MOSQUITOES are small, midge -like flies that constitute the family
CULICIDAE. Females of most species are ectoparasites , whose tube-like
mouthparts (called a proboscis ) pierce the hosts ' skin to consume
blood . The word "mosquito" (formed by mosca and diminutive -ito) is
Spanish for "little fly ". Thousands of species feed on the blood of
various kinds of hosts, mainly vertebrates , including mammals , birds
, reptiles , amphibians , and even some kinds of fish . Some
mosquitoes also attack invertebrates , mainly other arthropods .
Though the loss of blood is seldom of any importance to the victim,
the saliva of the mosquito often causes an irritating rash that is a
serious nuisance. Much more serious though, are the roles of many
species of mosquitoes as vectors of diseases. In passing from host to
host, some transmit extremely harmful infections such as malaria ,
yellow fever ,
West Nile virus
West Nile virus , dengue fever ,
Zika virus and other arboviruses , rendering it the
deadliest animal family in the world.
* 1 Taxonomy and evolution
* 1.1 Subfamilies
* 1.2 Genera
* 3 Lifecycle
* 3.1 Eggs and oviposition
* 3.4 Adult
* 4 Feeding by adults
* 4.1 Hosts of blood-feeding mosquito species
* 4.2 Mouthparts
* 4.4 Egg development and blood digestion
* 5 Distribution
* 5.1 Means of dispersal
* 7 Control
* 7.1 Repellents
* 8 Bites and treatment
* 9 In human culture
* 10 Notes
* 11 References
* 12 Further reading
* 13 External links
TAXONOMY AND EVOLUTION
The oldest known mosquito with an anatomy similar to modern species
was found in 79-million-year-old Canadian amber from the
An older sister species with more primitive features was found in
Burmese amber that is 90 to 100 million years old. Two mosquito
fossils have been found that show very little morphological change in
modern mosquitoes against their counterpart from 46 million years ago.
These fossils are also the oldest ever found to have blood preserved
within their abdomens. Despite no fossils being found earlier than
the Cretaceous, recent studies suggest that the earliest divergence of
mosquitoes between the lineages leading to
Anophelinae and Culicinae
occurred 226 million years ago.
The Old and New World
Anopheles species are believed to have
subsequently diverged about 95 million years ago.
Anopheles gambiae is currently undergoing speciation
into the M(opti) and S(avanah) molecular forms. Consequently, some
pesticides that work on the M form no longer work on the S form. Over
3,500 species of the Culicidae have already been described. They are
generally divided into two subfamilies which in turn comprise some 43
genera. These figures are subject to continual change, as more species
are discovered, and as DNA studies compel rearrangement of the
taxonomy of the family. The two main subfamilies are the Anophelinae
and Culicinae, with their genera as shown in the subsection below.
The distinction is of great practical importance because the two
subfamilies tend to differ in their significance as vectors of
different classes of diseases. Roughly speaking, arboviral diseases
such as yellow fever and dengue fever tend to be transmitted by
CULICINE species, not necessarily in the genus Culex. Some transmit
various species of avian malaria , but it is not clear that they ever
transmit any form of human malaria. Some species do however transmit
various forms of filariasis , much as many
Anopheline mosquitoes, again not necessarily in the genus Anopheles,
sometimes bear pathogenic arboviruses, but it is not yet clear that
they ever transmit them as effective vectors. However, all the most
important vectors of human malaria are Anopheline.
Mosquitoes are members of a family of nematocerid flies : the
Culicidae (from the
Latin culex, genitive culicis, meaning "midge" or
"gnat"). Superficially, mosquitoes resemble crane flies (family
Tipulidae ) and chironomid flies (family
Chironomidae ). In
particular, the females of many species of mosquitoes are blood-eating
pests and dangerous vectors of diseases , whereas members of the
Chironomidae and Tipulidae are not. Many species of
mosquitoes are not blood eaters and of those that are, many create a
"high to low pressure" in the blood to obtain it and do not transmit
disease. Also, in the bloodsucking species, only the females suck
blood. Furthermore, even among mosquitoes that do carry important
diseases, neither all species of mosquitoes, nor all strains of a
given species transmit the same kinds of diseases, nor do they all
transmit the diseases under the same circumstances; their habits
differ. For example, some species attack people in houses, and others
prefer to attack people walking in forests. Accordingly, in managing
public health, knowing which species or even which strain of mosquito
one is dealing with is important.
Over 3,500 species of mosquitoes have already been described from
various parts of the world. Some mosquitoes that bite humans
routinely act as vectors for a number of infectious diseases affecting
millions of people per year. Others that do not routinely bite
humans, but are the vectors for animal diseases, may become disastrous
agents for zoonosis of new diseases when their habitats are disturbed,
for instance by sudden deforestation.
Anatomy of a
Culex larva Image of pitcher plant mosquito
Wyeomyia smithii , showing segmentation and partial anatomy of
Like all flies, mosquitoes go through four stages in their
lifecycles: egg , larva , pupa , and adult or imago . In most species,
adult females lay their eggs in stagnant water; some lay eggs near the
water's edge; others attach their eggs to aquatic plants. Each species
selects the situation of the water into which it lays its eggs and
does so according to its own ecological adaptations. Some are
generalists and are not very fussy. Some breed in lakes, some in
temporary puddles. Some breed in marshes, some in salt-marshes. Among
those that breed in salt water, some are equally at home in fresh and
salt water up to about one-third the concentration of seawater,
whereas others must acclimatize themselves to the salinity. Such
differences are important because certain ecological preferences keep
mosquitoes away from most humans, whereas other preferences bring them
right into houses at night.
Some species of mosquitoes prefer to breed in phytotelmata (natural
reservoirs on plants), such as rainwater accumulated in holes in tree
trunks, or in the leaf-axils of bromeliads . Some specialize in the
liquid in pitchers of particular species of pitcher plants , their
larvae feeding on decaying insects that had drowned there or on the
associated bacteria; the genus
Wyeomyia provides such examples — the
Wyeomyia smithii breeds only in the pitchers of Sarracenia
However, some of the species of mosquitoes that are adapted to
breeding in phytotelmata are dangerous disease vectors. In nature,
they might occupy anything from a hollow tree trunk to a cupped leaf.
Such species typically take readily to breeding in artificial water
containers. Such casual puddles are important breeding places for some
of the most serious disease vectors, such as species of
transmit dengue and yellow fever. Some with such breeding habits are
disproportionately important vectors because they are well-placed to
pick up pathogens from humans and pass them on. In contrast, no matter
how voracious, mosquitoes that breed and feed mainly in remote
wetlands and salt marshes may well remain uninfected, and if they do
happen to become infected with a relevant pathogen, might seldom
encounter humans to infect, in turn.
The first three stages—egg, larva, and pupa—are largely aquatic.
These stages typically last 5 to 14 days, depending on the species and
the ambient temperature, but there are important exceptions.
Mosquitoes living in regions where some seasons are freezing or
waterless spend part of the year in diapause ; they delay their
development, typically for months, and carry on with life only when
there is enough water or warmth for their needs. For instance,
Wyeomyia larvae typically get frozen into solid lumps of ice during
winter and only complete their development in spring. The eggs of some
Aedes remain unharmed in diapause if they dry out, and
hatch later when they are covered by water.
Eggs hatch to become larvae , which grow until they are able to
change into pupae . The adult mosquito emerges from the mature pupa as
it floats at the water surface. Bloodsucking mosquitoes, depending on
species, sex, and weather conditions, have potential adult lifespans
ranging from as short as a week to as long as several months.
Some species can overwinter as adults in diapause.
EGGS AND OVIPOSITION
Mosquito habits of oviposition , the ways in which they lay their
eggs, vary considerably between species, and the morphologies of the
eggs vary accordingly. The simplest procedure is that followed by many
Anopheles ; like many other gracile species of aquatic
insects, females just fly over the water, bobbing up and down to the
water surface and dropping eggs more or less singly. The bobbing
behavior occurs among some other aquatic insects as well, for example
mayflies and dragonflies ; it is sometimes called "dapping ". The eggs
Anopheles species are roughly cigar-shaped and have floats down
their sides. Females of many common species can lay 100–200 eggs
during the course of the adult phase of their lifecycles. Even with
high egg and intergenerational mortality, over a period of several
weeks, a single successful breeding pair can create a population of
thousands. An egg raft of a
Culex species, partly broken, showing
individual egg shapes
Some other species, for example members of the genus
Mansonia , lay
their eggs in arrays, attached usually to the under-surfaces of
waterlily pads. Their close relatives, the genus Coquillettidia, lay
their eggs similarly, but not attached to plants. Instead, the eggs
form layers called "rafts" that float on the water. This is a common
mode of oviposition, and most species of
Culex are known for the
habit, which also occurs in some other genera, such as
Anopheles eggs may on occasion cluster together on the
water, too, but the clusters do not generally look much like compactly
glued rafts of eggs.
In species that lay their eggs in rafts, rafts do not form
adventitiously; the female
Culex settles carefully on still water with
its hind legs crossed, and as it lays the eggs one by one, it twitches
to arrange them into a head-down array that sticks together to form
Aedes females generally drop their eggs singly, much as
but not as a rule into water. Instead, they lay their eggs on damp mud
or other surfaces near the water's edge. Such an oviposition site
commonly is the wall of a cavity such as a hollow stump or a container
such as a bucket or a discarded vehicle tire. The eggs generally do
not hatch until they are flooded, and they may have to withstand
considerable desiccation before that happens. They are not resistant
to desiccation straight after oviposition, but must develop to a
suitable degree first. Once they have achieved that, however, they can
enter diapause for several months if they dry out. Clutches of eggs of
the majority of mosquito species hatch as soon as possible, and all
the eggs in the clutch hatch at much the same time. In contrast, a
Aedes eggs in diapause tends to hatch irregularly over an
extended period of time. This makes it much more difficult to control
such species than those mosquitoes whose larvae can be killed all
together as they hatch. Some
Anopheles species do also behave in such
a manner, though not to the same degree of sophistication.
The mosquito larva has a well-developed head with mouth brushes used
for feeding, a large thorax with no legs, and a segmented abdomen .
Larvae breathe through spiracles located on their eighth abdominal
segments, or through a siphon, so must come to the surface frequently.
The larvae spend most of their time feeding on algae , bacteria , and
other microbes in the surface microlayer .
They dive below the surface only when disturbed. Larvae swim either
through propulsion with their mouth brushes, or by jerky movements of
their entire bodies, giving them the common name of "wigglers" or
Larvae develop through four stages, or instars , after which they
metamorphose into pupae . At the end of each instar, the larvae molt,
shedding their skins to allow for further growth.
Mosquito larvae and pupa resting at water surface
Anopheles larva from southern Germany, about 8 mm long
Aedes aegypti larva
Culex larva and pupa
Culex larvae plus one pupa
Electron micrograph of a mosquito egg
As seen in its lateral aspect, the mosquito pupa is comma-shaped. The
head and thorax are merged into a cephalothorax , with the abdomen
curving around underneath. The pupa can swim actively by flipping its
abdomen, and it is commonly called a "tumbler" because of its swimming
action. As with 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. However, pupae do not
feed during this stage; typically they pass their time hanging from
the surface of the water by their respiratory trumpets. If alarmed,
say by a passing shadow, they nimbly swim downwards by flipping their
abdomens in much the same way as the larvae do. If undisturbed, they
soon float up again.
After a few days or longer, depending on the temperature and other
circumstances, the pupa rises to the water surface, the dorsal surface
of its cephalothorax splits, and the adult mosquito emerges. The pupa
is less active than the larva because it does not feed, whereas the
larva feeds constantly.
The period of development from egg to adult varies among species and
is strongly influenced by ambient temperature. Some species of
mosquitoes can develop from egg to adult in as few as five days, but a
more typical period of development in tropical conditions would be
some 40 days or more for most species. The variation of the body size
in adult mosquitoes depends on the density of the larval population
and food supply within the breeding water. Anatomy of an adult
Adult mosquitoes usually mate within a few days after emerging from
the pupal stage. In most species, the males form large swarms ,
usually around dusk, and the females fly into the swarms to mate.
Males typically live for about 5–7 days, feeding on nectar and
other sources of sugar . After obtaining a full blood meal, the female
will rest for a few days while the blood is digested and eggs are
developed. This process depends on the temperature, but usually takes
two to three days in tropical conditions. Once the eggs are fully
developed, the female lays them and resumes host-seeking.
The cycle repeats itself until the female dies. While females can
live longer than a month in captivity, most do not live longer than
one to two weeks in nature. Their lifespans depend on temperature,
humidity, and their ability to successfully obtain a blood meal while
avoiding host defenses and predators.
The length of the adult is typically between 3mm and 6mm. The
smallest known mosquitoes are around 2 mm (0.1 in), and the largest
around 19 mm (0.7 in). Mosquitoes typically weigh around 5 mg. All
mosquitoes have slender bodies with three segments: a head, a thorax
and an abdomen.
The head is specialized for receiving sensory information and for
feeding. It has eyes and a pair of long, many-segmented antennae . The
antennae are important for detecting host odors, as well as odors of
breeding sites where females lay eggs. In all mosquito species, the
antennae of the males in comparison to the females are noticeably
bushier and contain auditory receptors to detect the characteristic
whine of the females. Adult yellow fever mosquito
Aedes aegypti ,
typical of subfamily
Culicinae . Note bushy antennae and longer palps
of male on left vs. females at right.
The compound eyes are distinctly separated from one another. Their
larvae only possess a pit-eye ocellus. The compound eyes of adults
develop in a separate region of the head. New ommatidia are added in
semicircular rows at the rear of the eye. During the first phase of
growth, this leads to individual ommatidia being square, but later in
development they become hexagonal. The hexagonal pattern will only
become visible when the carapace of the stage with square eyes is
The head also has an elongated, forward-projecting, stinger-like
proboscis used for feeding, and two sensory palps. The maxillary palps
of the males are longer than their proboscises, whereas the females’
maxillary palps are much shorter. In typical bloodsucking species, the
female has an elongated proboscis.
The thorax is specialized for locomotion. Three pairs of legs and a
pair of wings are attached to the thorax. The insect wing is an
outgrowth of the exoskeleton. The
Anopheles mosquito can fly for up to
four hours continuously at 1 to 2 km/h (0.6–1 mph), traveling up to
12 km (7.5 mi) in a night. Males beat their wings between 450 and 600
times per second.
The abdomen is specialized for food digestion and egg development;
the abdomen of a mosquito can hold three times its own weight in
blood. This segment expands considerably when a female takes a blood
meal. The blood is digested over time, serving as a source of protein
for the production of eggs, which gradually fill the abdomen.
FEEDING BY ADULTS
Aedes aegypti , a common vector of dengue fever and yellow fever
Typically, both male and female mosquitoes feed on nectar and plant
juices, but in many species the mouthparts of the females are adapted
for piercing the skin of animal hosts and sucking their blood as
ectoparasites . In many species, the female needs to obtain nutrients
from a blood meal before it can produce eggs, whereas in many other
species, it can produce more eggs after a blood meal. A mosquito has a
variety of ways of finding its prey, including chemical, visual, and
heat sensors. Both plant materials and blood are useful sources of
energy in the form of sugars, and blood also supplies more
concentrated nutrients, such as lipids , but the most important
function of blood meals is to obtain proteins as materials for egg
The feeding preferences of mosquitoes include those with type O
blood, heavy breathers, those with a lot of skin bacteria, people with
a lot of body heat, and the pregnant. Individuals' attractiveness to
mosquitoes also has a heritable , genetically-controlled component.
When a female reproduces without such parasitic meals, it is said to
practice autogenous reproduction, as in
Toxorhynchites ; otherwise,
the reproduction may be termed anautogenous , as occurs in mosquito
species that serve as disease vectors, particularly
Anopheles and some
of the most important disease vectors in the genus Aedes. In contrast,
some mosquitoes, for example, many Culex, are partially anautogenous:
they do not need a blood meal for their first cycle of egg production,
which they produce autogenously; however, subsequent clutches of eggs
are produced anautogenously, at which point their disease vectoring
activity becomes operative. Here an
Anopheles stephensi female
is engorged with blood and beginning to pass unwanted liquid fractions
of the blood to make room in its gut for more of the solid nutrients.
With regard to host location, female mosquitoes hunt their blood host
by detecting organic substances such as carbon dioxide (CO2) and
1-octen-3-ol produced from the host, and through visual recognition.
Mosquitoes prefer some people over others. The preferred victim's
sweat simply smells better than others' because of the proportions of
the carbon dioxide, octenol and other compounds that make up body
odor. The most powerful semiochemical that triggers the keen sense of
Culex quinquefasciatus is nonanal . Another compound
identified in human blood that attracts mosquitoes is sulcatone or
6-methyl-5-hepten-2-one, especially for
Aedes aegypti mosquitoes with
the odor receptor gene Or4. 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. In Aedes, the search for a
host takes place in two phases. First, the mosquito exhibits a
nonspecific searching behavior until the perception of host
stimulants, then it follows a targeted approach.
Most mosquito species are crepuscular (dawn or dusk ) feeders. During
the heat of the day, most mosquitoes rest in a cool place and wait for
the evenings, although they may still bite if disturbed. Some
species, such as the
Asian tiger mosquito , are known to fly and feed
Prior to and during blood feeding, blood-sucking mosquitoes inject
saliva into the bodies of their source(s) of blood. This saliva serves
as an anticoagulant ; without it one might expect the female
mosquito's proboscis to become clogged with blood clots. The saliva
also is the main route by which mosquito physiology offers passenger
pathogens access to the hosts' interior. The salivary glands are a
major target to most pathogens, whence they find their way into the
host via the stream of saliva.
The bump left on the victim's skin after a mosquito bites is called a
wheal , which is caused by histamines trying to fight off the protein
left by the attacking insect.
Mosquitoes of the genus
Toxorhynchites never drink blood. This genus
includes the largest extant mosquitoes, the larvae of which prey on
the larvae of other mosquitoes. These mosquito eaters have been used
in the past as mosquito control agents, with varying success.
HOSTS OF BLOOD-FEEDING MOSQUITO SPECIES
Mosquitoes feeding on a reptile Play media Video of
Anopheline mosquito locating and feeding on a caterpillar
Many, if not all, blood-sucking species of mosquitoes are fairly
selective feeders that specialise in particular host species, though
they often relax their selectivity when they experience severe
competition for food, defensive activity on the part of the hosts, or
starvation. Some species feed selectively on monkeys, while others
prefer particular kinds of birds, but they become less selective as
conditions become more difficult. For example,
Culiseta melanura sucks
the blood of passerine birds for preference and such birds are
typically the main reservoir of the Eastern equine encephalitis virus
in North America. Early in the season while mosquito numbers are low,
they concentrate on passerine hosts, but as mosquito numbers rise and
the birds are forced to defend themselves more vigorously, the
mosquitoes become less selective in attacking their avian hosts. Soon
the mosquitoes begin attacking mammals more readily, thereby becoming
the major vector of the virus, and causing epidemics of the disease,
most conspicuously in humans and horses.
Even more dramatically, in most of its range in North America, the
main vector for the
Western equine encephalitis virus is Culex
tarsalis , because it is known to feed variously on mammals, birds,
reptiles, and amphibians. Even fish may be attacked by some mosquito
species if they expose themselves above water level, as mudskippers
Some species of blood-sucking flies, such as many of the
Ceratopogonidae , will attack large, live insects and suck their
haemolymph and others, such as the so-called "jackal flies"
Milichiidae ), will attack the recently dead prey of say, crab
Thomisidae ), but in the late 1960s it was reported that
some species of anautogenous mosquitoes would feed on the haemolymph
of caterpillars. Other observations include mosquitoes feeding on
cicadas, and mantids. More recently it has been shown that
malaria-transmitting mosquitoes will actively seek out some species of
caterpillars and feed on their haemolymph, and do so to the
caterpillar's apparent physical detriment.
Mosquito mouthparts are very specialized, particularly those of the
females, which in most species are adapted to piercing skin and then
sucking blood. Apart from bloodsucking, the females generally also
drink assorted fluids rich in dissolved sugar, such as nectar and
honeydew, to obtain the energy they need. For this, their
blood-sucking mouthparts are perfectly adequate. In contrast, male
mosquitoes are not bloodsuckers; they only drink sugary fluids.
Accordingly, their mouthparts do not require the same degree of
specialization as those of females.
Externally, the most obvious feeding structure of the mosquito is the
proboscis. More specifically, the visible part of the proboscis is the
labium , which forms the sheath enclosing the rest of the mouthparts.
When the mosquito first lands on a potential host, its mouthparts will
be enclosed entirely in this sheath, and it will touch the tip of the
labium to the skin in various places. Sometimes, it will begin to bite
almost straight away, while other times, it will prod around,
apparently looking for a suitable place. Occasionally, it will wander
for a considerable time, and eventually fly away without biting.
Presumably, this probing is a search for a place with easily
accessible blood vessels, but the exact mechanism is not known. It is
known that there are two taste receptors at the tip of the labium
which may well play a role.
The female mosquito does not insert its labium into the skin; it
bends back into a bow when the mosquito begins to bite. The tip of the
labium remains in contact with the skin of the victim, acting as a
guide for the other mouthparts. In total, there are six mouthparts
besides the labium: two mandibles , two maxillae , the hypopharynx ,
and the labrum .
The mandibles and the maxillae are used for piercing the skin. The
mandibles are pointed, while the maxillae end in flat, toothed
"blades". To force these into the skin, the mosquito moves its head
backwards and forwards. On one movement, the maxillae are moved as far
forward as possible. On the opposite movement, the mandibles are
pushed deeper into the skin by levering against the maxillae. The
maxillae do not slip back because the toothed blades grip the skin.
The hypopharynx and the labrum are both hollow.
anticoagulant is pumped down the hypopharynx to prevent clotting, and
blood is drawn up the labrum.
To understand the mosquito mouthparts, it is helpful to draw a
comparison with an insect that chews food, such as a dragonfly . A
dragonfly has two mandibles, which are used for chewing, and two
maxillae, which are used to hold the food in place as it is chewed.
The labium forms the floor of the dragonfly's mouth, the labrum forms
the top, while the hypopharynx is inside the mouth and is used in
swallowing. Conceptually, then, the mosquito's proboscis is an
adaptation of the mouthparts that occur in other insects. The labium
still lies beneath the other mouthparts, but also enfolds them, and it
has been extended into a proboscis. The maxillae still "grip" the
"food" while the mandibles "bite" it. The top of the mouth, the
labrum, has developed into a channeled blade the length of the
proboscis, with a cross-section like an inverted "U". Finally, the
hypopharynx has extended into a tube that can deliver saliva at the
end of the proboscis. Its upper surface is somewhat flattened so, when
pressed against it, the labrum forms a closed tube for conveying blood
from the victim.
For the mosquito to obtain a blood meal, it must circumvent the
vertebrate 's physiological responses. The mosquito, as with all
blood-feeding arthropods , has mechanisms to effectively block the
hemostasis system with their saliva, which contains a mixture of
Mosquito saliva negatively affects vascular
constriction , blood clotting , platelet aggregation, angiogenesis and
immunity , and creates inflammation . Universally, hematophagous
arthropod saliva contains at least one anti-clotting, one
anti-platelet, and one vasodilatory substance.
Mosquito saliva also
contains enzymes that aid in sugar feeding and antimicrobial agents
to control bacterial growth in the sugar meal. The composition of
mosquito saliva is relatively simple, as it usually contains fewer
than 20 dominant proteins . Despite the great strides in knowledge of
these molecules and their role in blood feeding achieved recently,
scientists still cannot ascribe functions to more than half of the
molecules found in arthropod saliva. One promising application is the
development of anti-clotting drugs, such as clotting inhibitors and
capillary dilators, that could be useful for cardiovascular disease.
It is now well recognized that feeding ticks , sandflies , and, more
recently, mosquitoes, have an ability to modulate the immune response
of the animals (hosts) on which they feed. The presence of this
activity in vector saliva is a reflection of the inherent overlapping
and interconnected nature of the host hemostatic and
inflammatory/immunological responses and the intrinsic need to prevent
these host defenses from disrupting successful feeding. The mechanism
for mosquito saliva-induced alteration of the host immune response is
unclear, but the data have become increasingly convincing that such an
effect occurs. Early work described a factor in saliva that directly
TNF-α release, but not antigen-induced histamine
secretion, from activated mast cells . Experiments by Cross et al.
(1994) demonstrated that the inclusion of Ae. aegypti mosquito saliva
into naïve cultures led to a suppression of interleukin (IL)-2 and
IFN-γ production, while the cytokines IL-4 and IL-5 are unaffected by
mosquito saliva. Cellular proliferation in response to IL-2 is
clearly reduced by prior treatment of cells with mosquito salivary
gland extract. Correspondingly, activated splenocytes isolated from
mice fed upon by either Ae. aegypti or Cx. pipiens mosquitoes produce
markedly higher levels of IL-4 and IL-10 concurrent with suppressed
IFN-γ production. Unexpectedly, this shift in cytokine expression is
observed in splenocytes up to 10 days after mosquito exposure,
suggesting natural feeding of mosquitoes can have a profound,
enduring, and systemic effect on the immune response.
T cell populations are decidedly susceptible to the suppressive
effect of mosquito saliva, showing increased mortality and decreased
division rates. Parallel work by Wasserman et al. (2004) demonstrated
that T and
B cell proliferation was inhibited in a dose dependent
manner with concentrations as low as 1/7 of the saliva in a single
mosquito. Depinay et al. (2005) observed a suppression of
T cell responses mediated by mosquito saliva and
dependent on mast cells and IL-10 expression.
A 2006 study suggests mosquito saliva can also decrease expression of
interferon −α/β during early mosquito-borne virus infection. The
contribution of type I interferons (IFN) in recovery from infection
with viruses has been demonstrated in vivo by the therapeutic and
prophylactic effects of administration of IFN inducers or IFN itself,
and different research suggests mosquito saliva exacerbates West Nile
virus infection, as well as other mosquito-transmitted viruses.
EGG DEVELOPMENT AND BLOOD DIGESTION
Female mosquitoes use two very different food sources. They need
sugar for energy, which is taken from sources such as nectar, and they
need blood as a source of protein for egg development. Because biting
is risky and suitable hosts may be difficult to find, mosquitoes take
as much blood as possible when they have the opportunity. Digesting
large volumes of blood takes time, requiring the use of energy from
sugars during the feeding process. To avoid this problem, mosquitoes
possess a digestive system which can store both food types, giving
access to both as needed. When the mosquito drinks a sugar solution,
it is directed to a crop . The crop can release sugar into the stomach
as it is required. At the same time, the stomach never becomes full of
sugar solution, which would prevent the mosquito taking a blood meal.
Blood is directed straight into the mosquito's stomach. In species
that feed on mammalian or avian blood, hosts whose blood pressure is
high, the mosquito feeds selectively from active blood vessels, where
the pressure assists in filling the gut rapidly.
Upon completion of feeding, the mosquito will withdraw her proboscis
, and as the gut fills up, the stomach lining secretes a peritrophic
membrane that surrounds the blood. This membrane keeps the blood
separate from anything else in the stomach. However, like certain
other insects that survive on dilute, purely liquid diets, notably
many of the
Hemiptera , many adult mosquitoes must excrete unwanted
aqueous fractions even as they feed. (See the photograph of a feeding
Anopheles stephensi: Note that the excreted droplet patently is not
whole blood, being far more dilute). As long as they are not
disturbed, this permits mosquitoes to continue feeding until they have
accumulated a full meal of nutrient solids. As a result, a mosquito
replete with blood can continue to absorb sugar, even as the blood
meal is slowly digested over a period of several days. Once blood is
in the stomach, the midgut of the female synthesizes proteolytic
enzymes that hydrolyze the blood proteins into free amino acids. These
are used as building blocks for the synthesis of egg yolk proteins.
In the mosquito
Anopheles stephensi Liston, trypsin activity is
restricted entirely to the posterior midgut lumen. No trypsin activity
occurs before the blood meal, but activity increases continuously up
to 30 hours after feeding, and subsequently returns to baseline levels
by 60 hours. Aminopeptidase is active in the anterior and posterior
midgut regions before and after feeding. In the whole midgut, activity
rises from a baseline of approximately three enzyme units (EU) per
midgut to a maximum of 12 EU at 30 hours after the blood meal,
subsequently falling to baseline levels by 60 hours. A similar cycle
of activity occurs in the posterior midgut and posterior midgut lumen,
whereas aminopeptidase in the posterior midgut epithelium decreases in
activity during digestion. Aminopeptidase in the anterior midgut is
maintained at a constant, low level, showing no significant variation
with time after feeding. Alpha-glucosidase is active in anterior and
posterior midguts before and at all times after feeding. In whole
midgut homogenates, alpha-glucosidase activity increases slowly up to
18 hours after the blood meal, then rises rapidly to a maximum at 30
hours after the blood meal, whereas the subsequent decline in activity
is less predictable. All posterior midgut activity is restricted to
the posterior midgut lumen. Depending on the time after feeding,
greater than 25% of the total midgut activity of alpha-glucosidase is
located in the anterior midgut. After blood meal ingestion, proteases
are active only in the posterior midgut. Trypsin is the major primary
hydrolytic protease and is secreted into the posterior midgut lumen
without activation in the posterior midgut epithelium. Aminoptidase
activity is also luminal in the posterior midgut, but cellular
aminopeptidases are required for peptide processing in both anterior
and posterior midguts. Alpha-glucosidase activity is elevated in the
posterior midgut after feeding in response to the blood meal, whereas
activity in the anterior midgut is consistent with a nectar-processing
role for this midgut region.
Female Ochlerotatus notoscriptus feeding on a human arm,
In the sense of the entire family Culicidae, mosquitoes are
cosmopolitan ; in every land region except for Antarctica and a few
islands, mainly in polar or subpolar climates , at least some species
of mosquito will be present.
Iceland is such an island, being
essentially free of mosquitoes. 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
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 and may take up to
300 ml of blood per day from each animal in a caribou herd.
The absence of mosquitoes from
Iceland and similar regions is
probably because of quirks of their climate, which differs in some
respects from mainland regions. At the start of the uninterrupted
continental winter of Greenland and the northern regions of Eurasia
and America, the pupa enters diapause under the ice that covers
sufficiently deep water. The imago ecloses only after the ice breaks
in late spring. In
Iceland however, the weather is less predictable.
In mid-winter it frequently warms up suddenly, causing the ice to
break, but then to freeze again after a few days. By that time the
mosquitoes will have emerged from their pupae, but the new freeze sets
in before they can complete their life cycle. Any anautogenous adult
mosquito would need a host to supply a blood meal before it could lay
viable eggs; it would need time to mate, mature the eggs and oviposit
in suitable wetlands. These requirements would not be realistic in
Iceland and in fact the absence of mosquitoes from such subpolar
islands is in line with the islands' low biodiversity;
fewer than 1,500 described species of insects, many of them probably
accidentally introduced by human agency. In
Iceland most ectoparasitic
insects live in sheltered conditions or actually on mammals; examples
include lice, fleas and bedbugs, in whose living conditions freezing
is no concern, and most of which were introduced inadvertently by
Some other aquatic Diptera, such as
Simuliidae , do survive in
Iceland, but their habits and adaptations differ from those of
Simuliidae for example, though they, like mosquitoes, are
bloodsuckers, generally inhabit stones under running water that does
not readily freeze and which is totally unsuited to mosquitoes;
mosquitoes are generally not adapted to running water.
Eggs of species of mosquitoes from the temperate zones are more
tolerant of cold than the eggs of species indigenous to warmer
regions. Many even tolerate subzero temperatures. In addition,
adults of some species can survive the winter by taking shelter in
suitable microhabitats such as buildings or hollow trees.
MEANS OF DISPERSAL
Worldwide introduction of various mosquito species over large
distances into regions where they are not indigenous has occurred
through human agencies, primarily on sea routes, in which the eggs,
larvae, and pupae inhabiting water-filled used tires and cut flowers
are transported. However, apart from sea transport, mosquitoes have
been effectively carried by personal vehicles, delivery trucks,
trains, and aircraft. Man-made areas such as storm water retention
basins , or storm drains also provide sprawling sanctuaries.
Sufficient quarantine measures have proven difficult to implement. In
addition, outdoor pool areas make a perfect place for them to grow.
Anopheles albimanus mosquito feeding on a human arm – this
mosquito is a vector of malaria , and mosquito control is a very
effective way of reducing the incidence of malaria. Main article:
Mosquitoes can act as vectors for many disease-causing viruses and
parasites . Infected mosquitoes carry these organisms from person to
person without exhibiting symptoms themselves. Mosquito-borne diseases
* Viral diseases, such as yellow fever , dengue fever , and
chikungunya , transmitted mostly by
Aedes aegypti .
Dengue fever is
the most common cause of fever in travelers returning from the
Caribbean, Central America, South America, and South Central Asia.
This disease is spread through the bites of infected mosquitoes and
cannot be spread person to person. Severe dengue can be fatal, but
with good treatment, fewer than 1% of patients die from dengue.
* The parasitic diseases collectively called malaria , caused by
various species of
Plasmodium , carried by female mosquitoes of the
Lymphatic filariasis (the main cause of elephantiasis ) which can
be spread by a wide variety of mosquito species
West Nile virus
West Nile virus is a concern in the United States, but there are
no reliable statistics on worldwide cases .
Eastern equine encephalitis virus
Eastern equine encephalitis virus is a concern in the eastern
Tularemia , a bacterial disease caused by Francisella tularensis,
is variously transmitted, including by biting flies.
Culiseta are vectors of tularemia, as well as arbovirus infections
such as West Nile virus.
* Zika , recently notorious, though rarely deadly. It causes fever,
joint pain, rashes and conjunctivitis. The most serious consequence
appears when the infected person is a pregnant woman, since during
pregnancy this virus can originate a birth defect called microcephaly
Potential transmission of
HIV was originally a public health concern,
but practical considerations and detailed studies of epidemiological
patterns suggest that any transmission of the
HIV virus by mosquitoes
is at worst extremely unlikely.
Various species of mosquitoes are estimated to transmit various types
of disease to more than 700 million people annually in Africa, South
America, Central America, Mexico, Russia, and much of Asia, with
millions of resultant deaths. At least two million people annually die
of these diseases, and the morbidity rates are many times higher
Methods used to prevent the spread of disease, or to protect
individuals in areas where disease is endemic, include:
Vector control aimed at mosquito control or eradication
Disease prevention, using prophylactic drugs and developing
* Prevention of mosquito bites, with insecticides , nets , and
Since most such diseases are carried by "elderly" female mosquitoes,
some scientists have suggested focusing on these to avoid the
evolution of resistance.
Indiscriminate eradication of mosquitoes is likely to have effects
undesirable to humans. Entomologist Phil Lounibos of Florida Medical
Entomological Laboratory (FMEL), Institute of Food and Agricultural
University of Florida
University of Florida , says that eradication "is
fraught with undesirable side effects", as mosquitoes are a
significant food source for birds and bats and, as larvae, fish and
frogs, parts of the food chain affecting many species. If mosquitoes
are eradicated, they may also be replaced by other species, possibly
Mosquito control World War II era pamphlet aimed
to discourage creation of stagnant water Mosquitofish Gambusia
affinis , a natural mosquito predator
Many measures have been tried for mosquito control , including the
elimination of breeding places, exclusion via window screens and
mosquito nets , biological control with parasites such as fungi and
nematodes, or predators such as fish, copepods , dragonfly nymphs
and adults, and some species of lizard and gecko . Another approach
is to introduce large numbers of sterile males . Genetic methods
including cytoplasmic incompatibility, chromosomal translocations, sex
distortion and gene replacement have been explored. They are cheaper
and not subject to vector resistance.
Insect repellents are applied on skin and give short-term protection
against mosquito bites. The chemical
DEET repels some mosquitoes and
other insects. Some CDC -recommended repellents are picaridin ,
eucalyptus oil (PMD ) and
IR3535 . Others are indalone, dimethyl
phthalate , dimethyl carbate , and ethyl hexanediol.
In 2015, researchers at
New Mexico State University
New Mexico State University tested 10
commercially available products for their effectiveness at repelling
mosquitoes. On the mosquito
Aedes aegypti, the vector of Zika virus,
only one repellent that did not contain
DEET had a strong effect for
the duration of the 240 minutes test: a lemon eucalyptus oil
repellent. All DEET-containing mosquito repellents were active.
There are also electronic insect repellent devices which produce
ultrasounds that were developed to keep away insects (and mosquitoes).
However, no scientific research based on the EPA's and many
universities' studies has ever provided evidence that these devices
prevent a human from being bitten by a mosquito. In 2005, the
British consumer magazine Holiday reported the results of its test of
a range of mosquito deterrents. The magazine's editor Lorna Cowan
described the four appliances that used a buzzer as "a shocking waste
of money" which "should be removed from sale" .
BITES AND TREATMENT
Play media Video of a mosquito biting on leg
Visible, irritating bites are due to an immune response from the
binding of IgG and IgE antibodies to antigens in the mosquito's saliva
. Some of the sensitizing antigens are common to all mosquito species,
whereas others are specific to certain species. There are both
immediate hypersensitivity reactions (types I and III) and delayed
hypersensitivity reactions (type IV) to mosquito bites. Both
reactions result in itching, redness and swelling. Immediate reactions
develop within a few minutes of the bite and last for a few hours.
Delayed reactions take around a day to develop, and last for up to a
week. Several anti-itch medications are commercially available,
including those taken orally, such as diphenhydramine , or topically
applied antihistamines and, for more severe cases, corticosteroids ,
such as hydrocortisone and triamcinolone .
IN HUMAN CULTURE
A still from
Winsor McCay 's pioneering 1912 animated film How a
Ancient Greek beast fables including "The Elephant and the Mosquito"
and "The Bull and the Mosquito", with the general moral that the large
beast does not even notice the small one, derive ultimately from
Winsor McCay 's 1912 film
How a Mosquito Operates was one of the
earliest works of animation, far ahead of its time in technical
quality. It depicts a giant mosquito tormenting a sleeping man.
The de Havilland
Mosquito was a high-speed aircraft manufactured
between 1940 and 1950, and used in many roles.
* ^ The plural can also be written "mosquitos".
* ^ Harbach, Ralph (November 2, 2008). "Family Culicidae Meigen,
Mosquito Taxonomic Inventory.
* ^ "mosquito".
Real Academia Española
Real Academia Española . Retrieved 24 July 2016.
* ^ Brown, Lesley (1993). The New Shorter Oxford English Dictionary
on Historical Principles. Oxford : Clarendon. ISBN 0-19-861271-0 .
* ^ "Mosquitoes of Michigan -Their
Biology and Control". Michigan
Mosquito Control Organization. 2013.
* ^ Gates, Bill. "The Deadliest
Animal in the World".
* ^ "Would it be wrong to eradicate mosquitoes? – BBC News". BBC
News. Retrieved 2016-02-01.
* ^ G. O. Poinar; et al. (2000). "Paleoculicis minutus (Diptera:
Culicidae) n. gen., n. sp., from
Cretaceous Canadian amber with a
summary of described fossil mosquitoes" (PDF). Acta Geologica
Hispanica . 35: 119–128.
* ^ Borkent A, Grimaldi DA (2004). "The earliest fossil mosquito
(Diptera: Culicidae), in Mid-
Cretaceous Burmese amber". Annals of the
Entomological Society of America . 97 (5): 882–888. doi
:10.1603/0013-8746(2004)0972.0.CO;2 . ISSN 0013-8746 .
* ^ "Discovery of new prehistoric mosquitoes reveal these
blood-suckers have changed little in 46 million years". Smithsonian
Science News. January 7, 2013. Retrieved October 27, 2015.
* ^ Briggs, D.E. (2013). "A mosquito\'s last supper reminds us not
to underestimate the fossil record" . Proceedings of the National
Academy of Sciences of the United States of America. 110 (46):
18353–4. doi :10.1073/pnas.1319306110 . PMC 3832008 . PMID
* ^ Greenwalt, D.E.; Goreva, Y.S.; Siljeström, S.M.; Rose, T.;
Harbach, R.E. (2013). "Hemoglobin-derived porphyrins preserved in a
Middle Eocene blood-engorged mosquito" . Proceedings of the National
Academy of Sciences of the United States of America. 110 (46):
18496–18500. doi :10.1073/pnas.1310885110 . PMC 3831950 . PMID
* ^ Reidenbach, K.R.; Cook, S.; Bertone, M.A.; Harbach, R.E.;
Wiegmann, B.M.; Besansky, N.J. (2009). "Phylogenetic analysis and
temporal diversification of mosquitoes (Diptera: Culicidae) based on
nuclear genes and morphology" . BMC Evolutionary Biology. 9 (1): 298.
doi :10.1186/1471-2148-9-298 . PMC 2805638 . PMID 20028549 .
* ^ Calvo E, Pham VM, Marinotti O, Andersen JF, Ribeiro JM (2009).
"The salivary gland transcriptome of the neotropical malaria vector
Anopheles darlingi reveals accelerated evolution of genes relevant to
BMC Genomics . 10 (1): 57. doi
:10.1186/1471-2164-10-57 . PMC 2644710 . PMID 19178717 .
* ^ Lawniczak MK, Emrich SJ, Holloway AK, Regier AP, Olson M, White
B, Redmond S, Fulton L, Appelbaum E, Godfrey J, Farmer C, Chinwalla A,
Yang SP, Minx P, Nelson J, Kyung K, Walenz BP, Garcia-Hernandez E,
Aguiar M, Viswanathan LD, Rogers YH, Strausberg RL, Saski CA, Lawson
D, Collins FH, Kafatos FC, Christophides GK, Clifton SW, Kirkness EF,
Besansky NJ (2010). "Widespread Divergence Between Incipient Anopheles
Species Revealed by Whole Genome Sequences" . Science. 330
(6003): 512–514. doi :10.1126/science.1195755 . PMC 3674514 .
PMID 20966253 .
* ^ Harbach, R.E. (2011).
Mosquito Taxonomic Inventory.
* ^ Walter Reed Biosystematics Unit. Wrbu.si.edu. Retrieved on
* ^ Jaeger, Edmund C. (1959). A Source-Book of Biological Names and
Terms. Springfield, Ill: Thomas. ISBN 0-398-06179-3 .
* ^ Society, National Geographic. "Mosquitoes,
Mosquito Facts – National Geographic". National Geographic.
* ^ Biological notes on mosquitoes. Mosquitoes.org. Retrieved on
* ^ Taking a bite out of mosquito research, Author Paul Leisnham,
University of Maryland. Enst.umd.edu (2010-07-26). Retrieved on
* ^ Molavi, Afshin (June 12, 2003). "Africa\'s
Malaria Death Toll
Still "Outrageously High"". National Geographic. Retrieved July 27,
* ^ "Mosquito-borne diseases". American
Association. Retrieved October 14, 2008.
* ^ World Health Organisation. Flooding and communicable diseases
* ^ Wilcox, B.A.; Ellis, B. (2006). "Forests and emerging
infectious diseases of humans". Unasylva. 57. ISSN 0041-6436 .
* ^ Wigglesworth V. B. (1933). "The Adaptation of
to Salt Water". J Exp Biol. 10 (1): 27–36.
* ^ Crans, Wayne J.;
Wyeomyia smithii (Coquillett). Rutgers
University, Center for Vector Biology.
* ^ Kosova, Jonida (2003) "Longevity Studies of Sindbis Virus
Aedes Albopictus". All Volumes (2001–2008). Paper 94.
* ^ Michigan
Mosquito Control Association; Michigan Mosquito
Manual, MMCA Edition. Pub. Michigan Department of Agriculture June
* ^ A B Spielman, Andrew; D'Antonio, M. (2001).
Mosquito : a
natural history of our most persistent and deadly foe. New York:
Hyperion. ISBN 978-0-7868-6781-3 .
* ^ Huang Juan; Walker Edward D; Vulule John; Miller James R
(2006). "Daily temperature profiles in and around Western Kenyan
larval habitats of
Anopheles gambiae as related to egg mortality".
Malaria Journal. 5: 87. doi :10.1186/1475-2875-5-87 .
* ^ Service, Mike (2012). Medical Entomology for Students (5th
ed.). Cambridge University Press. ISBN 978-1-107-66818-8 .
* ^ A B Harzsch S, Hafner G (2006). "Evolution of eye development
in arthropods: Phylogenetic aspects".
Arthropod Structure and
Development. 35 (4): 319–340. doi :10.1016/j.asd.2006.08.009 . PMID
* ^ Kaufmann C, Briegel H (2004). "Flight performance of the
Anopheles gambiae and
Anopheles atroparvus" (PDF).
Journal of Vector Ecology . 29 (1): 140–153. PMID 15266751 .
Archived from the original (PDF) on 2011-07-28. Retrieved June 21,
* ^ Frequency of
Mosquito Wings. Hypertextbook.com (2000-05-31).
Retrieved on 2013-04-01.
* ^ African Safari Travel Blog » Blog Archive » Facts you may not
know about mosquitoes. Safari.co.uk (2011-07-05). Retrieved on
* ^ Freudenrich, Craig. "
HowStuffWorks "How Mosquitoes Work"".
HowStuffWorks . Retrieved 7 September 2013.
* ^ Shirai Y, Funada H, Seki T, Morohashi M, Kamimura K (July
2004). "Landing preference of
Aedes albopictus (Diptera: Culicidae) on
human skin among ABO blood groups, secretors or nonsecretors, and ABH
antigens". Journal of Medical Entomology. 41 (4): 796–9. doi
:10.1603/0022-2585-41.4.796 . PMID 15311477 .
* ^ Bill Chappell (July 12, 2013). "5 Stars: A Mosquito\'s Idea Of
A Delicious Human".
* ^ G. Mandela Fernández-Grandon; et al. (22 April 2015).
"Heritability of Attractiveness to Mosquitoes".
PLOS ONE . 10:
e0122716. doi :10.1371/journal.pone.0122716 . Retrieved 24 July 2015.
* ^ Sawabe K, Moribayashi A (2000). "
Lipid utilization for ovarian
development in an autogenous mosquito,
Culex pipiens molestus
(Diptera: Culicidae)". Journal of medical entomology. 37 (5):
726–731. doi :10.1603/0022-2585-37.5.726 . PMID 11004785 .
* ^ Hallem EA, Nicole Fox A, Zwiebel LJ, Carlson JR (2004).
Mosquito receptor for human-sweat odorant". Nature . 427
(6971): 212–213. doi :10.1038/427212a . PMID 14724626 .
* ^ "Scientists identify key smell that attracts mosquitoes to
US News . October 28, 2009.
* ^ "Scientists have identified the gene that makes mosquitoes
crave human blood".
Richard Dawkins Foundation
Richard Dawkins Foundation . November 21, 2014.
* ^ Devlin, Hannah (February 4, 2010). "Sweat and blood why
mosquitoes pick and choose between humans". London: The Times.
Retrieved May 13, 2010.
* ^ Estrada-Franco, R. G.; Craig, G. B. (1995). Biology, disease
relationship and control of
Aedes albopictus. Technical Paper No. 42.
Washington, D.C.: Pan American Health Organization.
* ^ Wayne J. Crans (1989). Resting boxes as mosquito surveillance
tools. Proceedings of the Eighty-Second Annual Meeting of the New
Mosquito Control Association. pp. 53–57.
* ^ Maruniak, James E. (July 2014). "Asian tiger mosquito".
Gainesville, Florida :
University of Florida
University of Florida .
Retrieved October 2, 2014.
* ^ Huget, Jennifer (2007-07-31). "Will Nothing Stop That Infernal
Itch?". Washingtonpost.com. Retrieved 2013-10-15.
* ^ Jones, C.; Schreiber, E. (1994). "The carnivores,
Toxorhynchites". Wing Beats. 5 (4): 4.
* ^ "Site down for maintenance" (PDF). Pestscience.com. Retrieved
* ^ A B M. J. Lehane (9 June 2005). The
Biology of Blood-Sucking in
Insects. Cambridge University Press. pp. 151–. ISBN
* ^ Sloof R.; Marks E. N. (1965). "Mosquitoes (Culicidae) biting a
fish (Periophthalmidae)". Journal of Medical Entomology. 2: 16. doi
* ^ Holm, E., Scholtz, C. H. (1985). Insects of southern Africa.
London: Butterworths. ISBN 0-409-10487-6 . CS1 maint: Multiple names:
authors list (link )
* ^ Aldrich JR, Barros TM (1995). "Chemical attraction of male crab
spiders (Araneae, Thomisidae) and kleptoparasitic flies (Diptera,
Milichiidae, and Chloropidae)". J Arachnol. 23: 212–214.
* ^ Harris P, Cooke D (1969). "Survival and fecundity of mosquitoes
fed on insect haemolymph". Nature. 222 (5200): 1264–1265. doi
* ^ Howard LO, Dyar HG, Knab F (1912) The Mosquitoes of North and
Central America and the West Indies, Washington D.C. Carnegie
Institute of Washington. pp. 520
* ^ Matthews, Robert W. Foraging and Feeding. Springer Netherlands,
2010. ISBN 978-90-481-2388-9 . doi=10.1007/978-90-481-2389-6_4
* ^ George J, Blanford S, Thomas MB, Baker TC (2014). "Malaria
Mosquitoes Host-Locate and Feed upon Caterpillars" . PLoS ONE. 9 (11):
e108894. doi :10.1371/journal.pone.0108894 . PMC 4220911 . PMID
* ^ Martel V, Schlyter F, Ignell R, Hansson BS, Anderson P (2011).
Mosquito Feeding Affects Larval Behaviour and Development in a Moth"
. PLoS ONE. 6 (10): e25658. doi :10.1371/journal.pone.0025658 . PMC
3185006 . PMID 21991329 .
* ^ Wahid I, Sunahara T, Mogi M (2003). "Maxillae and mandibles of
male mosquitoes and female autogenous mosquitoes (Diptera:
Culicidae)". Journal of medical entomology. 40 (2): 150–158. doi
:10.1603/0022-2585-40.2.150 . PMID 12693842 .
* ^ A B C Mullen, Gary; Durden, Lance (2009). Medical and
Veterinary Entomology. London: Academic Press.
* ^ Richards, O. W.; Davies, R.G. (1977). Imms' General Textbook of
Entomology: Volume 1: Structure,
Physiology and Development Volume 2:
Classification and Biology. Berlin: Springer. ISBN 0-412-61390-5 .
* ^ A B Ribeiro JM, Francischetti IM (2003). "Role of arthropod
saliva in blood feeding: sialome and post-sialome perspectives".
Annual Review of Entomology
Annual Review of Entomology . 48: 73–88. doi
:10.1146/annurev.ento.48.060402.102812 . PMID 12194906 .
* ^ Grossman GL, James AA (1993). "The salivary glands of the
Aedes aegypti, express a novel member of the amylase
Biology . 1 (4): 223–232. doi
:10.1111/j.1365-2583.1993.tb00095.x . PMID 7505701 .
* ^ Rossignol PA, Lueders AM (1986). "Bacteriolytic factor in the
salivary glands of
Aedes aegypti". Comparative Biochemistry and
Physiology B . 83 (4): 819–822. doi :10.1016/0305-0491(86)90153-7 .
PMID 3519067 .
* ^ A B Valenzuela JG, Pham VM, Garfield MK, Francischetti IM,
Ribeiro JM (2002). "Toward a description of the sialome of the adult
Insect Biochemistry and Molecular
Biology . 32 (9): 1101–1122. doi :10.1016/S0965-1748(02)00047-4 .
PMID 12213246 .
* ^ Bissonnette EY, Rossignol PA, Befus AD (1993). "Extracts of
mosquito salivary gland inhibit tumour necrosis factor alpha release
from mast cells".
Parasite Immunology . 15 (1): 27–33. doi
:10.1111/j.1365-3024.1993.tb00569.x . PMID 7679483 .
* ^ A B Cross ML, Cupp EW, Enriquez FJ (1994). "Differential
modulation of murine cellular immune responses by salivary gland
Aedes aegypti". American Journal of Tropical Medicine and
Hygiene . 51 (5): 690–696. PMID 7985763 .
* ^ A B Zeidner NS, Higgs S, Happ CM, Beaty BJ, Miller BR (1999).
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".
Parasite Immunology .
21 (1): 35–44. doi :10.1046/j.1365-3024.1999.00199.x . PMID 10081770
* ^ Wanasen N, Nussenzveig RH, Champagne DE, Soong L, Higgs S
(2004). "Differential modulation of murine host immune response by
salivary gland extracts from the mosquitoes
Aedes aegypti and Culex
quinquefasciatus". Medical and Veterinary Entomology . 18 (2):
191–199. doi :10.1111/j.1365-2915.2004.00498.x . PMID 15189245 .
* ^ Wasserman HA, Singh S, Champagne DE (2004). "
Saliva of the
Yellow Fever mosquito,
Aedes aegypti, modulates murine lymphocyte
Parasite Immunology . 26 (6–7): 295–306. doi
:10.1111/j.0141-9838.2004.00712.x . PMID 15541033 .
* ^ Depinay N, Hacini F, Beghdadi W, Peronet R, Mécheri S (2006).
"Mast cell-dependent down-regulation of antigen-specific immune
responses by mosquito bites".
Journal of Immunology
Journal of Immunology . 176 (7):
4141–4146. doi :10.4049/jimmunol.176.7.4141 . PMID 16547250 .
* ^ Schneider BS, Soong L, Zeidner NS, Higgs S (2004). "Aedes
aegypti salivary gland extracts modulate anti-viral and TH1/TH2
cytokine responses to sindbis virus infection". Viral Immunology. 17
(4): 565–573. doi :10.1089/vim.2004.17.565 . PMID 15671753 .
* ^ Taylor JL, Schoenherr C, Grossberg SE (1980). "Protection
against Japanese encephalitis virus in mice and hamsters by treatment
with carboxymethylacridanone, a potent interferon inducer". The
Journal of Infectious Diseases . 142 (3): 394–399. doi
:10.1093/infdis/142.3.394 . PMID 6255036 .
* ^ Schneider BS, Soong L, Girard YA, Campbell G, Mason P, Higgs S
(2006). "Potentiation of West Nile encephalitis by mosquito feeding".
Viral Immunology. 19 (1): 74–82. doi :10.1089/vim.2006.19.74 . PMID
* ^ Schneider BS, Higgs S (2008). "The enhancement of arbovirus
transmission and disease by mosquito saliva is associated with
modulation of the host immune response" . Transactions of the Royal
Society of Tropical Medicine and Hygiene . 102 (5): 400–408. doi
:10.1016/j.trstmh.2008.01.024 . PMC 2561286 . PMID 18342898 .
* ^ Curic G, Hercog R, Vrselja Z, Wagner J (2014). "Identification
of person and quantification of human DNA recovered from mosquitoes
(Culicidae)". Forensic Sci Int Genet. 8 (1): 109–12. doi
:10.1016/j.fsigen.2013.07.011 . PMID 24315597 .
* ^ Billingsley PF, Hecker H (1991). "Blood digestion in the
Anopheles stephensi Liston (Diptera: Culicidae): activity
and distribution of trypsin, aminopeptidase, and alpha-glucosidase in
Journal of Medical Entomology . 28 (6): 865–871. PMID
* ^ A B "Vísindavefurinn: Af hverju lifa ekki moskítóflugur á
Íslandi, fyrst þær geta lifað báðum megin á Grænlandi?" (in
Icelandic). Visindavefur.hi.is. Retrieved 2013-10-15.
* ^ Fang J (July 21, 2010). "Ecology: A world without mosquitoes".
Nature. Nature. 466 (7305): 432–4. doi :10.1038/466432a . PMID
* ^ Peterson B.V. (1977). "THE BLACK FLIES OF ICELAND (DIPTERA:
SIMULIIDAE)". The Canadian Entomologist. 109: 449–472. doi
* ^ Gislason G. M.; Gardarsson A. (1988). "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". Verb. int. Ver. Limnol. 23:
* ^ Hawley WA, Pumpuni CB, Brady RH, Craig GB (1989).
"Overwintering survival of
Aedes albopictus (Diptera: Culicidae) eggs
Journal of Medical Entomology . 26 (2): 122–129. PMID
* ^ Hanson SM, Craig GB (1995). "
Aedes albopictus (Diptera:
Culicidae) eggs: field survivorship during northern Indiana winters".
Journal of Medical Entomology . 32 (5): 599–604. PMID 7473614 .
* ^ Romi R, Severini F, Toma L (2006). "Cold acclimation and
overwintering of female
Aedes albopictus in Roma". Journal of the
Mosquito Control Association. 22 (1): 149–151. doi
:10.2987/8756-971X(2006)222.0.CO;2 . PMID 16646341 .
* ^ WHO (2009). Dengue Guidelines for Diagnosis, Treatment,
Prevention and Control (PDF). Geneva: World Health Organization. ISBN
* ^ "Lymphatic Filariasis". World Health Organisation (WHO)
website. World Health Organisation (WHO). Retrieved 24 August 2011.
* ^ Muslu H, Kurt O, Özbilgin A (2011). "". Turkiye Parazitol Derg
(in Turkish). 35 (2): 100–4. doi :10.5152/tpd.2011.25 . PMID
* ^ "Can I get
HIV from mosquitoes?". CDC . October 20, 2006.
* ^ "Resistance is Useless". The Economist. April 8, 2009.
* ^ Claire Bates (28 January 2016). "Would it be wrong to eradicate
mosquitoes?". BBC News. Retrieved 28 January 2016.
* ^ "Fungus Fatal to
Mosquito May Aid Global War on Malaria". The
New York Times. 10 June 2005.
* ^ Kramer, J. P. (1982). "Entomophthora culicis(Zygomycetes,
Entomophthorales) as a pathogen of adultaedes aegypti(diptera,
culicidae)". Aquatic Insects. 4 (2): 73–79. doi
:10.1080/01650428209361085 . .
* ^ Shamseldean MM, Platzer EG (1989). "Romanomermis culicivorax:
Penetration of larval mosquitoes". Journal of
54 (2): 191–199. doi :10.1016/0022-2011(89)90028-1 . PMID 2570111 .
* ^ Krumholz, Louis A. (1948). "Reproduction in the Western
Gambusia affinis affinis (Baird & Girard), and Its Use
Mosquito Control". Ecological Monographs. 18 (1): 1–43. doi
JSTOR 1948627 .
* ^ edited by Kenneth T. MacKay. (1995). "PART III: INTERACTIONS31.
A Comparative Study of the Ability of
Fish to Catch
Rice-fish culture in China. International Development Research Centre.
ISBN 1-55250-313-5 . CS1 maint: Extra text: authors list (link )
* ^ Fradin MS (1 June 1998). "Mosquitoes and mosquito repellents: a
clinician\'s guide" (PDF).
Annals of Internal Medicine . 128 (11):
931–940. doi :10.7326/0003-4819-128-11-199806010-00013 . PMID
* ^ Marten GG, Reid JW (2007). "Cyclopoid copepods". Journal of the
Mosquito Control Association. 23 (2 Suppl): 65–92. doi
:10.2987/8756-971X(2007)232.0.CO;2 . PMID 17853599 .
* ^ Canyon DV, Hii JL (1997). "The gecko: An environmentally
friendly biological agent for mosquito control". Medical and
veterinary entomology. 11 (4): 319–323. doi
:10.1111/j.1365-2915.1997.tb00416.x . PMID 9430109 .
* ^ Jennifer Carpenter (8 August 2011). "Spermless mosquitoes hold
promise to stop malaria". BBC. Retrieved 5 August 2011. Scientists
have created spermless mosquitoes in an effort to curb the spread of
* ^ Webb, Jonathan (10 June 2014) GM lab mosquitoes may aid malaria
fight BBC News, Science and Environment, Retrieved 11 June 2014
* ^ Syed Z, Leal WS (2008). "Mosquitoes smell and avoid the insect
repellent DEET" .
Proceedings of the National Academy of Sciences
Proceedings of the National Academy of Sciences .
105 (36): 13598–13603. doi :10.1073/pnas.0805312105 . PMC 2518096
. PMID 18711137 .
* ^ CDC (2009). "Updated Information regarding
* ^ Rodriguez, Stacy D.; Drake, Lisa L.; Price, David P.; Hammond,
John I.; Hansen, Immo A. (2015). "The Efficacy of Some Commercially
Insect Repellents for
Aedes aegypti (Diptera: Culicidae) and
Aedes albopictus (Diptera: Culicidae)". Journal of
Insect Science. 15
(1): 140. doi :10.1093/jisesa/iev125 . ISSN 1536-2442 .
* ^ "
Mosquito repellents that emit high-pitched sounds don\'t
prevent bites" (Press release).
Eurekalert! . April 17, 2007.
* ^ William Kremer (11 December 2012)
repellents: Zapping the myth BBC News, Magazine, Retrieved 30 May
* ^ Clements, Alan (1992). The biology of mosquitoes – volume 1:
Development, Nutrition and Reproduction. London: Chapman & Hall. ISBN
* ^ Adrados, Francisco Rodríguez (1999). History of the
Latin Fable. BRILL. p. 324. ISBN 90-04-11454-8 .
* ^ Canemaker, John (2005). Winsor McCay: His Life and Art. Abrams
Books . p. 165. ISBN 978-0-8109-5941-5 .
* ^ "De Havilland Mosquito". The Aviation History Online Museum.
Retrieved 21 November 2015.
* Brunhes, J.; Rhaim, A.; Geoffroy, B.; Angel, G.; Hervy, J. P. Les
Moustiques de l'Afrique mediterranéenne French/English. Interactive
identification guide to mosquitoes of North Africa, with database of
information on morphology, ecology, epidemiology, and control. Mac/PC
Numerous illustrations. IRD/IPT 2000 CD-ROM. ISBN 2-7099-1446-8
* Davidson, Elizabeth W. (1981). Pathogenesis of invertebrate
microbial diseases. Montclair, N. J.: Allanheld, Osmun. ISBN
* Jahn, G. C., Hall, D. W. & Zam, S. G. (1986). "A comparison of the
life cycles of two Amblyospora (Microspora: Amblyosporidae) in the
Culex salinarius and
Culex tarsalis Coquillett". Journal of
the Florida Anti-
Mosquito Association. 57: 24–27. CS1 maint:
Multiple names: authors list (link )
* Kale, H. W., II. (1968). "The relationship of purple martins to
mosquito control" (PDF).
The Auk . 85 (4): 654–661. doi
JSTOR 4083372 . CS1 maint: Multiple names: authors
list (link )
Look up MOSQUITO in Wiktionary, the free dictionary.
Wikimedia Commons has media related to CULICIDAE .
Wikispecies has information related to