Tsetse fly

Tsetse (/ˈsiːtsi/ SEET-see,

US: /ˈtsiːtsi/ TSEET-see or
UK: /ˈtsɛtsǝ/ TSET-sǝ)
(sometimes spelled tzetze; also
known as tik-tik flies) are large,
biting flies that inhabit much of
tropical Africa.[1][2][3] Tsetse flies
include all the species in the
genus Glossina, which are

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 placed in their own family,
Glossinidae. The tsetse is an
obligate parasite, which lives by
feeding on the blood of
vertebrate animals. Tsetse has
been extensively studied
because of their role in
transmitting disease. They have a
pronounced economic impact in
sub-Saharan Africa as the
biological vectors of
trypanosomes, causing human
and animal trypanosomiasis.

Tsetse fly

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 Temporal range: Eocene - Recent

Glossina morsitans

Scientific classification

Domain: Eukaryota

Kingdom: Animalia

Phylum: Arthropoda

Class: Insecta

Order: Diptera

(unranked): Eremoneura

(unranked): Cyclorrhapha
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 (unranked): Cyclorrhapha

Section: Schizophora

Subsection: Calyptratae

Superfamily: Hippoboscoidea

Family: Glossinidae
Theobald,
1903

Genus: Glossina
Wiedemann,
1830

Species groups

Morsitans ("savannah"
subgenus)

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 Fusca ("forest" subgenus)
Palpalis ("riverine"
subgenus)

Range of the tsetse fly

Tsetse can be distinguished from

other large flies by two easily-
observed features: Primarily,
tsetse fold their wings over their
abdomens completely when they
are resting (so that one wing
rests directly on top of the other);

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 Secondly, tsetse also have a long
proboscis, extending directly
forward, which is attached by a
distinct bulb to the bottom of
their heads.

Fossilized tsetse has been

recovered from Paleogene-aged
rocks in the United States and
Germany. Twenty-three extant
species of tsetse flies are known
from the African continent as well
as the Arabian Peninsula.

Etymology
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 The word tsetse means "fly" in
Tswana, a Bantu language of
southern Africa.[4] As "tsetse fly"
is a pleonasm, (meaning, literally,
"fly fly"), recently, tsetse without
the "fly" has become more
common in English, particularly in
the scientific and development
communities.

The word is pronounced tseh-

tseh in the Sotho languages and
is easily rendered in other African
languages. During World War II, a
British de Havilland

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 antisubmarine aircraft was
known as the 'Tsetse' Mosquito.
[5]

Biology
The biology of tsetse is relatively
well understood by
entomologists. They have been
extensively studied because of
their medical, veterinary, and
economic importance, because
the flies can be raised in a
laboratory, and because they are
relatively large, facilitating their

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 analysis.

Morphology

Tsetse flies can be seen as

independent individuals in three
forms: as third-instar larvae,
pupae, and adults.

Tsetse first becomes separate

from their mothers during the
third larval instar, during which
they have the typical appearance
of maggots. However, this life
stage is short, lasting at most a

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 few hours, and is almost never
observed outside of the
laboratory.

Tsetse next develops a hard

external case, the puparium, and
become pupae - small, hard-
shelled oblongs with two
distinctively small, dark lobes at
the tail (breathing) end. Tsetse
pupae are under 1 centimetre
(1⁄2 in) long.[6] Within the puparial
shell, tsetse complete the last
two larval instars and the pupal
stage.

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 At the end of the pupal stage,
tsetse emerges as adult flies. The
adults are relatively large flies,
with lengths of 0.5–1.5
centimetres (1⁄4–5⁄8 in),[6] and
have a recognizable shape, or
bauplan, which makes them easy
to distinguish from other flies.
Tsetse have large heads,
distinctly separated eyes, and
unusual antennae. The thorax is
quite large, while the abdomen is
wider, rather than elongated, and
shorter than the wings.

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 Four characteristics collectively
separate adult tsetse from other
kinds of flies:

Tsetse have a distinct proboscis, a long and

Proboscis thin structure attached to the bottom of the
head, pointing forward.

Folded When at rest, tsetse fold their wings

wings completely, one-on-top of the other.

The discal medial ("middle") cell of the wing

Hatchet
has a characteristic hatchet shape, resembling
cell
a meat cleaver or a hatchet.

Branched
The antennae have arista with hairs which are,
arista
themselves, branched.
hairs

Anatomy

Like all other insects, tsetse flies

have an adult body comprising
three visibly distinct parts: the

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 head, the thorax, and the
abdomen.

The head has large eyes,

distinctly separated on each side,
and a distinct, forward-pointing
proboscis attached underneath
by a large bulb. The thorax is
large, made of three fused
segments. Three pairs of legs are
attached to the thorax, as are two
wings and two halteres. The
abdomen is short but wide and
changes dramatically in volume
during feeding.

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 Reproductive anatomy sketch by
[[es:User:Estefanía Alonso
Gómez]]

The internal anatomy of the

tsetse is fairly typical of the
insects; the crop is large enough
to accommodate a huge increase
in size during feeding, as tsetse
can take a blood meal equal in
weight to themselves. The
dipteran crop is heavily
understudied, with Glossina
being one of the few genera

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 having relatively reliable
information available: Moloo and
Kutuza 1970 for G. brevipalpis
(including its innervation) and
Langley 1965 for G. morsitans.[7]
The reproductive tract of adult
females includes a uterus, which
can become large enough to hold
the third-instar larva at the end of
each pregnancy. The article
"Parasitic flies of domestic
animals" has a diagram of the
anatomy of dipteran flies.

Most tsetse flies are, physically,

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 very tough. Houseflies, and even
horseflies, are easily killed with a
flyswatter, for example; a great
deal of effort is needed to crush a
tsetse fly.[8]

Life cycle

Glossina palpalis and G. morsitans from a 1920

lexicon

Tsetse has an unusual life cycle,

which may be due to the richness

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 of their blood food source. A
female fertilizes only one egg at a
time; she will retain each egg
within her uterus, the offspring
developing internally (during the
first three larval stages), in an
adaptation called adenotrophic
viviparity.[9] During this time, the
female feeds the developing
offspring with a milky substance
(secreted by a modified gland) in
the uterus.[10] In the third larval
stage, the tsetse larvae leave the
uterus and begin an independent

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 life. The newly-birthed larvae
crawl into the ground and
develop a hard outer shell (called
the puparial case), within which
they complete their
morphological transformations
into adult flies.

The larval life stage has a variable

duration, generally 20 to 30 days,
and the larvae must rely on
stored resources during this time.
The importance of the richness
and quality of blood to this stage
can be seen; all tsetse

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 development (prior to emerging
from the puparial case as a full
adult) occurs without feeding,
with only the nutrition provided
by the mother fly. She must get
enough energy for her own
survival (in addition to the needs
of her developing offspring), as
well as for the stored resources
that her offspring will require until
they emerge as adults.

Technically, these insects

undergo the standard
development process of insects,

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 beginning with oocyte formation,
ovulation, fertilization, and
development of the egg;
following egg development and
birth is the three larval stages, a
pupal stage, and the emergence
and maturation of the adult.

Hosts

Overall Suidae are the most

important hosts. By species,
bloodmeals are derived from:[11]

G. swynnertoni — 60–70%

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 from a warthog, ~8% from
giraffe
G. austeni — 50–60% from
bushpig, ~33% from Bovidae,
possibly 10% from various
duiker
G. fuscipleuris — 65% from
bushpig and giant forest hog,
up to 20% from hippopotamus
G. tabaniformis — 70% from
red river hog, >7% from
porcupines
G. morsitans — 30–45% from
warthog, 25–40% from various

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 Bovidae, especially kudu,
buffalo, bushbuck, and eland,
most especially domestic
cattle, ~2% from hartebeest
G. fusca — 55–90% from
bushbuck, 15% from red river
hog, ~12% from aardvark
G. brevipalpis — up to 40%
(high variability with
geography) from bushpig, up
to 36% from hippopotamus,
~25% from Bovidae, especially
buffalo and bushbuck
G. palpalis — ~3% from wild

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 Suidae, more substantial
amounts from domestic
Suidae when available, ~20–
40% from Bovidae (including
domestic cattle) depending on
geography, ~10% from
waterside birds including
cormorants, 25–30% from
Varanus and crocodile
(possibly higher in natural
settings, 50% from crocodile in
particular locations)
G. fuscipes — ~3% from wild
Suidae, ~20–40% from

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 Bovidae (including domestic
cattle) depending on
geography, ~10% from
waterside birds including
cormorants, 25–30% from
Varanus and crocodile
(possibly higher in natural
settings)
G. tachinoides — ~3% from
wild Suidae, more substantial
amounts from domestic
Suidae when available, ~20–
40% from Bovidae (including
domestic cattle) depending on

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 geography, >7% from
porcupines
G. pallidipes — 55–90% from
bushbuck
G. longipalpis — 55–90% from
bushbuck
G. longipennis — unusually
dependant (~60%) on
rhinoceros, also ~20% from
Bovidae, variably up to 12%
from elephant, up to 7% from
ostrich
G. m. submorsitans — ~6%
from various birds excluding

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 ostrich

Waterbuck (Kobus
ellipsiprymnus) are unmolested
by Glossina[11][12] because they
produce volatiles which act as
repellents. Waterbuck odor
volatiles are under testing and
development as repellents to
protect livestock.[13][14]: Suppl T1

Genetics

The genome of Glossina

morsitans was sequenced in

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 2014.[15]

Symbionts

Tsetse flies have at least three

known bacterial symbionts. The
primary symbiont is
Wigglesworthia (Wigglesworthia
glossinidia) within the fly's
bacteriocytes, the secondary
symbiont is Sodalis (Sodalis
glossinidius) intercellularly or
intracellularly, and the third is
some kind of Wolbachia.[16][17]

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 Diseases

The salivary gland hypertrophy

virus causes abnormal bleeding
in the lobes of the crop of G. m.
centralis and G. m. morsitans.[7]

Systematics
Tsetse are in the order Diptera,
the true flies. They belong to the
superfamily Hippoboscoidea, in
which the tsetse's family, the
Glossinidae, is one of four
families of blood-feeding

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 obligate parasites.

Up to 34 species and subspecies

of tsetse flies are recognized,
depending on the particular
classification used.

All current classifications place

all the tsetse species in a single
genus named Glossina. Most
classifications place this genus
as the sole member of the family
Glossinidae. The Glossinidae are
generally placed within the
superfamily Hippoboscoidea,

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 which contains other
hematophagous families.

Species

The tsetse genus is generally

split into three groups of species
based on a combination of
distributional, behavioral,
molecular and morphological
characteristics.[18] The genus
includes:

The 'savannah' flies:

(subgenus Morsitans,

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 occasionally named Glossina):
Glossina austeni
(Newstead, 1912) patr. of
Austen
Glossina morsitans
(Westwood, 1851)
Glossina morsitans
submorsitans[19]
Glossina pallidipes
(Austen, 1903)
Glossina swynnertoni
(Austen, 1922)[20][21] patr.
of Swynnerton[21]

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 The 'forest' flies: (subgenus
Fusca, previously named
Austenia):
Glossina fusca fusca
(Walker, 1849)
Glossina fuscipleuris
(Austen, 1911)
Glossina frezili (Gouteux,
1987)[22]
Glossina haningtoni
(Newstead and Evans,
1922)
Glossina longipennis
(Corti, 1895)

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 Glossina medicorum
(Austen, 1911)
Glossina nashi (Potts,
1955)
Glossina nigrofusca
nigrofusca (Newstead,
1911)
Glossina severini
(Newstead, 1913)
Glossina schwetzi
(Newstead and Evans,
1921)
Glossina tabaniformis
(Westwood, 1850)

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 Glossina vanhoofi
(Henrard, 1952)
The 'riverine' and 'lacustrine'
flies: (subgenus Palpalis,
previously named Nemorhina):
Glossina caliginea
(Austen, 1911)
Glossina fuscipes
(Newstead, 1911)
Glossina fuscipes
fuscipes (Newstead,
1911)[19]
Glossina fuscipes
martinii (Zumpt,

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 1935)
Glossina fuscipes
quanzensis (Pires,
1948)
Glossina pallicera
pallicera (Bigot, 1891)
Glossina pallicera
newsteadi (Austen, 1929)
patr. of Newstead
Glossina palpalis palpalis
(Robineau-Desvoidy,
1830)
Glossina palpalis
gambiensis (Vanderplank,

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 1911)
Glossina tachinoides
(Westwood, 1850)

Evolutionary history

Fossil glossinids are known from

the Florissant Formation in North
America and the Enspel
Lagerstätte of Germany, dating to
the late Eocene and late
Oligocene respectively.[23]

Range

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 Glossina is almost entirely
restricted to grassland and
forested areas of the Afrotropics.
Only two subspecies - G. f.
fuscipes and G. m. submorsitans
- are definitely present in the very
southwest of Saudi Arabia.
Although Carter found G.
tachiniodes in 1903 nearby, near
Aden in southern Yemen, there
have been no confirmations
since.[19]

Trypanosomiasis

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 Trypanosomes in a blood smear

Tsetse are biological vectors of

trypanosomes, meaning that in
the process of feeding, they
acquire and then transmit small,
single-celled trypanosomes from
infected vertebrate hosts to
uninfected animals. Some tsetse-
transmitted trypanosome
species cause trypanosomiasis,
an infectious disease. In humans,
tsetse transmitted

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 trypanosomiasis is called
sleeping sickness.[24] In animals,
tsetse-vectored
trypanosomiases include nagana,
souma (a French term which may
not be a distinct condition[25]),
and surra according to the animal
infected and the trypanosome
species involved. The usage is
not strict and while nagana
generally refers to the disease in
cattle and horses it is commonly
used for any of the animal
trypanosomiases.

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 Trypanosomes are animal
parasites, specifically protozoans
of the genus Trypanosoma.
These organisms are about the
size of red blood cells. Different
species of trypanosomes infect
different hosts. They range
widely in their effects on the
vertebrate hosts. Some species,
such as T. theileri, do not seem to
cause any health problems
except perhaps in animals that
are already sick.[26]

Some strains are much more

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 virulent. Infected flies have an
altered salivary composition
which lowers feeding efficiency
and consequently increases the
feeding time, promoting
trypanosome transmission to the
vertebrate host.[27] These
trypanosomes are highly evolved
and have developed a life cycle
that requires periods in both the
vertebrate and tsetse hosts.

Tsetse transmit trypanosomes in

two ways, mechanical and
biological transmission.

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 Mechanical transmission
involves the direct
transmission of the same
individual trypanosomes taken
from an infected host into an
uninfected host. The name
'mechanical' reflects the
similarity of this mode of
transmission to mechanical
injection with a syringe.
Mechanical transmission
requires the tsetse to feed on
an infected host and acquire
trypanosomes in the blood

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 meal, and then, within a
relatively short period, to feed
on an uninfected host and
regurgitate some of the
infected blood from the first
blood meal into the tissue of
the uninfected animal. This
type of transmission occurs
most frequently when tsetse
are interrupted during a blood
meal and attempt to satiate
themselves with another meal.
Other flies, such as horse-flies,
can also cause mechanical

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 transmission of trypanosomes.
[28]

Biological transmission
requires a period of incubation
of the trypanosomes within the
tsetse host. The term
'biological' is used because
trypanosomes must reproduce
through several generations
inside the tsetse host during
the period of incubation
(development within the fly is
known as the extrinsic
incubation period), which

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 requires extreme adaptation of
the trypanosomes to their
tsetse host. In this mode of
transmission, trypanosomes
reproduce through several
generations, changing in
morphology at certain periods.
This mode of transmission also
includes the sexual phase of
the trypanosomes. Tsetse are
believed to be more likely to
become infected by
trypanosomes during their first
few blood meals. Tsetse

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 infected by trypanosomes are
thought to remain infected for
the remainder of their lives.
Because of the adaptations
required for biological
transmission, trypanosomes
that can be transmitted
biologically by tsetse cannot
be transmitted in this manner
by other insects.

The relative importance of these

two modes of transmission for
the propagation of tsetse-
vectored trypanosomiases is not

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 yet well understood. However,
since the sexual phase of the
trypanosome life cycle occurs
within the tsetse host, biological
transmission is a required step in
the life cycle of the tsetse-
vectored trypanosomes.

The cycle of biological

transmission of trypanosomiasis
involves two phases, one inside
the tsetse host and the other
inside the vertebrate host.
Trypanosomes are not passed
between a pregnant tsetse and

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 her offspring, so all newly
emerged tsetse adults are free of
infection. An uninfected fly that
feeds on an infected vertebrate
animal may acquire
trypanosomes in its proboscis or
gut. These trypanosomes,
depending on the species, may
remain in place, move to a
different part of the digestive
tract, or migrate through the
tsetse body into the salivary
glands. When an infected tsetse
bites a susceptible host, the fly

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 may regurgitate part of a
previous blood meal that
contains trypanosomes, or may
inject trypanosomes in its saliva.
Inoculation must contain a
minimum of 300 to 450
individual trypanosomes to be
successful, and may contain up
to 40,000 cells.[26]

In the case of T. b. brucei

infecting G. p. gambiensis, during
this time the parasite changes
the proteome contents of the
fly's head. This may be the

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 reason/a reason for the
behavioral changes seen,
especially the unnecessarily
increased feeding frequency,
which increases transmission
opportunities. This may be due in
part to the altered glucose
metabolism observed, causing a
perceived need for more calories.
(The metabolic change, in turn,
being due to complete absence
of glucose-6-phosphate 1-
dehydrogenase in infected flies.)
Monoamine neurotransmitter

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 synthesis is also altered:
Production of aromatic L-amino
acid decarboxylase - involved in
dopamine and serotonin
synthesis - and α-methyldopa
hypersensitive protein was
induced. This is very similar to
the alterations in other dipteran
vectors' head proteomes under
infection by other eukaryotic
parasites of mammals, found in
another study by the same team
in the same year.[29]

The trypanosomes are injected

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 into vertebrate muscle tissue, but
make their way, first into the
lymphatic system, then into the
bloodstream, and eventually into
the brain. The disease causes the
swelling of the lymph glands,
emaciation of the body, and
eventually leads to death.
Uninfected tsetse may bite the
infected animal prior to its death
and acquire the disease, thereby
closing the transmission cycle.

Disease hosts and vectors

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 The tsetse-vectored
trypanosomiases affect various
vertebrate species including
humans, antelopes, bovine cattle,
camels, horses, sheep, goats, and
pigs. These diseases are caused
by several different trypanosome
species that may also survive in
wild animals such as crocodiles
and monitor lizards. The diseases
have different distributions
across the African continent, so
are transmitted by different
species. This table summarizes

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 this information:[26][30]

Species Trypanosoma Glossina

Disease Distribution
affected agents vectors

G. palpalis
Sleeping G.
sickness T. brucei Western tachinoides
humans
— chronic gambiense Africa G. fuscipes
form G.
morsitans

G.
Sleeping morsitans
sickness T. brucei Eastern G.
humans
— acute rhodesiense Africa swynnertoni
form G. pallidipes
G. fuscipes

G.
morsitans
G.
antelope
Nagana — swynnertoni
cattle T. brucei
acute Africa G. pallidipes
camels brucei
form G. palpalis
horses
G.
tachinoides
G. fuscipes

G. palpalis
G.
morsitans
Go
G. austeni
G.
Nagana — cattle swynnertoni

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 chronic camels T. congolense Africa G. pallidipes
form horses G.
longipalpis
G.
tachinoides
G.
brevipalpis

G. palpalis
G. fuscipes
G.
morsitans
G.
tachinoides
domestic pigs
Nagana — G.
cattle
acute T. simiae[31] Africa longipalpis
camels
form G. fusca
horses
G.
tabaniformis
G.
brevipalpis
G. vanhoofi
G. austeni

G.
morsitans
G. palpalis
G.
tachinoides
Nagana — cattle
G.
acute camels T. vivax Africa
swynnertoni
form horses
G. pallidipes
G. austeni
G. vanhoofi
G.

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 longipalpis

G. palpalis
G. fuscipes
G.
morsitans
domestic pigs
G.
warthog
tachinoides
—
Surra — G.
(Phacochoerus
chronic T. suis Africa longipalpis
aethiopicus)
form G. fusca
forest hogs
G.
—(Hylochoerus
tabaniformis
spp.)
G.
brevipalpis
G. vanhoofi
G. austeni

In humans

Human African trypanosomiasis,

also called sleeping sickness, is
caused by trypanosomes of the
species Trypanosoma brucei.
This disease is invariably fatal if

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 left untreated, but can almost
always be cured with current
medicines if the disease is
diagnosed early enough.

Sleeping sickness begins with a

tsetse bite leading to an
inoculation in the subcutaneous
tissue. The infection moves into
the lymphatic system, leading to
a characteristic swelling of the
lymph glands called
Winterbottom's sign.[32] The
infection progresses into the
blood stream and eventually

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 crosses into the central nervous
system and invades the brain
leading to extreme lethargy and
eventually to death.

The species Trypanosoma

brucei, which causes the disease,
has often been subdivided into
three subspecies that were
identified based either on the
vertebrate hosts which the strain
could infect or on the virulence of
the disease in humans. The
trypanosomes infectious to
animals and not to humans were

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 named Trypanosoma brucei
brucei. Strains that infected
humans were divided into two
subspecies based on their
different virulences:
Trypanosoma brucei gambiense
was thought to have a slower
onset and Trypanosoma brucei
rhodesiense refers to strains with
a more rapid, virulent onset. This
characterization has always been
problematic but was the best that
could be done given the
knowledge of the time and the

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 tools available for identification. A
recent molecular study using
restriction fragment length
polymorphism analysis suggests
that the three subspecies are
polyphyletic,[33] so the
elucidation of the strains of T.
brucei infective to humans
requires a more complex
explanation. Procyclins are
proteins developed in the surface
coating of trypanosomes whilst
in their tsetse fly vector.[34]

Other forms of human

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 trypanosomiasis also exist but
are not transmitted by tsetse. The
most notable is American
trypanosomiasis, known as
Chagas disease, which occurs in
South America, caused by
Trypanosoma cruzi, and
transmitted by certain insects of
the Reduviidae, members of the
Hemiptera.

In domestic animals

Animal trypanosomiasis, also

called nagana when it occurs in

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 bovine cattle or horses or sura
when it occurs in domestic pigs,
is caused by several
trypanosome species. These
diseases reduce the growth rate,
milk productivity, and strength of
farm animals, generally leading to
the eventual death of the infected
animals. Certain species of cattle
are called trypanotolerant
because they can survive and
grow even when infected with
trypanosomes although they also
have lower productivity rates

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 when infected.

The course of the disease in

animals is similar to the course of
sleeping sickness in humans.

Trypanosoma congolense and

Trypanosoma vivax are the two
most important species infecting
bovine cattle in sub-Saharan
Africa. Trypanosoma simiae
causes a virulent disease in
swine.

Other forms of animal

trypanosomiasis are also known

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 from other areas of the globe,
caused by different species of
trypanosomes and transmitted
without the intervention of the
tsetse fly.

The tsetse fly vector ranges

mostly in the central part of
Africa.

Trypanosomiasis poses a
considerable constraint on
livestock agricultural
development in Tsetse fly
infested areas of sub Saharan

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 Africa, especially in west and
central Africa. International
research conducted by ILRI in
Nigeria, the Democratic Republic
of the Congo and Kenya has
shown that the N'Dama is the
most resistant breed.[35][36]

Control
The conquest of sleeping
sickness and nagana would be of
immense benefit to rural
development and contribute to
poverty alleviation and improved

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 food security in sub-Saharan
Africa. Human African
trypanosomosis (HAT) and
animal African trypanosomosis
(AAT) are sufficiently important
to make virtually any intervention
against these diseases
beneficial.[37]

Tsetse fly from Burkina Faso

The disease can be managed by

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 controlling the vector and thus
reducing the incidence of the
disease by disrupting the
transmission cycle. Another
tactic to manage the disease is to
target the disease directly using
surveillance and curative or
prophylactic treatments to
reduce the number of hosts that
carry the disease.

Economic analysis indicates that

the cost of managing
trypanosomosis through the
elimination of important

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 populations of major tsetse
vectors will be covered several
times by the benefits of tsetse-
free status.[38] Area-wide
interventions against the tsetse
and trypanosomosis problem
appear more efficient and
profitable if sufficiently large
areas, with high numbers of
cattle, can be covered.

Vector control strategies can aim

at either continuous suppression
or eradication of target
populations. Tsetse fly

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 eradication programmes are
complex and logistically
demanding activities and usually
involve the integration of different
control tactics, such as
trypanocidal drugs, impregnated
treated targets (ITT), insecticide-
treated cattle (ITC), aerial
spraying (Sequential Aerosol
Technique - SAT) and in some
situations the release of sterile
males (sterile insect technique –
SIT). To ensure sustainability of
the results, it is critical to apply

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 the control tactics on an area-
wide basis, i.e. targeting an
entire tsetse population that is
preferably genetically isolated.

Control techniques

Many techniques have reduced

tsetse populations, with earlier,
crude methods recently replaced
by methods that are cheaper,
more directed, and ecologically
better.

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 Slaughter of wild animals

One early technique involved

slaughtering all the wild animals
tsetse fed on. For example, the
island of Principe off the west
coast of Africa was entirely
cleared of feral pigs in the 1930s,
which led to the extirpation of the
fly. While the fly eventually re-
invaded in the 1950s, the new
population of tsetse was free
from the disease.[39][40][41][42]

Land clearing

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 Another early technique involved
complete removal of brush and
woody vegetation from an area.
[43] However, the technique was
not widely used and has been
abandoned. Tsetse tend to rest
on the trunks of trees so
removing woody vegetation
made the area inhospitable to the
flies. Until about 1959 this was
done by hand and so was quite
time consuming. Glover et al
1959 describes the technique
which they call "chain clearing".

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 Chain clearing drags a chain
forward between two heavy
vehicles and thereby does the
same job much more quickly -
but still at some expense.[43]
Preventing regrowth of woody
vegetation requires continuous
clearing efforts which is even
more expensive,[43] and only
practical where large human
populations are present. Also, the
clearing of woody vegetation has
come to be seen as an
environmental problem more

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 than a benefit.

Pesticide campaigns

Pesticides have been used to

control tsetse starting initially
during the early part of the
twentieth century in localized
efforts using the inorganic metal-
based pesticides, expanding
after the Second World War into
massive aerial- and ground-
based campaigns with
organochlorine pesticides such
as DDT applied as aerosol sprays

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 at Ultra-Low Volume rates. Later,
more targeted techniques used
pour-on formulations in which
advanced organic pesticides
were applied directly to the backs
of cattle.

Trapping

Tsetse trap

Tsetse populations can be

monitored and effectively

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 controlled using simple,
inexpensive traps. These often
use electric blue cloth, either in
sheet or biconical form, since this
color attracts the flies. The traps
work by channeling the flies into
a collection chamber, or by
exposing the flies to insecticide
sprayed on the cloth. Early traps
mimicked the form of cattle, as
tsetse are also attracted to large
dark colors like the hides of cows
and buffaloes. Some scientists
put forward the idea that zebra

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 have stripes, not as a camouflage
in long grass, but because the
black and white bands tend to
confuse tsetse and prevent
attack.[44][45]

The use of chemicals as

attractants to lure tsetse to the
traps has been studied
extensively in the late 20th
century, but this has mostly been
of interest to scientists rather
than as an economically
reasonable solution. Attractants
studied have been those tsetse

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 might use to find food, like
carbon dioxide, octenol, and
acetone—which are given off in
animals' breath and distributed
downwind in an odor plume.
Synthetic versions of these
chemicals can create artificial
odor plumes. A cheaper
approach is to place cattle urine
in a half gourd near the trap. For
large trapping efforts, additional
traps are generally cheaper than
expensive artificial attractants.

A special trapping method is

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 applied in Ethiopia, where the
BioFarm Consortium (ICIPE,
BioVision Foundation, BEA,
Helvetas, DLCO-EA, Praxis
Ethiopia) applies the traps in a
sustainable agriculture and rural
development context (SARD).
The traps are just the entry point,
followed by improved farming,
human health and marketing
inputs. This method is in the final
stage of testing (as of 2006).

Sterile insect technique

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 The sterile insect technique (SIT)
is a form of pest control that uses
ionizing radiation (gamma ray or
X-ray) to sterilize male flies that
are mass-produced in special
rearing facilities. The sterile
males are released systematically
from the ground or by air in
tsetse-infested areas, where they
mate with wild females, which do
not produce offspring. As a result,
this technique can eventually
eradicate populations of wild
flies. SIT is among the most

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 environmentally friendly control
tactics available, and is usually
applied as the final component of
an integrated campaign. It has
been used to subdue the
populations of many other fly
species including the medfly,
Ceratitis capitata.

The sustainable removal of the

tsetse fly is in many cases the
most cost-effective way of
dealing with the T&T problem
resulting in major economic
benefits for subsistence farmers

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 in rural areas. Insecticide-based
methods are normally very
ineffective in removing the last
remnants of tsetse populations,
while, on the contrary, sterile
males are very effective in finding
and mating the last remaining
females. Therefore, the
integration of the SIT as the last
component of an area-wide
integrated approach is essential
in many situations to achieve
complete eradication of the
different tsetse populations,

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 particularly in areas of more
dense vegetation.

A project that was implemented

from 1994 to 1997 on the Island
of Unguja, Zanzibar (United
Republic of Tanzania),
demonstrated that, after
suppression of the tsetse
population with insecticides, SIT
completely removed the Glossina
austeni Newstead population
from the Island.[46][47] This was
carried out without any
understanding of the population

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 genetics of G. a., but future SIT
efforts can benefit from such
preparation. Population genetics
would help to select the Glossina
population to be deployed for
similarity to the target population.
[48] The eradication of the tsetse
fly from Unguja Island in 1997
was followed by the
disappearance of the AAT which
enabled farmers to integrate
livestock keeping with cropping
in areas where this had been
impossible before. The increased

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 livestock and crop productivity
and the possibility of using
animals for transport and traction
significantly contributed to an
increase in the quality of people's
lives.[49][50] Surveys in 1999,
2002, 2014, and 2015 have
confirmed this success -
continued absence of tsetse and
nagana on the island.[51]

In the Niayes region of Senegal, a

coastal area close to Dakar,
livestock keeping was difficult
due to the presence of a

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 population of Glossina palpalis
gambiensis. Feasibility studies
indicated that the fly population
was confined to very fragmented
habitats and a population
genetics study indicated that the
population was genetically
isolated from the main tsetse belt
in the south eastern part of
Senegal. After completion of the
feasibility studies (2006–2010),
an area-wide integrated
eradication campaign that
included an SIT component was

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 started in 2011, and by 2015, the
Niayes region had become
almost tsetse fly free. This has
allowed a change of cattle
breeds from lower producing
trypanotolerant breeds to higher-
producing foreign breeds.[52][53]

The entire target area (Block 1, 2

and 3) has a total surface of
1000 km2, and the first block
(northern part) can be
considered free of tsetse, as
intensive monitoring has failed to
detect since 2012 a single wild

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 tsetse fly. The prevalence of AAT
has decreased from 40 to 50%
before the project started to less
than 10% to date in blocks 1 and
2. Although insecticides are
being used for fly suppression,
they are applied for short periods
on traps, nets and livestock, and
are not spread into the
environment. After the
suppression activities are
completed, no more insecticide is
applied in the area. The removal
of trypanosomosis will eliminate

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 the need for constant
prophylactic treatments of the
cattle with trypanocidal drugs,
therefore reducing residues of
these drugs in the dung, meat
and milk.

The main beneficiaries of the

project are the many small holder
farmers, the larger commercial
farms and the consumers of
meat and milk. According to a
socio-economic survey and
benefit cost analysis,[54] after
eradication of the tsetse farmers

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 will be able to replace their local
breeds with improved breeds and
increase their annual income by
€2.8 million. In addition, it is
expected that the number of
cattle will be reduced by 45%,
which will result in reduced
environmental impacts.

Societal impact
In the literature of environmental
determinism, the tsetse has been
linked to difficulties during early
state formation for areas where

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 the fly is prevalent. A 2012 study
used population growth models,
physiological data, and
ethnographic data to examine
pre-colonial agricultural practices
and isolate the effects of the fly. A
"tsetse suitability index" was
developed from insect population
growth, climate and geospatial
data to simulate the fly's
population steady state. An
increase in the tsetse suitability
index was associated with a
statistically significant weakening

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 of the agriculture, levels of
urbanization, institutions and
subsistence strategies. Results
suggest that the tsetse
decimated livestock populations,
forcing early states to rely on
slave labor to clear land for
farming, and preventing farmers
from taking advantage of natural
animal fertilizers to increase crop
production. These long-term
effects may have kept population
density low and discouraged
cooperation between small-scale

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 communities, thus preventing
stronger nations from forming.

The authors also suggest that

under a lower burden of tsetse,
Africa would have developed
differently. Agriculture (measured
by the usage of large
domesticated animals, intensive
agriculture, plow use and female
participation rate in agriculture)
as well as institutions (measured
by the appearance of indigenous
slavery and levels of
centralization) would have been

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 more like those found in Eurasia.
Qualitative support for this claim
comes from archaeological
findings; e.g., Great Zimbabwe is
located in the African highlands
where the fly does not occur, and
represented the largest and
technically most advanced
precolonial structure in Southern
sub-Sahara Africa.[55]

Other authors are more skeptical

that the Tsetse fly had such an
immense influence on African
development. One conventional

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 argument is that the Tsetse fly
made it difficult to use draught
animals. Hence, wheeled forms of
transportations were not used as
well. While this is certainly true
for areas with high densities of
the fly, similar cases outside
tsetse-suitable areas exist. While
the fly definitely had a relevant
influence on the adoption of new
technologies in Africa, it has been
contended that it does not
represent the single root cause.
[56]

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 History

According to an article in the

New Scientist, the depopulated
and apparently primevally wild
Africa seen in wildlife
documentary films was formed in
the 19th century by disease, a
combination of rinderpest and
the tsetse fly. Rinderpest is
believed to have originated in
Asia, later spreading through the
transport of cattle.[57] In 1887,
the rinderpest virus was
accidentally imported in livestock

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 brought by an Italian
expeditionary force to Eritrea. It
spread rapidly, reaching Ethiopia
by 1888, the Atlantic coast by
1892 and South Africa by 1897.
Rinderpest, a cattle plague from
central Asia, killed over 90% of
the cattle of the pastoral peoples
such as the Masai of east Africa.
In South Africa, with no native
immunity, most of the population
– some 5.5 million domestic
cattle – died. Pastoralists and
farmers were left with no animals

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 – their source of income – and
farmers were deprived of their
working animals for ploughing
and irrigation. The pandemic
coincided with a period of
drought, causing widespread
famine. The starving human
populations died of smallpox,
cholera, and typhoid, as well as
African Sleeping Sickness and
other endemic diseases. It is
estimated that two-thirds of the
Masai died in 1891.[58]

The land was left emptied of its

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 cattle and its people, enabling the
colonial powers Germany and
Britain to take over Tanzania and
Kenya with little effort. With
greatly reduced grazing,
grassland turned rapidly to bush.
The closely cropped grass sward
was replaced in a few years by
woody grassland and thornbush,
ideal habitat for tsetse flies. Wild
mammal populations increased
rapidly, accompanied by the
tsetse fly. Highland regions of
east Africa which had been free

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 of tsetse fly were colonised by
the pest, accompanied by
sleeping sickness, until then
unknown in the area. Millions of
people died of the disease in the
early 20th century.[58]

Serengeti National Park, Tanzania

The areas occupied by the tsetse

fly were largely barred to animal
husbandry. Sleeping sickness
was dubbed "the best game
warden in Africa" by

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 conservationists, who assumed
that the land, empty of people
and full of game animals, had
always been like that. Julian
Huxley of the World Wildlife Fund
called the plains of east Africa "a
surviving sector of the rich
natural world as it was before the
rise of modern man".[58] They
created numerous large reserves
for hunting safaris. In 1909 the
newly retired president Theodore
Roosevelt went on a safari that
brought over 10,000 animal

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 carcasses to America. Later,
much of the land was turned over
to nature reserves and national
parks such as the Serengeti,
Masai Mara, Kruger and
Okavango Delta. The result,
across eastern and southern
Africa, is a modern landscape of
manmade ecosystems: farmland
and pastoral land largely free of
bush and tsetse fly; and bush
controlled by the tsetse fly.[58]

Although the colonial powers

saw the disease as a threat to

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 their interests, and acted
accordingly to bring transmission
almost to a halt in the 1960s,
[59]: 0174 this improved situation
led to a laxity of surveillance and
management by the newly
independent governments
covering the same areas - and a
resurgence that became a crisis
again in the 1990s.[59]: 0174
[59]: 0175

Current situation

Tsetse flies are regarded as a

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 major cause of rural poverty in
sub-Saharan Africa[9] because
they prevent mixed farming. The
land infested with tsetse flies is
often cultivated by people using
hoes rather than more efficient
draught animals because
nagana, the disease transmitted
by tsetse, weakens and often kills
these animals. Cattle that do
survive produce little milk,
pregnant cows often abort their
calves, and manure is not
available to fertilize the worn-out

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 soils.

Tsetse fly from Burkina Faso

The disease nagana or African

animal trypanosomiasis (AAT)
causes gradual health decline in
infected livestock, reduces milk
and meat production, and
increases abortion rates. Animals
eventually succumb to the
disease - annual cattle deaths
caused by trypanosomiasis are
estimated at 3 million, reducing

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 annual cattle production value by
US$600m-US$1.2b.[9] This has
an enormous impact on the
livelihood of farmers who live in
tsetse-infested areas, as infected
animals cannot be used to
plough the land, and keeping
cattle is only feasible when the
animals are kept under constant
prophylactic treatment with
trypanocidal drugs, often with
associated problems of drug
resistance, counterfeited drugs,
and suboptimal dosage. The

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 overall annual direct lost potential
in livestock and crop production
was estimated at US$4.5
billion[38][60]-US$4.75b.[9]

The tsetse fly lives in nearly

10,000,000 square kilometres
(4,000,000 sq mi) in sub-
Saharan Africa[9] (mostly wet
tropical forest) and many parts of
this large area is fertile land that
is left uncultivated—a so-called
green desert not used by humans
and cattle. Most of the 38
countries[9] infested with tsetse

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 are poor, debt-ridden and
underdeveloped. Of the 38[9]
tsetse-infested countries, 32 are
low-income, food-deficit
countries (http://www.fao.org/cou
ntryprofiles/lifdc/en/) , 29 are
least developed countries, and
30 or 34[9] are among the 40
most heavily indebted poor
countries. Eradicating the tsetse
and trypanosomiasis (T&T)
problem would allow rural
Africans to use these areas for
animal husbandry or the

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 cultivation of crops and hence
increase food production. Only
45 million cattle, of 172 million
present in sub-Saharan Africa,
are kept in tsetse-infested areas
but are often forced into fragile
ecosystems like highlands or the
semiarid Sahel zone, which
increases overgrazing and
overuse of land for food
production.

In addition to this direct impact,

the presence of tsetse and
trypanosomiasis discourages the

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 use of more productive exotic
and cross-bred cattle, depresses
the growth and affects the
distribution of livestock
populations, reduces the
potential opportunities for
livestock and crop production
(mixed farming) through less
draught power to cultivate land
and less manure to fertilize (in an
environment-friendly way) soils
for better crop production, and
affects human settlements
(people tend to avoid areas with

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 tsetse flies).

Tsetse flies transmit a similar

disease to humans, called African
trypanosomiasis, human African
trypanosomiasis (HAT) or
sleeping sickness. An estimated
60[9]-70[61] million people in 20
countries are at different levels of
risk and only 3-4 million people
are covered by active
surveillance.[9] The DALY index
(disability-adjusted life years), an
indicator to quantify the burden
of disease, includes the impact of

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 both the duration of life lost due
to premature death and the
duration of life lived with a
disability. The annual burden of
sleeping sickness is estimated at
2 million DALYs. Since the
disease tends to affect
economically active adults, the
total cost to a family with a
patient is about 25% of a year's
income.[62]

History of study
In East Africa, C. F. M.

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 Swynnerton played a large role in
the first half of the 20th century.
Swynnerton did much of the
earliest tsetse ecology research.
[63] For this E. E. Austen named a
patronymic taxon for him, G.
swynnertoni in 1922.[21]

Resistance to
trypanosomes
Tsetse flies have an arsenal of
immune defenses to resist each
stage of the trypanosome
infectious cycle, and thus are

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 relatively refractory to
trypanosome infection.[64]
Among the host flies' defenses is
the production of hydrogen
peroxide,[65] a reactive oxygen
species that damages DNA.
These defenses limit the
population of infected flies.

See also
David Bruce (microbiologist)
G.D. Hale Carpenter joined the
London School of Hygiene and
Tropical Medicine, and took the

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 DM in 1913 with a dissertation
on the tsetse fly (Glossina
palpalis) and sleeping
sickness. He published: A
Naturalist on Lake Victoria,
with an Account of Sleeping
Sickness and the Tse-tse Fly;
1920. T.F. Unwin Ltd, London;
Biodiversity Archive (https://ar
chive.org/details/naturalistonla
ke00carp)
Muriel Robertson, who
conducted early 20th century
research on the insect

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:
 Use of DNA in forensic
entomology
Horses in Botswana

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Further reading
Gerster, George (December
1986). "Tsetse". National
Geographic. Vol. 170, no. 6.
pp. 814–833. ISSN 0027-
9358 (https://www.worldcat.or
g/issn/0027-9358) .
OCLC 643483454 (https://w

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Page 173 of 184
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 ww.worldcat.org/oclc/643483
454) .
Gooding, R.H.; Krafsur, E.S.
(2005). "Tsetse genetics:
Contributions to Biology,
Systematics, and Control of
Tsetse Flies" (https://www.ncb
i.nlm.nih.gov/pmc/articles/PM
C1462949) . Annual Review
of Entomology. Annual
Reviews. 50 (1): 101–123.
doi:10.1146/annurev.ento.50.0
71803.130443 (https://doi.or
g/10.1146%2Fannurev.ento.50.

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 071803.130443) .
ISSN 0066-4170 (https://ww
w.worldcat.org/issn/0066-417
0) . PMC 1462949 (https://ww
w.ncbi.nlm.nih.gov/pmc/article
s/PMC1462949) .
PMID 15355235 (https://pubm
ed.ncbi.nlm.nih.gov/1535523
5) . S2CID 22834246 (https://
api.semanticscholar.org/Corpu
sID:22834246) .

Textbooks

Buxton, P. (1955). The Natural

History of Tsetse Flies: An

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 Account of the Biology of the
Genus Glossina (Diptera).
London, UK: H. K. Lewis & Co.
Ltd.
Ford, J. (1971). The Role of the
Trypanosomiases in African
Ecology. Oxford, UK:
Clarendon Press.
Glasgow, J. (1963). The
Distribution and Abundance of
Tsetse. International Series of
Monographs on Pure and
Applied Biology, No. 20.
Oxford, UK: Pergamon Press.

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 Leak, S. (1998). Tsetse Biology
and Ecology: Their role in the
Epidemiology and Control of
Trypanosomiasis. New York:
CABI Publishing. book site (htt
ps://web.archive.org/web/2011
0613040449/http://www.cabi
-publishing.org/bookshop/Boo
kDisplay.asp?SubjectArea=Hu
m&Subject=Public+Health+an
d+Communicable+Diseases&
PID=1406)
Maudlin, I., Holmes, P. H., and
Miles, M. A. (2004). The

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 Trypanosomiases. CAB
International.
McKelvey, J., Jr. (1973). Man
Against Tsetse: Struggle for
Africa. Ithaca, NY: Cornell
University Press.
Mulligan, H. & Potts, W. (1970).
The African Trypanosomiases.
London: George Allen and
Unwin, Ltd.

External links

Wikimedia Commons has media

related to Glossina.

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 Wikispecies has information
related to Glossina.
Programmes and information
to assist in the planning and
implementation of tsetse
control operations (http://www.
tsetse.org)
Programme Against African
Trypanosomiasis (http://www.f
ao.org/ag/paat.html)
PAN AFRICAN TSETSE AND
TRYPANOSOMIASIS
ERADICATION CAMPAIGN
(PATTEC) (https://web.archive.

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Page 179 of 184
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 org/web/20050409122056/h
ttp://www.africa-union.org/Stru
cture_of_the_Commission/dep
Pattec.htm)
Tsetse in the Transvaal and
Surrounding Territories - An
Historical Review (https://web.
archive.org/web/2012022704
0712/http://www.mosquitocata
log.org/files/pdfs/045050-0.p
df) —Claude Fuller (Division of
Entomology, 1923)
Leverhulme Trust Tsetse
Research Network (LTTRN) (ht

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 tps://web.archive.org/web/200
70606225734/http://www.lsh
tm.ac.uk/pmbu/lttrn/)
BITING FLIES - The NZI Trap (
https://archive.today/2013010
4081354/http://www.nzitrap.c
om/index.htm)
Distribution maps (https://web.
archive.org/web/2009030114
2911/http://ergodd.zoo.ox.ac.u
k/livatl2/tsetse.htm)
"The vector (tsetse fly)" (http
s://web.archive.org/web/2016
0929202859/http://www.who.

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 int/trypanosomiasis_african/di
sease/vector/en/) . World
Health Organization. 5 August
2016. Archived from the
original (http://www.who.int/try
panosomiasis_african/disease/
vector/en/) on 29 September
2016. Retrieved 4 December
2020.
STRATEGIC REVIEW OF
TRAPS AND TARGETS FOR
TSETSE AND AFRICAN
TRYPANOSOMIASIS
CONTROL - Training in

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 Tropical Diseases (http://www.
who.int/tdr/publications/docu
ments/tsetse_traps.pdf)
"Insect of the Month
(October): Tsetse fly, Glossina
morsitans" (http://www.icipe.or
g/news/insect-month-october
-tsetse-fly-glossina-morsitan
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Retrieved from
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