M i s s i o n E n t o m o l o g i q u e

0 . R .S .T .O .&5 o a u p r e s d e 1 O .C .C .G .E

Biology and C o n t r o l of T s e t s e F l i e s (1)

JeHamon , A , C h a l l i e r , J.Mouchet & J ,Rageau

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(1 ) C e document e s t bas4 s u r un cours f a i t l o r s du S6minaire 0.T'tPn.S.

. i n t e r + r 6 g i o n a l sur l e s m6thudes e n t o m o l o g i q u e s d e c o n t r b l e d e s

v e c t e u r s q u i a e u l i e u e n U . R . S . S . en S e p t e m b r e - O c t o b r e 1 9 6 5

I1 e s t diffus6 car il p o u r r a i t être u t i l e aux e n t o m o l o g i s t e s

d e v a n t f a i r e d e s cours sur l e s g l o s s i n e s d a n s l e s pays d'af- f e c t a t i o n . IC1 c o n v i e n d r a i t a lors d e compl6ter l a b i b l i o g r a -

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-I. J .Hamon , A . C h a l l i e r ' , J and J ,Rageau+'

M e d i c a l E n t o m o l o g i s t s , O f f i c e d e l a Wecherche L .

S c i e n t i f i q u e & T e c h n i q u e O u t r e Yer

I .

1. IlYTEODUCTIOR . For c e n t u r i e s human and a n i m a l t r y p a n o s o m i a s i s have r a v a g e d v a s t

a reas o f t r o p i c a l A f r i c a . A s r e c e n t l y as 2 0 y e a r s a g o s l e e p i n g - s i c k -

n e s s was c o n s i d e r e d o n e of t h e most t e r r i b l e of a l l endemic d i s e a s e s

a f f e c t i n g t h e A f r i c a n c o n t i n e n t , Animal t r y p a n o s o m i a s i s , r e n d e r s t o c k -

r a i s i n g d i f f i c u l t , i f n o t i m p o s s i b l e , i n *any a reas , and i n c a t t l e -

g r a z i n g arieaa b o r d e r i n g on t h e G l o s s i n a zones l i v e s t o c k i s weakened

and d o e s n o t s u p p l y any draugh$-anima$ (VAU@EL.~& a l . , f 9 6 3 - WILSON &

a l . , - FOED, 11963) . l i e s e a r c h e f f o r t s u n d e r t a k e n b o t h i n t h e L a b o r a t o r y and i n

t h e f i e l d have s u p p l i e d in r e c e n t y e a r s b e t t e r methods o f p r e v e n t i o n ,

d e t e c t i o n and t r e a t m e n t of t r y p a n o s o m i a s i s . Dur ing t h e s a m e p e r i o d

s p r e a d i n g p o s s i b i l i t i e s of t h e d i s e a s e h a v e i n c r e a s e d t h r o u g h comfiu-

n i c a t i o n improvemen t s and deve lopmen t ok w o r k e r s m i g r a t i o n s be tween

n e i g h b o u r i n g c o u n t r i e s ( A B E D I & I!AXLLBB, 1964).

Human s l e e p i n g - s i c k n e s s p r e v a l e n c e has been r e d u c e d t o a n

e x t r e m e l y low l e v e l , b u t the disease is f a r f rom e r a d i c a t e d and many

small f o c i e i t h e r r e m a i n ac t ive o r e v e n , a p p e a r i n p r e v i o u s l y c l e a r e d

a r e a s . The s i t u a t i o n i s s l i g h t l y / ... -----I--------_----____l_l________e____l-~----------------~,----------

. + Q.L<.S.T.O."F. Team n e a r O . C . C . G . E . , P . O . B . 171, Bobo-Dioulasso ,

Upper - V o l t a . ++ O . B . S . T . 0 . : ~ . 70-74, r o u t e d 'Au lnay , 93-Bondy, F r a n c e .

'. .

6 e t t e r i n West A f r i c a , where t h e p a r a s i t e Trypanosoma gambiense h a s

a p p a r e n t l y no a n i n a l r e s e r v o i r , t h a n in East A f r i c a where occur s a l s o

T . r h o d e s i e n s e w i t h b o t h human and a n i m a l h o s t s (ROFBRTSQN, 1 9 6 3 - WLL-

LETT, 1 9 6 3 ) .

G l o s s i n a - b o r n e a n i m a l t r y p a n o s o m i a s i s a re a l w a y s v e r y w i d e s -

p r e a d and h i n d e r econornic deve lopmen i of t h e m a j o r i t y o f A f r i c a n s t a t e s , ~

Chemotherapy c a n n o t be e x t e n d e d t o a l l d o m e s t i c a n i m a l s and h a s i t s

s e t b a c k s , i n c l u d i n g d r u g r e s i s t a n c e of t h e t r y p a n o s o m e s , Co t se t se f l y

c o n t r o l h a s a n i m p o r t a n t p a r t t o p l a y i n t h e deve lopmen t o i t h e b f r i -

caan c o n t i n e n t . 2. GEOGRAPHIC DISTRIBUTION OF TSETSE F E f i S .

.l

.;

- G l o s s i n a s p . i n f e s t a b o u t 16.000.000 km' on t h e Airrican

m a i n l a n d . The,ir p r e s e n t d i s t r i b u t i o n h a s been r e c e n t l y summarized by

POBP (1963) b u t more a c c u r a t e d a t a f o r f r e n c h - s p e a k i n g s t a t e s of \?est

Af r i ca c a n b e found i n MAILL6T i1952), RAGEAU & ADAPI (19SrS)j RICHEM-

3ACH (196%) and FINIBLLB i& a l . ( P 9 6 3 ) . T s e t s e f l y d i s t r i b u t i o n i s . , n o t e n -

t i r e l y s t a b l e , even i n a b s e n c e o f man i n t e r f e r e n c e . F o r exanlple seve- r a l s p e c i e s r e t r e a t e d f rom l a rge a reas of s o u t h - e a s t Afr ica d u r i n g t h e

Grea t R i n d e r p e s t o f 4896,' and s inc ' e t h i s p e r i o d d . p a l 1 i d i p e . s and G, m o r s i t a n s , f o l l o w e d by G . a u s t e n i , move s l o w l y s o u t h i n :!ozambique and

. R h o d e s i a , r e s t o r i n g t h e s i t u a t i o n i , r i o r t o t h e r i n ' d e r p e s t . I n West A f r i c a , d u r i n g t h e l a s t P5 y e a r s , G . s u b m o r s i t a n s h a s e x t e n d e d

n o r t h w a r d s i t s d i s t r i b u t i o n ('dILSOPd, 1958) . Such c h a n g e s i n d i s t r i b u -

t i o n are p r o b a b l y o f comnon o c c u r e n c e , b u t e s c a p e d e t e c t i o n by l a c k

of accu ra t e b a s e d a t a and s u r v e y s . i r

\ T s e t s e f1ie.s a r e c l a s s i f i e d i n t o t h r e e g r o u p s , k y p i f h e d i res- I

p i c t i v e l y by G . p a l p a l i s , C . m o r s i t a n s and G . f u s c a ; t h e i r d i s t r i b u t i o n s

a r e t h e f o l l o w i n g o n e s : I

ni , 2.1 . G l o s s i n a p a l p a l i s g r o u p (.. . .

The p a l p a l i s g r o u p i n c l u d e s f i v e s p e c i e s and f o u r sub-spe-

c i e s and i s ' r e s t r i c t e d t o idest and C e n t r a l A f r i c a , w i t b r e l i c s i n E-

t h i o p i a arLd A r a b i a . L

G . t a c h i n o i d e s o c c u r s i n t h e s u d a n e s e s a v a n n a h and i n t h e

s o u t h e r n p a r t o f t h e s a h e l i a n a rea o f West A f r i c a , w i t h r e s i d u a l f o c i

' i n n o r t h - e a s t e r n A f r i c a and i n t h e S o u t h - e a s t of t h e A r a b i a n p e n i n s u l a ,

It c a n w i t h s t a n d v e r y d r y c o n d i t i o n s , b u t c o n g r e g a t e s i n t h e d r y s e a 4

s o n a r o u n d water h o l e s , r e s i d u a l p o o l s of t e m p o r a r y rivers' , and so On. . .

3

G . p a l p a l i s and G . f u s c i p e s o c c u r i n t h e f o r e s t a s w e l l a s

i n t h e g u i n e a n and s u d a n e s e s a v a n n a h s . I n t h e s a v a n n a h s t h e y a r e res-

' t r i c t e d t o t h e i n m e d i a t e v i c i n i t y o f r i v e r s and r i v u l e t s . I n f o r e s -

t e d a reas a l s o t h e y c o n g r e g a t e a l o n g w a t e r c o u r s e s , b u t n o t as stric-

t l y a6 i n s a v a n n a h , u s i n g them more f o r t h e f l i g h t l i n e s t h a t t h e y e f -

f e r t h a n f o r t h e i r m i c r o c l i n a t e , 9

L , i G - p a l l i c e r a o c c u r s i n r a i n f o r e s t a r e a s , b u t c a n be encoun-

t e r e d sometimes in the g a l l e r y f o r e s t s of t h e g u i n e a n s a v a n n a h . *

G . c a l i g i n o s a is r e s t r i c t e d t o t h e c o a s t a l r a i n f o r e s t and

t o t h e mangrove a r e a s . G . t a c h i n o i d e s , G . p a l p a l i s arid G . f u s c i p e s are t h e main vec -

tors of t h e human s l e e p i n g - s i c k n e s s c a u s e d by T.gambiense . I

2.2. G . m o r s i t a n s g r o u p . The morsitans group i h c l u d e s s e v e n s p e c i e s and t h ree sub-

\

s p e c i e s , all i n h a b i t i n g s a v a n n a h s ,

G j m o r s i t a n s complex o c c u r s m a i n l y i n s u d a n e s e s a v a n n a h s ,

w i t h C . s u b m o r s i t a n s i n Nest and C e n t r a l A f r i c a , a n d G . m o r s i t a n s i n

E a s t A f r i c a ,

G . l o n g i p a l p i s i s r e s t r i c t e d t o guiniean s a v a n n a h s and t o sou,..

p a r t l y d e f o r e s t e d a r e a s of the f o r e s t zone o f + l e s t A f r i c a . G . a u s t e n i i s m a i n l y a lowlands and c o a s t a l s p e c i e s o f Eas t

A f r i c a . G , s w y n n e r t o n i and G . p a l 1 i d i p e s a r o h i g h l a n d s p e c i e s of 3 a s t

A f r i c a . T h e , f i r s t one is r e s t r i c t e d t o Kenya and T a n z a n i a . The s e c o n u

s p e c i e s o c c u r s from- B t h i o p i a t o Yozaanbique and i s p r e s e n t i n some

c o a s t a l a r e a s ; it h a s o c c u r e d i n t h e past i n ï l a t a 1 b u t h a s been e ra -

d i c a t e d there .

H

r

4

G . b o r g e s i h a s been r e c e n t l y d e s c r i b e d f rom ?"ozambique ( F I X E L

G . n i o r s i t a n s and G . swynner ton i are t h e normal v e c t o r s o f the .1 1960).

norma l v e c t o r s o f tile s e c o n d form o f human s l e e p i n g - s i c k n e s s , c a u s e d

by T . r h o d e s i e n s e ; t h e y a r e a l s o , w i t h G . p a l l i d i p e s and G . submors i tnn f

v e r y i m p o r t a i t t v e c t o r s o f ' s e v e r a l a n i n a l t r y p a n o s o m i a s i s ( c a u s e d by

T . b r u c e i , T . v i v a x , T . c o n g o l e n s e ) T . r h o d e s i e n s e h a s beeil a l s o rece-

vered f roin G , p a l l i d i p e s s i n c e 1940 (;-acK1CC'IiAM, 1944) and more r e c e n -

t l y froln G . f u s c i p e s ( 8 0 U T I 3 3 N & XOG'E&TSilPJ, 1961 - WICLETT & A1.,1964- I

WPEL3TT, 1965) ,

4

1

. . 4

2.3. G.fusca group, '

The f u s c a group i n c l u d e s 12 s p e c i e s and two s u b - s p e c i e s , "

Which i n h a b i t m a i n l y d e n s e l y f o r e s t e d a r e a s . Gknnsh i , G . s c h w e t z i , C , t a b n n i f o r m i s , C . h a n i n g t o n i , G.seve-

r i n i , G . v a n h o o f i a'nd G , f u s c i b l e u r i s o c c u r i n t h e Lower g u i n e a n f o r e s t :

s o n e t i m e s i n v e r y r e s t r i c t e d a rea ' s 1

G . f ~ s c a o c c u r s i n all f o r e s t e d a r e a s , f rom S i e r r a Leone t o

Uganda and i n some of t h e l a r g e r g a l l e r y f o r e s t s of t h e g u i n e a n sa-

vannah . G . n i g r o f u s c a i s w i d e s p r e a d i n t h e U p p e r g u i n e a n f o r e s t and

i n t h e n o r t h e r n g a r t of t h e Lower g u i n e a n f o r e s t , f rom Guinea t o Ugar

G.inedicoruiii i s r e s t r i c t e d t o t h e Upper g u i n e a n savnnnah anC - d a .

f o r e s t . G , b r e v i p a l p i s i s v e r y w i d e s p r e a d i n g a l l e r y f o r e s t s and sa-

v a n n a h s o f Prlali B a s t A f r i c a f rom N a t a l t o B a s t e r n Congo ( L e o p o l d v i l -

le) and t o S o u t h e r n E t h i o p i a and S o m a l i a . G . l o n g i p e n n i s is a l s o an e a s t e r n s a v a n n a h s p e c i e s ) b u t Pes-

C

t r i c t e d to Kenya, E t h i o p i a S o u t h e r n S o i n a l i a , S o u t h - e a s t e r n S u d a n ,

N o r t h - w e s t e r n Uganda ar,d N o r t h e r n T a n z a n i a . The most common s p e c i e s b e l o n g i n g t o t h e f u s c a g r o u p a re

i m p o r t a n t v e c t o r s of a n i m a l t r y p a n o s o m i a s i s caused b y ? ' .vivax and

T . c o n g o l e n s e ( J 0 2 D A N , P 9 6 1 & 1965 a ) . 3 . BIOLOGY OF TSETSE FLIES .

3.1. K e p r o d u c t i o n c y c l e . . T s e t s e f l i e s m a t e d u r i n g t h e days f o l l o w i n g e m e r g e n c e , when

t h e f e m a l e s are t e n e r a l . One i n s e m i n a t i o n is s u f f i c i e n t f o r t h e who-

l e l i f e of t h e f e m a l e , b u t ; '*ales ma te s e v e r a l t i m e s d u r i n g t h e i r l i f ~

, and p r o b a b l y some f e m a l e s do t h e same; r e c e n t i n v e s t i g a t i o n s c a r r i e d

o u t i n Upper Volta b y o n e of u s (A .C . ) on G . g a l p a l i s h a v e shown t h a t

n o n - t e n e r a l w i l d c a u g h t f e m a l e s a r e a l m o s t a l w a y s i n s e m i n a l e d ; howe-

v e r t h e p e r c e n t a g e o f i n s e m i n a t e d females w i t h i n c o m p l e t e l y f i l l e d - s p e r m a t h e c a e r e a c h e s a b o u t amongs t t e n e r a l f l i e s , a g a i n s t o n l y

51% i n f l i e s b e l o n g i n g t o . t h e n e x t two o l d e r a g e g r o u p s ; so i t i s v e r y p r o b a b l e t h a t f e m a l e s cf C . p a l p a l i s s e v e r a l t i s n e s d u r i n g

t h e i r f i r s t d a y s o f l i f e . Jemales y r e s e c t thsir.: naxinum a t t r a c t i v e -

n e s s f o r males When t h e y are only 3 days o l d , a n d i a r e less a t t r a c t i -

ve w i t h i n c r e a s i n g a g e , a l t h o u g h b e i n g a l w a y s a b l e t o m a t e (NkSEi,l95f

-

5

JOhDBN, 1958 * ) *

,Homologous m a t i n g s are t h e r u l e , b u t some t imes :na t ing o c c u r s i n na-

t u r e be tween c l o s e l y r e l a t e d s p e c i e s ; s u c h m a t i n g s a r e g e n e r a l l y n o t

f e r t i l e and c a n be l e t h a l for t h e f e m a l e

The two o v a r i e s are composed o f two o v a r i o l e g e a c h , and

e a c h o v a r i o l e d e v e l o p s i n t u r n (SBUHDERS, 1962 - CWALLIBW, 1963 b ) .

The f i r s t o v u l a t i o n o c c u r s S t o ' U days a f t e r t h e emergence of t h e

f e m a l e ; t h e m a t u r e o v o c y t e i s n o t l a i d , b u t p a s k e s i n t o t h e f e m a l e u t r

rus where f e c u n d a t i o n o c c u r s ; t h e n t h e l a r v a grows i n s i d e t h e u t e r u s ,

t h e f o o d b e i n g s u p p l i e d by t h e " m i l k g l a n d s " o f t h e f e a a l e ; t h e la-

t e t h i r d i n s t a r l a r v a is l a i d on t h e g round and p u p a t e s u s u a l l y two :..& t o f i v e c e n t i m e t e r s below t h e s u r f a c e , some hours l a t e r . L a r v a l de-

v e l o p m e n t , i n u t e r o , t a k e s a b o u t 10 d a y s , d u r i n g w h i c h t h e f e m a l e s

b i t e s a t l e a s t t h r e e times, b u t the f i r s t l a r v a i s l a i d when t h e f e -

male i s a b o u t 16 d a y s o l d , or moro. Yhen t h e f i r s t l a r v a i s l a i d t h e

secor ,d o v o c y t e p a s s e s i n t o t h e u t e r u s , and so o n . The d u r a t i o u ; of t h e

pupal l i f e v a r i e s f rom 2Q t o 60 d a y s , a c c o r d i n g to t e m p e r a t u r e ,

I

The a d u l t c a n f l y some h o u r s a f t e r i t s emergence f rom t h e

pupariun and is a b l e t o b i t e t h e f o l l o w i n g day ., 3.2. F e e d i n g h a b i , t s .

I . .

Both s e x e s o f t se tse flies suck b lood and t h e y do n o t u s e

aiiy o t h e r f o o d . They do n o t i n g e s t u s u a l l y e i t h e r water or n e c t e r ,

b u t seem somet imes a b l e t o p i e r c e p l a n t s for s u c k i n g sap,. They c a n

feed on i n s e c t s , a s o n e of u s ( J . R . ) h a s o b s e r v e d for G , p a l p a l i s w i t l -

c a t e r p i l l a r s . Bach s p e c i e s of t s e t se f l y e x h i b i t s d e f i n i t e f e e d i n g p r e f e -

r e n c e s , which a re d i r e c t l y r e l a t e d t o t h e i r ,neclical and v e t e r i n a r y

i m p o r t a n c e a s p a t h o g e n v e c t o r s ( J O L ~ D A I J , 1965 a ) , b u t t h e m a j o r i t y .

of G l o s s i n a sp. are a b l e t o f e e d on 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 kmown t r o p h i c p r e f e r e n c e s a r e t h e f o l l o w i n g (MBITZ, 2963 a ) , accc

d i n g t o 22,640 b lood meals i d e n t i f i e d :

- 6; . s w y n n e r t o n i , G . a u s t e n i , G , f u s c i p l e u r i s and G . t a b a n i f o r m i s f e e d

ma in ly ' on s u i d s , w i t h b o v i d s and o t h e r v e r t e b r a t e ( r h i n o c e r o s ,

h i p p o t m u s , p o r c u p i n e ) a s s e c o n d a r y h o s t s ;

- G . m o r s i t a n s and G . s u b . n o r s i t a n s f e e d e q u a l l y on s u i d s sild b o v i d s ,

w a r t h o g ( "hacochoe rus s p . ) b e i n g t h e p r e f e r r e d h o s t ; man is a l s o used

6

as h o s t , snainly i n Vest A f r i c a by G . sub :nor s i t ans ( J o r d a n , 1965 b ) ;

- G . p a l l i d i p e s , G . f u s c a and @ . l o n g i p e n n i s f e e d e x t e n s i v e l y OE bus-

hbuck ( T r a g e l a p h u s s c r i p t u s ) and t h e f i r s t and s e c o n d s p e c i e s b i t e

a l s o s u i d s t o soine e x t e n t i

- C . l o n g i p e n n i s f e e d s m a i n l y on r h i n o c e r o s j w i t h g i r a f f e , e l e p h a n t ,

b u f f a l o and o s t r i c h as s e c o n d a r y h o s t s ;

- G . b r e v l p a l p i s f e e d s m a i n l y 011 h i p p o p o t a m u s , s u i d s , b u f f a l e and

bushbuck ;

- Gopalpalis, G . f u s c i p e s and G o t a c h i n o i d e s f e e d e x t e i i s i v e l y 011 " z n ,

c r o c o d i l e s , m o n i t o r l i z a r d s and b o v i d s , b u t a l s o sometimes on w i l d

game (FOSTIIR, 3 - 9 6 4 ) ; @ . t a c h i n o i d e s i s p a r t i c u l a r y i n f e o d a t e d t o man

and c a t t l e . The b e h a v i o u r of each s p e c i e s is c h a r a c t e r i s t i c and i s n o t

e n t i r e l y depender , t on t h e a v a i l a b i l i t y o f d i f f e r e n t h o s t s ; t h i s view

is s u p p o r t e d by t h e f a c t t h a t commonly o c c u r i n g a n i m a l s are n o t f e d

don by G l o s s i n a , t h e b e s t example b e i n g z e b r a (Bquus b u r c h e l l i ) i n

G . m o r s i t a n s a reas o f B a s t A f r i c a , and w a t e r b u c k s ( l iobus s p .) (bd3ITZ,

1963 b ) . Few d a t a a re a v a i l a b l e a b o u t the a d a p t a b i l i t y of tsetse

f l i e s t o a v a r y i n g h o s t f a u n a . It i s l i k e l y t h a t f l i e s l i k e G . m o r s i -

t a n s and G . s u b m o r s i t a n s , the G . p a 1 p a l i s group and p o s s i b l y G.swynner-

- t o n i and G . p a l l i d i p e s would q u i c k l y a d a p t t o c h a n g e s i n f aunz ; howe-

v e r , f o l l o w i n g t h e r i n d e r p e s t p a n z o o t i o of 1890-1896 t h e G . m o r s i t n n s

f l y - b e l t s have r e c e s s e d i n 3 a s t A f r i c a , a f t e r a lmost e x t e r , n i n a t i a n

o f a l l f a v o u r a b l e h o s t s (FO>UD, l.965). The a u t h r o p o p h i l y of G.palpn-

- lis and cf G . t n c h i n o i d e s i n c r e a s e s when t h e w i l d f a u n a and t h e l i v e s -

teck d e c r e a s e (PAGE & KBCBOïÏktD, 1 9 5 9 - JOXDAII &: a l . , k96l. & 3.962 - LAITGEZIDGE & al., 1 9 6 3 ) . Seve ra l species w i t h v e r y r e s t r i c t e d f e e d i a g

h a b i t s , such as G . f u s c i f i l e u r i o , G . a u s t e n i - and G o t a b a n i f o r m i s ( i n f e o -

d a t e d t o s u i d s ) , G . l o n g i p a l p i s and G.fu:;ca ( i n f e o d a t e d t o bushbuck

and b u f f a l o ) , arid G . 1 o n g i p e n n i s ( fcedinE; o n r h i n o c e r o s , e l e p h a n t and

b u f f a l o ) , may be v e r y r a p i d l y a f f e c t e d by t h e remov-al o f o n l y a few

s p e c i e s .

7

3 . 3 . Bcology . T s e t s e f l y e c o l o g y and its p h y s i o l o g i c a l b a s i s have been

r e c e n t i y reviewed (LANGBIDGE & a l . , 1 9 6 3 - WARLBY, 1965 a-E'US3B'L.L, -2

1959, I960 & 1963) and have been t h e s u b j e c t i n a - r e c e n t p a s t o f ma-

ny p u b l i c a t i o n s , each o f them g e n e r a l l y d e v o t e d t o one or two G l o s s i -

na s p e c i e s . Bach s p e c i e s of t s e t se f l y h a s a s p e c i f i c d i s t r i b u t i o n a r e a ,

c h a r a c t e r i z e d by c l i m a t e , v e g e t a t i o n and s o i l , t h i s last e l e n e n t

b e i n g i m p o r t a n t f o r l a r v a l d e p o s i t and p u p a l s u r v i v a l . The r e c e n t

s t u d i e s of BURSEEL (see a b o v e ) have c l e a r l y shown t h a t one of t h e

m a j o r l i m i t i n g f a c t o r s f o r t h e d i s t r i b u t i o n of many s p e c i e s of t s e -

tse. f l J e s i s t h e r e l a t i v e hur i \ id i ty . A d u l t s , which f l y ancl cnn s e l e c t ;

t h e i r r e s t i n g - p l a c e s , c a n w i t h s t a n d l'ow r e l a t i v e h u m i d i t i e s of t h e

g e n e r a l e n v i r o n m e n t , b u t pupae o f ,nany c p e c i e s c a n n u t , and the d i s -

t r i b u t i o n i s under t h e dependence o f t h e p u p a l e u v e l o p p e r m e a b i l i t y .

The pupae of some s p e c i e s , s u c h a s t h o s e o f r w r s i t n n s , do n o t l o s e

zny w a t e r w i t h i n dry soils, w h e r e a s pup:.e of s e v e r a l o t h e r s p e c i e s ,

s u c h a s p a l p a l i s , a r e e a s i l y k i l l e d by d e s s i c a t i o n and r e q u i r e f o r

t h e i r deve lopmen t an a l m o s t s n t u r a $ e d a t m o s p h e r e , b u t w i t b o u t any

f r e e water. T e m p e r a t u r e may p l a y a p a r t i n 1irilitir:g t h e d i s t r i b u t i o n ,

b u t m a i n l y i n e x t r e m e c o n d i t i o n s

A t l e a s t i n t h e savannah z o n e s , i t i s o f t e n p o s s i b l e eo d i s t i n g u i s h , i n t h e d i s t r i b u t i o n a rea of e a c h s p e c i e s , t h r e e a r e a s

(MASII & PAGE, 1953 - PAGB, i961 a - MOUCHBT & a l . , 2-961 - JACKSON,

1946) . - t h e permanent b r e e d i n g - s i t e s , w h e r e f l i e s ocCTtlr a l l y e a r round and

where t h e y c o n g r e g a t e d u r i n g u n f a v o u r a b l e s e a s o n s , and u s u a l l y where

t h e y l a y l a r v a e ;

- t h e te inporary b r e e d l n g - s i t e s , where f l i e s o c c u r d u r i n g t he m o s t

f a v o u r a b l e s e a s o n s , u s u a l l y d u r i n g t h e r a i n y s e a s o n , b -u t where t h e y

c a n n p t s u r v i v e i n t h e d r y s e a s o n ;

- t h e f e e d i n g - g r o u n d s , wh ich a r e g e n e r a l l y open a r e a s , n o t v e r y f a r

from t,he b r e e d i n g - s i t e s , where f l i e s c a n e a s i l y d e t e c t t h e i r h o s t s

and f e e d upon them; f e e d i n g - g r o u n d s c a n e x t e n d a few l c i l o m e t e r s a-- '

round b r e e d i n g - s i t e s i n r a i n y sens-on , b u t are u s u a l l y r e s t r i c t e d t o

t h e i r inzmedliate v i c i n i t y d u r i n g t h e d r y season .

, .

8

I n e a c h a r e a r e s t i n g - p l a c e s o f t h e f l i e s ' v a r y a c c o r d i q t o

env i ronmen$ and t o p h y s i o l o g i c a l c o n d i t i o n of t h e f l y , b u t t h e y are

u s u a l l y c o n s t i t u t e d by l o g s , small' t r u n k s and r e d i u m - s i z e d b r a n c h e s ,

up t o some [fleters above t h e g r o u n d . Mowever soríe s p e c i e s , P i k e G.swyn--

n e r t o n i and G . p a l l i d i p e s , r e s t e x t e n s i v e l y on l e a v e s by n i g h t and

on t h e u n d e r s i d e o f b r a n c h e s by d a y (FiABLBY, ~ 9 6 0 Sc 1963 a ) . Sin WOG-

d e d s a v a n n a h a reas so:iie s p e c i e s of t r e e s a f f o r d b e t t e r r e s t i n g - p l a -

ces t h a n o t h e r s and a r e r e g u l a r l y s e l e c t e d by t h e majority of f l i e s .

T s e t s e , f l i e s are g e n e r a l 1 a c t i v e o n l y by d a y t ime, b u t some

s p e c i e s o f t h e f u s c a g r o u p a r e also a c t i v e by n i g h t and G .b rev ipa1-

p i s e x h i b i t s a v e r y c h s r q c t e r i s t i c c r e p u s c u l a r a c t i v i t y , i i m e d i a t e -

ly a f t e r s u n s e t and i ; n m c d i a t e l y b e f o r e s u n r i s e (HkitLEY, 1965 b ) , e G, palpalis and G . . submors i tans b i t e a l s o scrnetimes by n i g h t when d i s t u r -

* bed f rom t h e i r r e s t i n g p l a c e s (J . H . , p e r s o n a l o b s e r v a t i o n ) . P u p a l sites are s e l e c t e d by f e n a l e t s e t se f l i e s and , f o r

I a sane s p e c i e s and w i t h i n t h e same area, t h e i r l o c a t i o n and r e l a t i -

ve i m p o r t a n c e v a r y f rom s e a s o n t o seasor. (PTAS& & PAGE, 1953 - CLhS-

. GOY, 1961 - J O N U E , 1965 b ) . Pupae can w i t h s t a n d h i g h r e l a t i v e hutRi-

d i t i e s b u t n o t s u b m e r s i o n ; h i g h t e m p e r a t u r e s r e d u c e t h e l e n g t h of

t h e p u p a l period, b u t t o o h i g h . t e m p e r a t u r e s c a n k i l l t h e pupae i n

some d a y s and e v e n some h o u r s ; s o p u p a l s i t e s are u s u a l l y s i t u a t e d

i n w e l l - s h a d e s p l a c e s , unde r l o g s , . b u s h e s , i n ho l low t r u n k s , and s o

o n , For same s p e c i e s , s u c h a s G . t a c h i n o i d e s alid G.palpnlis, p u p a l s i -

t e s caz b e e n t i r e l y man-inade, b e i n g p r o v i d e d b j mango and baiznna p l n ~ -

t a t i o n s and e v e n somet imes by g r a n e r i e s , t h a t c h e n c l o s u r e s a n d o t h e r

a r t i f i c i a l s h e l t e r s . The d e p t h t o w h i c h t h e l a r v a e enter t h e soil. va-

r i es w i t h e n v i r o n m e n t a l c o n d i t i o n s ; i n t h e r a i n y s e a s o n t h e l a r v a c a n

p u p a t e o n t h e g r o u n d , or o n l y one o r t w o c e n t i m e t r e s be low; i n t h e d r y s e a s o n s t h e l a r v a c a n e n t e r f i v e t o e i g t h c e n t i m t r e s bellow /the g round

b e f o r e p u p a t i n g ,

. ' 3 . 4 . T s e t s e f l y s u r v i v a l and p o p u l a t i o n dynamics . Bxcept f o r t h e i r f i r s t days o f i m a g i n a l l i f e , d u r i n g w h i c h

t h e m o r t a l i t y r a t e i s h i g h (1lASH & TAGB, 1953 - I'JBSH & KqkNAGEAM,8965-

A.C. p e r s o n a l o b s e r v a t i o n ) I a d u l t s of G l o s s i n a have a h i g h s u r v i v a l

rate, i n t h e l a b o r a t o r y as i n n a t u r e . Average life o f two t o t h r e e

months seems t o be t h e r u l e (;LkL&BY, 3.963 b Oc 1965 a ) and s u r v i v a l

for 2-54 d a y s h a s been o b s e r v e d in G o p a l p a l i s ; howerve r a v e r y s m a l l

p r o p o r t i o n o f t h e p o p u l a t i o n i s l i k e l y t o a c h i e v e s u c h e x t r e m e age

8e ?AGE, 1 9 5 3 ) .

Y

9

L

Ø

t.

e

-

F

Causes of m o r t a l i t y o f l a r v a s and pupae a r e more iaportant

b e c a u s e p r e d a t o r s a r e nurilerous : i i i s e c t i v o ' r o u s b i r d s , i n s e c t i v o r o u s

i n s e c t s , and s o o n ; b e s i d e s mzny i n s e c t s p a r a s i t e t se t se pupae :iSip- Cera Eombyl idae , Hymenoptera C h a l c i d i d a e and N u t i l l i d a e . Fupae a re

also e a s i l y k i l l e d by u n f a v o u r a b l e e n v i r o n m e n t char iges . I n n a t u r e .

p r o b a b l y l e s s t h a n h a l f t h e l a i d l a r v a e reach the a d u l t s t a g e . T s e t s e f l y d e n s i t i e s do n o t f l u c t u a t e v e r y much f r o m one

y e a r t o a n o t h e r , a s l o n g a s t h e e n v i r o n m e n t a l c o n d i t i o n s roma in c o n s -

t a n t . G . swynner ton i h a s been s t u d i e d d u r i n g 23 c o n s e c u t i v e y e a r s i r r

S h i n y a n g a a rea , T a n z a n i a , d u r i n g w k i c h the h i g h e s t d e n s i t y v n l u e ' h a s

been 18 t i m e s t h e l o w e s t one . G . p a l p a l i s , wh ich h a s been- s t u d i e d du-

r i n g 16 y e a r s in Kaduna a rea , N i g e r i a , h a s e v e n more s t a b l e p o y u l a -

tions i n r i v e r i n e f o r e s t s , where the h i g h e s t r e c o r d e d d e n s i t y was on-

l y 3,B t i n e s t h e l o w e s t ane (G'LkSGQisJ & WBLCET, 1.962 - JOEDAN,f96!3 e).

I

3 . 6 , Sampl ing methods . The s a m p l i n g me thods u s e d must be r e l a t e d t o t h e l o c a l con-

d i t i o n s and t o t h e i n v e s t i g a t e d s p e c i e s , I

The m o s t w i d e s p r e a d method i s based on f l y r o u n d s , the f l y -

c a t c h e r s moving a long s t a n d a r d p.nths u n d e r no raa l i zed c o n d i t i o n s t h r o ~

the d i s t r i c t u n d e r i n v e s k i g a t i o n s ; results a re e x p r e s s e d i n f l y p e r

u n i t of l e n g t h of t h e p a t h s . The b a i t ,can b e human b e i n g s (if s o usual- l y the fly-catchers t h e m s e l v e s ) or a dornest$c aniri;al, o r e v e n i: c a r

moving a& a very low s p e e d , Catches d u r i n g f l y r o u n d s c a n also . b e . car-

r i e d o u t on r e s t i n g f l i e s . F i x e d c a t c h i n g s t a t i o n s , u s e d d u r i n g t h e who le a c t i v i t y pe-

r i o d o f t h e f l i e s ( s a g e n e r a l l y from d n w n i t o dtask) a r e a l s o w i d e l y u s e

b a i t s b e i n g huma.ns , o r t e t h e r e d a n i m a l s , or s p e c i a l l y d e s i g n e d traps.

Moving teams c o l l e c t n m i n l y u n f e d h u n g r y f l i e s on t h e b a i t s ,

W i t h a l a r g e e x c e s s o f m a l e s , and erieorged and n o t - h u n g r y f l i e s on

t h e i r r e s t i n g p l a c e s . S t a n d i n g teams c o l l e c t a h i g h e r p r o p o r t i o n of

females, m a i n l y when u s i n g traps. Moving c a r b a i t s are sometimes v e r y

e f f i c i e n t f o r a t t r a c t i n g non-man-bi t ing s p e c i e s . . Very low d e n s i t i e s o f t s e t se f l i e s a r e a l w a y s d i f f i c u l t t o

asse.ss and may e v e n e s c a p e e n t i r e l y t o d e t e c t i o n .

In many i n s t a n c e s tsetse c o n t r o l i s a v e r y e f f i c i e n t way

t o s t o p hunan t r y p a n o s o m i a s i s t r a n s m i s s i o n i n r e s t r i c t e d a r e a s , e i -

t h e r a l o n e o r aS a complement t o c h e m o t h e r a p i c neasUres. T s e t s e f l y

c o n t r o l o r ' b e t t e r , e r a d i c a t i o n , i s o f t e n t h e c h e a p e s t method f o r p r e -

v e n k i n g human or l i v e s t o c k t r y p a n o s o m i s s i s t r a n s m i s s i o n i n a r e a s o f

Sudanese and Guinean savai inzh z o n e s (~-1OCKIM3 & a l . , P 9 6 3 ) 4

Methods u s e d f o r t s e t se f l y c b n t r o l j u s t b e f o r e d n d ' a f t e r

t h e s e c o n d World War a r e g r a d u a l l y b e i n g r e p l a c e d by o t h e r o m s ba-

s e d o n r e s i d u a l i n s e c t i c i d e s a p p l i c a t i o n s , and c h e m o s t e r i l i z a t i o n is

b e i n g i n v e s t i g a t e d . E u t i n t h e p a s t d i r e c t d e s t r u c t i o n , i n d i s c r i m i n a -

te and s e l ec t ive c l e a r i n g s , and h o s t s e l i m i n a t i o n have been e x t e n s i -

v e l y u 5 s d , soli ietimes w i t h f a i r success (GZQ)V3;1., P96P) . 4.1. D i r e c t d e s t r u c t i o n of t s e t se f l i e s .

D i r e c t d e s t r u c t i o n o f t s e t s e f l i e s by n e t c o l l e c t i o n aird

g l u e d b n i t s h a s n o t been v e r y e f f i c i e n t i n t h e p a s t , e x c e p t i n P r i n -

c i p e I s l a n d where G.palpalis was a l m o s t e r a d i c a t e d . D e s t r u c t i o n by

t r a p p i n g ( H n r r i s t r a p s ) h a s been s u c c e s s f u l i n c o n t r o l l i n g G . i ; a P l i -

d i p e s i n a r e s t r i c t e d b r e e d i n g a r e a o f Z u l u l a n d , S o u t h A f r i c a , b u t

r e s u l t s have n o t been v e r y s a t i s f a c t o r y i n o t h e r a r e a s , Some a u t h o r s , '

l i k e KORBIS (1960, 1961) stress t h a t t r a p s g i v e t h e b e s t way f o r c a t -

c h i n g r e p r e s e n t a t i v e s a m p l e s of f l i e s , whereas o t h e r s , l i k e A B E D I (

1963) and FOSTER (1964) , c o n s i d e r t r a p s a s v e r y i n f e r i o r t o d i r e c t

c a t ' c h e s . R e s u l t s depends c e r t a i n l y o n the ' s p e c i e s i n v o l v e d and on t h e

e n v i r o n m e n t a l c o n d i t i o n s . 4.2. T s e t s e f l y c o n t r o l by game d e s t r u c t i o n . ~, ,

It h a s been o b s e r v e d i n t h e p a s t t h a t t s e t se f l i e s and & E -

m e a r e o f t e n a s s o c i a t e d , and t h a t game e l i m i n a t i o n , d u r i n g r i n d e r p e s t L.

e p i z o o t i c s f o r example , are f o l l o w e d by strong r e d u c t i o n o r e v e n disci:

p e a r a n c e o f G l o s s i n a p o p u l a t i o n s (GLkSGOb7 ¿3 WBLCH, 1962 - U00FF,L565) So gaine d e s t r u c t i o n h z s been used a s a r o u t i r i c method f o r tsetse f l y

c o n t r o l i n B a s t e r n and 3 o u t h - e a s t e r n A f r i c a , aiid hcs s u c c e e d e d i n

c l e a r i n g mmy t h o u s a n d s o f s q u a r e k i l o m e t r e s of f l i e s b e l o n g i n g t o

t h e m o r s i t a n s g r q u y , t h e b e s t known e x p e r i m e n t be ing t h e Sh inyanga

o n e , i n T a n z a n i a ( @ L k S G O % ~ , 1 9 6 0 ) . T h i s method, which i s a lways u s e d

i n so ine r e s t r i c t e d a r e a s , i s riot c h e a p ; f o r exemple fn S o u t h e r n

18

X h o d e s i a , d u r i n g t h e y e a r 1955, s l i g h t l y more t h a n 4P.000 h e a d s o?

game were d e s t r o y e d by 800 p a i d h u n t e r s , w i t h au e x p e n d i t u r e o f W 7 r 0 0 Q

r o u n d s o f ammuni t ion ( CBOEEBY, 1956 )

J u d i c i o u s l y a p p l i e d , game d e s t r u c t i o n c a n be a n e c o n o m i c e 1

and p r a c t i c a l means of t r y p a n o s o m i a s i s c o n t r o l , a s n o t o n l y G l o s s i n a h o s t s , b u t a l s o a n i m a l r e s e r v o i r s of t r y p a n o s d m e s a re d e s t r o y e d in on-

l y one o p e r a t i o n . IZowever, as s t r e s s e d by WBHTZ (1963) and b y GLQBriSP,

(%!264), t h e p a l p a l i s group o f t s e t se f l i e s c a n n o t be s t a r v e d e x c e p t if

r e p t i l e s and humans a r e a l s o e r a d i c a t e d , and t h e c o n t r o l by s t a r v a t i o n

o f many s p e c i e s , s u c h a s G . s w y n n e r t o n i , G . a u s t e n i , G . f u s c i p P e u r i s , G . I_

t a b a n i f o r m i s , G . ano r s i t ans and G . s u b m o r s i t a n s , s u p p o s e s t o t a l e1 i :n ina-

t i o n o f s u i d s w h i c h c o n s i t u t e a . g r o u p more d i f f i c u l t t o l o c a t e and

t b k i l l t h a n a n t e l o p e s and big game. B e s i d e s , a s u n d e r l i n e d by DASMAPJN

(1962) and by HQCIZXNG & al. ( P 9 6 3 ) , gaine farming car, b e more p r o f i t a -

b l e i n some m a r g i n a l a r eas t h a n c ~ . t t l e . - r a i q i n g a s a s o u r c e o f meat and

s k i n s and a l s o f o r t h e t o u r i s t i n 9 u s t r y . S o t s e t se c o n t r o l by gaine d e s -

q r u c t i o n is more and more r a r e l y u s e d now . i 3 . G . T s e t s e f l y c o n t r o l by bush c l e a r i n g .

The t s e t s e a b i l i t y t o w i t h s t a n d s e a s o n a l l y u n f a v o u r a b l e c l i -

mate d e p e n d s upon t h e p r e s e n c e o f a f a v o u r a b l e v e g e t a t i o n community

(MkSN,l935). Eushes and trees a r e w i d e l y u s e d by tsetse f l i e s a s res-

t i n g - p l a c e s ; moreover t h e y p r o v i d e a more o r l e s s c o n s t a n t m i c r o c l i m a -

t e e n s u r i n g t h e s u r v i v a l o f a t l e a s t a few f e m a l e s e v e n t h r o u g h t h e

most severe d r y s e a s o n s (PiAsII & 3 A G E , 1 9 5 3 ) ; t h e y c o n s t i t u t e also v e r y

o f t e n v i t a l s h e l t e r s f o r pupae w h i c h c a n o n l y s u r v i v e i f ' t h e soil i s

s u f f i c i e n t l y s h a d e d , p r o t e c t e d f rom d i r e c t i n s o l a t i o n and p r e s e r v e d

f rom r a p i d v a r i a t i o n s of h u m i d i t y ànd t e m p e r a t u r e . "Jan c a n n o t e a s i l y

a l t e r t h e c l i m a t e b u t h e c a n , a t l e a s t i n s a v a n n a h a r e a s , a l t e r t h e

v e g e t a t i o n c o m m u n i t i e s t h a t e n a b l e t h e t s e t se f l i e s t o w i t h s t a n d un-

f a v o u r a b l e p e r i o d s of t h e y e a r . The c l e i r i n g of a l l woody v e g e t a t i o n i n a f l y - i n f e s t e d a r e a

has 3 a r e l y been u s e d f o r t s e t s e f l y e x a d i c a t i o n , b u t h a s been e x t e n s i -

v e l y employed t o p r o v i d e b a r r i e r a r e a s o r t o r e d u c e t h e c o n t a c t b e t - , .

' w e e n f l i e s a n d humans a r o u n d v i l l n g e s and water h o l e s , d t r i v e r - c r o s -

s i n g s , and a long t h e main r o u t e s of communica t ion of humans and c a t t l e

( L E I tOUZIC, 1968) . Such c l e a r i n g s , c a l l e d " a g r o n o m i c a l $ r o p h y l a x i s "

i n f r e n c h - s p e a k i n g c o u n t r i e s , c a n be u s e d f o r c u l t i v s t i o n t o b e k e p t

12 . , c l e a r ' o f trees and b u s h e s o They c a n n o t be e x t e n d e d o n l a r ' g e a r e a s

i n s p a r s e l y i n h a b i t e d z o n e s b e c a u s e t h e y r e q u i r e a l a r g e r manpower

s u p p i y t h a n a v a i l a b l e and a r e v e r y c o s t l y i f c a r r i e d o u t by admin i s -

t r a t i v e a g e n p i e s . - .owever, f o r s p e c i a l p u r p o s e s , c l e a r i n g s and p r o t e c -

tive bar r ie rs have been done up t o 100 k i l o m e t r e s l o n g and th ree k i l o -

Metres w i d e , a s i n S o u t h e r n R h o d e s i a .

L - I

T I

I

S e l e c t i v e c l e a r i n g has been much more e x t e n s i v e l y u s e d f o r

tsetse f l y c o n t r o l . It i s b a s e d o n t h e t e n d e n c y o f f l i e s , i n unfavou-

r a b l e s e a s o n s , t o c o n g r e g a t e i n pe rmanen t b r e e d i n g or r e s t i r i g s i t e s

t h a t co inp r i se i d e n t i f i a b l e p l a n k communi t i e s and a c o m p a r a t i v e l y s m a l l

p r o p o r t i o n o f t h e b u s h - o r woodland a s a w h o l e , The r e q u i r e a e n t s of

t h e common tsetse f l y v e c t o r s of t r y p a n o s o m i a s i s have beall d e s c r i b e d

and t h e t y p e s of v e g e t a t i o n most f a v o u r a b l e as r e f u g e s a re known.They

must be l o c a t e d b e f o r e tsetse f l y c o n t r o l m e a s u r e s are u n d e r t a k e n h e -

l e c t i v e c l e a r i n g c a n be done d i r e c t l y , o r by t h e use o f c h e m i c a l s ( d e -

f i > l i a n t s , t r e e - t o x i c a n t s , . . ) ; i t s u f f e r s f rom t h e s a i n e d e f i c i e n c i e s

as t o t a l c l e a r i n g ; i t i s a c o s t l y measure, t h e r e g r o w t h of bush m u s t be k e p t down and i t is much more e f f i c i e n t i n d r y a r e a s t h a n i n humid

o n e s where i t d o e s n o e l p r e v e n t tsetse f l i e s o f t r a v e l l i n g and r e s t i n g

. .

(PAGE, 1961 b- FOSTER, 1964).

I n d r y c o u n t r i e s t e m p o r a r y c o n t r o l o f r i v e r i n e t se t se f l i e s

- h a s been a t t e m p t e d ; o n c e by u s e of " o b s t r u c t i v e c l e a r i n g " , by f e l l i n g

t h e trees f o r m i n g t h e o v e r h e a d canopy and b l o c k i n g t h e s t r e a n - b e d t o

o b s t r u c t t h e t se tse f l i g h t - l i n e . The f l i e s c o u l d n o t move f r e e l y un-

d e r s h a d e and e i t h e r were s t a r v e d o r f o r c e d i n t o t h e open where t h e

climate d u r i n g t h e d r y s e a s o n w a s i n t e l e r a b l e (NhSM ¿i STlBIBTBR, 11957),

W i t h t h e d i s c o v e r y of modern i n s e c t i c i d e s i t a p p e a r s t h a t

i t i s g e n e r a l l y more e f f i c i e n t eo s p r a y a r e s i d u a l t o x i c a n t on t h e tse

tse p e r m a n e n t r e s t i n g - s i t e s t h a n t o d e s t r o y t h e m , b e c a u s e tsetse can

change o f r e s t i n g - s i t e s when . t h e p : ?e fe r r ed o n e s a r e d e s t r o y e d , a s see21

i n Ankole d i s t r i c t o f Uganda (HAR@BY"& FILSOM,1961), w h e r e a s t h e y do

n o t abandon t h e m a f t e r s p r a v i n g (HOCIIIPITG, 1964) . 4.4. T s e t s e fly c o n t r o l by i n s e c t i c i d e s .

I

I n s e c t i c i d e s c a n be u s e d as aerosols of t e a p o r a r y e f f i c a c y

on huge> areas, o r a s r e s i d u a l s p r a y s o n r e s t i n g p l a c e s , t r a p s and

ba ' i t s

13

4,d.l. I n s e c t i c i d e a e r o s o l s . I n s e c t i c i d e a e r o s o l s havi2 a v e r y s h o r t r e s i d u a l . e f f e c t , i f

any a t all, and k i l l tsetse f l i e s 2 u r i n g some h o u r s o n l y a f t e r t h e i r I

. a p p l i c a t i o n , They c a n c o n t r o l or e r a d i c e t e t h e f l i e s o n l y i f t h e y a r e 1 ,

a p p l i e d on l a r g e a reas a t c o n v e n i e i i t i n t e r v a l s t o k i l l , b e f o r e t h e i r

- r e p r o d u c t i o n , a l l f l i e s h a v i n g emerged from pupae s i n c e t h e p r e v i o u s 4 a p p l i c a t i o n s . c o n v e n i e n t rhy thm seem t o be a b o u t s i x t o e i g h t a p p l i s -

c a t i o n s at i n t e r v a l s of two t o fou:. w e e k s (BURNBTT, 1962 b ) , e c c o r d i n i : I t o t h e b i o l o g y of t h e l o c a l p o p u l a C i o n cf tsetse f l i e s

A i r c r a f t a p p l i c a t i o n h a s t h e o b v i o u s a d v a n t a g e o f c o v e r i n g

q u i ç k l y l a r g e areas , and c a n be e f f i c i e n t a g a i n s q s a v a n n a h - i n h a b i t i n g i

s p e c i e s l i k e G . m o r s i t a n s , G . s w y n n e r t o n i and G . p a l l i d i p e s ; i t ' i s less

a d e q u a t e t h a n grounc? j p y l i c a t i o n a g a i n s t r i v e r i k e t s e t s e f l i e s l i k e '

- . G , p a l p a l i s and G ; $ u s c i p e s - (ÈURN;ET.T, 1962 b ) ; and i s a l m o s t i m p o s s i b l e

t o c a r r y o u t a g a i n s t s p e c i e s l i v i n g i n h i g h f o r e s t . Grou,rid a p p l i c a t i o n

c a n b e Idone e i t h e r . w i t h l o r r y - c a r r i e d . g e n e r a t o r s ; o r w i t h l i g h t p o r t a - .

I b l e m i s t - b l o w e r s (TWOPdSON & al., 1960 - CIIALLIBB & a l . , 19%) ,

I n s e c t i c i d e a e r o s o l s c a n o n l y be a p p l i e d some h o u r s a d a y ,

j u s l a f t e r dawn and before d u s k , a n d , sometimes by n i g h t , w h e n - a i r c u r -

r e n t s a r e I duwnwardG ;: dFzrir,g Othe - day - t ime p e r i o d s t h e i n s e C s t i c i d e

c l o u d d i s p e r s e s v e r y r a p i d l y a n d , if a p p l i e d by ai'roGaf t , does3 -no t

e v e n r e a c h i r o u n d l e v e l , Only a v e r y small amount o f ' i n s e c t i c i d e r e a - . .

. -, I ches e a c h i n d i v i d u a l f l y and g r a v i d f e m a l e s , l e s s s u s c e p t i b l e t o i n -

s ec t i c ides t h a n snales , and f e m a l e s of o t h e r p h y s i o l o g i c a l c o n d i t i o n s

(BURNETT, 1?6,2 a)., are n o t e a y i l y k i l l e d by c h l o r i n a t e d i n s e c t i c i d e s ;

some 0 . F . compounds l i l c r ; f e n t h i o n or m a l û t h i o n m i g h t p e r h a p s be m o r e -

e f f i c i e n t , GHQCHCING & a l . , 1983) . I

For e r a d i c a t i o n programmes d j e l d r i n , E.H.C. and t e l o d r i n i a e r o s o l s h a v e ' b e e n u s e d m a i n l y i n B a s t A f r i c a , w i t h v a r i a b l e r e s u l t s

i n c l u d i n g fair succes (-COCKBILL & a l . , 1963 - BUICKX, 1965) . Tech.ni-

c a l improvemen.ts i n - t h e s p r a y i n g - d e s i g n s have p e r m i t t e d t o u s e o n l y

v e r y s h a l l amounts o f i n s e c t i c i d e s , a v e r a g i n g i , 5 kg. of t e l o c l r i n , o r

4 ,5kg ,- of d i e l d r i n . ' b e r square kilometre. T h e s e t e c h n i c a l improvemen t s

and a b e t t e r k n o w l e d g e - o f tse%se e c o l o g y have decreased t h e c o s t of

s u c h p rogrammes ; 'wh ich w a s very heavy for t h e f i r s t e x p e r i m e n t s , t o

a r o u n d 860 t o '1.000 f r e n c h f r a n c s p e r s q - k i l o m e t r e now (BURNETT,1962 -. HOCICIHG & al,, 1.963 - EURNETT i$ a l , , 1964). The c o s t o f o p e r a t i o n s ;d5,

be r e d u c e d i f a e r o s o l s a re u s e d o n l y for s h o r t t e r m c o n t r o l ' o f tsetse,

f l y p o p u l a t i o n s i n s l e e p i n g - s i c k n e s s f o c i ,

c

.-

I I ( i , , I *

14 a . _

4.4 -2. R e s i d u a l t r e a t m e n t of v e g e t a t i o n , a ,

R e s i d u a l t r e a t m e n t of t s e t s e f l y r e s t i n g - p l a c e s must be l e -

t h a l f o r t h e f l y on s h o r t c o n t a c t f o r a l o n g e r p e r i o d t h a n t h e maxi-

mum d u r a t i o n of t h e p u p a l l i f e . I n s u c h c o n d i t i o n s o n l y one s p r ' a y i n g

may be s u f f i c i e n t t o c o n t r o l t h e s p e c i e s , and p e r h a p s e r a d i c a t e i t , i n . Y

. . an i s o l a t e d area . . The f i r s t r e s i d u a l a p p l i c a t i o n s have been aone a g a i n s t r i -

v e r i n e s p e ç i e s , l i k e G . p a l p a l i s and G . f u s c i p e s , w i t h h a b i t a t s r e s t r i c -

t e d t o water e d g e . I n l a r g e r g a l l e r y f o r e s t s i t Is somet imes p o s s i b l e

t o open pf i ths i n t h e f o r e s t , which will b e e x t e n s i v e l y used by moving

f l i e s ; t h e t r e a t m e n t o f sÚch p a t h s may be s u f f i c i e n t t o g e t t h e con-

t r o l o f t h e f l i e s . The same o b s e r v a t i o n s haGe , b e e n made i n dense b u s h ,

where s e v e r a l s p e c i e s , s u c h a s G . m c : r s & t a ~ s "- and G . p a l p a l i s , f r e q u e n t

thrdu 'gh th 'e budh; c o n d e n t r a t i o n s oi t h e f l i e s ' o c c u r a t ehe i n t e r s e c -

t i o n s ofl t h e p a t h s ; t h e c o r r e s p o n d 2 n g method of t se tse c o n t

s e d ' on t h e open ing o f p k r a l l e l p a t t s , s o le h u n d r e d s o f m e t r t h r o u g h d e n s e t h i c k e t s and wooded r r e a s , and by t r e a t i n g , n o t h i n g b u t

t h e p a t h s ; t h i s method h a s l b e e n f a i r l y s a c c e s s f b l i n s e v e r a l a r e a s o f

Kenya, a g a i n s t G . p a l l i d i y e s , G . f u s c i p e s , and G . b r o v i p a l p & (l!BOr!iS@N &

al., 1960 ) .

' p i t h s made by man and á n i n a l s and f o l l o w a v e n u e s and c h a n n e l s o c c u r i n g

' I

DUT s u s p e n s i o n s arid e m u l s i o n s , wh ich have been u s e d i n t h e

f i r s t e x p 6 r i m e n t s , have u s u a l l y been r e p La'ced by d i e l d r i n e m u b i a n s ,

which a r e assumed to b e e f f i c i e n t a l m o s t o n e y e a r , and somet imes more

t h a n one - y e a r i f a p p l i e d a t 4% (KBZNAGHAN, 1962) ' . However, a c c o r d i n g

t o BALDBY (1963 & 1964), DDT e m u l s i o n s have a l o n g e r r e s i d u a l e f f e c t

Chan d i e l d r i n and t e l o d r i n ( d s o b e n z a n ) o n e s when a p p l i e d a t ;he con-

c e r i t r a t i o n of 5% and e v e n , i n soglie c o n d i t i o n s , a t c o n c e n t r a t i o n s a s

low a s 2 , 5 % ( D B V I B S , L S 6 4 ) . Such c o n t r o l p r o c e d u r e s a re f i v e t a t e n

t i r n e s c h e a p e r t h a n bush c l e a r i n g and of a more pe rmanen t e f f e c t (Hoc- KPPaG, A 9 6 3 - '*acLEPJNAN & AHPCHZSON, 1 9 6 3 - CHkELIBG, 1965 a )

-

Improvement on t h e knowledge o f t s e t se f l y r e s t i n g - p l a c e s

h a s p e r m i t t e d t h e s u s e o f r e s i d u a l i n s e c t i c i d e s a g a i n s t s avannah spe -

c i e s l i k e G ,mors i t a r i s a n d . G , s w y n n e r t o n i , by s e l e c t i v e s g r a y i n g of t h e

$ i o d e r s i d e of t h e b r a n c h e s o f A c a c i a t recs i n f l y - c o n c e n t r a t i o n t a r e a s {

(He)CEING, 19611 - CBAlYbJICIC, 1964 - CI-IJXJbJIClf & $ a l , , 1564 - WOOFF,1965) The c o s t of t h e c o n t r o l i s t h e n s i m i l a r t o , or &lower t h a n , b y a i r c r a f t

f o g g i n g , b u t t h e work i s m o r e . d i f f i c a l t t o p l a n and s u p e r v i s e . I n t h e

I 1

15

d r y s e a s o n G . s u b m o r s i t a n s behave s o m e t i n e s a l m o s t l i k e a r i v e r i n e

s p e c i e s and c a n be c o n t r o l l e d by r e s t r i c t e d i n s e c t i c i d e s p r a y i n g s

(hlAt3LElYfdkP?, k965 - TE’*PLBTON, ,965 - JOL’,DAN, 2.965) . B e s i d u a l i n s e c t i c i d e s have been u s e d a l s o a g a i n s t G . t a c h i -

n o i d e s i n N o r t h e r n P f i g e r i n and a l o n g t h e Tchad-Cameroons b o r d e r , b u t

t h i s f l y i s n o t as c o n c e n t r a t e d as G . p a l p a l i a a l o n g w a t e r - p l a c e s ; s o

trees and b u s h e s must be t r e a t e d on a l a r g e r w i d t h on b o t h s i d e s o f

t h e r i ve r s and w a t e r - h o l e s (ï?OUCEET & a l . , 1 9 6 1 - DAVTBS, 1964 - &!!ac--

I-

LENNAN, 1965 - TIiYPLBTON,1965) * With t h i s s p e c i e s t h e main d i f f i c u l -

t y is t h e e x a c t s e l e c t i o n o f t h e v e g e t a l communi t i e s t o be s p r a y e d ,

wh ich i m p l i e s a v e r y c l o s e s u p e r v i s i o n of t h e s p r a y i n g - t e a m s , and t h z

a c c u r a t e t i m i n g o f t h e programme, be tween t h e end o€ f l o o d s and t h e

b e g i n n i n g of t h e r a i n s , f o r t h e b r e e d i n g - s i t e s n e i g h b o u r i n g t h e Chad

Lake b

R e s i d u a l i n s e c t i c i d e s a r e o n l y p r o m i s i n g when t se t se f l y

h a b i t a t s are r e s t r i c t e d , s o t h e y have n o t been u s e d a g a i n s t h i g h fo-

r e s t s p e c i e s . However t h e y have been s p r a y e d w i t h f a i r success a g a i n t s

G . fusca and G . f u s c i p e s i n l a r g e g a l l e r y f o r e s t s of t h e C e n t r a f r i c a n

H e g u b l i c (FINBLLE & a 1 . , 1 9 6 2 & 1963 - YVOiiB & a l . , 1 9 6 2 ) . 4.4.3. I n s e c t i c i d e - t r e a t e d t r a p s and b a i t s ,

R e s i d u a l i n s e c t i c i d e s have been s p r a y e d i n s i d e t s e t se fly t r a p s t o i n c r e a s e t h e i r e f f i c a c y w i t h l i m i t e d r e s u l t s . I n s e c t i c i -

des h a v e also been u s e d i n c o m b i n a t i o n w i t h a t t r a c t a n t s , on t r a p s , a n *

t h e method i s a l w a y s u n d e r i n v e s t i g a t i o n . ’ S t u d i e s c a r r i e d o u t i n A r u -

s c h a ; T a n z a n i a , have shown t h a t aqueous e x t r a c t s of pig s k i n s and

b r i s t l e s a r e v e r y a t t r a c t i v e f o r G . p a l l i d i p e s , b u t n o t f o r G.swgnner to

mi; t h e a c t i v e p r i n c i p l e i s s o l u b l e i n l i p i d s o l v e n t s ; t r a y s t r e a t e d wieh t h i s a t t r a c t a n t c a t c h two t i m e s more G 1 . p a l l i d i p e s t h a t u n t r e a -

t e d o n e s and c o u l d be u s e d i n c o m b i n a t i o n w i t h e i t h e r i n s e c t i c i d e s

o r c h e m o s t e r i l a n t s (kNONYYOUS, 1 9 6 5 ) . 4.5. B i o l o g i c a l c o n t r o l .

V a r i o u s a t t e m p s h a v e been made i n t h e p a s t t o m u l t i p l y and

r e l e a s e n a t u r e l t s e t se f l y p a r a s i t e s (JENKINS, 1964) t o c o n t r o l t h e -

se f l i e s ; b u t t h e r e s u l t s h a v e n o t been p r o m i s i n g . ‘Trials on h y b r i d i z a t i o n , by c r o s s - m a t i n g c l o s e l y r e l a t e d

s p e c i e s , t o p r o d u c e a h i g h d e g r e e of s k e r i l i t y i n a f l y p o p u l a t i o n ,

h a v e n o t been s u c c e s s f u l ( G Z A S 6 0 3 , 2960) and assume t h e p o s s i b i l i t y

of r a i s i n g l a r g e n u n b e r s of t s e t s e f l i e s .

16

S t e r i l i z a t i o n o f f l i e s i s p o s s i b l e , e - i t h e r by r a d i a t i o n

- o r w i t h c h e m o s t e r i l a n t s ( I I N I P L I N G , 3.965 - SN1TI-l & DAME, 1963), and

f i e l d and l a b o r a t o r y i n v e s t i g a t i o n s h a v e been c a r r i e d o u t . The p u b l i e

hed r e s u l t s are p a r t l y c o n t r a d i c t o r y . A c c o r d i n g t o POTTS ( f 9 6 4 ) , m n l e s

s t e r i l i z e d by gamwfia i r r a G i a t i o n a r e n o t c o m p e t i t i v e w i t h no rma l males

t h e s t e r i l i z i n g d o s e s b e i n g n e a r l y t h e safile a s t h e l e t h a l o n e s ; howe-

ver DAhIB & a l . (1965) have found t h e p n d i o s t e r i l i z e d m a l e s v e r y com-

p e t i t i v e , and h a v i n g a n o r m a l s u r v i v a l r a t e d u r i n g t h e i r f i r s t t h r e e -

weeks of l i f e CIIADNPCK (i964) h a s s t r e s s e d t h a t c h o m o s t e r i l a n t s a r e

n o t much more p r o m i s i n g t h a n r a d i a t i o n s ; i n h i s e x p e r i m e n t s the t r e a

fflent of males a l o n e r e d u c e s o n l y by 40% t h e a v e r a g e iîwnber o f produ-

c e d p u p a e , apd t h e s u r v i v a l r a t e of t r e a t e d males was c o n s i d e r a b l y

r e d u c e d ; t h e t r e a t m e n t of f e m a l e s was more e f f i c i e n t , b u t i t seems

d i f f i c u l t t o reach t h e f e m a l e component of w i l d t s e t s e f l y p o p u l a -

t i o n s . N e v e r t h e l e s s D A ' T i% a l . (1965) h a v e p u b l i s h e d t h a t Tepa - and

M e t e p a - t r e a t e d males a re a l m o s t a l l s t e r i l e and t h a t t h e i r s u r v i v a l

r a t e is a l m o s t n o r m a l , t h e c h e m i c a l s b e i n g a p p l i e d e i t h e r o n young

n a l e s o r on pupae . I f a n e f f i c i e n t c o n t r o 1 method i n v o l v i n g chemos-

t e r i l a n t u s e i s d i s c o v e r e d , t h e p r o b l e m o f l a b o r a t o r y m a s s r e a r i n g o f

t s e t s e f l i e s would h a v e t o be s o l v e d , w h i c h w i l l n o t b e an e a s y t a s k

(P!AILL,C)T, 1958 - MASH, i 9 6 3 - EVENS, 1964); w i d e s"ca1e e x p e r i m e n t s

are c a r r i e d o u t i n S o u t h Xhodes i a t o d e v e l o p more e f f i c i e n t methods

of tsetse m a s s - r e a r i n g t h a n these a c t u a l y a v a i l a b l e .

. . L I

L

. .

6 . HEARING FKOBLBfJIS B14D RESEARCH NEEDS . All tsetse ' f l y c o n t r o l me thods s u p p o s e a v e r y tho.rough knc

l e d g e of t h e phys i lo logy and e c o l o g y of t h e i n v o l v e d s p e c i e s . N o r e o v e r :

* i f c h e m o s t e r i l a n t s a r e p r o m i s i n g m a s s - r e a r i n g t e c h n i c s arid mat ing-ha-

b i t s mus t be s t u d i e d . I m p r o v e r e n t o f o u r knowledge on t h e b i o l o g y a n d

e c o l o g y of t se tse f l i e s r e q u i r e s mass m a r k i n g , r e l e a s e and r e c a p t u - re e x p e r i m e n t s wh ich also ; imp ly m a s s - r e a r i n g of s t a r i d a r d s p e c i , . vrlens .

The r e p r o d u c t i o n r a t e of . G l o s a i n a is v e r y low . ,Fqr example.

G . m o r s i t a n s d r o p s a l a r v a e v e r y e i g t h d a y a t 30°6., e v e r y e l e v e n t h

clay a t 2dQ and e v e r y t w e n t y f i f t h day a t ICo ( J A @ K , 1 9 3 9 ) . The b e s t eli-

matic c o n d i t k o n s , f o r G l o s s i n a c o l o n i e s h a v e been d e f i n e d by 7;IBLLkNB'IIT

(1937) ; t e m p e r a t u r e must be m a i n t a i n e d be tween 35O and 27OC. f o r almoe".

.. a l l s p e c i e s ; r e l a t i v e h u n i i d i t y must b e h i g h , r e a c h i n g a l m o s t s a t u r a -

t i o n , f o r t h e r i v e r i n e s p e c i e s , s u c h as G .pa lpa1 i . s . The main te ,nance of

a d u l t s h a s been done i n s e v e r a l t y p e s of c e n t a i n e r s , i n c l u d i n g test-

t u b e s (FOSTER, 1C35'9), b u t f o r m a s s r e a r i n g c a g e s are more h a n d a - b ,Le;

17 B r u c e ' s and Ploubaud's s m a l l - s i z e d c a g e s have been u s e d a s w e l l a s

b i g g e r P e t a n a ' s cage . P r e l i m i n a r y t r i a l s c a r r i e d o u t by one o f u s

( A .C. ) h a v e shown t h a t e n v i r o n m e n t a l c o n d i t i o n s s u c h a s l i g h t C n g

s h a p e and volume o f t h e c a g e s , and f l y . p o p u ï a t i o n . d e n s i t y , may be of

a p r i m a r y i m p o r t a n c e t o e u s u r e no rma l l o n g e v i t y , b e h a v i o u r . and r e p r o -

d u c t i o n r a t e ,. d High i n s e m i n a t i o n rates !nay be o b t a i n e d i n c o l o n i e s , rea-

c h i n g 97% for G , m o r s i t a n s ( FOSTEE, 1957) and 88% f o r G . i ~ a l ~ a l i s

(PJASB, f955), b u t t h e s tBr i le -male method s h a l l r e q u i r e a l s o a p e r -

f e c t knowledge of t h e making b e h a v i o u r . Some da ta are a l r e a d y a v a i - h a b l e f o r G . a o r s i t a n s and G . p n l p a l i s (FOSTER, l957 - NASR,195S-JOEDkT

.L95S), b u t s e v e r a l o t h e r e c o n o m i c a l l y i m p o r t a n t G l o s s i n a s y e c i e s ha-

ve n o t been s t u d i e d y e t .

'

F e e d i n g the a d u l t f l i e s i s a v e r y i m p o r t a n t p rob lem. Too

f r e q u e n t b l o o d m e a l s b r i n i on t h e g r o d u c t i o n o f s m a l l l a r v a e and i n d u -

ce a b o r t i o n ; t h e . no rma l f r e q u e n c y o f f e e d i n g i s t w o o r t h r e e t i n e s b e t -

ween two s u c c e s s i v e l n r v i p o s i t i o n s . G u i y i e a p i g s have been u s e d a s 1

. h o s t s f o r a l o n g t i m e , b u t s h e e p i s a b e t t e r h o s t , e n s u r i n g a , b e t t e r

l o n g e v i t y and a h i g h r e p r o d u c t i o n r a t e p f the f e m a l e f l i e s , aïid now

g o a t s are also u s e d w i t h s u c c e s s (NASH,.& IZEhlMAGHAN, 1964) . Xepea ted

tsetse f l y b i t i n g b r i n g s o u t a s k i n r e a c t i o n o f t h e h o s t s ; howover

HASII & KBRNkGHliM (f965) have develoRed a method a l l o w i n g t o f e e d 436

f l i e s e v e r y three d a y s on t h e same g e a t . I n v e s t i g a t i o n s are a l s o car

r i e d o u t i n - l h i n g t h e r e p l a c e m e n t of t h e h o s t by d e f i b r i n a t e d b l o o d :

t se tse f l i e s f e e d i n g t h r o u g h a aembrane (SOUTHON & COCBHNG,LS63~:a &

1963 b, BIKDBW & B k R L B Y , 1 9 6 5 ) . Wo may hope t h a t improvements of t h e

f e e d i n g p r o c e d u r e s w i l l e n a b l e t h e p r o d u c t i o n of l a r v a e a n d pupae of

no rma l w e i g h t , w i t h o u t any h i g h m o r t a l i t y of t h e newly emerged f l i e s : ,

, . which i s n o t a lways t h e case, i n t se tse f l y l a b o r a t o r y c o l o n i e s (H~ASIJ.

r & KBRNAGBAL'J, 1966 ) . Emergence r a t e f rom t h e pupae i s a l s o en i m p o r t a n t f a c t o r i r 1

t h e e s t a b l i s h m e n t o f a G l o s s i n a c o l o n y . k v e r y h i g h r a t e h a s been ob-

t a i n e d when l a r v i p o s i t i o n o c c u r s i n a l a y e r of d r y s a n d j u s t o v e r

w e t s a n d ( N A S H & al., 1950), for G o y a l p a l i s , b u t t h e p r a b l e m has t o

b e s o l v e d f o r t h e o t h e r s p e c i e s .

F

18

7. DXSCUSSION AND CONCLUSIOMS . !

S e v e r a l methods a r e a v a i l a b l e f o r c o n t r o l l i n g t s e t se f l i e s ,

b u t a l l a r e e x p e n s i v e and nane of th.em i s y e t a v a i l a b l e f o r d e n s e l y f o r e s t e d a r e a s . B e s i d e s t s e t s e f l i e s c a n r e i n v a d e c l e a r e d a r eas i f these

a reas a re n o t i s o l a t e d f rom n e i g h b o u r i n g t s e t s e f o c i , e i t h e r by n a t u -

rai b a r r i e r s o r by s p r a y e d o n e s . T s e f s e f l y c d n t r o l , t h e r e f o r o , m u s t . be p l a n n e d as a l o n g t e r m programme, on a s a w i d e t e r r i t o r i a l basis a s

' p o s s i b l e Pn many I n s t a n c e s e r a d i c a t i o n i s c h e a p e r t h a n c o n t r o l , on a

l o n g r a n g e , t h e new t r e a t e d a r e a s ' p r o t e c t i n g t h e p r e v i o u s l y t r e a t e d

o n e s . J t

S u r v e y s show t h a t w h e r e huaan p o p u l a t i o n d e n s i t y reaches 40

i n h a b i t a n t s p e r kin2 t h e a n i m a l h o s t s o f . t se t se f l i e s be l -unging t o t h e

f u s c a and m o r s i t a n s g r o u p s d i s a p p e a r , and most of t h e G l o s s i n a r e s t i n g

p l a c e s are c l e a r e d ; i n s u c h c o n d i t i o n s t s e t s e f l y v e c t o r s o f aïrimal t r i

p a n o s o m i a s i s a r e u s u a l l y no l o n g e r a p rob lem (FCEZD, 1962), e x c e p t if

"holy W0odlands" o c c u r , l i k e i n t h e Moss i c o n t r y of Nes t A f r i c a ; b u t

s u c h wood lands c a n be e a s i l y c l e a r e d from G l o s s i n a b y i n s e c t i c i d e s . C i -

a ,

I

v e r i n e s p e c i e s of G l o s s i n a , w h i c h t r a n s m i t m a i n l y human s l e e p i n g - s i c k -

- n e s s c a u s e d by T.gambiense , c a n s u r v i v e a l o n g r ive r s b u t a r e , v e r y e a s i

ly c o n t r o l l e d i n s u c h r e s t r i c t e d h a b i t a t s .

.When hunan d e n s i t y I s b e l o w 40 i n h a b i t a n t s p a r km2 t s e t se fl

c o n t r o l becomes more and more d i f f i c u l t and c o s t l y w i t h t h e s c a r c i t y

o f i n h a b i t a n t s . D e s p i t e our t e c h n i c a l a b i l i t y t o combat t se tse , f l i e s ,

. i t i s s t i l l n o t w o r t h w h i l e t o u n d e r t a k e l a r g e reclamation schemes , ex-

c e p t i n s p e c i a l i n s t a n c e s , where s o i l s and c l imate ax?e:&avourable t o

i n t e n s i v e a g r i c u l t u r e (COCKBILL & alr,L963), o r w,hen p u b l i c h e a l t h i s

, i n v o l v e d .such as i n r e s i d u a l f o c i of s l e e p i n g - s i c k n e s s (W.X:KJS,1962),

o r f o r p r a t e e t i o n o f c a t t l e d u r i n g s e a s o n a l m i g r a t i o n s (FINHELE & a l I I

Ce962 & 1965) I

I n o t h e r s i t u a t i o n s i n s u f f i c i e n t e x p l o i t a t i o n of t h e c o u n t r y

W i l l p e r m i t tsetse f l i e s t o r e p o p u l a t e t h e c l e a r e d a reas s o o n e r o r l a -

t e r , and t h e r e s s o u r c e s employed f o r t se t se fly c o n t r o l w i l l hav? been

, L

wast'eb . .

-. Y 9

ABEDI' ( Z .H , ) , 1963 .-' T s e t s e s n m p l h e me,thods . knn.Xpt . L i b e r i a n 1 n s t . a m e r i c a n Found . t rop.Ked . f o r 1 9 6 2 , 4 3 4 3 . .

k&Dl ( 2 .M. ) & WILLE11 ( L . J . ) ,1964 .- E o a d s i d e t s e t s e h a z a r d i n E i b e -

r i a . Amer .J . t ropOr ed.'%+yg., 1 3 499 - 504 . - 9

kNONYYOUS, 196P-1965.- B a s t A f r i c a n T r y p a n o s o m i a s i s E e s e a r c h Organ i -

F a t i o n Bepor ts , 1960. 3.9GP. f962-63 . & 1963-64, Pub1 . E a s t k f r .High Commissian and Bast Af r .Common Serv ices O r g a n i s a t i o n ,

N a i r o b i . mor?Ymus, i965 ,- T r o p i c a l P e s t i c i d e s R e s e a r c h I n s t i t u t e , Annual ' Repor'

- 0 1964 Pub1 , B a s t A f r .Common S e r v i c e s Ore., Arushrr, 4 5 pp,

BALDRY (D .A .T. ) I 1963 .- An e v a l u a t i o n by b i o a s s a y of the p e r s i s t e n c e

of DDT d e p o s i t s on r i v e r i n e v e g e t a t i o n i n t h e Mor the rn Gu inea

s a v a n n a h v e g e t a t i o n zone of N i g e r i a and o b s e r v a t i o n s on t h e EaL,

tors i n f l u e n c i n g t h e a v a i l a b i l i t y d f d e p o s i t s t o G l o s s i n a pal- p a l i s (R.D.), B u l l . e n t . R e s , , 54, 497 - 508 . . -

BALDRY (B .A ,T . ) , 1964,- An a s s e s s i n e n t by b i o a s s a y o f t h e c o m p a r a t i v e

t o x i c i t y and p e r s i s t e n c e of d i e l d r i n arid isobenzan d e p o s i t s on

r i v e r i n e v e g e t a t i o n i n t h e . U o r t h e r n Guinea.! savannah zone of I ?Ti

g e r i a i n r e l a t i o n t o t h e c o n t r o l of Glossina P a l P a l i s (R.D.). . . B u l l . e n t . R e s , , 55, 4 9 - 52 . I .

B U l C f X ( E a J ) , 2965 .- A p p l i c a t i o n s p a r v o i e a6 r i e r ine d e t k l o d r i n e d a n s

l a l u t t e c o n t r e G l o s s i n a m o r s i t a n s M e s t w . au B u g e s e r a (Rwanda)

c . C.SIH.Bo~r/C.C.~.k., PUb1.97, 1 4 5 - 155

BUHNETT ( G O F . ) , 3.962 a.- The s u s c e p t i b i l i t y of t se tse f l i e s t o t o p i c a l

a p p l i c a t i o n s of i n s e c t i c i ' d e s . 1x1,- The e f f e c t s of age and pre-

gnancy on t h e s u s c e p t i b i l i t y of a d u l t s of G l o s s i n a I i n o r s i t a n s ,

B u l l . e n t 0 R e s , 5 3 , 337 - 3 4 5 . BURNETT (G.P.), 1962 b.- E e s e a r c h i n B a s t A f r i c a o n t h e c o n t r o l of . .

t se t se f l i e s f rom t h e air: A g r i c u l t u r a l A v i a t i o n , 5, 79-87:

I._ ao 1 .

, ' .:t BURNETT (G.F.) , CHADWICK ( i ? . R . ) 3 YTLL5L7 (A.U.D.) & BEESLEU. ( J .S .S . ) ,

-i664.- k i r q r a f t a p p l i c a t i o n s o f i n s e c t i c i d e s i n B a s t A f r i c a .

XIV.- Very-low-volume ' a e r o s o l s of " d i e l d r i n and i s o b e r i z a n f o r

_. .

. .. . . .

the c o n t r o l of G l o s s i n a m o r s i t a n s W e s t w . B u l l . e n t . R e s . , . I

5 5 537 - 5 3 9 . - 9

BURSE,EL (a . ) , 1959 .- The w a t e r , b a l a n c e o f tsetse f l i e s . T r a n s .T: .3nt Sdc.Zond.,, 'kL1, 205 - 235 . .. . -

.. . . ..

BURSELL ( E . ) , í.960.- The e f f e c t o f t e m p e r a t u r e oil t h e c o n s u m p t i o n of

fat d u r i n g p u p a l deve lopmen t i n G l o s s i n a . . B u l l .ent .Zes . , 51 583 - 5 9 8 .

BUIX3EiCL ( B .) , 1965 .- T s e t s e - f l y p h y s i o l o g y : a r e v i e w o f . r e c e i i t advan-

c e s and c u r r e n t aims. Bull.Or6.mond.Saaté0, 2 8 , 1903 - 709, . . ,

( ' F . A . ) , 1955.- The n a t u r a l h i s t o r y o f t se t se f l i e s . Eond, Sch.ISyg. & t rop . iced . , Tliemoir n o i O , 816 p p .

C€-3ADb?ICK ( f . R . ) , i964 a,- iì s t u d y of t h e r e s t i n g sites of G l o s s i n a . ' s w y n n e r t o n i kust. i n 'F l o r t h e r n Tariganyika;

23 - 28 0 '

bull . e n t .Res'. , 55,

. * . > i

" .

CHADWICK (P .E . ) , 1964 b o - S f f e e t of two c h e m o s t e r i l a n t s . on G l o s s i n a i

m o r s i t a n s . N a t u r e , (Lond .) , e, 299 - 3 0 0 . , .

' ($<AD%JICK cl? .E. ) , B s E s L B Y ( J ,Ei .5. ) , WE<STB ( F .J ) & FlATHCZ%(PE .'I' ) ,1964 t

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