Aedes aegypti and Aedes albopictus: Life cycle,

biology and distribution.

(all diagrams, figures and photographs produced by D. Nimmo)

Section i - Introduction
Section ii - The life cycle (overview)
Section iii - The eggs
Section iv - The larvae
Section v - The pupae
Section vi - The adults
Section vii - Distribution

Appendix 1 References
Appendix 2 Bibliography

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D. Nimmo
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i The mosquitoes, or Culicidae, are a family of about three and a half thousand species
Introduction within the order Diptera, the two-winged flies. They are one of the more primitive
families of Diptera, being more closely related to midges, gnats and crane flies, for
example, than to houseflies and blowflies. Mosquitoes are found throughout the
world except in places that are permanently frozen. Three quarters of all mosquito
species live in the humid tropics and subtropics, where the warm moist climate is
favourable for rapid development and adult survival, and the diversity of habitats
permitted the evolution of many species. Mosquitoes are classified into three
subfamilies, the largest and most diversified of which is divided into a number of
tribes:

Family Culicidae:
Subfamily Toxorhynchitinae
Anophelinae
Culicinae:
Tribes Sabethini
Culicini;
Genus Coquillettidia
Culex
Culiseta
Deinocerites
Haemagogus
Mansonia
Orthopodomyia
Psorophora
Uranotaenia
Aedes;
Species aegypti
albopictus
stimulans
sierrensis etc...

Like other true flies, culicids exhibit 'complete metamorphosis', i.e. the juvenile form
passes through both larval and pupal stages. The larvae are anatomically different
from the adults, live in a different habitat and feed on a different type of food.
Transformation to the adult takes place during the non-feeding pupal stage.

Aedes aegypti (also known as Stegomyia aegypti (1)), commonly known as the yellow
fever mosquito (although more appropriately, because it transmits far more of this
virus, it should be known as the dengue mosquito), is a mosquito that can transmit
dengue fever, Chikungunya, yellow fever, West Nile virus and several other viruses.

Aedes albopictus, commonly known as the Asian tiger mosquito or forest day
mosquito, is characterized by its black and white striped legs and small, black and

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 white body. This species is able to survive in a wide range of habitats and conditions.
The Asian tiger mosquito has a rapid bite that allows it to escape most attempts by
people to swat it. This mosquito has become a significant pest in many communities
because it closely associates with humans (rather than living in wetlands), and
typically flies and feeds in the daytime rather than at night or at dusk and dawn. It is
a container and puddle breeder, needing only a few ounces of water to breed. Aedes
albopictus can transmit Eastern equine encephalitis virus and dengue fever.

ii
The life The mosquito goes through four separate and distinct stages of its life cycle: Egg
(1), Larva (2), Pupa (3), and Adult (4). In the case of Aedes eggs are laid one at a
cycle
time on a moist surface. Most eggs hatch into larvae within 48 hours if
(overview) submerged in water. But in the case of Aedes aegypti and Aedes albopictus the
eggs are resistant to desiccation and can survive out of water for up to a year.
Both Aegypti and
albopictus can
survive in very
small pools of water
Figure 1; like the rim of a
Aedes life plant container or a
cycle coke can. The larvae
diagram. grow and develop
through four instars
before developing
into pupae. Adult
mosquitoes then
emerge from the
pupae after about 48
hrs. This whole
process from egg to
adult can take as
little as 6-9 days,
dependent on
temperature. The
adult females, after they have mated, then search for a suitable bloodmeal (males
do not bloodfeed), aegypti has become specialised on feeding on humans,
albopictus does feed on humans but not as exclusively as aegypti. The females
develop eggs over the following 2-3 days and then lay them on a suitable moist
surface, completing the life cycle.

iii Female mosquitoes lay some 50 to 500 eggs at one time, depositing them on water or
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The eggs on sites that will be flooded. Aedes
aegypti and albopictus typically lay
Figure 2; between 50 and 120 eggs. Each egg is
Aedes aegypti protected by an egg shell.
eggs. Spermatozoa stored by the
inseminated female fertilize the
oocytes as they are ovulated, and
embryonic development starts almost
immediately after the eggs have been
laid. Within one to two days to a
week or more, depending on
temperature, the embryo develops
into a fully formed larva. In most species the larva hatches once it is formed, and can
survive for a few days at most in the absence of water. Mosquitoes of the tribe
Aedini (Aedes aegypti and albopictus) have water-proofed egg shells capable of
resisting desiccation, and fully-formed but unhatched aedine larvae can survive for
months or even years in the absence of free water. Aedine species lay their eggs in
places that may not be flooded for days, weeks or months. A fall of rain that
inundates oviposition sites, or a high tide flooding a salt marsh, stimulates hatching
and can lead to an apparent population explosion.

iv Mosquito larvae are legless, but they

The larvae retain a well-formed head and so do not
appear maggot-like. The larval habitats
are small or shallow bodies of water with
little or no water movement typically
shallow pools, sheltered stream edges,
Figure 3; marshes and water-filled tree holes, leaf
Aedes aegypti axils or man-made containers. The
larvae. habitats range in size from animal
footprints to marshes and rice fields. Most
species live in fresh water but a few are
adapted for a life in brackish or saline
water in salt marshes, rock pools or inland
saline pools. All aquatic animals have
problems of water and salt balance,
whether they live in fresh or salt water,
and the means by which mosquito larvae
have solved these problems has been the
subject of much investigation. In addition
to the usual internal organs of ion
regulation, mosquito larvae have four
external balloon-like anal papillae, which
are capable of ion uptake from very dilute solution. When it hatches from the egg the
young mosquito larva is fully adapted for living in water, and two features determine
its manner of life: use of atmospheric oxygen for respiration and use of water borne
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 particles as food.
The air breathing habit requires mosquito larvae either to live more or less
permanently at the air/water interface, as most anopheline and some culicine larvae
do, or to make frequent visits to the water surface. The only functional respiratory
openings are a pair of spiracles, near the end of the abdomen, from which air filled
tracheae extend to all parts of the body.
The spiracles of culicine and toxorhynchitine larvae are situated at the end of a tube
or siphon, and the larvae hang downwards from the surface membrane by their
siphons with their spiracles open to the air. The spiracles of anopheline larvae are
flush with the dorsal surface of the last abdominal segment, and the larvae lie
horizontally below the surface membrane, their spiracles opening through it. Larvae
of two culicine genera (Mansonia and Coquillettidia) are able to remain permanently
submerged. They live with their respiratory siphons, which have modified saw, like
tips, forced into the air-filled tissues that fill the stems and roots of certain aquatic
plants.
The characteristic food resource of mosquito larvae is 'particulate matter'. This
includes aquatic micro organisms, such as bacteria, diatoms and algae, and also, as
an important component, particles of detritus that are largely derived from decayed
plant tissues. Such particles provide food for diverse aquatic invertebrates, which
filter them from the water by a variety of mechanisms. Mosquito larvae, which live
mainly in still water, are exceptional in not relying on natural water currents to bring
the particles to them. Through the regular beating of their 'mouth brushes' mosquito
larvae generate water currents which flow towards the head, and in a manner that is
not well understood they separate particles of a certain size from the water.
Anopheline larvae typically feed at the water surface, in a particle-rich layer just
below the surface membrane. Culicine larvae feed on particles suspended in the
water column, and many supplement this feeding mode by abrading with their
mouthparts the layers of organic matter that cover submerged surfaces, so generating
new particles. Toxorhynchitine larvae are predatory on small invertebrates, as are a
very few species in the other two subfamilies.
The growing mosquito larva moults four times. On the first three occasions that it
leaves its cast cuticle the larva appears very much as before. During the period of the
fourth moult the imaginal disks develop rapidly, changing the form of the insect
crudely to that of an adult, and the organism that leaves the fourth larval skin is a
pupa. The rapid growth rates (5-10 days from hatched larvae to adult) of many
tropical species permit the exploitation of transient water bodies.

v The pupa remains an aquatic organism. The pupal stage is a resting, non-feeding
The pupae stage of development, but pupae are mobile, responding to light changes and moving
(tumble) with a flip of their tails towards the bottom or protective areas. That it has
assumed the form of an adult is largely concealed because the head and thorax, with
their elongate appendages are cemented together in the form of a cephalothorax. This
process is similar to the metamorphosis seen in butterflies when the butterfly
develops - while in the cocoon stage - from a caterpillar into an adult butterfly. The
abdomen, which now terminates in two large paddles, has retained the strong larval
Figure 4; musculature and is an effective organ of propulsion. An air bubble, which is enclosed
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Aedes aegypti between the appendages,
pupa provides buoyancy, and the
(female). pupa floats at the water
surface with the top of its
thorax in contact with the
surface membrane and its
abdomen hanging down.
The new form and posture
preclude use of the terminal
abdominal spiracles for
respiration. That function is
taken over by the
mesothoracic spiracles,
which open within large
'respiratory trumpets'. As the
pupa floats at the air/water
interface the hydrophobic
rims of the respiratory
trumpets protrude through
the surface membrane.
During the pupal stage certain larval organs are destroyed, e.g. the alimentary canal,
while replacement adult organs are constructed from undifferentiated embryonic
cells. Other organs, including the heart and fat body, are carried over to the adult
stage. These final stages of metamorphosis can be completed within one to two days
if the temperature is sufficiently high. When the adult is fully formed within the
pupal cuticle, the insect rests at the water surface and starts to swallow air. The
consequent increase in internal pressure forces a split along the midline of the pupal
thoracic cuticle, and the adult slowly expands out of the pupal cuticle and steps on to
the water surface.

vi Like many of the more primitive Diptera, adult mosquitoes have an elongate body
The adults and long wings and legs, which provide an aerodynamically stable form. The hind
wings are modified as small oscillating sense organs, or halteres, which assist flight
control. Like other Diptera, mosquitoes are fluid feeders. Exceptionally among the
more primitive Diptera, their mouthparts have evolved into an elongate composite
proboscis, half as long as the body, suitable for probing nectaries and, in the case of
the female, adapted for piercing skin and imbibing blood from peripheral blood
vessels. The outer sheath-like part of the proboscis, the labium, encloses the
remaining mouthparts, which have the form of needle-like stylets. Of these, the
female's mandibles and maxillae, which are flattened and toothed, can be driven

Figure 5;
Aedes aegypti
adult.
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 through tissue by the
muscles at their bases,
making a channel for other
styletized mouthparts which
contain canals for the
delivery of saliva and the
removal of blood. Both
males and females use the
sugar in plant juices as a
source of energy, usually
obtaining it from nectaries
but sometimes from other
sources such as rotting fruit
and honeydew. Anopheline and culicine females have a requirement for protein,
from which to develop large batches of eggs, and they engorge on vertebrate blood
for that purpose. T. oxorhynchitine females feed only on plant juices.
Body odour and carbon dioxide, carried on the wind stimulate sense receptors on the
antennae and palps of female mosquitoes, alerting them to the presence of a host. The
females respond by flying upwind, which takes them towards the host. Close to the
host, visual stimuli and the convection currents of warm moist air that rise from the
host provide additional cues. The females of some species are able to detect
individual vertebrate hosts at a distance. Anopheles males responded to one calf at a
distance of 14m, and to two calves at over 36m. The females of all blood-feeding
species show a degree of specificity in their choice of host, whether mammal, bird or
cold-blooded vertebrate. Some species are highly specific, feeding predominantly on
one or a few host species only, others are less specific. Individual human beings
differ in their attractiveness to mosquitoes; the cause of the difference has not been
elucidated. Once landed on an appropriate part of the host, the female of Anopheles
female drives the styletized feeding components of her mouth, parts into its skin. The
saliva that is injected as the mouth, parts penetrate contains a substance that prevents
haemostasis, the aggregation of blood platelets that is the host's first defence against
the laceration of small blood vessels. The saliva is also the source of immunogens
that are responsible for the characteristic skin reactions to mosquito bites. Sooner or
later the probing stylets pierce a blood vessel, and the presence of blood is identified
from its content of ADP and ATP. If the female is undisturbed, feeding continues
until abdominal stretch receptors signal repletion. Within a few minutes gorging
mosquitoes can imbibe up to four times their own weight of blood. This provides the
protein needed for egg production, but also inflicts upon the mosquito a water load,
which renders flight difficult, and potentially toxic amounts of sodium and
potassium. The adult excretory system is capable of rapid elimination of water and
salts, and diuresis commences while the female is still feeding.
Digestion of blood proteins yields amino acids which are reconstituted in the
mosquito's fat body as proteinaceous yolk. This is transported to the ovaries and
incorporated into the oocytes, which are matured in a number that matches the
provision of yolk. It is a feature of mosquito biology that eggs are not matured
continuously but in batches, following the periodic blood meals. Male mosquitoes
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 can be readily distinguished by their large and elaborate antennae, in which rings of
fibrils encircle the shaft. These antennae resonate in response to a pure tone of a
certain pitch. Female mosquitoes in flight produce a familiar whining sound, the
pitch of which reflects the wing beat frequency of the species. That sound activates
the antennae of conspecific males, and provides directional indicators which the
massive sense organs at the base of the antennae can resolve.
The role of adult male mosquitoes is insemination of females, and when not resting
the males are either feeding or exhibiting a behaviour pattern that is likely to bring
them into contact with females. One conspicuous manifestation of male behaviour is
swarming the localized assembly of from two or three individuals to many thousands
of individuals of a single species. Any conspecific female that enters a swarm will be
seized immediately by a male. Mating also occurs outside swarms. Inseminated
females store sufficient sperm in their spermathecae to fertilize a number of egg
batches. A factor called matrone that is transferred in semen renders females
unreceptive to males and refractory to further copulation, but its effectiveness may
not persist throughout the life of the female.
The behavioural activities of adult mosquitoes-emergence, mating, feeding, and
oviposition take place at particular times of day and night, which vary between
species. Adults of Anopheles gambiae, for example, emerge during the late afternoon
and, once mature, mate during a twenty-minute period at dusk. The females take
blood meals principally during the four hours after midnight. The timing is not
governed directly by light and dark but by endogenous or so-called circadian
rhythms, which are reset daily by the change from light to dark at sunset.
When a female has matured a batch of eggs she takes to the wing and responds to
stimuli from suitable oviposition sites. For most mosquitoes the oviposition site is a
water body with particular characteristics; odour, taste, flow and shade are known to
influence different species. The eggs may be dropped individually to float on the
water surface, as by females of Anopheles, or packed together to form a floating egg
raft, as by Culex. Aedine species deposit their eggs on moist surfaces, often at the
edge of a body of water or on an area of soil that will be flooded. By whatever
means, the female finds the appropriate habitat, and the larvae hatch into conditions
for which they are adapted.
In tropical regions the life span of adult mosquitoes ranges from a few days to
several weeks. In temperate regions it is frequently longer, and in species that
overwinter as adults the life span of females may approach one year. The females'
behavioural responses and physiological processes follow a pattern, the gonotrophic
cycle, which starts with response to the vertebrate host and feeding, continues with
the digestion of blood and formation of a batch of mature oocytes, and ends with
oviposition. Within an hour of completing one gonotrophic cycle a female may
commence another. At warmer temperatures, tropical Anopheles and Aedes oviposit
regularly every two or three days.

vii Aedes aegypti and Aedes albopictus have a global distribution (figures 6 and 7
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Distribution respectively). Aedes albopictus is a very recent invader of the New World; it was
first detected in America in 19852, being transported over from Asia in old used
tyres.

Figure 6;
Worldwide
distribution
of Aedes
aegypti
(CDC).

Figure 7;
Worldwide
distribution
of Aedes
albopictus
(adapted
from Parola
et al(3))

Appendix 1 References
1 Reinert, J. F., R. E. Harbach & I. J. Kitching (2004). Phylogeny and classification
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 of Aedini (Diptera: Culicidae), based on morphological characters of all life stages.
Zool. J. Linn. Soc. 142: 289–368.

2 NC Department of Environment and Natural Resources, Division of Environmental

Health, Public Health Pest Management Section (No date) "Asian Tiger
Mosquitoes... Some Facts." Information Pamphlet.

3 Philippe Parola,*†1 Xavier de Lamballerie,‡§1 Jacques Jourdan,¶ Clarisse

Rovery,* Véronique Vaillant,# Philippe Minodier,* Philippe Brouqui,*† Antoine
Flahault,** Didier Raoult,†‡ and Rémi N. Charrel‡§ Novel Chikungunya Virus
Variant in Travelers Returning from Indian Ocean Islands. Emerging Infectious
Diseases • www.cdc.gov/eid • Vol. 12, No. 10, October 2006

Appendix 2 Bibliography
Clements A.N. the Biology of Mosquitoes; development, nutrition and reproduction.
2000, Volume 1. CABI Publishing.

http://www.cdc.gov/

http://www.who.int/

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