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B.-7!9;:8.!7-&9;@K!8=@:=@86':!9-!A%8=!-.!:7-98-=K!P%@!%!B%78'8%.!@8<;9)!
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<D<!$@QBW!/Q@TOMTCL!UX!)MYMNK!@NS!%URMNK!ZMOP!OPC!2CTOUQ!
African societies have long coexisted with trypanosomiasis. In John Ford’s seminal work
The Role of the Trypanosomiases in African Ecology, Ford contended that pre-colonial societies
achieved resistance to the disease through limited, continuous exposure to the trypanosome (Ford
1971).

Accordingly, protection was acquired by modifying the environment to regulate

interactions between humans, domesticated animals, wild fauna, tsetse flies, and the
trypanosome.

Giblin (1990) reviewed Ford’s work, and presented alternative pre-colonial

methods of responding to the tsetse fly. One such method was found in Kjekshus’ Ecology
Control and Economic Development in East African History: the Case of Tanganyika, 18501950 (1977) in which avoidance of the tsetse fly was encouraged. Kjekshus’ method therefore
was one of evading the fly, while Ford found low levels of contact necessary in man’s
coexistence with tsetse. Torday (1910) seemed to agree with Kjekshus’ isolationist approach
when he described the people of the Kasai Basin in the Belgian Congo where sleeping sickness
was controlled by keeping populations away from the fly and through the practice of removing
sick villagers to isolated forests.!

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Whether pre-colonial Africans coexisted with trypanosomiasis and its persistent vector
through a process of limited but continuous exposure as suggested by Ford or if an isolationist

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approach was key to survival, the arrival of Europeans certainly disrupted the established
cohabitation practices. In discussing the colonial administration of Zambia, Vail (1977) listed
gun control laws, game control policies, village amalgamation policies, hut taxes, and labor
recruitment campaigns as a collection of policies that disrupted the pre-colonial coexistence of
man and the fly. Indeed, as early as 1908, David Bruce (1908), the Scottish microbiologist who
played the largest role in identifying the cause of sleeping sickness, noted the impact of colonists
in spreading sleeping sickness:
It cannot be forgotten that the introduction of sleeping sickness into Uganda was due to
England’s interference with existing conditions. The movement of large masses of men
or animals from the conditions to which they have become adapted is always attended
with danger. Civilized man presents the untutored savage… with what he calls the
dignity of labour with one hand, while with the other he scatters abroad the seeds of
tuberculosis, sleeping-sickness, and other pestilences which I need not enumerate.
In their influential work examining land degradation from the political ecology perspective,
Blaikie and Brookfield (1987) discussed the colonial role in the spread of the tsetse fly and
trypanosomiasis. Such an expansion in distribution occurred as the result of colonial policies
annexing land and forcing indigenous groups to move to areas previously avoided. These forced
relocations following the Europeans’ arrival contributed in no small part to the first sleeping
sickness epidemic (Vail 1977). Uganda and the Congo Basin experienced the worst of this
epidemic where, from 1896 to 1906, 200,000 people died from the disease (WHO 2010).

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2.3 Early Colonial Tsetse and Trypanosomiasis Control Practices
The early colonial control practices were implemented from the end of the nineteenth
century to the beginning of World War II (WWII). These included population evacuation, wild
host removal, tsetse habitat destruction, and the point-source techniques of traps and targets.

2.3.1 Population Evacuation
Knight (1971) suggested that the earliest form of active trypanosomiasis control was
population evacuation. This technique often involved moving entire villages to “safe areas.” In
1908, North-East Rhodesia, ruled by the British South Africa Company, pursued one such massevacuation from the eastern bank of the Luapula River (Figure 2.1). All villages along the bank
from Kabila to the Nsakaluba stream were moved to higher ground to avoid the tsetse population
at the river’s edge (Musambachime 1981). During this move, the abandoned villages were
burned in an effort to discourage the natives from returning. Those who were sick were sent into
quarantines from which they rarely returned.
Often during these evacuation events, the villagers were only allowed to carry basic
necessities, and rarely were their destinations adequately prepared for their arrival
(Musambachime 1981). Once relocated, it was common for villages to be densely resettled in
order to promote expedient collection of taxes by colonial officials and to allow for ease in
medically examining the resettled people (Vail 1977). The crowded conditions spawned
enormous overuse of land, which contributed to soil erosion, and the abandonment of previously
used land allowed tsetse habitat to regenerate, creating an environment ripe for the continued
spread of sleeping sickness (Vail 1977). Thus, the practice of population relocation was often
nothing short of disastrous. In fact, Musambachime (1981) presented an observation that, due to

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the disorderly nature of such relocations from the Luapula River, more people died from hunger
than died of sleeping sickness.
In addition to fostering conditions disastrous to the resettled population’s health, these
programs also disrupted lives both spiritually and economically. Relocation meant the
abandonment of ancestral resting places. According to Torday (1929), such displacement was
like “a tree cut off from its roots,” as relocated people were moved to environments deprived of
physical, cultural, and religious familiarity. Furthermore, relocation as well as the restrictions
placed on the resettled individuals’ movement disrupted rituals such as rain prayers and made
pilgrimages to sacred locations impossible (Musambachime 1981). Cattle, the economic core for
many African villages, often fared poorly in the newly settled areas due to concentrated
conditions that facilitated the spread of the disease (Soff 1969). As villagers lost their cattle and
other animals to AAT, it was not uncommon for livestock theft to increase, as occurred between
tribal groupings in British East Africa at the turn of the twentieth century (Soff 1969).
Additionally, population relocation further jeopardized economic prospects by eliminating the
use of more fertile land and disrupting local and regional trading of items such as salt, palm oil,
and fish (Musambachime 1981).

2.3.2 Wild Host Removal
Bruce (1905) presented an early discussion of the role wild animals play as a reservoir of the
disease. Bruce remarked that wild animals such as the buffalo and the wildebeest carried
trypanosomes in their blood, but that the parasite did not seem to hurt these animals. On the other
hand, when the parasite was introduced in domestic animals, AAT would result, often leading to
death. The practice of game destruction as a means of eliminating both the parasite reservoir and

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a food source for tsetse quickly followed Bruce’s discovery. In fact, Bruce (1905) personally
suggested the value of this control technique:
We have found that the reservoir of the disease exists in the wild animals, and that we can
blot out this disease from any particular tract of country by the simple expedient of
destroying or driving away the wild animals.
Elimination of wild species was widely debated following the discovery of their role in
harboring the parasite. At a meeting of The Royal African Society in 1913, Dr. Warrington
Yorke claimed that the only effective means of combating sleeping sickness was through the
elimination of the virus’ (today known to be a protist) reservoir. Dr. Yorke was specifically
addressing the situation in Rhodesia and Nyasaland where the population was troubled by the
tsetse species G. morsitans. As the local population could not be moved away from the infested
areas, Dr. Yorke advised “attempting to destroy the reservoir of the virus (Yorke 1913):”
It is obvious that the mere isolation of infected human beings is futile in view of the fact
that the main reservoir of the virus is the blood of the big game.
In addition to ridding the population of the parasite’s host, big game destruction was also
promoted as a means of eliminating the fly’s food supply, and thus causing the fly to disappear
(Yorke 1913).
Three years before Dr. Yorke’s call for the destruction of wild game, Alfred Sharpe, a British
colonial administrator, warned the Royal African Society of the difficulties and potential
unintended consequences of game destruction. He emphasized that certain conditions make
areas suitable for the existence of tsetse flies, and the presence of big game makes little
difference in their choice of habitat (Sharpe 1910). Additionally, it was cautioned that if the
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primary food source for the fly was removed, humans could be targeted as a secondary source,
thus making sleeping sickness more prevalent (Sharpe 1910). Arguments were also made for
more judicious slaughtering of game, acknowledging the successes of host destruction but
calling for better knowledge of the relationship between the fly and wild animals to avoid
ruthless mass killings (Swynnerton and Buxton 1938).
Ultimately, active host destruction declined as a control technique by the 1940s due to its
increasing social anathema and the rise of insecticides as a vector control option (Hargrove 2003;
Cox 2004). However, during its period of popularity and even after insecticides became more
broadly used, host destruction did offer effective, though modest, tsetse control. Jordan (1986)
described one such success that took place in Uganda from 1946 to 1966 (Figure 2.1). During
this period, active hunting of wild animals substantially reduced the populations of two tsetse
2

species from an area more than 20,000 km primarily due to its intensive culling of even the
most elusive animals (Jordan 1986).

2.3.2.1 Interference of Colonial Policies in Removing Wild Hosts
Often, host destruction is jeopardized when the level of hunting is not sufficient to achieve
the necessary degree of wild animal elimination. Reinvasion of the fly is common, and
sustainable elimination of hosts is therefore required for successful tsetse control. Vail (1977)
demonstrated how colonial policies limited the ability to sustain this necessary level of wildlife
suppression in eastern Zambia. During the first half of the twentieth century, the administration
of eastern Zambia, between the Luangwa River and the Zambia-Malawi border, was not unlike
many colonial regions of its time. Policies concerned with the capturing of resources and the
promotion of European interests frequently increased the vulnerability of Africans and left them

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to cope with foreseen and unforeseen consequences of these policies. Such policies included gun
control laws stemming from the Brussels Act of 1890, the conservation of wild animals from the
London Convention of 1900, hut taxes, and labor recruitment programs (Vail 1977).
Throughout Central Africa, the British, fearing confrontation with the vastly more numerous
local populations, implemented gun control laws (Vail 1977). While largely successful in
sterilizing the chances of conflict, gun control, coupled with the protection of wildlife at sites
such as the Mweru Marsh Game Reserve, increased tsetse food supplies, and simultaneously
increased the number of reservoirs for the parasite. With infected wildlife existing in greater
numbers and the means for checking their growth becoming increasingly limited,
trypanosomiasis was able to sweep across eastern Zambia with ease in the early 1900s.
Furthermore, as early as 1898, the North-East Rhodesia administration had been imposing a five
shilling hut tax within its villages (Vail 1977). Unable to pay the tax due to limited employment
opportunities, the men of North-East Rhodesia often journeyed, and were recruited, to Southern
Rhodesia where employment opportunities were more widespread, further reducing the available
labor pool most capable of managing wildlife populations.

2.3.3 Tsetse Habitat Destruction
The discovery that many sleeping sickness cases existed along the shores of rivers and lakes
led to discussions of tsetse habitat clearing. In Bruce (1908), the distribution of sleeping sickness
cases was presented to the Royal African Society: along the west coast of Africa, along the
shores of Lake Tanganyika and Lake Victoria, in parts of western Uganda, and at Wadelai on the
Nile. Additionally, these sites were shown to coincide with the existence of G. palpalis. With
such reports clearly locating the presence of sleeping sickness to such confined areas inhabited

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by G. palpalis, bush clearing practices became more frequent, and often were employed
alongside game destruction to curb sleeping sickness cases caused by the more dispersed G.
morsitans (Harcourt 1912).
Shrub and bush clearing occurred around Central and East African lakes in the early 1900s,
and these practices were met with some success; however, the projects were often costly and
required the movement of populations (Harcourt 1912). In Bruce (1908) the daunting task of
habitat destruction was made clear:
If we picture the hundreds of miles of lake and island shore, with huge trees and dense
undergrowth up to the water’s edge, we must come to the conclusion that the wholesale
destruction of the fly is impossible.
Soff (1969) highlighted the inefficiencies often found in bush clearing when recounting
Ugandan Protectorate’s Governor Hesketh Bell’s belief that “all bush harbored tsetse.” Such
views frequently led to total destruction of lakeshore vegetation when, in reality, only certain
species of bush, based on physiological structure, provide habitat for the fly. Additionally,
unless the land was populated after clearing or the bush kept from regenerating, the tsetse fly
would return. As a result, habitat destruction typically included the encouragement of villagers
to live closer together, as their routines of building, collecting firewood, and practicing
agriculture would discourage the return of tsetse habitat (Fight Tsetse Fly 1927). The increase in
agriculture that followed the process of concentrating villagers was promoted as an opportunity
to increase the standard of living for those in the fly belts. This in turn was believed to have a
positive cyclical effect: as the economic status of an area increased, so too would the public
health (Gilks 1935).

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2)/3,%87##!6)#'&-#1(.-4&8.#36)#-)%2)/3&5)#"0#3%))N2">)%#&2%"..#36)#2"/3(/)/3#GO",%2)<#P)=%().#
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Despite its inefficiencies, the practice of habitat destruction, when coupled with additional
control techniques, has proven to be very effective at combating the tsetse fly (Hargrove 2003).
Moreover, the practice of removing all habitat believed to be harboring tsetse, employed in the
infancy of tsetse habitat removal, was discovered to be only one of several effective forms of
habitat clearing, with the others (i.e., partial, selective, and discriminative clearing) being far less
damaging. As an example, using a discriminative clearing scheme where only the woody
vegetation from a plant community was removed, Mbala, Zambia (then Abercorn, Northern
Rhodesia) was able to effectively control the tsetse fly in the early 1950s after more than a
decade’s worth of discriminative clearing (Figure 2.1). Remarkably, this achievement resulted
2

from clearing only 3 percent of the vegetation in a 725 km area (Hargrove 2003). Other
successes achieved by discriminative clearing have occurred in Nigeria and Ghana (Hargrove
2003).

2.3.3.1 Interference of Colonial Policies in Destroying Tsetse Habitat
As stated above, the success of habitat removal requires the settlement or use of an area to
discourage the vegetation from re-growing. During the colonial period, continuous and intensive
use of the land was often hampered by official policies (Vail 1977; Musambachime 1981). The
unfortunate result was the expansion of tsetse habitat during the early twentieth century and the
continued pervasiveness of sleeping sickness cases.
It has already been demonstrated that man and tsetse coexisted for centuries in Africa prior
to the arrival of Europeans. Kjekshus (1977), among others, suggests that tsetse were cleared
from areas leading to tsetse-free zones. Such zones were created by the intensive use of land,
which eliminated the fly’s preferred habitat. Unfortunately, several of the European policies

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already mentioned as allowing reservoir host populations to increase also led to tsetse habitat
expansion; the hut tax is one such policy. With the imposition of the North-East Rhodesia hut tax
in 1898, men moved to areas offering better employment opportunities (Southern Rhodesia),
and, consequently, land management of the fields left behind declined (Vail 1977). Accordingly,
agricultural output decreased and the fields were allowed to revert to habitat conducive to the fly.
The conclusion of World War I provided another opportunity for the fly to reclaim lost
habitat. In the early 1920s, the administration of North-East Rhodesia was eager to move
Africans off from the most fertile lands in order to attract European settlers intent on growing
2

tobacco. This resulted in approximately 3,500 mi of inferior land set aside for African reserves,
2

and 6,500 mi of the most fertile land allocated to Europeans and those settlers expected to
follow (Vail 1977). The collapse of the tobacco market in the 1920s discouraged settlers from
moving to the ‘European’ land, and after a fallow period the area was reclaimed by the tsetse fly.
Sleeping sickness then became a greater risk for the Africans living in the nearby reserves and in
one village a death rate of sixty-six per thousand was reported each year throughout the 1930s
(Vail 1977).

2.4 Post World War II Tsetse and Trypanosomiasis Control Practices
Contemporary control practices have been implemented since WWII, which include ground
and aerial spraying of tsetse habitat with insecticides, the sterile insect technique (SIT), and the
point-source techniques of traps, targets, and insecticide-treated cattle.

2.4.1 Insecticides to Control Vectors

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In 1874, DDT was first synthesized. It was not until 1936, however, that its insecticidal
properties were realized, and not until the conclusion of WWII were insecticides widely used to
control vector populations (Garnham 1967). With the use of insecticides, vector eradication (i.e.,
the complete removal of all wild populations of a species) finally seemed possible, and critically
the process by which it could be achieved was often easier than previous control measures
(Garnham 1967). In DDT Can Wipe Out Plagues (1945), optimism was expressed that DDT
could send disease-carrying insects to “join the dodo and the dinosaur,” and thereby end “these
particular plagues for all time.”

2.4.1.1 Ground Spraying
Application of insecticides initially occurred most commonly in the form of ground spraying
(Allsopp 2001). These operations typically included large teams of trained staff dispatched over
the control area. The staff, consisting of control officers and laborers equipped with pressurized
and non-pressurized sprayers carried on their backs, applied insecticides to vegetation frequented
by tsetse. Spraying operations could only be successful if the control area was made
uninhabitable for both the adult flies and the tsetse puparia buried in the soil (Jordan 1986). This
was, and still is, achieved in one of two ways: through the use of residual insecticides that remain
lethal long enough to control tsetse after they have emerged from the puparia (i.e., about 22 to 25
days) (Hargrove 2003), or through reapplication of a non-residual insecticide.
Currently, ground spraying is used infrequently due to its high costs, dependence on large
numbers of well-trained technicians, susceptibility to reinvasion, and regular dependence on
residual insecticides (Hargrove 2003). In fact, the presence of residual insecticides in West
Africa in the late 1970s and early 1980s led to a decline in aquatic arthropod populations (FAO

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1992). Other creatures that saw population numbers drop after sprayings included the planthoppers and silverfish in Zimbabwe at the beginning of the 1990s and the little bee-eaters during
the 1980s (FAO 1992). Due in no small part to the devastating effects it had on non-target
species, such indiscriminate, high-dose sprayings have largely been replaced by more selective,
low-dose sprayings.

2.4.1.2 Aerial Spraying
In the mid and late 1940s, South African Air Force pilots flying over Zululand, South Africa
participated in the first widespread aerial spraying of tsetse habitat (DDT War on African Flies
2

1947) (Figure 2.1). During the operation, pilots applied DDT to an area of 100 mi , and ground
teams set off DDT grenades to target habitat missed by the aerial spraying. The campaign
effectively controlled G. pallidipes in the sprayed area (Hargrove 2003), and optimism that the
fly could be removed from the continent began to grow. Unfortunately, limited funding,
environmental concerns, poor coordination between countries, and the ability and efficiency of
the fly reinvading cleared areas have limited the successes of control efforts (PATTEC 2001).
The large scale 1973 to 1991 control campaign over Botswana’s Okavango Delta is one such
example (Figure 2.1). During this operation, extensive and repeated aerial sprayings occurred
over the vast delta, but in order to entirely remove any opportunity for reinvasion, pilots also
needed to spray the portion of the fly belt that extended into neighboring Namibia (Hargrove
2003). Unfortunately, permission was not granted for such cross-boundary flights, and a corridor
of reinvasion was made available to the fly. The inability to spray along the Namibian border,
however, was by no means the sole culprit for the operation’s ultimate failure. When conducting
aerial sprayings it is critical that the spray zones are of sufficient size and that barriers are erected

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to inhibit the fly’s movement between the successively sprayed zones. In the Okavango Delta
campaign, spray zones were far too small and movement between sprayings was not impeded to
any great degree allowing the fly to avoid treatments (Hargrove 2003). In fact, spray blocks
were so small that, over the course of a year, approximately 30 percent of a spray zone’s tsetse
population was capable of avoiding treatments by passing into successively sprayed areas
(Hargrove 2003). Thus, despite nearly two decades of active suppression, fly invasions into the
sprayed areas and the persistence of flies already in the control region continued the Okavango
Delta’s tsetse-infestation. In 2001, determined to eliminate the fly from the Delta, aerial
spraying operations were renewed (Figure 2.1).
2

The second spraying operation, a campaign spanning 16,000 km , took place during two
periods (Torr, Hargrove, and Vale 2005). The first treatment occurred in the northern half of the
Delta from June to September 2001, and the second treatment in the southern half from May to
August 2002 (Kgori, Modo, and Torr 2006). The sheer size of the separate spray regions was an
improvement upon the 1973 to 1991 operation, as it was much more difficult for flies to seek
refuge in untreated areas. Hargrove (2003) discussed an experiment that found female tsetse
flies capable of traveling 1,000 meters per day, but even a fly traveling at this rate would have
had difficulty reaching the Delta’s untreated area. What is more, the 2001 to 2002 campaign
importantly included a barrier of 12,000 deltamethrin-treated targets to separate the northern
spray zone from the southern zone (Kgori, Modo, and Torr 2006). Since the completion of this
operation, no tsetse flies have been caught in the Delta, allowing for the claim that the Okavango
Delta is now tsetse-free. Such claims must be made with great caution, though, since detecting
and trapping fly populations at low densities is difficult and inefficient. Nevertheless, the 2001 to
2002 campaign in the Okavango Delta demonstrates how improvements can be made to previous

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control efforts, and how the fly’s ability to reinvade or persist in sprayed regions can be
challenged.

2.4.2 The Sterile Insect Technique
The sterile insect technique (SIT) is another large-scale and internationally popular method
for control. This technique, which relies on the use of radiation to sterilize male flies, has been
advanced as a less destructive means of achieving control since only the target species is harmed
(SIT, however, is used in conjunction with other techniques that may cause damage to non-target
species). In the 1950s, the release of artificially sterilized male screw-worm flies over the island
of Curacao successfully eliminated the screw-worm from the island and led to the use of
sterilized male flies to eliminate the pest from the southern United States (Simpson 1958).
Elsewhere, SIT eradicated the melon fly from Okinawa and the Mediterranean fruit fly from
Mexico, Chile, and southern Peru (Townson 2009). Such successes naturally spurred interest in
the use of the sterilized male technique to confront the tsetse fly. Simpson (1958) suggested that
the sterilized male technique would be most effective when eliminating a low-density population
especially after the area had been treated with insecticide. The tsetse fly particularly lends itself
to SIT due to its unusual reproductive behavior. Female flies rarely mate more than once during
the course of their lives; in fact, they vigorously resist males after they have once mated (Jordan
1986). Therefore, by exploiting this mating practice, fly populations can be sent crashing as
female flies increasingly mate with sterile males.
In Tanga, Tanzania in the 1970s, researchers supported by the U.S. Agency for International
Development (USAID) set up a “fly factory” that produced thousands of sterile male tsetse flies
per week. In this operation, unhatched male pupae were sterilized with small doses of Cesium

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137, and then released, sometimes 10,000 each week, into the wild where it was determined that
the sterilized males led to a decrease in the tsetse population (Broad 1978). More recently, the
island of Zanzibar was declared tsetse-free in 1997 following the release of nearly 8 million
sterile male flies over the island from 1994 to 1997 (Figure 2.1) (Tsetse fly eliminated on
Zanzibar 1998). Other countries have also explored SIT, such as Ethiopia which recently spent
roughly $12 million on a “fly factory” (Enserink 2007) expected to assist in tsetse eradication
2

from 25,000 km of Ethiopia’s Southern Rift Valley by 2017 (Figure 2.1).
The costs of SIT are a barrier to implementation. In the successful use of sterile males over
Zanzibar, nearly $6 million was spent from 1994 to 1997. This figure, while large, does not
include the costs of establishing a facility to rear the sterile flies, nor does it include the costs of
previous suppression work on the island (Molyneux 2001). Furthermore, Zanzibar, as an island,
is an anomaly due to its natural barriers to reinvasion and presence of only one tsetse species. On
the African mainland, SIT success is questionable with few natural barriers to reinvasion and the
frequent presence of several tsetse species in the same area. The rearing of more than one sterile
species, which would be required across much of Africa, for any SIT campaign would lead to a
substantial increase in costs (Hargrove 2003). Furthermore, SIT is only successful if the sterile
males outnumber wild males ten to one (Enserink 2007), and in order to achieve this, traditional
insecticidal techniques are still required. On the island of Zanzibar, the wild tsetse population
had to be suppressed by 90 percent using traps and other techniques before SIT could achieve its
goal. This has caused some skeptics to suggest that continued use of the control technique that
has achieved 90 percent control should be able to clear the remaining flies (Rogers and Randolph
2002). In this way, the economic burden of rearing sterile males would be avoided.

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2.5 Point-Source Techniques
As mentioned above, the apparent success of the control effort in the Okavango Delta was
largely due to the use of barriers to restrict reinvasion of sprayed areas. In that operation, tsetse
attempts to reinvade the northern zone after treatment were prevented by a barrier of targets
(screens sprayed with insecticides) set up between the two spray zones. This method of using
targets and traps (the target’s 3-dimensional counterpart; Figure 2.2) to impede the fly’s
reinvasion efforts has frequently been used alongside large-scale control efforts (IAEA 1997;
Kuzoe and Schofield 2004). More recently, traps and targets have been used alone due to the
rise of community participation in control campaigns. These two devices as well as insecticidetreated cattle constitute the point-source techniques. The success of point-source techniques
hinges on their ability to provide an attractive visual and/or aromatic stimulant to lure in the fly.

2.5.1 Visual and Aromatic Attractants for Traps and Targets
The use of visual stimuli to attract tsetse to control devices has taken a variety of creative
forms since the early 1900s. Several of these traps include the animal trap by Morris and Morris
(Jordan 1986) and the “moving staircase” trap (Swynnerton 1933). However, the first successful
use of visual stimuli to capture tsetse quite possibly took place on the island of Principe, off the
west coast of Africa (Figure 2.1). Bulhões Maldonado, the estate manager of a cocoa and coffee
plantation on the island, noticed that G. p. palpalis were attracted to the backs of the plantation’s
laborers (Maldonado 1910). In an innovative strategy to limit the fly’s presence in and around
the plantation, Mr. Maldonado ordered the laborers to wear black cloths covered with a
“glutinous substance” on their backs. The strategy was successful: between April 1906 and the

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end of 1907, 133,778 tsetse were captured (Maldonado 1910), and in the process, the use of
visual stimuli to attract tsetse to control devices was born.

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The visual responses of tsetse had not been extensively studied at the time of Mr.
Maldonado’s control campaign; however, according to Steverding and Troscianko (2004), the
tsetse fly searches for shaded areas when seeking daytime resting sites. Thus, by using dark

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cloths to attract the flies, the laborers were effectively mimicking the darkness of daytime
shadows. Today, nearly all trapping and targeting devices rely on black and blue surfaces to
provide a visual attractant for the fly.
In terms of the aromatic attractants used when deploying traps and targets, a progression of
ideas similar to the development of visual stimuli occurred. In the 1930s, observations that tsetse
were particularly attracted to traps baited with a live animal kept out of view of the fly indicated
that efficiency could be improved by using host scents (Kuzoe and Schofield 2004). In Charles
Swynnerton’s 1933 work, Some Traps for Tsetse-Flies, a list of potential host scents to act as
baits was recorded, including urine, dung, blood, and hides. Since discovering the potential
benefits of animal scents, much effort has gone into improving the effectiveness of attractive
odors. Today, odor attractants largely consist of acetone, octenol, and synthetic phenols, or
blends of these attractants, placed in sachets and attached to the trap or target (Leak, Ejigu, and
Vreysen 2008).

2.5.2 Differences between Traps and Targets
The dissimilarities between traps and targets, though subtle, may result in one device
preferred to the other depending on the goals and/or limitations of the control operation. As
stated earlier, targets are two-dimensional insecticide-impregnated screens utilizing visual and
aromatic stimuli to attract tsetse. Traps, on the other hand, are three-dimensional devices of
various shapes. And while both traps and targets utilize visual and aromatic attractants, traps do
not necessitate the use of insecticides since a cage is used to retain the attracted flies. Thus, there
are potential benefits to choosing one form of attractive device over the other. Ecologically, the
trap may be preferred due to the optional use of insecticides. From a cost perspective, however,

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Lindh et al. (2009) found a large savings when using targets to control fly populations.
Additionally, the simpler design of targets makes the task of maintenance much easier (Leak,
Ejigu, and Vreysen 2008). Despite the differences between traps and targets, they are both less
expensive and less environmentally damaging than many of their counterparts (Day and Sjogren
1994; Hargrove 2003), and combined with their favorable use in community-centered control
efforts, support for these devices is strong and growing.

2.5.3 Insecticide-Treated Cattle
The third form of point-source control, the deployment of insecticide-treated cattle, has also
witnessed a recent surge in popularity. In this method, also referred to as cattle dipping, cattle are
commonly sprayed with pyrethroids (Hargrove 2003) along the parts of the body where the tsetse
feed, typically the legs and belly (Torr, Hargrove, and Vale 2005). As a point-source control,
using the host is an inexpensive and simple alternative to trapping and targeting devices. This
method also finds allies in those advocating for smaller-scale, community-driven control efforts.
With cattle dipping, often the livestock owner, rather than the government or a donor agency, is
responsible for applying the insecticide and determining the frequency of applications (Torr,
Maudlin, and Vale 2007). Thus, the individuals investing in the spraying of the livestock are
also those receiving the direct benefit of control.
Like other tsetse control efforts the deployment of insecticide-treated cattle presents
efficiency challenges. The tendency of cattle to roam differentiates them from other point-source
methods of control. While traps, targets, and cattle can all be used at prescribed densities to
achieve control, the mobility of cattle does not ensure that the entire control area will be equally
served. In the Tanga Region of North-East Tanzania, such cattle mobility contributed to a failed

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control effort as the treated cattle did not penetrate areas of tsetse habitat, thereby allowing fly
populations to persist (Hargrove et al. 2000). In fact, the Tanga control effort demonstrated that
if tsetse are present in high densities, the use of insecticide-treated cattle is entirely ineffective
(Hargrove et al. 2000) as the cattle avoid these large residual populations of tsetse. From a
financial perspective, ability and willingness to pay may also become an issue when expecting
stockowners to purchase the insecticides. This financial burden was calculated to be roughly
$.20/animal/year (Torr, Maudlin, and Vale 2007), which could be a considerable obstacle in subSaharan Africa where it is not uncommon for 70 percent of a country’s population, or more, to
live on less than $1.25 per day (World Bank 2010).

2.5.4 Community Participation
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!
2.6 Trypanotolerant Cattle
A second livestock-centered method of “controlling” AAT within tsetse regions is the keeping
of trypanotolerant breeds. While such a technique does not rid an area of the tsetse fly, and, thus,
should not be considered a tsetse population control technique, it does present an insecticide-free
method by which land inhabited by tsetse can be put to productive use. Unfortunately, not all
cattle are trypanotolerant; resistance is found in the N’dama cattle of West Africa as well as a
number of breeds of dwarf West African shorthorn, including Lagune, Baoule, Simba, and
Muturu (Ormerod 1976). However, despite their ability to exist alongside tsetse, trypanotolerant
cattle are not often the first choice of many pastoralists due to their smaller relative size when
compared to zebu cattle (Rogers and Randolph 1988). In 2005, the Food and Agriculture
Organization (FAO), reporting on trypanotolerant livestock, summarized a 1979 study by the
International Livestock Centre for Africa, which disputed the widely-held belief by pastoralists

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that trypanotolerant livestock are substantially less productive than those that are
trypanosusceptible due to their smaller size. In fact, by constructing a productivity index,
created by combining several production and viability traits, the zebu were found to be only 7
percent more productive than the smaller trypanotolerant breeds under nominal disease burden
conditions (FAO 2005).
The ability of certain breeds of domestic livestock to remain productive in tsetse-infested
areas has long been recognized. Murray et al. (1984) listed a summary from 1906 that offered an
early account of the ability of West African cattle to survive in tsetse belts; however, Roberts and
Gray (1973) pointed out that the nature of the resistance, despite numerous studies, was not clear.
Today, more remains to be learned about the responses in trypanotolerant breeds, but evidence
exists that resistance is the result of at least two mechanisms. Specifically, non-hemopoietic
tissues are responsible for efficient control of parasitemia, the content of parasites in the blood,
and hemopoietic tissues allow for resistance to anemia in trypanotolerant cattle (Naessens, Teale,
and Sileghem 2002). This natural resistance has generated interest in the ability to cross the
trypanosomiasis-resistant phenotype into improved cattle. However, much work remains to be
done in this area as the mechanisms that allow trypanotolerant livestock to maintain their health
under challenging conditions are complex and may differ between trypanotolerant breeds (Black,
Seed, and Murphy 2001).

2.7 Control versus Eradication
The aforementioned methods of ground spraying, aerial spraying, and SIT comprise the
group of large-scale techniques. These were, and are, the methods that have given hope to the
idea of Africa-wide tsetse eradication, which is defined as it was earlier: The complete removal

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of all wild populations of a species. However, since the 1970s there has been an ongoing decline
in spending by African governments on tsetse and trypanosomiasis control (Hargrove 2000).
This decline in funding has partly been the result of a reduction in donor support due to
increasing donor concerns regarding the environmental consequences of large-scale control
efforts and impatience that has resulted from witnessing only minimal improvements from large
investments (Hargrove 2000). As donor and government support for control projects has waned,
community participation has become a common phrase in aid projects (Catley and Leyland
2001). Under this approach, direct involvement from those benefitting from the aid program is
encouraged. Predictably, this shift from government-centered or donor-centered control to local
control has also brought about a shift from large-scale to small-scale control techniques
(Hargrove 2000; Hargrove et al. 2000; Torr, Hargrove, and Vale 2005). The consequences of
this shift have yet to be fully realized, but Torr, Hargrove, and Vale (2005) offered that if
eradication of tsetse is to occur, a return to large-scale campaigns must occur.
The feasibility of eradication has been questioned by some (e.g., Molyneux 2001; Hargrove
2003; Torr, Hargrove, and Vale 2005), and outright refuted by others (Rogers and Randolph
2002), with issues of financial resources, coordination between countries, and fly reinvasion
raised as potential problem areas (Rogers and Randolph 2002). Still, there are others who claim
that eradication is the best solution, due to the heavy economic burden that tsetse exact in
endemic areas (e.g., Kabayo 2002; Kamuanga 2003). If eradication is to be achieved on the
African continent, the Pan-African Tsetse and Trypanosomiasis Eradication Campaign
(PATTEC) will need to play a large role.

2.7.1 PATTEC

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PATTEC is responsible for coordinating continent-wide tsetse eradication across political
boundaries, providing technical guidance to member countries, and obtaining financial and
material support when possible (PATTEC 2001). PATTEC includes such members as the FAO,
International Atomic Energy Agency (IAEA), and World Health Organization (WHO), as well as
the African Union Inter-African Bureau for Animal Resources (AU-IBAR) (Taverne 2001). It
promotes a variety of control methods, including aerial spraying and SIT, to attack tsetse in
distinct fly belts across the continent (Kabayo 2002). In a direct appeal to the donor community,
PATTEC (2001) offered that the environmental impact of all eradication campaigns will be
considered before implementation, and that, despite the substantial initial cost of the large-scale
efforts, eradication is a “once-and-for-all cost,” while control costs recur indefinitely. Even with
such announcements, the large initial cost of eradication may simply be too great for donors and
governments given the long legacy of eradication and control failures. Consequently, the
transition to cheaper, smaller, and more environmentally benign forms of control seems
irreversible (Brightwell et al. 2001).

2.8 Tsetse Ecology
Tsetse rely on the presence of suitable conditions for their survival. These conditions consist
of proper land cover types, climate conditions, and food sources (Pollock 1982a, 1982b).
Preferred conditions vary by tsetse species. However, regarding land cover types, irrespective of
the tsetse species considered, it is the geometry of the vegetation that makes land cover suitable,
including the vegetation’s ability to mitigate higher temperatures that may be damaging to the fly
(e.g., temperatures above 32°C for the morsitans group) (Cecchi et al. 2008; Leak, Ejigu, and
Vreysen 2008).

!
_c!

Climatically, depending on the species, temperatures between 19 and 28°C are preferred
with conditions ranging from very dry to highly humid (Challier 1982; Pollock 1982b). When
temperatures rise above the preferred levels, tsetse seek shelter that helps to mitigate the higher
temperatures (Leak 1999). As temperatures drop below the preferred levels, a “chill coma” sets
in, which prevents tsetse from flying and eventually leads to starvation (Knight 1971; Hargrove
1980; Terblanche et al. 2008).
Regarding food sources, tsetse take a blood meal from a host every two to three days
(Schofield and Torr 2002). Tsetse primarily feed on wild ungulates and ruminants, including the
warthog, bushpig, kudu, and bushbuck (Pollock 1982a; Jordan 1986). Tsetse also feed upon
livestock, including cattle, sheep, and goats.

2.8.1 Seasonal Fluctuations in Tsetse Distributions
Seasonal fluctuations of fly distributions have been extensively recognized and recorded
(e.g., Nash 1933; Bursell 1957; Leak 1999; Bett et al. 2008). As moisture levels, temperatures,
and the availability of food sources fluctuate, so too do tsetse distributions. In Kenya, these
fluctuations in fly distributions often show a bimodal or unimodal pattern corresponding with the
changing seasons (DeVisser et al. 2010). Generally, distributions contract during the hot dry
season, expand during the long rains, contract once again during the prolonged cool dry season,
and expand during the short rains. Refer to section 1.2 for a review of the timing of each of these
seasons.
Davies (1964) and Glover (1967) both recognized the importance of understanding the
ecological traits of tsetse and their responses to seasonal events. Both stressed the potential for
reduce fly control costs if these traits were understood. In this study, the seasonal fluctuations in

!
_M!

tsetse fly distributions will be examined to reduce tsetse management costs by controlling fly
populations when they are spatially and temporally constrained. This will be explained further in
the next chapter.

2.9 Costing Tsetse Control
Concern regarding the cost of tsetse control using the above described control techniques has
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3.1.1 Introducing Control Reservoirs and Tsetse Zones
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