Rev. Sci. Tech. Off. Int. Epiz.

, 2015, 34 (1), 83-96

Insects as vectors: systematics and biology

F. Rodhain
Académie Nationale de Médecine, Académie Vétérinaire de France, 132, boulevard du Montparnasse, 75014
Paris, France
E-mail: f.rodhain@noos.fr

Summary
Among the many complex relationships between insects and microorganisms
such as viruses, bacteria and parasites, some have resulted in the establishment
of biological systems within which the insects act as a biological vector for
infectious agents. It is therefore advisable to understand the identity and biology
of these vectors in depth, in order to define procedures for epidemiological
surveillance and anti-vector control. The following are successively reviewed in
this article: Anoplura (lice), Siphonaptera (fleas), Heteroptera (bugs: Cimicidae,
Triatoma, Belostomatidae), Psychodidae (sandflies), Simuliidae (black flies),
Ceratopogonidae (biting midges), Culicidae (mosquitoes), Tabanidae (horseflies)
and Muscidae (tsetse flies, stable flies and pupipara). The authors provide a rapid
overview of the morphology, systematics, development cycle and bio-ecology of
each of these groups of vectors. Finally, their medical and veterinary importance
is briefly reviewed.

Keywords
Epidemiology – Insect – Vector – Vector ecology – Vector-borne disease – Zoonosis.

Introduction
Over a period of tens of millions of years, the arthropods,
on the one hand, and viruses, bacteria and parasites, on
the other, have had ample time to come together and
develop a variety of relationships. As each evolved, some
of these relationships led to the establishment of ‘vector
systems’ involving vertebrates, arthropods (insects, mites)
and microorganisms. Within these complex biological
systems, arthropods act as a ‘vector’ for the transmission,
from one vertebrate to another, of microorganisms, which
may or may not be pathogenic for the vertebrate concerned.
In some cases, this vector can also serve as a reservoir or
disseminator for the microorganism.
A distinction is made between ‘mechanical vectors’, which
are only carriers of infectious agents, and ‘biological vectors’,
within which the agent lives for part of its life cycle. Only
biological vector insects are considered in this article.
The different groups of vector
insects: systematics and biology
The great majority of arthropods acting as biological vectors
for infectious agents belong to the classes of insects and
mites. Only the former are considered here. To tackle the
epidemiology of disorders transmitted by insects, a basic
understanding of their bio-ecology is needed. More details
can be found in the specialised literature (1, 2, 3, 4, 5, 6).
The species involved in the transmission of infectious agents
belong to four orders of insects: Phthiraptera, Siphonaptera,
Heteroptera and Diptera.
Phthiraptera: Anoplura (lice)
The order of Phthiraptera contains two suborders: Anoplura
(lice) and Mallophaga. Both are permanent ectoparasites,
but only Anoplura are haematophagous and only this
suborder is considered here.
Anoplura: morphology and systematics
The general appearance of Anoplura is well known in the
form of human lice: a wingless insect, grey or brown in
colour, flat-bodied and 0.5  to 8 mm in length. Their biting
mouthparts take the form of a short retractable proboscis;
their legs are equipped with strong claws enabling them to
cling to bristles or hair. The legs are stubby and the inner
edges of the tibia have strong teeth which, together with
the powerful tarsal claws, form very effective pincers. The
abdomen is in nine segments, only seven of which are
separate (Fig. 1).
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The suborder of Anoplura contains more than 500 species,
usually divided into three families (or more, according
to some authors): Pediculidae, which includes human
parasites; Haematopinidae (parasites of various mammals)
and Haematomyzidae (parasites of elephants and
rhinoceros).
Anoplura: development cycle and bio-ecology
Anoplura are heterometabolous and haematophagous
insects at all stages and in both sexes.
The female begins to lay eggs 24 h after fertilisation and will
lay a hundred or so eggs during her lifetime. At a pace of
around ten a day, these eggs, or nits, are enveloped in a sticky
secretion that attaches them to hair. The incubation period
of these eggs is usually eight to ten days. The pre-imaginal
stages (larvae or nymphs) have the same general appearance
as adults, but are smaller. There are three successive stages,
with a total duration of eight to twelve days. The adult
lifespan is approximately one month.
Anoplura are permanent obligatory ectoparasites of
mammals. They generally have strict host specificity. These
insects must feed on blood at short intervals, otherwise they
can barely survive for more than 48 h. They rarely leave
their host animal, but they can be transmitted from one host
to another by direct contact (7).
Fig. 1
Louse (Pediculus humanus)
Photo: F. Rodhain
Rev. Sci. Tech. Off. Int. Epiz., 34 (1)
Human lice, Pediculus humanus (body lice), proliferate
under favourable conditions: a cool climate, poor socio-
economic conditions and overcrowding, poor individual
and collective hygiene, etc., as found in poor war-torn
countries, in prisons and in refugee camps.
Anoplura: medical and veterinary importance
The permanent presence of Anoplura on the surface of their
hosts’ bodies leads to disorders known as pediculosis and
phthirosis. In animals, these disorders are characterised
by scaling, itching and lesions due to scratching; they are
cosmopolitan infections and are especially common in
livestock in winter.
Human body lice, P. humanus, are vectors for the bacteria
responsible for epidemic typhus (Rickettsia prowazekii) and
for trench fever and its various clinical variants (Bartonella
quintana), which are transmitted in lice faeces. It is also
a vector for Borrelia recurrentis, the causative agent of
cosmopolitan relapsing fever, which is transmitted through
the haemolymph.
Siphonaptera (fleas)
Siphonaptera, or fleas, are a very uniform order in terms of
their morphology and bio-ecology.
Siphonaptera: morphology and systematics
Adult fleas are small insects (0.5–8 mm), brown in colour,
with thin, flat bodies. They are always wingless and
characterised by biting mouthparts; simple eyes; antennae
in three segments; three pairs of legs, the third being highly
developed for jumping; and an abdomen of ten segments,
at least the last three of which are transformed into the
reproductive organs. In males the morphology of this unit is
used in systematics, as are the spermathecae in females, the
number (one or two) and form of which vary. In addition,
there are numerous bristles on the integument and, in some
species, one or more rows of spines (combs or ctenidia) in
the cephalus and/or thorax (Fig. 2).
The order of Siphonaptera currently includes around
2,500 species and subspecies, which are difficult to identify
until they reach the adult stage. Most authors group these
different species into 15 or so families (8).
Siphonaptera: development cycle and bio-ecology
Fleas exist on all continents and are holometabolous insects
(complete metamorphosis).
Depending on the species, the eggs are laid inside dwellings
in animal bedding and, more rarely, in the coat/fur of the
host, without being attached. The eggs hatch four to six
Rev. Sci. Tech. Off. Int. Epiz., 34 (1)
Fig. 2
Flea (Xenopsylla cheopis)
Photo: F. Rodhain
days later. A female can lay between 200 and 500 eggs or
more (several thousand for Tunga) during its lifetime.
Flea larvae are vermiform, legless and eyeless, but armed
with a cephalus with chewing mouthparts. Their thoracic–
abdominal integument is whitish in colour. The last ten
abdominal segments have finger-like appendages. The
larvae go through three successive stages of development in
the host habitat. The last larval stage measures 5 to 10 mm.
Flea larvae usually feed on detritus such as squamae,
hair or feathers, and the faeces of adult fleas containing
haemoglobin, degraded to varying degrees.
The nymph, immobilised in a cocoon of silk and various
agglutinated debris, already has most of the morphological
characteristics of the adult. At the end of the nymph stage,
the adult is enclosed in its cocoon and will only emerge
in response to a stimulus, generally the perception of
vibrations caused by an approaching animal or person.
In the absence of such a stimulus, the insect may remain
inactive in its cocoon for months. It is common for the
nymphal diapause to last for several months. When
circumstances are favourable and no diapause occurs, the
entire pre-imaginal development will take between three
weeks and one month.
In adult fleas, both sexes are haematophagous and bite
homeotherms, mainly mammals (rodents, carnivores,
lagomorphs, bats), and sometimes birds. The blood-feeding
rate varies depending on the species, from four times a day
for those living on the host to once every four days for those
confined to a den or nest. They can fast for several months
if the temperature and humidity fall or in the absence of a
suitable host. The average lifespan of adults is around ten
85
months. In temperate climates there is probably only one
generation a year.
In biological terms, these insects are characterised by close
contact with their hosts. Although certain species live in the
nest or den of the host and are only parasitic when sucking
their blood, others are permanent ectoparasites, living
constantly in their fur or plumage. Yet others live attached,
sometimes permanently, to the integument of the vertebrate
host. The host specificity of fleas is nevertheless rather
weak; in certain cases it can be phyletic (an association
with a particular zoological group, such as bats) or, more
frequently it would seem, of ecological origin, i.e. governed
more by a given microclimate than a specific host (various
potential hosts frequenting this environment are then
susceptible to infestation).
Siphonaptera: medical and veterinary importance
Flea bites are usually pruritic and often cause an erythematous
and oedematous reaction. Repeated bites from these insects
result in pulicosis and sometimes dermatitis, due to the
hypersensitivity of the host. Female chigoe fleas (Tunga)
remain attached, gradually embedding themselves in the
integument of their hosts (animal paws, lower members of
humans); this results in inflammation of the tissues around
a small wound prone to secondary infection.
Siphonaptera can also act as intermediate hosts for different
cestodes by allowing the development of their cysticercoid
larvae (Dipylidium, Hymenolepis).
Fleas act as vectors for several types of microorganisms (9)
and use different transmission mechanisms, depending
on the individual case, i.e. transmission via saliva, faeces,
regurgitation and ingestion during grooming. (Grooming
is individual or mutual behaviour in mammals and birds,
consisting of smoothing the coat or plumage and scratching
or biting the skin to remove squamae and dislodge
ectoparasites, or removing them by killing or eating them.)
Some viruses are commonly transmitted by these insects,
such as myxomatosis virus in lagomorphs (mechanical
transmission). Fleas also act as natural vectors for rodent
trypanosomes. Finally, fleas play an occasional role in
the dissemination of the bacillus of tularemia (Francisella
tularensis). However, their medical importance is largely
due to the transmission of certain rickettsioses (such as
murine typhus due to Rickettsia typhi), bartonelloses (such
as cat scratch fever due to Bartonella henselae) and, above
all, the plague (infection by Yersinia pestis). Several other
microorganisms have been isolated in fleas, although their
role as a vector remains unproven.
Heteroptera (bugs)
The great majority of insects of the Heteroptera order
are sap-sucking (meaning that they can transmit plant
86 Rev. Sci. Tech. Off. Int. Epiz., 34 (1)

viruses). Others are predators. Three families include

haematophagous species that bite humans and animals:
Cimicidae, Polyctenidae (permanent ectoparasites of bats
with no known importance in human medicine) and
Reduviidae. Certain aquatic bugs, mainly belonging to the
Belostomatidae family, are suspected of playing a role in the
circulation of mycobacteria.

Cimicidae
Cimicidae are flat-bodied, brown insects with no functional
wings, around 5 mm long in the adult stage. They usually
have five pre-imaginal stages during which the morphology
is largely a smaller version of that of the adult. These insects,
which have biting mouthparts, are haematophagous in both
sexes and at all stages. They live in the habitat of the host.
They are active at night and take refuge during the daytime,
sheltering in fissures in cave walls and, for species associated Fig. 3
with humans, in cracks in walls, roofs, woodwork, etc. Triatomine
(Rhodnius prolixus mutant with red eyes)
Almost 100 species have been identified in the Cimicidae Photo: J.-F. Pays
family; although some are associated with birds, most are
associated with bats, and only two with other mammals. wings are entirely membranous. The legs are long as these
insects are mainly runners and are poor flyers. The abdomen
Without being truly specific to humans, two domestic is flat in fasting insects but becomes distended and globular
species of cimicids (‘bed bugs’) are habitual human when gorged.
parasites: Cimex lectularius and C. hemipterus, which are
well known for their annoying bites. A third, Leptocimex The larvae (or nymphs) have the same general morphology
boueti, associated with bats in Africa, has been known to as the adults but they are smaller and the wings only start to
bite humans. These bugs do not transmit any pathogenic take shape during the fourth stage.
agents and they are of little medical importance (sometimes
causing allergic reactions to repeated bites). Similarly, The Reduviidae family contains more than 6,000 species
Cimicidae are parasites of poultry that can cause severe worldwide. A few of them, belonging to the Triatominae
dermatitis, anaemia and weight loss in their hosts, but sub-family (around 120 species), are haematophagous; the
most do not transmit pathogens. Nevertheless, a few majority are exclusively neotropical. It is generally easy to
Cimicidae are known to transmit various infectious agents
identify the species, at least in its adult form.
to animals (bat trypanosomes and possibly several Oeciacus-
transmitted arboviruses of minor importance in birds in the
United States). Triatominae: development cycle and bio-ecology
The eggs are 1.5 to 2.5 mm in length with an operculum;
Reduviidae they are laid in cracks in walls, floors and roofs, either
singly or in batches of between ten and twenty. Incubation
This article reviews only the members of the sub-family
lasts around ten days if the temperature and humidity are
Triatominae (triatomines) within this large family, which are
conducive. Females lay between 100 and 600 eggs during
haematophagous (while other reduviids are generally insect
their lifetime.
predators) (Fig. 3).

There are five successive stages, separated by moults. Each of

Triatominae: morphology and systematics
Adult triatomines are large bugs (measuring between 5 mm
and 45 mm, depending on the species), brown in colour,
with red or yellow marks on the thorax and abdomen.
Their biting mouthparts form an elongated proboscis that
is folded onto the underside of the body when at rest. There
are two pairs of wings on the thorax: the forward pair has
a coriaceous base and a membranous apex; the posterior
these stages includes at least one blood meal, but the larvae
can survive without feeding for several months. Under ideal
conditions, the development cycle of triatomines lasts from
three to six months.
The many sylvatic species of triatomines in the neotropical
region live in birds’ nests, palm trees, under the bark and
in the hollows of trees, in burrows, etc. They are associated
Rev. Sci. Tech. Off. Int. Epiz., 34 (1)
with a wide variety of wild animals (birds, rodents,
opossums, armadillos, etc.) and do not normally come into
contact with humans. They are sedentary insects, but can
fly in search of new homes if necessary (10).
A few species have secondarily adapted to a peri-domestic
or domestic environment. These species live in close
association with humans and/or domestic animals in
dwellings and stables, even though some of them have
retained sylvatic populations to varying degrees.
Triatomines are haematophagous in both sexes and at
all stages and probably live one or two years, or possibly
more. They are generally nocturnal and unfussy feeders,
with domestic species feeding on the blood (once every
four to nine days) of sleeping humans or animals, retiring
during the day to the shelter of cracks in the floors and
walls (adobe dwellings and stables), or in roofs (palm
leaves, thatch, etc.). They can be found in large numbers
in certain dwellings (sometimes several thousand). Feeding
(2–3 ml for an adult) is painless, lasting around 20 min,
and is generally accompanied by the release of liquid faeces
onto the host.
Triatominae: medical and veterinary importance
Where there are large numbers of triatomines in human
dwellings, the blood loss resulting from their feeding can be
sufficient to cause chronic anaemia.
In addition, in tropical America, this family of bugs acts as a
vector for Trypanosoma cruzii, the agent of the dreaded Chagas
disease, or human American trypanosomosis (transmission
via insect faeces). Three genera are particularly important
in the transmission of this parasite: Panstrongylus, Triatoma
and Rhodnius. The sylvatic species can nevertheless ensure
the circulation of trypanosomes among many vertebrate
reservoirs, thereby playing a significant epidemiological
role. Certain species of Rhodnius are also involved in the
transmission of Trypanosoma rangeli, a seemingly non-
pathogenic parasite of wild and domestic mammals.
Belostomatidae
Belostomatidae are large aquatic bugs (up to 5 cm), found
in tropical regions of Africa and Asia. They are predators
and can inflict very painful bites on humans (and no
doubt animals). They are suspected of playing a role
in the circulation – and possibly the transmission – of
mycobacteria; in particular Mycobacterium ulcerans, the
agent of Buruli ulcer disease.
Diptera (flies)
Numerically, Diptera are a very important order of insects.
The order contains around 120,000 described species. The
87
haematophagous members of these species that could act
as vectors for infectious agents belong to seven groups
(sandflies, black flies, mosquitoes, midges, horseflies, tsetse
flies, pupipara), belonging to different families.
Psychodidae
The large Psychodidae family includes insects with fairly
wide-ranging lifestyles. Within this family, only the sub-
family Phlebotominae (sandflies) is of interest to doctors
and veterinarians because the females are haematophagous,
and so this is the sub-family reviewed in this article.
Phlebotominae: morphology and systematics
Adult sandflies are very small insects (1–4 mm), pale,
hairy and humpbacked. They are also characterised by
their biting mouthparts, forming quite a short proboscis,
lanceolate wings with no transversal veins but covered in
bristles, and highly developed external genitalia in males
(Fig. 4).
The vermiform larvae look like small caterpillars, around
8 mm in length (fourth stage), with chewing mouthparts.
The integument of the thorax and the abdomen is whitish,
with short, stumpy bristles on each segment; they also have
long bristles, usually four in number, on the last segment.
The nymphs, also whitish in colour, have a cephalothorax
and an abdomen, the last two segments of which are
generally inserted in the larval skin, which keeps the
nymphs attached to the substrate.
There are around 700 species of sandflies (11). Identifying
them is often tricky because there are many sibling species
that cannot be distinguished by their morphology, but they
can nevertheless have very different vector characteristics.
Several classifications have been proposed. Most species
of medical or veterinary interest belong to the genera
Phlebotomus (vectors of Eurasian and African leishmanioses)
and Lutzomyia (vectors of American leishmanioses).
Phlebotominae: development cycle and bio-ecology
The eggs, laid singly by the female in sheltered places, hatch
after one or two weeks (although some Phlebotominae
can hibernate in the form of diapause eggs). The larvae
and nymphs are soil-dwelling, confined to locations
characterised by certain permanent environmental factors:
dark, quiet, rather humid places (tree hollows, crevices
in floors and walls, rodent burrows, etc.). The four larval
stages of Phlebotominae are saprophagous, feeding on plant
debris. The entire pre-imaginal development, from hatching
through to the emergence of the adult, lasts 20 to 75 days,
depending on local climatic conditions.
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Fig. 4
Sandfly
Photo: Nil Rahola, ©IRD
The adults also frequent quiet, dark and humid
microhabitats, often near the habitats of vertebrate hosts,
which therefore vary according to the feeding preferences of
the females, which alone are haematophagous. Most species
live in the burrows of rodents or reptiles, in tree hollows,
termite mounds, dead leaves in forests, caves, etc. Some
are domestic or peri-domestic, living in or near human
dwellings (cellars, crevices in old walls, etc.). Phlebotominae
are generally active at dusk and during the night and do
not travel far as they are poor flyers. Many remain near the
ground; however, in forest habitats, some species can live
at height, as far up as the foliage of trees. Some even make
daily journeys between the canopy and ground level, which
could be of major importance in epidemiological terms.
The interval between blood meals is around three to ten
days. In temperate regions, most Phlebotominae produce
only one generation per year, which results in a single period
for the transmission of infectious agents. Nevertheless, some
are bivoltine (two generations a year). Many species prefer
to bite mammals (rodents, carnivores), while others prefer
birds or even reptiles; relatively few species are attracted
by humans. Feeding preferences often appear to be linked
to environment, insofar as the environment determines
contact with whatever vertebrate hosts are found in the
biotope. This results in a distribution of natural foci for
diseases transmitted by such insects. Certain species only
enter dwellings to gorge on humans. In addition to females
feeding on blood, sandflies of both sexes feed on sugary
juices and sometimes on aphid honeydew. Studying the
biology of sandflies in the field is always tricky, meaning that
much remains unknown in terms of their bio-ecology and
hence the ways in which these Diptera transmit pathogenic
agents (12).
Rev. Sci. Tech. Off. Int. Epiz., 34 (1)
The majority of sandfly species live in intertropical zones,
but some are found in temperate regions, especially in the
Mediterranean Basin and central and eastern Asia. They
have been observed at altitudes of up to 2,800 m. There are
no sandflies in the Pacific Islands.
Phlebotominae: medical and veterinary importance
Under certain circumstances, abundant sandflies can be a
major nuisance and, because of their painful and pruritic
bites, they can sometimes cause severe urticarial reactions
(called ‘harara’).
Above all, sandflies are responsible for the transmission
of leishmanioses, various arboviroses and one form of
bartonellosis. In total, around 70 species are involved in
human and animal medicine.
Sandflies transmit, by regurgitation, various leishmanioses
of carnivores (especially canines), rodents and humans,
mainly in subtropical and temperate, often dry, zones in
the Old World and in the wet intertropical regions of the
Americas.
The saliva-borne viruses transmitted by sandflies belong
largely to three genera in three different families: Phlebovirus
(family: Bunyaviridae), Vesiculovirus (family: Rhabdoviridae)
and Orbivirus (family: Reoviridae). These viruses infect
mainly rodents but a few, especially Phlebovirus, can infect
humans and are then responsible for ‘sandfly fever’.
Finally, one species of Bartonella is transmitted by sandflies:
Bartonella bacilliformis, the agent of Carrion’s disease or
‘Peruvian warts’ (also called verruga peruana), which occurs
at altitude in certain Andean countries.
Simuliidae (black flies)
This family is very homogeneous and practically
cosmopolitan. Some species of Simuliidae, which are
small Diptera, are of considerable medical and veterinary
importance in a number of African and Latin American
countries (13).
Simuliidae: morphology and systematics
Adult black flies are small stubby insects (1–6 mm), usually
dark in colour. The eyes are very large, joined in males
and separate in females. Although short, the antennae are
generally formed of 11 cylindrical segments, and the biting
mouthparts form a short proboscis. The highly developed
thorax holds a pair of rather short but broad wings,
supported by a small number of veins, and short, strong
legs. The abdomen, comprising ten segments, is also short;
the last two segments constitute the genital organs. The
female has a single spermatheca (Fig. 5).
Rev. Sci. Tech. Off. Int. Epiz., 34 (1)
The aquatic larvae are elongated, with a swollen abdomen,
measuring 5 to 12 mm in the last stage. They have a cephalic
capsule with two antennae, chewing mouthparts and two
premandibles with long bristles, arranged in a fan shape when
deployed. The thorax has a pseudopod with apical hooks
in the ventral position. The main feature of the abdomen
is the crown of hooks at the extremity, used to attach the
larva firmly to the substrate. The nymphs, also aquatic, are
housed in a triangular cocoon of silk threads, open on the
head side and attached to a support. A pair of gills in the
form of branched filaments is housed on the thorax.
There are around 1,700 known species of black flies, divided
into around 25 genera. The Simuliidae family contains
many sibling species complexes, in particular the Simulium
damnosum complex, which may contain some 40 species.
As we shall see, these are true species that are difficult to
distinguish between just by looking at their morphology.
They do, however, have different biologies.
Simuliidae: development cycle and bio-ecology
The females tend to lay their eggs on a totally or partially
immersed support. During their lifetime, they lay between
800 and 1,000 eggs in four or five clutches. After an
average of one week, the eggs hatch and release the first-
stage larvae. In the great majority of species, the larvae and
nymphs live in fresh running water. Black-fly larvae are
attached by abdominal hooks to immersed substrates, such
as stones or branches; certain species remain attached to
the carapace of crustaceans or aquatic insects. The larvae
can also weave a silk web to which they remain anchored
when moving from one support to another, to prevent
them from being carried away by a strong current. Larval
Fig. 5
Black fly
© OMS/WHO
89
habitats are found at sites where the currents run strong
and the water is highly oxygenated: fords, rapids, waterfalls,
inverts, slopes of dams, irrigation canals, etc. These larvae
feed on live prey and plankton that they capture using
the bristle fans of their premandibles, which act as filters.
The number of successive larval stages varies, even within
a given species: usually between six and nine (sometimes
up to 11), which last in tropical environments for five to
fifteen days. In cold climates, certain species can hibernate
for several months during the larval stages. The nymphs are
also attached by their cocoon in the same breeding sites.
They are immobile and do not feed. This stage lasts for
two to ten days. The adult emerges underwater during the
daytime and immediately rises to the surface.
The bio-ecology of adult black flies is still not well
understood. Only the females are haematophagous; blood
sucking, which lasts 3 to 6 min, always takes place outdoors
during the daytime on a mammal or a bird, depending on
the feeding preferences of the species concerned. Black
flies exist throughout the world, from the equator to the
Arctic. They have been observed at an altitude of 4,500 m
in the Andes. Whatever the region, there are major seasonal
variations in the dynamics of black fly populations.
Extraordinary pullulations are observed during summer
in the Arctic, which can cause a considerable nuisance. In
certain species their dispersal can be extensive; powerful
flight enables black flies to travel tens (or even hundreds)
of kilometres. Little is known about the lifespan of adult
females; in tropical regions it seems to be in the order of
two to four weeks, and the gonotrophic cycle is usually
three to seven days. In the S.  damnosum complex, 15 to
20 generations a year have been observed. In contrast,
univoltine species (one generation per year) persist in
Arctic areas.
Simuliidae: medical and veterinary importance
The medical importance of black flies lies mainly in
their role as vectors of onchocerca. In humans, the filaria
of Onchocerca volvulus is the agent of onchocercosis, a
potentially disabling disease that attacks the eyes, which
is spread by rural foci in sub-Saharan Africa and Latin
America. Other species of onchocerca, also spread by black
flies, are found in wild and domestic animals, especially
cattle and ducks. Black flies are also vectors of the human
filaria of Mansonella ozzardi in tropical America, as well as of
bird protozoa, such as Leucocytozoon, in turkeys, geese and
ducks. Moreover, large numbers of bites inflicted by black
flies can cause severe reactions in both humans and animals.
Finally, black flies may be involved in the transmission of
viruses, such as the one responsible for vesicular stomatitis.
Ceratopogonidae
The Ceratopogonidae family contains many species of small
Diptera, the biology of which is poorly understood. They
90
are sometimes referred to under the rather imprecise name
of ‘midges’.
Ceratopogonidae: morphology and systematics
Adult Ceratopogonidae, which measure 0.6 to 5 mm in
length, have a short proboscis, short stubby legs and wings
that are often speckled (Fig. 6).
The aquatic or semi-aquatic larvae are vermiform with a
sclerotised cephalic capsule and chewing mouthparts. The
nymph has two prothoracic respiratory horns.
The vast and complex Ceratopogonidae family is widely
dispersed around the world. There are more than
4,000 species, divided into 60 or so genera, including
one, the genus Culicoides, which alone contains more
than 1,000 species and is of real medical and veterinary
importance.
Ceratopogonidae: development cycle and bio-ecology
The females lay between 60 and 200 eggs in each clutch,
grouped together in clusters, which hatch two to fifteen days
later. The aquatic or semi-aquatic larvae live in silt or mud
on the banks of shallow bodies of fresh, brackish or even
salt water. Some develop in decomposing plants (severed
banana tree trunks, manure, dead leaves, tree hollows).
Depending on the species and temperature, larval
development may last from two weeks to seven months or
more (larvae hibernate in cold temperate countries). They
are succeeded by a nymph, with poor mobility, from which
the adult emerges after two to ten days.
The adult females are generally haematophagous, biting
hot- or cold-blooded vertebrates that are usually exophagic
and exophilic. Some are diurnal while others are nocturnal.
Fig. 6
Biting midge (Culicoides nubeculosus)
Photo: J-B. Ferré © EID Méditerranée
Rev. Sci. Tech. Off. Int. Epiz., 34 (1)
Very little is known about the ecology of Ceratopogonidae,
which appears to vary considerably, depending on the
species. Dispersal is low, approximately 500 m, but on
several occasions they have been observed to travel over
large distances (several hundred kilometres), carried by the
wind. The lifespan of females is around one month. In the
few cases where it has been observed, the gonotrophic cycle
was in the order of two to four days. In temperate regions,
most species are univoltine, although others have three or
four successive generations in one year.
Ceratopogonidae: medical and veterinary importance
These insects, mainly of the genus Culicoides, are responsible
for the transmission of many viruses (14). Only one of
them is truly pathogenic for humans: the Oropouche virus
in South America. As for veterinary pathology, several
important viroses use Ceratopogonidae as vectors and are
of considerable economic importance:
– bovine ephemeral fever, caused by a Rhabdovirus, which is
found in Asia, Australia and Africa
– bluetongue, caused by an Orbivirus, which is a very severe
disease of ruminants, especially sheep (15, 16)
– African horse sickness, also caused by an Orbivirus, which
is prevalent in Africa and the Middle East and can result in
mortality levels of 90%
– Schmallenberg disease, caused by an Orthobunyavirus,
which has spread in Europe since its emergence in 2011,
where it causes abortion and malformation in newborn
lambs and calves.
These insects may also be involved in the transmission of
the virus responsible for vesicular stomatitis.
In humans, three filarian species of the genus Mansonella
are also transmitted by Culicoides: Mansonella ozzardi in
Latin America and the Antilles; M. perstans in West and
Central Africa and tropical America; and M. streptocerca in
the forests of West and Central Africa. Filariae of animals
are also transmitted, such as Onchocerca of horses and cattle
and other filariae of birds and amphibians.
Finally, several protozoa of monkeys and birds use
Ceratopogonidae as vectors.
Culicidae (mosquitoes)
This Diptera family is of considerable medical and veterinary
importance. Mosquitoes are among the most dangerous
vectors for humans and are involved in the transmission of
many animal pathogens (Fig. 7).
Rev. Sci. Tech. Off. Int. Epiz., 34 (1)
Fig. 7
Mosquito (Aedes)
© Institut Pasteur
Mosquitoes: morphology and systematics
Adult mosquitoes are winged insects, 3 to 20 mm in
length, with biting mouthparts forming a long proboscis,
protected when at rest inside a flexible sheath called the
labium. The globular head has two composite eyes, a
pair of maxillary palpi and a pair of antennae. The thorax
has a pair of membranous wings, a pair of balancers (a
second pair of vestigial wings) and three pairs of legs. The
abdomen comprises ten segments, the last two of which
are morphologically modified to form the external genital
organs. In males, the very complex morphology of the last
abdominal segments is used in taxonomy.
Mosquito eggs take various forms, depending on the genera,
and measure between 0.5 and 1 mm. The eggs of certain
species have lateral or apical floats.
Mosquito larvae have a distinct chitinous head with two
antennae and chewing mouthparts, a broad thorax and
an abdomen in nine visible segments. Mosquitoes other
than anophelines have spiracles opening at the apex of a
respiratory siphon, a single organ in the form of a truncated
cone. Each of these organs has features that vary greatly
from one species to another.
The mosquito nymph comprises a cephalothorax and an
abdomen with nine visible segments, the last of which
bears a pair of racquet-shaped paddles. At rest, the nymph
remains below the surface of the water, breathing through
its two prothoracic horns.
The Culicidae family contains around 3,600 known species.
This family is generally subdivided into three sub-families:
Toxorhynchitinae, Anophelinae and Culicinae, within
which there are 40 or so different genera, including Culex,
Aedes, Ochlerotatus, Mansonia, Anopheles and Toxorhynchites.
91
Mosquitoes: development cycle and bio-ecology
Mosquitoes undergo a complete metamorphosis, i.e. during
its lifetime an individual passes through several successive
stages: egg, larva, nymph and adult, the first three of which
are aquatic (17, 18).
The eggs are generally laid singly by the female on the
surface of a body of water (Anopheles), sometimes adhering
to an inert support (Aedes), and some lay all their eggs in
one clutch that either floats like a raft (Culex) or remains
clustered in one place (Mansonia). This stage usually lasts
from two to three days, but the eggs of certain mosquitoes
(Aedes, Haemagogus) have a waterproof chorion enabling
them to withstand desiccation for several weeks or months
(‘durable’ eggs) and to await the first rains before hatching.
Each clutch contains around 50 to 500 eggs, meaning that,
during her lifetime, a female can lay between 1,000 and
2,500 eggs in several clutches.
When the egg hatches, still in water, the larva is released.
Mosquitoes undergo four successive larval stages, each
of which is separated by a moult, which allows a sudden
increase in size: from 1 mm for larvae in the first stage to
around 2.5 cm for larvae in the fourth stage. These highly
mobile larvae feed on plankton-like microorganisms:
bacteria, algae, protozoans, and sometimes large live prey,
such as the larvae of other insects or mosquito species.
When the weather conditions are conducive, the total
duration of the larval stage is around 8 to 12 days. When
the weather is unfavourable, as during the winter in Eurasia,
larval development may take several months.
At the end of the fourth larval stage, a moult transforms
the insect into a nymph: a mobile aquatic organism that
does not feed for the two or three days it takes to become
an adult. In a phenomenon known as ‘emergence’ the adult
becomes ‘free’ and passes from an aquatic existence to an
aerial existence. Emergence must take place away from any
disturbance; during this phase the insect is often defenceless
against surface predators.
The ecology of the pre-imaginal stages is governed
entirely by the type of breeding site (19). Each species has
preferences that vary in their stringency. The different types
of breeding sites can therefore be characterised by a range
of factors, such as:
– water movement, temperature and pH
– mineral salt content
– sunshine levels
– size
– type (permanent or temporary, natural or artificial)
– type of vegetation
92
– agricultural practices (in the case of irrigated crops)
– levels of organic or chemical pollution.
The first distinction that needs to be made in adults is
between males and females. Males are not haematophagous
and have a short lifespan and low dispersal. Females are
mostly haematophagous with an average lifespan of around
two months (sometimes much more), and their dispersal
is governed by the search for a vertebrate host (mammals,
birds, reptiles or amphibians, depending on the species)
and a breeding site. For mosquitoes, flight dispersal ranges
from a few hundred metres to several kilometres but, in
addition to this active dispersal, there is passive dispersal
by winds and human means of transport, which may
pose problems for epidemiologists because they can lead
to unexpected emergencies. Certain species are strict or
preferential zoophiles or anthropophiles, with the latter
including endophagic mosquitoes that enter dwellings to
bite and exophagic mosquitoes that feed outdoors (20).
Each blood meal is usually followed by egg-laying two or
three days later. The timing of bites varies, depending on
the species. Anophelines are mostly nocturnal, while many
Aedes spp. bite during the day. It is important to ascertain
the frequent resting places used by mosquitoes between
feeding sessions because in many cases they can be attacked
there, using residual insecticides.
These insects are found worldwide, where they colonise
a variety of environments, from rainforests to semi-arid
zones, as well as urban areas. During periods not conducive
to activity (winter in temperate regions and the dry season
in tropical zones), mosquitoes persist either in the form of
fertilised females at rest (not sucking blood or with non-
developing ovaries), or in the form of durable eggs that will
hatch with the first rains.
Mosquitoes: medical and veterinary importance
Mosquitoes can not only be a major nuisance (21), they
are of considerable importance for human and veterinary
medicine because of their role in the transmission of
diseases causing major public health problems: arboviroses,
malaria and filarioses.
Arboviruses transmitted by mosquitoes are extremely
numerous; they belong mainly to the genera Alphavirus,
Flavivirus, Phlebovirus and Orthobunyavirus. The majority
are zoonotic agents and some are among the most
dangerous for humans, including yellow fever, dengue fever
and Japanese encephalitis, as well as Chikungunya virus,
several American equine encephalitis viruses, the viruses of
Murray Valley, Saint Louis and California encephalitis, West
Nile virus, Rocio virus, Ross River virus, Rift Valley fever
virus, and many others. Mosquitoes can act as reservoirs
for viruses in these various arboviral systems, at least
Rev. Sci. Tech. Off. Int. Epiz., 34 (1)
temporarily, and, depending on their feeding preferences,
they can act as vectors for transmission to humans or to
certain wild or domestic animals. As usual, it all depends on
ecological contacts between natural populations.
Of the Plasmodium sporozoans, four are responsible for
malaria in humans. All Plasmodium species affecting primates
are transmitted by mosquitoes of the genus Anopheles.
Finally, all three species of nematode responsible for
lymphatic filariasis in humans are transmitted by mosquitoes.
They are Wuchereria bancrofti (causing Bancroft’s filariasis),
by far the most widespread and specific to humans; Brugia
malayi (agent of Malaysian filariasis), specific to Asia, which
can also infect wild and domestic animals; and Brugia
timori (causing Timor filariasis), which exists only in some
of the Lesser Sunda Islands. Many animal filariae are also
transmitted by mosquitoes, including Brugia, Dirofilaria,
Setaria and Wuchereria.
Tabanidae (horseflies)
Tabanidae, or horseflies, are Diptera that are found
throughout the entire world; in most cases the females are
haematophagous. They are well known for their often very
unpleasant bite.
Horseflies: morphology and systematics
Adult horseflies are quite large Diptera (5–25 mm in length).
They are large, dark-coloured flies. The head has two large
eyes, sometimes featuring brightly coloured bands. The
mouthparts form a short, strong proboscis, at least in the
blood-sucking species. The wings are hyaline and they are
either entirely transparent or marked with brown spots.
Horsefly larvae are elongated and grey or brown in colour.
They have a retractable cephalic capsule, a thorax in three
segments and an abdomen with eight segments, some of
which have pseudopodia; the posterior end of the body
has an organ similar to a respiratory siphon in the dorsal
position.
The large Tabanidae family, which contains some 4,000
species, is generally subdivided into three main sub-
families:
– Pangoniinae (non-haematophagous and therefore of no
medical or veterinary interest)
– Chrysopsinae, which include the genus Chrysops,
characterised by yellow marks on the thorax and abdomen,
and wings with smoky marks
– Tabaninae, which include genera of medical and, above
all, veterinary importance, such as Atylotus, Haematopota,
Rev. Sci. Tech. Off. Int. Epiz., 34 (1)
Tabanus and Hybomitra. Tabanus are large insects with
completely hyaline or smoky wings; Haematopota, on the
other hand, are small with smoky wings, marked with
many small, light spots.
The larvae and nymphs of many species are still unknown;
identification of these stages is often very difficult, or even
impossible.
Horseflies: development cycle and bio-ecology
The eggs are laid in compact clusters, deposited on a support
(plant, rock, etc.) near water. After a few days, they hatch
into larvae which fall onto the substrate. The larvae are
aquatic or semi-aquatic and develop very slowly (six months
to three years) through seven to thirteen successive stages.
During the final stage, they may grow to 3 cm in size.
The insect then moults and transforms into an immobile
nymph that does not feed. One or two weeks later the adult
emerges.
The larvae develop in the hollows of trees, in silt or mud
bordering bodies of water in wet grasslands, sometimes in
salt marshes or even in areas with heavy organic pollution.
Many are carnivores, or even predators of arthropods, but
others are saprophytic. The nymph stage takes place in the
same habitats.
Horseflies are not found in urban environments as they
frequent environments with little ecological change: prairies
and grasslands, forest canopies and swamps. Most females
feed on blood and are attracted by wild mammals or domestic
cattle. They are almost always exophagic and exophilic and
generally feed several times during the day (20–200 mg in
total). Although they are strong flyers, these insects do not
travel far (a few kilometres). The lifespan of the females
appears to be in the order of two to six weeks. A female
generally lays five or six clutches of eggs. Most horseflies
living in temperate regions produce only one generation a
year and are sometimes forced to undergo a diapause.
Horseflies: medical and veterinary importance
Many tabanids are mechanical vectors for different
trypanosomes of mammals; in particular, Trypanosoma
evansi, which is responsible for surra in horses, camels and
other animals in Africa and tropical Asia, and has been
introduced into South America. They can also mechanically
transmit Besnoitia besnoiti, which causes elephant skin
disease in cattle, and various bacteria, such as Anaplasma
and Francisella tularensis (the agent of tularemia) and,
no doubt, the anthrax agent. Horseflies also transmit
retroviruses, such as the virus of equine infectious anaemia.
Finally, this family includes biological vectors of various
filariae of mammals, including Loa loa, an African filaria
which is pathogenic for humans.
93
Muscidae
Muscidae are a vast family of Diptera, but only two sub-
families have haematophagous species that could transmit
infectious agents: Glossininae (tsetse flies) and Stomoxyinae
(stable flies).
Tsetse flies: morphology and systematics
Adult tsetse flies are biting insects measuring 6 to 16 mm in
length, with a fly-like appearance, brown or grey in colour.
They are nevertheless easily recognised by their antennae
with a single branched hair (arista), the specific venation
of their wings and, above all, their mouthparts, which form
a long proboscis, pointing forwards when at rest (Fig. 8).
The morphology of the tsetse fly larva is very similar to that
of a maggot (5–8 mm), but at the posterior end it has two
dark, sclerified protuberances with respiratory spiracles.
The nymph (or pupa) is enclosed in a puparium, a sort
of rigid, brown, barrel-shaped husk, which also has two
protuberances at one end.
Tsetse flies are a small group of Diptera with a very specific
and uniform morphology and biology (some entomologists
believe it is justified to group these insects into a separate
family of Glossinidae). There are 31 known species or
subspecies, all of which fall within the genus Glossina,
specific to sub-Saharan Africa (two or three species also exist
in a small area of the Arabian Peninsula), with the exception
of the temperate south of the continent, and, of course, the
high-altitude zones, which are too cold. Three sub-genera
are generally recognised within the genus Glossina:
– the sub-genus Nemorhina (a fairly small, hygrophilic
species, with a preference for inhabiting pre-forest or forest
zones, and sometimes mangroves or plantations)
Fig. 8
Tsetse fly (Glossina morsitans)
Photo: Jean-Paul Hervy © IRD
94 Rev. Sci. Tech. Off. Int. Epiz., 34 (1)

– the sub-genus Glossina (a medium-sized xerophilic
species, found in wooded grassland, thickets and dense
bushes, and sometimes in woodlands)
– the sub-genus Austenina (a large species, especially
widespread in dense rainforest and sometimes in mixed
woodland and grassland or in large and dense gallery
forests).
Tsetse flies: development cycle and bio-ecology
Female tsetse flies usually mate only once in their lifetime
and their average lifespan is at least two to three months
(even as much as nine months for certain females). Tsetse
flies are larviparous Diptera. The three larval stages take
place in the uterus of the female, which therefore gives birth
to a larva at the end of stage three. A single larva is laid at
a time and, over its lifetime, the same female lays only 6 to
13 larvae. Once laid by the female, the larva quickly buries
itself in the soil a few centimetres deep, where it transforms
into larval stage four and then the pupa (nymph). The pupae
are found in the ground, concentrated in pupa breeding
grounds in shady locations at the base of trees and bushes.
This pupal stage lasts around one month.
Tsetse flies: medical and veterinary importance
Tsetse flies are responsible for transmitting several species
of trypanosomes that are parasitic on wild and domestic
vertebrates. Trypanosomoses of cattle cause significant
economic losses as they are responsible for some of the most
important vector-borne diseases on the African continent.
Humans also act as hosts for two forms of Trypanosoma
brucei, which can cause the dreaded disorder known as
sleeping sickness.
Stomoxes (stable flies)
These biting insects constitute the sub-family Stomoxyinae
within the Muscidae family. Although they are of little
interest for human medicine, stable flies are of major
importance for veterinarians.
Stable flies: morphology and systematics
The morphology of these small haematophagous flies
(3–7 mm) includes recognisable mouthparts forming a
proboscis similar in appearance to that of tsetse flies.

The adults also have rather strict environmental The Stomoxyinae sub-family contains around 50 species.
requirements. Many tsetse flies have been observed in Only two genera are of interest: Stomoxys and Haematobia,
ecotones (an interface zone between two ecosystems; for which are distinguished by the morphology of their palpi
example, the edge of a forest), such as the fringes of wooded (23) (Fig. 9).
zones, forest trails, and gallery forests (22). Some species
have been known to frequent the outskirts of villages, or Stable flies: development cycle and bio-ecology
even the green spaces within towns. The various types
include: The larvae of these flies develop in all types of high-humidity
organic matter; in particular, in dung and faecal matter or in
– grassland species, such as Glossina morsitans, which live
in wooded grassland and sometimes in more arid areas,
concentrated in the dry season in more wooded areas near
waterholes

– riparian species associated with water courses and

therefore located in gallery forests, forested islands,
mangroves and on the banks of lakes (G. palpalis,
G. tachinoides)

– forest species that require a high level of humidity, but

also venture out to the forest edges, alongside tracks or in
plantations (G. fusca).

These flies fly rapidly in search of their hosts, with which

brief contact takes place in preferred sites: alongside tracks,
stockyards, on the outskirts of villages and plantations,
close to water points, etc. Both tsetse fly sexes are
haematophagous. They suck blood during the daytime
from wild animals (antelope, buffalo, warthogs, crocodiles,
etc.) or domestic animals, and sometimes humans. At rest,
tsetse flies can be found on vegetation (trunks, branches, Fig. 9
leaves), at varying heights, depending on the species, season Stable fly (Stomoxys calcitrans)
and time. Photo: G. Duvallet
Rev. Sci. Tech. Off. Int. Epiz., 34 (1)
rotting vegetation. The females lay their eggs in such matter,
the larval stage lasts six to ten days and the pupal stage an
average of five to seven days. In total, the life cycle varies
from two to six weeks, depending on the temperature.
The adults of both sexes are haematophagous, feeding on
domestic or wild mammals and only occasionally attacking
humans. They are found in abundance around herds of
cattle in pasture land, near cattle stockyards and in stables.
For the species associated with domestic animals, in
particular the cosmopolitan species Stomoxys calcitrans, the
main hosts are cattle, horses and, more rarely, dogs and
humans. High densities of these flies can sometimes be seen
on animals; they rarely leave their host and the female must
have a blood meal two or three times a day. These meals are
frequently interrupted by the defensive behaviour of their
hosts.
Stable flies: medical and veterinary importance
Incessant biting by these insects can be a considerable
nuisance for animals, and sometimes for humans. Moreover,
some species are intermediate hosts for nematodes, such
as Habronema, horse stomach parasites, or cattle filariae
(Stephanofilaria). Owing to their frequent, and often
interrupted, feeding, these Diptera may occasionally act
as mechanical vectors of various viruses (agents of equine
infectious anaemia, lumpy skin disease of cattle, African
swine fever, etc.) and bacteria (24).
Pupipara
Pupipara are a very curious set of haematophagous Diptera
grouped into three families, characterised by their highly
specific morphology: Hippoboscidae, Nycteribiidae and
Streblidae. The latter two contain only species restricted
to bats, while Hippoboscidae are ectoparasites of birds and
mammals (and sometimes humans).
Hippoboscidae are large, flat-bodied flies with strong claws
on their feet, and biting mouthparts reminiscent of tsetse
flies; their wings are of normal development, but sometimes
smaller or, in certain species, only temporarily present or
95
totally absent. These Diptera are ‘pupiparous’, meaning that
the egg and the larva develop inside the body of the female,
which therefore lays an older larva that transforms into a
pupa a few minutes later (25).
They are parasitic on many species of birds, as well as
on wild mammals (e.g. deer) and domestic mammals
(e.g. horses, cattle, dogs, camels, sheep). The veterinary
importance of Hippoboscidae lies in the weight loss and
anaemia that can accompany a very heavy infestation, as
well as the transmission of parasites, such as trypanosomes
of ruminants or Haemoproteus of birds.
Conclusion: the importance of
insect vectors for surveillance
and control
Clearly, an understanding of the functioning of vector
systems is a prerequisite for defining surveillance
procedures (constantly bearing in mind the ever-changing
epidemiological situation) and for determining appropriate
methods of anti-vector control as a part of disease prevention
programmes.
To understand the functioning of a vector system, it is
necessary to understand the biology of the vector(s), the
vertebrate(s) and the microorganism in parallel. For this,
the first step is to identify the vectors and the second is
to understand all aspects of their bio-ecology (geographical
distribution, population density, environmental
preferences, feeding preferences, rate of attack, duration of
gonotrophic cycle, lifespan, hibernation/aestivation habits,
etc.) in order to be in a position to correctly evaluate their
vectorial capacity. Such is the task of medical-veterinary
entomologists.
96
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