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Biological Conservation 48 (1989) 311-319

The Status , Dis tr ibut ion and Conservat ion o f the TigerP a n t h e r a t i g r i s in Bhutan

D a s h o P a l j o r D o r j iMinistry of Social Services, Tashichhodzong, Thimphu, Bhutan

&Charles Sant iapi l la i*

WWF Indonesia Programme, PO Box 133, Bogor, Java Barat, I n d o n e s i a(Received 13 October 1988; accepted 10 November 1988)

A B S T R A C TThe t i ge r P a n t h e r a t i g r i s i s t he t op car n i vor e in Bhu t an , wher e i t occur s f r o mt h e f o o t h i l l s o f t h e H i m a l a y a s a t a n a l t it u d e o f 2 O O m in t h e s o u th t o o v e r3000 m in t he nor th . I t ha s a w i de d i s lr i bu t i on and i t s r ange i nc l udes s avanna hg r a s sl a n d s, s u b - t ro p i c a l a n d t e m p e r a t e f o r e s t s . I t f e e d s o n a v a r i e t y o f p r e ys pec i e s r ang i ng i n s i z e f r o m t he bar k i ng de e r Muntiacus muntjak t o t he gaurBos gaurus. A t t a c k s o n d o m e s ti c y a k a r e n o t u n c o m m o n a n d a re t he c a us e o fcon f l ic t w i t h he r ds men , w ho t r ea t t he t ige r a s v e r mi n a nd e l i mi na t e i t u s ingpo i s on . Ther e a r e e s t i ma t ed t o be be t w een 150 and 250 t i ge r s in Bhu t an . I np r i m e h a b i ta t s, a s i n t h e R o y a l M a n a s N a t i o n a l P a r k , t h e t ig e r ca n e x i s t a t ad e n s i t y o f 4 " 5 1 0 0 k m - 2 . A s w i t h o t h e r l a r g e p r e d a t o r s, t h e b a s is f o rc o n s e r v a ti o n m u s t b e a s e p a ra t io n o f h u m a n s e t t l e m e n t s a n d w i l d l if e re s er v e s.G i v e n t h is b a c k g r o u n d , t h e e m p h a s i s m u s t b e t o m a i n t a i n f o r e s t c o v e r o v erl a r ge a r eas w her e r emo t enes s , d i f f i cu l ty o f te r r a i n and dens i t y o f cover p r ov i den a t u r a l p r o t e c ti o n . N o t a l l s m a l l p o p u l a t io n s a r e n e c e s s a r il y d o o m e d . T h eo v e r a ll p r o s p e c t s f o r t he t i g e r a p p e a r g o o d in B h u t a n p r o v i d e d d e f o r e s t a ti o n iscar e f u l l y con t r o l led . I n o r der t o succeed , p r o t ec t i on o f cons e r va t i on a r eass h o u l d b e a c c o m p a n i e d b y m e a s u r e s t o i m p r o v e t h e l i v i n g s t a n d a r d s o f t h epeople .

* Author to whom reprint requests should be addressed.311

Biol. Conserv. 0006-3207/89/$0350 1989 Elsevier Science Publishers Ltd, England. P r i n t e din Great B r i t a i n

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312 Da sho Pal jor Dor j i, Char les San t iap i l la i

INTRODUCTIONThe tiger P a n t h e r a t i g r i s is the largest carnivore in Bhutan. The widegeographical dist ribut ion (Fig. 1) even within Bhutan underlines itsversatility and ability to adapt to a variety of environmental conditions. Itsonly requirements for survival appear to be some form of vegetative cover,water supply and adequate prey (Schaller, 1967). In Bhutan, it is largelynocturnal and elusive. While these characters are of survival value andtherefore to be welcomed by the conservationist, they also make it difficult tostudy scientifically. There has so far been no attempt at serious research onthe species in Bhutan, where much of what is known of its ecology has beenderived from incidental observations by naturalists and herdsmen livingnear tiger habitats. The present account is an attempt to collate theinformation which is available, given the need to know its currentdistribution if measures aimed at its conservation in Bhutan are to beappropriate and effective.

STUDY AREAThe Kingdom of Bhutan lies in the Eastern Himalayas between longitudes8845 ' and 92 10'E and latitudes 2640 and 28 15'N. With an area of46600km 2, Bhutan is comparable to Switzerland both in size andtopography. Given its altitudinal range of 200-7500 m from south to nor th,it has an ecological diversity paralleled in few other countries. The unusualrichness and diversity of its fauna and flora are due to the fact that Bhutanlies in an area where two of the world's major zoogeographical realms, thePalaearct ic and the Oriental, meet (Blower, 1986). Several rivers such as theTorsa, Mo (Sankosh), Pho and Manas originate in the Himalayas and flowsouthward to disgorge eventually into the Brahmaputra river in India. Theclimate is variable, being hot and humid in the south and alpine in the north.The western part of Bhutan receives the highest mean annual rainfall of3954mm (Sargent, 1985). The human population was estimated to be 1.2million in 1982, giving an average overall densi ty of 25 km-2 . However , theproductive land covers only about 19000 km 2, and the 'true' popula tiondensi ty there may be as high as 50-60kin -2 (Fisher, 1976).

METHODSMuch of the information on the numbers and distribution of the tiger wascollected in surveys carr ied out in Bhutan in 1988. Two kinds o f survey were

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c o n d u c t e d : o n a c o u n t r y - w i d e s ca le , b y v i s it in g th e a r e a s a n d i n t e r v i e w i n gl o ca l p e o p l e in J u n e 19 88, a n d m o r e i n t e n s iv e l y i n th e R o y a l M a n a s N a t i o n a lP a r k i n M a r c h 1 98 8 b y s t u d y i n g t h e s p o o r a l o n g tr a n se c ts . F o o d h a b i t s o ft h e ti g er w e r e d e t e r m i n e d f r o m t h e o b s e r v a t i o n o f c ar c as s es a n d s c at s f o u n di n t h e s tu d y a r e a ( M a n a s ) a n d f r o m i n t er v i e w i n g t ra c k e rs , g a m e g u a r d s a n dl iv e s to c k h e r d s m e n . P r e y r e m a i n s w e r e i d e n ti fi ed f r o m t h e h a i r a n d h o o ff r a g m e n t s .

R E S U L T S A N D D I S C U S S I O NDistr ibut ion

T i g e rs o c c u r f r o m a l t it u d e s o f o v e r 3 0 0 0 m in t h e n o r t h t o 2 0 0 m i n t h eH i m a l a y a n f o o t h i l ls in t h e s o u t h a l o n g t h e b o r d e r w i t h I n d i a (F ig . 1 ). O f t h e1 8 D z o n g k h a g s ( D i s t r i c t s ) i n B h u t a n , i t i s k n o w n f r o m a l l b u t t h e C h i r a n gD z o n g k h a g . A c c o r d i n g t o s o m e h e r d s m e n a n d t r ac k e rs , t h e r e a p p e a r t o b es o m e a l t i t u d i n a l d i f f e re n c e s i n t h e t i g er s : t h o s e i n t h e h i g h e r e l e v a t i o n s h a v e al a rg e r h e a d , s h o r t e r b o d y a n d l o n g e r ta il c o m p a r e d t o t h o s e t h a t o c c u r i n th el o w l an d s . T h i s m i g h t b e a n i n d i c a t io n o f l o ca l a d a p t a t i o n t o m e e t t h ed e m a n d s o f a h a r s h e r t e r r a i n i n t h e h i g h a l t i tu d e h a b i t a ts . I n t h e c e n t r a l b e l t,w h i c h r a n g e s i n a l t it u d e b e t w e e n 2 0 0 0 a n d 3 5 0 0 m , t h e t ig e r h a s b e e n r e l i ab l yr e p o r t e d i n t h e g a l l e r y f o r e s t s b o r d e r i n g t h e r i v e r s T o r s a , H a , P a r o , M o ,M a n g d e , K u r u , B u m t h a n g a n d M a n a s . I n t h e s o u t h , t h e t ig e r is r e p o r t e df r o m a n u m b e r o f p r o t e c te d a re a s s u c h a s t h e R o y a l M a n a s N a t i o n a l P a r k( w h i c h i n c l u d e s t h e N a m g y a l W a n g c h u k W i l d l i f e S a n c t u a r y ) , P h i p s o oR e s e r v e d F o r e s t a n d N e o l i W i ld l if e S a n c t u a r y ( B l ow e r , 1 98 6). It a ls o o c c u r si n th e K h a l i n g R e s e r v e d F o r e s t i n t h e e x t re m e s o u t h - e a s t o f th e c o u n t r y . I t ss t r o n g h o l d a p p e a r s t o b e th e c e n t r a l b e lt b e t w e e n M o r iv e r in t h e w e st a n dK u l o n g r iv e r in t h e e as t. G i v e n t h e c o n s i d e r a b l e e x t e n t o f a r ea s w h e r e t h et ig e r m a y i n f ac t b e p r e s e n t b u t n o t y e t s e a r c h e d f o r, it c o u l d b e m a i n t a i n e dt h a t i t s c o n s e r v a t i o n s t a t u s i s n o t t h r e a t e n e d .Hab i t a t

T h e h a b i t a t o f th e t ig e r ra n g e s f r o m t h e s u b - t ro p i c a l f o r es ts a n d s a v a n n a hg r a s s la n d s i n t h e s o u t h a l o n g t h e b o r d e r w i t h I n d i a t o t e m p e r a t e f o re s ts a t3 0 0 0 m a n d a b o v e . T h e r e is a b u n d a n t f o o d a t a l l l ev e ls a n d i n a v a r i e t y o fh a b i t a ts . T h e a n i m a l t h r i v e s i n s e v e ra l s ta g e s o f v e g e t a t i o n a l s u c c e s s io nw h i c h c an s u p p o r t a h i g h e r u n g u l a t e b i o m a s s t h a n c l im a x v e g e t a ti o n . I nE a s t e r n B h u t a n , l a r g e a r e a s o f f o r e s t h a v e b e e n d e s t r o y e d b y s la s h a n d b u r na g r i c u l t u r e ( S a r g e n t , 1 98 5), w h i c h a r e u t i l iz e d i n t h e l o n g p e r i o d s o f f a l l o w b ya v a r i e t y o f w i l d l i f e i n c l u d i n g t h e t i g e r . L o w l a n d f o r e s t s w i t h t h e i r r i c hu n g u l a t e p r e y a r e t h e o p t i m a l h a b i t a t o f th e t ig e r in B h u t a n .

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The tiger in Bhutan 315

Food habitsT i g e r s r a n g e o v e r l a rg e a r e a s a n d n e e d m e d i u m - s i z e d o r l a r g e h e r b i v o r e s a sp r e y ( A s h b y & S a n t i a p i l l a i , 1 98 7). I t i s c a t h o l i c i n i ts d i e t , f e e d i n g o n a v a r i e t yo f p r e y s p ec ie s i n B h u t a n , f o r e x a m p l e , b a r k i n g d e e r M u n t i a c u s m u n t j a k ,w i l d p i g S u s s c r o fa , h o g d e e r A x i s p o r c i n u s , s a m b a r Cervus un ico lor , a n d e v e ng a u r B o s g a u r u s a n d b u f f a lo Bubalus bubal i s a t l o w e r a l t i t u d e s . O n e c a r c a s so f a n a d u l t b u l l g a u r k il le d b y a ti g e r w a s f o u n d i n M a n a s n e a r a d r y r i v e rb e d . T h e t ig e r f e d o n t h e h i n d q u a r t e r s a n d l ef t t h e r e s t o f t h e c a r c a s s i n ta c t .

A t h i g h e r a l t i t u d e s , t h e t i g e r p r e y s o n a n i m a l s s u c h a s t h e s e r o wCapr icorn i s sumatraens i s , t h e t a k i n B u d o r c a s t a x i c o lo r a n d o t h e r m o u n t a i nu n g u l a t e s . T a k i n u s u a l l y i n h a b i t t h e d e n s e b a m b o o f o r e st s a t a lt i tu d e s o f1 5 0 0 t o 3 0 0 0 m ( P r a t e r , 1 96 5) a n d a s c e n d t o e v e n h i g h e r a l t i t u d e s i n s u m m e r .T h e t ig e r is k n o w n t o f o l lo w t h e m o v e m e n t o f t h e t a k i n h e r ds . A t h i g h e ra l ti t u d es , i t a l so s o m e t i m e s p r e y s o n t h e d o m e s t i c y a k , w h i c h i s o n e o f t h ec a u s e s o f c o n f l ic t w i t h t h e h e r d s m e n .Tiger numbers and densityE s t i m a t i n g t i g e r n u m b e r s i n t h e d e n s e a n d t a n g l e d v e g e t a t i o n o f t h e t r o p i c a lr a i n f o r e s t h a b i t a t s is d if fi cu l t. T h e r e s u l t s o f th e i n t e r v i e w s i n d i c a t e t h ep r e s e n c e o f a b o u t 1 50 ti g e r s i n B h u t a n ( T a b l e 1 ). H o w e v e r , g i v e n t h e f a c t t h a tt h e t i g e r c a n i n h a b i t a n a r e a w i t h o u t b e t r a y i n g i t s p r e s e n c e , t h i s i s am i n i m u m e s t im a t e . In N e p a l , w h i c h is v e r y m u c h s i m i l ar to B h u t a n i nt o p o g r a p h y , c l i m a t e a n d v e g e t a t i o n , a d u l t t ig e r d e n s it ie s c a n r e a c h 6 7i n d i v i d u a l s 1 00 k m - 2 i n p r i m e h a b i t a t s , a n d 1 - 2 10 0 k i n - 2 i n le ss o p t i m u ma r e a s ( M c D o u g a l , 1 97 7; S m i t h , 1 9 78 ; S u n q u i s t , 1 98 1). I n t h e 6 5 8 k m 2 R o y a lM a n a s N a t i o n a l P a r k in B h u t a n , a b o u t 3 0 t ig e rs a r e k n o w n t o b e p r e s e n t( T o b g e y D u k p a , p e rs . c o m m . ) , g i v in g a d e n s i t y o f 4-5 1 00 k m - 2 . T h i s v a l u ec o m p a r e s f a v o u r a b l y w i t h th e e s t im a t e d t ig e r d e n s i t y o f 4"3 1 0 0 k i n 2( K a r a n t h , 19 87 ) i n t h e a d j o i n i n g M a n a s T i g e r R e s e r v e i n I n d i a. H o w e v e r ,n o t a l l t h e a r e a s i n B h u t a n a r e p r i m e t i g e r h a b i t a t - - t h e t o t a l a r e a g iv e nc o n s e r v a t i o n s t a t u s is 9 50 5 k m 2 ( B l o w e r , 1 9 86 ). T h e e x i s t in g c o n s e r v a t i o na r e a s in B h u t a n , a s s u m i n g t h e i r st a b il i ty r e m a i n s a s s u r e d , w o u l d p r o t e c t n om o r e t h a n a b o u t 2 50 t ig e rs a t a n a v e r a g e d e n s i t y o f o n e t ig e r 4 0 k m - 2

C O N S E R V A T I O N A I M S A N D P R O S P E C T SG i v e n t h e n e e d f o r l a r g e h o m e r a n g e a n d a b u n d a n t p r e y s p e c i e s , t h e t i g e rc a n n o t b e m a i n t a i n e d a t h ig h d e n s i t y e v e n i n t h e m o s t f a v o u r a b l e h a b i t a t( S a n ti a p il la i & W i d o d o , 1 98 5). M a n y o f t h e p r o t e c t e d a r e a s i n B h u t a n c a no n l y s u p p o r t a b o u t 2 0 - 3 0 t ig e rs . S u c h s m a ll p o p u l a t i o n s a r e m o r e

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316 Dasho Paljor Dorji, Charles Santiapillai

T A B L E 1N u m b e r o f T i ge r s Panthera tigris i n B h u t a n ( b a s e d o n t h e t i g e r

c o u n t c a r r i e d o u t i n 1 9 88 b y D a s h o P a l j o r D o r j i )Locality District (Dzongkhag) M inim um number

S h e lg a n a P u n a k h a 2T s a c h i p h u g 3 "S u i L a 1G a s a 3R i d a h W a n g d i p h o d r a n g 2R u k u b j i 1B a l i n g / D a n g l i n g T o n g s a 3N g a L a B u m t h a n g 2D u b i T s a c h u 3D o c h u L a T h i m p h u 1D r u g y a l D z o n g P a r o 1J a b a n a 2C h a p c h a C h h u k h a 1C h u k h a 1C h i m a k o t h i 1C h a s i l a k h a 1G e d u 1D a m t h a n g H a 2 aS i b s u S a m c h i 5bP h i p s o o G e y l e g p h u g 7M a n a s 3 0 bK a l i k h o l a 2K a n a M a k r a 4G a y l e g p h u g 4 6D a g a D a g a 6P e m a g a t s e l P e m a g a t s e l 2 0B a r s h a m T a s h i ga n g 3T a s h i Y a n g t s i 5 bS h e m g a n g S h e m g a n g 3K h e n 4A r j a n a M o n g a r 5K u r t e y L h u n t s h i 2M i n g j i 3D a i f a m / B a n g t a r S a m d r u p J o n g k h a r l 0 bN g a l h a m 5D e c h h e l i n g 2T o t a l ( m i n i m u m ) 151

a P r e s e n c e o f a n a d u l t f e m a l e a n d c u b .b T h e s e a n i m a l s a r e s h a r e d w i t h I n d i a .

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The t iger in Bhutan 317

v u l n e r a b l e th a n l a r g er o n e s to r a n d o m c h a n g e s (s u c h a s m a r k e d f l u c t u a t i o n si n se x ra t io ) , i n b r e e d i n g d e p r e s s i o n a n d l o c a l c a t a s t r o p h i c e v e n t s ( B e r t r a m ,1 98 6). T h e s e p r o b l e m s c a n t o s o m e e x t e n t b e m i n i m i s e d i f t h e p r o t e c t e d a r e a sc a n b e l in k e d t o o n e a n o t h e r t h r o u g h t h e e s t a b l is h m e n t o f f o r es t c o r r i d o r s i no r d e r t o f a c il i ta t e g e n e t ic e x c h a n g e . I f t h e o b j e c t i v e is to m a i n t a i n a s m a n yt ig e r s a s p o s s i b l e i n a s w i d e a n a r e a a s is f ea s ib l e, p r o p e r m e a s u r e s m u s t b et a k e n n o w t o m i n i m i s e ti g er d e p r e d a t i o n s o n d o m e s t i c y a k a n d c a tt le . S o m el o s s o f l i v e s to c k i s i n e v i t a b l e i f t h is s t o c k g r a z e s i n s i d e a b i g p r e d a t o r ' s r a n g e ,a n d f a r m e r s a l m o s t w i t h o u t e x c e p t i o n g r o s sl y e x a g g e r a te t h e e c o n o m i ce ff ec ts th a t a n y s u c h l o s s m a y c a u s e .

T i g e r a t t a c k s o n d o m e s t i c y a k h a v e b r o u g h t t h e a n i m a l i n d ir e c t c o n f li ctw i t h t h e h e r d s m e n , w h o a r e s til l k n o w n t o p o i s o n m a r a u d i n g a n i m a l s d e s p it ea d e c r e e b y t h e R o y a l G o v e r n m e n t . T h i s h a s b a n n e d v i l l a g e r s f r o m u s i n gp o i s o n t o k i ll ti g e r s i n B h u t a n f o r t h e l a s t 1 3 y e a r s . T h i s is d u e l a r g e l y to t h ef ac t th a t h e r d s m e n h a v e n o t b e e n c o m p e n s a t e d f o r t h ei r lo s s e s - - t h e m a i nc a u s e o f re s e n t m e n t . I n a d d i t io n , a n y p l a n f o r c o n s e r v a t i o n m u s t t a k e i n t oa c c o u n t t h e h i g h v u l n e r ab i l it y o f a l ar g e p r e d a t o r t o p o i s o n i n g a n d d i r ec th u n t i n g . H o o g e r w e r f (1 97 0) a t t r ib u t e d t h e e x t i n c ti o n o f t h e J a v a n t ig e rP a n t h e r a t i g r i s s o n d a i c a f o l lo w i n g t h e f r a g m e n t a t i o n o f it s h a b i t a t p r i m a r i l yt o t h e u s e o f p o is o n .

T r a d e i n t i g e r s k i n s i s h i g h l y i p r o f i t a b l e b u t p o a c h i n g i s n o t y e t a m a j o rc o n s e r v a t i o n p r o b l e m i n B h u t a n . T h e B u d d h i s t a v e r s i o n t o k i l l i n g h a sm e a n t t h a t t h e r e i s l i t t l e d i r e c t p r e s s u r e o n w i l d a n i m a l s ( S a r g e n t , 1 9 8 5 ) .

T h e c o n s e r v a t i o n p o l i c y m u s t m a i n t a i n f o r e s t c o v e r o v e r l a r g e a r e a su n i n t e r r u p t e d b y h u m a n s e t tl e m e n t s a n d r o a d s , w h e r e r e m o t e n e s s , d i ff ic u lt yo f te r r a in a n d d e n s i ty o f c o v e r p r o v i d e n a t u r a l p r o t e c t io n . E v e n q u i t e s m a l ll o c a l p o p u l a t i o n s a r e v a l u a b l e a n d t h e s e s h o u l d b e p r o t e c t e d w h e r e v e rp r a c t i c a b l e a n d n o t a b a n d o n e d s i m p l y f o r g e n e t i c r e a s o n s ( S c h a l l e r , 1 9 8 6 ;A s h b y & S a n t i a p i l l a i , 1 9 8 7 ) . I t w o u l d b e a m i s t a k e t o r e l y o n t h e s p e c i e sb e i n g s a fe in p e r p e t u i t y i n o n e o r t w o l a rg e r e s er v e s o n l y - - s m a l l e r r e se r v e sa l so have a v i t a l ro l e .T h e r e i s a n u r g e n t n e e d f o r s u r v e y s t o b e d o n e t o d e t e r m i n e t h ew h e r e a b o u t s o f v ia b le p o p u l a t i o n s o f ti ge r, p a r ti c u l ar l y i n t h e r e m a i n i n gu n f r a g m e n t e d f o r e s t s i n t h e B l a c k m o u n t a i n s . A s B l o w e r ( 1 9 8 6 ) r e c o m -m e n d s , t h e t i g e r s h o u l d b e l i s t e d a m o n g t h e s p e c i e s t o b e d e s i g n a t e d a sR o y a l G a m e 'w h i c h c o u ld o n l y b e h u n t e d o r c a p t u r e d w i t h th e p e r m i s s i o n o ft h e K i n g a n d i ll eg a l k il li n g o f w h i c h w o u l d i n c u r e s p e c ia l ly s e v e re p e n a l t ie s '.

T h e m a i n t h r e a t s t o t h e t i g er in t h e f u t u r e w o u l d a ri se n o t m e r e l y f r o m t h ed e m a n d s o f th e g r o w i n g h u m a n p o p u l a t i o n f o r t h e r e so u r ce s o f t h e l a n d b u ta ls o t h r o u g h t h e l e g it im a t e a s p i r a ti o n s o f t h e B h u t a n e s e f o r a n i m p r o v e ds t a n d a r d o f li vi n g. A s M y e r s ( 19 75 ) p o i n t s o u t , 'a s p i r a t i o n s t o t h e e n h a n c e ds t a n d a r d s o f li vi n g o f t e n c a u se g r e a t e r p r e s su r e o n u n d i s t u r b e d n a t u r a l

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318 Da sho Paljor D orji, Charles Santiapillaienvironments than sheer growth of human numbers'. Already in thehighlands large tracts of forest have been burnt and cleared to provideadditional grazing for the yak herds (Sargent, 1985).

All the current efforts to conserve the tiger, whether in the wild or incaptivity, are unlikely to succeed if they do not have the support of the localpeople. The herdsmen must believe that conservation measures are beingtaken in their long-term interests. Protection of conservat ion areas should beaccompanied by measures that would improve the living standards of thepeople without further disrupting the environment (Schaller et al. , 1987).Land-use pa tterns in areas adjacent to the wildlife reserves inhabited by tigermust be designed in such a way as to make them compatible with tigerconservation and thus minimise the human-wildlife conflicts as much aspossible.

ACKNOW LEDGEM ENTSThis work is the result of a collaborative research programme betweenthe Department of Forestry in Bhutan and the World Wildlife Fund. Wewish to express our thanks to the Director General of the Department ofForestry, Dasho Dorji Tenzing and to the Director, Asia Programme of theWorld Wildlife Fund, Dr Bruce W. Bunting.

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The tiger in Bhutan 319M ye r s , N . ( 1975). The C he e ta h A c i n o n y x j u b a t u s in Afr ica , I U C N M o n o g r . , No. 4 .Prater, S. H. (1965). The Bo ok o f Ind ian An imals . B o m b a y N a t u r a l H i s t o r y S o c ie ty ,B o m b a y .Sant iapi l la i , C . & Widodo, S . R . (1985) . Tiger numbers and habi ta t eva lua t ion in

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