas.0805312105 |doi-access=free }}</ref> Some [[Centers for Disease Control and Prevention|CDC]]-recommended repellents are [[picaridin]], [[eucalyptus oil]] ([[P-Menthane-3,8-diol|PMD]]), and [[ethyl butylacetylaminopropionate]] (IR3535).<ref>{{cite web |publisher=[[Centers for Disease Control and Prevention]] |year=2009 |title=Updated Information regarding Insect Repellents |url=https://www.cdc.gov/ncidod/dvbid/westnile/repellentupdates.htm |access-date=2017-09-10 |archive-date=2013-05-12 |archive-url=https://web.archive.org/web/20130512034920/http://www.cdc.gov/ncidod/dvbid/westnile/repellentupdates.htm |url-status=live }}</ref> [[Pyrethrum]] (from ''[[Chrysanthemum]]'' species, particularly ''C. cinerariifolium'' and ''C. coccineum'') is an effective plant-based repellent.<ref>[[Rachel Nuwer|Nuwer, Rachel]], ''[https://www.scientificamerican.com/article/natural-mosquito-repellents-powers-finally-decoded/ Natural Mosquito Repellent's Powers Finally Decoded] {{Webarchive |url=https://web.archive.org/web/20210812180532/https://www.scientificamerican.com/article/natural-mosquito-repellents-powers-finally-decoded/ |date=2021-08-12 }}'', Scientific American 325, 2, 23 (August 2021)</ref> [[Electronic insect repellent]] devices that produce [[ultrasound]]s intended to keep away insects (and mosquitoes) are marketed. No [[United States Environmental Protection Agency|EPA]] or university study has shown that these devices prevent humans from being bitten by a mosquito.<ref>{{cite web |url=http://www.evidenceaid.org/wp-content/uploads/2013/08/Electronic_mosquito_repellents_for_preventing_mosquito_bites_and_malaria_infection.pdf |title=Electronic mosquito repellents for preventing mosquito bites and malaria infection |access-date=2018-09-19 |archive-date=2017-08-08 |archive-url=https://web.archive.org/web/20170808175758/http://www.evidenceaid.org/wp-content/uploads/2013/08/Electronic_mosquito_repellents_for_preventing_mosquito_bites_and_malaria_infection.pdf |url-status=live }}</ref>
=== Bites ===
{{further|Mosquito bite allergies}}
Mosquito bites lead to a variety of skin reactions and more seriously to [[mosquito bite allergies]].<ref name="Sawada Inoue Kawa 2017">{{cite journal |last1=Sawada |first1=Akihisa |last2=Inoue |first2=Masami |last3=Kawa |first3=Keisei |title=How we treat chronic active Epstein–Barr virus infection |journal=[[International Journal of Hematology]] |volume=105 |issue=4 |date=2017 |doi=10.1007/s12185-017-2192-6 |pages=406–418 |pmid=28210942 |s2cid=35297787 }}</ref>  Such [[hypersensitivity]] to mosquito bites is an excessive reaction to mosquito saliva proteins.<ref name="pmid24364549">{{cite journal |last=Juckett |first=G. |title=Arthropod bites |journal=American Family Physician |volume=88 |issue=12 |pages=841–847 |date=December 2013 |pmid=24364549 }}</ref> Numerous species of mosquito can trigger such reactions, including ''[[Aedes aegypti]]'', ''[[Aedes vexans|A. vexans]]'', ''[[Aedes albopictus|A. albopictus]]'', ''[[Anopheles sinensis]]'', ''[[Culex pipiens]]'',<ref name="Tatsuno Fujiyama Matsuoka Shimauchi 2016">{{cite journal |last1=Tatsuno |first1=Kazuki |last2=Fujiyama |first2=Toshiharu |last3=Matsuoka |first3=Hiroyuki |last4=Shimauchi |first4=Takatoshi |last5=Ito |first5=Taisuke |last6=Tokura |first6=Yoshiki |title=Clinical categories of exaggerated skin reactions to mosquito bites and their pathophysiology |journal=Journal of Dermatological Science |volume=82 |issue=3 |date=2016 |doi=10.1016/j.jdermsci.2016.04.010 |pages=145–152 |pmid=27177994 }}</ref> ''[[Aedes communis]]'', ''[[Anopheles stephensi]]'',<ref name="Peng Simons 2007">{{cite journal |last1=Peng |first1=Z. |last2=Simons |first2=F.E. |title=Advances in mosquito allergy |journal=[[Current Opinion in Allergy and Clinical Immunology]] |volume=7 |issue=4 |pages=350–354 |date=August 2007 |pmid=17620829 |doi=10.1097/ACI.0b013e328259c313 |s2cid=45260523 }}</ref> ''[[Culex quinquefasciatus|C. quinquefasciatus]]'', ''[[Culex tritaeniorhynchus|C. tritaeniorhynchus]]'',<ref name="pmid17169531">{{cite journal |last=Asada |first=H. |title=Hypersensitivity to mosquito bites: a unique pathogenic mechanism linking Epstein-Barr virus infection, allergy and oncogenesis |journal=[[Journal of Dermatological Science]] |volume=45 |issue=3 |pages=153–160 |date=March 2007 |pmid=17169531 |doi=10.1016/j.jdermsci.2006.11.002 }}</ref> and ''[[Ochlerotatus triseriatus]]''.<ref name="pmid23352522">{{cite journal |last1=Crisp |first1=H.C. |last2=Johnson |first2=K.S. |title=Mosquito allergy |journal=[[Annals of Allergy, Asthma & Immunology]] |volume=110 |issue=2 |pages=65–69 |date=February 2013 |pmid=23352522 |doi=10.1016/j.anai.2012.07.023 }}</ref> Cross-reactivity between salivary proteins of different mosquitoes implies that allergic responses may be caused by virtually any mosquito species.<ref name="pmid23442453">{{cite journal |last1=Singh |first1=S. |last2=Mann |first2=B.K. |title=Insect bite reactions |journal=Indian Journal of Dermatology, Venereology and Leprology |volume=79 |issue=2 |pages=151–164 |date=2013 |pmid=23442453 |doi=10.4103/0378-6323.107629 |doi-access=free }}</ref> Treatment can be with [[Antipruritic|anti-itch]] medications, including some taken orally, such as [[diphenhydramine]], or applied to the skin like [[antihistamine]]s or [[corticosteroids]] such as [[hydrocortisone]]. [[Aqueous ammonia]] (3.6%) also provides relief.<ref name="Ammonium">{{cite journal |last1=Zhai |first1=Hongbo |last2=Packman |first2=Elias W. |last3=Maiback |first3=Howard I. |title=Effectiveness of Ammonium Solution in Relieving Type I Mosquito Bite Symptoms: A Double-blind, Placebo-controlled Study |journal=Acta Dermato-Venereologica |date=21 July 1998 |volume=78 |issue=4 |pages=297–298 |doi=10.1080/000155598441918|pmid=9689301 |doi-access=free }}</ref> Both topical heat<ref>{{cite journal |last1=Müller |first1=C. |last2=Großjohann |first2=B. |last3=Fischer |first3=L. |title=The use of concentrated heat after insect bites/stings as an alternative to reduce swelling, pain, and pruritus: an open cohort-study at German beaches and bathing-lakes |journal=[[Clinical, Cosmetic and Investigational Dermatology]] |volume=4 |pages=191–196 |date=2011-12-15 |pmid=22253544 |pmc=3257884 |doi=10.2147/CCID.S27825 |doi-access=free }}</ref> and cold may be useful as treatments.<ref>{{Cite web |url=https://www.nhs.uk/conditions/insect-bites-and-stings/treatment/ |title=Treatment of Insect bites and stings |website=nhs.uk |access-date=2018-10-31 |date=2017-10-19 |archive-date=2018-10-31 |archive-url=https://web.archive.org/web/20181031173837/https://www.nhs.uk/conditions/insect-bites-and-stings/treatment/ |url-status=live }}</ref>
== In human culture ==
=== Greek mythology ===
[[File:Rackham-gnat and bull.jpg|thumb|[[Arthur Rackham]]'s illustration of the [[fable]] of "[[The Gnat and the Bull|The Bull and the Mosquito]]", 1912]]
[[Ancient Greek]] beast fables including "The Elephant and the Mosquito" and "[[The Gnat and the Bull|The Bull and the Mosquito]]", with the general moral that the large beast does not even notice the small one, derive ultimately from [[Mesopotamia]].<ref name="Adrados1999">{{cite book |last=Adrados |first=Francisco Rodríguez |author-link=Francisco Rodríguez Adrados |title=History of the Graeco-Latin Fable |url=https://books.google.com/books?id=093Gl8KEktMC&pg=PA324 |year=1999 |publisher=[[Brill Publishers]] |isbn=978-90-04-11454-8 |page=324 |access-date=2016-02-18 |archive-date=2016-05-28 |archive-url=https://web.archive.org/web/20160528074423/https://books.google.com/books?id=093Gl8KEktMC&pg=PA324 |url-status=live }}</ref>
=== Origin myths ===
The peoples of [[Siberia]] have [[origin myth]]s surrounding the mosquito. One [[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 [[Samoyedic peoples|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 Mountains|Altai]] had a variant of the same myth, involving the fragments of the dead giant, ''Andalma-Muus'', becoming mosquitoes and other insects.<ref>{{citation |volume=4 |year=1927 |first=Uno |last=Holmberg |author-link=Uno Holmberg |title=Finno-Ugric and Siberian |url=https://archive.org/details/mythologyofallra41gray |work=The Mythology of All Races |at=IX. "The Origin of the Mosquito", p.386 |publisher=Marshall Jones Company }}</ref>
[[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 [[Hungry ghost#In Japan|pretas]]".<ref>{{cite book |last=Hearn |first=Lafcadio |author-link=Lafcadio Hearn |chapter=Mosquitoes |title=Kwaidan: Stories and Studies of Strange Things |title-link=Kwaidan: Stories and Studies of Strange Things |publisher=[[Dover Publications]] |orig-year=1968 |year=2020 |isbn=978-1420967517 |pages=72–74}}</ref>
=== Modern era ===
[[File:Winsor McCay (1912) How a Mosquito Operates.webm|thumb|alt=Silent animated black-and-white film of a giant mosquito drinking blood from a sleeping man.|thumbtime=00:36|''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.<ref>{{cite book |last=Webster |first=Chris |title=Action Analysis for Animators |url=https://books.google.com/books?id=jylUYRFfcywC |year=2012 |publisher=[[Focal Press]] |isbn=978-0-240-81218-2 |access-date=2022-09-04 |archive-date=2021-11-04 |archive-url=https://web.archive.org/web/20211104181628/https://books.google.com/books?id=jylUYRFfcywC |url-status=live}}</ref> It depicts a giant mosquito tormenting a sleeping man.<ref>{{cite book |last=Canemaker |first=John |author-link=John Canemaker |title=Winsor McCay: His Life and Art |url=https://books.google.com/books?id=vs82AQAAIAAJ |year=2005 |publisher=[[Abrams Books]] |isbn=978-0-8109-5941-5 |page=165 }}</ref>
Twelve ships of the [[Royal Navy]] have borne the name HMS ''Mosquito'' or the archaic form of the name, HMS ''Musquito''.<ref>{{Cite Colledge2006}}, "Mosquito" and "Musquito".</ref>
The [[de Havilland Mosquito]] was a high-speed aircraft manufactured between 1940 and 1950, and used in many roles.<ref>{{cite web |title=De Havilland Mosquito |url=http://www.aviation-history.com/dehavilland/mosquito.html |publisher=The Aviation History Online Museum |access-date=21 November 2015 |archive-date=11 January 2017 |archive-url=https://web.archive.org/web/20170111050938/http://www.aviation-history.com/dehavilland/mosquito.html |url-status=live }}</ref>
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.<ref>{{Cite news|url=https://www.bbc.com/news/blogs-news-from-elsewhere-33525015|title=Russia: Mosquitoes honoured at annual festival|work=BBC News |date=July 14, 2015}}</ref>
{{Clear}}
== References ==
{{reflist}}
== Further reading ==
* {{cite book |last=Winegard |first=Timothy Charles |title=The mosquito: a human history of our deadliest predator |isbn=978-1524743413 |oclc=1111638283 |year=2019 |publisher=Penguin Random House |ref=none}}
== External links ==
{{Commons category}}
* {{wikiquote-inline|Mosquitoes}}
* [https://fmel.ifas.ufl.edu/mosquito-guide/ Mosquito at IFAS]
* {{Internet Archive film clip|id=life_cycle_of_the_mosquito|description= describing The Life Cycle of the Mosquito}}
* [https://en.wikibooks.org/wiki/Parasitic_Insects,_Mites_and_Ticks:_Genera_of_Medical_and_Veterinary_Importance/Mosquitoes_and_similar Parasitic Insects, Mites and Ticks: Genera of Medical and Veterinary Importance] Wikibooks
{{Diptera|1}}
{{Taxonbar|from=Q7367}}
{{Authority control}}
{{DEFAULTSORT:Mosquito}}
[[Category:Culicidae|.]]
[[Category:Insect vectors of human pathogens]]
[[Category:Articles containing video clips]]
[[Category:Ectoparasites]]
[[Category:Extant Jurassic first appearances]]
[[Category:Hematophages]]
[[Category:Mosquito genera|.]]
[[Category:Insects in culture]]
[[Category:Aquatic insects]]
[[Category:Hazards of outdoor recreation]]