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family: culicidae
common names: mosquitoes
antennae characters: 1 pr.
specialties: medical importance
hosts: mammals, birds
natural enemies: gambusia fish, dragonflies, purple martins, notonectids,
etc.
cultural control methods: vegetable oil (or other monofilm) on
top of breeding sites
physical control methods: drain water recepticles, put excelsior
into latrine tanks
notes: Small, fragile, elongate,long legged insects
with scales or dense hairs on veins of wings, with long beak containing
piercing-sucking
mouthparts, and with weakly plumose antennae. Larvae (wrigglers)
aquatic 85 spp of Culicidae transmit malaria (55 for human parasites).
Plasmodia spp life cycle: blood sucked up by Anopheles (45 degree
body angle while feeding) carrying gametocytes (sexual stage of
plasmodia) which develop into sperm and egg cells in the mosquito's
body and undergo sexual reproduction then forming an oocyst, which
upon maturing, bursts and releases sporozites which migrate to
the salivary glands of the mosquito where they are released into
the next host on which it feeds. Once established in the vertebrate
host, the sporozoites (spore stage) migrate to liver and send out
merazooites to infect blood cells that can continue to infect blood
cells (in the typical 48 hour life cycle marked by regular periods
of chills and fever in the host) or may give rise to gametocytes
that can be sucked up by mosquitoes. Filaria worms (in the microfilariae
stage) are active in the peripheral circulatory system only between
10pm and 4am when they may be picked up by night-flying skeeters.

To protect you and your family from mosquito bites:
Wear baggy, long-sleeved shirts and pants.
- Wear light-coloured
clothing since dark colours attract mosquitoes.
- Use mosquito
repellents that contain DEET. The percentage of DEET in repellents
should not exceed 30% for adults or 10% for
children. DEET should not be used on children under 6 months of age.
- Ensure
you read the instructions before application.
- Make sure
that doors and windows have tight-fitting screens
- Make sure
your lawn is mowed to decrease the area where mosquitoes
can rest during the hot part of the day.
- Avoid forested
areas during the day
It doesn’t take
much time, or water, for mosquitoes to develop from eggs into
adults. Therefore, any thing that can hold water
is a potential development site. To destroy common mosquito breeding
sites around your home, consistent yard maintenance is the best
defense. We can all reduce risk by doing the following:
- Empty saucers
under flower pots.
- Change water
in bird baths and pet water/food dishes weekly or more frequently.
- Empty wading
pools when not in use or drain and refill each week; drain tarps
and trampolines that fill with rainwater.
- Make sure
roof gutters drain properly.
- Remove used
tires and other debris where rainwater may collect.
- Repair leaky
outdoor faucets.
- Use fine mesh
to cover rain barrels and containers that cannot be dumped.
If there is a large area of stagnant water near your house, but
not on your property, that you are concerned about, please contact
your local engineering or public works department. Arrangements
may be made to drain it or prevent mosquitoes from breeding in
it.

Info on mosquitoes and their control:
http://www.hc-sc.gc.ca/pmra-arla/english/pdf/pnotes/mosquitos-e.pdf
http://www.hc-sc.gc.ca/pmra-arla/english/mosquito/mosquito-e.html
http://www.epa.gov/pesticides/factsheets/mosquito.htm
http://wlapwww.gov.bc.ca/epd/epdpa/ipmp/Brochures/goodbugs.htm
Info on insect repellents:
http://www.epa.gov/pesticides/factsheets/insectrp.htm
http://www.hc-sc.gc.ca/pmra-arla/english/pdf/pnotes/deet-e.pdf
http://www.epa.gov/pesticides/factsheets/chemicals/deet.htm
Info on Bacillus thuringiensis subspecies israelensis (Bti), a
biological pesticide effective against mosquito larvae, is available
at:
http://www.hc-sc.gc.ca/pmra-arla/english/pdf/fact/fs_bti-e.pdf
http://www.epa.gov/pesticides/biopesticides/ingredients/factsheets/factsheet_006476.htm
(Please note that, under provincial regulations, pesticide treatment
of any bodies of water other than those that are human-made and
self-contained on private land can proceed only under permit or
similar provincial authorization.)
Info on products that can be legally used in Canada (most of
them only by a person possessing a valid Pesticide Applicators
Certificate) by typing in the word "mosquito" in the
'free text' field at:
http://www.eddenet.pmra-arla.gc.ca/4.0/4.0.asp

Agriculture and Agri-Food Canada Insect Identification Sheet No.
86 1983
Mosquitoes
About 60 species
of mosquitoes occur in Canada, and because females require a
blood meal to develop eggs, they are a serious pest of
humans, livestock and wildlife during spring and summer.
Injury:
Mosquitoes cause much annoyance and discomfort to humans and animals.
Their attacks reduce beef and milk production, and lessen the efficiency
of agricultural, forestry and other workers, They also interfere
with recreation and spread diseases.

Life Cycle:
Mosquitoes
develop only in stagnant water, and pass through four distinct
life stages — egg, larva, pupa and adult. A female may
lay over a hundred eggs these are deposited either singly on the
soil among grass roots and organic matter in low places where water
collects, or in rafts on the surface of
stagnant water. The larvae, often known as "wrigglers",
hatch and feed on tiny animals and plants in the water. They breathe
through a tube at the end of the body that lets them remain just
below the surface. When fully grown, the larvae transform to comma-shaped
pupae, often called "tumblers". Pupae do not feed but
move actively in the water. When the adult is ready to emerge,
the pupa floats to the surface, its skin splits down the back and
the adult emerges. The adult rests on the pupal skin long enough
for its body to harden, then flies away to vegetation or moist
and shady places. Both sexes feed on plant juices. A few
days after emergence, the mosquitoes mate. The females then search
for a blood meal which is necessary to produce eggs. Males live
only a few days, whereas females can live for several months.
Most common mosquitoes (Aedes spp.) pass the winter as eggs that
hatch when flooded with water. The less common mosquitoes (Culex spp., Culiseta spp. and Anopheles spp.) pass the winter as fertilized
females in cellars, animal burrows, hollow trees, basements and
unheated buildings; they appear early in the spring and the eggs
of these species hatch shortly after they are laid.
Pest Management:
Mosquito control
is difficult because of the diverse habits and breeding places
of the different species, Consult your provincial Departments
of Health, Environment and Agriculture for control procedures
recommended for your area.
last updated
on the 16th of September 1995 by G.J.L.Ramel@exeter.ac.uk
Mosquitoes (Culicidae, suborder Culicinae) 1,500+ species Worldwide,
about 42 in the UK. These are the most important group of flies
in this suborder. The word mosquito is Spanish and means 'little
fly'. Mosquito adults can be recognized because they are the only
family of flies which have both the veins of their wings covered
in scales (very rare in any flies at all) and a long projecting
proboscis. Flies in general are lovers of the light but not mosquitoes
most of whom are true denizens of the night. Not all species of
mosquitoes are blood suckers and in those species which are it
is only the females which do; this is because they need the protein
to develop their eggs. One genus of mosquitoes called Harpagomyia have learned how to rob ants of the genus Crematogaster.
The mosquito
alights in front of an ant returning to its nest beating its
wings whereupon the ant opens its jaws allowing the mosquito
to use it proboscis to remove the contents of the ants crop.
Mosquitoes have aquatic larva and one species or another lays
its eggs in most areas of still water from large lakes to puddles
and knot holes in trees. Two very interesting groups are the
genus Trypteroides which lays its eggs in the urns of the Pitcher
plants of Indo-Malaya and Australia despite these being filled
with digestive enzymes used by the plant to consume the insects
trapped within and the genus Megarhinus whose larva are carnivorous
and live in the same pitcher urns feeding on the larval Trypteroides.
The strangest place any fly lays its eggs is on its own legs,
this is done by the mosquitoes of the genus Amigeres the females
carefully places each of her eggs onto her legs where they diapause
(a kind of hibernation) until the female immerses her legs in water,
the stimulus of the water causes the eggs to fall off their mothers
legs and to come out of diapause so that they can hatch in the
water in the normal way.
Rearing mosquitoes — pets or public enemies?
Sarah A. Corbet and G. W. Danahar
Department of Zoology, Downing Street, Cambridge, CB2 3EJ
Sarah Corbet is a lecturer, and Gary Danahar is a technical assistant,
in the Department of Zoology, University of Cambridge
Pithy statement: Mosquitoes can be reared in the laboratory, with
little trouble or expense, and offer endless opportunities for
experimental studies that can contribute to non-insecticidal methods
of control.
Running title: Enjoying mosquitoes

Abstract
The mosquito Culex
pipiens form molestus is easy to rear because
the adults do not need a blood meal. This paper describes methods
for maintaining a stock culture and for producing cohorts
of uniformly-aged larvae for experiments. Mosquitoes are major
vectors of disease. There is a growing need for control methods
that do not rely on insecticides, but these methods depend on an
understanding of mosquito biology and of the natural history and
biology of predators. Some research avenues that may contribute
to this understanding are outlined.
Key words
Mosquitoes, Culex, rearing, surface film, predators, control.
Introduction
Mosquitoes are important
vectors of disease, especially in the tropics. Resistance to
conventional insecticides is a serious and
growing problem (Brookfield, 1991), and increasingly attention
is being paid to alternative methods of control. Among the weapons
available for use against the aquatic stages of mosquitoes are
oils and insoluble surfactants such as Arosurf 'Monomolecular Surface
Film', insect juvenile hormone analogues such as methoprene, and
natural enemies that can be introduced as biological control agents
(Laird & Miles, 1985). Microbial control agents include bacteria
such as Bacillus thuringiensis israelensis, which is commercially
available for application to mosquito larval habitats, and the
fungi Coelomomyces and Culicinomyces. Parasitic nematodes have
also been
suggested for biological control
of mosquito larvae. Potentially useful predators include dragonfly
larvae (Sebastian et al., 1990), the water boatman Notonecta, the
magnificent predatory mosquito Toxorhynchites, and fishes such
as the mosquito fish Gambusia affinis or the guppy Poecilia
reticulata.
Research is needed to discover how effective such methods may
be in different conditions, and to explore the possible interactions
between them. For example, if oils act by preventing mosquito larvae
or pupae from gaining access to atmospheric air at the water surface,
are these oils ineffectual in well-aerated water where the immature
mosquitoes can get oxygen from solution in the water? If mosquito
larvae in water butts in Trinidad are controlled with a film of
an insoluble surfactant such as Arosurf 'Monomolecular Surface
Film', might control be improved by adding a water boatman or a
guppy to each container, or would these predators be harmed by
the surfactant? If dragonfly larvae are introduced into domestic
water cisterns in Myanmar (Burma) to control mosquito larvae, might
control be improved by adding a guppy, or would the guppies and
the dragonfly larvae eat each other? There are endless opportunities
for potentially useful research on mosquitoes, using relatively
simple equipment.
The world's
most notorious vector species do not occur naturally in Britain,
and we do not recommend them as experimental subjects
in schools and colleges, but useful work can be done on related
species that occur in Britain or that can easily be reared in the
laboratory. Snow (1990) describes the ecology and identification
of British mosquito species and suggests approaches to field studies.
A British species easily reared in the laboratory is Culex pipiens
form molestus, and in this paper we outline a simple method
for keeping it in culture. The London School of Hygiene and Tropical
Medicine, Keppel Street, London WC1E 7HT is usually willing to
supply eggs for starting cultures.
C. pipiens form molestus has several features that make it easy
to rear. First, whereas most other mosquitoes need a large space
in which to swarm and mate, this form can mate in a confined
space. Secondly, whereas some species have an obligate period of
delayed development, or diapause, usually in the egg or larval
stage, this form does not: successive generations develop without
delay in the laboratory (although in nature some adults overwinter
in diapause in underground cavities such as cellars). Lastly, and
most importantly, whereas adult females of most other mosquitoes
need a meal of blood before they can lay the first batch of eggs,
this form does not: it is autogenous. This means
that the adult females do not need to be fed on vertebrate blood.
It does not mean that they will not bite if given the opportunity.
If they escape, they will occasionally bite. It is essential to
take care that pupae are never left to emerge in open bowls. These
should be securely covered with glass or gauze
if there is any risk of adults emerging.
The methods we describe are not infallible. Rearing mosquitoes
is as much an art as a science, and adjustments will have to be
made to meet individual circumstances.

Beer barrel technology
We keep a source culture ticking over with little attention through
the year in three large containers. We use translucent 22.5-litre
(40-pint) barrels designed for making home-made beer (Boots; 22.49
each in February 1993, complete with tap) (fig. 1) in a 27oC room
(12 hr light: 12 hr dark). Each barrel is filled to about one-quarter
to one-third of its depth with water. Cheaper and less robust alternatives
are available, such as the Boots 25-litre winemaker's fermenting
barrel, or a translucent polythene fermentation bin (15-litre bins
cost 3.55) with a fitted lid from which an area can be cut away
and replaced by a glued-on sleeve of netting. In either case, a
tap (Boots barrel tap, 7.25) needs to be fitted in a hole drilled
in the wall of the bin about 2.5 cm above the base. It can be sealed
in with aquarium sealant.
The beer barrel mouth is wide enough to admit a hand and arm for
cleaning. There are two holes in the screw cap. The smaller hole
(diameter about 0.5 cm) is exposed by unscrewing the pressure relief
valve and the larger one (diameter about 1.5 cm) by unplugging
the rubber stopper in the centre of the lid.
Through the smaller hole we run a 0.5-cm-diameter air pipe, ending
inside the barrel in an air stone (available from shops that sell
aquarium accessories) which allows air from the air pump (Rena
Aquarium Air Pump 301, @ 24.49 in October 1992) to bubble air into
the water. If an aquarium air line plastic tap is introduced between
the pump and the air stone, the flow of air can be regulated. It
is usually best to adjust the bubbler so that there is only a gentle
trickle of air. If the bubbles are fierce, they make it difficult
for the larvae and pupae to break the water surface for gaseous
exchange, they disturb adults that are emerging or standing on
the surface to lay eggs there, and they capsize the egg rafts,
breaking the eggs' contact with the water and preventing many of
them from hatching.
The larger hole in the cap is the right size to support a free-standing
plastic funnel with a 15-cm-diameter mouth. A wad of cotton wool
in the funnel prevents the escape of adult mosquitoes, and can
be removed when water, food (in suspension or in solid form) or
livestock is to be introduced into the barrel through the funnel.
We also drilled a series of 30-50 holes, 0.2 cm in diameter, in
the screw cap. These holes are small enough to prevent adult mosquitoes
from escaping, but large enough to allow some ventilation of the
barrel contents. They are restricted to the lid because this can
be replaced cheaply if the barrel needs to be used for another
purpose later.
The Boots
beer barrel has a tap at the bottom through which the aquatic
component of the culture may be removed, without allowing
the adults to escape. Larvae and pupae are easily drained out through
the tap, but the egg rafts float on the water surface, and to
withdraw these it is necessary to tilt the barrel, bringing
the water surface level with the tap and allowing the surface,
with its egg rafts, to dribble out. While the liquid from the barrel
is run out into a washing-up bowl, the fate of the egg rafts can
be monitored by looking through the peep hole ordinarily occupied
by the funnel.

Food
C. pipiens
molestus larvae feed when hanging below the surface
by filtering out small particles suspended in the water, or swim
down to browse on the bottom of the container. There are several
possible food sources. A mixture of desiccated liver powder (catalogue
No. L. 26 from Unipath Ltd, Wade Road, Basingstoke, Hampshire,
RG24 0PW at 13.01 for 250 gm), dried yeast (Yesta 20B at 44.25
per 25Kg from BFI-The tastemakers, Beesham Food Ingredients, P.O.
Box 18, Wellington Road, Burton-on Trent, Staffordshire, DE14 2AB;
or in small quantities from health food shops) and sugar, in the
ratio of 3:10:2 by weight, at one level teaspoonful per barrel
every 4-5 days, has sustained our colony for eight years. But the
liver powder is expensive, and the soupy bacterial community that
this food supports, and on which the mosquito larvae feed, is unpredictable
and somearial foul-smelling. If the temperature is too high or
there is too much food, undesirable anaerobic micro-organisms develop.
If the soup becomes really foul, a stronger blast of air bubbles
for an hour or so may help to drive out any noxious gases.
Because of the unpredictability
of the liver/yeast/sucrose diet and the cost of the liver, we
have explored alternatives. One of
our cultures has survived for six months solely on yeast and sugar,
in the ratio of 5:1 by weight (one level teaspoonful per barrel
at 4-5 day intervals), and is still flourishing. Some people feed
mosquito cultures on Farex baby food. Perhaps the most convenient
food is that recommended by the London School of Hygiene and Tropical
Medicine: Guinea Pig Food (RGP pellets code 678, manufactured by
Grain Harvesters Ltd., The Old Colliery, Wingham, Nr. Canterbury,
Kent CT3 1LS, who will also supply on request a breakdown of the
composition of the guinea pig food). This is available in small
quantities from pet shops, or in much larger quantities from Wm.
Lillico & Son, Wonham Mill, Betchworth, Surrey RH3 7AD, at
13.18 per 25-kg pack plus a surcharge for small orders and a delivery
fee. We drop ten pellets into each barrel every 4-5 days.
The need for frequent feeding can cause problems over a holiday
period. It has been suggested that these might be overcome by using
a commercially-available device designed to deliver a daily food
ration to aquarium fish.
Kept at a mean temperature of 27oC (range 26-28oC) and a mean
humidity of about 50% (range c. 35-65%), mosquitoes in the barrels
take up to three weeks to complete their life cycle. Development
is much slower at room temperature.

Mass-rearing larvae for experiments
Cultures can be maintained in barrels for long periods with little
effort, so that a few mosquitoes of any stage are available whenever
they are needed; but when large numbers of larvae are required,
all of the same age and in uniformly good condition, we rear separate
cohorts in washing-up bowls.
Ten egg rafts
taken from a barrel culture are placed in a plastic washing up
bowl (40 cm long by 30 cm wide and 20 cm deep) one-third
full of water. We use white bowls so that the young larvae can
be seen easily. The larvae have usually hatched by the next day
and can be seen as tiny, light grey threads. At this stage they
can acquire oxygen through the cuticle and no air supply is required.
The next day a little food is added and a gently-bubbling air stone
placed in the bowl. The rate of air supply can be increased gradually
as the larvae grow, but the bubbles should never be vigorous
enough to agitate the larvae.
If the liver/yeast/sucrose diet is used, a pinch is added to each
bowl every day, and each cohort will develop from egg to pupa in
about five days. Underfeeding results in uneven, delayed larval
development and small adults that do not lay eggs; overfeeding
produces a foul-smelling brew.
On the guinea pig food diet the larvae seem to develop a little
more slowly but the amount of food is less critical. One advantage
of the guinea pig food diet over the liver/yeast/sucrose diet is
that there is no need for an air line into the bowls, although
it may be advantageous to supply one if available. In the absence
of an air line a surface scum somearial develops on the water but
this can be removed by laying a paper towel over the water surface,
allowing it to come into contact with the scum and then pulling
the towel away sideways, avoiding contact with the side of the
bowl.
If the cohort is well fed, any larvae that have not been used
in experiments will all begin to pupate within a day or so of one
another. If further egg rafts are needed to keep a continuous supply
of cohorts in production, any pupae remaining unused in the cohort
bowls are transferred into an 'oviposition pot', a one-pint disposable
plastic beer glass. Each beer glass should contain about 150-200
individuals. With such a high density of adults in a confined space,
mating is frequent and large numbers of egg rafts will be laid
in the pot. Overcrowding should be avoided because too much contamination
of the water surface with cast pupal cuticles and other debris
may discourage egg-laying.
For transfer
to the beer glass, pupae are removed from the cohort bowl by
pouring its contents through a tea strainer, and they are
put in the beer glass together with about 20 ml of their rearing water.
By pouring fresh water through the tea strainer to dislodge the
larvae and pupae, the glass is filled about half full. A single
sheet of white toilet tissue is hung over the edge of the glass
with one end in the water, to give the adult mosquitoes a surface
on which to rest and to mate. A saucer (preferably a glass one)
resting on top of the beer glass prevents the adults from escaping
(fig. 2).
If the age structure of the cohorts is uneven (and this happens
somearial with all the diets used), it is necessary to select individuals
uniform in age as parents for the next generation, because if small
larvae are kept with the developing pupae they will die, decompose,
and foul the contents of the beer glass, discouraging egg-laying.
When the age structure is not uniform, we use a plastic Pasteur
pipette (with the tip cut off to give it a wide mouth), or a 50-ml
syringe with a short length of polythene tubing attached to the
end, to pick out pupae individually from the cohort bowl, leaving
the larvae behind. The pupae are put into a plastic beer glass
as before, together with pupae from several other cohorts, to produce
a batch in which good numbers of adults emerge simultaneously.
Adults emerge after 2-3 days. Males (which have feathery antennae)
emerge first, and females usually emerge about a day later. The
period between adult emergence and egg-laying is about 3-4 days.
Some people allow the adults to feed on 10% glucose solution or
soaked sultanas. This is not essential but it may help to encourage
egg-laying. If the sugar solution is left for more than a day or
two it will ferment.
If the water
in the oviposition pot becomes foul, it may be necessary to change
it after most of the adults have emerged but before the
eggs are laid. The trick of doing this without releasing the adults
is to tilt the beer glass (with the tissue side uppermost), keeping
the liquid in by holding the saucer firmly in place. The tilted
glass is then partially immersed in a bowl of water, and the
dirty water, plus any remaining larvae,
is let out by sliding the saucer and slowly raising the beer glass.
So long as any gaps are below the water surface, no adult mosquitoes
can escape. The glass can be refilled by submerging it in a bowl
of clean water and again displacing the saucer a little. With practice
considerable control can be exercised and no adults escape.
The egg rafts
are white when first laid, but within an hour or two they darken
and eventually they become charcoal black. The
egg rafts can be removed daily by the method described above for
changing the water in the beer glasses; the glass is rocked gently
from
side to side to prevent the egg rafts from adhering to the
sides of the glass, and care is taken to keep the rafts floating
upright, because shipwrecked rafts do not hatch properly. After
transfer from the beer glass to a bowl, the rafts can be scooped
off the surface of the water in a white plastic spoon and used
to set up new cohorts. A single beer glass usually
produces about 50 rafts in all; our record is total of 160 egg
rafts in one glass over 3 days.

Mosquitoes and research
With large numbers of uniformly-aged last-instar larvae or pupae,
it is possible to conduct experiments on the effectiveness of control
agents less toxic than conventional insecticides.
Since the early use of oil films on water to control mosquito
larvae, insoluble surface-active agents have been developed further
and are among the more environmentally acceptable methods of control
for the aquatic stages of mosquitoes. The mode of action is uncertain,
but there are three ways in which these agents might kill mosquito
larvae: by interfering with the surface tension effects on which
gas exchange at the water surface depends; by flooding the tracheal
system; or by direct toxicity.
We have tried
to find cheap, readily available materials that form a surface
film on water and that could be used to treat water
butts and other small water bodies in and near houses in regions
where technically elaborate control operations are not practicable.
Plant-derived oils, such as eucalyptus oil, have proved promising.
Using disposable 40-ml pots (clear plastic 'one-ounce' tubs with
lids, sold by Concept Catering, 1 Duncock Lane, Elsworth, Cambridge
CB3 8JL, @ 67.50 per thousand), each containing 20 ml tapwater
giving a surface area of about 16 cm2, we set up an experiment
with
ten last-instar mosquito larvae per
pot and five pots per dose, testing a range of doses (1,2,4,8 16
microlitres) of oil run carefully onto the surface of the water
after the
larvae had been put in. The pots, with their lids on,
were kept in an incubator at 30oC for 24 h, at the end of which
larvae in each pot were scored as 'dead' (motionless, even when
disturbed by gently jogging the pot) or 'alive' (capable of moving).
From the count of dead larvae in each pot we could calculate the
24-h LD50, the dose that kills 50% of larvae in 24h, by plotting
the mean number dead at each dose on a probability scale (corresponding
to a cumulative normal distribution scale) against dose of oil
on a logarithmic scale. This is easily done using special graph
paper known as probit paper, with a logarithmic scale on the x
axis and a probability scale on the y axis. The mathematical basis
is explained by Garvin (1986) and Sokal and Rohlf (1981). The plot
produces a straight line from which the dose corresponding to 50%
mortality can be read off. This graphical method generally gives
an adequate estimate of the LD50, but if a more accurate estimate
is required the LD50 can be calculated from a probit regression
using a computer programme
such as Generalised Linear Interactive Modelling (GLIM) (Atkin
et al., 1989) or by a time-consuming manual calculation (Busvine,
1971).
On the basis
of the LD50, expressed as microlitres per square centimetre,
we can compare a range of readily-available oils (somearial
spiced with surface-active compounds to increase their ability
to spread on the water surface) with surface films marketed commercially
for mosquito control, such as Arosurf MSF. Experiments might be
designed along similar lines to see how susceptibility to these
control agents varies through the life cycle of the mosquito. Our
aim has been to find a control agent that is inexpensive, readily
available in rural areas, and of low toxicity to humans and other
non-target organisms.
Some oils
can be seen to enter the gas exchange system of the mosquito
larvae, flooding the tracheae and giving them a translucent
appearance under a stereoscopic microscope, instead of the opaque
silvery look they have when filled with air. Mosquito larvae
in oil-treated pots frequently nibble at the snorkel-like
siphon at the tip of the abdomen, as if attempting to remove a
contaminant. If these oils kill larvae by interfering with gas
exchange between the atmosphere and the tracheal system, we might
expect that they would kill mosquito larvae faster in stagnant
waters rich in decomposing organic material, where dissolved oxygen
is in short supply, than in well-oxygenated waters. This possibility
might be explored by comparing the effectiveness of a chosen oil
on water deoxygenated by boiling with its effectiveness on well-oxygenated
water. The difference may be more obvious with young larvae, which
derive more of their oxygen from solution in the water, than with
large larvae, or pupae, that depend more on atmospheric oxygen.

Mosquito larvae
and pupae divide their time between resting, tail up, at the
surface, and diving down below the surface. They alter the allocation
of time between these two activities in response to certain compounds
applied to the surface of the water, changing the mean frequency
or the mean duration of dives below the surface. Detailed behavioural
effects of this kind are difficult to monitor directly in a crowd
of active mosquito larvae, but can easily be quantified if a
group of, say, five or ten mosquito larvae is filmed on video,
preferably with a time signal shown on the tape. The film can
be made and analyzed using
ordinary home video equipment. We use a clear plastic rectangular
box in which we have improvised five vertical compartments so that
individual mosquitoes are kept apart but all five can be filmed
at the same time through the flat wall of the box. One might, for
instance, film the larvae for five minutes, then add a surface
oil, then continue filming for a further five minutes. By scoring
the number of larvae at the surface, the number of dives and the
mean duration of a dive for each successive minute, it is possible
to see how surface treatments affect the probability of diving
(number of dives, per mosquito at the surface, per minute) and
the mean duration of a dive (or of the period at the surface between
dives). Sih (1986) has suggested that mosquito larvae alter their
dive timing in the presence of
certain predators, perhaps reducing their vulnerability by spending
less time in the region where predators are most likely to find
them; and that this predator effect is mediated by a water-borne
chemical. Video filming could be used to explore effects of this
kind.
If insoluble surface-active agents disrupt the surface forces
that enable mosquito larvae to hang at the water surface, these
agents may be expected to be equally damaging to other insects
that depend on surface forces. Guthrie (1989) describes the natural
history of some surface-dwelling insects and outlines methods for
exploring the effects on them of contaminants that alter surface
forces. Among these surface-dwelling insects are predators such
as the water boatman or backswimmer, Notonecta, that feed readily
on mosquitoes, and are regarded as important natural control agents
in, for example, rice fields in the southern United States. If
these predators are harmed by surface-active mosquito control agents,
the natural control exercised by their predation may be sacrificed.
Studies on the predatory behaviour of such insects, in clean water
and in the presence of surface-active contaminants, can be valuable.
The bacterium Bacillus
thuringiensis israelensis is used as a
mosquito control agent, somearial in combination with a surface-active
agent that helps to hold the bacteria at the surface where the
mosquito larvae will contact them. These bacteria are effective
when eaten by mosquito larvae. It is therefore important to study
feeding behaviour. Information available so far is reviewed
by Merritt, Dadd and Walker (1992) and Clements
(1992), but much remains to be discovered about the factors that
determine the rate of feeding and the way a larva allocates
its time between feeding at the surface (where it may
pick up bacterial spores), filtering in the water column and browsing
on the bottom. Some compounds have already been shown to increase
feeding rate; perhaps others may be found to decrease it, causing
starvation.
One way to
investigate the rate of feeding is to feed larvae on a suspension
of coloured particles (such as Indian ink, or carmine
particles) for known periods of time, and monitor the movement
of this coloured material along the gut by examining larvae under
a stereoscopic microscope at intervals. The position of the visible
boundary between the original food and the coloured particles can
be recorded in terms of the number of segments along the body.
It would be interesting to see whether adding, for instance, dried
powdered yeast to the water will stimulate feeding, making this
boundary move faster down the body.
If the larvae are non-selective filter feeders, ingesting particles
of a given size regardless of composition, it might be possible
to control them by adding to the water a suspension of inert, non-nutritious
particles of the appropriate size, preventing the larvae from obtaining
a nutritious diet. Even if inert particles are not normally eaten,
they may be taken if flavoured with a compound that acts as a feeding
stimulant. Marmite-flavoured kaolin or talcum powder would be a
novel control agent!
Safe, effective mosquito control is a matter of life and death
to many people in the tropics. Particularly needed are control
methods that can be operated by local communities on a small scale
without the intervention of professional agents or expensive spraying
machinery (see Curtis, 1991). The development of cheap, simple,
practicable community control methods of this kind depends on careful
exploration of mosquito biology, coupled with imaginative lateral
thinking.
A disadvantage of some of the investigations suggested here is
that they involve killing mosquitoes, and some people may feel
reluctant to do this, especially after studying their exquisite
complexity under the microscope. On the other hand, in countries
where mosquito-borne diseases cause serious human suffering and
death, mosquito control is considered essential; conventional control
may be ineffective because so many races of mosquitoes have developed
resistance to insecticides, which may cause high mortality among
non-target organisms. Serious, careful research that provides the
foundation for more specific and biologically based control methods
can benefit both people and the environment.
The study of mosquitoes is not only worthwhile, but also fascinating,
and the books by Clements (1993) and Snow (1991) have recently
made the field accessible by providing a firm base and making it
possible to set individual studies into the context of existing
knowledge.

Acknowledgements
We are very grateful to Barbara Sawyer (London School of Hygiene
and Tropical Medicine) for her valued help and advice, Stuart Green
for guidance, Philip Corbet and Michael Reiss for commenting on
the text, and many students whose ideas and enthusiasm have sustained
the mosquito project over the years.
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