Arthropod vectors  Tsetse flies

Tsetse flies
Author: Dr Reginald De Deken
Licensed under a Creative Commons Attribution license.

TABLE OF CONTENTS
INTRODUCTION............................................................................................................................................................2

IMPORTANCE ............................................................................................................................................................... 4

DISEASE TRANSMISSION ...........................................................................................................................................5

ANATOMY/MORPHOLOGY ..........................................................................................................................................8

IDENTIFICATION/DIFFERENTIAL DIAGNOSIS......................................................................................................... 16

DISTRIBUTION............................................................................................................................................................ 22

The subgenus Morsitans: ....................................................................................................................................... 23

The subgenus Palpalis: .......................................................................................................................................... 23

The subgenus Fusca: ............................................................................................................................................. 23

SAMPLING AND COLLECTION METHODS .............................................................................................................. 24

Sampling methods .................................................................................................................................................. 24

Fly-round ..................................................................................................................................................... 24

Traps ........................................................................................................................................................... 25

The electric screen ...................................................................................................................................... 26

ECOLOGY/LIFE CYCLE ............................................................................................................................................. 28

Ecology................................................................................................................................................................... 28

Life cycle ................................................................................................................................................................ 29

CONTROL ................................................................................................................................................................... 32

Ecological control ................................................................................................................................................... 33

Chemical control ..................................................................................................................................................... 34

Biological control .................................................................................................................................................... 41

Genetic control ....................................................................................................................................................... 42

PATTEC ................................................................................................................................................................. 43

REFERENCES............................................................................................................................................................. 44

1|Page
Arthropod vectors  Tsetse flies

INTRODUCTION
Tsetse flies, being the vector of human and animal trypanosomosis, constitute one of the major health
and agricultural livelihood constraints of sub-Saharan Africa. It had long been known that domestic
stock could not be kept where tsetse flies occurred and that disease was associated with the flies.
However, it was not until Bruce, working in Zululand in 1895, demonstrated the transmission of
pathogenic trypanosomes of livestock by tsetse flies that the reason for livestock deaths became
known. The role of tsetse flies as vectors of human trypanosomosis was also demonstrated by Bruce
(Bruce et al., 1909).

The presence of tsetse flies has been a major obstacle to the development of much of the continent.
There are those, even now, who look upon the tsetse fly as the guardian of the natural ecosystems of
Africa, and who would like this fly to remain until humans have learnt to manage the land in a
sustainable manner. At present, though, the human population of Africa is expanding rapidly, and the
economic situation is such that the livelihood of the burgeoning population can, for the most part, be
accommodated only as subsistence farmers. The importance of livestock to such people is very high,
not only as a source of food, draught power, and money, but also for the important role that livestock,
especially cattle, play in cultural affairs. Under these circumstances it is essential to try to eliminate, or
at least control, the diseases of livestock, amongst which trypanosomosis plays a major role in Sub-
Saharan Africa. If tsetse flies wouldn‟t be capable of spreading African animal and human
trypanosomosis, their impact on the African continent would be minimal. Therefore it is also important
to have knowledge of the factors (environmental, physiological, host/vector interactions, etc.) that can
influence the transmission of the pathogen by the vector.

For those, who aren‟t familiar with the morphology of tsetse flies (see Dr. Livingstone‟s tsetse
collection), the morphological differences between tsetse and other haematophagous diptera will be
explained as well as the morphological and habitat characteristics of the three distinct tsetse fly
groups.

2|Page
Arthropod vectors  Tsetse flies

Dr. Livingstone’s Tsetse collection

In July 2000 the Lomé meeting of Heads of State of the Organization of African Unity decided to
eradicate the tsetse fly from the African continent. The Pan African Trypanosomosis and Tsetse
Eradication Campaign (PATTEC) were borne. At first glance, it may be surprising why tsetse flies
aren‟t eradicated already, since the tsetse fly produces at most a single larva per week and is
extremely susceptible to insecticides. However, some important issues have to be solved before
tsetse eradication may be accomplished.

Available multimedia

In depth information and analysis on specific issues related to the

problem of tsetse & trypanosomosis, its management and
intervention strategies can be found at the Programme Against
African Trypanosomosis (PAAT) website:

http://www.fao.org/Ag/againfo/programmes/en/paat/papers.html

3|Page
Arthropod vectors  Tsetse flies

IMPORTANCE
Tsetse flies represent a key constraint for food security in the 37 sub-Saharan African countries where
tsetse flies are present. Tsetse flies are the biological vectors of several trypanosomes affecting
livestock and of two trypanosome species, Trypanosoma brucei gambiense and T.b.rhodesiense,
causing respectively in eastern and southern Africa a chronic form and in Western and Central Africa
an acute form of human sleeping sickness. The distribution of sleeping sickness areas has a focal
nature and the localisation of the actual areas fluctuates over the course of time. Important vectors for
human African trypanosomosis are: all subspecies of Glossina fuscipes and of G.palpalis, G.pallicera
pallicera, G.swynnertoni, G. morsitans centralis, G. morsitans morsitans and G.pallidipes.

Contrary to human African trypanosomosis, almost all tsetse flies can transmit the trypanosomes
responsible for animal trypanosomosis and the disease has a wide geographic distribution.

Distribution of tsetse and cattle

Through the disease they transmit, these trypanosomes hamper livestock keeping or genetic
improvement of indigenous breeds and an efficient use of animal traction. The economic cost of
trypanosomosis will largely depend on the livestock production system (the type of animal bred;
trypanotolerant or susceptible – draught, dairy or beef animals) and is inversely related to the
endemicity of the disease (epidemics occurring essentially when livestock is encroaching on
undeveloped areas). According to Kristjanson P.M. (1999) the disease costed African livestock
producers and consumers at that time an estimated $1340 million annually, without including indirect

4|Page
Arthropod vectors  Tsetse flies

livestock benefits such as manure and traction. The most striking direct effects of the disease are
observed in the field of milk production, calving rate and mortality among calves and draught animals.
Apart from these direct economic effects of the disease, there are also considerable indirect losses
due to unbalanced land use and settlement patterns, difficulties to integrate livestock breeding and
agriculture, reduced use of draught power and massive expenditures on trypanocidal drugs in tsetse
infested areas. On top of this, resistance to trypanocidal drugs is spreading. The highest losses from
animal trypanosomosis currently occur in areas with high cattle population densities on the margins of
the tsetse distribution and where animal traction is an important component of farming systems
(Shaw, 2009).

Available multimedia

Videos: A 45 min lasting video “Survival – Deadliest disease”

produced by rockhopper TV for BBC can be visualised or
downloaded at

http://www.rockhopper.tv/films/detail/survival-the-deadliest-disease

Some videos on the zoonotic form of the disease caused by T.brucei

rhodesiense may be consulted on the ICONZ website:

http://www.iconzafrica.org/videos/sleeping_sickness_march_2010.mov or
http://www.iconzafrica.org/videos/sleeping_sickness_long.mov

DISEASE TRANSMISSION
The transmission of trypanosomes by the tsetse fly occurs cyclically. This means that the vector,
when he feeds on an infected host, is not immediately infectious but that the pathogen during “the
extrinsic incubation period” has to pass through a developmental cycle inside the vector. During this
period the trypanosome undergoes substantial morphological and metabolic changes. The
developmental cycle of Trypanosoma vivax is restricted to the mouthparts and lasts 5 to14 days.
Trypanosoma congolense develops in the midgut and the proboscis and its development takes
generally around 12 - 18 days. Trypanosoma brucei species have the most complicated cycle with
trypanosomes migrating from the midgut to the salivary glands of the tsetse fly. The cycle takes
between 17 and 45 days.

The basic reproductive rate of an infection is the mean number of secondary cases caused by an
infected individual into a population with no pre-existing immunity to the disease and in the absence of
control interventions. It is often denoted R0. However, what is often claimed to be R0 is, in fact, often
simply the assessment of a threshold (the disease spreading or not spreading), not of the average
number of secondary infections. This is particularly true in case of a vector-borne disease because
calculations to assess the R0 of a vector-borne disease are very complex (since both the vector and

5|Page
Arthropod vectors  Tsetse flies

the host are responsible for the secondary cases and since several vectors and hosts may be
present) and the exact value of several contributing parameters are often unknown. Thresholds will
merely demonstrate whether a disease will die out (if R0 < 1) or will be able to spread in a population
(if R0 > 1).

The important parameters governing the spread of a disease may be demonstrated by the formula
(based on the work of Macdonald (1957) and Smith et al. (2007)) to assess the basic reproductive
rate of a vector-borne infection:

2 n
ma bcp

R0 = ---------------

r g

Where:

m = rate of vectors to hosts (amongst others this will depend on the space utilization of the host and
alternative hosts in relation to tsetse mobility and distribution).

a = feeding rate of the vector on the host (depends on the feeding rate “f” at that moment, and also
on the feeding preference “q” of tsetse flies for a particular host) so a = fq

b = probability that a host may become infected once bitten by an infected vector

c = the probability that the vector becomes infected from a bite on an infected host

-g
p = daily survival rate of the vector, so p = e and the death rate g = -ln(p)

g = death rate of the vector

n = extrinsic incubation period (number of days before the vector becomes infectious)

r = the rate with which the host cures from the infection.

Regarding m:

 The rate of vectors to hosts (m) may decrease as a result of: vector control, disappearance or
fragmentation of tsetse habitat, or a massive immigration of cattle in an area (e.g. temporarily
available pasture land)
 The rate of vectors to hosts (m) may increase as a result of:
- livestock movements (e.g. cattle moving into game reserves),
- particularly beneficial climatic conditions for tsetse (at the end of the rainy season the
tsetse fly population density is generally at its top because of an increased
reproduction in the previous months and an increased longevity at that time),
- tsetse flies seeking refuge in forest galleries during the hot dry season resulting in a
temporarily increased transmission risk for cattle at watering points. Notwithstanding

6|Page
Arthropod vectors  Tsetse flies

the tsetse population is not at its highest density at that time high trypanosomosis
incidences may be observed,
- sudden immigrations of tsetse flies (these may be particularly high in case of tsetse
control operations when the control area is not isolated making the control effort often
unsustainable).

Regarding a: This important parameter depends on the number of feeds taken by the vector per unit
of time, which will increase when the climate becomes dry and hot, but it depends also on the feeding
preference (see the biology section).

Furthermore, it was demonstrated that upon colonization of the tsetse salivary glands with
Trypanosoma brucei, the anti-haemostatic activity of the saliva changes resulting in a prolonged
probing/feeding time, which may result in an increase of interrupted feeding and thus contribute to an
increased parasite transmission (Van den Abbeele et al., 2010). On the other hand, increased probing
activity was observed in T. congolense infected G. morsitans flies, which is believed to be caused by
physical interference of the parasite with phagoreceptors in the proboscis in combination with a
reduced diameter of the tsetse labrum due to the presence of parasite rosettes (Livesey et al., 1980).

Regarding b: This may depend on the “trypanotolerance” or “susceptibility” to trypanosome infection

of the host. Trypanotolerant animals are more difficult to infect (smaller value of “b”) and once infected
they may clear the infection more rapidly, resulting in a larger value for the parameter “r”. The
parameter b will also be reduced when prophylactic trypanocidal drugs are used.

Regarding c: This will depend on the vectorial competence (ability to become infected) and vectorial
capacity (the ease with which the development cycle in the vector is completed) of the vector. In most
tsetse flies taking up trypanosomes, these protozoa will be killed by the combined action of proteases,
lectins, immunopeptides and oxidative stress (Lehane, 2004; MacLeod, 2007). Young flies infect
themselves more easily with trypanosomes than old ones. However, it was shown that even older flies
can show a similar vectorial capacity in case the vector undergoes a stress of some kind which affects
the fly‟s immune gene expression (Akoda, 2009). The morsitans group of tsetse flies has a higher
infection rate than those of the palpalis group (except for T.vivax in West Africa). The fusca group is
generally a poor vector for the T.brucei group of trypanosomes but experiments showed it to be a
good vector for T.vivax and T.congolense.

Regarding p and g: these depend on the longevity of the vector, which will be influenced by
environmental conditions (climate, host availability, tsetse density dependent mortality) and possibly
by local tsetse control operations.

Regarding n: to become infectious the longevity of an infected tsetse fly must be superior to the
extrinsic incubation period of the trypanosome. This period varies according to the trypanosome
species and the environmental conditions (e.g. when temperatures are rising, the pathogen will
develop more rapidly in the vector).

It is obvious that the formula above is too simple to give a correct reflection of what happens in the
field; for example some trypanosomes can infect several livestock species (e.g. T.congolense infects
ruminants as well as pigs or dogs) and thus the vector can become infectious by feeding on any one
7|Page
Arthropod vectors  Tsetse flies

of those infected hosts. Feeding rate will differ depending on the particular host, its density and the
tsetse fly species involved.

Available multimedia

On the website of TDR animations can be found on the life-cycle of a

Trypanosoma brucei sp. In the human host

http://www.who.int/tdr/publications/documents/t-brucei-human-
host.swf

and the complex development cycle of T. brucei in the tsetse fly

http://www.who.int/tdr/publications/documents/t-brucei-tsetse-fly.swf

ANATOMY/MORPHOLOGY
The head bears a long piercing proboscis, which has a distinct basal bulb and points forward when at
rest. The mouthparts consist of 3 parts (the upper lip or labium, the hypopharynx and the lower lip or
labrum) surrounded by a pair of maxillar palps.

Lateral aspect of a female Glossina morsitans. Note

forward-pointing proboscis

Glossina morsitans

8|Page
Arthropod vectors  Tsetse flies

Glossina proboscis
Lateral aspect of the head of a tsetse fly with the
haustellum lowered to the feeding position

Lateral aspect of the antenna of a tsetse fly

Both sexes are hematophagous. The third segment of the antenna bears an arista with hairs (on the
upper surface only) which are themselves branched (feathered). Tsetse flies can detect odours by
means of sensilla situated on the antennae. Kairomones produced in the host's breath (e.g. carbon
dioxide, acetone or 1-octen-3-ol) and in the urine of African buffaloes and cattle (phenols) attract
tsetse (Vale et al., 1985 & 1988). The odours of humans and some animal species (e.g. waterbuck)
repel some species of tsetse flies (Gikonyo, 2002).

9|Page
Arthropod vectors  Tsetse flies

Glossina antenna and palps

The compound eyes are widely separated by the frons and are important in short-range host location.
The eyes can also distinguish between light and dark, a useful attribute for seeking out shaded
microclimates when ambient temperatures are at lethal levels.

The wings at rest are folded scissor-like, so that they are fully overlapping one another. The wing
venation is typical with the discal cell (comprised between 4th and 5th longitudinal vein and the
posterior cross-vein) being “hatchet” shaped (shaped like a butcher's cleaver).

Lateral aspect of a female Glossina morsitans.

Note forward-pointing proboscis

Dorsal aspect of a female Glossina morsitans

showing the position of the wings when the fly
is at rest. Note the forward-pointing proboscis

10 | P a g e
Arthropod vectors  Tsetse flies

Glossina wing

„Hatchet‟ cells

Wing of a tsetse fly, with the ‘hatchet’ cell indicated by stippling

Abdomen: The male presents at the ventral side of the posterior abdomen a tumefaction: the
hypopygium that is in fact the folded male terminalia.

Glossina male and female

11 | P a g e
Arthropod vectors  Tsetse flies

Ventral aspect of the terminal portion of the abdomen of

both sexes of Glossina morsitans

During copulation the hypopygium is deployed and reveals the superior claspers and penis.

12 | P a g e
Arthropod vectors  Tsetse flies

Hypopygium folded out

The claspers are used for withholding the female during copulation and their form is characteristic for
each subgenus.

13 | P a g e
Arthropod vectors  Tsetse flies

Glossina superior claspers

The internal anatomy of tsetse flies is characterized by 2 extremely long salivary glands which end in
the anterior part of the abdomen, an alimentary tract subdivided in an oesophagus, a crop, which
receives the major part of the blood meal before the meal is diverted to the midgut, the proventriculus
which controls the passage of blood between the crop and the midgut and secretes the peritrophic
membrane (a semi-permeable membrane protecting the midgut cells), a midgut responsible for water
absorption, digestion of the meal and absorption of the nutrients and behind the insertion of the
Malphigian tubules (with a function comparable to kidneys) the hindgut.

14 | P a g e
Arthropod vectors  Tsetse flies

Glossina proventriculus and gut

Glossina salivary glands

The female reproductive system consists of paired ovaries each consisting of 2 ovarioles, the uterus
and two spermathecae.

Glossina spermatheca

Glossina female reproductive organs

As the 4 ovarioles are producing a single follicle in turn in a predictable order, this sequence can be
used to determine the age of an adult female tsetse (Saunders, 1960; Challier, 1965).

15 | P a g e
Arthropod vectors  Tsetse flies

IDENTIFICATION/DIFFERENTIAL DIAGNOSIS
Tsetse flies of the fusca group are large (10.5 to 15.5 mm) and the abdomen is generally of dark and
uniform colour. The male genital armature comprises divergent claspers not joined by an intercalated
membrane.

Superior claspers Fusca group

Either all or only the two last segments of the hind tarsi are uniformly dark dorsally. On the
pteropleuron (a plate on the side of the thorax under the insertion point of the wing) short, thin hairs
are intermingled with longer, stronger hairs.

Fusca pteropleuron

Silks, that cover the fringe of the thoracic squama are long and corrugated and appear muddled and
disordered.

16 | P a g e
Arthropod vectors  Tsetse flies

Thoracic squama

Flies of the fusca group are primarily forest dwelling species (except G. longipennis and G.
brevipalpis), are most active at dusk and are mostly feeding on vertebrate animals.

Tsetse flies of the palpalis group are of small to average size (6.5 to 11 mm). Their abdomen is
uniformly brown or grey with some dark spots (except G.tachinoides of which the abdomen often
resembles that of a morsitans fly.

17 | P a g e
Arthropod vectors  Tsetse flies

Glossina fuscipes Glossina tachinoides

The male genital armature comprises divergent superior claspers joined by an intercalated
membrane. The tarsi of the hind legs are uniformly dark.

Tarsus of hindleg G.fuscipes

Bristles on the pteropleuron are composed of rather small, uniform hairs, which are much shorter and
weaker than those on the sternopleuron. Straight hairs cover the fringe of the thoracic squama.
Palpalis flies are mainly species living in the forest or in riverine or lakeside habitats. They are diurnal;

18 | P a g e
Arthropod vectors  Tsetse flies

apart from domestic animals they often feed on reptiles and birds and are more or less
anthropophagic.

Tsetse flies of the morsitans group are small or medium sized (7.5 to 11 mm) and the abdomen is
usually coloured with dark spots on a yellow background. Glossina austeni has a small size compared
to the other flies of the group.

Glossina pallidipes and austeni

Male genital armature comprises convergent claspers with an intercalated membrane. Only the last
two segments of the hind tarsi are darker than the more proximal segments, except in G.austeni
where the tarsi may be uniformly dark.

G.m. morsitans (Chipopela, 2007), C Mweempwa G. pallidipes (Lusandwa, 2007), C Mweempwa

Superior claspers Morsitans

19 | P a g e
Arthropod vectors  Tsetse flies

Hindleg Glossina morsitans

Bristles on the pteropleuron are composed of rather small, uniform hairs, which are much shorter and
weaker than those on the sternopleuron.

Pteropleuron and squama of Glossina morsitans

20 | P a g e
Arthropod vectors  Tsetse flies

Straight hairs cover the fringe of the thoracic squama. The morsitans species are primarily found in
savannah woodland and evergreen thickets. They are diurnal, feed on wildlife or livestock and are
more or less anthropophagic.

For identification of Glossina spp. appeal can be made to an identification key edited by FAO
(ftp://ftp.fao.org/docrep/fao/011/i0535e/i0535e03.pdf) and to specific software (Brunhes J et al., 2009).
For more information on this software see also:

http://www.cnev.fr/index.php/publications-et-outils/outils-didentification/identiciels

Larvae and nymphs of all Glossina spp are characterised by the presence of 2 polypneustic lobes.

Larva and pupa of tsetse fly

The tsetse fly may be confounded with a stable fly but a stomoxys at rest holds its wings spread,
while a resting tsetse folds its wings one on top of the other.

Stomoxys calcitrans

21 | P a g e
Arthropod vectors  Tsetse flies

Furthermore tsetse flies differ from stable flies by their feathery arista (in a stable fly the hairs on the
arista are without secondary branching) and the presence of the “hatchet cell” on the wing (which is
lacking in the stable fly).

Tsetse flies can be distinguished from tabanids by the antenna (projecting forwards in tabanids), the
discal cell (oval to hexagonal in tabanids), the mouthparts (adapted for both blood-sucking and
lapping in tabanids).

Haematopota pluvialis

Head Tabanus bovinus

Available multimedia

http://bioinfo-prod.mpl.ird.fr/identiciels/glossines/java/glossines.html

DISTRIBUTION
The distribution area of the genus Glossina is restricted to lowland rainforest and wooded savannah
regions of Africa south of the Sahara. Some species were also observed in south-west Saudi-Arabia.
Roughly speaking tsetse flies can be found between the 15th degree of latitude in the Northern
hemisphere and the 30th parallel south (eastern part) or the 20th parallel south (western part). The
distribution is however not uniform but often patchy. Country lying below the 500 mm isohyet does not
support tsetse flies unless there are watercourses with some vegetation along them (forest galleries).

22 | P a g e
Arthropod vectors  Tsetse flies

Mountains exceeding 1600 m of height generally form an insuperable obstacle for tsetse flies.
However it is not the altitude itself which plays a role, but rather the low temperatures prevailing at
these heights. Recently, however, G.m.submorsitans has been observed in Ethiopia and Cameroon
at a height of 2000 m, corroborating the hypothesis of global warming of the earth.

The subgenus Morsitans:

It is closely associated with open woodland and wooded savannah with Brachystegia (miombo) or
Colophospermum (mopane) trees in East and Central Africa (e.g. G.m.morsitans and G.pallidipes) or
Isoberlinia (doko) trees in West Africa (e.g. G.m.morsitans). These « savannah tsetse flies» penetrate
during the rainy season in arid areas, while during the dry season they concentrate in denser
vegetation along drainage lines or in better watered woodlands. The flies of the morsitans group are
usually very sensitive to agricultural development, degradation of natural habitats and reduced wildlife
density (especially G.swynnertoni, G.pallidipes and G.longipalpis), although they will feed readily on
domestic livestock when available.

The subgenus Palpalis:

It is found in gallery forests (e.g. G.p.gambiensis and G.tachinoides), near lakes and river systems,
flowing either into the Atlantic Ocean or in the Mediterranean (e.g. G.f.fuscipes and G.p.palpalis), or in
rain forest (e.g. G.p.palpalis, G.calliginea and G.pallicera). The majority of these “riverine” species
need a relatively high atmospheric humidity and shade although G.tachinoides can support great
climatic variations and is found from arid zones in Niger to humid degraded forest in Nigeria. The best
conditions for “palpalis” flies are a temperature of approximately 25°C and an atmospheric humidity of
80 to 85%. These “riverine” tsetse flies cope better with increased human occupation of the land and
environmental changes than the “savannah” flies.

The subgenus Fusca:

All the tsetse flies pertaining to the Fusca group, except for G.longipennis and G.brevipalpis, are
forest flies, including the humid tropical forest. As forest isn‟t particularly suited for livestock breeding
their importance as a vector of animal trypanosomosis is generally low. G.longipennis lives in the arid
and semi-arid savannas of East Africa. G.brevipalpis lives in the more humid parts of wooded
savannas of Central, South and East Africa. This species can fly over relatively long distances and
therefore is also observed in open grassland. It has some importance as a vector of animal
trypanosomosis.

Available multimedia

At the website of Program Against African Trypanosomosis (PAAT)

distribution maps of the different Glossina species may be consulted
and downloaded as a pdf or in GIS-format.

http://www.fao.org/Ag/againfo/programmes/en/paat/maps.html

23 | P a g e
Arthropod vectors  Tsetse flies

The maps, showing the predicted areas of suitability for the tsetse fly
species, were produced by Environmental Research Group Oxford
(ERGO Ltd) in collaboration with the Trypanosomosis and Land Use
in Africa (TALA) research group at the Department of Zoology,
University of Oxford.

SAMPLING AND COLLECTION METHODS

Sampling of tsetse flies is often used to assess the apparent density, which is related to:

1. The true density: the number of tsetse flies per surface unit. True density depends on the renewal
of the population (influenced by the development rate and the number of females one or two
months before), the mortality (particularly important during harmful climatic conditions and
shortage of hosts) and the rate of immigration and emigration.
2. The availability of the tsetse fly which varies according to:
- the behaviour of the tsetse flies with respect to the method used to capture the flies. Each
capture method gives a more or less significant bias while capturing only among certain parts
of the population of tsetse flies (e.g. fly rounds generally give an over-estimate of the number
of males and non-teneral males present).
- the activity of the tsetse flies which varies according to the moment the density is monitored
(influenced by climatic conditions); the place where the sampling is carried out (depending on
the season and their physiological state flies prefer some places and types of vegetation); and
the sex, the age and some cyclic phenomena such as hunger or pregnancy.

Thus longitudinal surveys will assess essentially the population dynamics and behaviour of the tsetse
fly with respect to the spatial (ecological zones) and seasonal variations of the environment.

Cross-sectional sampling of adult tsetse flies is often carried out to study their distribution, determine
their trypanosome infection rate or the effectiveness of control measures.

Sampling methods

Both vision and odour detection is used by the tsetse fly to locate its host. These visual and olfactory
stimuli attract tsetse flies respectively at short (<15m) and long range (<100m for the odour of a single
ox). The different sampling methods make use of these stimuli.

Fly-round

Two fly catchers carry a black coloured flag on a tour which is representative for the tsetse fly
habitat. The black screen is often baited with a sachet containing a tsetse fly attractant
(butanone or octenol).

24 | P a g e
Arthropod vectors  Tsetse flies

Fly round Zambia

The fly catchers move along this path, stop at well-defined points and capture during a fixed
period the tsetse flies which are attracted. This method gives by no means an idea of the true
fly density but makes it possible to observe fluctuations in the density of the flies over time. This
method captures much more males than females (notwithstanding female tsetse flies live
longer than males and thus a population will comprise more females than males). With this
method teneral males are often underrepresented and teneral females relatively
overrepresented. Instead of a black flag to attract the flies an ox can be used as a bait to
increase the catch.

Traps

They attract and capture tsetse flies. The colour of the trap is important: blue attracts tsetse
flies while black encourages them to land.

Different tsetse traps

Several types of traps were developed of which the efficiency may vary according to the
tsetse fly species. The biconic trap of Challier Laveissière is often considered as the golden
standard.
25 | P a g e
Arthropod vectors  Tsetse flies

Biconic trap

Performance of the trap will depend on its location, visibility and the use of chemical odours
(CO2, acetone, octenol and phenols) or natural odours (cow urine) which attract tsetse,
especially those of the savannah group. In general teneral flies (especially nulliparous females)
are underrepresented in the samples coming from the traps, but the ratio males/females in the
sample is generally more representative than in case of fly rounds. Most flies captured by a trap
are hungry and have low fat reserves.

The electric screen

The electric screen: It is made up of a metal frame and parallel electric wires between which a
high voltage is applied. Any fly colliding with the screen is electrocuted and falls into the
collecting device. Electric screens are used to evaluate the effectiveness of traps, target
screens, odours etc.

26 | P a g e
Arthropod vectors  Tsetse flies

Electric screen to study tsetse flies

If the tsetse population would be in equilibrium and no immigration or emigration would occur,
an estimate of the real population density may be obtained using mark / release / recapture
experiments. Hereby, a random sample of the population is marked, released and recaptured
at a later date. The population density (X) can then be calculated since

Where Y = the number of all recaptured flies (marked and unmarked), M = number of all
marked flies, R = number of marked flies recaptured.

Marked tsetse fly

27 | P a g e
Arthropod vectors  Tsetse flies

ECOLOGY/LIFE CYCLE
Ecology

Significant interactions between tsetse flies and their environment, especially regarding climate,
vegetation and fauna, have been reported. A comprehensive review on the biology and ecology of
tsetse flies has been recorded by Leak, (1999).

Climate (essentially temperature and vapour pressure deficit) governs the spread of the fly over the
African continent. A humid atmosphere makes it possible for tsetse to move away from protected
habitats where it survives during harsh conditions (e.g. the rainy season will enable G.tachinoides and
G.palpalis gambiensis in the Sudanian savanna to leave the gallery forests and G.pallidipes to leave
the Lambwe Valley National Park in Kenya and reach the surrounding arable land).

Temperatures above 38°C and below 17°C are risky for adult flies. They will seek shelter as soon as
the temperature reaches 35°C (negative phototactism). At these inadequate temperatures the fat
reserve of the pupa becomes also easily exhausted.

The development rate of the various stages of the tsetse fly is directly proportional to the temperature.
High temperatures will shorten the interlarval period, the pupal stage (20 days at 30°C but 100 days at
16°C), the lifespan of the adult (although the correlation with the vapour pressure deficit is better) and
the period between successive feeds.

The vegetation is determined by the climate and soil. Apart from grasslands, which do not support
tsetse flies, all forms of woodland, from savannah to rain forest, can provide a suitable habitat for
some species of these flies, but no single vegetation type is suitable for all species. Some species
(e.g. G.palpalis) adjust easily to artificial biotopes, such as plantations of coffee, cocoa and palm oil.
Thickets (sometimes comprising chiefly Lantana camara), which develop on abandoned agricultural
land, are also often good habitats for them. As the type of vegetation, its photosynthetic activity over
the course of the year (Normalized Difference Vegetation Indices) and the rate of fragmentation of the
natural habitat can easily be monitored by satellite data; these are often used to predict the
distribution and density of a Glossina sp. in an area.

A third important environmental factor for a tsetse fly is the availability of hosts. Both vision and odour
detection are used by the tsetse fly to locate its host, vision is used in short-range recognition (about
15m) while odours are responsible for attracting flies from greater distances. The choice of a host will
depend primarily on the host preference of the fly but, as tsetse flies are opportunistic feeders, host
availability will also play an important role (e.g. in the unspoiled forest G.p.palpalis feeds mainly on
reptiles. However in West and Central Africa the most important densities of this fly are met around
villages where pigs are bred. For tsetse flies of the morsitans group such peridomestic habits are
seldom observed.

28 | P a g e
Arthropod vectors  Tsetse flies

G. f. quanzensis feeding on a pig ear

It was shown that tsetse flies, which have fed on a specific host species, have the tendency to return
to the same host species for their following meals (Bouyer, 2005). Repeated feeding on the same
host species by a disease vector is likely to increase the within-species disease-transmission risk, but
to decrease it between species.

In the laboratory, feeding sterile, warm defibrinated blood through a membrane is generally used to
maintain colonies of tsetse flies.

Life cycle

Tsetse flies have an unusual reproduction method. They are adenotrophic viviparous, the female
feeding the larva inside the uterus and producing one fully grown third instar larva (L3) at each
reproduction cycle. The deposited larva burrows in the soil prior to transform in a pupa. Consequently,
more than half of the total tsetse population lives under the ground as pupae.

29 | P a g e
Arthropod vectors  Tsetse flies

g
h

a
f

e
c

The life cycle of a tsetse fly

a = blood-engorged female e = fly emerging from the puparium

b = larvipositing female f = lateral aspect of a tsetse fly
c = third instar larva g = dorsal aspect of a tsetse fly
d = puparium containing the pupa h = some hosts of the morsitans group of tsetse flies
e = fly emerging from the puparium
f = lateral aspect of a tsetse fly
g = dorsal aspect of a tsetse fly
h = some hosts of the morsitans group of tsetse flies

Ovulation takes place about 4 days after the first meal of the female. The egg passes into the uterus
where it is fertilised by sperma from the spermatheca. After 3 to 4 days the egg hatches and gives
rise to a first stage larva (L1), which is nourished with secretions produced by the milk glands. At 25°C
the first instar lasts 24hrs the second 36hrs and the third 60hrs. The L3 larva once deposited burrows
down into the soil. Then it moults to form the pupa, but remains within the shed third instar cuticle
which hardens to form the puparium. Simultaneously with the larval deposition a new egg ovulates.
The female fly is between 16 and 20 days old at the moment of the first larval deposition. The
subsequent larvipositions generally occur with intervals of 8 to 12 days depending on the species and
ambient temperature (for example G.morsitans at 30°C - 8 days, at 18°C – 25 days). The length of the
pupal period varies according to the sex (shorter for females) and the ambient temperature (on
average 30 to 35 days, but pupal periods from 17 up to 88 days have been observed). The young
adult emerges from the puparium and the soil using its ptilinum. Young flies of which the exocuticle is
not yet hardened and the muscles not well formed, are called teneral flies.

30 | P a g e
Arthropod vectors  Tsetse flies

Third instar larva of a tsetse fly

Puparium of Glossina morsitans

Tsetse flies are relatively long lived, up to 8 to 12 weeks for females, with males having shorter lives
of around 4 to 6 weeks. Around 10 larvae are deposited during the lifetime of a single female. Hence
the rate of reproduction is extremely slow compared to other diptera.

During mating the male with its superior claspers seizes the female. Repeated mating can cause the
female to abort and even die. Therefore, the female generally accepts the male only once and one
insemination is sufficient to fertilize the female throughout its life.

Both sexes of tsetse flies feed on blood, mainly from mammals but also from reptiles and birds. Blood
meals are taken with intervals of 2 to 3 days. However, the bloodmeal does not cover the totality of

31 | P a g e
Arthropod vectors  Tsetse flies

the nutritional needs of the tsetse. Therefore tsetse flies maintain intracellular symbiotic micro-
organisms to supplement their nutrition (Wigglesworthia spp).

Tsetse flies gain energy for flight through the partial breakdown of proline, an amino acid gained from
the blood meal, into alanine. Then alanine is reconverted in proline by using the triglycerides which
are stored in the fat reserve of the fly. Considering the reserve of proline must constantly be renewed,
the tsetse fly is able to fly for short periods only and on the whole less than an hour per day. Flight
speed is around 20 km/h.

Available multimedia

The life cycle of the tsetse fly

http://film.wellcome.ac.uk:15151/mediaplayer.html?0055-0000-3674-0000-
0-0000-0000-0

CONTROL
Tsetse fly control is only one of the trypanosomosis control methods and not always the most
adequate. Unsubsidised, effective and sustainable vector control managed by local communities is
often unrealistic. Situations that justify the use of vector control are:

 a trypanosomosis risk which remains high (e.g. prevalence >10%) notwithstanding regular
chemotherapy or chemoprophylaxis,
 the protection of valuable trypanosomosis-free areas
 when the aim is to eradicate the disease.

Notwithstanding the poor reproductive capacity of the tsetse fly and its unusual high susceptibility to
insecticides, results of past tsetse control campaigns were often unsustainable. Major reasons for
these failures were:

 conflicting objectives between governments, local farmers and donors,

 campaigns aiming control instead of eradication, whereby barriers to stop reinvasion of tsetse
flies from neighboring uncontrolled regions were abandoned because too costly to maintain,
 vector control campaigns not foreseeing participation from the local communities resulting in a
complete stop of the activities once the program ended, and
 insufficient concerted effort among affected countries to control the trans boundary problem of
trypanosomosis.

In those countries, where eradication is aimed for, complete isolation of the area by natural or artificial
barriers as well as political and economic stability is essential in order to reach the objective. The size
of the eradication area has to be carefully evaluated through assessment of habitat suitability, risk
analysis and entomologic surveys (Bouyer, 2010). When isolation of the eradication zone is

32 | P a g e
Arthropod vectors  Tsetse flies

impossible, one can try to eradicate the fly in areas where the environment is the most hostile to the
tsetse fly (at the edge of the fly‟s distribution area).

When, instead of eradication, control of tsetse and/or trypanosomosis is the main objective it is
important to prioritise the areas that are most suitable for an intervention on the basis of agricultural
potential and carrying capacity of the soil, human and livestock densities, socio-economic impact of
the disease as well as environmental considerations.

Tsetse control may be ecological, chemical, biological or genetic.

Ecological control

It consists of the modification of the biotope in order to render the biotope less suitable for the tsetse
fly. Total vegetation clearance (see Bush-clearing tsetse.jpg), partial discriminative vegetation clearing
(mainly used against riverine species), game elimination and game fences, or combinations of these
strategies have all proven to be effective in eliminating tsetse.

Complete bush-clearing to prevent tsetse to enter Ruanda-Urundi

from the Akagera National Park

Since these methods are costly and nowadays unacceptable from an environmental point of view,
they are no longer in use. However, the gradual expansion of the human population has similar
effects since arable land also does constitute an environment unfavourable for tsetse, especially for
those of the fusca and morsitans group. The influence of humans on the palpalis group of flies is not
always adverse, and these flies can often exist in close contact with people and their domestic stock.

Traditional ecological methods such as using smoke and a continuous wooden or netting wall, 1.5 m
high, to protect cattle reduce significantly the numbers of tsetse flies feeding on the animals (Torr,
2011).

33 | P a g e
Arthropod vectors  Tsetse flies

Chemical control

When chemical agents are used to combat tsetse flies, it must be taken into account that tsetse flies
spend about 50% of their lifespan under the ground as pupae. Therefore, either insecticides must be
used, which remain active for at least the maximal pupal period, or repeated treatments with a short-
acting product must be foreseen.

Another issue is the possible reinvasion by flies coming from untreated areas. This requires the
creation of barriers (natural or chemical) between the infested and control zones which constitute an
unbridgeable gap for the tsetse fly. In case of artificial barriers, the larger the control zone surface, the
smaller the proportion of barrier costs in the total costs.

Different methods of chemical tsetse control have been applied:

1. Ground spraying: Method: Preferably during the dry season, a residual insecticide is applied to
the resting and refuge sites of the flies. Spraying is discriminating (only certain types of vegetation
are treated, generally 7 to 15% of the total surface) and selective (only the parts of trees and
bushes are treated where the tsetse flies rest during the day). Persistent insecticides (DDT 3%
2
and dieldrin 2% at 15 to 60 kg a.i./km ) were used for this purpose. Deltamethrin (w.p.) 0.1% (1 to
3 kg/km²) and alpha-cypermethrin are also effective and less harmful to the environment, but
these products are more expensive. Ground spraying may be used to create chemical barriers.
Because of environmental concerns and organizational difficulties this method is nowadays
seldom used.
2. Sequential aerial spraying: This method includes the application of a non-residual insecticide by
airplane (see Aerial spraying plane.jpg) as an ultra-low volume aerosol (droplets of 20 to 50µ)
generated by a rotary atomiser.

Aerial spraying plane Aerial spraying atomizer for ULV

One droplet has to contain sufficient insecticide to kill the least susceptible tsetse fly (some
Glossina spp. and especially pregnant females can tolerate higher insecticidal doses). However
doses used during aerial spraying must be low enough to have no residual action and negligible

34 | P a g e
Arthropod vectors  Tsetse flies

toxicity for non-target organisms. Four to seven treatments are carried out. Timing of the spray
sequence depends on the duration of the first interlarval period and of the puparial period (and
thus environmental temperature). Care must be taken that newly emerged flies are sprayed
before they have the opportunity to deposit their first larva. Conventionally spray cycles are
scheduled two days short of the first interlarval period and stopped only once two sprays
subsequent to the emergence of the last pre–spray pupae (one puparial duration) have been
carried out. The airplane flies at a height of approximately 20 to 30 m above the tree canopies
(therefore a flat terrain is required) and at a distance of 200 to 400 m from the preceding flight
line. To reduce aerosol drift, applications are made under conditions of temperature inversion
early in the morning or late in the evening (1 h before sunset or up to 2 h after sunrise). This
method was shown to be very effective against the morsitans group of tsetse flies in wooded
savannah on flat terrain. Major problems with this method are that, unless the sprayed area is
isolated or other techniques are used in combination, there is nothing to stop flies invading from
beyond the sprayed area. Equally, if there are any survivors left after the completion of the final
treatment, there is no hindrance to the growth of the population.

Insecticides that are used are Endosulfan (e.g. 20% E.C.) at 10 to 20 g a.i./ha and Deltamethrin
0.2 to 0.5 g a.i./ha.

The technique is fast (a surface of 50 km² is treated in 1 h time), environmentally acceptable and
effective but demands a lot of expertise and accurate navigation systems (see: Aerial spraying
instrumentarium.jpg) (Kgori 2009).

Aerial spraying instrumentarium

3. Aerial spraying by helicopter: An insecticide is applied on the nocturnal resting sites of the tsetse
flies. Using this method the most inaccessible sites can be sprayed but the helicopter is the most
expensive way to apply insecticides. Therefore generally persistent insecticides are used, in order
to obtain a residual effect and thus to reduce costs. Harmful effects on terrestrial and water fauna
are to be expected.

35 | P a g e
Arthropod vectors  Tsetse flies

Insecticides that are used are Endosulfan (100 to 1000 g/ha), Dieldrin (800 to 1000 g/ha), and
Deltamethrin (12.5 to 30 g/ha)

4. Insecticide target screens or impregnated traps: This method of reduction of the population of
adult tsetse flies make use of systems to capture flies (traps or screens, which act as traps
because they are treated with "Temo-o-cid" glue) or to kill flies with persistent insecticides applied
to screens or traps.

Target screen spraying

Target screens consist of a blue, black or blue and black cloth screen that has been sprayed with
insecticide. In some versions the cloth is flanked by bands of insecticide treated black mosquito
netting, which is not readily visible to tsetse flies.

Various designs of traps have been developed for use against particular target species in
particular environments.

36 | P a g e
Arthropod vectors  Tsetse flies

Biconical and Monoconical trap

Trap Epsilon and NZI

37 | P a g e
Arthropod vectors  Tsetse flies

Table: Comparison between traps and target screens to control tsetse flies

Traps Target screens

Manufacturing and price more complicated and costly easier to produce and cheaper than traps

Insecticide application not essential essential

even very small screens may have a relatively
Size large
good efficacy (Lindh 2009)
less than half the number of flies attracted by a
Attraction higher than target screen
trap
variable according to species, usually ± 30% if
variable according to species, usually ±50% of
Efficacy trap is untreated, and > 50% if trap is treated
attracted flies alight on screen and are killed
with insecticide
Vulnerability high low

Efficacy visualised yes (visible capture) no (flies die not immediately)

Impact on non-target organisms low to very low if untreated low

Knowledge of tsetse species
essential to ensure good results an asset
behaviour

The major advantages of screens and traps are that:

 the techniques are rather simple and adapted to the requirements of community participation,
 it is relatively cheap, easy to produce and with almost no pollution of the environment,
 it allows the spreading of the operation over time and is easily combined with other methods,
 it allows some errors without causing the failure of the entire operation but eradication is
difficult to attain,
 the efficacy of the technique especially against flies of the group morsitans can substantially
be improved using attractive odours,
 it can be used to produce barriers to prevent reinvasion of tsetse flies.

Target screen barrier

38 | P a g e
Arthropod vectors  Tsetse flies

The major disadvantages of screens and traps are:

 its effectiveness, which is variable according to the species (e.g. G.pallidipes is 4 times more
susceptible to a control by targetscreens that G.m.morsitans which generally requires 4
2
screens/km ). Far more traps and screens are necessary against riverine tsetse than against
savannah species as riverine species are less attracted to odours,
 the larger the control area the better the results but the more difficult the management of traps
and screens,
 sometimes many access roads have to be created (pernicious in game reserves),
 maintenance of the traps and screens is essential if the control has to be continued.
However, community participation fades once control has diminished disease impact,
 with the current knowledge these methods do not always lead to eradication,
 the type of landscape which does not always allow the use of the screens or traps,
 problems of theft (necessity to inform the community) and of degradation of the materials
(e.g. fires, wildlife, swelling of rivers).

Insecticides: pyrethroids (deltamethrin, alpha-cypermethrin, lambda-cyhalotrin) are usually used

to spray screens or traps. Immersion of the screens in an insecticidal solution for 15 minutes and
afterwards drying them on a horizontal surface is preferred to simple spraying. Glossinex ®
(deltamethrin 19% + UV-absorber) is used at concentrations up to 1g deltamethrin/m² in order to
reduce the number of sprayings (once a year) and avoid maintenance. The frequency, with which
the insecticidal spray on the screen must be renewed, depends on the insecticide (type,
formulation and dose) but also on the quality of the fabric used to manufacture the trap or screen
and on the season.

The chitin synthesis inhibitor “triflumuron” (Langley, 1995) and the macrocyclic lactone "spinosad"
(De Deken et al., 2004) are alternative insecticides should it become undesirable to use
pyrethroids.

Attractants: Kairomones that are used in the field to attract tsetse flies to traps or targets are
acetone, methyl ethyl ketone (butanone), 1-octen-3-ol (octenol), 4-methylphenol, 3-n-
propylphenol and 3-methylphenol. Not all of these substances are effective against all species
and not all need to be included in a bait for it to be effective (Vale et al; 1988; Vale & Torr, 2004;
Rayaisse et al, 2010).

The way in which tsetse flies move through the habitat has an important bearing on where traps
and targets should be positioned so as to be most effective (Vale, 1998; Kuzoe & Schofield, 2004;
Bouyer et al, 2010).

Density of baits: The use of olfactory products makes it possible to reduce considerably the
density of the baits for the control of tsetse flies of the morsitans group. A density of 2 to3
screens/km² for G.pallidipes and 4 to 5 screens/km² for G.morsitans is sufficient to control these
flies. In Burkina Faso 33 biconical traps or 25 blue-colored screens/km² were used against
G.m.submorsitans, but by adding olfactory products the density could be reduced to 5 traps or
screens/km².

39 | P a g e
Arthropod vectors  Tsetse flies

For riverine tsetse flies (e.g. G.p.gambiensis and G.tachinoides) the screens (or traps) are posted
along the river banks and in isolated forest galleries at a rate of 10 to 12 screens/km or 3 to 10
traps per km. It is often necessary to withdraw the traps or screens during the rainy season in
order to prevent that they are carried away by the flood.

However, if the biotope of flies belonging to the palpalis or fusca groups is not strictly riparian,
screens and traps have to be used at much higher densities, since these flies are poorly attracted
by odours and some species in this group (e.g. G.fuscipes) are not very mobile. In the human
trypanosomosis focus of Vavoua 180 blue screens/km² were used to control G.palpalis in its
biotope (primarily cocoa and coffee plantations). The screen density could be decreased to 32
screens/km² by using screens with blue and black bands side by side. In Uganda to control
G.f.fuscipes 8 to 10 pyramidal traps/km² were set up at the transition between forest and other
biotopes (Lancien, 1993).

Creation of barriers using target screens: The barrier surrounding the control area to prevent
invasion of tsetse flies must be created before the start of the control campaign (for example
control by aerial spraying). Broad barriers (4 to 8 km), having a width of up to 8 times the distance
covered by the tsetse species in 1 day and made up of a number of screens or traps/km² equal to
or twice the usual density, are to be preferred on small barriers with higher screen densities
(Hargrove, 1993).

5. Insecticide treated hosts: In livestock breeding areas tsetse flies can be controlled by dipping the
cattle regularly (e.g. every two weeks) in a solution of 0, 0038% to 0,005% of deltamethrin
(Decatix 5% SC), applying up to 20 times/year a pour-on insecticide (Cylence 1%, SpotOn 1%,
Renegade 1, 5%, Bayticol 1%), or spraying the animals every two weeks with 0,005% of
deltamethrin (Decatix) using a portable compressed air sprayer. Such treatment frequencies are
too expensive for most African cattle owners. Treatment frequency may be decreased when the
area is not subjected to invasion.

However, since 75 - 90% of tsetse flies feed on the legs or belly of cattle, spraying may be
restricted to these body parts of all animals in the herd or only of the larger animals in the herd
(Torr, et al, 2007). This strategy reduces insecticide costs by 40 to 80%, with only a 20% to 30%
reduction in efficacy and makes restricted application affordable for poor livestock owners. These
treatments will in most situations have no or little impact on endemic stability against local tick-
borne diseases. Treatment of only a part of the herd is cost beneficial as the percentage of tsetse
fly feeds from an animal is correlated to its live weight. However, depending on the region and its
local customs the benefits of this particular strategy may vary heavily as Masaai graze their adult
cattle often separately from the calves while small stock of the Shona in Zimbabwe graze all their
livestock together.

If the application of insecticides on cattle is used as the sole method to control the vector, the
zone, in which the cattle have to be treated simultaneously, must be significant. The effectiveness
of the method can be reduced in case of alternative hosts, such as an abundant wild fauna and
village herds not treated simultaneously, or when the treated cattle do not penetrate all the
infested tsetse biotopes (Van den Bossche et al, 2004). In absence of behavioral resistance the

40 | P a g e
Arthropod vectors  Tsetse flies

applied insecticide has no influence on the number of tsetse flies attracted towards the host or on
the percentage of tsetse flies taking a meal on the treated animal. Thus insecticide treated
animals are not completely protected from trypanosomosis.

Through the years chemical control of tsetse changed gradually from area-wide application of
insecticides towards a more targeted control using insecticide treated hosts, traps or target
screens. These devices attract and kill tsetse with limited environmental impact and allow
participation of local communities in the control. However, it is still not sure that these devices
may eradicate all tsetse flies. Especially in case of riverine species the “Sterile Insect Technique”
may be required to obtain complete eradication after reduction of the tsetse population by
insecticides.

Biological control

Predators: Many animals (e.g. lizards, snakes, birds, bats, mongoose, rodents, insects, spiders, ants)
predate on tsetse flies but it is very difficult to evaluate their impact.

Parasitoids: some deposit their eggs in tsetse fly pupae, but usually the percentage of parasitized
pupae fluctuates strongly according to the time of the year.

Parasitoids belonging to the Hymenoptera

- Nesolynx spp. (Syntomosphyrum spp.) do not attack the pupae of tsetse flies exclusively.
Establishment of breeding colonies of this parasite is possible, but significant releases in
Eastern Africa were not successful while this insect is not able to dig the ground while
searching for the pupae of tsetse flies.
- Chrestomutilla spp. (Mutilla spp.) are able to reproduce in a parthenogenetic way, the not-
fertilized eggs giving rise to males. The female lays 2 to 3 eggs in the same pupa, but only
one adult parasitoid will emerge. The development of the immature stages takes
approximately a month and half. Breeding of this insect in the laboratory has not been
successful.

Parasitoids belonging to the Diptera

- Twelve different species of Exhyalanthrax (Thyridanthrax) were found to parasitize tsetse flies
as well as other flies; 10 in Eastern Africa and 2 in Western Africa. The cycle from egg to
adult generally lasts 20 to 35 days, but sometimes there is a kind of hibernation of the larvae
so that the adults hatch only after several months (up to 14 month). The breeding of these
diptera could not been achieved.

Pathogens: a microbe (Bacterium mathisi) several fungi and a Baculovirus (salivary gland hypertrophy
virus) are pathogenic for tsetse flies but none of these pathogens can be used as a biological control
agent yet.

41 | P a g e
Arthropod vectors  Tsetse flies

Genetic control

This method of control aims to alter the reproductive potential of the vector or its vectorial
competence.

The sterile insect technique via irradiation: This method considers primarily the eradication of isolated
populations of tsetse flies. It involves the breeding of thousands of tsetse flies of each subspecies
present in the area and the male tsetse are then sterilised using gamma (or X-) radiation and released
at regular intervals, thus swamping the population with males that are unable to fertilise females
successfully. Glossina austeni has been successfully eradicated from Zanzibar using this method
(Vreysen et al., 2000). During this campaign 5.5 million sterile males were released in total. Before
flooding the area with the sterile males the local wild tsetse population must be largely reduced by
other control methods. It is estimated that a proportion of 10 to 30 sterile males per fertile male is
needed in order to assure that most couplings with the few remaining wild females will occur with
sterile males and, since most female tsetse fly does copulate only once with a male, the population
will be eradicated. Sterilisation by irradiation is possible for adult male or female tsetse as well as for
pupae. Before being released the sterile males are nourished with blood from animals treated with
trypanocides in order to decrease as such as possible the chances of transmission by the sterile flies.
For riverine tsetse fly species it seems easier for the sterilised, released male to find wild flies of the
opposite sex than for savannah species.

The sterile insect technique via chemical sterilisation: Mass breeding followed by gamma ray
sterilization of males could possibly be replaced by the direct sterilization of the males or/and females
in the field thanks to systems of traps or screens allowing a sufficient contact between the insect and
a sterilizing chemical product (e.g. bisazir, tepa, hempa). As these products are relatively dangerous
and toxic to mammals, they may be substituted by analogues of the juvenile hormone (e.g.
pyriproxyfen: (Langley et al., 1994). An amount of 0.02 µg of pyriproxyfen applied on a tsetse fly
female inhibits any hatching of pupae by this female. Pyriproxyfen being relatively stable under
tropical conditions, screens treated with a mixture of oil (Cerechlor S45) and of pyriproxyfen 2 mg/cm²
were efficacious for a period of 9 months. This juvenile hormone mimics the chitin synthesis inhibitor,
triflumuron, and may provide a safe way to effectively autosterilize female tsetse flies. These methods
may eliminate the need for the costly artificially maintained tsetse fly colonies.

Genetic control via transgenic tsetse fly symbionts: Current research on genetic control of tsetse flies
involves the development of transgenic symbionts of the tsetse fly (Aksoy, 2005). The 3 symbionts of
the tsetse fly (Sodalis glossinidius, Wigglesworthia glossinidium and a strain of Wolbachia) are all
maternally transmitted to progeny. The research aims to alter the genome of the endosymbiont,
Sodalis glossinidius, so that the symbiont expresses trypanolytic substances into the fly. Wolbachia
will be used to introduce cytoplasmic incompatibility in the natural tsetse population. Cytoplasmic
incompatibility is achieved when wild-type females are mated with males infected by a Wolbachia
strain that is non-existent in the female. The intracellular Wolbachia will then cause embryonic
mortality. When both methods are combined it must, in theory, be possible to replace the wild-type,
trypanosome-susceptible tsetse population with the engineered, trypanosome-refractory line.

42 | P a g e
Arthropod vectors  Tsetse flies

PATTEC

Recently the Organisation of African Unity decided to launch the Pan-African Tsetse and
Trypanosomosis Eradication Campaign (PATTEC). PATTEC promotes an implementation policy
designed to select feasible intervention areas, suppress tsetse populations using any appropriate
combination of trap and target deployment, insecticide-treated cattle, and/or Sequential Aerial
Technique, to be followed by the Sterile Insect Technique if necessary for definitive elimination of the
target population. The first phase of this project attempts to free isolated zones or zones close to the
limit of the current fly distribution. Parts of Burkina Faso, Ghana and Mali in West Africa and parts of
Kenya, Ethiopia and Uganda in East Africa will be concerned. Whether or not it will be possible to
eradicate tsetse from Africa is the subject of much debate. Some are sceptical about the chances of
success of such a campaign due to the extent, complexity and trans boundary nature of the problem.
Furthermore, many environmentalists are concerned about the indirect impact tsetse fly control may
have on the environment. They fear that eradicating tsetse may open the path to unsustainable
pastoral encroachment of national parks and game reserves and are overtly opposed to any
eradication attempt. It is correct that many tsetse control campaigns are carried out because of
scarcity of arable land or pasture and in these situations it may be difficult to prevent movement of
people into tsetse-cleared area, if the future use of the tsetse cleared land is not well planned in
advance and implemented afterwards. Otherwise, the benefit of control campaigns can easily be lost
by the damage inflicted to the ecological system. Therefore, simultaneously with the planning of the
control campaigns, future land-use must be laid down in consultation with potential users, senior civil
servants and experts in environmental conservation.

Now that vector control in the framework of PATTEC has started, it is vital that not only the
entomological efficacy of the different control techniques is examined, but also their relative cost.
Such cost estimations are not simple to be determined as was demonstrated by Shaw et al., 2007 &
2009 and most African countries haven‟t the resources to eliminate flies within their borders.

Available multimedia

Available multimedia

Research on tsetse fly control (Wellcome Foundation):

http://film.wellcome.ac.uk:15151/mediaplayer.html?0055-0000-3886-0000-
0-0000-0000-0

43 | P a g e
Arthropod vectors  Tsetse flies

REFERENCES
1. Akoda, K., Van den Bossche, P., Marcotty, T., Kubi, C., Coosemans, M., De Deken, R. & Van
den Abbeele, J. (2009) Nutritional stress affects the tsetse fly's immune gene expression, Medical
and Veterinary Entomology 23 (3), 195-201.

2. Aksoy S., Rio R.V.M. (2005) Interactions among multiple genomes: Tsetse, its symbionts and
trypanosomes, Insect Biochemistry and Molecular Biology 35, 7, 691-698.

3. Bouyer J., Cuisance D., Messad S., Guerin P., Learning affects host preference in tsetse flies,
Rev Elev Med Vet Pays Trop 58, 27-29.

4. Bouyer J., Seck M.T., Sall B., Ndiaye E.Y., Guerrini L., Vreysen M.J.B., (2010) Stratified
entomological sampling in preparation for an area-wide integrated pest management program:
The example of Glossina palpalis gambiensis (Diptera: Glossinidae) in the Niayes of Senegal,
Journal of Medical Entomology 47, 4, 543-552.

5. Bruce D., Hamerton A.E., Bateman H.R., Mackie F.P. (1909) The development of Trypanosoma
gambiense in Glossina palpalis. Proceedings of the Royal Society (B) 81, 405–414.

6. Brunhes J., Cuisance D., Geoffroy B., Hervy J.P. (1998) “Glossina or the tsetse fly: Software for
identification and training”. Editions ORSTOM, Montpellier, France.

7. Challier A. (1965) Amélioration de la méthode de détermination de l'âge physiologique des

glossines. Etudes faites sur Glossina palpalis gambiensis Vanderplank. Bull. Soc. Path. exot. 58,
250-259.

8. De Deken R., Speybroeck N., Gillain G., Sigue H., Batawi K., Van den Bossche P. (2004) The
macrocyclic lactone 'spinosad,' a promising insecticide for tsetse fly control. Journal of Medical
Entomology 41, 5, 814-818.

9. Gikonyo N.K., Hassanali A., Njagi P.G.N., Gitu P.M., Midiwo J.O. (2002) Odour composition of
preferred (buffalo and ox) and nonpreferred (waterbuck) hosts of some savanna tsetse flies, J
Chem Ecol 28, 969-981.

10. Hargrove J.W. (1993) Target barriers for tsetse flies (Glossina spp.) (Diptera Glossinidae): quick
estimates of optimal target densities and barrier widths. Bulletin of Entomological Research 83,
197–200.

11. Kgori P.M., Orsmond G. & Phillemon-Motsu T.K. (2009) Integrating GIS and GPS-assisted
navigation systems to enhance the execution of an SAT -based tsetse elimination project in the
Okavango delta (Botswana)In “Geospatial datasets and analyses for an environmental approach
to African trypanosomiasis”, editors Cecchi G. & Mattioli R.C., PAAT Technical and Scientific
Series 9, FAO, p. 61-67.

44 | P a g e
Arthropod vectors  Tsetse flies

12. Kristjanson P.M., Swallow B.M., Rowlands G.J., Kruska R.L., Deleeuw P.N. (1999) Measuring the
costs of African animal trypanosomosis, the potential benefits of control and returns to research.
Agr Syst 59, 79-98.

13. Kuzoe F.A.S., Schofield C.J. (2004) Strategic review of traps and targets for tsetse and African
trypanosomiasis control, http://apps.who.int/tdr/publications/tdr-research-publications/strategic-
review-traps-targets-tsetse/pdf/tsetse_traps.pdf.

14. Langley P.A., Hargrove J.W., Mauchamp B., Royer C., Oouchi H. (1994) Prospects for using
pyriproxyfen treated targets for tsetse control. Entomologia Experimentalis Applicata 153, 159-
166.

15. Langley P.A. (1995) Evaluation of the chitin synthesis inhibitor triflumuron for controlling the tsetse
Glossina morsitans morsitans (Diptera : Glossinidae), Bulletin of Entomological Research 85,
495-500.

16. Leak S.G.A. (1999) Tsetse biology and ecology: Their role in the epidemiology and control of
trypanosomosis. Oxford: CABI Publishing. 568 pp.

17. Lehane M. J., Aksoy S. & Levashina E. (2004) Immune responses and parasite transmission in
blood-feeding insects. Trends Parasitol 20, 9, 433-439.

18. Lindh J.M., Torr S.J., Vale G.A. & Lehane M.J. (2009) Improving the cost-effectiveness of artificial
visual baits for controlling the tsetse fly Glossina fuscipes fuscipes, Plos Negl Dis 3, 7, e474. doi:
10.1371/journal.pntd.0000474.

19. Livesey J.L., Molyneux D.H., Jenni L. (1980) Mechanoreceptor-trypanosome interactions in the
labrum of Glossina: fluid mechanics. Acta Tropica 37, 151–161.

20. Macdonald G. (1957) The epidemiology and control of malaria. Oxford: Oxford University Press.
201 pp.

21. Macleod E.T., Maudlin I., Darby A.C. & Welburn S.C. (2007) Antioxidants promote establishment
of trypanosome infections in tsetse. Parasitology 134, 6, 827-831.

22. Rayaisse J.B., Tirados I., Kaba D., Dewhirst S. Y., Logan J. G., Diarrassouba A., Salou E.,

23. Omolo M., Solano P., Lehane M.J., Pickett J. A., Vale G.A., Torr S.J. and J. Esterhuizen (2010)
Prospects for the development of odour baits to control the tsetse flies Glossina tachinoides and
G. palpalis s.l. PLoS Negl Dis 4, 3: e632. doi:10.1371/journal.pntd.0000632

24. Saunders, D.S. (1960) The ovulation cycle in Glossina morsitans Westwood (Diptera: Muscidae)
and a possible method of age determination for female tsetse flies by examination of their ovaries.
Transactions of the Royal Entomological Society of London 112, 221-238.

25. Shaw A, Torr S, Waiswa C, Robinson T, (2007) Comparable Costings of Alternatives for Dealing
with Tsetse: Estimates for Uganda, Working Paper of the Pro-Poor Livestock Policy Facility,
http://www.fao.org/ag/againfo/programmes/en/pplpi/docarc/wp40.pdf
45 | P a g e
Arthropod vectors  Tsetse flies

26. Shaw, A (2009) Assessing the economics of animal trypanosomosis in Africa-history and current
perspectives, Onderstepoort Journal of Veterinary Research 76, 27-32.

27. Smith D.L., McKenzie F.E., Snow R.W. & Hay, S.I. (2007) Revisiting the basic reproduction
number for malaria and its implications for malaria control. PLOS Biol 5, 3: e42.
doi:10.1371/journal.pbio.0050042.

28. Torr S.J., Maudlin I., Vale G.A. (2007) Less is more: restricted application of insecticide to cattle
to improve the cost and efficacy of tsetse control. Med Vet Entomol 21: 53–64.

29. Torr S.J., Mangwiro T.N.C., Hall D.R. (2011) Shoo fly, don't bother me! Efficacy of traditional
methods of protecting cattle from tsetse, Medical and Veterinary Entomology 25, 2, 192-201.

30. Vale G.A. & Hall D.R. (1985) The role of 1- octen-3-ol, acetone and carbon dioxide in the
attraction of tsetse flies, Glossina spp. (Diptera: Glossinidae) to ox odour. Bulletin of
Entomological Research, 75, 209–217.

31. Vale G.A., Hall D.R & Gough A.J.E. (1988) The olfactory responses of tsetse flies Glossina spp.
(Diptera: Glossinidae) to phenols and urine in the field. Bulletin of Entomological Research 78,
293–300.

32. Vale G.A. & Torr S.J. (2004) Development of bait technology to control tsetse. In: Maudlin I,
Holmes PH, Miles MA, editors. The trypanosomiases. Wallingford: CABI Publishing; 2004. p. 509-
523.

33. Van Den Abbeele J., Caljon G., De Ridder K., De Baetselier P., Coosemans M. (2010)
Trypanosoma brucei modifies the tsetse salivary composition, altering the fly feeding behaviour
that favours parasite transmission. PLoS Pathog 2010; 6 (6): e1000926.

34. Van den Bossche, P. & De Deken R. (2004) The application of bait technology to control tsetse
In: Maudlin I, Holmes PH, Miles MA, editors. The trypanosomiases. Wallingford: CABI Publishing;
2004. p. 525-532.

35. Vreysen M.J.B., Saleh K.M., Ali M.Y., Abdulla A.M., Zhu Z.R., Juma K. G., Dyck V.A., Msangi
A.R., Mkonyi P.A., Feldmann H.U. (2000) Glossina austeni (Diptera: Glossinidae) eradicated on
the Island of Unguja, Zanzibar, using the sterile insect technique, Journal of Economic
Entomology 93, 123-135.

46 | P a g e