1.

WORLD HEALTH ORGAN ISATION MONDIALE

Cl RGAN I EAT1 ON DE LA SANTE
WHO I"R-mGIONAL SEMINAR .... VC/ENT.
........ SEM/WP/4.65
..<.~, .. . . . .. . ., .
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ON ENTOMOLOGICAL METHODS I .
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IN .VECTOR CONTROL . . .
. . . .
ENGLISH ONLY
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USSR,' '20Sep-tembe'r. 16 October i.965.'.. . . . . . '-

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BIOLOGY AID CONTROL OF TSETSE FLIES . 8 ,

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....... .VI.
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.> ' . by- .

J. Hamon, P. Challier, J. Mouchet and J. Rageau,

Medical. Entomologists, ORSTOM, Paris9 France

1. INTRODUCTION

For centuries human and animal trypanosomiasis have ravaged vast areas of
tropical Africa. As recently as 20 years ago sleeping-sickness was considered one
of "ce most terrible of all endemic diseases affecting the African continent.
Animal trypanosomiasis render stock-raising difficult, if not impossible, in many
areas, and in cattle-grazing areas bordering on the Glossina zones, livestock is
weakened and does not supply any draught-animal (Vaucel et al., 1963; Wilson et al.,
I

1963; Ford, 1963).

Research efforts undertaken both in the laboratory and in the fSeld have supplied
in recent years bet$er.met@ods of prevention, d-etectionand treatment of trypano-
somiasis.
i

Human sleeping sickness prevalence has been reduced to an extremely low level,
but the disease is far from eradicated and many small foci either remain-activeor
appear in previously cleared areas. The situation is slightly better-in West
Africa, where the parasite Trypanosoma gambiense has apparently no animal reservoir,
than cin-&ast:)Afr.iga where occurs also T. rhodesiense with both human and animal h o s t s
(Robertson, 1963; Willett, 1963).

Glossina-borne animal trypanosomiasis are always very widespread and hinder

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economic development o f the majority of African states. Chemotherapy cannot be ex-

tended to all domestic animals and has its setbacks, including drug-resistance of the
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1- trypanosomes. So tsetse fly control has an important part to play in the devglopment
1
of the African continent. r
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2. GEOGRAPHIC DISTRIBU~IÓN
OF TSETSE FLIES ,
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2
Glossina sp. in?est about 10 O00 O00 ltm on t h e African mainland. Their present
d i s t r i b u t i o n has bean r e c e n t l y summarized by Ford (1963) but more sccurate d a t a f o r .,
. . I

French-speaking s t a t e s of West Africa can be found i n Rickenbach (1961), Maillo’t’ (1952)>

Rageau 8s Adam (1953) and F i n e l l ë e t a l . (1963).
Tsetse f l i e s a r e c l a s s i f i e d i n t o three groups t y p i f i e d r e s p e c t l v e l y by G. p a l p a l i s ,
G. morsitans and G. fusca.
--Y_
IIU-

2.1 G. p a l p a l i s group

The p a l p a l i s group includes f i v e species and four sub-species and i s r e s t r i c t e d

t o West and Central Africa, wfth r e l i c s i n Ethiopia and Arabia.

-G. tachinoides occurs i n t h e Sudanese savannah anci i n t h e southern p a r t of t h e

Sahelian area o f West Africa with r e s i d u a l f o c i i n North-Eastern Africa, and can with-
stand very dry conditions but congregates i n t h e dry season around water holes, r e s i -
dual pools of temporary r i v e r s , and so on.

G . p a l p a l i s and G. f u s c i p e s occur i n t h e m f o r e s tas well a s i n t h e Guinean and

--.,I_-
Sudalnese savannahs, but i n these l a s t environments they a r e r e s t r i c t e d t o r i v e r s and
rLvu.lqts i n ‘che neighbourhood.

----
G , p a l l i c e r a and G . caliginea occur i n high f o r e s t areas, but t h e f i r s t can be
encountered sometires i n t h e g a l l e r y f o r e s t s of t h e Guinean savannah.
8 ‘ ~

2.2 G. morsitans group

I

The m o r s i t a s group includes seven species and t h r e e sub-species, a l l inhabiting

savannahs.

-
G. m o r s i t 2 complex occurs mainly i n Sudanese savannahs, with G. submorsitans i n
West ard Central Africa and -.-
G. morsitans i n East Africa.

G. 1ongiQalpis i s r e s t r i c t e d t o Guifiean savannahs of West Africa.

G. a u s t e n i i s mainly a c o a s t a l spectes of EasL Africa.

G. swynnertoni and G. p a l l i d i p e s are highland species o f East Africa, t h e f i r s t

being r e s t r i c t e d t o Kenya and Tanzania and t h e second species occurring from Ethiopia
t o Mozambique and being prasent i n some c o a s t a l a r e a s o
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f h 4

8. borgesi .has. been ,re:cently.de.scribed .from

. . Mozambique.
. . . . . . . . .
, * ' ...... . / <

2.3 ' G. fusca group ' '' . ... I

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.... _ . /

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~: .. .. . . ." . . :, . . . . . I
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.,.
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The fusca group includes 12 species and two sub-species, which i n h a b i t mainiy
.,;, . . . .
densely f o r e s t e d areas.
... ,." .. . . ... . . . .....
. .
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G. nashi, G. schwetzi, G. tabaniformis, G. haningtoni, G. Severini, G. va&oofi'

;, . ~ ( . t i.

and G. f u s c i p l e u r i s occur i n t h e Lower Guinean f o r e s t , sometimes i n very' r e s t r . i c t e d

areas. %*.
'. \..."ar. :, -,-
J a
"..".
.!) . .
k; fusca occurs i n " a l 1 forested areas, from Siel-ra Leone-to Uganda,: and G. n
fusca is widespread i n t h e Upper Guinean f o r e s t ' and i n t h e northern p a r t of 'che' Lower
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Guinean f o r e s t , from Guinea t o Uganda.

, <
G. medicorum i s r e s t r i c t e d t o Upper Guinean savannah and f o r e s t .

G. brevipalpis i s very widespread i n g a l l e r y f o r e s t s and savannahs of East- AfPica

from Natal t o Eastern Congo (Léopoldville) and t o Southern Ethiopia and Somalia.
..... . . ~. . . . .. .,. ,
-.. . .......-.,... . _I.__ . . . . . . . ._-.-..-.I. . . ... __.,.
"..-....... . . . !
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G. longipennis, i. s .a l s o an e a s t e r n . savannah
. species,
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but^-.re-s~riicFe-d~".to
, . . .. Kë-riyä;-.-
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Ethiopia, Southern Somalia, South-Eastern Sudan, North-Western Uganda' and N

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Tanzania.
i ~. .... ... . . . .. . .. .. . .
~
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3. '. *TsETsE.-."FLIE.s
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...... _.-...._. . .I .. . - ,..

3.1 Reproduction
. . cycle .
I : : ,I. . ..
*..... .. . . . . .! -__,
I - -_._ -..
1 .

Tsetse f l i e s mate. during the.,day<s,following: emergence, when

One inseminat,ion i s s u f f , i c i c n t f o r t h e w h o l e , l i f e of t h e female,
-. c. I , ' , .
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times during t h e i r l i f e and perhaps some females do t h e same. N
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females a r e always inseminated. Homologous matings a r e t h e r u l e but 'sometimes mating

occurs i n nakure 'between closely'.'rel;zted:'specie's; such matings' a r e g e n . e r a ~ l ~ , , . . ~ ~ t ' . , f e " r .
t i l e and .can 'be lethai.:.,4,or,
the.:female. . - ' .; i..

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The two ovaries are':'composed ' o f two' ovaeidles each,. 2nd each ovar?ole develops i n
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.Challier, 1963b);
turn (Saunders, 1962;::. :. ! The...first ovul.ation occurs t h r e e. t. o , 11 days
a f t e r t h e emergence of the fanale; ., , .-the. mature o:vo,cgte. i s . n o t. .. Laid, but passes. into, I

. . . .uterus.
t h e female .. where fecundation, oc,curs, then,.,the.;
..n/..
......-......- .
l a r v a s o w s inside t h e. uterus.,,
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the food bein6 .supplied:!by.

..-.~.-. ..- the. "milk glands" . the female; the. l. a .t e . t h i r d i,n$ar..
/....-........... _ . * .........
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.-
@@p%&@.
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~ @Fj ~ ~ I ~ ~ J ~ ~ ~
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l a r v a is l a i d on t h e ground and pupates about two t o f i v e centimetres below t h e sur-

face, some hours l a t e r . -.~.. during
Larval development,":"in utero, 'X&l<e.S;-ab;out10 days, . . . .which . /

. l a. i d when the, female i s

t h e female b i t e s a t l e a s t three.times, but t h e f i r . & l a r v a:.-.L s ..:. ~

- .
about 16 days old. When t h e f i r s t l a r v a i s l a i d t h e second ovOcyte passes i n t o t h e
uterus, and s o on. The duration of t h e pupal l i f e v a r i e s from 20 t o 60. days,. acdosding
t o temperature. . . .

The a d u l t can f l y some hours a f t e r i t s emergence from t h e pupàriwn, and i s abl&'t¿?

b i t e tha..n.fallo.wing.day )... . . . . .. . . . .
..................... . .~ .. -..

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3.2 Feeding h a b i t s ..

Both sexes of t s e t s e f l i e s suck blood, and they do not use any other food. They
, .- .. .!.L!
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do nöt?TÏïgSSt usually e i t h a r water o r n e c t a r but seem sometimes able t o pieree;;j32ants
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for sucking sap.

Each species of t s e t s e f l y e x h i b i t s defini-te feeding pref?rences, but t h e majority

of Glossina a r e able ' t o b i t e 'a g r e a t v a r i e t y ' o f v e r t e b r a t e s . The known tF0phi.c..pre.-
ferehdës a r e - t h e following (Weitz, 1963) according t o 22 640 blood meals .i".d a .......
n t i f i e d.: . ^"
I-...

G. swynnertoni, G. austeni, G . f u s c i p l e u r i s and G. tabaniformis feed mainly on suids,

............ .secondary
with bovîds and other v e r t e b r a t e s (rhinoceros; .hippopotamus, porcupine).as . . .
hosts. ' . . . . i. .: . '',,'< :,. .

G. morsitans s.1. feeds equally on suids and bovids, warthog (Phacochoerus sp.) being
t h e preferred host; man i s a l s o used as host, mainly by submorsitans of West Africa.

G . p a l l i d i p e s , G. fusca and G. l o n g i p a l p i s feed extensively on bushbuck (Tragelaphus

s c r i p t u s ) , and t h e f i r s t and second species b i t e a l s o suids t o some extent.

G. longipennis feeds mainly on rhinoceros, with g i r a f f e , elephant, buffalo and o s t r i c h

a s secondary hosts.

G. brevipalpis feeds mainly on hippopotamuses, suids, buffaloes and bushbuck.

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G. p a l p a l i s , G. fuscipes and G. tachinoides feed extensively on man, crocodiles, l i z a r d s

and bovids, t h e l a s t species being p a r t i c u l a r l y infeodated t o man and c a t t l e .

The behaviour of each species i s c h a r a c t e r i s t , i c and i s not e n t i r e l y dependant on

t ,

t h e availabil.itv; of d i f f e r e n t hosts; t h i s view i s supported by t h e f a c t t h a t commonly

occurring,aQim a r e not fed on by Glossina, t h e best example being zebra (Equus
b u r c h e l l i ) i n G. morsitans areas of East Africa, and waterbucks (Kobus sp.). * I
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-,..-.... .<.?.
Few data are ávaïlable about the::.gaaQt-abi&ity of.....tsetse :,flies
". to-a yasying...ho&,,. :;

fauna. It i8'?P%kelyth& flies like G. morsitans s.1.; the :G: palpalis group and.
possibly G. swynnërtoni and G.. .pallidipeswould quickly adapt to changes in fauna.
The anthropophily of G. palpalis alld. G . tacliinoides increases when the wild.fauna and
the livestock decrease (Page Be McDonald, 1-959; Jordan et al., 1961 and 1962; ,.
......., .- . . . .. ..
Langridge et
..
al., 1963).
...
However some species with very restricted feeding habits,
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- G. austeni and G. tabaniformis'(infeodated to"suids),
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such as G. fuscipleuris,
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G. longipalpis and.G. fusca (infeodated to bushbuck and buffalo). and G. longiperhs

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(rhinoceros, elephant, buffalo), may be very rapidly affected by the remova1"'ofonly '

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a few species.

3.3 Ecology

The subject ,,as been recently reviewed by Langridge et al., (1963) and by Marley,

3.3.1 Adult flies

Each species of tset'se '.flyhas a_specxfic distribution

. . . . . . . . . . . . . .=ea,
.. characterized by
climate, vegetation and soil, this last element being important.for.larva1deposit and..
pupal survival. ,
, . .
... ... ,,.-.. --.. .... ........ ..-
I... ...................
In each .distributionarea it is often
. . possible,
. . . to;,distinguishthree.,zones (Mouchet

et al., 1961): .,: k '

. . . . . . . . . . .
. . . . . .

q
the 'permanentbreedingtsites, where flies occur all year round and where they
' ' congregate in unfavo'urable seasons, and iusually where they lay larvae; ;

the temporary breeding-sites, where flies occur during the m o s i favourable

usually in the rain:: season;hut where they cannot surv5ve in the ,dry
season; ' . . . . . . .. : .... ....

thé . ~ ' ~ e e ~ ï n ~ - g r o u n'are

~ ~ -generally
. w h ~ c ~ open. areas," not v a r y far'from the breeding-
. .
sites, where flies'can easily detect their hosts, and feed upon them; ,fecd%ng-
~..- .
grounds can extend a few kilometres around breeding-sites, but are usually re-
,.
stricted to their immediate vicinity. I

In each zone resting-places of the flies vary according to environmental condi-

tions and physiological condition of the fly, but they are usually constituted by
poles, small trunks and medium sized branches, up to some metres above the ground.
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But some species,.like. G. swynnertoni and G. pallidipes rest extensively on leaves by

night and on the underside of branches by day (Harley, 1960-1963). In wooded savannah
- . ... . . . .~~.,
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areas some species of trees afford. better resting-places than others and are regularly
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selected by the majority of flies.

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3.3.2 Larvae and pupae

The depth to which the larvae enter the soil varies with environmental conditions;
ìn the rainy season the larva can pupate almost on the ground or only one o r two centi-
metres-below.’’ In the dry season the larva can enter five to eight centimetres belaw
the ground before pupating.

The pupae cannot move and so are susceptible to wide variations of temperature
and humidity. They can withstand high relative humidities but neither submersion by
water nor dryness (Bursell, 1963). High temperatures reduce the length of pupal Life,
but too high temperatures can k i l l the pupae in some days and even some hours, and
pupal sites are usually situated in well-shaded places, under logs, bushes, and so on.
.”- - _” -
For some species, as G. tachinoides and G. palpalis, breeding-sites c.an be entirely
man-made, such as mango and banana plantations.

Tsetse fly survival and death-causes . .

3.4
Adults of Glossina have a high survival rate, in the laboratory as in nature;
survival for 154 dags has been observed in G. palpalis, and average lives of two to
three months seem to be the rule. Flies can die from aging, but also from starvation,
unfavourable environment changes, or be killed by predators (birds, spiders, insects)
or, more rarely, by parasite fungus and protozoa.

Causes of mortality of larvae and pupae are more important because predators are
numerous; insectivorous birds, insectivorous insects, ants, and so on. Many insects
parasite tsetse pupae; Diptera bombylidae, Hymenoptera chalcididae and Mutillidae.
Pupae are also much more easily killed by unfavourable environment changes than adult
flies which can move away, In nature probably less than half the laid larvae reach
the adult stage.
4. i TSETSE FLY, CONTROL :. 'I '
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In-inmy iisiancks tsetse cbntrd++s-.a very efficiènt way to stop ;Gan trypano- -'

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somiasis transmission in restricied areas as a complement to chemotherapic 'measures,.'
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and tsetse control o r eradicatlun'ia often the clleapast illethod for preventini livestock '
trypanosomiasis transmiss'ionin''cattle-grazingareas of Sudanese and Guinean'. Savannah
, ,. * .
i . 1963).-.. - . ' . ..
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zb'&s'.',(&,cki& et .. .. . . ". ./ - ' ' .-..d ,

, 3 .
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Metbods
:. used for.tsetse fly control just before and after the second'W o r l d War ..
.i.:. , ' I

. .. . . .being replaced by other ones based on residual insecticide applications,

are .graduallx
.
. . . ..
and ,chemosterili,zation
.. is being investigated.
I But in the past direct destruction,
indiscriminate
.. '. .
I.. and.sele,ctivzclearings, and hosts elimination have been extensively
used, sometimes with fair success (Glover, 1951).

4.1 Direct destruction of tsetse flies

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Direct destruction of tsetse flie,sby net collection and glued baits has not been
very promising in the past, except in Princi Island where G. palpalis was almost
eradicated. Destruction by trapping (Harris traps) has been successful in controlling
G. pallidipes ia a restricted breeding area of Zululand, South Africa, but results have
not been very satisfactory in other areas. Some authors, like Morris, (1960-1961)
stress that traps give the best way for catching representative samples of flies,
_.
whereas others, like Abed:, (1963), consider traps as useless.

4.2 Tsets,e,fly control by game destruction

It.has been observed in the past that tsetse flies and game are often associated,
and that game elimination, during rinderpest epidemics for example, are followed by
strong reduction or even disappearance of 'Glossina populations. So game destruction
has been used as a routine method of tsetse fly control in Eastern and South-Eastern
Africa., and has succeeded in clearing many thousands of square kilometres of flies
belonging to the - - group, the best known experiment being the Shinyanga one,
morsitans
in Tanganyika (Glasgow, 1960). The method was not cheap; for example in Southern
Rhodesia, during the year 1955, slightly more than 41 O00 head of game were destroyed
by 800 paid hunters, with an expenditure of lo7 O00 rounds of 3
ition (Chorley, ,

1956) - I

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Judiciously applied game destruction can be an economical and practical means sf

trypanosomiasis control, as not only hosts of Glossina, but also animal reservoirs of
trypanosomes are destroyed in only one operation. However, as stressed by Weitz,
(1963) and by Glover, (1364), the -palpalis
-- group of tsetse flies cannot be starved
except if reptiles and humans a l s o are eradicated, and the control by starvation o f
3 ,

many tsetse species (such as swynnertoni, austeni, fuscipleuris, tabaniformis, morsitans

'
and submorsitans) supposes total elimination of suids, which constitute a group more
difficult t o locate and to kill than antelopes and big game. Besides, as underlined
by Dasmann,' (1962) and Hocking ,?t al., (1963), game farming can be more profitable in
some marginal areas than cattle-raising as a source of meat and skins and also for the
tourist' inc%.&my. So tsetse control by game destruction is almost entirely abandoned
now.

4.3 Tsetse fly control by bush clearing . .. .. , * ......, .

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Bushes and trees are widely used by tsetse flies as resting-places, but they are
. , . .
vital as shelters *kor pupae. Pupae can only survive if the soil is sufficiently
; , I , ..i,: ;,'y, I. 1;;
sh&ed, prdtected from kirect insolation and preserved from rapid variations of humidity
, 1.: ,.. .I

I..,.
... , "

and 'temperature* . .
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The clearing of all woody vegetation in a fly-infested area has rarely been used
for tsetse fly eradication, but has been extensively employed to provide barrier areas
or to reduce the contact between flies and humans around villages and water holes, at
river-crossings, and along the main routes of communication of humans and cattle
(Le Rouzic, 1948). ' Such clearings, called "agronomical prophylaxis" in French-speaking
countrses, can be used f o r cultivation to be kept clear of trees and bushes. They
cannot be extended on large areas in sparsely inhabited zones because they require 8

larger manpower supply than avaflable, and are very costly if carried out by adininis-
trative agencies. However, f o r special purposes, clearings and protective barriers
have been done up to 100 kilometres long and three kilometres wide, as in Southern ,

Rhodesia a

Selectivn cleax-ing has been much more extensiv?ly used for tse-tsefly control.
It is based on the tendency of Tlies, in unfavourable seasons, to concentrate in per-
manent breeding or restiag sites that comprise identil'iable plant communities and a
comparatively small proportion of the bush or woodland as a whole. The requirements
of the common tsetse fly vectors of trypanosomiasis have been described and the types
of vegetation most favourable as 'refugesare known. They inust be 1ocated:before
tsetse fly control measures are uindertaken. Selective clearing can be done directly,
o r by use of chemicals; it suffers from the same deficiencies as total clearing; it
is a costly measure, the regrowth of bush must be kept down and it is much more effi-
cient in dry areas than in humid-ones.

dry cauntPieS%emporary control ofIriverine tsetse flies can be got by use of

"obstructive clearing"; by felling the trees forming the overhead canopy and blocksing
the stream-bed to obstruct .the tsetse flight-line,." The flies cannot move fieelYiunder ,

shade and either are starved o r forced into the open where -the climate in the dry. ,

season is intolerable (Nash & Steiner, 1957).

With th'e."discovery of modern insecticides it appears that it is ver,y ofken. more

efficienk to spray a residual toxicant on the tsetse permanent restilqpsites than to
destrÓy them, because tsetse can change resting-sites when the preferred ones are des- . r
troyed,' but they do' not abandon them after spraying (Hocking, 1964). 'I

.
. . a
.
4.4 Tsetse fly control by insecticides'
_' ..
Insecticides can be used as aerosols,of
. . . . temporary efficacy on huge . .areas,
, or as
.... , ;

residual sprays on resting places, in tráps and on baits.

. .
............ . ... -........ "__*.I_. ........ I .
.( i l *

4.4.1 Insecticide
.).
aerosols
: ' 2
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Ipsectïcide aerosols have a-very short

. .
residua1,effectand ki1l:tsetse ',.: : . . .
flies dur- .,
.
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ing some~hours,only.after
their application. They can control
, . or eradicate
. . the.flies
only if they are applied on large areas at convenient
.
intervals,to
., .
. . . . . kill, before their
. . . 4 .
reproduction, all flies emerged from pupae since the previous application. A con-
. . . :. . .
venient rhythm see&
.....
to-be about cight app1ications"at''intervais"op'twô to four weeks
. . .
."' :. ! . ,. , i
' :
I _

. . ". i, ..,
................ , . , " ' . . . s .

(Burnett, lg62b) ....

.:. ._.. ~

: . .! ._ ':. - .<
. . . . . ..: ..
I .
. . . . . ,. . .. : i

1 . ;Ail?cr,aftapplicatiop,
, . . . .has.
. . . . the
. . obviqus,
. . advantage
. . of cqvering large areas qui
L .
, . . .
and can.be,.effic&ent
. >,aga@st.
. . s;avaiVah-inhabi,%ing
.
.. species
".
. , like morsitans,
, . swynnertoni
. . . I t I' .:
. I

and pallidipes; it,is.. less.adcqupte than

. < . . ,ground applica$ion aga,instriverine
.. tsetse. . . _ I I . I . I s.:.,;
, - . . .
flies.;like palpalis.;(Burnett,
.. :1962@$ ... a
. . . . ,:. I is 'almosti.ypossible . carry out against
. .t o ... 1 . .

in h,igh.forest..
specie.s-:living , . . .
. .
:.. . . . . . . . . .
. . I

. .
, . . I . . . . ..
: I
- . . , '. , : . I ! ' .. . . : .. , , .. ; I . _ *
!i.
I .
 . 65::,,
VC/ENT SEM/WP/IC.
page 10

Ground applicatiop can be done e i t h e r with lorry-carried generators, o r with l i g h t

generators, l i k e Swingfog (Challier z t a l . , 1964).

Insecticide aerosols can only Se applied some hours a day, j u s t a f t e r dawn and
bofore dusk, and sometimes by night, when a i r currents a r e downwards; during other
day-time periods the i n s e c t i c i d e cloud disperses very rapidly and, i f applied by a i r -
c r a f t , does not reach ground l e v e l . Only a very small amount o f i n s e c t i c i d e reaches
each individual f l y , and gravid females, l e s s susceptible t o i n s e c t i c i d e s than males
and. females of other physiological conditions (Burnett, 1962a), are not e a s i l y k i l l e d
by chlorinated insecticides; some OP compounds l i k e fenthion might perhaps be more
e f f i c i e n t (Hocking e t a l . , 1963).

for.^ eradication progrrames d i e l d r i n and 3HC .aerosols have been,used, maiFly i n

East Africa, with- v a r k b l e r e s u l t s including f a i r success (Cockbill e t al., 1963).
The cost o f . such programmes was very heavy i n the first experiments, but has decreased
with technical improvements and bette,r knowledge of t s e t s e ecology, and i s now around
2
800 t o 2000 French francs per km (Burnett, 1962; Hocking e t a l . , 1963; Burnett e t
1 . . ..
al., 1964). The cost of operations can Ise reduced i f aero$Öls"We used. o n l y :for short2
term control'.'&f" t s e t s e f l i e s i n sleeping-sickness f o c i .

4-4-2 Residual treatment of vegetation

Residual treatment of t s e t s e f l y resting-places must be l e t h a l f o r the f l y on

short contact f o r a longer period than the m a x i m u m duration of t h e pupal l i f e . In
such conditions only one spraying inay be s u f f i c i e n t t o control the species, and per-
haps eràdicate it, i n an i s o l a t e d area.

The f i r s t r e s i d u a l applications have been done against r i v e r i n e species, l i k e

p a l p a l i s and fuscipes, with h a b i t a t s r e s t r i c t e d t o water edge. In larger gallery
f o r e s t s it i s sometimes possible t o open paths i n the f o r e s t , which w i l l be exten-
s i v e l y used by moving f l i e s , and t o t r e a t thein f o r controlling f l i e s . DDT suspensions
and emulsions, which have been used i n the first experiments, have usually been r e -
placed by d i e l d r i n emulsions, which are assumed t o be e f f i c i e n t almost one year,,"a,Qd
sometimes more than one year i f applied a t 47; (Kernaghan, 1962). However, according
t o Baldry (1963 and 1964), DDT emulsions have a longer r e s i d u a l e f f e c t than d i e l d r i n
and t e l o d r i n (= isobenzan) when applied a t the concentration of 5% and even, i n some
conditions; at.lower conc&t*ations. Such control procedwis are'five"to 10'.times
I .
. .
cK&ap.er than bush clear%& 'And of a more permanent effect (Hocking,' 63;' Challieb', !" '

.. .. ,.!. .'
1965a; ha'cLennan& Aitchison, ,1963). . '
'

. .. ,-.~.
. . ' -. L

Improvement o n the knowledge of tsetse fly resting-places ha permitted the use

. .
of residual insecticides against savannah species like G . morsitans and G. swynnertoni,
, > .' ,.
by selective spraying of the lower side of the branches of Acacia trees in fly-
. <:
concentration areas (Hocking,'
. . 1961;
. Chadwick, 1964; ChadwTck et al., 1964)'. The
1. ,
. ... I

cost of the control

. .
is similar to or lower than by
,.
aircraft. foggipg but the work."
'
is I !.'
, ! '.. i '

more difficult to supervise.

Residual insecticides have been used also against G. tachinoides in Northern

Nigeria, and along Tchad-Cameroons border, but that fly is nof a centrated as
II

palpalis along water-places, and trees and bushes must be treated on a large width on
both sides of the rivers and water holes (Moucln'et et al., 1961). *'Theyhave also been
used with fair success against G. fusca and G. fuscipes in large gallery forests of
the Centrafrican Republic (Fine112 et al., 1962 and 1963; Yvore et al., 1962).

Tsetse fly control by residual insscticides has not been carried out against high
forest species and is only promising when the fly habitats are restricted.

4.4.3 Insecticide treated traps and baits

_. . 1

Residual insecticides have been sprayed inside tsetse fly,traps to increase their
efficacy with limi+ed results, Insecticides have also been used in combination with
attractants, on traps, and the method is always under investigation. ..- consti-
Cattle
tute natural attractants and both systematic and residual conventional insecticides -_ . I

applied to cattle are under experimentation for killing tsetse flies in restricted
areas where wild game is scarce or absent.

4.5 Biological control '

Various attempts have been ma& I I

in the past to multiply and release natural tsetse
fly parasites (Jenkins, 1964) to control these flies, but the results have not been
promising.

Trials on hybridization by cross-mating closely related species to produce a high

degree of st,erility in a fly-population have not been successful (Glasgow, 1960) and . a

assume the possibility of raising large numbers of tsetse flies.

 .
VC/ENT SEM[W~/~.65
page 12

Sterilization of flies is possible, either by radiation

. .
or with chemosterilantg.
(Knipling, 1963;. Smith .& Dame, 1963) and field and laboratory investigations
. have
. been
carried out. Tsetse fly males sterilized by gamma irradiation are not competitive
- . ... .,
with normal males, the sterilizing doses being nearly the same as the lethal ones
_.
(Potts, 1964). Chemosterilants are not much more promising; the treatment of males
'

, . '. .
aloriëY5ducës only by- 407; 'Xh '&veragc
.... . number of produced pupae) the treatment of
"
females is moTe efficient büt'.'i-i
seems.difficult to treat the female cömponent of wild
tsetse fly populations; the survTGal rate of treated individuals is considerably
reduced (Chadwick, 1964). If an efficient control method by chemosterilants is dis-
covered the problem of laboratory mass-rearing of tsetse flies would have to be solved,
and will"l?t*%e an.ea.syi.?ask. (Marillot, 1958; Nash, 1963; Evens, 1964).

5. DISCUSSION AND CONCLUSIONS

Several methods are available Por controlling tsetse flies, but all are expensive
and assume a good knowledge of the physiology and ecology of the species to assume
control under the local conditions. No method is yet available for densely forested
areas.
2
Surveys show that where human population density reaches 40 inhabitants per km
their animal hosts disappear, and their resting places are cleared; in such conditions
tsetse fly vectors of animal trypanosomiasis are usually no longer .a-problem-(Ford,
1962) (Bxcept if "holy woodlands" occur, like in the Mossi country of West Africa, but
such woodlands'can be easily cleared from tsetse flies by insecticides). R.iverine
species bf Glossina, which transmit mainly human sleeping-sickness caused by.
T. gambiense, can survive along rivers but are very easily controlled in such restricted
habitats. When human density is below 40 inhabitants per km2 tsetse fly control be-
comes more and more difficult and costlg with the scarcity of inhabitants. Despite
our technical ability to combat 'isetse flies, it is still not worthwhile to undertake
large reclamation schemes exc?pt in special instanczs, where soils and climate are
favourable to intensive agriculture as in the Lubu valley of Southem Rhodesia (Cockbill
-, 'T
et al., 1963) repopulated by inhabitants from the part of the Zambezi valley flooaed
7
:
b the Kariba Dam, when public health is involved such as in residual foci of sleeping-
sickness (Morri's, 1962), or to protect cattle during seasonal migrations (Finelle et
al., 1962-1963). I the$ situations insufficient exploitation of the country will
permit tsetse flies to repot&late the cleared areas sooner or later and the resources
employed for tsetse fly control will have been wasted.
 .-T&t.s.eLrsanpling, m,e.t"hods, A n n . Rpt . Liberian I n s t . American Founa.
.:.
... . .<,. . . . 1.962,; _ . q2-43 .. . ". ._
. ,,~-.:.:.:-.~~~--,:~~
---....____.._.._.__
.
.--
,A
. . .._. .
,, .
'
L;:. .. -.
.' . . . .... . c ....
, I '

. . . . . . ~.-.---....._:._L,." ....._.. . ! ....-

Baldry, D. A. T, (1963) An l u a t i o n by bioassay o svewce' .6f
-DEQ-"&e.posits
on r i v e r i n e vegetation i n the Northern Guinea savannah vegetation zone o f Nigeria
and observations on the f a c t o r s influencing the a v a i l a b i l i t y of deposits
. . . to
. . .. . . .
Glossina p a l p a l i s (R.D. ) B u l l . ent. .Res ,,
................
'497-508 2,
Baldry, D. A . T. (1961C) An assessment by bioassay o f the comparative t o x i c i t y and per-
sistence o f ' d i e l d r i n and isobinzan deposits on r i v e r i n e vegetation i n
Guinea savannah zone of Nigeria i n r e l a t i o n t o the 'cokit'rol of ' G l o s s i
...
(R.D.), B u l l . ent. Res., 55, 49-52 . . . . .. . . . . . . .. . ..... ... ..- . '
'
.1,:z:.;.

Burnett, G . F. (1962~~). The s u s c e p.t. i. .b. i. .l.i.t y of t s e t s e f l i e s t o t o p i c a l applications of .

insecticides. III. The effects. o f age and pregnancy' on the s u s c e p. .t l.. b.. .i. .l..f .t y of i '

a d u l t s of Glossina morsitans, ' B u l l . ' e n t . Res., 53, 337-345 .- .

Research i n East África on the control o f t s e t s e f .l .i e. s from'the

Burnett, G:. F'; -,.(1962@.) :.. , ,

. . ..
a i r , Agr3kultural Aviation, 79-87
..
5, . . .
.I
.I- ,

Burnett, -G;:F.. et, a l . (1964) Aircraft applications of i n s e c t i c i d e s ..in East Africa. '

X W . Very-low-volume aerosols of d i e l d r i n and isobenzan f o r t h e còn$roi ,., ~ " .

of
.... _.
Glossina morsitans Westw., B u l l . ent. Res., 55, 527-539 ... .-.
# .,
., I , .f. I .
.
, *_. . . .,.. .. / i.

Bursell, E. (1963) Tsetse f l y physiology: a review o i recent

.. advances .and
. cG
B u l l . Org. mond. Santé, 28, 703-709
I
..... .-__ .
- . , . ' ..
.............

' .'
Buxton, P. A. (1955) The n a t u r a l h i s t o r y of t s e t s e f l i e s , 'Lond. Sch.. . Byg. &,.trop,
. ._ 'Mecl., ,

. . . . _..
..!,. Memoir No. 10, 816 pp.
. .
, - :...i l'i.:,..:. : ' ., i i '. .... .,
. .~
Chadwick, P. R. (1964) A study o f the r e s t i n g s i t e s of Glossiha. swmnertoni. . . . Aus?. 'in
.. --....+.. .............
Northern
. . . . .
Tanganyika, B u l l .
:
ent. Res., 55,
., . .
23-28
. ;. . . . . .. ..

Chadwick, P. R. e t a l . (1964) An éxperiment o.n.&he e r a d i c a t i o n o f GlossSna' swynnertoni

. . . .
.
Aust by i n s e c t i c i d a l treatment of i t s r e s t i n g s i t e s ; Bull.. ent .-Rk~&:-,- .55, 411-419
., ..
-
. .
Challier; .A.. ( ~ . g . ~ ~ . ~ - ..... __ nGlossina
~ .-Sde~ ~ ~ : b , p~a ~l p~a lti sé gambiensis Vande?plank '1949 au
DDT e t & l a dieldrine, déterminée au mcyen de l a trousse standard OMS pour "
moustiques adu1tes;:Bull:. Soc. :Pa.th. exo-L., 56, 5l:g-533 . , .
, . i

E .
...
Challier, A . (1965a) Campagne de l u t t e contre Glossina palpa?ï&- g a ~ b ~ e n s . i s . . ~ a ~ d e
dans-.-le...foyer de Bamako, (Rgpublique du Mali ) Septième enquête en s a i w n. .s.kche,:
. . . . I..

du 8 au 22 Février 1965, document rondotyp6 . .

O.C.C.G.E., .Bobo-Dioulasso: ' .
. . i
...... ......_.. -- .- ..............
Challie&,---.A.(lg65b) L' évaluation de 1'^age physiologique chez Glossina p a l p a l i s
gambiensis Vanderplank 1949, Bull. Soc. Path. exot ( I n press) . -.-..-"_ - --
 _.
VC/ENT. SEM/WP/~.65
page 14

REFERENCES (Continued)
'

....B.,. (1964) Etude de 1'effet .deJ'HCH nébulisé

Challier; A;, -Eyraud,.M.... &. .D.e.dewano.u,
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galerie forestière (Kankalaba, République de Haute-Volta), Document ronéotypé
/,.,
O.C.C.G.E., Bobo-Dioulasso
. .
. .
Chorley, J. K. (1955) Annual report of the director 3 f tsetse fly operatlons for the
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Cockbill, G. F., Lovemore, D. F. & Phelps, R. J. (1363) The control of tsetse flies.,
(Glossina, Diptera, Muscidae) in a heavily infested',areaof. Southern Rhodesia by
.......
means"of--insecticidedischarged from aircraft, followed'.
by settlement of indi,-
genous people, Bull. ent. Res., 93-106 2, . . -.- .. : _.__ . -. ": . . .
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...
Finelle, P. et al. (1962) Essai de lutte contre Glossina fusca, par pulvérisation de ...

dieldrin, en République Centrafricaine, R ~ vElev., 3, '24.7-253 .'.

. .
. - ......._._....I.._ .__"I" . %
..... .
Finella, P., Laurent, J. & Raynaud, J. P. (1963)Note complémentaire sur un essai de
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417-418 .... ._.-_ .

Ford, J. (1962) African'wildlife'and the tsetse fly-borne dTséases, Bull. epiz. B s .

-
Afr., 10, 9-12
- , ,:".
... - -..-.- .....___... .. . ...,
-,_.. ............
Ford, J., (,1963)
The distribution of the vectors of African pathogenic trypanosom&s,
Bull. Org. mond. Santé, g,
653-669 ' "
. .!.' ,. . 8 . ,$.,

...............
Gaschen, H. (1945) Les Glossines de 1'Afrique occidentale française, Acta trop.
.?
(Basel), suppl. 2, 127 pp.

Glasgow, J. P. (1960) Shinyanga: a review of the work of the tsetse research labora-
tory, E. Afr. agric. Forest J., %, 22-34
...... . . ................... " .
Glover, P. E. (1964)'A review of recent informations on game tsetse relationship.
..
€'Foc.' 1st i n t . Congr. Parasit., Rome, (In p r e s s )

Harley, J. M. B. (1960) A review of entomological research carried out by East African

Trypanosomiasis Research 0rganiza.ticm. ..Rpk,..
7th Commonwealth ent. Conf., London

Harley, J. M. B. (1963)New trends in ent,omologicalresewch,-intrypanosomiasis,

Bull. Org.. mond. Santé, -
28, 825-830
, . . .. ........
. .-.........
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fX.
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Hocking, K. S . (1961). . , , i n s $ & ~ i c i d e ~ . ; ; a gGlossina
Discriminative.:,application._of ~~nst
morsitans West., Bull. ent. Res., 52, 17-22 . . .................
. . . .

Hocking, K. S. (1964) Recent advances in the control of-tsetse flies, Proc. 12th int.
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Hocking, K. S., Lamerton, J. F. & Lewis, E. A. (1963)Tsetse fly control and eradi-
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. . . .. . . . .
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Jordan, A. M. (1961) An assessment of the economic importance -Ö tsetse species
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..... ., .,,;:,z,
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. ,
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1. ' ;

Jordan, A. M., Lee-Jones, F. & Weitz, B. (1962) The natural h o s t

. . . . . r ? ;. . . .. . .
i?-- 'tse tgë- YI,,.-
.
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Northern Nigesia, Ann. trop... Med. Parasit.,
. .
,s,
430-441 . . . . . . . . . s..: I .

. J. ..
(1962) An experiment in the'control of Glossina palpalis. over. . a
...... ....._.........._..I
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I. ..
limited area, J. trop. Med. Hyg., 65, 146-150
!. , .. I
. .i : . .
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Knipling, , E . F. 41963) La stérilisation d' insec es comme moyen poqsible d'obtenir

1 6radicat~o-n-'-de--'fa
'. tsg-tsé, Fnr6;/Tte ÖSz27, Geneva'::.
...... -.. ........ .....................

Langridge, :W. P., . Kernaghan, R. J. & Glover, ; P. E .!,. (1965

.)
. ...:
.A
review
. o f recent. knowledge
. .. ,

of the ecology of the main vectors of trypanosomiasis, Buzl. Org. mond. Santé,
-.-- .. .-,.
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"___"
28, 671-701
.. . . .. . . . . :
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_.
,.- . . ., . .
Le Rouzic (1948) La prophylaxie agronomique. . Un moyen de l.utte pour 1'assainissement ,
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des régions infestées, Bull. méd'. A f r . ' occia';"-franç., - 5;-117-122
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MacLeman: K. J. RT'.&.''Aitkh?son, I?? J: (1'463) Simultaneous control' of three species

'
-

of Glossina by the selective application of insecticid?, Bull.,ent. Res., - 54, I . -

199-212 .... . , . - ......_-..-.I..
. . .
. . ... . . . *. . A
- I

Morris, K. R. S.,.(1960). Trapping as a means ing the game .tsetse, Glossina. 1. .

..;..'::
pallidìpes
_... . . . Au , Bul.1, ent. Res.,,* ,gli ~ / . . , I

.
,.:
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Morris, K. R . S. (1961) Effectiveness of traps in tsetse surveys in the ,Li%ëriärain-
forest; Amer. J. trop. Med..Hyg.,, 10, 905-:913 _./ ; < , , . : " :
. .
... -. . .
i .
I
, i .

~. . .. ,,..:
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. . - ..
-
,
\I,_

Morris, K. R. S." (1962')~~0cal.-'na~-~e'

of epidemic disease, Nature (Lond. );-2?3z-.-*-'
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Mouchet, J., Delas, P. & Yvore, P. (1961) La campagne expérimentale de lutte contre
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Nash, T. A. M. (1963) Progress and problems in the establishment and maint&nañce"-of

labpra$ory
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.. ......
~
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. . . -
8 '.
'Nash, T. A. M. & Steiner, J. O. (1957) The effect of obstructive clearing &ï"Gloss&m
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:I ..1: .

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du Comité OMS d'experts d,es

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-
265, 179-181
.... .I... . . .
 1

n
-,
.
VC/ENT SEM/W/4.65
page 16

. .
REFERENCES (continued)

Page, W. A. & McDonald, W. A. (1959) An assessment of Lhe degree of man-fly contact

zxhibited by Glossina palpalis at water-holes in Northern and Southern Nigeria,
Ann. trop. Med. Parasit., 53, 162-165

Rickenbach, A. (1961) Cartes de répartition des Glossines.en.Afrique odclden$ale

d'expression française, O.S.T.O.M. éd., Paris
t.:. i

Robertson, D. H. H. (1963)Human trgp~nosomiasis_,ip south-east Uganda: a further study

of the. epidemio1og;r of the disease among fishermen and peasant cultivators, Bull.
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........... --.I ....-........

Saunders,,D. S. (1962) Age determination for female tsetse flies and the age compo-
'-
., ..
'sitio66k samples of Glossina pallidipes:Aust., G . piilpalis fuscipes.Rewst. and
G. brevipalpis Nëwst., Bull. erit, Res., .Q, 579-595 ....

Smith, C. N. '& Dame, D. A . (1963) Chemosterilization - 'ä new.field of:xesearch,.in '

Bull. epiz. Dis. Afr., 2, 403-41.4

tseE.se... flyl._c.~nt.rol, . .
; ;., : i.
. . .. , ...
"

Vaucel, M. A. et al. (1963) Répartition de la trypanosomiase africaine chez l'ho&ë

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, ..
545-594 .
. .
, . ,

. . ,
28,
........... .-. ..................-__I__. ......
. . .
Weitz, B. (1963) The feeding-habits of Glossina, Bull. Org. mond. Santé, 28, 711-729
, ,
-
. . . .
Willett.,K: e-...
(1.96.3).Some principles of the epfdemiology'of human trypanosomiasis
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Africa, Bull. Org. mond. Santé, - 28, 645-652
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Wilson, S. ..IL..
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Santé, -28,
.
595-613
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Yvore, P. et al. (1962)Essai d'assainissement une.zong,,,in$e,stée,, par. Glossina fuscipes ;,s.'
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........ .I .......... < .

ADDENDA . i
.
~. . . , . .
. . . . . .. . .
. . . . ,. . .,
.
,
...................
Chadwick, P. R. (1964) Effect of two chemosterilants o n Glossina marsitans, ,Mature
(Lona.), 2,299-300 . -

Dasmann, R. F. (1962) Game...-ranchïng,-,.in,

A&:ican land-use planning, Bull. epiz Dis.;. .
-
Afr., 10, 13-17
r,:
, .' . _ i ..L

. -31, 52c&-%
Evens, F. (1964) L'élevage des glossinas, Bull. Org. ïïond. Santé,. ..._*

Finelle, P. et al. (1963) Rgpartition des glossines en République Centrafricaine.

Etat actuel des connaissan-c-es,Rev. Elev., 337-348 16,
r

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6

ADDENDA (continued)

Glover, P. E. (1961) The t s e t s e problem i n Northern Nigeria, Patwa News Agency (E.A.)
Limited, Nairobi, 383 pp.

Jznkins, D. W. (1964) Pathogens, p a r a s i t e s a.nd predators o f medically important anthro-

pods. Annotated l i s t and bibliograpl-Gr, B u l l . O r g . mond. Santé, 30 Suppl., 17-21

Jordan, A. M, Lee-Jones, F. & Weitz, B. (1962) The n a t u r a l h o s t s of t s e t s e f l i e s i n

-
Northern Nigeria, Ann. t r o p . Med. P a r a s i t . , 56, 430-441

Maillot, L. (1952) Carte de r é p a r t i t i o n géographique des glossines en A.E.F.,

O.R.S.T.O.M., Paris

Maillot, L. (1958) Elevage de Glossina fuscipes quanzensis P i r e s à Brazzaville,

Bull. I.E.C., 15-16, 85-90

P o t t s , W, H. (1964) Gamma-irradiation of Glossina puparia1 stages and control, Proc.

12th I n t . Congr. Ent., London ( I n press)

Rageau, J. & Adam, J. P. (1953) Répartition des glossines au Cameroun f r a n ç a i s , W.

-
Elev., 5,
73-78

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