Daftar isi

Chapter 2

Biology of Mosquitoes
Regarding their special adaptational mechanisms,
mosquitoes are capable of thriving in a variety of environments.
There is hardly any aquatic habitat anywhere
in the world that does not lend itself as a breeding site
for mosquitoes. They colonise the temporary and permanent,
highly polluted as well as clean, large and
small water bodies; even the smallest accumulations
such as water-filled buckets, flower vases, tyres, hoof
prints and leaf axes are potential sources.
In temporarily flooded areas, along rivers or lakes
with water fluctuations, floodwater mosquitoes such as
Aedes vexans or Ochlerotatus sticticus develop in large
numbers and with a flight range of several miles,
become a tremendous nuisance even in places located
far away from their breeding sites (Mohrig 1969;
Becker and Ludwig 1981; Schfer etal. 1997).
In swampy woodlands, snow-melt mosquitoes such
as Oc. cantans, Oc. communis, Oc. punctor, Oc. hexodontus,
and Oc. cataphylla, encounter ideal conditions
for development in pools that are formed after the
snow melts or after heavy rainfall.
In floodplains along coastal areas, the halophilous
species (preferring brackish or salt water habitats) such
as Oc. taeniorhynchus, Oc. sollicitans, Oc. vigilax, Oc.

caspius, Oc. detritus, develop in huge numbers. Larvae

of the Anopheles can be found in association with other
mosquito species in fresh- or salt-water marshes, mangrove
swamps, rice fields, grassy ditches, edges of
streams as well as in small temporary water collections.
Many species prefer habitats either with or without
vegetation.
Tree-holes are the habitat of arboreal species such
as Oc. geniculatus, Ae. cretinus, Anopheles plumbeus
and Orthopodomyia pulcripalpis.
Species like Cx. p. pipiens, Ae. aegypti [St. aegypti]
Ae. albopictus [St. albopicta] or Oc. j. japonicus can
even breed in a variety of small water containers such
as rain-water drums, tyres, cemetery vases, small clay
pots, etc.
Furthermore, their capacity to adapt to various
climatic factors or changing environmental conditions
is fascinating. For instance, Ae. albopictus, the Asian
tiger mosquito is originally a tropical species. In the
course of a climate-related evolutionary adaption it
developed a photoperiodic sensitivity. When days are
shorter, the photoperiodically sensitive female inhabiting
a temperate climate, lays eggs that are different
from the eggs that she lays when days are longer. Eggs
laid during shorter days, are dormant and do not hatch
until the following season, ensuring the species survival
through the winter.
This ability to adapt to moderate climatic conditions
and the fact that the eggs are resistant to desiccation
and survive for more than a year, including the
capability of adaptation to artificial breeding sites
such as tyres and flower pots, make Ae. albopictus
or Oc. j. japonicus successful species. This has contributed
to its spread globally via international trade
in plants like Dracaena spp. (lucky bamboo) and
tyres. Within hours or days they can be transported
from one country or continent to another by cars, aircrafts
or trans-oceanic containers (Madon et al.
2002).
These are but a few examples, which illustrate the
tremendous ecological flexibility of mosquitoes successfully
adapted to.
Like all Diptera, mosquitoes exhibit complete metamorphosis.
All mosquitoes need aquatic habitats for
their development, although Aedes/Ochlerotatus spp.

can lay their eggs in moist soil. After hatching they pass
through four larval instars and a pupal stage when the
transformation into an adult takes place. Most species
are unautogenous: following copulation, the females have
to take a blood-meal to complete the egg development.
Only a few species have populations that are autogenous.
They first develop egg batches without a bloodmeal
(e.g. Cx. p. pipiens biotype molestus).

2.1 Oviposition
Female mosquitoes lay between 50 and 500 eggs, 24
days (or longer in cool temperate climates) after the
blood-meal. In general, the mosquitoes can be divided
into two groups depending on their egg-laying behaviour
(Barr 1958) and whether or not the embryos
enter into a period of dormancy (externally triggered
resting period) or diapause (genetically determined
resting period).
In the first group, females deposit their eggs onto the
water surface either singly (Anopheles) or in batches
(e.g. Culex, Uranotaenia, Coquillettidia, Orthopodomyia
and subgenus Culiseta, Fig.2.1).
The Culex females lay their eggs in rafts comprising
several hundred eggs locked together in a boat-shaped
structure. During oviposition, the females stand on the
water surface with the hind-legs in a V-shaped position.
The eggs are released through the genital opening
and grouped together between the hind-legs, forming a
raft where the eggs stand vertically on their anterior
poles attached together by chorionic protrusions
(Clements 1992). The anterior pole of each egg has a
cup-shaped corolla with a hydrophilic inner surface,
which lies on the water surface, the outer surface is
hydrophobic. The resulting surface tension helps to
keep the egg-raft in position and when rafts drift to
their aquatic boundaries they tend to remain there.
Immediately following oviposition, the eggs are soft
and white, but they sclerotize and darken within
12h.
Anophelines lay single eggs while standing on the
water surface or hovering above it. The eggs of this
subfamily are adapted for floating and can easily be
damaged by desiccation.
The embryos of the first group do not enter dormancy
or diapause and hatch when the embryonic development
is completed. Species producing nondormant eggs usually
have several generations each year. Their developing
stages are found for the most part in more permanent
waters where one generation succeeds another during
the breeding season. The number of generations depends
on the length of the breeding season, as well as the
abiotic and biotic conditions, and most importantly, it
is the temperature which influences the speed of
development.
The parameters that determine the choice of a
breeding site by the females laying their eggs onto the
water surface are still unknown for many species.
Factors such as water quality, incidence of light, existing
eggs, available food, and local vegetation are decisive

factors in selecting a favourable breeding site.

For Cx. p. pipiens, it is known that the content of
organic material in the water plays an important role in
attracting the females about to lay eggs. Apparently,
gaseous substances such as ammonia, methane, or carbon
dioxide, which are released when organic material
decomposes, create an effect of attracting the females
of Cx. p. pipiens (Becker 1989b). They recognise that
such a site has adequate food and that favourable conditions
prevail for the development of their brood.
A few other examples illustrate that the egg-laying
behaviour reflects the ecological conditions in the breeding
site. The submerged larvae and pupae of Coquillettidia
obtain the oxygen they need by inserting their siphon
into the aerenchyma (air filled tissue) of certain plants
under water. Therefore, in order to ensure the development
of the larvae, the females must recognise the appropriate
aquatic plants at the time they lay their eggs in
order to ensure the development of the larvae.
The second group lays eggs which do not hatch
immediately after oviposition (Fig.2.2). The egg-laying
behaviour of the floodwater mosquitoes (e.g. Ae. vexans), and the subgenus Culicella of
genus Culiseta, which lay their eggs singly, not on the water surface but into the
moist soil, which is subsequently flooded when water
levels rise, is most interesting. The eggs are laid into
small depressions or between particles of moss with a
high degree of soil moisture which protect the sensitive
eggs from drying-out during the embryogenesis (Barr
and Azawi 1958; Horsfall et al. 1973). For Ae. vexans
and Oc. caspius, which breed in flooded areas where the
levels fluctuate frequently, the appropriate egg-laying
behaviour is crucial to ensure successful development of
the immature stages. A suitable egg-laying site for floodwater
species must meet the following prerequisites:
(a) The substrate must be wet enough at the time the
eggs are laid in order to ensure that the freshly laid
eggs which are very sensitive to any water loss, do not
desiccate before their impermeable endochorion has
been tanned and the wax layer of the serosal cuticle is
formed (Horsfall etal. 1973; Clements 1992);
(b) There must be a subsequent and sufficient flooding
of the soil where the eggs have been laid, so
that the complete process can take place from
hatching all the way to the imago emergence;
(c) The water body for subsequent breeding should
have as few mosquito predators as possible, to
ensure that the larvae are not preyed upon by natural
enemies when they hatch.
The ability of a floodwater mosquito female to find
appropriate places for egg-laying which guarantees
maximum breeding success is not yet fully understood.
However, respect is due to these tiny insects which have
adapted their behaviour to overcome the hostile conditions
in their breeding sites. If the females chose to lay
their eggs in low-lying areas with almost permanent
water-flow, they would encounter crucial disadvantages:
low-lying areas are flooded for long periods of
time and have, therefore, a very unfavourable alternating
sequence of dry and flooding spells, therefore, allowing

only very small populations of floodwater mosquitoes

to develop. Areas with almost permanent water-flow
generally have a high concentration of natural enemies,
such as fish, so that the risk of being preyed upon would
be very high for the mosquito larvae.
However, flooded areas with a very short period of
water-flow are also unfavourable egg-laying sites,
because they entail the risk of premature drying-out.
This kind of terrain becomes flooded for a short period
of time and only in the years when water is abundant,
thus the wet and dry sequences are not favourable for
the development of several consecutive generations.
These areas dry very rapidly following a flood so that
the Aedes or Ochlerotatus eggs run the risks of desiccation
since the developing embryos are very sensitive
to water loss. Figure 2.3, shows the preferred egg-laying sites of
the floodwater mosquitoes (e.g. Ae. vexans). Usually,
these areas consist of dense vegetation and silty soil,
with the reed (e.g. Phragmites australis) being highly
attractive to ovipositing female mosquitoes.
Usually, the reed zone precisely demarcates the
mid-water level in riparian systems, since reeds need a
good amount of water.
The fluctuating curve of the Rhine river, for example,
makes it quite clear how important this egg-laying
behaviour is for the development of the floodwater
mosquitoes (Fig.2.4). In a zone of between 4 and 5m,
there is the ideal timing of dry and wet periods, with
optimum water-flow to guarantee the development of a
large number of individuals in a population. Further
more, predators, especially fish are usually low in
numbers or absent.
But how do these female floodwater species find
the optimum oviposition site? Gravid females obviously
recognise the wet, silty, riparian clay soil as an
appropriate egg-laying substrate and are attracted to it.
However, these criteria alone are not enough, given
that during rainy periods there would be many other
places which would apparently appear adequate due to
their high moisture content.
It is likely that the floodwater mosquitoes are able
to differentiate between various soil types. The soil in
most floodplains consists of a high percentage of clay
and low percentage of humus or organic materials, in
contrast to many other soils (Ikeshoji and Mulla 1970;
Strickman 1980a,b; Becker 1989b).
It is also possible that flooded areas along the riverbanks
produce pheromone-like odours, which may be
recognised by the female mosquitoes and which induce
them to lay their eggs. These odours could come from
eggs, which have already been laid in the soil, or from
particular associations of plants which are indicators of
a specific moisture level in the soil, and the occurrence
of regular floods. As a whole, the ovipositional behaviour
of the floodwater mosquitoes shows an astonishing

Mosquitos
Mosquitos are important vectors of several tropical diseases, including malaria,
filariases, and numerous viral diseases, such as dengue, Japanese encephalitis and
yellow fever. In countries with a temperate climate they are more important as
nuisance pests than as vectors.

There are about 3000 species of mosquito, of which about 100 are vectors of human
diseases. Control measures are generally directed against only one or a few of the
most important species and can be aimed at the adults or the larvae.

Life cycle

Mosquitos have four distinct stages in their life cycle: egg, larva, pupa and adult (Fig.
1.2). The females usually mate only once but produce eggs at intervals throughout
their life. In order to be able to do so most female mosquitos require a blood-meal
(Fig. 1.3). Males do not suck blood but feed on plant juices. The digestion of a bloodmeal and the simultaneous development of eggs takes 2 - 3 days in the tropics but
longer in temperate zones. The gravid females search for suitable places to deposit
their eggs, after which another blood-meal is taken and another batch of eggs is laid.