Tsetse fly

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"Tsetse" redirects here. For other uses, see Tsetse (disambiguation).

Tsetse fly

Temporal range: Eocene - Recent 34–

0 Ma 

PreꞒ

Pg

Glossina morsitans

Scientific classification
Kingdom: Animalia

Phylum: Arthropoda

Class: Insecta

Order: Diptera

(unranked): Eremoneura

(unranked): Cyclorrhapha

Section: Schizophora

Subsection: Calyptratae

Superfamily: Hippoboscoidea

Family: Glossinidae
Theobald, 1903

Genus: Glossina
Wiedemann, 1830

Species groups

 Morsitans ("savanna
h" subgenus)
 Fusca ("forest"
subgenus)
 Palpalis ("riverine"
subgenus)
 Range of the tsetse fly

Tsetse (/ˈsiːtsi/ SEET-see, US: /ˈtsiːtsi/ TSEET-see or UK: /ˈtsɛtsə/ TSET-sə), sometimes
spelled tzetze and 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 placed in their own family, Glossinidae. The tsetse are obligate parasites that
live by feeding on the blood of vertebrate animals. Tsetse have been extensively
studied because of their role in transmitting disease. They have a prominent
economic impact in sub-Saharan Africa as the biological vectors of trypanosomes,
which cause human sleeping sickness and animal trypanosomiasis. Tsetse
are multivoltine and long-lived, typically producing about four broods per year, and
up to 31 broods over their lifespans.[4]
Tsetse can be distinguished from other large flies by two easily observed features.
Tsetse fold their wings completely when they are resting so that one wing rests
directly on top of the other over their abdomens. Tsetse also have a long proboscis,
which extends directly forward and is attached by a distinct bulb to the bottom of
their heads.
Fossilized tsetse have been recovered from Paleogene aged rocks in the United
States and Germany. Twenty-three extant species of tsetse flies are known from
Africa and Arabia.
Contents

 1Etymology
 2Biology
o 2.1Morphology
o 2.2Anatomy
o 2.3Life cycle
o 2.4Hosts
o 2.5Genetics
o 2.6Symbionts
o 2.7Diseases
 3Systematics
o 3.1Species
o 3.2Evolutionary history
 4Range
 5Trypanosomiasis
o 5.1Disease hosts and vectors
o 5.2In humans
o 5.3In domestic animals
 6Control
o 6.1Control techniques
  6.1.1Slaughter of wild animals
 6.1.2Land clearing
 6.1.3Pesticide campaigns
 6.1.4Trapping
 6.1.5Sterile insect technique
 7Societal impact
o 7.1History
o 7.2Current situation
 8History of study
 9Resistance to trypanosomes
 10See also
 11References
 12Further reading
o 12.1Textbooks
 13External links

Etymology[edit]
The word tsetse means "fly" in Tswana, a Bantu language of southern Africa.
[5]
 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 antisubmarine
aircraft was known as the 'Tsetse' Mosquito.[6]

Biology[edit]
The biology of tsetse is relatively well understood by entomologists. They have been
extensively studied because of their medical, veterinary, and economic importances,
because the flies can be raised in a laboratory, and because they are relatively large,
facilitating their analysis.
Morphology[edit]
Tsetse flies can be seen as independent individuals in three forms: as third-
instar larvae, pupae, and adults.
Tsetse first become 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 few hours, and is almost never observed outside of the laboratory.
Tsetse next develop a hard external case, the puparium, and become pupae—small,
hard-shelled, oblongs with two distinctive, small, dark lobes at the tail (breathing)
end. Tsetse pupae are under 1 centimetre (1⁄2 in) long.[7] Within the puparial shell,
tsetse complete the last two larval instars and the pupal stage.
At the end of the pupal stage, tsetse emerge as adult flies. The adults are relatively
large flies, with lengths of 0.5–1.5 centimetres (1⁄4–5⁄8 in),[7] 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 wide rather than elongated and shorter than the
wings.
Four characteristics definitively separate adult tsetse from other kinds of flies:
 Tsetse have a distinct proboscis, a long
Proboscis thin structure attached to the bottom of
the head and 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 has a characteristic hatchet
Hatchet cell
shape resembling a meat cleaver or a
hatchet.

Branched The antennae have arista with hairs

arista hairs which are themselves branched.

Anatomy[edit]
Like all other insects, tsetse flies have an adult body comprising three visibly distinct
parts: the 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.
Reproductive anatomy sketch by [[es:User:Estefanía Alonso Gómez]]

The internal anatomy of tsetse is fairly typical of the insects. The crop is large

enough to accommodate a huge increase in size during the bloodmeal since tsetse
can take a bloodmeal equal in weight to themselves. The dipteran crop is heavily
understudied and Glossina is one of the few genera with relatively good information
available: Moloo and Kutuza 1970 for G. brevipalpis (including its innervation) and
Langley 1965 for G. morsitans.[8] 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 very tough. Houseflies are easily killed with a
flyswatter, but a great deal of effort is needed to crush a tsetse fly. [9]
Life cycle[edit]

Glossina palpalis and G. morsitans from a 1920 lexicon

Tsetse have an unusual life cycle which may be due to the richness of their food
source. A female fertilizes only one egg at a time and retains each egg within her
uterus to have the offspring develop internally during the first three larval stages, a
method called adenotrophic viviparity.[10] During this time, the female feeds the
developing offspring with a milky substance secreted by a modified gland in the
uterus.[11] In the third larval stage, the tsetse larva leaves the uterus and begins its
independent life. The newly independent tsetse larva crawls into the ground, and
develops a hard outer shell called the puparial case, in which it completes its
morphological transformation into an adult fly.
The larval life stage has a variable duration, generally 20 to 30 days, and the larva
must rely on stored resources during this time. The importance of the richness of
blood to this development can be seen, since all tsetse development before it
emerges from the puparial case as a full adult occurs without feeding, based only on
nutritional resources provided by the female parent. The female must get enough
energy for her needs, for the needs of her developing offspring, and for the stored
resources which her offspring will require until it emerges as an adult.
Technically, these insects undergo the standard development process of insects,
which consists of oocyte formation, ovulation, fertilization, development of the egg,
three larval stages, a pupal stage, and the emergence and maturation of the adult.
[citation needed]

Hosts[edit]
Overall Suidae are the most important hosts. By species, bloodmeals are derived
from:[12]

 G. swynnertoni — 60–70% from 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 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 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 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 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 ostrich
Waterbuck (Kobus ellipsiprymnus) are unmolested by Glossina[12][13] because they
produce volatiles which act as repellents. Waterbuck odor volatiles are under testing
and development as repellents to protect livestock.[14][15]: Suppl T1 
Genetics[edit]
The genome of Glossina morsitans was sequenced in 2014.[16]
Symbionts[edit]
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.[17][18]
Diseases[edit]
The salivary gland hypertrophy virus causes abnormal bleeding in the lobes of
the crop of G. m. centralis and G. m. morsitans.[8]

Systematics[edit]
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 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, which contains other hematophagous families.
Species[edit]
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The tsetse genus is generally split into three groups of species based on a
combination of distributional, behavioral, molecular and morphological
characteristics.[19] The genus includes:
 The 'savannah' flies:  The 'forest' flies:  The 'riverine' and
(subgenus Morsitans, (subgenus Fusca, 'lacustrine' flies:
occasionally previously (subgenus Palpalis,
named Glossina): named Austenia): previously
o Glossina o Glossi named Nemorhina):
austeni ( na o Glossina
Newstead, fusca caliginea (
1912) patr.  fusca ( Austen,
of Austen Walker, 1911)
o Glossina 1849) o Glossina
morsitan o Glossi fuscipes (N
s (Westwo na ewstead,
od, 1851) fuscipl 1911)
 euris ( 
G Austen, Gl
1911)
o Glossina o Glossi 
pallidipe na Gl
s (Austen, frezili 
1903) (Goute 
o Glossina ux,
Gl
1987)
swynnert [23]

oni (Auste o Glossi o Glossina

n, 1922)[21] pallicera
[22]
 patr. of  na
Swynnerto hanin pallicera (
gtoni ( Bigot, 1891)
n[22]
Newste o Glossina
ad and pallicera
Evans, newsteadi (
1922) Austen,
o Glossi 1929) patr. of 
na Newstead
longip o Glossina
ennis ( palpalis
Corti, palpalis (R
1895) obineau-
o Glossi Desvoidy,
na 1830)
medic o Glossina
orum ( palpalis
Austen, gambiensis 
1911) (Vanderplan
o Glossi k, 1911)
na o Glossina
nashi ( tachinoide
Potts, s (Westwood
1955) , 1850)
o Glossi
na
 nigrof
usca
nigrof
usca (
Newste
ad,
1911)
o Glossi
na
severi
ni (Ne
wstead,
1913)
o Glossi
na
schwe
tzi (Ne
wstead 
and
Evans,
1921)
o Glossi
na
tabani
formis 
(Westw
ood,
1850)
o Glossi
na
vanho
ofi (He
nrard,
1952)

Evolutionary history[edit]
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.[24]

Range[edit]
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.[20]

Trypanosomiasis[edit]
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 trypanosomiasis is called sleeping sickness. 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.
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 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.

 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 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 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
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 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 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 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[dubious  –  discuss] host, the fly 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 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 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 into vertebrate muscle tissue, [citation needed] 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[edit]
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 this information:[26][30]

Disease Species affected Trypanosoma agents Distribution Glossina vectors

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

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

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

G. palpalis
G. morsitans
G. austeni
cattle
Nagana — G. swynnertoni
camels T. congolense Africa
chronic form G. pallidipes
horses
G. longipalpis
G. tachinoides
G. brevipalpis

Nagana — domestic pigs T. simiae[31] Africa G. palpalis

 G. fuscipes
G. morsitans
G. tachinoides
cattle G. longipalpis
acute form camels G. fusca
horses G. tabaniformis
G. brevipalpis
G. vanhoofi
G. austeni

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

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

In humans[edit]
Main article: African trypanosomiasis
Human African trypanosomiasis, also called sleeping sickness, is caused by
trypanosomes of the species Trypanosoma brucei. This disease is invariably fatal
unless treated, 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 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 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 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][clarification needed]
Other forms of human 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[edit]
Main article: Animal trypanosomiasis
Animal trypanosomiasis, also called nagana when it occurs in 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 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 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 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[edit]
The conquest of sleeping sickness and nagana would be of immense benefit to rural
development and contribute to poverty alleviation and improved 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 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 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 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 the control tactics on an area-wide
basis, i.e. targeting an entire tsetse population that is preferably genetically isolated.
Control techniques[edit]
Many techniques have reduced tsetse populations, with earlier, crude methods
recently replaced by methods that are cheaper, more directed, and ecologically
better.
Slaughter of wild animals[edit]
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[edit]
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.[citation needed] 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". 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 than a benefit. [citation needed]
Pesticide campaigns[edit]
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
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[edit]

Tsetse trap

Tsetse populations can be monitored and effectively controlled using simple,

inexpensive traps. These often use electric blue cloth, since this color attracts the
flies. Early traps mimicked the form of cattle but this seems unnecessary and recent
traps are simple sheets or have a biconical form. The traps can kill by channeling the
flies into a collection chamber or by exposing the flies to insecticide sprayed on the
cloth. Tsetse are also attracted to large dark colors like the hides of cow and
buffaloes. Some scientists put forward the idea that zebra 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 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 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[edit]
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 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 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, 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 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 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 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 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 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 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
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[edit]
See also: African trypanosomiasis §  History
In the literature of environmental determinism, the tsetse has been linked to
difficulties during early state formation for areas where 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 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 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 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 sub-Sahara Africa. [55]
Other authors are more skeptical that the Tsetse fly had such an immense influence
on African development. One conventional 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]
History[edit]
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 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 – 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][additional citation(s)
needed]

The land was left emptied of its 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 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][additional citation(s) needed]

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
conservationists[citation needed], 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 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][additional citation(s) needed]
Although the colonial powers saw the disease as a threat to their interests, and acted
accordingly to bring transmission almost to a halt in the 1960s, [Sim 1] 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.[Sim 2][Sim 3]
Current situation[edit]
Tsetse flies are regarded as a major cause of rural poverty in sub-Saharan
Africa[10] 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 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 [citation needed], reducing annual cattle
production value by US$600m-US$1.2b.[10] 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 overall annual direct lost potential in livestock and crop production was
estimated at US$4.5 billion[38][59]-US$4.75b.[10]
The tsetse fly lives in nearly 10,000,000 square kilometres (4,000,000 sq mi) in sub-
Saharan Africa[10] (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[10] infested with tsetse are poor, debt-ridden and
underdeveloped. Of the 38[10] tsetse-infested countries, 32 are low-income, food-
deficit countries, 29 are least developed countries, and 30[citation needed] or 34[10] 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 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 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 tsetse flies).
Tsetse flies transmit a similar disease to humans, called African trypanosomiasis,
human African trypanosomiasis (HAT) or sleeping sickness. An estimated 60 [10]-
70[60] million people in 20 countries are at different levels of risk and only 3-4 million
people are covered by active surveillance.[10] The DALY index (disability-adjusted life
years), an indicator to quantify the burden of disease, includes the impact of 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.[61]

History of study[edit]
This section needs expansion. You
can help by adding to
it. (December 2021)

In East Africa, C. F. M. Swynnerton played a large role in the first half of the 20th
century. Swynnerton did much of the earliest tsetse ecology research. [62] For this E. E.
Austen named a patronymic taxon for him, G. swynnertoni in 1922.[22]

Resistance to trypanosomes[edit]
Tsetse flies have an arsenal of immune defenses to resist each stage of the
trypanosome infectious cycle, and thus are relatively refractory to trypanosome
infection.[63] Among the host flies' defenses is the production of hydrogen peroxide,
[64]
 a reactive oxygen species that damages DNA. These defenses limit the population
of infected flies.

See also[edit]
 David Bruce (microbiologist)
 G.D. Hale Carpenter joined the London School of
Hygiene and Tropical Medicine, and took
the 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
  Muriel Robertson, who conducted early 20th
century research on the insect
 Use of DNA in forensic entomology

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1. ^ p. 0174, "Systematic screening, treatment, and follow-

up of millions of individuals in the whole continent led to
transmission coming to a near halt by the 1960s."
2. ^ p. 0174,
   "With the advent of independence in most countries where HAT
was endemic, the newly independent authorities had other
priorities to deal with. The rarity of HAT cases, and a decline in
awareness of how the disease could return, led to a lack of interest
in disease surveillance. Over time the disease slowly returned, and
some thirty years later, flare-ups were observed throughout past
endemic areas (Figure 1).
   Since 1995, the World Health Organization (WHO) has on many
occasions expressed its concern about the rise in HAT cases."

3. ^ p. 0175, Figure 1. New Cases of Sleeping Sickness

Reported for All Africa between 1927 and 1997

Further reading[edit]
 Gerster, George (December 1986).
"Tsetse". National Geographic. Vol. 170, no. 6.
pp. 814–833. ISSN 0027-9358. OCLC 6434834
54.
 Gooding, R.H.; Krafsur, E.S. (2005). "Tsetse
genetics: Contributions to Biology, Systematics,
and Control of Tsetse Flies". Annual Review of
Entomology. Annual Reviews. 50 (1): 101–
123. doi:10.1146/annurev.ento.50.071803.13044
3. ISSN 0066-4170. PMC 1462949. PMID 1535
5235. S2CID 22834246.
Textbooks[edit]
 Buxton, P. (1955). The Natural History of Tsetse
Flies: An Account of the Biology of the Genus
 Glossina (Diptera). London, UK: H.K. Lewis &
Co.
 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.
 Leak, S. (1998). Tsetse Biology and Ecology:
Their role in the Epidemiology and Control of
Trypanosomiasis. New York: CABI
Publishing. book site
 Maudlin, I., Holmes, P. H., and Miles, M. A.
(2004). The 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[edit]
Wikimedia Commons has
media related to Glossina.

Wikispecies has
information related
to Glossina.

 Programmes and information to assist in the

planning and implementation of tsetse control
operations
 Programme Against African Trypanosomiasis
 PAN AFRICAN TSETSE AND
TRYPANOSOMIASIS ERADICATION
CAMPAIGN (PATTEC)
 Tsetse in the Transvaal and Surrounding
Territories - An Historical Review—Claude
Fuller (Division of Entomology, 1923)
 Leverhulme Trust Tsetse Research Network
(LTTRN)
 BITING FLIES - The NZI Trap
 Distribution maps
  "The vector (tsetse fly)". World Health
Organization. 2016-08-05. Archived from the
original on September 29, 2016. Retrieved 2020-
12-04.
 STRATEGIC REVIEW OF TRAPS AND
TARGETS FOR TSETSE AND AFRICAN
TRYPANOSOMIASIS CONTROL - Training in
Tropical Diseases
 "Insect of the Month (October): Tsetse
fly,  Glossina morsitans". icipe (International
Centre of Insect Physiology and Ecology).
Retrieved 2021-10-09.
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Extant Diptera families

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Human interactions with insects

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ssinidae

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6841

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333

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