The Feeding Biology Relation to F©od Soyircss of Mect'afiv/©iToy£ flats

ti i t o p t © if o: M s c f © 'H I© ss y ft s !«? i'i/! Msifaysia

A R f'O M f M. 'START

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1 d e c l a r e t h a t t h i s t h e s i s s com posed b y m y s e l f an d em bodying

work done by m y s e l f h a s n o t b e e n a c c e p t e d i n a n y p r e v i o u s

a p p l i c a t i o n f o r a h i g h e r d e g re e , . A l l s o u r c e s o f r e f e r e n c e

and a l l q u o t a t i o n s h a v e b e e n d u e l y a c k n o w le d g e d .

M f t p ,

A n to n y . N . S t a r t .

/Ilo

CONTENTS.

I n t r o d u c t i o n

S tu d y a r e a s

P e n i n s u l a r v M a l a y s i a

E a s t M a l a y s i a

M ethods

C h i r o p t e r o p h i l o u s p l a n t s

M a c r o g l o s s i n e h a t s

O c c u r r e n c e o f p o l l e n

R e s u l t s

C h i r o p t e r o p h i l o u s p l a n t s

M o n o c o ty le d o n s

1 . M u saceae

2 . Fa lm ae

3 ) i c o ty l e d o n s

1 . A n a c a r d i a c e a e

2 . B i g n o n i a c e a e

3® B om b acaceae

if. L e c y t h i d a c e a e

3® L eg u m in o sae

6« M y r ta c e a e

7® R h i s o p h o r a c e a e

8 . S o r m e r a t i a c e a e

9® M o rac eae

M a c r o g lo s s u s m in im us

1 ® E c o l o g i c a l d i s t r i b u t i o n

p a g e

3® Food s o u r c e s 92

4* A n a l y s i s o f p o l l e n s a m p le s 100

5o I t e m s b e s i d e s p o l l e n i n f a e c e s 1 04

6 „ F l i g h t 107

7o A bundance a t Bako N a t i o n a l P a r k

8 . R i n g i n g r e s u l t s 110

9 . Movements 111

1 0 . R e p ro d u c t i o n 11 4

M a c r o g l o s s u s s o b r i n u s

1 . E c o l o g i c a l d i s t r i b u t i o n 123

2 . R o o s t i n g 123

3* Food s o u r c e s 1 24

4 . A n a l y s i s o f p o l l e n s a m p le s 129-

5 . I t e m s b e s i d e s p o l l e n i n f a e c e s 1 3 2

6 . F l i g h t 133

7® A c t i v i t y p a t t e r n 134

8 . R i n g i n g r e s u l t s 1 3 5

9* A bundance a t K a ja n g i n r e l a t i o n t o

a b u n d a n c e o f f l o w e r i n g b a n a n a s 137

1 0 . R e p r o d u c t i o n 139

E o n y c t e r l s s p e l a e a

1 . E c o l o g i c a l d i s t r i b u t i o n 140

2 . R o o s t 140

3 ® Guanb c o l l e c t e d from B a tu C av es 1 4 6

4 . Food s o u r c e s 149

5 . P o l l e n t y p e s i n guano from B a tu C aves 150

6 . A n a l y s i s o f p o l l e n i n guano from B a tu

c a v e s

p ag e

7 . P r e s e n t a t i o n and l i m i t a t i o n o f d a t a 1 65

8o S u b f o s s i l d e p o s i t s 1 69

9 . P o l l e n s a m p le s from t h e b a t s '170

1 0 . F l i g h t 172

1 1 . D i s t a n c e t r a v e l l e d f o r fo o d 174

12* A ctivity tiinGS &X iveui CGU jrcuij3.ng

an d e l s e w h e r e 1 76

13° R i n g i n g r e s u l t s 178

1 4 . F o r a g i n g 1 79

1 5 . R e p r o d u c t i o n 1 85

D i s c u s s i o n

1 . E c o l o g i c a l d i s t r i b u t i o n 191

2 . Food s o u r c e s 195

3 . R o o s t i n g b e h a v i o u r i n r e l a t i o n to

fo o d s o u r c e s 210

4 . F o r a g i n g m e th o d s - 219

5 . F l i g h t 2.23

6 . R e p r o d u c t i o n 226

7 . B a t f l o w e r s and f l o w e r b a t s 230

Summary 245

A ck no w led g e m en ts 246

B i b l i o g r a p h y ' . 248

A p p en d ix 1 « Taxonomic s t a t u s o f M a c r o g lo s s u s s p e c i e s

i n M a l a y s i a .

2 , D a ta t e s t i n g a d e q u a c y o f p o l l e n a n a l y s i s

m e t h o d s .

3 . D a ta r e l a t e d t o g uano c o l l e c t i o n s from B a tu

Caves,

ifo D e t a i l s o f p o l l e n c o u n t s from guano c o l l e c t e d

i n B a tu C a v e s ,

INTRODUCTION.

The o r d e r C h i r o p t e r a ( b a t s ) i s r e a d i l y d i v i s i b l e i n t o two s u b ­

o r d e r s , t h e MEGACHIROPTERA and t h e MICROCHIROPTERA. The Mega-

c h i r o p t e r a o n l y o c c u r i n t h e O ld W o r ld ; t h e y a r e p r i n c i p a l l y

f r u g i v o r o u s . The M i c r o c h i r o p t e r a a r e fo u nd b o t h i n t h e O ld and

t h e New W o r ld ; t h e y a r e p r i n c i p a l l y i n a e c l i v o r - o u s b u t some

New W orld s p e c i e s a r e p a r t l y o r c o m p l e t e l y f r u g i v o r o u s . A few

members o f b o t h s u b o r d e r s h a v e a d o p t e d a d i e t o f n e c t a r and

p o l l e n an d p o l l i n a t e t h e f l o w e r s t h e y f e e d f r o m .

D u r in g t h e l a s t two d e c a d e s t h e r e h a s b e e n an

i n c r e a s i n g i n t e r e s t i n c h i r o p t e r o p h i l y ( t h e p o l l i n a t i o n o f

f l o w e r s b y b a t s ) e s p e c i a l l y i n t h e New W orld ( e . g . A l v a r e z

an d Q u i n t e r o , 1 9 70 ; B a k e r , 1 9 73 ; H e i t h a u s , O p l e r an d B a k e r ,

1 97 4 ; H o w e l l , 1 9 74 ; V o g e l , 1 9 6 8 -6 9 ) and t o a l e s s e r e x t e n t i n

A f r i c a ( e . g . B a l te r an d H a r r i s , 1957> 1 9 3 9 ; H a r r i s and B a k e r ,

1 9 5 8 , 19 59 ; J a e g e r , 1 9 5 4 ) .

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

S o u th E a s t A s i a w as t h a t o f v a n d e r P i j l (1 9 3 6 ) i n w h ic h t h e

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

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

o r s u s p e c t e d o f b e i n g c h i r o p t e r o p h i l o u s . H i s o b s e r v a t i o n s w e re

made i n J a v a b u t h e w as a b l e t o i n c l u d e some e x o t i c s p e c i e s

p l a n t e d i n t h e B o t a n i c G ard en i n B o g o r . He r e c o r d e d w h ich

s p e c i e s o f b a t s v i s i t e d some o f t h e p l a n t s an d i n c l u d e d a

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

t h e i r d i e t . He l a t e r a d d e d t o and r e v i s e d t h i s w ork ( v a n d e r

P i j l , 1 9 4 1 5 1956) b u t o t h e r w i s e v e r y few o b s e r v a t i o n s r e l a t i n g

t o S o u th E a s t A s i a h a v e b e e n p u b l i s h e d s i n c e t h e n .

B a k e r and H a r r i s (1 9 5 7 ) su m m a r i s e d t h e e a r l y

J a v a n e s e r e c o r d s q u o te d by van d e r P i j l ( 1 9 3 6 ) ; N'ur ( 1 9 5 8 ,

q u o te d f rom P u r s g l o v e , 1972) c o n f i r m e d t h a t hemp, Musa t e x t i l i s

( M u s a c e a e ) , was p o l l i n a t e d by b a t s i n J a v a ; B e rw ic k (1 9 5 0 )

p u b l i s h e d a p h o t o g r a p h o f E o n y c t e r l s s p e l a e a D obson (M a c ro -

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

M u l l . ( P t e r o p o d i n a e , P t e r o p o d i d a e ) v i s i t i n g t h e f l o w e r s o f

C e ib a p e n t a n d r a ( L . ) G a e r t n . (B o m baca ceae ) b u t f a i l e d t o

r e a l i s e i t s s i g n i f i c a n c e ; Lira (1 9 7 3 ) and Tan (1 9 6 5 ) b o t h

r e p o r t e d p o l l e n i n t h e s to m a c h s o f M a c r o g lo s s u s s p p . B eck and

Lira (1 9 7 3 ) s t u d i e d t h e r e p r o d u c t i o n o f E o n y c t e r i s s p e l a e a .

The o b j e c t o f t h e p r e s e n t s t u d y w as t o exam ine

t h e d i e t s a n d some r e l a t e d a s p e c t s o f t h e b i o l o g y o f t h r e e

s p e c i e s o f n e c t a r i v o r o u s b a t s i n P e n i n s u l a r M a l a y s i a : i . e .

M a c r o g l o s s u s m in im us ( G e o f f r o y ) , M a c r o g lo s s u s s o b r l n u s

( A n d e r s e n ) an d E o n y c t e r i s s p e l a e a D o bson . A l l t h r e e b e l o n g t o

t h e s u b f a m i l y M a c r o g l o s s i n a e w h ic h i s d i s t i n g u i s h e d from o t h e r

P t e r o p o d i ^ d a e b y a d a p t a t i o n s t o a d i e t o f p o l l e n and n e c t a r ,

e s p e c i a l l y by t h e p r e s e n c e o f l o n g , f i l i f o r m p a p i l l a e t h a t

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

The taxonom y o f M a c r o g lo s s u s i s c o n f u s e d and

i s c u r r e n t l y b e i n g r e v i s e d ( S t a r t , ’ i n p r e p . ) . The names u se d

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

M a l a y s i a n t a x a ( e . g . b y A n d e r s e n , 1 91 2 ; C h a s e n , 1 9 4 0 ; Medway,

1 9 6 9 ) . M a c r o g l o s s u s m in im u s ( G e o f f r o y ) was f o r m e r l y M .l a g o c h l l u s

M a t s c h i e . M .s o b r i n u s ( A n d e r s e n ) was f o r m e r l y M.m in im us s o b r i n u s

A n d e r s e n . D e t a i l s o f t h e ta x o n o m ic p o s i t i o n o f t h e M a l a y s i a n

t a x a a r e g i v e n i n A p p en d ix 1 .

3

M a c r o g lo s s u s m in im us i s v e r y w i d e l y d i s t r i b u t e d ,

f rom S u m a t r a t h r o u g h o u t M a l a y s i a , I n d o n e s i a , P h i l i p p i n e s and

New G u in e a t o t h e Solom on I s l a n d s . I t a l s o e x t e n d s i n t o S o u th

V ie tn a m , T h a i l a n d and n o r t h e r n A u s t r a l i a ( S t a r t , i n p r e p . ) b u t

M a c r o g lo s s u s s o b r i n u s an d E o n y c t e r i s s p e l a e a a r e c o n f i n e d t o

m a i n l a n d S o u th E a s t A s i a ( i n c l u d i n g Burma, T h a i l a n d , t h e Khmer

R e p u b l i c and S o u th V ie tn am ) t h r o u g h t h e M alay P e n i n s u l a r and

S u m a t r a t o J a v a an d B a l i . E . s p e l a e a a l s o o c c u r s i n t h e P h i l i p ­

p i n e s and B o rn e o ( W a l k e r , 1 96 4 ; S t a r t , i n p r e p . ) .

The two s p e c i e s o f M a c r o g lo s s u s a r e v e r y s m a l l ;

M* m in im us w e ig h s 1 6 t o 24 g , M. s o b r i n u s 1 8 t o 26 g b u t

£ • s p e l a e a i s a medium s i z e d b a t w e ig h in g 40 t o 70 g . The

m a le s o f a l l t h r e e s p e c i e s a v e r a g e h e a v i e r t h a n f e m a l e s , e x c e p t

p r e g n a n t f e m a l e s a p p r o a c h i n g p a r t u r i t i o n . A d u l t m a le E . s p e l a e a

may h a v e a s m a l l r u f f on t h e t h r o a t and a d u l t m a le M. m in im us

h a v e a d e r m a l g l a n d u l a r s t r u c t u r e on t h e lo w e r t h r o a t . T h i s i s

n o r m a l l y a b s e n t from M. s o b r i n u s i n P e n i n s u l a r M a l a y s i a . T h e re

i s no o t h e r m ark ed s e x u a l d im o r p h i s m . A l l t h r e e s p e c i e s h a v e

b e e n a d e q u a t e l y d e s c r i b e d by v a r i o u s a u t h o r s , e . g . A n d e rse n

(1 9 1 2 ) and Medway ( 1 9 6 9 ) .

The s t u d y w as u n d e r t a k e n b e tw e e n m i d - J a n u a r y 1972

a n d e a r l y J a n u a r y 1974 i n t h e s t a t e s o f S e l a n g o r an d P e r a k ,

P e n i n s u l a r M a l a y s i a b u t v i s i t s w e re made t o m o s t M a l a y s i a n

s t a t e s . , n o t a b l y t o S a b a h a a d S a ra w a k i n J a n u a r y an d May 1973

( F i g s . 1 an d 2 ) .

I n i t i a l l y , t h e s t u d y was r e s t r i c t e d t o two

s p e c i e s , M a c r o g l o s s u s m in im us and E o n y c t e r i s s p e l a e a . E f f o r t s

Fig.1 P a r t of S o u th East Asia s h o w i n g loca l i t i e s m e n t i o n e d in t h e t e x t .

Mt . K i n a b a l u a

S A B A HBRUN

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F i g . 2 . W e s t e r n P e n i n s u l a r M a l a y s i a s h o w i n g l o c a l i t i e s m e n t i o n e d in t h e t e x t .

F .R . = F o r e s t R e s e r v e G. = G u n o n g ( M o u n t a i n )

F . S . C . = F ie ld S t u d i e s C e n t r e

K u a l a G u l aP E R A K /

A M a x w e l l Hill

• G u a P o n d o k

\ k e l a n t a n

P e r a kR i v e r

d p o h ( N ,^-^G. B r i n c f i a n g 1 C a m e r o n

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K u a l a S e l a n g o r S P Ulu G o m b a k / m ' ^e\ _ r » 5 * v » -

\ B a tu C a v e s *

\ 1 X R a n t a u P a n j a n g I _ _ ® K u a l a L u m p u rv n i i W © M e r u F .R .K a m p o n q D e l e k O ^ \

< 2 / k ? * K l a n g i© K a j a n g g®,

Te lok F.R.

101° 3 0 'E

w e re d i r e c t e d t o w a r d s e s t a b l i s h i n g t h e i r d i e t s t h r o u g h o u t t h e

y e a r b e c a u s e many o f t h e p l a n t s from w h ich t h e y w e re t h o u g h t

t o f e e d (v a n d e r F i j i , 1 9 3 6 , 1 9 4 1 , 1956) w e re known t o f l o w e r

s e a s o n a l l y . The f o r a g i n g m e th o d s and r o o s t i n g h a b i t s o f b o t h

s p e c i e s w e re a l s o s t u d i e d i n r e l a t i o n t o t h e i r fo o d s o u r c e s

t o d i s c o v e r how two s p e c i e s o f b a t s t h a t w e re t h o u g h t t o u s e

t h e same fo o d s o u r c e s c o u l d c o e x i s t . R e p r o d u c t i v e d a t a was

g a t h e r e d i n c i d e n t a l l y so t h a t c a u s e s o f an y s e a s o n a l v a r i a t i o n

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

t h e p o s s i b i l i t y t h a t some d i e t a r y f a c t o r m ig h t a f f e c t s e a s o n ­

a l i t y o f b r e e d i n g . The s t u d y was n o t c o n f i n e d t o t h e b a t s b u t

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

w h ic h t h e y f e d , e . g . t h e i r d i s t r i b u t i o n , m o r p h o lo g y , f l o w e r i n g

b i o l o g y and p h e n o l o g y .

D u r in g t h e s e c o n d y e a r , l e s s a t t e n t i o n was p a i d

t o M a c r o g l o s s u s m in im us t o p e r m i t more t im e f o r t h e s t u d y o f

E o n y c t e r j s s p e l a e a , e s p e c i a l l y t h e a n a l y s i s o f p o l l e n e x t r a c t e d

f ro m guano o f t h i s s p e c i e s . A p r e l i m i n a r y s t u d y o f t h e d i e t

an d o t h e r a s p e c t s o f t h e b i o l o g y o f M. s o b r i n u s w e re a l s o

u n d e r t a k e n d u r i n g t h e ' s e c o n d y e a r . The p r i m a r y a im o f t h i s

l a s t s t u d y was t o i d e n t i f y t h e fo o d s o u r c e s o f M. s o b r i n u s

and t o d i s c o v e r hov; t h a t s p e c i e s w a s ' a b l e t o c o e x i s t w i t h

M. m in im u s an d E . s p e l a e a .

5

STUDY AREAS.

PENINSULAR MALAYSIA.

P h y s i c a l f e a t u r e s .

The l a n d i s m o s t l y b e lo w 1000 m a . s . l , b u t a

r a n g e o f m o u n t a in s r i s i n g t o 2183 m a . s . l . r u n s s o u t h from

t h e T h a i l a n d b o r d e r a l o n g two t h i r d s o f t h e p e n i n s u l a r n e a r

i t s w e s t c o a s t . A s m a l l e r r a n g e r u n s n o r t h - s o u t h f o r 2/+0 km

a l o n g t h e n o r t h - e a s t c o a s t . T h e re a r e s e v e r a l enorm ous

i n s e l b e r g s r i s i n g a b o v e 1000 m a . s . l . , t h e m o s t n o t a b l e

b e i n g Gunong Tahan ( 2 1 8 6 m) an d Gunong Benom (2 1 0 7 m ) . I n

t h e n o r t h e r n p a r t o f t h e c o u n t r y a r e n um erous l i m e s t o n e

o u t c r o p s , some o f w h ic h r i s e more t h a n 500 m a b o v e t h e

s u r r o u n d i n g c o u n t r y : Many c o n t a i n hu ge c a v e s . T h e i r d i s t r i b u t i o n

was mapped b y G o b b e t t (1 9 6 5 )* They a r e m os t n u m ero u s on t h e

w e s t e r n c o a s t a l s t r i p from P e r l i s , w h ere t h e y p r o d u c e a

t y p i c a l k a r s t l a n d s c a p e , ’ t o P D rak ; I s o l a t e d o u t c r o p s a l s o

o c c u r i n n o r t h e r n a r e a s to t h e e a s t o f t h e m ain r a n g e b u t

t h e r e a r e n o n e s o u t h o f B a tu C aves i n S e l a n g o r ( F i g . 2 ) .

C l i m a t e .

The c l i m a t e o f P e n i n s u l a r M a l a y s i a , w h ic h i s

h o t and w e t , i s d e t e r m i n e d b y i t s e q u a t o r i a l m a r i t i m e

p o s i t i o n . The a n n u a l t e m p e r a t u r e v a r i a t i o n o f 1 0 - 3°C i s

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

(L ockw ood, 1 9 7 1 ) . Much o f t h e r a i n f a l l i s o f o r o g r a p h i c

o r i g i n and v e r y h e a v y l o c a l s t o r m s a r e c h a r a c t e r i s t i c b u t

t h e o n s e t o f t h e n o r t h e a s t monsoon i n O c to b e r o r November

ca n a l s o b r i n g w id e s p r e a d h e a v y r a i n f a l l t o t h e e a s t c o a s t .

6

The m o s t i m p o r t a n t s e a s o n a l f e a t u r e i n t h i s n o r m a l l y w e t

e c l i m a t e i s t h e o c c u r r e n c e o f d r y s e a s o n s . D a le (1 9 6 0 ) h a s

r e c o r d e d t h e number o f c o n s e c u t i v e d r y d a y s i n e a c h c a l e n d e r

month a t v a r i o u s l o c a l i t i e s . T h i s d a t a s u g g e s t s two d r i e r

s e a s o n s e a c h y e a r i n m o s t l o c a l i t i e s . They a r e v e r y i n d i s t i n c t

i n t h e s o u t h b u t t e n d t o become more p ro n o u n c e d i n t h e n o r t h .

H o w ever , t h e o n l y a r e a w i t h a d e p e n d a b l e a n n u a l d r y s e a s o n

i s P e r l i s , i n t h e n o r t h w e s t o f t h e c o u n t r y .

V e g e t a t i o n .

The m a j o r v e g e t a t i o n t y p e s i n P e n i n s u l a r M a l a y s i a

a r e d e s c r i b e d b y W y a t t - S m i t h ( 1 9 5 2 ) . The c l i m a x v e g e t a t i o n

o v e r m o s t o f t h e c o u n t r y i s t r o p i c a l lo w la n d r a i n f o r e s t .

This gives way to lower montane rainforest at around 1000 m a.s.l. and upper montane rainforest at around 2000 m a.s.l. (e.g. Burgess, 1969; Whitmore and Burnham, 1969) and to mangrove forest along most of the west coast and along the tidal reaches of all rivers.

Much o f t h e l o w la n d f o r e s t h a s b e e n lo g g e d and

a l l o w e d t o r e g e n e r a t e o r c l e a r e d and r e p l a c e d e i t h e r b y mono­

c u l t u r e ( p r i n c i p a l l y r i c e . O ry za s a t i v a L . G ra m in a e ; r u b b e r ,

H evea b r a s i l i e n s i s M u e l l e r E u p h o r b i a c e a e o r o i l p a lm , ' 'E l a e i s

g u i n e e n s i s J a c q . P a lm a e ) o r by m ixed c u l t i v a t i o n ( i n w h ich

v a r i o u s f r u i t t r e e s , c o c o n u t p a lm s and b a n a n a s a r e among

t h e m o s t i m p o r t a n t p l a n t s ) . L a rg e a r e a s h a v e b e e n a l i e n a t e d

f o r m in in g o f a l l u v i a l t i n d e p o s i t s o r f o r i n d u s t r i a l

d e v e lo p m e n t .

The i n r o a d s i n t o m o n tane f o r e s t a r e l e s s s e v e r e

7

e x c e p t n e a r t h e h i l l s t a t i o n s , p a r t i c u l a r l y t h e Cameron

H ig h l a n d s i n P e r a k and P a h an g ( F i g . 2 ) . Much o f t h e m angrove

f o r e s t h a s t e e n r e c l a i m e d and a l m o s t a l l t h e r e m a i n d e r h a s

b e e n lo g g e d a nd a l l o w e d t o r e g e n e r a t e .

S tu d y a r e a s .

F i g . 2 shows a l l l o c a t i o n s i n P e n i n s u l a r M a l a y s i a

a t w h ich o b s e r v a t i o n s w ere m ade . A t many l o c a l i t i e s t h e

o b s e r v a t i o n s w ere b r i e f o r i r r e g u l a r b u t t h e y w ere more

d e t a i l e d i n t h e f o l l o w i n g s t u d y a r e a s ,

1 , R a n ta u P a n j a n g , S e l a n g o r .

( 3 ° 0 2 «N‘f' 1 0 1 ° 2 5 * E; a l t . s e a l e v e l ; F i g . 3 . )

T h i s s t u d y a r e a w h ic h c o v e r s 2 . 6 s q . km o f f l a t

l a n d l i e s on t h e n o r t h b a n k o f t h e r i v e r K l a n g . I t c a n b e

d i v i d e d i n t o two p a r t s m angrove f o r e s t an d m ixed c u l t i v a t i o n .

The m angrove f o r e s t w h ic h was f o r m e r l y v e r y e x t e n s i v e h a s

b e e n r e d u c e d by l a n d r e c l a m a t i o n t o a f r i n g e v a r y i n g from

100 m t o 200 m w id e a l o n g e a c h b a n k o f t h e r i v e r . I t c o n t a i n s

an e x c e p t i o n a l l y w id e d i v e r s i t y o f m angrove s p e c i e s i n c l u d i n g

t h r e e s p e c i e s o f S o n n e r a t i a L . ( S o n n e r a t i a c e a e ) ( s e e p . ~jS

f o r d e t a i l s o f S o n n e r a t i a d i s t r i b u t i o n w i t h i n t h e s t u d y a r e a ) ,

two s p e c i e s o f A v i c e n n i a L . ( V e r b e n a c e a e ) , a t l e a s t two s p e c i e s

R h iz o p h o r a L . and o f E r u g u i e r a Lam. ( R h i z o p h o r a c e a e ) , one

s p e c i e s e a c h o f L u m n i t z e r a W i l l d . ( C o m b r e t a c e a e ) , X y lo c a r p u s

K o e n ig ( M e l i a c e a e ) and D o l i c h a n d r o n e F e n z l . , ( B i g n o n i a c e a e )

M angrove f e r n s , A c r o s t i c h u m s p p . L . a r e a l s o v e r y common.

I n t h e c u l t i v a t e d a r e a , c o c o n u t s ( Cocos n u c i f e r a

L . ) and be^-ow them e d i b l e b a n a n a s (Musa s p p . L .M u sa c e a e )

F ig .3. Study a re a a t Rantau Panjang.

~ . ’ f.’V -o 'y

'ir'rs.c

s m m

v\>-0 6 cm = 1 k m

—y d y k e □ n e t s i t e

V3* ^ m a n g r o v et l i .5 '*.* I r o a d s

viLV>H>3'£ f e . , K | v e r

.-%X ;V^ «.%**

F i g . 4 . H i s t o g r a m s sh o w in g t h e m o n th ly r a i n f a l l a t M a r d i ,

K la n g (3 0 0 ' N, 101 30 * E) an d a t Rantau-P-an-jjan'g

( 3 ° 1 4 ’N, 101 °li-1 1E) S e l a n g o r . D a ta from t h e M e t e o r ­

o l o g i c a l O f f i c e , M a l a y s i a .

B a t u C a v e s . 1 3 7 2

B a t u C a v e s . 1973

aco

os M a r d i , K l a n g . 1972

M a r d i , K l a n g . 1973".'W . I . .

M o n t h s

( s e e p . 3b f o r n o t e s on t h e c l a s s i f i c a t i o n o f e d i b l e b a n a n a s )

w e re u b i q u i t o u s e x c e p t w h e re c o f f e e , C o f f e a r o b u s t a L .

( R u b ia c e a e ) o r o i l p a lm , E l a e l s g u l n e e n s l s . s o m e t im e s r e p l a c e d

t h e b a n a n a s . V a r i o u s c u l t i v a t e d t r e e s , m o s t l y f r u i t ^>erbs7

w e re p l a n t e d among t h e c o c o n u t s n e a r h o u s e s . They i n c l u d e d

iil&ugOGS ( ncin&lLf 63Tcl 3pp • L* AnciC&i"u.Xa.CGc3.s) $ j&iuuUS ( E u g e n ia

a q u e a Burm. and l e s s commonly E . m a l a c c e n s i s L . M y r t a c e a e ) ,

n a n g k a and chem pedak ( A r t o c a r p u s h e t e r o p h y l l u s Lam. and A.

i n t e g e r ( T h u n . ) Merr." M o rac eae ) , k a p o k ( C e ib a p e n t a n d r a

G a e r t n . B o m b acaceae ) and one m a tu re d u r i a n ( D u r io z i b e t h i n u s

L . B o m b a c a c e a e ) .

R a i n f a l l d a t a d u r i n g t h e s t u d y r e c o r d e d a t M a r d i ,

K la n g ( 3 ° OO'N, 1 0 1 ° 3 0 ' E, 3 m a . s . l , ) , 14 km s o u t h e a s t o f

t h e s t u d y a r e a i s g i v e n i n F i g . 4*

The d r i e s t m on th s i n 1972 w ere May, J u n e and

A u g u s t and i n 1973 J a n u a r y , J u n e and J u l y . T h e re w e re 129

an d 171 r a i n y d a y s and 8 1 . 0 3 and- 115*78 i n c h e s o f r a i n

r e s p e c t i v e l y i n e a c h y e a r . A mean o f 0 . 6 5 5 i n c h e s o f r a i n

f e l l on e a c h r a i n y d a y o v e r t h e w h o le p e r i o d ( c f . d a t a f o r

B a tu C aves b e l o w ) .

2 . K a ja n g , S e l a n g o r .

( 2 ° 5 9 'N , 1 0 1 ° 5 V E ; a l t . 50 - 120 m a . s . l . ; F i g . 5 . )

The s t u d y a r e a was l o c a t e d i n a r u b b e r (H evea

b r a s i l i e n s i s ) p l a n t a t i o n i n w h ic h s e v e r a l s m a l l s t r e a m s

o r i g i n a t e d . Each w a t e r c o u r s e was l i n e d b y w i l d b a n a n a s

( Musa m a l a c c e n s i s R i d l e y and M. v i o l a s c e n s R i d l e y ) f o r a t

l e a s t p a r t o f i t s l e n g t h b u t t h e y had b e e n e l i m i n a t e d from

Fi g . 5 . S t u d y a r e a at K a j a n g

0-5 km

0 = N e t s i t e sm = f l o w e r i n g M u s a m a l a c c e n s i s v = ” M . v i o l a s c e n sc s ” c u l t i v a t e d b a n a n a s

9

some s e c t i o n s b y t h e p l a n t a t i o n o w n e r s . A l l t h e l a n d a d j a c e n t

t o t h e s t u d y a r e a was p l a n t e d w i t h r u b b e r and t h i s was th e

p r i n c i p a l v e g e t a t i o n t y p e f o r s e v e r a l k i l o m e t e r s a ro u n d i t .

V/ild b a n a n a s h ad b e e n ' e l i m i n a t e d from a d j a c e n t e s t a t e s . V ery

l i t t l e b e s i d e s . f e r n ? , g r a s s , a r o i d s and a few h e r b a c e o u s

p l a n t s grew b e lo w t h e r u b b e r b u t t h e r e w e re a few r a m b u ta n

( N e p h e l lu m la p p a c e u m L . S i p i n d a c e a e ) f r u i t t r e e s and some

c u l t i v a t e d b a n a n a s n e a r t h e c a r e t a k e r ' s h o u s e . A d e t a i l e d

d e s c r i p t i o n o f t h e a r e a h a s b e e n g i v e n b y Kiew ( 1 9 7 4 ) .

3 . B a tu C a v e s , S e l a n g o r .

( 3 ° 1 4 'N , 1 0 1 ° 1+1 1E ; a l t . 20 m a . s . l . ; F i g . 6 . )

B a tu C aves w e re f u l l y d e s c r i b e d b y M c C lu re ,

Lim an d Winn ( 1 9 6 8 ) . H o w ever , s i n c e t h e i r s t u d y c o n c r e t e

p a t h s and f l o o d l i g h t s h a v e b e e n p l a c e d i n c a v e r n s A, B , D

an d t h e G r e a t Room. T hese s e c t i o n s a r e now open t o t h e

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

s t u d y , r e m a in e d c l o s e d t o t h e p u b l i c and was r e l a t i v e l y

u n d i s t u r b e d .

M o n th ly r a i n f a l l d a t a f o r B a tu C av es f o r 1 9 7 1 , '

1972 and 1973 a r e g i v e n i n F i g . 4 . The d r i e s t m o n th s w e re

J a n u a r y o r F e b r u a r y and J u n e o r J u l y . The num ber o f r a i n y

d a y s p e r y e a r w as s i m i l a r t o K la n g ( 1 3 3 , 147 an d 18 8 d a y s

r e s p e c t i v e l y ) b u t t h e q u a n t i t y o f r a i n f a l l p e r y e a r was

m a r k e d ly low a t 3 3 . 3 8 , 3 2 . 7 5 and 4 8 . 9 9 i n c h e s r e s p e c t i v e l y

w i t h a mean o f 0 . 2 5 i n c h e s f a l l i n g p e r r a i n y d a y o v e r t h e

t h r e e y e a r s .

F i g . 6 M a p o f B a t u C a v e s . ( A f t e r M c C l u r e , Lim & W i n n ( 1 9 6 7 ) )

T r a y IV.

C a v e r n C

T r a y III

T r a y IL

T r a y I.

G r e a t R o o m

Hole in r o o f of G r e a t R o o mC a v e r n D

C a v e r n B

C a v e r n A

100m e t r e sT e m p le

C a v e& = c h i m n e y

E n t r a n c e

10

fhif . U n i v e r s i t y o f M alay a F i e l d S t u d i e s C e n t e r ( F . S . C . ) ; 15

m i l e , U lu Gombak, S e l a n g o r .

( 3 ° 20»N, 1 0 1 ° i f5 ’ E; a l t . 260 ra a . s . l . )

The F i e l d S t u d i e s C e n t e r h a s b e e n d e s c r i b e d b y

Medway ( 1 9 6 6 ; . ) . The h e a d q u a r t e r s , w h e re a l l o b s e r v a t i o n s

r e p o r t e d h e r e w ere m ade, i s on t h e e a s t b a n k o f t h e Gombak

R i v e r a t t h e 1 5i- m i l e s t o n e from K u a la Lumpur an d l i e s w i t h i n

an a r e a o f w e l l r e g e n e r a t e d s e c o n d a r y f o r e s t . W ild g i n g e r s

( Z i n g i b e r a c e a e ) a n d bamboo ( G i g a n t o c h l o a s c o r t e c h l n i i Gamble

G ra m in a e ) w e re p r e s e n t . W ild b a n a n a s ( Musa m a l a c c e n s i s ,

i n c l u d i n g M. m a l a c c e n s i s x M. t r u n c a t a R i d l e y h y b r i d s and

v i o l a s c e n s ) grow i n c l e a r i n g s and t h e r e i s a g r o v e o f

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

C e n t e r - i n c l u d i n g f i v e l a r g e d u r i a n t r e e s , D u r io a i b e t h i n u s „

D uabanga g r a n d i f l o r a ( S o n n e r a t i a c e a e ) , P a r k i a s p e c i o s a H a s s k . ,

P . . l a v a n i c a ( L a n k . ) M e r r . (L e g u m in o sa e ) and Bo mb ax v a l e t o n i i

H o c h r . (B o m b acaceae ) a r e common i n t h e s u r r o u n d i n g f o r e s t .

5 . T a p a h - R i n g l e t R o a d , P e r a k .

T h i s r o a d r u n s from Ta£>ah, P e r a k (if0 1 2 'N , 1 0 1 ° l 6 f E;

200 m a . s . l . ) t o R i n g l e t , P a h a n g (if0 2 5 'N , 1 0 1 ° 23*E ; 1100 m

a . s . l . ) ( F i g . 2 ) . The i n i t i a l s e v e n m i l e s p a s s e s t h r o u g h

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

s t e a d i l y a l o n g t h e s i d e o f a v a l l e y t h r o u g h a m i x t u r e o f

s e c o n d a r y f o r e s t and a b a n d o n e d r u b b e r p l a n t a t i o n s i n w h ich

f o r e s t t r e e s w e re r e g e n e r a t i n g . A t v a r i o u s p l a c e s s m a l l

c l e a r i n g s h a d b e e n made b y a b o r i g i n a l v i l l a g e r s and t h e r e

was a s m a l l p l a n t a t i o n o f i n t r o d u c e d P i n u s s p p , a t t h e 1 7 -

11

m i l e . C h i r o p t e r o p h i l o u s t r e e s w ere v e r y common a l o n g s i d e

t h e r o a d . The f o l l o w i n g s p e c i e s w e re r e c o r d e d : Musa t r u n c a t a ,

m a l a c c e n s i s ( a n d h y b r i d s o f t h e s e two s p e c i e s ) an d c u l t i v a t e d

Musa s p p . (M u sac eae ) ; C ocos n u c i f e r a (P a lm a e ) ; M a n g i f e r a s p p .

( A n a c a r d i a c e a e ) ; O roxy lum in d ic u m V e n t , and p a . j a n e l l a

-P/N 1 4 ^ T> n f TiA ^ ^ ^ N . n «*r *-\ 1 /-v 4-^*4 4 A ^ V ~j l x u x x » xj • • v j j x ^ u u u x a u v a c ; / , j J u m u a A v a x o u c j , u a

p e n t a n d r a G a e r t n . a n d D u r io z i b e t h i n u s (B o m b a c a c e a e ) ; P a r k i a

. l a v a n i c a an d P . s p e c i o s a (M im osaceae ) and D uabanga g r a n d i f l o r a

( S o n n e r a t i a c e a e ) .

EAST MALAYSIA.

1 . T e lo k Asam, Bako N a t i o n a l P a r k , S a ra w a k .

( 1 ° 4 2 'N , 1 1 0 ° 2 7 1E; F i g . 7 )

The o n l y s i t e i n B o rn eo a t w h ic h a d e t a i l e d

s t u d y w as u n d e r t a k e n w as a t T e lo k Asam i n Bako N a t i o n a l

P a r k , S a r a w a k . A a c c o u n t o f t h e p h y s i c a l f e a t u r e s o f Bako

h a v e b e e n g i v e n b y W i l f o r d (19 6 5 )> and A sh to n (1 9 7 1 ) h a s

d e s c r i b e d t h e v e g e t a t i o n .

T e lo k Asam i s a b a y l y i n g b e tw e e n two r o c k y

h e a d l a n d s on t h e e a s t c o a s t o f t h e p e n i n s u l a r . B e h in d t h e

b a y a s a n d s t o n e c l i f f r i s e s t o a p l a t e a u on w h ic h k e r i n g a s

f o r e s t g r o w s .

The s h o r e - l i n e i s b i s e c t e d by a s m a l l s t r e a m .

To t h e s o u t h o f i t i s a s a n d y b e a c h b o r d e r e d b y a n a r r o w

b a n d o f v e g e t a t i o n c h a r a t e r i s e d b y common r h u , C a s u a r i n a

e q u i s e t i f o l i a L . ( C a s u a r i n a c e a e ) , b e h i n d w h ic h i s s e c o n d a r y

s c r u b a n d a l i t t l e p r i m a r y lo w la n d f o r e s t . The p a r k b u i l d i n g s

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

. 7 M a p o f T e lo k A s a m , B a k o N a t i o n a l P a r k , S a r a w a k .

Telok Paku

s a n d y b e a c h

yO' Sonnerr.tia a lba

12

t h e r e i s an e x t e n s i v e s t a n d o f m a tu re S o n n e r a t i a a l b a J . . J . S m i t h .

A f o o t - b r i d g e , 1 $1+ m l o n g , h a s b e e n b u i l t i n a s t r a i g h t l i n e

3 m a b o v e t h e mud a c r o s s p a r t o f t h e m a n g ro v e s ( F i g . 7 and

8 an d s e e p . 76 ) . A few i s o l a t e d t r e e s o f o t h e r m angrove

s p e c i e s , n o t a b l y R h iz o p h o r a s p p . grow a l o n g t h e u p p e r t i d a l

r e a c h e s o f t h e s t r e a m . The o n l y o t h e r kno>vn c h i r o p t e r o p h i l o u s

p l a n t s r e c o r d e d t h e r e w e re a few c o c o n u t p a lm s ( Cocos n u c i f e r a )

b u t D u r io c a r i n a t u s M a s t , w h ic h t h e b a t s may f e e d on grew

i n t h e f o r e s t b e h i n d t h e b a y .

13

METHODS.

T h i s s tud y , was p r i m a r i l y c o n c e r n e d w i t h t h e

t h r e e / r w ie ro g lo s s in e b a t s fo u n d i n P e n i n s u l a r M a l a y s i a . Most

o f t h e p l a n t s 011 w h ic h t h e y f e e d h av e b e e n d e s c r i b e d and

f i g u r e d i n s t a n d a r d t e x t b o o k s an d e xam ined i n r e l a t i o n t o

c h i r o p t e r o p h i l y by v a r i o u s w o r k e r s e i t h e r i n J a v a o r e l s e w h e r e

( e . g . A f r i c a o r A m e r i c a ) . D e s p i t e t h i s , e v e r y o p p o r t u n i t y

was t a k e n t o exam in e t h e p l a n t s a s w e l l a s t h e b a t s . The

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

o f t h e b a t s 1 fo o d s o u r c e s f a c i l i t a t e s an u n d e r s t a n d i n g o f

t h e i r f e e d i n g b i o l o g y .

CHIROPTEROPHILOUS PLANTS.

The e c o l o g i c a l d i s t r i b u t i o n , m o r p h o lo g y s .

f l o w e r i n g b i o l o g y an d p h e n o lo g y o f a l l t h e known c h i r o p t e r o p h i l o u s

p l a n t s w e re s t u d i e d ' w h e r e v e r p o s s i b l e .

E c o l o g i c a l d i s t r i b u t i o n .

N o te s w e re k e p t o f t h e h a b i t a t s i n w h ich e a c h

s p e c i e s g r e w , w i t h p a r t i c u l a r a t t e n t i o n b e i n g p a i d t o an y

s p e c i a l i s e d r e q u i r e m e n t s w i t h i n t h o s e h a b i t a t s e . g . a l t i t u d i n a l

z o n a t i o n , z o n a t i o n i n m angrove f o r e s t s and p r e f e r e n c e f o r

open o r s h a d e d c o n d i t i o n s . The e f f e c t s o f man on t h e a b u n d a n c e

an d d i s t r i b u t i o n o f e a c h s p e c i e s was a l s o r e c o r d e d .

The a l t i t u d i n a l z o n a t i o n o f c h i r o p t e r o p h i l o u s

p l a n t s a l o n g t h e r o a d from Tapah t o Tanah R a t a v i a R i n g l e t

was i n v e s t i g a t e d by d r i v i n g t h e 39 m i l e s from T anah R a t a t o

T apah on 1 5 S e p te m b e r 1972 a t b e tw e e n 20 and 3 5 m . p . h . At

e v e r y m i l e p o s t f rom T anah R a t a (1*1-33 m a . s . l . ) t h e a l t i t u d e

/

I

n

r e g i s t e r e d on an. a n e r o i d a l t i m e t e r was r e c o r d e d . (The a l t i m e t e r

w h ich was c a l i b r a t e d i n 100 f t d i v i s i o n s b u t c o u l d b e r e a d

t o 2 5 f t , h a d b e e n a d j u s t e d t o r e a d J+ 7 5 0 f t (= 1 ^ -5 0 m) a tI'd.Tanah R a t a an d i t s r e a d i n g a t t h e 23 m i l e f rom T apah c h e c k e d

a g a i n s t a s i g n b o a r d a l o n g s i d e t h e r o a d w h ich r e c o r d e d t h e

a l t i t u d e a s 2500 f t (= ye>2 m) a . e . i . a t t h a t p o i n t . ) The

num ber o f t r e e s o f e v e r y known c h i r o p t e r o p h i l o u s s p e c i e s s e e n

w i t h i n a b o u t 5 0 m o f t h e r o a d b e tw e e n e a c h m i l e p o s t from

Tanah R a t a t o t h e b e g i n n i n g o f t h e w e l l managed r u b b e r e s t a t e s

a t t h e 7*"*1 m i l e from T apah (75 0 f t = 229 m a . s . l . ) was n o t e d .

B a n a n a s (Musa s p p . ) w ere n o t c o u n t e d b u t a r e c o r d w as k e p t

o f w h ich s p e c i e s w as s e e n i n e a c h one m i l e s e c t i o n .

The d i s t r i b u t i o n o f t h e t h r e e S o n n e r a t i a s p e c i e s

( a l l m angrove t r e e s ) a l o n g t h e R i v e r K la n g , p a r t o f t h e R i v e r

L a n g a t , t h e c o a s t f ro m P o r t K la n g t o Kampong B a t u Empat (o n e

km n o r t h o f R a n ta u P a n j a n g ) an d i s l a n d s o f f t h e S e l a n g o r c o a s t ,

w e re m apped . M ost o f t h e r i v e r b a n k s w e re a c c e s s i b l e b y r o a d

o r on f o o t b u t a s m a l l b o a t was u s e d t o e x p l o r e t h e i s l a n d s

and t h e c o a s t l i n e i n c l u d i n g a l l t h e l a r g e r c r e e k s .

F lo w e r m o r p h o lo g y .

The m o rp h o lo g y o f t h e f l o w e r s a nd t h e s t r u c t u r e

o f t h e i n f l o r e s c e n c e s o f m o s t s p e c i e s w e re e x am in ed i n t h e

f i e l d a n d , i n g r e a t e r d e t a i l , i n t h e l a b o r a t o r y . P a r t i c u l a r

a t t e n t i o n was p a i d t o t h o s e f e a t u r e s w h ich r e l a t e d t o

c h i r o p t e r o p h i l y , T h e se i n c l u d e d

1 . The p o s i t i o n o f t h e f l o w e r o r i n f l o r e s c e n c e s on

t h e p l a n t .

15

2 , - The s t r u c t u r e o f t h e i n f l o r e s c e n c e .

3 . The s t r u c t u r e o f i n d i v i d u a l f l o v / e r s i n c l u d i n g t h e '

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

o f t h e v a r i o u s f l o r a l m em bers .

' Zf. The q u a n t i t y o f n e c t a r p ro d u c e d b y t h e f l o w e r s .

( T h i s was m e a s u re d b y d ra w in g i t i n t o a 2 , 0 ml

d i s p o s a b l e s y r i n g e g r a d u a t e d i n 0 .1 ml d i v i s i o n s .

I n t h e c a s e o f Musa s p p . t h e n e c t a r f rom f o u r f l o w e r s

was draw n i n t o t h e s y r i n g e and t h e t o t a l q u a n t i t y

m e a s u re d e a c h t i m e . The mean vo lum e w as e a s i l y c a l c u l a t e d

an d t h e m e th od r e d u c e d t h e e r r o r i n v o l v e d i n r e a d i n g

t h e v o lu m e . The q u a n t i t y o f n e c t a r i n a f l o w e r was

m e a s u re d d i r e c t l y a f t e r a n t h e s i s b u t b e f o r e a n y

v i s i t o r s h a d l a n d e d on t h e f l o w e r s . I f n e c e s s a r y ,

t h e b u d s w e re p r o t e c t e d b y p a p e r b a g s t o e n s u r e t h a t

no v i s i t o r s f e d from t h e f l o v / e r s a f t e r t h e y o p e n e d .

5» The n a t u r e o f t h e p o l l e n i . e . w h e t h e r i t was p o w d e ry ,

s t i c k y o r i n t e r m e d i a t e : The names a r e s e l f e x p l a n a t o r y .

6 . The s c e n t .

F l o w e r i n g b i o l o g y .

F i e l d o b s e r v a t i o n s v/ere made on t h e t im e and

d u r a t i o n o f a n t h e s i s . A l l a n i m a l s s e e n v i s i t i n g t h e f l o v / e r s

w e re r e c o r d e d an d t h e i r b e h a v i o u r on t h e f l o w e r s w as c a r e f u l l y

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

a s p o l l i n a t o r s . The a p p r o x i m a t e number o f f l o w e r s t h a t s e t

f r u i t was r e c o r d e d .

P h e n o l o g y .

I n d i v i d u a l s t r e e s o f m o s t s p e c i e s w e r e , o b s e r v e d

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

a b s e n c e o f f l o w e r s a nd f r u i t , a s w e l l a s e v i d e n c e o f l e a f

c h a n g e , w e re r e c o r d e d e a c h t i m e . Any i n c i d e n t a l b u t r e l e v a n t

o b s e r v a t i o n s made a t a n y t im e on an y known c h i r o p t e r o p h i l o u s

t r e e s w ere r e c o r d e d .

B e c a u s e t h e p o l l e n o f P a r k l a j a v a n i c a was

i n d i s t i n g u i s h a b l e from t h a t o f P . s p e c i o s a i t w as o f p a r t i c u l a r

i m p o r t a n c e t o t h e i n t e r p r e t a t i o n o f t h e a n a l y s i s o f p o l l e n

r e c o v e r e d from t h e b o d i e s o r gu an o o f b a t s t o know i f t h e s e

two t r e e s f l o w e r e d s i m u l t a n e o u s l y . S i n c e t h e t r e e s a r e d i f f i c u l t

t o i d e n t i f y f rom a d i s t a n c e i n t h e f i e l d 24 P . . j a v a n i c a and

til til32 P . s p e c i o s a t r e e s g ro w in g b e tw e e n t h e 7 m i l e an d 14

m i l e on t h e T a p a h - R i n g l e t r o a d w e re i n d i v i d u a l l y i d e n t i f i e d

f ro m l e a f l e t s and n u m b e re d . (S e e C o r n e r , 195?- o r W h itm o re , 1972

f o r t h e i d e n t i f i c a t i o n o f t h e s e P a r k i a s p e c i e s from l e a f l e t s . )

Once a month from J u n e 1972 t o D ecem ber 1973 e a c h t r e e was

ex am in ed and t h e p r e s e n c e o f f l o w e r s o r f r u i t an d a n y e v i d e n c e

o f l e a f c h a n g e w e re r e c o r d e d . R e g u l a r o b s e r v a t i o n s w e re a l s o

m a i n t a i n e d on t h e P . s n e c i o s a t r e e a t t h e 2 1 mi l e , U lu Gombak

w h ich h a d b e e n o b s e r v e d o v e r n i n e y e a r s from 1 96 1 t o 1 9 6 9

by Medway (1972c).

R e p r o d u c t i v e c o m p a t a b l l i t y . -

P r e l i m i n a r y e x p e r i m e n t s w ere c o n d u c te d t o d e t e r m i n e

w h e th e r S o n n e r a t i a a l b a w as s e l f c o m p u ta b le o r n o t . The s t a m e n s

w e re rem ov ed from 2 0 f l o w e r s j u s t b e f o r e t h e b u d s o p e n e d and

17

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

w ere h a n d p o l l i n a t e d w i t h p o l l e n o b t a i n e d from open f l o w e r s .

Ten w e re p o l l i n a t e d v / i th p o l l e n from o t h e r f l o v / e r s on t h e

same t r e e t h a t t h e y grew on an d t e n w e re p o l l i n a t e d v / i th

p o l l e n from f l o w e r s on n e i g h b o u r i n g t r e e s . The f l o v / e r s w ere

a g a i n c o v e r e d w i t h p a p e r b a g s f o r two d a y s and t h e num ber

t h a t s e t f r u i t w as r e c o r d e d two w eeks l a t e r .

MAGROGLOSSINE BATS.

F i e l d o b s e r v a t i o n s .

B a t s w e re observed;, i n t h e f i e l d e i t h e r b y moon­

l i g h t . o r b y a r t i f i c i a l l i g h t . M o o n l ig h t was b r i g h t enough t o

a l l o w o b s e r v a t i o n s f o r a b o u t one w eek e i t h e r s i d e o f f u l l

moon; h o w e v e r , f o l l o w i n g f u l l moon t h e moon d o e s n o t r i s e

u n t i l a f t e r s u n s e t . T h e r e f o r e , t h e o n ly w eek i n e a c h l u n a r

month d u r i n g w h ic h b a t s c o u l d b e o b s e r v e d b y m o o n l i g h t

d i r e c t l y a f t e r s u n s e t v/as t h e w eek p r e c e d i n g f u l l moon.

An e l e c t r i c f l a s h - l i g h t ( w i t h n e a r l y dud b a t t e r i e s )

was p l a c e d on t h e g ro u n d t o i l l u m i n a t e f l o w e r s f o r o b s e r v a t i o n .

The t h r e e s p e c i e s o f b a t s b e i n g s t u d i e d w e re a p p a r e n t l y

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

was d im an d m o t i o n l e s s . They w ere f r i g h t e n e d i f t h e beam was

b r i g h t o r moved, e v e n when i t was c o v e r e d b y a d o u b le l a y e r

o f d e e p r e d c e l l o p h a n e p a p e r .

By e i t h e r o f t h e s e m e th o d s i t v/as u s u a l l y

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

t h e t i m e , a p p r o x i m a t e d u r a t i o n and f r e q u e n c y o f v i s i t s , b e s i d e s

n o t i n g an y o t h e r l a r g e v i s i t o r s , e . g . s p h i n g i d m o t h s . I t was

18

n o t p o s s i b l e t o d e t e r m i n e t h e b a t ' s a g e , s e x n o r d e t a i l s o f

i t s b e h a v i o u r on t h e f l o w e r . I n h e a v y shad ow , e . g . i n a

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

a n d , c o n s e q u e n t l y , l e s s d a t a c o u l d b e g a t h e r e d .

P h o t o g r a p h y .

B a t s w e re p h o t o g r a p h e d a s t h e y v i s i t e d v a r i o u s

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

from t h e f l o w e r s a s w e l l a s t h e i r f e e d i n g p o s t u r e s on t h e

f l o w e r s c o u l d b e s t u d i e d . E q u ip m e n t u s e d v/as a P e n t a x SP500

ca m e ra b o d y w i t h a P e n t a x t e l e p h o t o l e n s ( f = 200 mm) m oun ted

on a t r i p o d ; a n i n e i n c h c a b l e r e l e a s e ; a N a t i o n a l E l e c t r i c

F l a s h m o de l PE 1 82S ( d u r a t i o n 1 / 1 0 0 0 s ) m oun ted d i r e c t l y ab o v e

t h e c a m e ra ; 400 ASA b l a c k an d v /h i t e Kodak f i l m o r 80 ASA

( " E k t a c h r o m e " ) c o l o u r f i l m . The a p p a r a t u s was a s s e m b l e d and

fo c u s e d on an a p p r o p r i a t e f l o w e r b e f o r e d a r k . The s h u t t e r

was r e l e a s e d w h e n e v e r a b a t a p p r o a c h e d t h e f l o w e r .

A l l o t h e r p h o t o g r a p h s ( e . g . o f i n d i v i d u a l f l o w e r s

o r h a b i t a t s ) w e re t a k e n w i t h t h e same ca m e ra a n d f l a s h

a t t a c h m e n t b u t w i t h a s t a n d a r d P e n t a x l e n s ( f = 55 mm).

S h o o t i n g .

One b a t v/as s h o t w h i l e v i s i t i n g a f l o w e r so t h a t

i t s i d e n t i t y c o u l d b e v e r i f i e d .

M i s t n e t t i n g .

M i s t n e t s w e re u s e d t o c a t c h b a t s i n t h e f i e l d .

M ost n e t s w ere 10 m lo n g a l t h o u g h 20 m n e t s w e re o c c a s i o n a l l y

u s e d . N e t s w e re u s u a l l y s e t w i t h t h e lo v / e s t s h e l f a p p r o x i m a t e l y

1 0 cm a b o v e t h e g ro u n d a nd a d j a c e n t t o o r b e lo w f l o w e r i n g

,/Ii

19

p l a n t s t h a t w ere known o r s u s p e c t e d t o b e c h i r o p t e r o p h i l o u s .

I n Bako N a t i o n a l P a r k t h e p o l e s s u p p o r t i n g t h e n e t s w e re t i e d

t o t h e r a i l i n g s o f t h e b r i d g e t h a t t r a v e r s e d ' t h e m a n g ro v e s

( p . 11 and F i g , 8 ) . The l o w e s t s h e l v e s w ere t h e n l e v e l w i t h

t h e h a n d r a i l , k m ab o v e t h e mud a t low t i d e o r 2 m a b o v e

t h e w a t e r s u r f a c e a t hxgh t x d e . One 2 . 5 x £j-»0 m n e t was

m o d i f i e d so t h a t i t c o u l d b e r a i s e d i n t o t h e c a n o p y from

g ro u n d l e v e l ( F i g . 1 0 ) . F i v e 10 m n e t s ( o r o c c a s i o n a l l y f o u r

1 0 m n e t s an d one 2 0 m n e t ) w ere s e t e a c h t im e u n l e s s o t h e r ­

w i s e s p e c i f i e d . They w e re i n s p e c t e d a t l e a s t once e v e r y h a l f

h o u r u n t i l m i d n i g h t a f t e r w h ic h t h e y w ere i n s p e c t e d a t l e a s t

o nce e v e r y h o u r .

I n t h e s t u d y a r e a s a t R a n ta u P a n j a n g an d K a j a n g ,

r e g u l a r n e t s i t e s w e re e s t a b l i s h e d , nu m bered an d mapped' ( F i g s .

3 and 5 ) • H o w ever , t o p r e v e n t b a t s l e a r n i n g t h e i r w h e r e a b o u t s ,

n e t s w e re n e v e r s e t more t h a n t w i c e i n p r e c i s e l y t h e same

p o s i t i o n s a t an y n e t s i t e .

R i n g i n g .

A lm o s t a l l M a c r o g lo s s u s s p p . a nd E o n y c t e r i s s p e l a e a

c a u g h t i n t h e s t u d y a r e a s w e re r i n g e d . M onel b a t r i n g s , s e r i e s

2 0 - f o r M a c r o g l o s s u s s p p . an d 3 0 - f o r E . s p e l a e a w e re p r o v i d e d

by C . S . I . R . O . A u s t r a l i a . T h e i r u s e on E . s p e l a e a was d i s c o n t i n u e d

a f t e r f o u r m on th s b e c a u s e o f t h e damage t h e y i n f l i c t e d on th e

b a t s . S e r i e s 060 b i r d r i n g s s u p p l i e d by t h e U n i v e r s i t y o f

M alay a B i r d B a n d in g Scheme w ere u s e d i n s t e a d . ' R in g s w ere p l a c e d

on t h e r i g h t ' f o r e a r m ,a b o u t 0 . 5 mm from t h e w r i s t , t h r o u g h a

s m a l l i n c i s i o n made i n t h e a n t e p a t a g i u m w i t h t h e p o i n t o f a

s i z e 10 S w ann-M orton s c a l p e l b l a d e ( F i g . 10 ) . I f a r e c a p t u r e d

Fig. 8 . A section of the bridge at Telok Asam, Bako National Park.

Fig. 9 . Guano collecting Tray III in Cavern C , BatuCaves. Note the small pool of water from drips from the ceiling.

20

b a t showed a n y s i g n s o f i n f l a r a a t i o n o r damage c a u s e d by t h e

r i n g t h e l a t t e r w as rem oved an d a new one p l a c e d on t h e o t h e r

f o r e a r m . B e c a u s e t h e n o r m a l m e th od o f c l o s i n g b i r d r i n g s

w i t h p l i e r s i s n o t f e a s i b l e on a b a t * s w in g , b i r d r i n g s had

t o b e r e s h a p e d b y c l o s i n g and p r i s i n g them open a g a i n b e f o r e

p l a c i n g them o n t o t h e b a t ' s f o r e a r m ( f i g . 1 0 ) .

M ost M a c r o g l o s s u s s p p . c a u g h t o u t s i d e s t u d y a r e a s

w e re n o t r i n g e d b u t m o s t E . s p e l a e a ( e x c e p t t h o s e c o l l e c t e d

f o r l a b o r a t o r y s t u d y ) w e re r i n g e d b e f o r e r e l e a s e . Where

p o s s i b l e , t h e f o l l o w i n g d a t a w ere r e c o r d e d f o r a l l b a t s a t

t h e t im e o f c a p t u r e .

L o c a l i t y , an d i n s t u d y a r e a s a t R a n ta u P a n j a n g an d K a ja n g t h e

n e t n u m b e r .

D a t e . To a v o i d c o n f u s i o n v/hen a n a l y s i n g r e t r a p d a t a a l l b a t s

c a u g h t i n one n i g h t , w h e t h e r b e f o r e o r a f t e r m i d n i g h t , h a v e

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

2 3 .0 0 h r s on 1 6 November 1972 and a t 0 1 . 0 0 h r s 17 November

1972 a r e a l l r e f e r r e d t o 1 6 November 1 9 7 2 .

T im e .

Species.

W e ig h t i n g ram s b y u s e o f a P e s o l a b a l a n c e ( a 0 - 30 g b a l a n c e

f o r M a c r o g l o s s u s s p p . a n d a 0 - 300 g b a l a n c e f o r E . s p e l a e a ) .

F o re a rm l e n g t h i n mm m e a s u re d t o 0 . 5 mm on t h e c l o s e d w in g •

w i t h v e r n i e r c a l i p e r s .

A g e . F r e e f l y i n g b a t s w e re r e c o r d e d a s s u b a d u l t s o r a d u l t s .

R e p r o d u c t i v e m a t u r i t y was t h e d e f i n i t i v e c h a r a c t e r u s e d

t o d i s t i n g u i s h a d u l t s f rom s u b a d u l t s . I n a l l t h r e e s p e c i e s

a d u l t m a le s w e re d i s t i n g u i s h e d from s u b a d u l t s by t h e

s u p p o r t i n g r o p e

F i g . 10.

< >

« i

MI ST NET DES I GNED FOR USE IN TREES

s t a b i l i s i n g r o p e

BAT W I N G TO S H O W P O S I T I O N O F R I N G

S H A P E O F BIRD RINGS M O D I F I E D FOR USE O N BATSA . u n m o d i f i edB. c l o s e dC. r e o p e n e d

21

p r e s e n c e o f w e l l d e v e lo p e d s c r o t a l t e s t e s i n t h e fo r m e r

( a n d i n M a c r o g l o s s u s m in im u s b y t h e d e g r e e o f d e v e lo p m e n t

o f t h e d e r m a l t h r o a t g l a n d , s e e b e l o w ) . F e m a le s w e re r e c o r d e d

a s a d u l t s i f t h e y w e re p r e g n a n t ( s e e b e l o w ) , l a c t a t i n g o r

i f e n l a r g e m e n t o f t h e n i p p l e s i n d i c a t e d e a r l i e r l a c t a t i o n s .

( C a re i s n e e d e d a s t h e n i p p l e s may r e g r e s s a l m o s t t o t h e

s i z e o f t h o s e o f s u b a d u l t s i f t h e i n t e r v a l b e tw e e n l a c t a t i o n s

i s a b n o r m a l l y l o n g . ) S u b a d u l t s u s u a l l y w e ig h e d l e s s t h a n

a d u l t s b u t t h e f o r e a r m l e n g t h s o f a l l b u t t h e y o u n g e s t f r e e

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

T h e r e f o r e w e i g h t was a u s e f u l ' . s u p p l e m e n t a r y g u i d e t o

m a t u r i t y b u t f o r e a r m l e n g t h v/as n o t . The e p i p h y s e s o f t h e

m e t a c a r p a l s o f s u b a d u l t s w e re i n c o m p l e t e l y f u s e d . T h i s

c o n d i t i o n p e r s i s t e d i n t o e a r l y a d u l t h o o d . Thus c o m p l e t e

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

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

w e re r e c o r d e d a s j u v e n i l e s .

R e p r o d u c t i v e s t a t u s .

The s i z e o f t h e t e s t e s o f m a le s w as r e c o r d e d a s s m a l l ( l e s s

t h a n 2 mm lo n g ) o r l a r g e ( o v e r 8 mm l o n g ) . I f t h e y w ere

i n t e r m e d i a t e t h e i r l e n g t h was r e c o r d e d . A l l m e a s u re m e n ts

w e re made w i t h v e r n i e r c a l i p e r s t o t h e n e a r e s t mm th r o u g h

t h e s k i n o f t h e s c r o t u m . P r e g n a n c y i n a d u l t f e m a le s v/as

r e a d i l y d e t e c t e d b y p a l p a t i o n some t im e a f t e r t h e i m p l a n t a t i o n

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

an d was g r a d e d i n t o one o f t h e c a t e g o r i e s l i s t e d b e lo w .

L a c t a t i o n w as o b v i o u s from t h e g r e a t l y e n l a r g e d n i p p l e s

f rom w h ic h m i l k c o u l d b e e x t r a c t e d b y g e n t l e p r e s s u r e .

22

i . N e i t h e r l a c t a t i n g n o r p r e g n a n t .

ii. L a c t a t i n g , n o t p r e g n a n t .

iii.Early pregnancy. Uterine swelling palpable, firm, round, up to 5 mm diameter.

iv. 1 / 3 pregnant. Uterine swelling over 3 ram, firm, round. v; 2/3 pregnant. As in 1/3 pregnant but uterine swelling

l a r g e r an d no l o n g e r f i r m . A b d o m in a l s w e l l i n g may b e

n o t i c e a b l e i n t h e l a t t e r p a r t o f t h i s p h a s e ,

v i . 3 / 3 p r e g n a n t . Embryo n e a r l y f u l l y fo r m e d , i t s s h a p e

d i s c e r n a b l e b y b y p a l p a t i o n , a b d o m in a l s w e l l i n g n o t i c e a b l e .

Mammary g l a n d s e n l a r g e d u r i n g t h i s p h a s e .

G l a n d u l a r c o l l a r .

The p r e s e n c e o r a b s e n c e o f a d e r m a l g l a n d u l a r c o l l a r on t h e

t h r o a t was r e c o r d e d i n a l l M a c r o g lo s s u s s p p . a s f o l l o w s

1 . A b s e n t .

2 . P r e s e n t , s m a l l , c o l o u r t h e same a s a d j a c e n t s k i n .

3 . P r e s e n t , s m a l l , r e d .

k* P r e s e n t , l a r g e , d e e p r e d t o c h o c o l a t e b ro w n .

C a p t i v e b a t s .

A t t e m p t s w e re made t o k e e p M. m in im us and

s p e l a e a i n c a p t i v i t y i n a c a g e t h a t m e a s u re d 1 m x 1 m x

1 . 3 ra h i g h . I t h a d a wooden b a s e and p l a s t i c g a u z e s i d e s and

r o o f . A s m a l l p a lm t r e e i n a f l o w e r p o t was p r o v i d e d f o r t h e

b a t s t o h a n g f r o m . The c a g e was k e p t i n t h e s h a d e . The b a t s

w e re f e d on a s o l u t i o n o f ,,Complanu and s u g a r i n w a t e r , a s

d e s c r i b e d b y M a r s h a l l a n d Lim ( 1 9 6 8 ) . A l th o u g h t h e y f e d

r e a d i l y a nd b ecam e r e m a r k a b l y tame t h e y l o s t c o n d i t i o n r a p i d l y

II

23

and d i e d w i t h i n two w e e k s .

Wing c h a r a c t e r i s t i c s .

The m e th o d s d e v i s e d h y F i n d l e y , S t u d i e r and

W i l s o n (1 9 7 2 ) t o d e t e r m i n e i n d i c e s f o r t h e l e n g t h o f t h e w in g

t i p r e l a t i v e t o t h e r e s t o f t h e w in g , t h e w in g a r e a , t h e w ing

l o a d i n g an d t h e a s p e c t r a t i o o f b a t s w ere a d o p t e d b e c a u s e

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

m e n ts c o u l d b e o b t a i n e d q u i c k l y i n t h e f i e l d from b a t s t h a t

w ere l a t e r r e l e a s e d . The m e a s u re m e n ts r e q u i r e d f o r t h e

c a l c u l a t i o n o f t h e i n d i c e s w e re t h e l e n g t h o f t h e f o r e a r m ,

I'd . t h .3 f d i g i t , § d i g i t a n d , w h e re a p p l i c a b l e , t h e w e i g h t o f t h e

b a t . The i n d i c e s w e re c a l c u l a t e d b y : -

w ing t i p i n d e x3 r d d i g i t

f o r e a r m

w in g a r e a i n d e x = ( f o r e a r m + 3 r d d i g i t ) x ( ^ ^ d i g i t ) x 2

w e i g h tw in g l o a d i n g i n d e x = ••••—----------- --------------

w in g a r e a i n d e x

a s p e c t r a t i o i n d e x =( f o r e a r m + 3 r<* d i g i t )

5 t h d i g i t

A f i f t h i n d e x , t h e w ing l e n g t h i n d e x , h a s b e e n

d e v i s e d t o r e l a t e t h e r e l a t i v e l e n g t h o f t h e b a t s * w in g s t o

t h e i r s i z e ( u s i n g w e i g h t a s a m e a s u re o f s i z e ) b y

( f o r e a r m + 3 r d d i g i t ) w in g l e n g t h i n d e x = —

w e i g h t

2/+

( F i n d l e y _et. a l . ( o p p . c i t . ) d i s c u s s t h e i m p l i c a t i o n s o f

r e l a t i v e w in g l e n g t h on t h e f l i g h t c a p a b i l i t i e s o f b a t s b u t

do n o t g i v e a s t a n d a r d m ethod f o r d e t e r m i n i n g t h e r e l a t i v e

l e n g t h o f b a t s * w i n g s . )

OCCURRENCE OF POLLEN.The e x i n e o f p o l l e n g r a i n s i s d u r a b l e an d i n e r t

( e . g . W e s t , 1 9 7 1 )* I t i s n o t d i g e s t e d b y t h e b a t s an d may b e

r e c o v e r e d from f r e s h f a e c e s o r (m o s t s p e c i e s ) from f r e s h

guano a s w e l l a s from t h e f u r a n d / o r t h e m em branes o f b a t s .

P o l l e n e x i n e s a r e i d e n t i f i a b l e t o v a r y i n g d e g r e e s , some t y p e s

t o s p e c i e s l e v e l , o t h e r s o n ly t o g e n u s o r f a m i l y l e v e l ( T a b l e 3 3 ) .

Preparation of pollen samples.A c e t o l y s i s , t h e s t a n d a r d m eth od f o r p r e p a r i n g

p o l l e n f o r m i c r o s c o p i c a l a n a l y s i s (E rd tm a n , 195 2 ) w as se ld o m

u s e d b e c a u s e i t i s t im e c o n su m in g and o n l y f e a s i b l e i n t h e

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

D u r io s p p . and Musa s p p . , b o t h i m p o r t a n t i n t h e d i e t s o f b a t s .

I n s t e a d , p o l l e n was s t a i n e d w i t h b a s i c f u s c h i n i n an a q u e o u s

s o l u t i o n o f p a r t i a l l y h y d r o l y s e d p o l y v i n a l a l c o h o l ( " G e lv a to l* *

g r a d e ifO - 20 m a n u f a c t u r e d b y S h a w i n i g a n L t d . , L o n d o n ) . The

s o l u t i o n was made by a d d i n g a few g r a i n s o f c r y s t a l l i n e b a s i c

f u s c h i n t o a p p r o x i m a t e l y e q u a l v o lu m es o f P .V .A . p e l l e t s and

c o l d w a t e r . The m i x t u r e was s t i r r e d f r e q u e n t l y u n t i l i t v/as

o f a u n i f o r m t r e a c l e - l i k e c o n s i s t e n c y . I t u s u a l l y t a k e s two d a y s

t o d i s s o l v e a l l t h e P .V .A . The s t a i n i s e f f e c t i v e w i t h i n

m i n u t e s . The a d v a n t a g e s o f t h i s medium w e r e ; t h a t i t d i d n o t

25

d e s t r o y a n y p o l l e n g r a i n s ; t h e m ethod em ployed t o make s l i d e s

( s e e b e lo w ) v/as s u f f i c i e n t l y s i m p l e and f a s t t o p e r m i t s a m p le s

t o b e m ou n ted i n t h e f i e l d a t n i g h t ; i t was c h e a p a n d , b e i n g

w a t e r s o l u b l e , g l a s s w a r e i n c l u d i n g s l i d e s and c o v e r s l i p s

w a s . e a s i l y w a sh e d f o r r e u s e . As l o n g a s t h e s l i d e s an d t h e

c o v e r s l i p s a r e f r e e from g r e a s e t h e p r e p a r a t i o n s a r e semx^

p e r m a n e n t . They w e re l a b e l l e d w i t h a f e l t t i p m a r k in g p e n .

Each t im e a b a t was c a u g h t w i t h p o l l e n on i t s

f u r o r m em bran es , a s a m p le was c o l l e c t e d by d r a w in g a s c a l p e l

b l a d e g e n t l y a c r o s s t h e a r e a . The p o l l e n w h ic h a d h e r e d t o t h e

l e a d i n g edg e o f t h e b l a d e was t r a n s f e r r e d t o a g l a s s s l i d e ,

m ixed w i t h a d r o p o f b a s i c f u s c h i n and p o l y v i n a l a l c o h o l and

a c o v e r s l i p a d d e d . The d i s t r i b u t i o n o f t h e p o l l e n on t h e

b a t was r e c o r d e d . W h ile b e i n g h a n d l e d , t h e b a t s f r e q u e n t l y

d e f a e c a t e d . S a m p le s o f t h e f a e c a l p e l l e t s w e re m o u n ted on

g l a s s s l i d e s b y t h e same m e th o d .

C o l l e c t i o n o f gu ano from E. s p e l a e a .

E . s p e l a e a o f t e n r o o s t s i n d e n s e c lu m p s on t h e

r o o f s o f r e a v e s . I n an a t t e m p t t o m o n i t o r t h e i r fo o d s o u r c e s ,

g uano was c o l l e c t e d f o r p o l l e n a n a l y s i s f rom c o l o n i e s i n

B a tu C aves and Gua S a n d in g ( F i g . 2 ) . I n B a tu C av es one s a m p le

was c o l l e c t e d e v e r y week from b e lo w e a c h o f f o u r r o o s t i n g

g r o u p s o f E . s p e l a e a ( F i g . 6) f rom 19 F e b r u a r y 1972 t o

28 J a n u a r y 1 97*f ( e x c e p t on two o c c a s i o n s when c o l l e c t i o n s

w e re two w eeks a p a r t , v i z . 8 - 2 2 May 1972 an d 27 A u g u s t -

10 S e p te m b e r 1 9 7 3 ) . Kach s a m p le c o n s i s t e d o f t h e gu ano f a l l i n g

on a t r a y c o n s i s t i n g o f a s h e e t o f p o l y t h e n e m e a s u r in g 60 cm

26

x 75 cm s u p p o r t e d on a wooden f r a m e w o rk '3 0 cm a b o v e t h e c a v e

f l o o r ( F i g . 9 ) . The p o l y t h e n e was r e p l a c e d e v e r y a l t e r n a t e

week and t u r n e d o v e r on t h e i n t e r v e n i n g w eeks : T h u s , e a c h

w e e k ly s a m p le was c o l l e c t e d on a c l e a n s h e e t .

Each s a m p le was s c r a p e d from t h e t r a y i n t o a

p o l y t h e n e b a g w i t h a c l e a n wooden s p a t u l a , n u m bered and b r o u g h t

b a c k t o t h e l a b o r a t o r y w h e re i t v/as w e ig h e d , t h o r o u g h l y mixed

and a s u b s a m p le t a k e n ( a p p r o x i m a t e l y 30 g ) . 0 . 5 g o f KOH

p e l l e t s w e re d i s s o l v e d i n a 5 g p o r t i o n o f e a c h s u b s a m p le

and a f t e r th o r o u g h m ix in g t h e s o l u t i o n was a l l o w e d t o s t a n d

f o r two d a y s . (T he KOH e n s u r e d t h e b reak d o w n o f an y s m a l l

c lu m ps o f p o l l e n g r a i n s t h a t r e m a i n e d . ) I t was t h e n sh a k e n

v i g o r o u s l y and a few d r o p s m ixed w i t h b a s i c f u s c h i n i n P .V .A .

i n a p e t r i d i s h . T h i s v/as t r a n s f e r r e d t o a g l a s s s l i d e and

a c o v e r s l i p a d d e d .

From 22 March 1972 t o 8 November 1 973 a s e c o n d

s u b s a m p le was c o l l e c t e d i n a p r e w e ig h e d g l a s s c o n t a i n e r ( w t . =

W ^ ) . .T h e c o n t a i n e r + gu ano v/as r e w e ig h e d ( w t . = W^) and t h e

gu ano d r i e d i n an oven a t 105°C t o c o n s t a n t w e i g h t (= W ^) .

A l l w e i g h t s v/ere t o 0 .01 g on a M e t t l e r Beam B a l a n c e .

The % d r y w eight(DW ) was c a l c u l a t e d by

(W, - W. )% m = —-L- X 100

(w2 - )

■ from w h ich an e s t i m a t e d r y w e i g h t o f t h e w h o le s a m p le was

o b t a i n e d .

V a r i o u s d i f f i c u l t i e s w ere e n c o u n t e r e d w i t h t h e

27

c o l l e c t i o n o f g uan o from B a tu C a v e s . On s e v e r a l o c c a s i o n s

t h e t r a y s w e re t o r n from t h e wooden f r a m e s when l a r g e t o a d s

( B ufo a s p e r G r a v e n h . ) o r f e r a l c a t s ( b o t h o f w h ic h i n h a b i t

t h e c a v e ) jum ped o n to th e m . O c c a s i o n a l l y i n w e t w e a t h e r t h e

a c c u m u l a t i o n o f g ua n o and w a t e r d r i p from t h e r o o f b u r s t t h e

s h e e t s , o r e l s e t h e e x c e s s w a t e r o v e r f l o w e d . (When t h e s h e e t s

w ere f l o o d e d w i t h w a t e r t h e g u a n o , b e i n g more d e n s e t h a n w a t e r ,

s a n k t o t h e b o t t o m o f t h e p o o l t h a t h ad a c c u m u l a t e d and v/as

n o t l o s t i f e x c e s s w a t e r o v e r f l o w e d . ) D u r in g w e t w e a t h e r d r i p s

from t h e r o o f s p l a s h e d c a v e s o i l ( c o n t a i n i n g p o l l e n o f unknown

a g e ) from t h e f l o o r o n to t h e s a m p l e . C o n ta m in a te d s a m p le s

w e re d i s c a r d e d . To r e d u c e s p l a s h , t h e o ld p o l y t h e n e s h e e t s

w e re p l a c e d on t h e f l o o r u n d e r t h e d r i p s .

E ach week t h e r e l a t i v e a b u n d a n c e o f E . s p e l a e a

i n B a tu C aves was r e c o r d e d t o g e t h e r w i t h a n o t e o f t h e am oun t

o f f r e e w a t e r e n t e r i n g t h e c a v e . E . s p e l a e a was c a t e g o r i s e d

a s ( a ) " v e r y a b u n d a n t" (V A ) , ( b ) " a b u n d a n t " ( A ) o r ( c ) I ' . le ss

a b u n d a n t " ( L A ) . T h ese c a t e g o r i e s a r e s u b j e c t i v e , i n d i c a t i n g

c h a n g e s i n t h e num b ers o f b a t s p r e s e n t , n b t a b s o l u t e num bers

o f b a t s p r e s e n t . The c a v e was r e c o r d e d a s ( a ) " v e r y w e t

v/hen a p o o l i n C a v e rn C w as f u l l o f v / a t e r . T h i s p o o l f i l l e d

o n l y a f t e r a week d u r i n g v /h ich h e a v y r a i n f e l l a l m o s t d a i l y .

F o l l o w i n g d r y w e a t h e r , i t d r a i n e d i n a few d a y s . (The w a t e r

e n t e r e d t h e ;c a v e b y way o f c r a c k s i n t h e r o o f f rom w h ich i t

d r i p p e d o r r a n a s s m a l l s t r e a m s down t h e w a l l s . ) (b ) " w e t"

when w a t e r f l o w e d o v e r t h e f lo w s t o n e i n C a v e rn B . V /a ter

f lo w e d o v e r t h i s , . c o l u m n a f t e r a few d a y s o f i n t e r m i t t e n t

h e a v y r a i n when w a t e r a l s o d r i p p e d from s e v e r a l p a r t s o f t h e

/

■/

28

r o o f i n C av e rn C o r ( c ) " d r y " when t h e r e was no v / a t e r on t h e

f lo w s t o n e . "D ry" i s u s e d a s a r e l a t i v e , n o t a n a b s o l u t e

t e r r a .

I n Gua S a n d i n g , c o l l e c t i o n s w e re made once a

m o n th . I t was n o t p o s s i b l e t o l e a v e e q u ip m e n t i n t h e c a v e

b e c a u s e o f v a n d a l i s m and s o e a c h sa m p le v/as made up from tv/o

h u n d re d f r e s h f a e c a l p e l l e t s c o l l e c t e d from t h e c a v e f l o o r

w i t h f o r c e p s . On t h e f i r s t o c c a s i o n , 400 p e l l e t s w e re c o l l e c t e d

i n two g r o u p s o f 200 e a c h and a n a l y s e d s e p a r a t e l y i n o r d e r

t o c h e c k t h e a d e q u a c y o f t h e s a m p le s i z e . (The f l o o r o f t h e

c a v e was r e l a t i v e l y c l e a n a s v e g e t a b l e g a r d e n e r s f r e q u e n t l y

s w e p t up t h e gu ano f o r f e r t i l i s e r . The c a v e was s h a l l o w and

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

f a e c e s d e p o s i t e d on t h e r o c k y f l o o r . ) F a e c a l p e l l e t s v/ere

c o n s i d e r e d f r e s h i f t h e i r s u r f a c e was s t i l l m o i s t . S l i d e s

w e re p r e p a r e d e x a c t l y a s t h o s e o f guan o from E a t u C aves e x c e p t

t h a t t h e w h o le s a m p le and n o t a s u b s a m p le was t r e a t e d w i t h

KOH.

C o l l e c t i o n o f s u b f o s s i l gu ano d e p o s i t s .

I n t h e ho p e o f f i n d i n g p o l l e n p r e s e r v e d i n guano

t h a t ha d b e e n d e p o s i t e d b e f o r e man a l t e r e d t h e e n v i r o n m e n t ,

c o r e s a m p le s w e re c u t from c a v e s o i l d e p o s i t s i n B a tu C aves

and Gua S a n d i n g . I n a l m o s t a l l c a v e s i n P e n i n s u l a r M a l a y s i a

m i n e r s h a v e rem o ved o r s e r i o u s l y d e p l e t e d and d i s t u r b e d ca v e

s o i l d e p o s i t s f o r f e r t i l i s e r ( P e a c o c k , 1 9 6 5 ) . The o n l y r e m a i n i n g

d e p o s i t i n Gua S a n d in g fo rm s t h e b ed o f a p o o l i n t h e b a s e

o f t h e c a v e . I n B a tu C aves t h e o n l y d e p o s i t n e v e r m ined

29

i s i n t h e P e n n y Room (M c C lu re , Lira and W inn, 1 9 6 7 , p . A 0 7 ) .

T h e s e , t h e r e f o r e , w e re t h e s i t e s from w h ich c o r e s w ere c o l l e c t e d .

The c o r e r w h ich v/as o f a s t a n d a r d d e s i g n w i t h

an i n t e r n a l l e n g t h 50 cm and an i n t e r n a l d i a m e t e r o f A ♦2cm

was l o a n e d b y t h e G e o g ra p h y D e p a r t m e n t , The U n i v e r s i t y , H u l l ,

w h e re i t was d e s i g n e d a n d c o n s t r u c t e d . C o re s w e re c u t f rom

th e s u r f a c e o f t h e d e p o s i t t o t h e b e d r o c k . E ach c o r e s e c t i o n

was c u t f ro m a s l i g h t l y d i f f e r e n t p o i n t s i n c e a n y s o i l f a l l i n g

i n t o t h e h o l e from w h ich a s e c t i o n . h a d a l r e a d y b e e n rem oved

w o u ld c o n t a m i n a t e t h e n e x t o n e . E ach s e c t i o n was e j e c t e d

from t h e c o r e r o n t o .a c l e a n p o l y t h e n e s h e e t i n w h ich i t was

w r a p p e d . The s e c t i o n number and t h e d e p t h from w h ich i t came

w e re w r i t t e n on t h e p o l y t h e n e w r a p p in g w i t h a f e l t t i p pen

c o n t a i n i n g w a t e r p r o o f i n k . The p r o x i m a l and d i s t a l e n d s w e re

a l s o m arked a p p r o p r i a t e l y .

I n t h e l a b o r a t o r y , n o t e s w ere made on t h e c o l o u r

and t h e t e x t u r e o f t h e c o r e s and s a m p le s w e re t a k e n e v e r y two

i n c h e s f o r p o l l e n a n a l y s i s . The s a m p le s w e re s t a i n e d i n b a s i c

f u s c h i n i n P .V .A . Some w ere a l s o t r e a t e d w i t h d i l u t e h y d r o c h l o r i c

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

A p r o f i l e o f t h e B a tu C aves d e p o s i t from t h e

s u r f a c e t o b e d r o c k was c u t i n t h e s i d e o f a f l o o d v /a t e r

c h a n n e l i n t h e P en n y Room. S a m p le s w ere c o l l e c t e d e v e r y two

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

d e p o s i t .

P o l l e n a n a l y s i s .

A l l p o l l e n s a m p le s w ere e xam ined a t 10 x AO

30

m a g n i f i c a t i o n t h r o u g h a m i c r o s c o p e f i t t e d w i t h an a d j u s t a b l e

b u i l t i n l i g h t s o u r c e , a b i n o c u l a r h e a d an d a m o b i l e s t a g e .

500 p o l l e n g r a i n s f rom e a c h s a m p le w ere i d e n t i f i e d a s f a r a s

p o s s i b l e and c o u n t e d by t r a v e r s i n g t h e s l i d e from s i d e t o

s i d e a t ra n d o m ly ; c h o s e n p o i n t s and c o u n t i n g e a c h g r a i n t h a t

p a s s e d t h r o u g h t h e f i e l d o f v iew u n t i l t h e r e q u i r e d num ber

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

w e re r e q u i r e d t o o b t a i n 500 g r a i n s . The p o s i t i o n o f e a c h

t r a n s e c t was d e t e r m i n e d from t h e r e a d i n g o f t h e v e r n i e r s c a l e

on t h e m o b i l e s t a g e and no t r a n s e c t was c o u n t e d t w i c e . G r a i n s

l y i n g p a r t l y o u t o f t h e f i e l d o f v iew w ere c o u n t e d i f h a l f

o r more o f t h e g r a i n was v i s i b l e . The d e c i s i o n t o i n c l u d e o r

e x c l u d e s u c h g r a i n s was n e c e s s a i l y som ewhat s u b j e c t i v e s i n c e

i t was s o m e t im e s d i f f i c u l t t o d e c i d e w h a t c o n s t i t u t e d h a l f

a g r a i n ( e s p e c i a l l y i n g r a i n s s u c h a s Bo mb ax w h ic h a r e t r i a n g u l a r ) .

I n c l u d i n g o r e x c l u d i n g a l l g r a i n s t h a t w ere in i ly p a r t l y i n t h e

f i e l d o f . v i e w w o u ld h a v e i n t r o d u c e d a f a r more s e r i o u s b i a s

t h a n t h e m ethod a d o p t e d s i n c e a h i g h e r p r o p o r t i o n o f l a r g e

g r a i n s t h a n o f s m a l l g r a i n s a r e p a r t l y o u t o f v i e w . O c c a s i o n a l l y

s m a l l c lu m p s o f p o l l e n g r a i n s w e re e n c o u n t e r e d e . g . i n an

a n t h e r . T h ese w e re r a r e and t h e y w ere n o t i n c l u d e d i n t h e

c o u n t .

P o l l e n i d e n t i f i c a t i o n .

M ost p o l l e n ' t y p e s r e c o v e r e d ' f r o m t h e b a t s o r

t h e i r gu ano w e re i d e n t i f i e d b y c o m p a r i s o n w i t h p o l l e n s a m p le s

v c o l l e c t e d f rom p l a n t s known o r s u s p e c t e d t o b e c h i r o p t e r o p h i l o u s .

M u l l e r (1 9 6 9 ) h a s p u b l i s h e d an e x c e l l e n t a c c o u n t o f t h e p o l l e n

II

31

o f S o n n e r a t i a h u t t h e r e a r e few o t h e r c o m p r e h e n s i v e s t u d i e s

o f c h i r o p t e r o p h i l o u s g e n e r a . A l th o u g h t h e p o l l e n o f S o n n e r a t i a

a l b a c o u l d ;.b e d i s t i n g u i s h e d from t h a t o f S . o v a t a B a c k e r ,

t h e l a t t e r was v e ry : uncommon i n t h e s a m p le s c o l l e c t e d d u r i n g

t h i s s t u d y . They w e r e , t h e r e f o r e , n o t d i s t i n g u i s h e d i n t h e

p o l l e n a n a l y s e s and b o t h a r e i n c l u d e d u n d e r S . a l b a t h r o u g h ­

o u t t h e t h e s i s . S a m p le s o f unknown p o l l e n w e re s e n t t o v a r i o u s

p a l y n o l o g i s t s f o r e x a m i n a t i o n . Each unknown p o l l e n t y p e was

a l l o c a t e d a r e f e r e n c e number and s k e t c h e d . The s a m p le i n w h ic h

i t was f i r s t fo u n d becam e t h e " t y p e s p e c im e n " f o r t h a t p o l l e n

t y p e . Any i d e n t i f i a b l e o b j e c t s b e s i d e s p o l l e n g r a i n s w ere

a l s o r e c o r d e d .

C o n f id e n c e l i m i t s f o r t h e p o l l e n c o u n t s w e re

e s t i m a t e d from nomograms d e v i s e d b y Maher ( 1 9 7 2 ) . T h e i r a c c u r a c y

was a l s o t e s t e d b y two e x p e r i m e n t s (A p p e n d ix 2 ) .

1 . S l i d e s w e re p r e p a r e d from e a c h o f t h r e e d i f f e r e n t s u b ­

s a m p le s o f B a tu Caves s a m p le # F 6 , 1000+ g r a i n s c o u n te d

on e a c h and t h e r e s u l t s w e re c o m p a r e d .

2 . Two c o u n t s o f 500+ g r a i n s w ere made on one s l i d e from

Gua S a n d i n g ( c o l l e c t e d 1^ May 1972) and s e v e n s l i d e s

from B a tu C aves (# F 7 , F 8 , F 9 , F 1 0 , Fh9 , F53 and F57)

an d t h e r e s u l t s c o m p a re d .

R e s u l t s o f t h e s e e x p e r i m e n t s show t h a t t h e r e w as no s i g n i f i c a n t

d i f f e r e n c e ( p > 0 , 0 5 ) b e t w e e n t h r e e i s l i d e s p r e p a r e d i n d e p e n d e n t l y

from one s a m p le an d t h a t w h e re two c o u n t s w e re made from one

s l i d e t h e r e s u l t s o f e a c h c o u n t w e re v e r y s i m i l a r ( n o n e w ere

s i g n i f i c a n t l y d i f f e r e n t a t t h e p = 0 . 0 5 l e v e l an d a l l w ere w i t h ­

i n t h e 95% c o n f i d e n c e l i m i t s com p u ted from M a h e r ' s nomograms

(M a h e r , 1 9 7 2 ) . The m e th o d s u s e d t o p r e p a r e t h e s l i d e s and

c o u n t t h e p o l l e n p r e s e n t on them w e re t h e r e f o r e a ssu m ed t o

a d e q u a t e .

33

RESULTS.

CHIROPTEROPHILOUS PLANTS,.

I n P e n i n s u l a r M a l a y s i a , E o n y c t e r i s s p e l a e a f e d

from a l l p l a n t s from w h ic h b o t h s p e c i e s o f M a c r o g l o s s u s w e re

r e c o r d e d f e e d i n g , b e s i d e s many o t h e r s . T h e r e f o r e , T a b le 33

l i s t s a l l t h e c h i r o p t e r o p h i l o u s p l a n t s r e c o r d e d d u r i n g t h i s

s t u d y i n P e n i n s u l a r M a l a y s i a .

A l l t h e o b s e r v a t i o n s made on t h e s e s p e c i e s t h a t

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

a c c o u n t s a l r e a d y e x i s t s . . A l l d a t a r e f e r s t o P e n i n s u l a r M a l a y s i a

u n l e s s o t h e r w i s e s t a t e d .

Many o f t h e p l a n t s w e re c u l t i v a t e d s p e c i e s w h i l e

s e v e r a l w i l d s p e c i e s p a r t l y owed t h e i r a b u n d a n c e t o man, e i t h e r

t h r o u g h h i s a c t i v e p r o t e c t i o n ( e . g . P a r k i a s p e c i o s a ) o r t h r o u g h

h i s c r e a t i n g s u i t a b l e h a b i t a t s ( e . g . w i l d Musa s p p . ) . The

r e m a i n d e r w e re m ang ro ve o r p r i m a r y f o r e s t t r e e s . The m a j o r i t y

grew i n t h e l o w l a n d s b e lo w 1000 m ( e . g . T a b le 1 ) .

MONOCOTYLEDONS.

1 .. . .M usaceae .

One g e n u s o f M u s a c e a e , M usa, ( t h e b a n a n a s ) g row s

i n P e n i n s u l a r M a l a y s i a . Two w i l d s p e c i e s , M. t r u n c a t a and

M* r o a l a c c e n s l s ( b o t h s e c t i o n Eumusa) w ere c h i r o p t e r o p h i l o u s

an d c u l t i v a t e d b a n a n a s w e re r e g u l a r l y v i s i t e d b y b a t s f o r

n e c t a r b u t s i n c e t h e y a r e p a r t h e n o c a r p i c and r e l e a s e no

p o l l e n t h e y c an h a r d l y b e t e r m e d c h i r o p t e r o p h i l o u s ! C o m p a ra t iv e

o b s e r v a t i o n s w e re a l s o made on one o r n i t h o p h i l o u s s p e c i e s ,

v l o l a s c e n s ( s e c t i o n C a l l i m u s a ) .

T a b le 1 .

T a b le s h o w in g t h e num ber o f t r e e s o f f i v e g e n e r a

c o u n t e d w i t h i n a b o u t 50 m o f t h e r o a d b e tw e e n e a c h m i l e p o s t

and t h e a l t i t u d e r e c o r d e d a t e a c h m i l e p o s t from T anah R a ta

(P a h a n g ) t o T apah ( P e r a k ) .

D uab. = D uabanga g r a n d ! f l o r a

O ro x . = O roxylum in d i c u m

Bomb. = Bombax v a l e t o n i i

P a r k . = P a r k i a s p p .

B u r . = D u r io s p p .

M iles from Tapah

A l t i t u d e i n ra a . s . l .

No . o f t r e e s p r e s e n t •

Comments.Duab. O rox . Tomb. F a rk . D ur .

38-37 1433-1372 F o r e s t c l e a r e d

37-36 1372- 736-35 ? -11 73 1

35-34 1173-1097 17 1. *7 *7 i O

C -

33-32 1097 2

32-31 1097 *' R e s e r v o i r , f l a t

0K'l1 1097 530-29 1097 R in g le t town29-28 1097-1128 F o r e s t c l e a r e d28-27 1128-1074 1827-26 1 0 7 4 -1 0 0 6 4726-25 1006-945 3625-24 945-892 8

24-25 892-81 5 132 3 -2 2 81 5 -7 6 2 30 A l t . s ig n 2200*22-21 762-71 6 8 10 4 121 -2 0 716-685 13 320-19 685-632 7 3 11 9 -1 8 632-594 5 2 7 4 1

18-17 594-541 1 6 4 p a r t l y c l e a r e d1 7 -1 6 541 -533 4 1 3 13 15

16-15 533-495 4 13 11 12

15-14 495-450 4 3 1 20 10

14-13 450-404 4 11 1 5 6

13-12 404-396 * 5 18 212-11 396- 8 13 6

1 1 -1 0 396-351 1 7 2

10-9 351 -305 1 21 13 P a . la n e l ia p r e s e n t9-8 305-267 19 2 11 n

8-7 267-229 1 1 20 13

34

(T he taxonom y o f Musa i s v e r y c o m p l i c a t e d , ,

t r u n c a t a and M. m a l a c c e n s i s b e l o n g t o t h e M. a c u m i n a t a

C o l l a s p e c i e s c o m p le x . Cheesman (1 9 4 8 ) r e d u c e d them to

v a r i e t i e s o f M, a c u m i n a t a and Simmonds (19 5 5 ) r e g a r d e d them

s a s s u b s p e c i e s o f M. a c u m i n a t a . F o l l o w i n g Chen (1 9 7 2 ) t h e y

a r e t r e a t e d a s f u l l s p e c i e s h e r e . C u l t i v a t e d b a n a n a s h a v e

b e e n d e r i v e d from M. a c u m i n a t a s e n s u l a t o o r h y b r i d s b e tw e e n

i t and M. b a l b i s i a h a C o l l a (C h e esm an , 1 947 ; S im m onds, 1955>

1 9 6 2 ) . I n t h e p a s t t h e r e h a s b e e n c o n s i d e r a b l e c o n f u s i o n o v e r

t h e i r n o m e n c l a t u r e an d M. p a r a d i s l a c a L. o r M. s a p l e n t l u m L.

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

a r e n o t a d i s t i n c t s p e c i e s i n t h e b i o l o g i c a l s e n s e and i t i s

t h e r e f o r e i n c o r r e c t t o r e f e r t o them b 3r t h e s e nam es ( Cheesmamt,1 947) •

E c o l o g i c a l d i s t r i b u t i o n .

Musa m a l a c c e n s i s was a b u n d a n t i n s u i t a b l e h a b i t a t s

t h r o u g h o u t t h e l o w la n d s o f S e l a n g o r , P e r a k and w e s t e r n P a h an g

up t o 200 t o 3 0 0 m a . s . l . ; i t was n o t lo o k e d f o r i n o t h e r

s t a t e s . Above 500 m a . s . l . on Gunong U lu K a l i ( S e l a n g o r ) , a t

F r a s e r ' s H i l l ( P a h a n g ) , i n t h e Cameron H i g h l a n d s ^ ( P e r a k and

P ah an g ) and p r o b a b l y e l s e w h e r e i t was r e p l a c e d b y M. t r u n c a t a

b u t b e tw e e n 300 m and 500 m a . s . l . n a t u r a l h y b r i d s o f t h e s e

two s p e c i e s p r e d o m i n a t e d . H y b r i d s so m e t im e s e x t e n d e d down

t o 200 m a . s . l . an d up t o 600 m a . s . l . C u l t i v a t e d b a n a n a s

w e re p l a n t e d a r o u n d m o s t r u r a l d w e l l i n g s ( a n d many .u rb a n

o n e s to o ! ) i n t h e l o w l a n d s b u t t h e y w ere grown l e s s a t h i g h e r

a l t i t u d e s . M. v i o l a s c e n s was fo u n d i n a l l s u i t a b l e h a b i t a t s

35

and a t a l l a l t i t u d e s up t o 2000 m a . s . l . on Gunong B r i n c h a n g .

B a n a n a s r e q u i r e h i g h l i g h t i n t e n s i t y , a b u n d a n t '

w a t e r and w e l l d r a i n e d s o i l . N a t u r a l h a b i t a t s o f w i l d s p e c i e s ,

w h ich i n c l u d e d r e c e n t l a n d s l i p s and r i v e r b a n k s ( p a r t i c u l a r l y

w h ere f l o o d i n g c l e a r e d t h e b a n k s o f o t h e r v e g e t a t i o n ) a r e

r a r e ; h o w e v e r , man h a s c r e a t e d i n n u m e r a b l e s i t u a t i o n s i n w h ich

w i l d b a n a n a s p r o l i f e r a t e , e . g . a l o n g w a t e r c o u r s e s i n c u l t i v a t e d

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

r o a d m a r g in s a nd on t h e f o r e s t edg e w h e r e v e r c l e a r i n g s h a v e

b e e n made ( F i g . 1 1 ) . Musa v i o l a s c e n s was u s u a l l y more s h a d e

t o l e r a n t t h a n M. m a l a c c e n s i s o r M. t r u n c a t a b u t o t h e r w i s e

showed l i t t l e d i f f e r e n c e i n i t s c h o i c e o f h a b i t a t .

M o rp h o lo g y ( F i g . 3 9 ) .

The g r o s s m o rp h o lo g y o f b a n a n a s h a s b e e n d e s c r i b e d

by s e v e r a l a u t h o r s ( e . g . C heesm an , 19^-7; S im m onds, 1955) They

a r e g i g a n t i c , r h i z o m a t o u s p l a n t s w i t h e n t i r e l e a v e s an d lo n g

p e t i o l e s , t h e b a s a l p o r t i o n s o v e r l a p p i n g t o fo rm a p s e u d o s te m

up t o k m h i g h i n M. t r u n c a t a b u t so m e t im e s h i g h e r , u p t o 5 m,

•*-n JI* m a l a c c e n s i s an d c u l t i v a t e d b a n a n a s . The p s e u d o s t e m s o f

v i o l a s c e n s se ld o m e x c e e d e d 3 m i n h e i g h t . The a b a x i a l

s u r f a c e s o f t h e l e a v e s o f M. m a l a c c e n s i s and c u l t i v a t e d

b a n a n a s a r e c o v e r e d i n a f i n e , w h i t e , waxy pow der w h ic h

g i v e s them a s i l v e r y a p p e a r n a c e .

The i n f l o r e s c e n c e i s t e r m i n a l on a t r u e s te m

w h ich i s f o r c e d up t h e c e n t r e o f t h e p s e u d o s t e m . The r a c h i s

r e m a i n s v e r t i c a l i n M. v i o l a s c e n s ( F i g . 11) b u t i n M. t r u n c a t a ,

M. m a l a c c e n s i s an d c u l t i v a t e d b a n a n a s i t f l e x e s t o g i v e a

F ig . 11. Musa spp.

a . A grove of wild bananas, Musa malaccensis. on recently cleared forest land, Selangor.

Musa violascens Musa borneensisb. Both ornithophilous species of Musa, section Callimusa.

'-Vi.-

'

p e n d u l o u s o r s e m i “p e n d u l o u s i n f l o r e s c e n c e h a n g i n g b e lo w t h e

free p o r t i o n o f t h e p e t i o l e s ( F i g s . 12 an d 13).As i n a l l b a n a n a s , t h e f l o w e r s a r e a r r a n g e d i n

h a n d s . The f l o w e r s i n t h e b a s a l h a n d s a r e f u n c t i o n a l l y f e m a l e ,

t h e r e m a i n d e r a r e f u n c t i o n a l l y m a l e . Each h a n d i s s u b t e n d e d b y

a l a r g e , b r i g h t l y c o l o u r e d b r a c t . The b r a c t c o l o u r v a r i e s frum

d e e p b l a c k i s h - p u r p l e on t h e o u t e r ( a b a x i a l ) s u r f a c e a n d w h i t e

on t h e i n n e r ( a d a x i a l ) s u r f a c e i n M. t r u n c a t a ( F i g . 13) t o

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

( F i g . 13) to b r i c k r e d on b o t h s u r f a c e s i n M. m a l a c c e n s i s

( F i g . 1 2 ) . ( I n one g r o v e o f M. m a l a c c e n s i s n e a r B e n to n g ,

P a h a n g , t h e b r a c t s o f a l l t h e f l o w e r i n g p l a n t s w e re s t r e a k e d

y e l l o w on t h e a b a x i a l s u r f a c e ( F i g . 1 2 ) . T h i s fo rm was d e s c r i b e d

a s M. f l a v a R i d l e y b u t i s now r e g a r d e d a s a v a r i e t y o f

M. m a l a c c e n s i s (S im m onds, 1935). The b r a c t s o f M. v i o l a s c e n s

a r e w h i t e a t t h e b a s e g r a d i n g i n t o l i l a c a t t h e a p e x on b o t h

s u r f a c e s b u t t h e i n n e r ( a d a x i a l ) s u r f a c e i s p a l e r a t t h e a p e x .

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

a r e c o v e r e d i n a w h i t e , waxy pow der o r b loom on t h e a b a x i a l

s u r f a c e . T h e re i s no n e on M. m a l a c c e n s i s o r M. v i o l a s c e n s .

The m o r p h o l o g i c a l c h a r a c t e r s o f h y b r i d s a r e

m o s t l y i n t e r m e d i a t e b e tw e e n t h e p a r e n t s p e c i e s (C h e n , 197?-; Simmonds, 1 9 55 ; p e r s , o b s . ) e x c e p t f o r t h e c o l o u r o f t h e

a d a x i a l s u r f a c e o f t h e b r a c t o f t h e h y b r i d s i n v o l v i n g M. t r u n c a t a

Above 600 m a . s . l . , w h e re o n ly M. t r u n c a t a g r o w s , t h e a d a x i a l

s u r f a c e s o f i t s b r a c t s a r e w h i t e . H ow ever , b e tw e e n 300 m and

600 m a . s . l . , w h e re h y b r i d i s a t i o n w i t h M. m a l a c c e n s i s o c c u r s ,

many p l a n t s w h ic h a r e v e r y s i m i l a r t o M. t r u n c a t a i n a l l

F i g . 12. Musa spp.

a . Musa m a la c c e n s is pho to g rap h ed i n m id-m orning . N o te : - 1 . The s c r a t c h marks made by th e f e e t o f

th e b a t s .2 . The p u n c tu re marks made by b i r d c law s .3 . The m elaponid b e e .

b . Musa m a la c c e n s is v a r f l a v a . B en tong , Pahang.

F ig . 13* Musa spp.

a. Musa spp. cultivated. Two openbracts are unusual.(From Grounds, S.R.,1974)

b. Musa truncata photographed at about 07.00 hrs. Note the white adaxial and very dark abaxial surfaces of the bracts.

o t h e r r e s p e c t s h a v e t h i s s u r f a c e r e d . No i n t e r m e d i a t e c o l o u r s

w ere fo u n d and j . t i s a ssu m ed t h a t t h e w h i t e a d a x i a l b r a c t

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

p l a n t s w i t h a r e d a d a x i a l s u r f a c e a r e d e r i v e d f rom h y b r i d

a n c e s t o r s , (C a re m u s t b e t a k e n when r e c o r d i n g t h e b r a c t c o l o u r

from f a l l e n b r a c t s a s t h e w h i t e a d a x i a l s u r f a c e o f M. t r u n c a t a

b r a c t s may t u r n r e d a f t e r f a l l i n g . )

There is little variation in the structure 6f the flowers of these species of bananas and the general descriptions of Musa flowers given by Cheesman (1947) and

Simmonds (1955) are adequate (see also Fig. 14)* However, there is variation in the orientation, the colour and the size of the flowers of the different species.

As a r e s u l t o f t h e o r i e n t a t i o n o f t h e w h o le

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

t h o s e o f t h e o t h e r s p e c i e s d o w nw ards . The f l o w e r s o f M. t r u n c a t a

a r e w h i t e , w i t h p a l e y e l l o w t e r m i n a l l o b e s t o t h e compound

t e p a l . T ho se o f M. m a l a c c e n s i s an d c u l t i v a t e d b a n a n a s a r e

p a l e y e l l o w w i t h b r i g h t e r l o b e s and t h o s e o f M. v i o l a s c e n s

s l i g h t l y d a r k e r b u t w i t h v e r y b r i g h t y e l l o w l o b e s , t h e c o l o u r

e x t e n d i n g o n to t h e f u s e d , d i s t a l p a r t o f t h e compound t e p a l .

The compound t e p a l s o f t h e m a le f l o w e r s o f

M* M a l a c c e n s i s an d M. t r u n c a t a a r e a b o u t 30 mm l o n g . I n

c u l t i v a t e d b a n a n a s t h e y v a r y from one c u l t i v a r t o a n o t h e r .

b u t a r e u s u a l l y l o n g e r , up t o 50 mm, and i n M. v i o l a s c e n s

t h e y a r e s h o r t e r , a b o u t 25 mm. The f r e e t e p a l i s a b o u t h a l f

t h e l e n g t h o f t h e compound t e p a l i n a l l e x c e p t M. v i o l a s c e n s

Fig. 14. A) D I A G R A M S O F M U S A SPP. FLOWERS TO S H O W

THE M O V E M E N T OF NECTAR W H E N THE TFPALS

ARE PARTED X 2

n e c t a r

B) DI AGRAM TO S H O W POS I TI ON O F THE BRACT O F MUS A TRUNCATA DUR I N G O N E NI G HT

17-00 18 00 hrs

20 00 hrs

38

i n w h ic h i t i s b e tw e e n two t h i r d s and t h r e e q u a r t e r s t h e l e n g t h

o f t h e compound t e p a l . I n a l l o f them t h e l e n g t h o f t h e f l o w e r s

d e c r e a s e s t o w a r d s t h e a p e x o f t h e i n f l o r e s c e n c e , ,

The l e n g t h o f t h e t e p a l s i n f e m a le f l o w e r s i s

a p p r o x i m a t e l y e q u a l t o o r s l i g h t l y l o n g e r t h a n t h o s e o f t h e

m a l e s ! H o w ev er , t h e fo r m e r a r e a s much a s t w i c e a s b r o a d a s

t h e l a t t e r i n a l l s p e c i e s . The r e c e p t a c l e on t h e w e l l d e v e lo p e d

i n f e r i o r o v a r y o f f e m a le f l o w e r s i s a l s o much l o n g e r t h a n

t h a t o f m a le f l o w e r s . (The o v a r y o f m a le f l o w e r s i s non

f u n c t i o n a l an d c o n s i d e r a b l y s m a l l e r t h a n t h a t o f f e m a le f l o w e r s . )

N e c t a r .

N e c t a r i s s e c r e t e d from t h e n e c t a r i e s i n t h e

r e c e p t a c l e . I n t h e s p e c i e s w i t h a p e n d u lo u s o r s e m i - p e n d u l o u s

i n f l o r e s c e n c e i t i s r e t a i n e d i n t h e p r o x i m a l p o r t i o n o f t h e

c o r o l l a t u b e b y s u r f a c e t e n s i o n ( F i g . 1 i f ) . H o w e v e r , i f t h e

compound and f r e e t e p a l s a r e f o r c e d a p a r t i t r u n s down t h e

f r e e t e p a l f o r m in g a l a r g e d r o p a t i t s a p e x ( F i g . 1 4 ) .

The q u a n t i t y o f n e c t a r e x t r a c t e d from f l o w e r s

v a r i e d f rom 0 . 1 5 ml ( 0 . 1 0 t o 0 . 4 9 , n = 1 1 5 ) i n c u l t i v a t e d

b a n a n a s t o 0 .1 1 ml ( 0 . 0 7 t o 0 . 2 0 , n = 1 1 ) i n M. t r u n c a t a .

T h u s , one h a n d o f c u l t i v a t e d b a n a n a f l o w e r s c o n t a i n e d a b o u t

2 . 4 ml o f n e c t a r and t h o s e o f M. t r u n c a t a and M. m a l a c c e n s i s

a b o u t 1 . 8 m l . ( T h e s e s p e c i e s o f Musa n o r m a l l y h a d 1 6 f l o w e r s ,

a r r a n g e d i n two row s o f e i g h t p e r h a n d . ) The f l o w e r s may

c o n t i n u e t o s e c r e t e h e c t a r t h r o u g h o u t p a r t o r a l l o f t h e

n i g h t b u t no m e a s u re m e n ts w e re made t o c o n f i r m t h i s . I n

f l o w e i ' s t h a t w ere n o t v i s i t e d b y b a t s ( e . g . b e c a u s e a m i s t

39

n e t was s e t b e s i d e them ) , e x c e s s n e c t a r d ro p p e d from t h e

f l o w e r s a n d r a n down t h e u n d e r l y i n g b r a c t a f t e r a b o u t 2 2 <,00

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

t h e n . No m e a s u re m e n ts o f t h e n e c t a r p ro d u c e d by M. v i o l a s c e n s

w e re made.

No a n a l y s i s o f t h e c o m p o s i t i o n o f t h e n e c t a r

was made e x c e p t t h a t t h e d r y w e i g h t o f 1 .1 ml c o l l e c t e d from

11 f l o w e r s o f c u l t i v a t e d b a n a n a s was 2 1 . 8 % o f t h e t o t a l w e i g h t .

P o l l e n .

D e s c r i b e d an d f i g u r e d f o r M u saceae b y E rd tm an

( 1 9 5 2 ) . A s t e n o p a l y n o u s f a m i l y , t h e s p e c i e s c o u l d n o t b e

s e p a r a t e d p a l y n o l o g i c a l l y . P o l l e n was r e l e a s e d b y t h e a n t h e r s

o f m a le f l o w e r s o f a l l s p e c i e s e x c e p t c u l t i v a t e d b a n a n a s

w h ich w e re o n l y v e r y r a r e l y fo u n d t o r e l e a s e p o l l e n . The

p o l l e n g r a i n s a r e s p h e r i c a l w i t h a v e r y t h i n e x i n e ( F i g . ) .

The d i a m e t e r o f 201 M. t r u n c a t a g r a i n s v a r i e d f rom 51 p t o

90 ju (mean 73 p + 0 , 9 p ) • The d i a m e t e r o f M. m a l a c c e n s i s

p o l l e n was n o t m e a s u re d b u t was o f a s i m i l a r s i z e .

A s t i c k y s u b s t a n c e fo rm ed an i r r e g u l a r l a y e r

on t h e o u t e r s u r f a c e o f t h e g r a i n s c a u s i n g them t o a d h e r e

t o e a c h o t h e r and t o a n y t h i n g t h e y came i n c o n t a c t w i t h .

S c e n t .

B an an a f l o w e r s h a v e a d i s t i n c t i v e m u s ty s m e l l .

I t i s n o t s t r o n g t o t h e human n o s e .

F l o w e r i n g b i o l o g y .

One b r a c t on e v e r y M. t r u n c a t a i n f l o r e s c e n c e

i n th e . m a le s t a g e b e g i n s t o r i s e a t d u s k e a c h n i g h t . W i t h i n

9 0 m i n u t e s i t h a s o p en ed enough t o a l l o w b a t s t o l a n d and

f e e d from th e n e w ly e x p o s e d f l o w e r s . The b r a c t c o n t i n u e s

r i s i n g thw ugh a b o u t 1 0 0 ° t o i 1 0 ° when t h e t i p b e g i n s t o r e c u r v e ,

and t h e n i t r o l l s up s l o w l y u n t i l b y dawn a l m o s t t h e w h o le

b r a c t h a s r o l l e d up ( F i g . 1 *+) . T h e re a r e 110 o b v i o u s c h a n g e s

i n t h e f l o w e r s t h e m s e l v e s f rom t h e o p e n in g o f t h e b r a c t u n t i l

dawn ( e x c e p t t h e r e l e a s e o f p o l l e n and t h e s e c r e t i o n o f n e c t a r ) .

By m id -d a y on t h e day f o l l o w i n g a n t h e s i s b o t h t h e b r a c t s and

t h e f l o w e r s h a v e f a l l e n o f f . O c c a s i o n a l l y b i r d s k n o c k them

o f f ( s e e b e l o w ) .

The same s e q u e n c e was o b s e r v e d f o r M, m a l a c c e n s i s

an d c u l t i v a t e d b a n a n a s e x c e p t t h a t t h e i r b r a c t s d i d n o t a lw a y s

r i s e a t d u s k , p a r t i c u l a r l y i n M. m a l a c c e n s i s i n w h ic h t h e

b r a c t s o f many p l a n t s b e g a n t o open i n t h e e a r l y m o r n in g .

The b r a c t s and f l o w e r s o f M. m a l a c c e n s i s o f t e n r e m a in on t h e

p l a n t f o r c o n s i d e r a b l y l o n g e r t h a n 12 h o u r s b u t s t i l l a new

o ne o p en ed e v e r y 2i|- h o u r s . Two open b r a c t s on one i n f l o r e s c e n c e

w ere v e r y o c c a s i o n a l l y s e e n on c u l t i v a t e d b a n a n a s , b u t n e v e r

on w i l d s p e c i e s .

A . s c a r r e m a i n s on t h e r a c h i s a f t e r e a c h h an d o f

f l o w e r s and i t s s u b t e n d i n g b r a c t f a l l o f f . The num ber o f d a y s

f o r w h ic h a n i n f l o r e s c e n c e h a s b e e n f l o w e r i n g i s e q u a l ( o r

a l m o s t e q u a l ) t o t h e num ber o f s c a r s on t h e r a c h i s and can

b e d e t e r m i n e d b y c o u n t i n g t h e l a t t e r . The f o l l o w i n g num ber

o f s c a r s w e re c o u n t e d on i n f l o r e s c e n c e s t h a t w e re n e a r l y

e x h a u s t e d ; 3 8 t o 1 6 6 (n = 11 ) on M. t r u n c a t a ; i+5 t o 1 51 (n = 15)

on £■[• m a l a c c e n s i s an d 28 t o 2 0 7 (*i = 2 3 ) on c u l t i v a t e d b a n a n a s .

(More e x p o s e d p l a n t s and t h o s e g ro w in g i n m a r g i n a l h a b i t a t s

w e re s m a l l e r and h a d fe w e r h a n d s . )

The f e m a le s e c t i o n o f t h e i n f l o r e s c e n c e d i f f e r e d

i n t h a t s e v e r a l b r a c t s . o p e n e d more o r l e s s s i m u l t a n e o u s l y

an d d i d n o t r o l l up b u t r e m a in e d i n s i t u f o r f o u r o r f i v e

d a y s b e f o r e f a l l i n g . The f e m a le f l o w e r s w e re n o t d e c i d u o u s .

The b r a c t s o f Musa v i o l a s c e n s w e re n o t d e c id u o u s

an d t h e y a lw a y s o p e n e d i n t h e e a r l y m o rn in g s t o e x p o s e t h e

f l o w e r s t h r o u g h t h e d a y . Some o b s e r v a t i o n s s u g g e s t e d t h a t

t h e m a le h a n d s r e m a in e d on t h e p l a n t l o n g e r t h a n 2 4 h o u r s .

b u t t h i s was n e v e r c o n f i r m e d .

V i s i t o r s b e s i d e s m a c r o g l o s s i n e b a t s .

S m a l l m e l a p o n id b e e s ( T r ig o n a s p p . ) w e re c o l l e c t e d

from t h e f l o w e r s o f M. m a l a c c e n s i s n e a r B e n to n g and on t h e

T a p a h - R i n g l e t r o a d and w e re s e e n on t h e f l o w e r s o f t h i s s p e c i e s

w h e r e v e r t h e y w e re e x a m in ed d u r i n g t h e d a y . The b e e s do n o t

p e n e t r a t e t h e f l o w e r s b u t g a t h e r p o l l e n from t h e t i p s o f t h e

a n t h e r s w h ich p r o j e c t b e y o n d t h e compound t e p a l . ( P o l l e n m a s se s

t a k e n from t h e t i b i a s o f s e v e r a l s p e c im e n s c o n t a i n e d o n l y Musa

s p p . p o l l e n . ) M e la p o n id b e e s w ere f r e q u e n t l y s e e n on t h e

f l o w e r s o f c u l t i v a t e d b a n a n a s and M. v i o l a s c e n s .

M oths ( u n i d e n t i f i e d b u t a l l a p p a r e n t l y o f t h e

same s p e c i e s ) g a t h e r e d b e lo w t h e f l o w e r s o f M. t r u n c a t a a s

so o n a s t h e b r a c t s b e g a n t o o p e n . As many a s 17 w e re r e c o r d e d

s i m u l t a n e o u s l y on one i n f l o r e s c e n c e b u t b y a b o u t 2 1 . 0 0 h r s

m o s t h a d d e p a r t e d ; a few r e m a in e d a l l n i g h t . They moved a b o u t

42

on t h e b r a c t s b e lo w t h e f l o w e r s f e e d i n g from d r o p l e t s o f

l i q u i d t h a t g a t h e r e d t h e r e , ( t h e n a t u r e o f t h e l i q u i d i s n o t

knbwn) an d s o m e t im e s p r o b i n g t h e m o u ths o f t h e f l o w e r s , b u t

t h e y w e re u n a b l e t o f o r c e t h e t e p a l s a p a r t ,

• B i r d s w e re a l s o f r e q u e n t v i s i t o r s t o Musa f l o w e r s .

A t R a n ta u P a n j a n g , s u n b i r d s , m o s t l y L ep tocom a , j u g u l a r i s L.

( N e c t a r i n i d a e ) , f r e q u e n t l y v i s i t e d c u l t i v a t e d b a n a n a s b y d a y ;

e l s e w h e r e t h e m o s t commonlyseen v i s i t o r s w e re t h e s p i d e r h u n t e r s ,

A r a c h n o t h e r a s p p . ( N e c t a r i n i d a e ) ,

On t h e s p e c i e s w i t h p e n d u lo u s i n f l o r e s c e n c e s t h e

b i r d s h an g u p s i d e down from t h e open b r a c t p r o b i n g e a c h f l o w e r

i n t u r n . T h e i r c la w s l e a v e s m a l l d a r k m arks on t h e b r a c t s

( m o s t l y on t h e d o r s a l s u r f a c e n e a r t h e b a s e u n l e s s t h e b r a c t

h a d r o l l e d up c o m p l e t e l y ) w h e re t h e y p u n c t u r e i t . Where t h e

f l o w e r s a r e r e a d y t o f a l l t h e i r p r o b i n g f r e q u e n t l y d i s l o d g e s

them ; e . g . on Gunong B r i n c h a n g s p i d e r h u n t e r s w ere w a tc h e d a s

t h e y f e d from t h e f l o w e r s o f M. t r u n c a t a a t a b o u t 0 7 , 0 0 h r s .

One b i r d p ro b e d e a c h o f t h e 1 6 f l o w e r s on one h an d i n t u r n

an d e v e r y f l o w e r f e l l o f f a s a r e s u l t o f t h e b i r d ' s a c t i v i t y .

When v i s i t i n g f l o w e r s o f M. v i o l a s c e n s t h e

b i r d s p e r c h on t h e a p e x o f t h e i n f l o r e s c e n c e r e a c h i n g down

t o p r o b e t h e f l o w e r s . The t i p s o f a l l i n f l o r e s c e n c e s a r e

d i s f i g u r e d b y t h e p u n c t u r e m a rk s made by t h e b i r d s ' c l a w s .

( F i g . 11 show s t h e i n f l o r e s c e n c e o f a n o t h e r t y p i c a l o r n i t h o -

p h i l o u s s p e c i e s , M. b o r n e e n s i s B e c c a r i ( s e c t i o n C a l l i r n u s a )

p h o to g r a p h e d a t Tav/au, S a b a h . An t h e s i s was d i u r n a l ; t h e c o l o u r

was y e l l o w a n d , a l t h o u g h t h e i n f l o r e s c e n c e w as p e n d u l o u s , t h e

/

I

k3

s i n g l e row o f f l o w e r s p r o j e c t e d a t an a c u t e a n g l e from t h e

u n d e r l y i n g b r a c t . The c law m arks o f b i r d s a r e c l e a r l y v i s i b l e

bn t h e u n o p e n e d b r a c t s . )

.2 , Palmae 0

Two g e n e r a c o n t a i n c h i r o p t e r o p h i l o u s s p e c i e s ,

C ocos (o n e i n t r o d u c e d s p e c i e s , C., n u c i f e r a L . ) an d ' A ren g a

( f o u r s p e c i e s i n M a la y a , W h itm o re , 1973) b u t o n l y two a r e

l i k e l y t o b e c h i r o p t e r o p h i l o u s , A. p i n n a t a (Wurinb.) M e r r i l l

a n d A. w e s t e r h o u t i i G r i f f . . The o t h e r two a r e s m a l l p l a n t s o f

t h e f o r e s t u n d e r g r o w t h .

2 . i , Cocos "n u c i f e r a .

E c o l o g i c a l d i s t r i b u t i o n .

I n M a l a y s i a c o c o n u t p a lm s a r e o n l y fo u n d u n d e r

c u l t i v a t i o n . They a r e common an d w i d e s p r e a d i n t h e l o w l a n d s ,

e s p e c i a l l y i n c o a s t a l d i s t r i c t s w h e re t h e y a r e o f t e n grown

i n p l a n t a t i o n s w i t h o t h e r c r o p s p l a n t e d b e lo w th e m .

M o rph o log y ( F i g . 3 9 ) .

D e s c r i b e d b y many a u t h o r s ( e . g . B l a t t e r , 1 9 26 ;

W h itm o re , 197 3 )*

N e c t a r .

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

volum e was n o t m e a s u r e d .

P o l l e n .

P o l l e n h a s b e e n d e s c r i b e d by T h a n ik a im o n i ( 1 9 6 6 ) .

The l a r g e s u l c a t e g r a i n s h a v e a sm ooth e x i n e b u t t h e s u l c u s

i s c h a r a c t e r i s t i c and d i s t i n g u i s h e s them from r u p t u r e d D u r io

kk

s p e c i e s g r a i n s w i t h w h ich t h e y m ig h t h av e o t h e r w i s e h a v e

b e e n c o n f u s e d . T h e i r mean l e n g t h was 9 7 . 5 /u an d t h e i r mean

d i a m e t e r 76 .1 / i (n = 2 0 ) .

S c e n t .

No d a t a .

F l o w e r i n g b i o l o g y .

I t h a s b e e n d e s c r i b e d by B l a t t e r ( 1 9 2 6 ) . No

f u r t h e r o b s e r v a t i o n s w ere made e x c e p t t h a t many i n s e c t s

e s p e c i a l l y m e l a p o n i d b e e s ( T r ig o n a s p p . ) were- commonly

a t t r a c t e d t o t h e f l o w e r s . S u n b i r d s , N e c t a r i n i a s p p . ( N e c t a r i n i d a e )

w ere v e r y o f t e n o b s e r v e d f e e d i n g among t h e f l o w e r s .

P h e n o l o g y .

Once m a t u r e , c o c o n u t p a lm s f l o w e r e d c o n t i n o u s l y ,

one i n f l o r e s c e n c e d e v e l o p i n g i n t h e a x i l o f e a c h f r o n d .

2 . i i ' . , - A r e n g a ,

N o t e . No b a t s w e re s e e n v i s i t i n g a n y s p e c i e s

o f A r e n g a , h o w e v e r , p o l l e n commonly fo u n d i n t h e gu ano o f

E o n y c t e r i s s p e l a e a i n Gua S a n d in g (a n d v e r y r a r e l y i n B a tu

C a ves) was i d e n t i f i e d i n d e p e n d e n t l y by D r . J . M u l l e r , R e ^ k s -

h e r b a r i u m , L e id e n and D r . T h a n i k a i m o n i , P o n d i c h e r r y I n s t i t u t e ,

I n d i a , a s b e i n g o f t h i s g e n u s . W hitm ore (1 9 7 3 ) r e c o r d e d f o u r

s p e c i e s o f A re n g a i n P e n i n s u l a r M a l a y s i a , two (A. h a s t a t a

( B e c c . ) W hitm ore and _A. h o o k e r i a n a ( B e c c . ) W h itm o re ) a r e

p l a n t s o f t h e f o r e s t u n d e r g r o w t h . They h av e v e r y s m a l l

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

The r e m a i n i n g two a r e A. p i n n a t a and A. w e s t e r h o u t i i ; A. p i n n a t a

b5

t h e s u g a r p a lm , i s i n t r o d u c e d and was a t one t i m e grown w id e ly ,

f o r s u g a r p r o d u c t i o n . I t i s s t i l l grown i n P e r a k (A b d u l A z iz ,

1 9 7 ^ ) ; A* w e s t e r h o u t i i i s a w i l d S p e c i e s . T h ese two s p e c i e s

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

a t t r a c t r a a c r o g l o s s i n e b a t s .

E c o l o g i c a l d i s t r i b u t i o n .

B o th s p e c i e s a r e o c c a s i o n a l i n t h e l o w l a n d s ;

A* P i n n a t a i n c u l t i v a t i o n and A. w e s t e r h o u t i i i n t h e v / i l d .

W hitm ore (1 9 7 3 ) d i s c u s s e s t h e i r e c o l o g i c a l p r e f e r e n c e s . Few

p l a n t s w e re n o t e d d u r i n g t h i s s t u d y ,

M o r^ho lo gE ( F i g . 3 9 ) .

The h u g e i n f l o r e s c e n c e s o f A. p i n n a t a and A.

w e s t e r h o u t i i a r e p r o d u c e d i n t h e l e a f a x i l s ( F i g . 1 5 ) . The

f l o w e r s a r e d e s c r i b e d an d f i g u r e d b y W hitm ore (1 9 7 3 )

N e c t a r .

No d a t a .

P o l l e n .

The s m a l l s p i n u l i f e r o u s , s u l c a t e p o l l e n g r a i n s

a r e d i s t i n g u i s h e d from t h o s e o f a l l o t h e r p l a n t s b y t h e s w o l l e n

b a s e s t o t h e s p i n e s ( F i g . 3 6 ) . They a r e t h u s r e a d i l y i d e n t i f i e d

( J . M u l l e r a n d G .T h a n ik a im o n i i n l i t t . , o p i n i o n s e x p r e s s e d

i n d e p e n d e n t l y ) . They h a v e a l s o b e e n f i g u r e d a nd d e s c r i b e d

b y ' T h a n ik a im o n i ( 1 9 6 6 ) . The p o l l e n o b t a i n e d from guano s a m p le s

an d a t t r i b u t e d to. A re n g a was i n d i s t i n g u i s h a b l e from s a m p le s

o f p o l l e n from A re n g a s p p . l o a n e d to me b y D r . R u th K iew . The

mean l e n g t h o f g r a i n s from g uan o s a m p le s was 5 9 .8 p and t h e

mean d i a m e t e r Z+6.2 p (n = 2 0 ) .

Fig. 15. Arenga pinnata, the sugar palm.

Fig. 16. Inflorescences of Oroxylum indicum.Note 1 . How th e y a r e h e l d above th e canopy.

2. One or two buds on each are enlarged. They will open at night.

I

if6

S c e n t *

No d a t a .

F l o w e r i n g b i o l o g y .

No d a t a .

P h e n o l o g y .

They f l o w e r i n a l l m o n th s o f t h e y e a r .

DICOTYLEDONS.

1 . A n a c a r d i a c e a e .

The f l o w e r s o f one g e n u s , M a n g i f e r a (m angos)

a r e v i s i t e d b y m a c r o g l o s s i n e b a t s .

1 . i , M a n g i f e r a ( F i g . 39 )

N o t e . B a t s w e re n o t s e e n t o v i s i t t h e f l o w e r s

o f M a n g i f e r a an d t h e i n c l u s i o n o f t h i s g e n u s i s b a s e d on t h e

i d e n t i f i c a t i o n o f p o l l e n i n g uan o o f E o n y c t e r i s s p e l a e a and

M a c r o g lo s s u s m i n im u s . C o rn e r (1 9 5 2 ) r e c o r d e d a b o u t 20 s p e c i e s

o f M a n g i f e r a i n P e n i n s u l a r M a l a y s i a o f w h ic h n i n e w ere i n

c u l t i v a t i o n , f i v e commonly s o . The o c c u r r e n c e o f M a n g i f e r a

s p p . p o l l e n i n t h e f a e c e s o f M. m in im us a t R a n ta u P a n j a n g f a r

from an y w i l d mangos s t r o n g l y s u g g e s t s t h a t t h e y , a t l e a s t ,

w e re f e e d i n g f rom a c u l t i v a t e d s p e c i e s . The i d e n t i t y o f t h i s

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

w e re c o m p l e t e d and no s t u d i e s w e re made o f t h e mangos a t

R a n ta u P a n j a n g o r a n y w h e re e l s e .

P u r s g l o v e ( 1 9 6 8 ) d e s c r i b e s and i l l u s t r a t e s t h e

f l o w e r s o f M a n g i f e r a i n d i c a w h i l e C o rn e r (1 9 5 2 ) f i g u r e s t h e

f l o w e r s o f s e v e n s p e c i e s o f M a n g i f e r a . i n c l u d i n g M. i n d i c a .

k?

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

P e n i n s u l a r M a l a y s i a .

P o l l e n .

The p o l l e n g r a i n s ( F i g . 3 6 ) a r e u s u a l l y tr i™

c o l p a t e , s p h e r i c a l w i t h a c h a r a c t e r i s i t c e x i n e r e t i c u l a t i o n .

The mean d i a m e t e r o f 10 g r a i n s r e c o v e r e d from g ua n o s a m p le s

was 3 8 . 0 p..

2 . B i g n o n l a c e a e .

Two c h i r o p t e r o p h i l o u s g e n e r a w e re o b s e r v e d ,

O roxylum and P a . j a n e l l a . B o th a r e m o n o s p e c i f i c g e n e r a . The

s p e c i e s , 0 . i n d i c u m and p . l o n g i f o l i a a r e i n d i g e n o u s t o

P e n i n s u l a r M a l a y s i a .

2 » i . O roxylum i n d i c u m .

E c o l o g i c a l d i s t r i b u t i o n .

A common w i d e s p r e a d s p e c i e s r e c o r d e d i n P e r a k ,

P a h a n g , S e l a n g o r , T re n g g a n u a n d K e l a n t a n . I n K e l a n t a n i t i s

o f t e n grown a l o n g s i d e h o u s e s o r p a d i f i e l d s . E l s e w h e r e , i t

i s o c c a s i o n a l l y grown n e a r v i l l a g e s b u t i s more f r e q u e n t on

t h e f o r e s t ed g e and i n young S e c o n d a r y f o r e s t . I t was

p a r t i c u l a r l y common on s t e e p s l o p e s a l o n g t h e T a p a h - P i n g l e t

r o a d w h e re i t p r e f e r r e d f u l l s u n l i g h t b u t was s e ld o m fo u n d

w h e re t h e s o i l d r i e d o u t e x t e n s i v e l y .

M o rp h o lo g y ( F i g . 3 9 ) •

C o r n e r (1 9 5 2 ) an d v an d e r P i j l (1 9 3 6 ) g av e

good a c c o u n t s o f t h e s t r u c t u r e and g ro w th h a b i t o f t h e t r e e .

48

The i n f l o r e s c e n c e i s t e r m i n a l . I t c o n s i s t s o f

a co m p ac t g r o u p o f b u d s a t t h e end o f a l o n g u n b r a n c h e d r a c h i s

w h ic h p r o j e c t s a b o v e t h e c a n o p y ( F i g . 1 6 ) . The b u d s tbpen

a c r o p e t a l l y and t h e r a c h i s c o n t i n u e s t o grow a s new b u d s

d e v e l o p a t i t s m e r i s t e m a t i c a p e x u n t i l i t i s f rom 1 . 5 t o 2 . 0 m

l o n g .

The f l o w e r i s i l l u s t r a t e d i n F i g s . 17 and 1 8 .

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

t h i c k e s t r e g i o n (u p t o 2 . 9 mm t h i c k ) b e i n g on t h e d o r s a l s i d e

f rom k t o 7 cm from t h e a n t e r i o r m a r g in ( F i g . 1 7 ) . The p e t a l s

a r e t h i c k , w r i n k l e d an d f l e x i b l e . The t e r m i n a l p o r t i o n o f t h e

p e d u n c l e , t h e b u d s a nd a l l t h e o u t e r s u r f a c e s o f t h e open

f l o w e r a r e m aroon t o l i v e r c o l o u r e d b u t t h e i n n e r s u r f a c e

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

t u b e a r e c rea m c o l o u r e d , t i n g e d i n p l a c e s w i t h g r e e n o r p u r p l e

( F i g . 1 7 ) .

The f i v e a n t h e r s ( F i g s . 1-7 and 1 8 ) l i e c l o s e t o

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

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

w h ic h p r o j e c t s b e y o n d t h e a n t h e r s , i s b i l o b e d ; t h e two s p a d e

s h a p e d l o b e s a r e c l o s e l y p r e s s e d a g a i n s t one a n o t h e r e x c e p t

a n t e r i o r a l l y w h e re t h e t i p s d i v e r g e s l i g h t l y ( F i g . 18 ) . The

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

o f e a c h l o b e .

P o l l e n ( F i g . 3 6 ) .

The p o l l e n ' . is t y p i c a l o f t h e B i g n o n i a c e a e , The

l a r g e t r i - c o l p a t e g r a i n s a r e n e a r l y s p h e r i c a l when f u l l y

Fig. 17. Oroxylum indicum.

a. Inflorescence with two flowers in anthesis. One has been bisected longitudinally to show the arrangement of anthers. Note the colour scheme,(maroon and white) and the remaining parts of the flower that was in anthesis the previous night; the corolla has fallen off.

b. A fallen corolla to show the marks made by1 . the thumb claws of bats in the corolla,2 . the feet of bats on the petals.

Fig. 18,

O R O X Y L U M I N D I C U M ( Xl J

L.S, w h o l e f l o w e r in a n t h e s i s

S T I G M A

> X2- 5ANTHER

N X 2*5

D i a g r a m s to show h o w po l l en is d e p o s i t e d on t h e h e a d of Eonycter i s

s p e l a e a a n d how t h e b a t * o p e n s * the s t i g m a lobes wh e n f e e d i n g .

49

t u r g i d b u t when d e h y d r a t e d t h e y c r e a s e l o n g i t u d i n a l l y a l o n g

t h e l i n e s o f t h e c o l p i , b ec o m in g t r i l o b e d i n t r a n s v e r s e s e c t i o n .

The e x i n e i s f i n e l y s c u l p t u r e d i n a c h a r a c t e r i s i t c r e t i c u l a t e

p a t t e r n . . I t i s s e m i - s t i c k y an d g r a i n s a g g r e g a t e i n t o e a s i l y

d i s r u p t e d c lu m p s . The mean l e n g t h o f f u l l y t u r g i d g r a i n s was

i ^ u x u m v / I ‘- ^ > 4 \*J- “ c _ w / #

N e c t a r .

N e c t a r was e x t r a c t e d from two f l o w e r s w h ic h

o p en ed on a b r a n c h t h a t h a d f a l l e n o f f one t r e e on t h e

U n i v e r s i t y o f M a lay a campus on t h e a f t e r n o o n o f 14 A u g u s t 1 9 7 3 .

(T he b r a n c h f e l l o f f a t a b o u t 1 6 .0 0 h r s and t h e f l o w e r s , s t i l l

on t h e b r o k e n b r a n c h , o p en ed a t a b o u t 2 2 .0 0 h r s when t h e

q u a n t i t y o f n e c t a r was m e a s u r e d . ) One c o n t a i n e d 0 . 6 m l , t h e

o t h e r 1 .2 m l . The d r y w e i g h t o f a sam ple , o f t h e n e c t a r was

15 .05% o f t h e t o t a l w e i g h t .

S c e n t .

The s c e n t i s m u s ty b u t I fo u n d i t n e i t h e r u n d u e l y

s t r o n g n o r n a u s e a t i n g . I was u n a b l e t o s m e l l t h e f l o w e r u n l e s s

i t was h e l d w i t h i n a b o u t 1 m .

F l o w e r i n g b i o l o g y .

A l l o b s e r v a t i o n s on f l o w e r i n g b i o l o g y w e re

made on t h e two t r e e s g ro w in g on t h e U n i v e r s i t y o f M alay a

campus u n l e s s o t h e r w i s e s t a t e d .

The d a y b e f o r e a n t h e s i s t h e b u d s a p p r o x i m a t e l y

d o u b le t h e i r s i z e . On t h e n i g h t o f a n t h e s i s t h e y b u r s t open

b e tw e e n 2 0 .3 0 and 2 2 . 0 0 h r s . The f l o w e r s w e re o f t e n s e e n t o

open w i t h o u t an y s t i m u l a t i o n from b a t s . H o w e v e r , on f i v e

50

o c c a s i o n s o p e n in g was a p p a r e n t l y s t i m u l a t e d by th em : Each

t im e a b a t ( E o n y c t e r i s s p e l a e a ) was s e e n t o l a n d on an

u n o p en ed b u d , b u t when i t f l e w away a s e c o n d o r two l a t e r ,

t h e b u d h a d b u r s t o p e n . The m echan ism b y w h ic h t h i s was

a c h i e v e d w as unknow n. O b s e r v a t i o n s so m e t im e s c o n t i n u e d u n t i l

02*00 h r s i n t h e m o rn in g when t h e f l o w e r s w ere s t i l l i n t a c t

on t h e t r e e s . By 0 8 . 0 0 h r s , t h e c o r o l l a s o f f l o w e r s t h a t w ere

open t h e n i g h t b e f o r e , h a d d e t a c h e d from t h e r e c e p t a c l e s and

f a l l e n . They t h u s f e l l b e tw e e n 0 2 . 0 0 and 0 8 .0 0 h r s b u t t h e

p r e c i s e t im e i s unknow n. The p e d i c e l s c a r r y i n g t h e r e m a i n i n g

p a r t s o f t h e f l o w e r s , f e l l o f f d u r i n g t h e n e x t 48 t o 60 h o u r s

u n l e s s f r u i t p o d s d e v e l o p e d .

On s i x n i g h t s , b e tw e e n 5 and 19 A u g u s t 197 3 ) t h e

num ber o f f l o w e r s p e r i n f l o r e s c e n c e on t h e two t r e e s was c o u n t e d .

One t r e e h a d two i n f l o r e s c e n c e s , t h e o t h e r f i v e u n t i l 14 A u g u s t

when t h e b r a n c h c a r r y i n g one i n f l o r e s c e n c e f e l l o f f . On 39

o f t h e 40 i n f l o r e s c e n c e - n i g h t s r e c o r d e d e a c h i n f l o r e s c e n c e

h a d two op en f l o w e r s ; t h e r e m a i n i n g one had t h r e e open f l o w e r s .

H o w ev er , on o t h e r o c c a s i o n s so m e t im e s o n ly one f l o w e r op ened

on an i n f l o r e s c e n c e , e s p e c i a l l y when t h e l a t t e r h a d j u s t b e g u n

o r h a d n e a r l y f i n i s h e d f l o w e r i n g .

The r a c h i s on t h e b r a n c h w h ic h b r o k e o f f on

14 A u g u s t h a d 89 p e d i c e l s c a r s , 4 d e v e l o p i n g f r u i t s , 3 c a l y c e s

s t i l l a t t a c h e d t o t h e p e d u n c l e ( t h e s e may h a v e d e v e lo p e d f r u i t s

h a d t h e b r a n c h n o t b r o k e n ) , two f l o w e r s and 49 b u d s . More b u d s

w e re d e v e l o p i n g a t i t s a p e x . E x c l u d i n g t h e t h r e e c a l y c e s t h e r e

w ere t r a c e s o f 97 f o r m e r f l o w e r s b u t o n l y 4 (4 .3% ) h a d s e t

51

f r u i t . T h i s c o m p a re s w i t h k r a c h i s e s t h a t h a d f i n i s h e d

f l o w e r i n g w h ic h w e re c u t o f f a t r e e on t h e T a p a h - R i n g l e t

r o a d on 17 November 1 9 7 2 . They c a r r i e d t h e f o l l o w i n g

R a c h i s n o . 1 2 3 h

P e d i c e l s c a r s 1.21 1 56 159 1 6 7

F r u i t s 1 7 3 h

T o t a l 122 163 162 1 71

% f l o w e r s s e t t i n g 0 .82% 4.29% 1 .85% 2.3*i.%

f r u i t

The l a r g e s t num ber o f f r u i t s r e c o r d e d on an y

r a c h i s d u r i n g t h e s t u d y was 1^ on a t r e e on t h e T a p a h - R i n g l e t

r o a d on 15 S e p te m b e r 1 9 7 2 .

The p a u c i t y o f f r u i t s was n o t a lw a y s due t o a

l a c k o f b a t v i s i t o r s t o t h e f l o w e r s . 1 2ii- f a l l e n c o r o l l a s w ere

c o l l e c t e d b e lo w t h e t r e e s on t h e U n i v e r s i t y o f M a lay a cam p u s .

A l l h a d c la w marks made on them b y b a t s ( s e e b e lo w ) an d y e t

o n ly t h r e e t o s i x f r u i t s d e v e lo p e d on an y r a c h i s on t h e s e

t r e e s .

When b a t s l a n d on t h e f l o w e r s t h e y b u r y t h e

c la w s on t h e i r thum bs i n t o t h e c o r o l l a t u b e , l e a v i n g

c h a r a c t e r i s t i c t r i a n g u l a r p u n c t u r e s i n t h e t h i c k t i s s u e

( F i g . 1f) a nd g r a s p t h e l a t e r a l p e t a l s w i t h t h e i r f e e t , t h e

f i v e t o e s l e a v i n g c h a r a c t e r i s t i c ro w s o f s m a l l p u n c t u r e m arks

on th e m . The p u n c t u r e m a rk s w e re u s u a l l y fo u n d a l o n g t h e

I

52

u p p e r m a r g in s o f t h e p r o m i n e n t w r i n k l e s i n t h e p e t a l s ( F i g , 1 7 ) .

2 5 f r e s h l y f a l l e n c o r o l l a s w ere c o l l e c t e d and

d i v i d e d i n t o e i g h t t r a n s v e r s e b a n d s , 1 cm a p a r t on t h e d o r s a l

s i d e ( F i g , 1 9 ) , The num ber o f c law m arks i n e a c h b a n d was

c o u n t e d . The t h i c k n e s s o f t h e c o r o l l a a t t h e m i d p o i n t i n e a c h

s e c t i o n on t h e m i d - d o r s a l an d m i d - v e n t r a l p o s i t i o n s w e re

m e a s u re d on s i x o f them ( F i g , 19 )* Of 564 c la w m a r k s , 531

( 9 4 . 1 %) w e re on t h e d o r s a l h a l f 0 |f t h e c o r o l l a an d 8 6 . 4 2 %

w e re b e tw e e n 4 an d 7 cm from t h e b a s e ; i . e . i n s e g m e n ts 5>

6 an d 7 . ( A l l o f s e g m e n t 1 and m o s t o f s e g m e n t 2 h a d b e e n

p r o t e c t e d b y t h e c a l y x when t h e f l o w e r s w e re i n s i t u . ) The

t h i c k e s t p o r t i o n o f t h e c o r o l l a w as a l s o b e tw e e n 4 an d 7 cm

from t h e b a s e on t h e d o r s a l s i d e .

F ig . . 1 8 shows how t h e p o s i t i o n o f t h e a n t h e r s

r e s u l t s i n p o l l e n b e i n g d e p o s i t e d on t h e h e a d an d n a p e o f

t h e b a t s a s t h e y f e e d from t h e f l o w e r s . To t r a n s f e r p o l l e n

o n to t h e i n n e r , r e c e p t i v e s u r f a c e o f t h e s t i g m a , i t s lo w e r

l o b e m u s t b e b e n t b a c k b y t h e b a t s h e a d r u b b i n g o v e r i t . a s

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

s t i g m a ’ s r e c e p t i v e s u r f a c e t o b e b r o u g h t i n t o c l o s e c o n t a c t

w i t h t h e h e a d an d n a p e o f t h e b a t , t h u s c o l l e c t i n g p o l l e n

p r e v i o u s l y d e p o s i t e d t h e r e b y o t h e r f l o w e r s . W i t h d r a w a l o f

t h e b a t ’ s h e a d r e t u r n s t h e s t i g m a l o b e s t o t h e i r o r i g i n a l

p o s i t i o n p r e v e n t i n g s e l f p o l l i n a t i o n by p o l l e n j u s t d e p o s i t e d

on t h e b a t ’ s h e a d ( n o t e , t h e s t i g m a p r o j e c t s b e y o n d t h e a n t h e r s ) .

( T h i s m ech an ism was o b s e r v e d when f l o w e r s w e re p r e s e n t e d t o

E o n y c t e r i s s n e l a e a i n c a p t i v i t y . I t was n o t o b s e r v e d i n t h e

f i e l d . )

F i g . 19* A) H i s t o g r a m sh o w in g t h e d i s t r i b u t i o n o f E o n y c t e r i s

s p e l a e a thumb c law m arks on 2 5 f a l l e n c o r o l l a s

o f O roxylum i n d i c u m c o l l e c t e d on t h e U n i v e r s i t y

o f M a la y a cam p u s . Each c o r o l l a was d i v i d e d i n t o

8 s e c t i o n s ( a s shown i n t h e d ia g r a m ) and t h e

num ber o f c law m arks i n e a c h c o u n t e d . A t o t a l

o f 564 c law m arks w e re c o u n t e d .

B) H i s t o g r a m s sh o w in g t h e t h i c k n e s s o f t h e c o r o l l a

i n t h e m i d - d o r s a l (A-A) an d m i d - v e n t r a l (B-B)

r e g i o n s o f e a c h s e c t i o n . M e a s u re m e n ts w ere made

i n mm, from 6 o f t h e c o r o l l a s u s e d i n ( A ) .

THICKNESS (in ram)

at B - B at A - AN> r o co

ro

CO

jyrt

Ul cr»

C5 o>

CO CO

% claw marks per section

[tluw.

P h e n o l o g y ,

The num ber o f Oroxylum in d i c u m t r e e s s e e n i n

f l o w e r , o r f r u i t o r s t e r i l e b e tw e e n m i l e 5 t o m i l e 1k a l o n g

t h e T a p a h - R i n g l e t r o a d w e re c o u n t e d once a m onth from J u n e 1972

t o D ecem ber 1973* The num ber o f t r e e s c o u n t e d e a c h m onth v a r i e d

and no a t t e m p t was made t o f o l l o w t h e h i s t o r y o f i n d i v i d u a l

t r e e s . The p e r c e n t a g e o f t r e e s i n e a c h c o u n t t h a t h a d b u d s

a r e shown i n F i g . 2 0 . I n b o t h y e a r s t h e r e was one d i s t i n c t

f l o w e r i n g s e a s o n l a s t i n g from J u n e t o S e p te m b e r . H ow ever,

n o t a l l t r e e s k e p t t o t h i s p a t t e r n ; t h e two on t h e U n i v e r s i t y

o f M a la y a campus f l o w e r e d from May t o O c to b e r 1 9 7 2 , F e b r u a r y

t o J u n e 1973 (o n e ) a nd F e b r u a r y t o O c to b e r 1973 ( t h e o t h e r ) .

The f r u i t s a r e a t t a c h e d t o t h e r a c h i s f o r t h r e e

t o f o u r m o n th s from f e r t i l i s a t i o n t o t h e t im e t h e y f a l l . A f t e r

t h e l a s t f r u i t s h a v e f a l l e n f rom t h e r a c h i s t h e l e a v e s on t h e

p a r e n t b r a n c h a l s o d i e and a new s h o o t d e v e l o p s from a l a t e r a l

b u d . The b r a n c h e s on one t r e e a r e u s u a l l y , b u t n o t i n v a r i a b l y ,

s y n c h r o n i s e d

V i s i t o r s b e s i d e s m a c r o g l o s s i n e b a t s .

Hawk m o th s ( S p h i n g i d a e ) . so m e t im e s v i s i t e d t h e

f l o w e r s , h o v e r i n g i n f r o n t o f t h e c o r o l l a m ou th b u t t h e y w ere

n o t s e e n t o l a n d , and p r o b a b l y f a i l e d t o p o l l i n a t e a n y f l o w e r s .

2 « i i . P a . j a n e l i a l o n g i f o l i a .

C o r n e r (1 9 5 2 ) g i v e s an a c c o u n t o f t h i s t r e e .

D u r in g t h e p r e s e n t s t u d y i t was o n ly fo u n d once when two t r e e s ,

b o t h i n f l o w e r , w e re l o c a t e d a t m i l e 8 from T apah on t h e r o a d

tree

s w

ith

infl

ore

sce

nc

es

20. Histogram showing the % of Oroxylum indicum trees seen each month on the Tapah to Ringlet road that were flowering. Data were collected between May 1972 and January I97*t.

1 0 0 - j

9 0 -

1972

5h

t o R i n g l e t on 16 F e b r u a r y 1 9 7 3 . They w e re i n t h i c k s e c o n d a r y

f o r e s t t h a t was r e c o l o n i s i n g an o l d r u b b e r p l a n t a t i o n .

M o rp h o lo g y ( F i g . 3 9 ) .

The h u g e , t e r m i n a l i n f l o r e s c e n c e s a r e on

b r a n c h i n g p e d u n c l e s , 1 - 1 , 5 m l o n g , p r o j e c t i n g a b o v e th e

c an o p y ( F i g , 2 1 ) . A t t h e t i m e o f o b s e r v a t i o n i n m i d - a f t e r n o o n

t h e r e w e re no open f l o w e r s on e i t h e r o f t h e t r e e s b u t s i x

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

f l o w e r s w e re g a t h e r e d from b e lo w one t r e e . The c o r o l l a s w e re

s i m i l a r t o t h o s e o f O roxylum i n d i c u m b u t s m a l l e r and l e s s

r o b u s t . The t h i c k n e s s o f t h e m i d - d o r s a l r e g i o n o f t h e c o r o l l a

o f one o f them was a b o u t 1 ,2 mm when i t was m e a s u re d t h a t

e v e n i n g ; The r e m a i n d e r w h ic h v/ere som ewhat w i l t e d v /ere n o t

m e a s u r e d .

The c o l o u r o f t h e f l o w e r s was p u r p l e and t i n g e d

w i t h g r e e n on t h e o u t e r s u r f a c e s . The i n n e r s u r f a c e v/as c ream

c o l o u r e d and t i jn g e d w i t h g r e e n ( F i g . 2 1 ) . T h e re w ere f o u r

a n t h e r s l y i n g a j g a i n s t t h e r o o f o f t h e c o r o l l a . The s t ig m a ;

was b i l o b e d a s i n _0. i n d i c u m b i i t t h e l o b e s w e re n a r r o w e r ( F i g . 2 2 ) .

P o l l e n . \.fl

The p o l l e n i s s i m i l a r t o b u t s m a l l e r t h a n t h a t f

Oroxylum in d ' i c u m . I t was a l s o s l i g h t l y s t i c k y , f o r m in g l o o s e

a g g r e g a t e s

N e c t a r .

No d a t a .

S c e n t .

No d a t a .

Fig. 21 . Pa.janelia longifolia.

<-

a. Inflorescence. Note how it is held above thecanopy.

b. Fallen corolla. Note the white petals and dark corolla tube. The colour of the outer side of the corolla tube is greenish purple, darker than is suggested by the photograph.

Fi g . 22 ,

p a j a n e l i a l o n g i f o l i a ( X I )

W H O L E F L O W E R

V E N T R A L A S P E C T

L. S. L A TE R AL A S P E C T

S T I G M A (X 4 )

A . l a t e r a l

B. v e n t r a l

L .S . V E N T R A L A S P E C T

F l o w e r i n g b i o l o g y .

The f l o w e r s a r e p r o b a b l y n o c t u r n a l ( C o r n e r , 1 9 5 2 ) .

The c o r o l l a t u b e s c o l l e c t e d i n F e b r u a r y 1973 a l l h a d b a t c law

m arks s i m i l a r t o t h o s e o b s e r v e d on c o r o l l a s o f O roxy lum i n d i c u m .

The two t r e e s f l o w e r i n g i n F e b r u a r y 1 973 d i d

n o t f l o w e r a g a i n b e f o r e J a n u a r y 1974 when t h e s t u d y e n d e d ,

3 . B o m b a c a c e a e .

T h re e g e n e r a w ere shown t o c o n t a i n c h i r o p t e r o p h i l o u s

s p e c i e s : Bomb ax ( tw o i n d i g e n o u s s p e c i e s b u t o n l y o n e , B . v a i e t o n i j .,

was o b s e r v e d ) ; C e ib a (o n e i n t r o d u c e d s p e c i e s , C. p e n t a n d r a ) and

D u r io ( t h i r t e e n s p e c i e s r e c o r d e d i n M a l a y s i a * Kochummen ( 1 9 7 2 ) ,

b u t o n l y two w ere shown t o b e c h i r o p t e r o p h i l o u s ; t h e common

d u r i a n , D. z i b e t h i n u s and a w i l d s p e c i e s , D. g r a v e o l e n s B e c c .

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

b u t some e . g . I), g r i f f i t h i ( M a s t . ) B ak h . a r e d e f i n i t e l y n o t s o . )

3 . i . Bombax v a l e t o n i i .

f o r e s t s , i t i s common a l o n g t h e m a in r a n g e i n S e l a n g o r , P a h a n g

and s o u t h e r n P e r a k . I t s s t a t u s e l s e w h e r e was n o t r e c o r d e d .

C o r n e r ( 1 9 5 2 , u n d e r S a l m a l i a ) an d Kochummen (1 9 7 2 )

d e s c r i b e t h e t r e e and v an d e r P i j l ( 1 9 3 6 , u n d e r G o s s a m p in u s )

An e m e r g e n t o f t h e l o w la n d p r i m a r y an d s e c o n d a r y

d e s c r i b e s t h e f l o w e r s - f rom t h e p o i n t o f v iew o f c h i r o p t e r o p h i l y

The f l o w e r i s i l l u s t r a t e d i n F i g s . 23 and 2 4 * A l l t h e b u d s

and f l o w e r s t h a t w e re s e e n w e re s o l i t a r y and a x i l l a r y .

Fig. 23. Bombax valetonii.

a. Portion of a flowering tree. Perak.Note 1:- It flowers while leafless (the leaves

on the left are of a creeper growing up the trunk of the tree).

2:- The huge buds.

b. Single flower in anthesis collected at 21.00 hrs. The stamens are white, the stigma red and the recurved petals are cream coloured.

Fig. 2 4 . B o m b a x v a l e t o n i i

a n t h e s i s

The f l o w e r s a r e h u g e , t h e u n o p e n ed b u d s from one t r e e on t h e

T a p a h - R i n g l e t r o a d w e re 14 cm lo n g . M ost o f t h o s e f r o m ' t r e e s

i n t h e U lu Gombak w e re s h o r t e r , a b o u t 10 cm l o n g ( F i g . 2 4 ) .

The p e t a l s , w h ich r e c u r v e d u r i n g a n t h e s i s ( F i g . 2 3 ) , a r e

g r e e n i s h - w h i t e ; t h e s t a m e n s w h i t e ; t h e a n t h e r s b ro w n ; t h e

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

l o b e s .

T h e re w ere 400 t o 560 (mean 4 52 ) s t a m e n s on

e i g h t f l o w e r s ( tw o from one t r e e a t m i l e 15 on t h e T a p a h -

R i n g l e t r o a d an d s i x from t h r e e t r e e s a t m i l e 15 i n t h e Ulu

Gombak; b o t h e x t r e m e s w ere from t h e l a t t e r l o c a l i t y ) . Each

s ta m e n c a r r i e s a n a n t h e r a b o u t 2 mm l o n g . The s t a m e n s a r e

a r r a n g e d i n s i x g r o u p s . The b a s e s o f a l l s i x f u s e i n t o a

s i n g l e s t r u c t u r e w h ic h e n c i r c l e s t h e s t i g m a an d w h ic h i s a l s o

f u s e d t o t h e b a s e o f t h e p e t a l s . F iv e o f t h e s i x g r o u p s a r i s e

from t h e p e r i p h e r y o f t h i s s t r u c t u r e o p p o s i t e t h e f i v e p e t a l s .

The s i x t h s u r r o u n d s t h e s t i g m a ( F i g . 2 4 ) .

P o l l e n .

The p o l l e n i s d r y an d p o w d e ry . The t r i - c o l p a t e

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

midway a l o n g e a c h s i d e . They a r e c o m p r e s s e d d o r s o v e n t r a l l y .

The e x i n e i s f i n e l y r e t i c u l a t e ( F i g . 3 7 ) • The mean l e n g t h o f

t h e p e r p e n d i c u l a r from one s i d e t o t h e a p e x o f t h e o p p o s i t e

a n g l e was 9 7 . 8 p (n = 2 0 ) .

N e c t a r .

No d a t a was o b t a i n e d on t h e q u a n t i t y o r n a t u r e

o f n e c t a r i n t h e f l o w e r s . The n e c t a r y i s l o c a t e d on t h e r e c e p t a c

57

b e tw e e n t h e s u p e r i o r o v a r y and t h e i n n e r m a r g in o f t h e f u s e d

s t a m i n a l m ass w h ich e n c i r d l e s i t .

S c e n t .

The s c e n t l i k e t h a t o f S o n n e r a t i a s p p . was

r e m i n i s c e n t o f s o u r m i l k b u t i t was n o t p a r t i c u l a r l y s t r o n g . -

F l o w e r i n g b i o l o g y .

The f l o w e r s a r e n o c t u r n a l , op en ir ig a t d u s k . By

e a r l y m o rn in g t h e f u s e d p e t a l s an d s t a m i n a l m ass h a v e f a l l e n .

A t a n t h e s i s t h e p e t a l s c u r l b a c k b u t i n t h e ' f a l l e n c o r o l l a s

t h e y h a v e s t r a i g h t e n e d o u t a g a i n . The i l l u s t r a t i o n i n Kochummen

(1 9 7 2 ) i s c l e a r l y d raw n from a f a l l e n c o r o l l a .

P h e n o l o g y .

Bombax v a l e t o n i i was d e c id u o u s i n S e l a n g o r and

P e r a k an d i t f l o w e r e d w h i l e i t was l e a f l e s s . When t h e s t u d y

b e g a n i n J a n u a r y 1 9 7 2 , t h e t r e e s i n S e l a n g o r wePe l e a f l e s s

an d f l o w e r i n g . New l e a v e s w e re grow n i n F e b r u a r y and March 1972

a n d a g a i n i n 1 9 7 3 . They w ere s h e d i n O c to b e r an d November 1972

a n d a g a i n i n 1 9 7 3 . F l o w e r i n g was r e c o r d e d from November 1972

t o F e b r u a r y 1973 an d o v e r t h e same p e r i o d f o r 1973 - 7 4 . No

d a t a was c o l l e c t e d a f t e r t h e f i r s t week i n F e b r u a r y 1 9 7 4 .

3 » i i i . C e ib a p e n t a n d r a .

T h i s was a common v i l l a g e t r e e i n t h e l o w l a n d s .

I t was s e e n i n a l l t h e s t a t e s i n P e n i n s u l a r M a l a y s i a ( e x c e p t Kedah,

and P e r l i s w h e re I d i d n o t go ) and i n S a b a h , S a ra w a k and

S i n g a p o r e , i t i^ liow a common p a n t r o p i c a l c u l t i v a t e d t r e e

( P u r s g l o v e , 1 9 7 2 ) .

Morp ho log y ( F i g . 3 9 ) .

As t h e t r e e i s c u l t i v a t e d i n a l m o s t a l l . t r o p i c a l

a r e a s , i t h a s b e e n d e s c r i b e d b y many a u t h o r s ( e . g . C o r n e r , 1 95 2 ;

J a e g e r , 1 95 4 ; P u r s g l o v e , 1 9 6 8 ) . The f l o w e r s h a v e b e e n d e s c r i b e d

from a c h i r o p t e r o p h i l o u s s t a n d p o i n t b y B a k e r and H a r r i s ( 1 9 5 9 ) 9

J a e g e r (1 9 5 4 ) a n d van d e r P i j l (1 9 3 6 ) .

P o l l e n ( F i g , 3 7 ) .

The p o l l e n i s d r y a nd p o w d e ry . The g r a i n s a r e

s i m i l a r i n s t r u c t u r e an d s i z e t o t h o s e o f Bomb ax v a l e t o n i i b u t

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

c o l p i a t t h e a n g l e s , i n t h e more d i s t i n c t , c o a r s e r r e t i c u l a t i o n

o f t h e e x i n e . The mean e q u a t o r i a l d i a m e t e r w as 9 7 . 8 / 1 and t h e

p o l a r l e n g t h 4 2 . 3 /1 ( n = 2 0 ) .-

N e c t a r .

No d a t a .

S c e n t .

The s c e n t i s n o t s t r o n g b u t i t i s c h a r a c t e r i s t i c a l l y

r e m i n i s c e n t o f s o u r m i l k .

P h e n o l o g y .

C e ib a p e n t a n d r a f o l lo w e d t h e same p a t t e r n a s

Bombax v a l e t o n i i : I t f l o w e r s a f t e r s h e d d i n g i t s l e a v e s arid

b e f o r e g ro w in g new o n e s . When t h e s t u d y b e g a n i n J a n u a r y 1 972

m o s t t r e e s i n S e l a n g o r w ere l e a f l e s s and f l o w e r i n g b u t I h av e

no r e c o r d s f o r t r e e s e l s e w h e r e . New l e a v e s w e re grov/n i n

F e b r u a r y and M a rc h , They w e re l o s t i n November and December

1972 i n S e l a n g o r an d P e r a k an d new o n e s grown i n F e b r u a r y and

59

a n d M arch 1973* F l o w e r i n g o c c u r r e d d u r i n g t h i s i n t e r v a l . The

l e a v e s w e re l o s t a g a i n i n b o t h t h e s e s t a t e s i n O c t o b e r and

November 1973 a n d f l o w e r i n g o c c u r r e d from t h e n u n t i l t h e

s t u d y e n d e d i n e a r l y F e b r u a r y 1 9 7 ^ . U n l ik e Bombax v a l e t o n i i ,

o c c a s i o n a l f l o w e r i n g t r e e s w e re fo u n d i n a l l m o n th s o f t h e

year- i n S e l a n g o r . I n P e r a k r e c o r d s o f f l o w e r i n g t r e e s w e re

c o n f i n e d t o t h e m o n th s s t a t e d a b o v e b u t no c a r e f u l s e a r c h e s

v /ere made i n o t h e r m o n th s .

3 . i i i . D u r io z i b e t h i n u s .

E c o l o g i c a l d i s t r i b u t i o n .

A common t r e e i n an d a ro u n d v i l l a g e s i n t h e

l o w l a n d s t h r o u g h o u t p e n i n s u l a r M a l a y s i a . I t a l s o g ro w s i n

S a b a h , S a ra w a k a n d e l s e w h e r e (Kochummen, 1 9 7 2 ) .

M o rp h o lo g y ( F i g . 3 9 ) •

T h i s t r e e h a s b e e n d e s c r i b e d b y many a u t h o r s

( e . g . C o r n e r , 1 9 5 2 ; Kochummen, 1 9 7 2 ) . I t i s c a u l i f l o r o u s o r

r a m i f l o r o u s , t h e f l o w e r s c l u s t e r e d on t h e u n d e r s i d e s o f l a r g e r

b r a n c h e s . The f l o w e r s h a v e b e e n d e s c r i b e d f ro m a c h i r o p t e r o p h i l o u s

p o i n t o f v ie w a n d f i g u r e d b y v a n d e r F i j i ( 1 9 3 6 ) . The p e t a l s

an d s t a m e n s a r e w h i t e ; t h e a n t h e r s c re am t o p a l e y e l l o w ( F i g . 25

an d 2 6 ) .

N e c t a r .

The n e c t a r i e s a r e l o c a t e d on t h e r e c e p t a c l e

be tw een , t h e p e t a l s an d t h e i n n e r c a l y x . N e c t a r i s r e t a i n e d i n

t h e s w o l l e n b a s e o f t h e i n n e r c a l y x by s u r f a c e t e n s i o n ( F i g . 26 )

an d an y e x c e s s w h ic h d r i p s o u t i s t r a p p e d b y t h e

F i g . 25 . Durio z i b e t h i n u s .

a . S e c t i o n o f a f lo w e r in g t r e e . Note: Most buds i n some c l u s t e r s a r e c lo s e d w h i le i n o t h e r s th ey a r e open. Many f lo w e rs i n a c l u s t e r i n a n t h e s i s s i m u l t a n e o u s l y enhances t h e i r v i s u a l im p a c t . Pho tographed a t abou t 2 1 .00 h r s .

b. F lo w ers . Pho tographed a t abou t 22.00 h r s .

L.S. OF DURI O ZI BETHI NUS

FLOWER I N A N T H E S I S T O

S H O W LOCATI ON O F NECTAR

. . 60

r e c u r v e d p e t a l s . The p e t a l b a s e s do n o t o v e r l a p and so b a t s

c an e a s i l y r e a c h b e tw e e n them w i t h t h e i r t o n g u e s f o r t h e n e c t a r .

From 0 . 1 0 t o 0 . 6 5 ml (mean 0 . 4 9 + 0 . 1 4 ml) o f

n e c t a r was e x t r a c t e d from 1 ? f l o w e r s a t t h e F . S . C . , U lu Gombak

b e tw e e n 1 9 . 0 0 an d 1 9 .3 0 h r s on 5 A p r i l 1973* 15 o f t h e f l o w e r s

c o n t a i n e d o v e r 0 . 4 0 mi o f n e c t a r a nd n e c t a r may h a v e b e e n l o s t

f rom t h e o t h e r two w h i l e r e m o v in g t h e p a p e r b a g w i t h w h ich

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

f e e d i n g from th e m .

A f t e r t h e n e c t a r was e x t r a c t e d t h e b a g was r e p l a c e d

and t h e e x p e r i m e n t r e p e a t e d a t 2 2 . 5 0 h r s . No m e a s u r e a b l e volum e

o f n e c t a r was o b t a i n e d t h e s e c o n d t i m e .

On 12 O c to b e r 1972 a t 1 9 .3 0 h r s n e c t a r was

e x t r a c t e d from f o u r f l o w e r s on a t r e e on t h e U n i v e r s i t y o f

M a lay a cam p u s . T h ese f l o w e r s h a d n o t b e e n p r o t e c t e d from

v i s i t o r s . . . s e e k i n g n e c t a r . They c o n t a i n e d 0 . 2 ml (1 ) , 0 . 4 5 ml (2 )

an d 0 . 5 5 ml ( 1 ) r e s p e c t i v e l y (mean 0 .41 m l ) . No a n a l y s i s o f

t h e n e c t a r ’ s c o m p o s i t i o n was made.

P o l l e n ( F i g . 3 7 ) .

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

t o t h e f i n g e r s and f o r m in g i r r e g u l a r , e a s i l y d i s r u j ) t e d c lum ps

on t h e a n t h e r s . The e x i n e i s s m o o th . The s p h e r i c a l g r a i n s a r e

u s u a l l y t r i - p o r a t e b u t o c c a s i o n a l l y t e t r a - p o r a t e , One h e x a -

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

t h i c k e n i n g o f t h e w a l l s a r o u n d t h e i r m a r g i n s . The mean d i a m e t e r

was 1 3 0 .4 p. ( n = 2 0 ) . A m u t a n t fo rm , a b o u t h a l f t h e n o rm a l

s i z e , s e m i t r i a n g u l a r i n t r a n s v e r s e s e c t i o n and w i t h e x c e p t i o n a l l y

t h i c k e n e d w a l l s w as fo u n d i n m os t s a m p l e s . I t w a s ,h o w e v e r ,

r a r e . The p r o p o r t i o n o f t h e s e g r a i n s i n e a c h s a m p le was. n o t

c a l c u l a t e d b u t w e re p r o b a b l y < 0 . 1 %.

S c e n t .

D u r ia n f l o w e r s h a d . a c h a r a c t e r i s t i c a n d f a i r l y

s t r o n g s c e n t r e m i n i s c e n t o f s o u r m i l k b u t som ew hat more f r a g r a n t .

F l o w e r i n g b i o l o g y .

On a n y one f l o w e r i n g t r e e t h e r e a r e u s u a l l y

many c l u s t e r s o f f l o w e r s . M ost o f t h e f l o w e r s i n e a c h c l u s t e r

m a t u r e s i m u l t a n e o u s l y and o p e n ' w i t h i n a few d a y s o f o ne a n o t h e r

( F i g . 2 5 ) . H o w e v e r , d i f f e r e n t c l u s t e r s a r e l e s s s y n c h r o n i s e d

so t h a t t h e f l o w e r i n g s e a s o n l a s t e d two o r t h r e e w e e k s .

The f l o w e r s a r e n o c t u r n a l and a n t h e s i s l a s t s one

n i g h t o n l y , " T h e f o l l o w i n g s e q u e n c e was n o rm a l : The o u t e r c a l y x

s p l i t s i n t o two l o b e s more t h a n 2 if h o u r s b e f o r e a n t h e s i s ; t h e

i n n e r c a l y x s p l i t s on t h e m o rn in g b e f o r e a n t h e s i s a n d t h e s t a m e n s

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

The p e t a l s do no.t r e c u r v e u n t i l a f t e r d a r k a l t h o u g h t h e a n t h e r s

may d e h i s c e d u r i n g t h e a f t e r n o o n . The m o rn in g f o l l o w i n g a n t h e s i s

t h e c a l y x , p e t a l s and a n t h e r s f a l l , l e a v i n g t h e g y n a e c iu m i n

s i t u b u t many o f t h e s e a l s o f a l l o f f i n t h e s u c c e e d i n g few d a y s .

( I t was n o t known i f t h e y w e re U n f e r t i l i s e d o r w h e th e r t h e y

w e te a b o r t e d . ) T h ose t h a t r e m a in s w e l l t o form s m a l l f r u i t s

b u t m os t f a l l o f f b e f o r e t h e y a r e more t h a n 3 cm i n d i a m e t e r .

V ery se ld o m d o e s more t h a n one f r u i t d e v e l o p from e a c h c l u s t e r

o f f l o w e r s and manyc c l u s t © r , s 7p ro d u e e n n o r f r u t t s .

62

V i s i t o r s b e s i d e s m a c r o g l o s s i n e b a t s .

The f r u i t b a t s , C y n o p te r u s b r a c h y o t i s M u l l e r ,

w ere c a u g h t b e lo w f l o w e r i n g d u r i a n t r e e s w i t h D u r io s p p . p o l l e n

on t h e i r b o d i e s and a w h i t e s u b s t a n c e i n t h e i r s t o m a c h s , -

i n c l u d i n g a few D u r io s p p . p o l l e n g r a i n s . They a p p e a r e d t o

h a v e b e e n e a t i n g t h e f lo v /e r s . . The f l y i n g f o x , p t e r o p u s vam pyrus L . ,

a l s o a t e t h e f l o w e r s and . i s - r e p u t e d b y v i l l a g e r s t o b e a s e r io v i s

p e s t o f d u r i a n s .

D u r in g t h e d ay m e l a p o n id b e e s a r e v e r y common

on p a r t i a l l y open d u r i a n f l o w e r s . I n t h e m o r n i n g s , t h e y c o l l e c t e d

p o l l e n from t h e a n t h e r s o f f l o w e r s t h a t h a v e o p en ed t h e n i g h t

b e f o r e b u t w h ic h h a v e n o t y e t f a l l e n o f f and i n t h e a f t e r n o o n s ,

t h e y s e a r c h f o r p o l l e n i n t h e a e w ly o p e n in g f l o w e r s . They a r e

n o t p r e s e n t on t h e f l o w e r s a f t e r d a r k . A l th o u g h t h e y move a l l

o v e r t h e a n t h e r m ass t h e y w e re n e v e r o b s e r v e d t o c o n t a c t t h e

s t i g m a a t t h e a p e x o f t h e p r o j e c t i n g s t y l e ( F i g , 2 5 ) •

H a n g in g p a r r o t s ( L o r i c u l u s g a l g u l u s L , P s i t t a c i d a e )

an d v a r i o u s s p e c i e s o f s u n b i r d s ( N e c t a r i n i d a e ) v/ere t h e m o s t

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

t h e F . S . C . , U lu Gombak. S u n b i r d s w ere a l s o commonly s e e n t o

v i s i t d u r i a n f l o w e r s e l s e v / h e r e . I was n o t a b l e t o d e t e r m i n e

t h e a t t r a c t i o n t o t h e p a r r o t s b u t t h e s u n b i r d s p r o b e d t h e

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

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

f a l l o f f , They c b u l d h a v e t r a n s f e r r e d p o l l e n from one f l o w e r

t o a n o t h e r o r one t r e e t o a n o t h e r .

A t n i g h t m o th s o f many s p e c i e s (n o n e w e re i d e n t i f i e d )

v/ere s e e n t o v i s i t t h e f l o w e r s . They al} . l a n d e d on t h e c a l y x

o r t h e r e c u r v e d p e t a l s and i n s e r t e d t h e i r p r o b o s c e s b e tw e e n

t h e b a s e s o f t h e p e t a l s i n s e a r c h o f n e c t a r . None w ere s e e n

t o come i n t o c o n t a c t w i t h e i t h e r t h e a n t h e r s o r t h e s t i g m a .

The b row n n i g h t w a s p s , P r o v e s p a ano m a le ( V e s p i d a e ) , w e re a l s o

f r e q u e n t l y s e e n c l a m b e r i n g o v e r t h e a n t h e r s a t n i g h t b u t l i k e

t h e m e la p o n iu b e e s t h e y d i d n o t come i n t o c o n t a c t w i t h t h e

s t i g m a .

P h e n o l o g y .

The t r e e s a r e n o t d e c i d u o u s . T h e re w ere two

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

c o n s i d e r a b l y i n i n t e n s i t y . I n S e l a n g o r t h e f i r s t f l o w e r i n g

s e a s o n was f rom e a r l y March u n t i l mid A p r i l 1 972 an d from

mid March t o mid A p r i l 19 7 3 ; t h e s e c o n d from mid S e p te m b e r

t o mid O c to b e r 1972 b u t c o n f i n e d t o S e p te m b e r i n 1 9 7 3 . Fewer

d a t a w ere c o l l e c t e d i n o t h e r s t a t e s b u t a l l f l o w e r i n g r e c o r d s

w e re i n t h e same m o n th s a s t h o s e i n S e l a n g o r . N o te s v/ere k e p t

on t h e a b u n d a n c e o f r o a d s i d e s t a l l s s e l l i n g d u r a i n s . F r u i t

was a b u n d a n t i n a l l s t a t e s from December t o F e b r u a r y an d from

J u n e t o A u g u s t i n 1972 an d 1 9 7 3 . H ow ev er , t h e r e w e re a lv /ays

a few o u t o f s e a s o n f r u i t s on s a l e i n d i c a t i n g t h a t v e r y

o c c a s i o n a l l y t r e e s f l o w e r e d o u t o f s e a s o n . O n ly one t r e e , a

c u l t i v a r ( t h e " S ia m e s e d u r i a n " ) was fo u n d f l o w e r i n g o u t o f

s a e s o n . I t was i n f l o v /e r n e a r P o r t D ic k s o n , N e g r i S e m b i la n

on 23 A p r i l 1972 a b o u t t h r e e w eeks a f t e r t h e l a s t f l o w e r i n g

r e c o r d f o r o t h e r d u r i a n t r e e s .

T ho se t r e e s t h a t f l o w e r e d d u r i n g t h e s e c o n d

f l o w e r i n g s e a s o n h a d u s u a l l y a l s o f l o w e r e d d u r i n g t h e f i r s t .

6k

T h u s , i n d i v i d u a l t r e e s c a n f l o w e r t w i c e i n a y e a r . One' t r e e

on t h e U n i v e r s i t y o f M a lay a campus and two a t t h e F . S . C . , U lu

Gombak f l o w e r e d t w i c e i n 1972 and once ( i n M arch a n d A p r i l )

i n 1 9 7 3 , i . e . i n t h r e e s u c c e s s i v e s e a s o n s .

I n b o t h y e a r s f l o w e r i n g i n t h e f i r s t (M arch -

A p r i l ) s e a s o n was h e a v i e r t h a n t h e s e c o n d (A u g u s t - S e p t e m b e r ) .

T h i s i s n o rm a l a n d t h e f i r s t f r u i t s e a s o n i s f r e q u e n t l y r e f e r r e d

t o a s t h e "m a in s e a s o n " . I n 1973 t h e d i f f e r e n c e was v e r y

m a rk e d . The f i r s t f r u i t s e a s o n t h a t y e a r was " t h e l o n g e s t and

m o s t p l e n t i f u l " f o r 25 y e a r s ( S t r a i t s T im e s , 19 J u l y 1 9 7 3 ) .

T h i s f o l l o w e d an e x c e p t i o n a l l y h e a v y f l o w e r i n g s e a s o n . The.

s e c o n d , h o w e v e r , was r e m a r k a b l y p o o r . Many t r e e s f a i l e d t o

f l o w e r a l t o g e t h e r an d i n some a r e a s , no t r e e s f l o w e r e d ( e . g . th. t l ib e tw e e n t h e 11 a n d 15 m i l e , U lu Gombak, a r e g i o n fam ous

f o r i t s d u r i a n s ) .

3 . i v . D u r io g r a v e o l e n s .

E c o l o g i c a l d i s t r i b u t i o n -.

No d a t a was c o l l e c t e d on t h i s t r e e i n t h e w i l d

b u t one was ex am in ed i n t h e a r b o r e t u m o f t h e F o r e s t r y R e s e a r c h

I n s t i t u t e , K epong , S e l a n g o r . I t i s an i n d i g e n o u s p r i m a r y

f o r e s t t r e e an d i s n o t u s u a l l y c u l t i v a t e d i n P e n i n s u l a r M a l a y s i a .

M o rp h o lo g y ( F i g . 3 9 ) ,

I t was d e s c r i b e d an d f i g u r e d b y Kochummen ( 1 9 7 2 ) . .

The f l o w e r s a r e v e r y s i m i l a r t o t h o s e o f D u r io z i b e t h i n u s b u t

l a r g e r . On t h e t r e e e x a m in e d t h e y v/ere c l u s t e r e d a l o n g t h e

p r o x i m a l e n d s o f a l m o s t a l l t h e b r a n c h e s o v e r .3 cm t h i c k and

65

a l o n g t h e t r u n k t o g ro u n d l e v e l . The t r e e was n o t very , o l d ,

a b o u t 1 0 m t a l l an d w 1 t h a g i r t h o f a b o u t 9 0 ciiu

P o l l e n ( F i g . 3 7 ) .

I t was s i m i l a r i n g r o s s t e x t u r e t o an d i n d i s t i n g u i s h ­

a b l e m i c r o s c o p i c a l l y from t h a t o f D u r io z i b e t h i n u s .

N e c t a r .

No n e c t a r was e x t r a c t e d from t h e f l o w e r s .

S c e n t .

I t was s i m i l a r t o t h a t o f D. z i b e t h i n u s .

F l o w e r i n g b i o l o g y ,

g r a v e o l e n s was n o c t u r n a l , t h e f l o w e r s o p e n in g

f u l l y i n t h e e v e n i n g an d l a s t i n g o n ly one n i g h t . The s e q u e n c e

l e a d i n g up t o a n t h e s i s was s i m i l a r t o t h a t - r e p o r t e d ' . f o r ,

D. z i b e t h i n u s . a b o v e .

V i s i t o r s b e s i d e s m a c r o g l o s s i n e b a t s .

L ik e D. z i b e t h i n u s , m e la p o n id b e e s c o l l e c t e d

p o l l e n from t h e f l o w e r s d u r i n g t h e d ay an d m o th s s o u g h t n e c t a r

a t n i g h t . N e i t h e r w e re l i k e l y t o c o n t a c t t h e s t i g m a . No o t h e r

a n i m a l s w ere o b s e r v e d b u t .o b s e r v a t i o n s v/ere v e r y l i m i t e d .

H ow ever , s e v e r a l u n o p e n e d b u d s an d many f l o w e r s h a d b e e n

p a r t i & l l y o r t o t a l l y consum ed b y s q u i r r e l s .

P h e n o l o g y .

The t r e e f l o w e r e d i n March 1 972 a n d f r u i t s

r i p e n e d i n J u l y and A u g u s t t h a t y e a r . I t f l o w e r e d a g a i n i n

March 1973 a nd f r u i t s r i p e n e d i n J u l y t h a t y e a r . The d a t e s

on w h ich e l e v e n f l o w e r i n g s p e c im e n s i n t h e h e r b a r i u m o f t h e

66

F o r e s t E e a s e a r c h I n s t i t u t e w ere c o l l e c t e d i n d i c a t e a v e r y w ide

f l o w e r i n g s e a s o n , v i a .

' Month No. s p e c im e n s S t a t e

March 3 N o r th P e r a k ( l ) , S e l a n g o r (2 )

A p r i l 1 , N o r th P e r a k

May t S e l a n g o r

J u l y 1 P a h a n g

A u g u s t 2 S e l a n g o r ( 1 ) , N e g r i S e m b i la n (1 )

S e p te m b e r 2 P e r a k ( 1 ) , T re n g g a n u (1 )

O c to b e r 1 T re n g g a n u

T o t a l 11

3 » v . O th e r s p e c i e s o f D u r i o .

No f i e l d w ork w as u n d e r t a k e n on a n y o t h e r s p e c i e s

o f D u r io ( e x c e p t D u r io g r i f f i t h i , w h ic h i s n o t c h i r o p t e r o p h i l o u s )

and t h e i r p o l l i n a t i n g a g e n t s a r e unknown h u t b a t s may be

i m p o r t a n t t o some s p e c i e s . F l o w e r i n g s p e c i m e n s o f e i g h t s p e c i e s . :

w e re ex a m in ed i n t h e h e r b a r i u m o f t h e F o r e s t R e s e a r c h I n s t i t u t e ,

K epong an d T a b le 2 .show s t h e i r c o l l e c t i o n d a t e s .

A l th o u g h t h e d a t a a r e v e r y s c a n t ' t h e y

s u g g e s t t h a t m o s t s p e c i e s f l o w e r b e tw e e n A p r i l an d J u l y a n d / o r

b e tw e e n S e p te m b e r and N ovem ber. T h ese d a t e s a r e a l i t t l e l a t e r

t h a n t h e f l o w e r i n g s e a s o n s o f D_. z i b e t h i n u s r e c o r d e d i n 1 972

and 1 9 7 3 .

Table 2 . A r e c o r d o f th e months i n which f lo w e r in g spec imens o f

some Dur i o s p p . were c o l l e c t e d i n P e n i n s u l a r M a la y s i a . Data from

specimens d e p o s i t e d i n t h e h e rb a r iu m o f th e F o r e s t Resea rch

I n s t i t u t e , Kepong, S e l a n g o r .

S p e c ie s J F M A M J J A S 0 N D T o ta l

D. a f f i n u s 3 3

D. c a r i n a t u s 1 1 2 1 2 7

D. k u t . i e n s i s 0

D. lowianus 2 2 1 1 6

D. ox leyanus 1 1 2

D. m a c ro rh y l lu s 2 T 1 1 5

D. ra a laccen s is 6 2 2 1 0 ‘

D. p in a n g ia n u s 1 2 1 k

T o t a l 1 - - 5 7 7 3 - k 5 5 - 37

^ e c y t i i i d a c e a e ( F i g . 3 9 )*

P o l l e n o f one g e n u s , B a r r i n g t o n i a , was v e r y r a r e

i n g uan o o f E . s p e l a e a c o l l e c t e d from B a tu C aves a nd no b a t s

w e re s e e n t o v i s i t e i t h e r B . r a c e m o s a L . o r B . a s i a t i c a K u r t z ,

d u r i n g many i n t e r m i t t e n t o b s e r v a t i o n s on b o t h s p e c i e s . The

p o l l e n o f B a r r i n g L u i i i a i s c h a r a c t e r i s t i c and e a s i l y i d e n t i f i e d

( F i g . 3 8 ) ( E r d t m a n , 1 9 52 ; P a y e n s , 1 9 6 ? ) . F r e s h p o l l e n c o l l e c t e d

from f l o w e r s was n o t b r o k e n down by a c e t o l y s i s an d i t c a n t h u s

b e r e g a r d e d a s i n e r t and u n l i k e l y t o b r e a k down r a p i d l y i n

g u a n o . F u r t h e r m o r e , t h o s e g r a i n s t h a t w ere fo u n d i n g uano

showed no s i g n s o f d i s i n t e g r a t i o n . The s c a r c i t y o f B a r r i n g t o n i a

spp. p o l l e n i n t h e g u an o s u g g e s t s t h a t e i t h e r t h e b a t s f e d o n ly

f rom a r a r e - s p e c i e s ( o r one t h a t f l o w e r e d r a r e l y d u r i n g t h e

s t u d y ) o r t h a t t h e y se ld o m f e d from an y s p e c i e s o f B a r r i n g t o n i a

( P a y e n s t (1 9 6 7 ) r e v i s e d B a r r i n g t o n i a and s u g g e s t e d t h a t some

c a u l i f l o r o u s s p e c i e s m ig h t b e c h i r o p t e r o p h i l o u s b u t t h a t m o s t

a r e e n t o m o p h i l o u s . C o rn e r (1 9 5 2 ) h a s d e s c r i b e d t h e s p e c i e s

i n h a b i t i n g P e n i n s u l a r M a l a y s i a . )

5 . L e g u m in o s a e .

One g e n u s , P a r k i a , was fo u n d t o c o n t a i n two

c h i r o p t e r o p h i l o u s s p e c i e s , P . .- javanica and P . s p e c i o s a . The

r e m a i n i n g two s p e c i e s o f P a r k i a i n P e n i n s u l a r M a l a y s i a ,

P . s i n g u l a r i s M iq . and P . s u m a t r a n a M iq . a r e r a r e an d w e re

n o t s t u d i e d i n t h e f i e l d b u t t h e s t r u c t u r e o f t h e i r i n f l o r e s c e n c e s

s u g g e s t t h a t t h e y a r e c h i r o p t e r o p h i l o u s .

E c o l o g i c a l d i s t r i b u t i o n .

P a r k i a . j a v a n ic a and P . g r e c i o s a a r e common t r e e s

i n lo w la n d p r i m a r y and - s e c o n d a ry f o r e s t s i n S e l a n g o r , P e r a k

a s v / e l l a s p a r t s o f w e s t e r n P ah an g and N e g r i S e ra b i la n ( e . g .

T a b le 1 ) . T h e i r s t a t u s e l s e w h e r e was no't c h e c k e d . B o th s p e c i e s

a r e f r e q u e n t l y l e f t s t a n d i n g d u r i n g f o r e s t c l e a r a n c e o p e r a t i o n s

a s t h e i r f r u i t a r e e a t e n b y some p e o p l e ; T h i s a p p l i e s

p a r t i c u l a r l y t o P . s p e c i o s a , w h ich i s a l s o p l a n t e d i n some

v i l l a g e s . P . s i n g u l a r i s an d P . s u m a t r a n a a r e a l s o lo w la n d

f o r e s t t r e e s (W h i tm o re , 1 9 7 2 ) .

M 'orphology ( F i g . 3 9 ) .

The t r e e s h a v e b e e n d e s c r i b e d b y W hitm ore (1 972)

an d C o rn e r ( 1 9 5 2 ) . Van d e r P i j l (1 9 3 6 ) d e s c r i b e d t h e i n f l o r e s c e n c e

£ • s p e c i o s a from t h e p o i n t o f v iew o f c h i r o p t e r o p h i l y . The

i n f l o r e s c e n c e s o f P . j a v a n i c a , P . s i n g u l a r i s and P . s u m a t r a n a

a r e v e r y s i m i l a r t o t h a t o f P . s p e c i o s a w h ic h i s i l l u s t r a t e d

i n F i g s . 2.7 and 2 8 . They a r e a l l p a l e y e l l o w b u t t h e c o l o u r

i s i n t e n s i f i e d by t h e b r i g h t y e l l o w a n t h e r s d u r i n g a n t h e s i s .

N e c t a r .

No d a t a was o b t a i n e d on n e c t a r p r o d u c t i o n .

P o l l e n ( F i g , 3 7 ) .

The p o l l e n o f t h r e e s p e c i e s ( P . j a v a n i c a , P . s p e c i o s a

a n d P . s i n g u l a r i s ) w ere e x a m in e d . The s p e c i e s w e re fo u n d t o

be p a l y n o l o g i c a l l y i n s e p a r a b l e . They fo rm ed ' , ,p o ly a d s .with 1 6

o r , r a r e l y , 3 2 g r a i n s i n e a c h p o l y a d . The o u t e r s u r f a c e s o f

t h e g r a i n s w ere v e r y f i n e l y r e t i c u l a t e , t h e d e g r e e o f r e t i c u l a t i o n

v a r i e d from d i s t i n c t t o a l m o s t i n v i s i b l e .

Single inflorescence , whole and bisected longitudinally. Note the many small flowers that make up one inflorescence. Those in the collar region (l) produce nectar. The remainder are fertile. The n furry" looking structures are anthers and filaments.

F ig . 27 . .Parkia s p e c io s a .

Portion of a tree in flower. Note the pendulous inflorescences in various stages of development.

1

F i j . 1 8 -

S c e n t*

The s c e n t o f f r e s h f l o w e r s on t h e t r e e s a t a n t h e s i

was n o t r e c o r d e d b u t f a l l e n i n f l o r e s c e n c e s h ad a s o u r s m e l l .

F l o w e r i n g b i o l o g y .

No d a t a was o b t a i n e d on t h e f l o w e r i n g b i o l o g y

w h ich h a s b e e n d e s c r i b e d f o r P . s p e c i o s a by v an d e r - P i j l ( 1 9 3 6 ) .

V i s i t o r s b e s i d e s m a c r o g l o s s i n e b a t s .

I n s e c t s , e s p e c i a l l y m e la p o n id b e e s w ere i n v a r i a b l y

a t t r a c t e d t o t h e i n f l o r e s c e n c e s o f P . . j a v a n i c a a n d P . s t ? e c io s a

d u r i n g t h e d a y . No o t h e r v i s i t o r s w ere s e e n .

P h e n o l o g y .

32 P . s p e c i o s a a nd 2 if P . . j a v a n i c a t r e e s g ro w in g

b e tw e e n 7 a n d 1 /+ m i l e s from T apah on t h e r o a d t o R i n g l e t w ere

n um bered and ex am in ed e a c h m onth from J u n e 1 972 t o J a n u a r y 197A-.

F i g . 29 show s t h e p e r c e n t a g e o f e a c h s p e c i e s f l o w e r i n g e a c h

m o n th . ^P. s p e c i o s a h ad a m a jo r p e a k i n t h e m id d le o f e a c h y e a r

and a m in o r p e a k a t t h e end o f e a c h y e a r . H ow ev er , o n l y one

t r e e (3*2%) f l o w e r e d f o u r t i m e s d u r i n g t h e s t u d y ; e i g h t ( 2 5 . 8 %)

f l o w e r e d t h r e e t i m e s and t w e n ty - o n e (67*7%) f l o w e r e d t v / i c e .

One (3 .2% ) f l o w e r e d once and non e f a i l e d t o f l o w e r a l t o g e t h e r .

( T h e re was a ga p i n t h e r e c o r d o f one t r e e ; I t h a s b e e n e x c lu d e d

from t h e a n a l y s i s . ) At l e a s t one t r e e was i n f l o w e r e v e r y m o n th .

£ • J a v a n i c a h ad o n ly two f l o w e r i n g p e a k s d u r i n g

t h e s t u d y , i . e . from S e p te m b e r t o November e a c h y e a r a n d , t h e r e ­

f o r e , o u t o f p h a s e w i t h t h e P . s p e c i o s a t r e e s . T h e re was a gap

i n t h e r e c o r d o f one t r e e and a n o t h e r was c u t down d u r i n g t h e

s t u d y . D i s r e g a r d i n g t h e s e two t r e e s , t h r e e t r e e s (1 3 .6 % ) f a i l e d

F i g . 2 9 . G raph sh o w in g t h e % o f 2k- m arked P a r k i a . j a v a n i c a

and 32 m arked P a r k i a s p e c i o s a t r e e s f l o w e r i n g and

c h a n g in g l e a f e a c h month from J u n e 1972 t o J a n u a r y

197^* D a ta c o l l e c t e d a l o n g t h e T a p a h - R i n g l e t r o a d ,

P e r a k .

I- - - - - - - - - - - 1J F

A - A P. j a v a n i c a

© © P . s p e c i o s a

i r i i i 11 | iJ A S O N q J

1974

▲

1974

70

t o f l o w e r t h r o u g h o u t t h e s t u d y ; e i g h t e e n ( 8 1 . 8 %) f l o w e r e d i n •

1972 b u t o n l y t e n (45*5%) f l o w e r e d i n 1 9 7 3 . O n ly one t r e e t h a t

f l o w e r e d i n 1973 f a i l e d t o do so i n 1 9 7 2 . P . , j a v a n i c a was n o t

fo u n d i n f l o w e r e v e r y m o n th .

O b s e r v a t i o n s i n S e l a n g o r c o n f i r m e d t h a t t h e

f l o w e r i n g s e a s o n s o f b o t h s p e c i e s was v e r y p o o r i n 1 973 WhGft-'

5 4 . 5 % o f t h e n u m b ered P . j a v a n i c a t r e e s on t h e T a p a h - R i n g l e t

r o a d f a i l e d t o f l o w e r a t a l l . Many t r e e s o f b o t h s p e c i e s t h a t

d i d f l o w e r i n 1973 p r o d u c e d o n ly a h a n d f u l o f i n f l o r e s c e n c e s .

The p a u c i t y o f f l o w e r s was n o t c o n f i n e d t o t h i s r e g i o n b u t was

v e r y n o t i c e a b l f e i n S e l a n g o r w h e re e . g . t h e P . s p e c i o s a t r e e a t

S "t Jt h e 21 m i l e , U lu Gombak w h ic h f l o w e r e d e v e r y y e a r from 1961

t o 1969 ( M e d w a y 1 972c) and i n 1972 ( p e r s . o b s . ) f a i l e d t o

f l o w e r a s d i d t h e n e i g h b o u r i n g t r e e s i n 1 9 7 3 .

B o th P . s p e c i o s a and P . j a v a n i c a a r e d e c i d u o u s

b u t t h e y grow new l e a v e s a s so o n a s t h e o l d o n e s a r e s h e d .

Among t h e nu m bered t r e e s on t h e T a p a h - R i n g l e t r o a d , p e a k s i n

t h e number o f b o t h s p e c i e s c h a n g in g l e a f o c c u r r e d i n F e b r u a r y

and O c to b e r 1 9 7 3 ; l e s s t h a n h a l f t h e t r e e s w e re i n v o l v e d e a c h

t i m e . T h e re was a l s o a s m a l l p e a k i n t h e num ber o f P . j a v a n i c a

t r e e s c h a n g in g l e a f i n O c to b e r 1 9 7 2 . T h u s , l e a f c h a n g e d i d n o t

c o i n c i d e w i t h f l o w e r i n g o f P . s p e c i o s a b u t a few P . j a v a n i c a

t r e e s w ere i n f l o w e r and w i t h o u t l e a v e s s i m u l t a n e o u s l y . I n d i v i d u a l

t r e e s showed c o n s i d e r a b l e v a r i a t i o n i n t h e f r e q u e n c y o f l e a f l o s s ;

among 22 P . j a v a n i c a 13 (5 9 .1 % ) t r e e s l i s t t h e i r l e a v e s once

d u r i n g t h e s t u d y , 7 (31 .8% ) t w i c e an d 2 (9 .1% ) t h r i c e ; among

£ • s p e c i o s a 13 (41 .9% ) l o s t t h e i r l e a v e s o n c e , 1 6 (51 .6% )

tw ice and 2 (6 ,k% ) t h r i c e .

Leaf l o s s r e c o r d s i n S e langor c o i n c i d e d w i th th o se

i n P e rak .

There were fou r f lo w e r in g specimens and- t e n

f r u i t i n g spec imens of _P_. s i n g u l a r i s i n the h e rb a r iu m o f the

F o r e s t kesearc 'h I n s t i t u t e , Kepong. The c o l l e c t i o n d a t e s and

l o c a l i t i e s a r e a s f o l lo w s

Flow er ing December 1 S e la n g o r

F eb ru a ry 2 P erak ( 1 ) , Jo h o re (1 )

March 1 J o h o re

*+ «

F r u i t i n g J a n u a ry 1 S e la n g o r

F eb ru a ry 2 Perak ( 1 ) , S e la n g o r (1)

March 3 P erak ( 1 ) , S e la n g o r ( 1 ) , Pahang

A p r i l 3 P e rak (1 ) , J o h o re ( 1 ) , ? (1)

May l S e la n g o r

1 0 •

These r e c o r d s s u g g e s t a f lo w e r in g season from

December to March i n most of P e n i n s u l a r M a la y s ia . Th is was

a t th e time when ve ry few P . s p e c i o s a or P. . javanica were

f lo w e r in g i n S e la n g o r and Pahang d u r in g 1 972/73 and 1973/7*+.

6 . M y r t a c e a e .

One g e n u s , E u g e n i a , was shown t o ' c o n t a i n a t l e a s t

one c h i r o p t e r o p h i l o u s s p e c i e s , E. m a l a c c e n s i s , h u t i t p r o b a b l y

c o n t a i n s o t h e r c h i r o p t e r o p h i l o u s s p e c i e s t o o . E . jam bos L . ,

w h ic h h a s l a r g e , w h i t e , f r a g r a n t , n o c t u r n a l f l o w e r s , s u p e r ­

f i c i a l l y r e m i n i s c e n t o f S o n n e r a t i a a l b a ( S o n n e r a t i a c e a e ) d id

no a t t r a c t b a t s , ( D u r i n g one .hour o f o b s e r v a t i o n no b a t s

v i s i t e d a t r e e i n f u l l b lo o m , o n ly 50 rn from a f l o w e r i n g

D u r io z d b e t h i n u s w h ic h a t t r a c t e d many f l o c k s o f E . s n e l a e a

d u r i n g t h e same t i m e . )

6 . i . E u g e n ia m a l a c c e n s i s .

E c o l o g i c a l d i s t r i b u t i o n .

A common v i l l a g e t r e e t h r o u g h o u t t h e lo w la n d s

o f S e l a n g o r ; . i t was a l s o s e e n i n P e r a k , N e g r i S e m b i la n and

M a la c c a and i t i s p r o b a b l y grown e l s e w h e r e . I t g row s i n f u l l

s u n l i g h t , i t s d e n s e ca n o p y c a s t i n g a .d e e p shadow u n d e r w h ic h

v e r y l i t t l e e l s e g r o w s .

M o r p h o lo g y ( F i g . 3 9 ) .

T h i s s p e c i e s h a s b e e n d e s c r i b e d b y C o r n e r ( 1 9 5 2 ) .

The f l o w e r s w h ic h a r e c l u s t e r e d on t h e b r a n c h e s b e h i n d t h e

l e a v e s , a r e o f a d e e p a r e d c o l o u r .

N e c t a r .

No d a t a was o b t a i n e d .

P o l l e n . ( F i g . 36)

The p o l l e n i s t y p i c a l o f t h e M y r t a c e a e w h ic h i s

a v e r y s t e n o p a l y n o u s f a m i l y ( E r d tm a n , 1 9 5 2 ) . The l e n g t h o f t h e

73

p e r p e n d i c u l a r j o i n i n g one a n g l e t o t h e o p p o s i t e s i d e was 2 7 .2 ju .

I t s g r o s s t e x t u r e was d r y , a l m o s t p o w d e ry .

S c e n t .

N o t s t r o n g , s l i g h t l y f r a g r a n t .

F l o w e r i n g b i o l o g y .

F l o w e r s w ere n o c t u r n a l , l a s t i n g one n i g h t o n l y .

The h e a v y s e p a l s t h a t p r o t e c t e d t h e hud p a r t i a l l y op en ed t h e

day b e f o r e a n t h e s i s t o r e v e a l t h e f o l d e d s t a m e n s b e l o w . O th e r

d e t a i l s o f t h e o p e n in g o f t h e f l o w e r s w e re n o t o b s e r v e d . The

m o rn in g a f t e r a n t h e s i s , t h e s t a m e n s f e l l o f f a t a n a b s c i s s i o n

l a y e r fo rm ed a t t h e i r b a s e s .

V i s i t o r s b e s i d e s m a c r o g l o s s i n e b a t s .

The f r u i t b a t , C y n o p te r u s b r a c h y o t i s , d e v o u re d

a l m o s t a l l t h e f l o w e r s p r o d u c e d on t r e e s a t R a n ta u P a n ja n g

b u t no o t h e r a n i m a l s w ere r e c o r d e d v i s i t i n g t h e f l o w e r s .

P h e n o l o g y .

The t r e e s w ere n o t d e c i d u o u s . I n S e l a n g o r t h e r e

w e re s h o r t f l o w e r i n g s e a s o n s l a s t i n g f r o m ' t e n ' d a y s t o two w eeks

i n March an d A u g u s t 1972 a n d 1 9 7 3 .

7 . R h i z o p h o r a c e a e .

The p o l l e n o f one g e n u s , R h iz o p h o r a ( a l l m angrove

t r e e s ) , w as v e r y r a r e i n t h e gu an o o f E o n y c t e r i s s p e l a e a

c o l l e c t e d from B a tu C aves a nd i n f a e c e s o f M a c r o g l o s s u s min im us

c a u g h t a t R a n ta u P a n j a n g .

N o te , R h i z o p h o r a ' m u c r o n a ta Lam. P o l l e n i d e n t i f i e d b y D r . J . M u l l e r ,

R e j k s h e r b a r i u m , L e i d e n . F lo w e r s w e re o f t e n w a tc h e d b u t no b a t s

w e re s e e n t o v i s i t th e m . F r e s h p o l l e n was n o t b r o k e n down by

a c e t o l y s i s an d g r a i n s fo u n d i n guan o showed no s i g n s o f

d i s i n t e g r a t i o n . S i n c e R h iz o n h o r a s p p . v/ere v e r y common i n t h e

m a n g ro v e s a t R a n ta u P a n j a n g a n d e l s e w h e r e o f f t h e c o a s t o f

S e l a n g o r and s i n c e th e y , f l o w e r e d f r e e l y , R h iz o p h o r a i s , t h e r e f o r e ,

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

R h iz o p h o r a h a s b e e n d e s c r i b e d b y many a u t h o r s

b u t t h e m o s t r e l e v a n t a c c o u n t t o M a l a y s i a i s t h a t o f W atson (1 92 8 )

He g i v e s an e x c e l l e n t d e s c r i p t i o n o f t h e e c o l o g i c a l r e q u i r e m e n t s

o f t h e s p e c i e s a s w e l l a s t h e i r m o r p h o lo g y .

R h i z o p h o r a p o l l e n h a s b e e n d e s c r i b e d by

E rd tm an ( 1 9 5 2 ) . The g r a i n s a r e v e r y s m a l l . No m e a s u re m e n ts

a r e a v a i l a b l e b u t i n s i z e t h e y a r e a p p r o x i m a t e l y e q u a l t o o r

s l i g h t l y s m a l l e r t h a n t h o s e o f A r t o c a r p u s s p p . ( M o r a c e a e ) .

8 . S o n n e r a t i a c e a e .

Two g e n e r a i n P e n i n s u l d r M a l a y s i a ; S o n n e r a t i a

(3 s p e c i e s , S . a l b a , S . c a s e o l a r i s (L) and S . o v a t a ) and

D uabanga (1 s p e c i e s , D. g r a n d i f l o r a ; a s e c o n d s p e c i e s ,

D. m o l u c c a n a , g row s i n B o r n e o ) . A l l a r e c h i r o p t e r o p h i l o u s .

8 . i . S o n n e r a t i a .

Taxonomic n o t e . N o m e n c la tu r e f o l l o w s B a c k e r an d v an S t e e n i s (1 9 5 4 )

I n h i s w e l l known b o o k on M alayan m a n g r o v e s , W atso n (1 9 2 8 )

c o n f u s e d t h e n o m e n c l a t u r e o f t h e s p e c i e s . H is e r r o r s h a v e

b e e n i n c o r p o r a t e d i n t o many s u b s e q u e n t p u b l i c a t i o n s . The

75

f o l l o w i n g t a b l e e q u a t e s t h e names u s e d h e r e w i t h t h o s e u s e d

b y W a ts o n .

B a c k e r and van S t e e n i s , 1954

S . a l b a

S . o v a t a

S . c a s e o l a r i s

E c o l o g i c a l d i s t r i b u t i o n .

A l l t h r e e s p e c i e s a r e m angrove t r e e s . S . a l b a

g ro w s i n t h r e e e c o l o g i c a l s i t u a t i o n s , e a c h i n d u c i n g a d i f f e r e n t

g ro w th fo r m . F i r s t l y and m os t commonly, i t i s a p i o n e e r s p e c i e s

on a c r e t i n g s h o r e s a l o n g t h e w h o le S e l a n g o r c o a s t , i n c l u d i n g

t h e o f f - s h o r e i s l a n d s , e x c e p t w h e re i t i s s o m e t im e s r e p l a c e d

A v i c e n n i a a l b a ( F i g , 3 0 ) . (The u s e o f m o to r d r i v e n b o a t s ,

e s p e c i a l l y a ro u n d P o r t K la n g a nd t h e i s l a n d s , h a s e r a d i c a t e d

S . a l b a from many a r e a s b e c a u s e t h e bow w aves from t h e b o a t s

wash away t h e mud b a n k s , t h u s u n d e r c u t t i n g t h e r o o t s o f t h e

t r e e s . )

I n t h i s s i t u a t i o n i t fo rm s a n a r r o w s t r i p b e tw e e n

t h e s e a an d t h e more e x t e n s i v e s t a n d s o f o t h e r m a n g ro v es e . g .

B r u g u t g r a s p p . an d A v i c e n n i a s p p . T h i s s t r i p e x t e n d s up t h e

m angrove c r e e k s an d t h e lo w e r t i d a l r e a c h e s o f r i v e r s w h e r e v e r

s u i t a b l e mud bank s ' Occur* e . g , w h e re r i v e r s c h a n g e d i r e c t i o n s

i t i s o f t e n p r e s e n t on t h e i n s i d e o f t h e b e n d s w h e re mud b a n k s

W a ts o n , 1 928

S . g r i f f i t h i

S . a l b a

S . a c i d a

F ig . 30. S on n era tia a lb a .

a. Narrow strip of small trees growing on the bank of the Klang river near its mouth.Note:- 1. the stilt roots of Rhizophora spp. just

behind the S. alba trunks.2 . the high tide level.

b. Grove of mature trees growing at Telok Asam, Bako National Park. Photographed at low tide.

I ■ '

' . ! ■ '

76

b u i l d u p . The t r e e s w e re s m a l l , up t o 10 m h i g h b u t u s u a l l y

l e s s , w i t h n a r r o w c ro w n s an d l e a f y tw ig s from b e lo w o r a t

h i g h t i d e l e v e l u p w ard s ( F i g . 3 0 ) . R e g e n e r a t i o n t a k e s p l a c e

on t h e s e a w a rd s i d e o f t h e s t r i p and t h e o l d e r t r e e s a r e

u s u a l l y o v e r r u n by t h e a d v a n c e o f t h e s p e c i e s g ro w in g i m m e d i a t e l y

b e h i n d th em .

S e c o n d l y , an d v e r y r a r e l y , a t r e e t h a t was o n ce

p a r t o f a p i o n e e r s t r i p w i t h s t a n d s t h e a d v a n c e o f t h e o t h e r

m angrove s p e c i e s an d d e v e l o p s i n t o a hu ge e m e r g e n t , 2 0 t o 3 0 m

t a l l , w i t h a s t r a i g h t t r u n k . B r a n c h e s a r e m o s t l y ’c o n f i n e d t o

t h e u p p e r p a r t o f t h e t r u n k , a b o v e t h e c an o p y o f t h e o t h e r

s p e c i e s . As t h e m ang ro ve f o r e s t a d v a n c e s , s u c h t r e e s become

s t r a n d e d i n c r e a s i n g l y f a r from t h e s e a f r o n t .

T h i r d l y , a t M o r ib , S e l a n g o r and i n Bako N a t i o n a l

P a r k , S a ra w a k l a r g e m a t u r e S . a l b a t r e e s form e x t e n s i v e mono-

s p e c i f i c s t a n d s ( F i g s . 8 and 3 0 ) . I n b o t h s i t u a t i o n s , t h e

s t a n d s a r e open t o t h e s e a on one s i d e and t h e y grow on mud

f l a t s w h ic h a r e c o v e r e d b y a t l e a s t 1 m and o c c a s i o n a l l y ,

( e s p e c i a l l y a t Bako) 2 m o r m<?re o f w a t e r a t mean h i g h t i d e .

The t r e e s i n t h e s e f o r e s t s grow t o a b o u t 1 5 m h i g h , w i t h r o u n d e d

c ro w n s . They a r e w i d e l y s p a c e d , t h e t r u n k s a r e g n a r l e d and

m a jo r b r a n c h e s d e v e l o p a t a b o u t t h e h ig h t i d e l e v e l .

S . c a s e o l a r i s i s a l s o a p i o n e e r , g ro w in g on t h e

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

r i v e r s . I t was n e v e r s e e n g ro w in g on t h e s e a f r o n t . On t h e

K la n g r i v e r , i t i s a b u n d a n t fro ln B a tu "Tig’a" ' tb"K iang ' ( F i g s . 3 2 . and 6 ?)

b u t t h e r e a f t e r , i t i s g r a d u a l l y r e p l a c e d by S . a l b a s o t h a t

F i g . 31• S o n n e r a t i a ov a ta on th e bank o f a sm a l l r i v e r n e a r Kemaman, Trengganu. Note th e f e r n s , g r a s s and Pandanus s p p . growing around i t .

F i g . 32 . S o n n e r a t i a c a s e o l a r i s growing on th e banks o f th e Klang r i v e r a t Rantau P a n ja n g . Note th e s c a n t l e a f and pendu lous t w i g s .

77

a t R a n ta u P a n j a n g b o t h s p e c i e s o c c u r r e d i n a p p r o x i m a t e l y e q u a l

n um b ers b u t n e i t h e r g row s w e l l and few grow d o w n s tr e a m o f

R a n ta u P a n j a n g . A t B a tu T ig a t h e w a t e r i s f r e s h ; t h e t i d a l

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

o f r i v e r w a t e r u n a b l e t o f lo w o u t t o t h e s e a when t h e t i d e

^D9 'Ho apex's* cciiiEi•) * S • c o c G o lo x i s £rcy ,r2 w e l l li x* o i n

a s s o c i a t i o n w i t h f r e s h w a t e r p l a n t s , e . g . t h e w a t e r h y a c i n t h ,

E i c h h o r h i a c r a s s i p e s ( P o n t e d e r i a c e a e ) .

iL* o v a ' a was o n l y fo u n d on t h e la n d w a r d m a r g in s

o f t h e m a n g r o v e s . On t h e w e s t c o a s t o f P e n i n s u l a r M a l a y s i a

t h e r e , a r e e x t e n s i v e m angrove f o r e s t s b u t t h e la n d w a r d m a r g in s

h a v e f r e q u e n t l y b e e n r e c l a i m e d a nd S . o v a t a h a s b e e n e r a d i c a t e d

from m o s t a r e a s . A few t r e e s w e re fo u n d g ro w in g n e a r t h e dyke

on b o t h b a n k s o f t h e r i v e r K la n g a t R a n ta u P a n j a n g . On t h e

e a s t c o a s t , w h e re m a n g ro v e s m o s t l y grow i n n a r r o w f r i n g e s a l o n g

th e r i v e r s , t h e r e h a s b e e n l e s s c l e a r a n c e and t h i s s p e c i e s was

fo u n d more commonly on t h e m a r g in s o f t h e m a n g ro v e s f r e q u e n t l y

g ro w in g i n a s s o c i a t i o n w i t h t e r r e s t r i a l p l a n t s s u c h a s g r a s s

and P a n d a n u s ( F i g . 3 ) .

Mo r p h o l o g y , ( p i g . 3 9 ) .

A l l t h r e e s p e c i e s h a v e b e e n d e s c r i b e d b y B a c k e r

and va n S t e e n i s (195^+, 1972) and W atson ( 1 9 2 8 ) . The m o rp h o lo g y

o f h y b r i d s o f S . a l b a and S . o v a t a a s w e l l a s S . a l b a and

c a s e o l a r i s w e re d e s c r i b e d by M u l l e r and H o u -L iu ( 1 9 6 6 ) .

The f l o w e r s o f a l l t h r e e s p e c i e s a r e v e r y s i m i l a r t o one a n o t h e r

i n s i z e and i n g r o s s m o r p h o lo g y ( F i g . 3 3 ) b u t t h e y d i f f e r i n

c o l o u r ; S . a l b a and S . o v a t a h a v e t h e s t a m e n s e n t i r e l y w h i t e

/I

Fig. 33.

L.S. OF GENERALI SED

S O N N E R A T I A FLOWER

IN AN THESIS

( X 2 ) /

n e c t a r y

S O N N E R A T I A SPP. BUDS ( x i )

A d e v e l o p i n g (whol e )

B " (I.s.)C o n t h e d a y p r e c e e d i n g

a n t h e s i s (I.s.)

■ ■ 78

:?.• c a s e o l a r i s h a s t h e p r o x i m a l h a l f o f t h e s t a m e n s r e d

( F i g , 34 ) • i n S . c a s e o l a r i s t h e f l o w e r s a r e t e r m i n a l on lo n g

p e n d u l o u s t w i g s , i n t h e o t h e r two s p e c i e s t h e y a r e t e r m i n a l

b u t on s h o r t e r , l e s s f l e x i b l e t w i g s .

H y b r i d s o f S . a l b a an d S . o v a t a fo u n d a t Kampong

D e ie k ( s i t e 1 0 , F i g . 3 ) w ere i n t e r m e d i a t e i n a l l f e a t u r e s

b e tw e e n t h e i r p a r e n t s p e c i e s c o n f i r m i n g M u l l e r an d H o u -L iu * s

(1 9 6 6 ) o b s e r v a t i o n s .

N e c t a r .

N e c t a r was e x t r a c t e d from t w e lv e f l o w e r s o f

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

u n t i l 2 0 .0 0 h r s on t h e n i g h t o f a n t h e s i s when t h e volum e o f

n e c t a r i n e a c h w as m e a s u re d t o 0 .1 m l . The v o lu m e s r e c o r d e d

w e re S , a l b a 0 . 2 t o 0 .^ ^ (m e a n 0 . 3 4 + 0 . 0 6 m l ) ; S , o v a t a 0 . 2

t o 0 .4 ml (mean 0 . 3 2 +. 0 . 0 7 m l ) ; S . c a s e o l a r i s 0 .1 t o 0 . 4 ml

(mean 0 . 3 0 + 0 . 0 8 m l ) . A f t e r t h e n e c t a r was e x t r a c t e d , t h e

c l o t h b a g s w ere r e p l a c e d . M ost o f t h e f l o w e r s w e re e xam ined

a g a i n a t 2 2 . 0 0 h r s b u t no m e a s u r e a b l e volum e o f n e c t a r was

o b t a i n e d from a n y . No f u r t h e r e x a m i n a t i o n s w e re m ade.

P o l l e n ( F i g . 3 8 ) .

The p o l l e n o f a l l t h r e e s p e c i e s an d t h e h y b r i d s

o f two o f them h a s b e e n ve ry , f u l l y d e s c r i b e d b y M u l l e r ( 1 9 6 9 ) .

M e a s u re m e n ts o f ' 20 g r a i n s o f e a c h s p e c i e s c o l l e c t e d a t R a n ta u

P a n j a n g an d m o u n ted i n P .V .A . w ere

Mean d i a m e t e r

5 4 . 3 jn

5 2 . 0 /x

5 4 . 3 / i

Mean l e n g t h

S- a l b a 8 7 . 0 fi

£* o v a t a 7 7 . 4 p

S_. c a s e o l a r i s 7 6 , 1 ^

a . S o n n e r a t i a c a s e o l a r i s

b . S o n n e r a t i a a l b a .

79

S c e n t .

D i s t i n c t i v e , r e m i n i s c e n t o f s o u r m i l k b u t n o t

p a r t i c u l a r l y s t r o n g t o t h e human s e n s e o f s m e l l . I was n o t

a b l e t o s m e l l f l o w e r s h e l d more t h a n 50 cm away from my n o s e

an d was u n a b l e t o d e t e c t a f l o w e r i n g t r e e by s m e l l a l o n e .

F lo w e r i n g b i o l o g y .

A l l t h r e e s p e c i e s b e h a v e s i m i l a r l y . I n t h e v e r y

young b u d s t h e s t i g m a and s t a m e n s a r e s t r a i g h t b u t t h e y e l o n g a t e

a s t h e b ud m a t u r e s , t h e fo r m e r b e co m in g f o l d e d i n t o t h e a p e x

o f t h e b ud w h i l e t h e l a t t e r b en d d o u b le ( F i g . 5 3 ) • B e tw een

m i d n i g h t and 0 3 . 0 0 h r s on t h e d ay p r e c e d i n g a n t h e s i s , t h e c a l y x

l o b e s s p l i t an d t h e s t i g m a s t r a i g h t e n ^ b u t t h e s t a m e n s r e m a in

b e n t i n w a r d s ( F i g . 3 3 ) . D u r in g t h e a f t e r n o o n p r e c e d i n g a ' n t h e s i s ,

t h e s t a m e n s s l o w l y u n f o l d b u t t h e a n t h e r s r e m a in h i d d e n u n t i l

a t d u sk ( i . e . b e tw e e n 19*00 and 19*^5 h r s ) t h e y s u d d e n l y b u r s t

o u t a s t h e s t a m e n s s t r a i g h t e n ( F i g . 33 ) • The f i n a l o p e n in g

i s v e r y r a p i d an d s p e c t a c u l a r . P o l l e n i s n o t r e l e a s e d u n t i l

t h i s moment. ( A n t h e r s rem o ved 10 m i n u te s p r i o r t o t h e f i n a l

o p e n in g h ad n o t d e h i s c e d b u t t h o s e l e f t i n t a c t on t h e f l o w e r

h ad done so w i t h i n two o r t h r e e m i n u t e s o f t h e f l o w e r o p e n i n g . )

A n t h e s i s l a s t s o n ly one n i g h t . E ach a n t h e r fo rm s

an a b s c i s s i o n l a y e r a t i t s b a s e and b r e a k s o f f o r i s d i s l o d g e d

b y a n i m a l s a r o u n d dawn.

A t R a n ta u P a n ja n g (w h e re e v e r y S o n n e r a t i a s p p .

f l o w e r t h a t was w a tc h e d , e i t h e r d u r i n g an e x p e r i m e n t ( s e e p . 110

o r i n c i d e n t a l l y w h i l e w o rk in g n e a r b y , was v i s i t e d by b a t s )

+8 (96%) o f 51 m ark ed £>. a l b a f l o w e r s s e t f r u i t an d f a l l e n

80

f l o w e r h e a d s o f any s p e c i e s w e re v e r y r a r e . I n a n o t h e r e x p e r i m e n t

t h e s t i g m a was rem oved from t e n ,3. a l b a f l o w e r s d u r i n g t h e

a f t e r n o o n p r e c e d i n g a n t h e s i s . None h a d f a l l e n a f t e r f i v e d a y s

b u t a f t e r e l e v e n d a y s n i n e h a d f a l l e n and. t h e t e n t h had fo rm ed

an a b s c i s s i o n l a y e r an d f e l l when h a n d l e d .

At Bako w here many o f t h e f l o w e r s w ere a p p a r e n t l y

n o t v i s i t e d by b a t s ( s e e p . 110) , f a l l e n f l o w e r h e a d s o f S . a l b a

w e re v e r y common on t h e mud b e n e a t h t h e t r e e s an d w ere w ashed

up a l l a l o n g t h e s h o r e s a t h i g h t i d e . The c a l y c e s o f t e n S . a l b a

f l o w e r s m arked d u r i n g a n t h e s i s a t Bako on 9 J a n u a r y 1973 w ere

s t i l l i n s i t u on 13 J a n u a r y ( f o u r d a y s l a t e r ) b u t s e v e n o u t

o f t e n m arked on 9 May 1973 h a d f a l l e n by 16 May (7 d a y s l a t e r ) .

The r e s u l t s o f t h e e x p e r i m e n t i n v o l v i n g t h e r e m o v a l o f t h e

s t i g m a o f S . a l b a f l o w e r s a t R a n ta u P a n ja n g m u s t b e k e p t i n

mind when a s s e s s i n g t h e d a t a a t B ak o . The p e r i o d o f o b s e r v a t i o n

i n J a n u a r y (i+ d a y s ) was p r o b a b l y t o o s h o r t f o r a n y c o n c l u s i v e

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

V i s i t o r s b e s i d e s m a c r o g l o s s i n e b a t s .

Most o b s e r v a t i o n s r e f e r t o S . a l b a . B e f o r e a n t h e s i s

v i s i t o r s w ere r a r e . C r e p u s c u l a r s k i p p e r s ( H e s p e r i i d a e ) f r e q u e n t l y

l a n d e d on t h e c a l y x o f m a tu re b u d s t h a t w ere a b o u t t o open ( a s

i n F i g . 33 ) an d i n s e r t e d t h e i r p r o b o s c e s b e tw e e n t h e s t a m e n s

t o o b t a i n n e c t a r . They n e v e r , damaged t h e f l o w e r s i n an y o b v io u s

way and w ere u n l i k e l y t o e f f e c t p o l l i n a t i o n o r t o e x t r a c t much

n e c t a r . D u r in g a n t h e s i s , s m a l l , u n i d e n t i f i e d - m o th s ( L e p i d o p t e r a )

l a n d e d i n t h e same way and i n s e r t e d t h e i r p r o b o s c e s b e tw e e n

t h e b a s e s o f t h e a n t h e r s t o o b t a i n n e c t a r . T h ey , a l s o , w ere

u n l i k e l y t o h a v e e f f e c t e d p o l l i n a t i o n an d w e re n o t fo u n d t o

damage t h e . f l o w e r s . %

On t h e m o rn in g f o l l o w i n g a n t h e s i s , s u n b i r d s ,

e s p e c i a l l y t h e brow n t h r o a t e d - s u n b i r d ( A n t h r e p t e s m a l a c c e n s i s

( S c o p ) , N e c t a r i n i d a e ) , a common s p e c i e s i n t h e m a n g r o v e s ,

■Pv'A/in ttrt *tr tr*? <•.«•? 4* n -P”] HIV) v» n n tf*' 4 4 inT n J ma 4 «** v v j U v u w x j J X u l w v u vnvy AXWHOA U« i-UOXJ. CXV U-L. V J_ U Jf UXOXUUgjOU CUJ.jf

s t a m e n s t h a t h ad n o t y e t f a l l e n , ^'hey l a n d e d on t h e p e d i c e l

and i n s e r t e d t h e i r b i l l s b e tw e e n t h e c a l y x l o b e s i T h u s , t h e y

w e re u n l i k e l y t o h a v e come i n t o c o n t a c t w i t h t h e s t i g m a b u t

t h e y c o u l d c o n c i e v a b l y t r a n s f e r p o l l e n from one f l o w e r t o t h e

s t i g m a o f a n o t h e r . The s u n b i r d s w e re a l s o s e e n t r y i n g t o p r o b e

t h e u n o p en ed b u d s on t h e d ay p r e c e d i n g a n t h e s i s .

A f t e r t h e s t a m e n s h ad b e e n d i s l o d g e d , i n s e c t s ,

e s p e c i a l l y m e l a p o n id b e e s an d a wasp ( V espa t r o p i c a L . , V e s p i d a e ) ,

f r e q u e n t l y s o u g h t n e c t a r i n t h e c a l y x t u b e . They d i d n o t come

i n c o n t a c t w i t h t h e s t i g m a . The a n t s , O ec o p h .y l la s m a r a g d in a L . ,

w h ich a b o u n d e d on a l m o s t e v e r y S o n n e r a t i a t r e e , w ere u s u a l l y

fo u n d " g u a r d i n g " t h e c a l y x , p e d i c e l o r tw ig l e a d i n g t o them

b u t t h e y w ere n e v e r s e e n t o v e n t u r e among t h e a n t h e r s even

when p r o v o k e d .

P h e n o l o g y .

S o n n e r a t i a a l b a .

S . a l b a f l o w e r s i n f l u s h e s . a n d f l o w e r i n g i n

m o s t t r e e s i n one a r e a was s y n c h r o n i s e d . New l e a f g ro w th

o c c u r s b e tw e e n f l o v / e r i n g f l u s h e s b u t t h e t r e e s a r e n e v e r

d e c i d u o u s . O ld l e a v e s a r e l o s t c o n t i n o u s l y b u t e s p e c i a l l y

d u r i n g f l o w e r i n g f l u s h e s . A t R a n ta u P a n j a n g , r e c o r d s o f t h e

82

d u r a t i o n o f f l u s h e s w ere m a i n t a i n e d t h r o u g h o u t t h e s t u d y

( T a b l e 3 ) ♦. M ost o b s e r v a t i o n s w ere made a t s i t e s 1 , 2 a n d 10

( F i g . 3 ) . A f l u s h was d e f i n e d a s s t a r t i n g on t h e f i r s t d a t e

r e c o r d e d on w h ic h two o r more f l o v / e r s w ere o pen on two o r more

£ r e e s a t a n y o f . t h e s e s i t e s an d when m os t o r a l l t h e r e m a i n i n g

1*i1 o 0 s h-?„d. b u d s a p p r o a c h i n g m a t u r i t y v Xt was r o c o r d o d a s 0 n d 0 d

when t h e a b o v e c o n d i t i o n s no l o n g e r a p p l i e d . S i n c e e x c u r s i o n s

t o R a n ta u P a n j a n g w ere n o t made d a i l y and s i n c e , i n p r a c t i c e ,

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

n e c e s s a r i l y an a p p r o x i m a t i o n o f t h e d u r a t i o n o f f l u s h e s .

I r r e g u l a r s p o t c h e c k s a t o t h e r l o c a l i t i e s showed

t h a t t h e s y n c h r o n i s a t i o n o f f l o w e r i n g f l u s h e s e x t e n d e d a l o n g

t h e w h o le S e l a n g o r c o a s t from C a re y I s l a n d , s o u t h o f K la n g ,

t o K u a la S e l a n g o r i n t h e n o r t h ( F i g . 2 ) . B eyond t h i s , i n s u f f i c i e n t

d a t a was o b t a i n e d t o show how f a r t h e t r e e s f f l o w e r e d t o g e t h e r .

T a b le 3 show s t h a t t h e d u r a t i o n o f t h e f l u s h e s

l a s t e d a l i t t l e o v e r a month and t h e r e w ere f o u r f l u s h e s i n

b o t h y e a r s . The l o n g e s t f l u s h , f rom 17 A p r i l t o 1 J u l y 1 9 7 2 ,

v/as e x c e p t i o n a l b e c a u s e d u r i n g t h e two v e r y , d r y m o n th s o f May

and J u n e t h a t y e a r ( F i g . i|. ) t h e g ro w th o f many o f t h e b u d s

v/as R e t a r d e d . F l o w e r i n g t h r o u g h o u t t h a t f l u s h was l i g h t and

s p o r a d i c .

I n J a n u a r y an d May 1 9 7 3 , a l l t h e t r e e s a t T e lo k

Asam, Bako N a t i o n a l P a r k , v/ere i n s y n c h r o n y , i . e . i n t h e m id d le

o f a f l o w e r i n g f l u s h i n J a n u a r y and n e a r t h e end o f a f l u s h .

i n May. T h e se f l u s h e s v/ere o u t o f p h a s e w i t h t h o s e i n S e l a n g o r .

Oh a 3 Km b o a t j o u r n e y down t h e B o m b a la i r i v e r

Table 3«

Occurrence of flowering flushes of Sonneratia alba and S. ovata at Rantau Panjang, Selangor in 1972 and 1973*

S . a l b a f lo w e r in g S. a l b a n o t f lo w e r in g

F e b . t o 20 Mar. 20 Mar. to 17 A p r i l

17 A p r i l 11 1 J u l y 1 J u l y " 6 Aug.

1 972 6 Aug. 11 11 S e p t . 1 972 1 1 S e p t . " 30 O c t .

30 O c t . " k Dec. k Dec. " 5 Feb. 1973

5 Feb. to 1 Mar. 1 Mar. to 30 A p r i l

30 A p r i l " 18 June 1 8 June ** 1 6 J u l y1973 1973

16 J u l y " 20 Aug. 20 Aug. " 2 k S e p t .

2k S e p t . " 3 Dec. 3 Dec. " ------------

S . o v a t a f lo w e r in g S . o v a ta n o t f lo w e r in g

1 May to 6 June 6 June to 13 J u l y

13 J u l y " 28 S e p t . 28 S e p t . " 28 Nov.1972 1972

28 Nov. 11 ? (D ec . o r ? - »» 13 F e b . 1973(Jan .1 973)

1 3 F eb . to 17 Mar. 17 Mar. to 7 May

? May *' 25 June 25 June 11 13 Aug.1973 1973

13 Aug. " 15 S e p t . 1 5 S e p t . " 15 Nov.

1 5 Nov. M 15 Dec. ■ 15 Dec. M ? J a n . 197**

83

t o i t s m outh n e a r Tawau, S a b a h , on 28 December 1972 many S . a l b a

t r e e s w ere e x a m in e d . O t h e r s w e re s e e n on t h e Imam r i v e r i n t h e

same a r e a . No t r e e s w ere fo u n d i n f l o w e r b u t o l d f r u i t s w ere

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

f l u s h e s and w e re o u t o f p h a s e w i t h t h o s e o f Bako w h ic h w ere

f l o w e r i n g t h e n .

S o n n e r a t l a o v a t a .

L ik e S . a l b a , S . o v a t a f l o w e r s i n f l u s h e s and

m o s t l e a f g ro w th t a k e s p l a c e b e tw e e n f l u s h e s . On 7 November 1972

t h r e e t r e e s on t h e n o r t h b a n k o f t h e r i v e r K la n g a t R a n ta u

P a n j a n g h a d s h e d a l l t h e i r l e a v e s w h i l e many o f t h o s e a t

Kampong D e le k , on t h e s o u t h b a n k - o f t h e r i v e r , h a d l o s t m o s t

o l d l e a v e s b u t h a d many new l e a v e s d e v e l o p i n g . T h i s was th e

o n l y o c c a s i o n on w h ic h an y S o n n e r a t i a s p p . t r e e s w ere s e e n

l e a f l e s s . The l e a f l o s s o c c u r r e d a t t h e end o f one f l o w e r i n g

f l u s h an d g ro w th o f new l e a v e s was r e m a r k a b l y r a p i d , ( D e v e lo p i n g

f i ' u i t s r e m a in e d on t h e t r e e s . ) T o t a l l e a f l o s s may h av e e s c a p e d

n o t i c e a t o t h e r t i m e s b u t i t w as n o t a r e g u l a r o c c u r r e n c e

b e tw e e n e a c h f l u s h o f f l o w e r s .

The o c c u r r e n c e o f f l o w e r i n g f l u s h e s o f S . o v a t a

a r e g i v e n i n t a b l e 3 • The d a t a a r e b a s e d p r i m a r i l y on o b s e r v a t i o n s

a t Kampong D e le k ( s i t e 1 0 , F i g . 3 ) °n 'the s o u t h b a n k o f t h e

r i v e r K la n g o p p o s i t e t h e m ain s t u d y a r e a b u t a l s o on t r e e s on

t h e n o r t h b a n k ( s i t e 1 , F i g . 3 ) . The d e f i n i t i o n o f a f l o w e r i n g

f l u s h an d l i m i t a t i o n s o f t h e d a t a a r e t h e same a s f o r S . a l b a

( s e e a b o v e ) The f l u s h e s w ere s i m i l a r i n c h a r a c t e r t o t h o s e

a ^ha b u t u s u a l l y l a g g e d a l i t t l e b e h i n d t h a t s p e c i e s .

8it

H ow ever , m a j o r d i f f e r e n c e s o c c u r r e d when t h e f l u s h e s o f e i t h e r

s p e c i e s w ere p r o l o n g e d , e . g . t h e p r o l o n g e d f l u s h o f S . a l b a

from 17 A p r i l t o 1 J u l y 1973 (2-J m o n th s ) was n o t p a r a l l e M b y

S . o v a t a w h ic h b e g a n f l o w e r i n g two w eeks l a t e r on 1 May and

s t o p p e d on 6 J u n e ( 1 ^ m o n t h s ) . The r e v e r s e o c c u r r e d when

S . o v a t a f l o w e r e d from 13 J u l y t o 8 S e p te m b e r 1972 (2-j m o n th s )

and jS. a l b a f l o w e r e d from 6 A u g u s t t o 11 S e p te m b e r (1 + month)

t h a t y e a r .

S o n n e r a t l a c a s e o l a r i s .

On t h e r i v e r K la n g S . c a s e o l a r i s f l o w e r e d c o n t i n o u s l y ,

n o t i n f l u s h e s . On t h e v e r y b i g t r e e s ( up t o 15 m h i g h ) a t

K la n g a s many a s 17 o pen f l o w e r s w ere , c o u n t e d on one t r e e one

n i g h t b u t t h e r e w e re u s u a l l y from 5 t o 10 f l o w e r s . I n t h e more

s a l i n e a w a t e r a t R a n ta u P a n j a n g w h e re t h e t r e e s w e re s m a l l e r

(u p t o 3 m h i g h ) t h e y p ro d u c e d from one o r two f l o w e r s p e r

n i g h t t o a p p r o x i m a t e l y one f l o w e r p e r weelf, d e p e n d i n g on t h e

s i z e o f t h e t r e e , t h r o u g h o u t t h e y e a r . T h e re w e re m in o r

f l u c t u a t i o n s , e . g . so m e t im e s one t r e e m ig h t h a v e t h r e e f l o w e r s

p e r n i g h t an d n o n e f o r a n o t h e r t e n n i g h t s .

U p s t r e a m o f K la n g , t h e t r e e s r e m a in e d f a i r l y

l a r g e w i t h a l u x u r i a n t l e a f g ro w th b u t f l o w e r s becam e p r o g r e s s i v e l y

f e w e r . A t t h e B a tu T ig a b r i d g e , more t h a n 12 t r e e s ( a l l o v e r

7 m h i g h ) w e re ex am in e d s e v e r a l t i m e s b u t t h e o n l y e v i d e n c e

o f f l o w e r i n g was t h e p r e s e n c e once on one t r e e o f a h a l f

d e v e lo p e d f r u i t .

A l l S . c a s e o l a r i s t r e e s t h a t w e re e x am in ed e l s e ­

w h e re i n p e n i n s u l a r M a l a y s i a h a d b u d s an d f r u i t s i n a l l s t a g e s

o f d e v e lo p m e n t .

85

R e p r o d u c t i v e c o m p a t a b l l i t y .

'Seven o f t h e t e n S . a l b a f l o w e r s t h a t w ere

e x p e r i m e n t a l l y c r o s s p o l l i n a t e d s e t f r u i t b u t no n e o f t h o s e

t h a t w ere e x p e r i m e n t a l l y s e l f p o l l i n a t e d d i d s o . T h i s s t r o n g l y

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

8 . i i . D uabanga g r a n d i f l o r a .

E c o l o g i c a l d i s t r i b u t i o n .

I n S e l a n g o r , P e r a k , P ah an g an d N e g r i S e m b i la n

t h i s s p e c i e s i s v e r y common i n t h e v a l l e y s a l o n g t h e main

r a n g e . I t w as m o s t f r e q u e n t l y s e e n i n l o w la n d s e c o n d a r y f o r e s ts tb u t was a l s o fo u n d i n l o w la n d p r i m a r y f o r e s t ( e . g . 21 m i l e ,

U lu G om bak), a r o u n d t h e b a s e s o f l i m e s t o n e o u t c r o p s ( e . g . B a tu

C aves a nd Gunong G a ja h , n e a r Gua S a n d in g ) a n d , l e s s commonly,

i n m on tane f o r e s t s ( e . g . T a b le 1 ) . I t was n o t l o o k e d f o r i n

o t h e r s t a t e s .

M o rp h o log y ( F i g . 3 9 ) .

T h i s s p e c i e s h a s b e e n d e s c r i b e d and f i g u r e d by

B a c k e r and van S t e e n i s ( 1 9 5 ^ ) ; C o rn e r (1 9 5 2 ) g i v e s an a c c o u n t

o f i t i n P e n i n s u l a r M a l a y s i a and van d e r P i j l ( 1 9 3 6 , 1956)

d i s c u s s e s a s p e c t s r e l a t i n g t o c h i r o p t e r o p h i l y . The w h i t e f l o w e r s ,

w h ich a r e v e r y s i m i l a r t o S o n n e r a t j a e x c e p t i n t h e p r e s e n c e

o f l a r g e , w h i t e p e t a l s , a r e t e r m i n a l , i n c l u s t e r s o f one t o

f i v e , o r r a r e l y more ( F i g . 3 5 ) . !

N e c t a r .

A t n i g h t , l a r g e d r o p l e t s o f n e c t a r can be s e e n

s p a r k l i n g i n t h e f l o w e r s when a t o r c h i s s h i n e d i n t o them from

IiI

Fig. 35. Duabanga grandiflora.

5 a . A young t r e e , Ulu Gombak v a l l e y .

b . I n f l o r e s c e n c e . Note on the l o w e s t i n f l o r e s c e n c e ; -

1 . immature bud,2 . bud t h a t w i l l open a t dusk

( c a l y x s p l i t ) .3 . r em ains o f f lo w e rs t h a t have

opened p r e v i o u s l y from which a n t h e r s and p e t a l s have f a l l e n .

M

86

b e lo w . H o w ever , a t t e m p t s t o m e a s u re t h e vo lum e o f n e c t a r I n

t h e f l o w e r s f a i l e d b e c a u s e no f l o w e r s w ere c o l l e c t e d w i t h o u t

d i s l o d g i n g t h e n e c t a r .

P o l l e n .

The p o l l e n g r a i n s h av e b e e n d e s c r i b e d by

T h a n ik a i m o n i an d J a y a w e e r a (1 9 6 6 ) and b r i e f l y b y M u l l e r (1 9 6 9 )*

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

The mean p o l a r l e n g t h o f 20 g r a i n s was 3 2 . 6 p and t h e i r mean

e q u a t o r i a l d i a m e t e r p . The g r o s s t e x t u r e o f t h e p o l l e n ,

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

n o t a g g r e g a t e i n t o c lu m p s a s d o e s O ro x y lu m r

S c e n t .

I t was s i m i l a r t o t h a t 6 f S o n n e r a t i a b u t s l i g h t l y

s t r o n g e r .

■F l o w e r i n g b i o l o g y .

A n t h e s i s i s n o c t u r n a l an d l a s t s o n l y one n i g h t .

The f l o w e r s open b e tw e e n 1 7 .0 0 and 1 9 .0 0 h r s , i . e . e a r l i e r t h a n

S o n n e r a t i a s p p . a nd t h e s t a m e n s and a n t h e r s f a l l o f f a t dawn-.

V i s i t o r s b e s i d e s m a c r o g l o s s i n e b a t s .

None r e c o r d e d .

P h e n o l o g y .

L e a f g ro w th and f l o w e r i n g a r e s i m u l t a n e o u s and

c o n t i n o u s . No s e a s o n a l d i f f e r e n c e i n t h e r a t e o f e i t h e r was

o b s e r v e d b u t no p r e c i s e d a t a shov /ing t h e number o f f l o w e r s

p r o d u c e d p e r t r e e p e r u n i t t i m e w e re r e c o r d e d . As many a s 16

open f l o w e r s w ere c o u n t e d on one t r e e i n one n i g h t b u t t h e r e

w e r e - u s u a l l y t h r e e t o f o u r p e r t r e e on t r e e s xxp t o 15 m h i g h .

T h e re may h a v e b e e n more on l a r g e r t r e e s w h e re a c c u r a t e c o u n t s

w e re d i f f i c u l t t o o b t a i n .

8 . i i i . D uabanga m o l u c c a n a .

E c o l o g i c a l d i s t r i b u t i o n .

T h i s ' t r e e was s e e n o n l y i n S a b a h ( a l t h o u g h i t

i s a l s o common i n S a ra w a k and e l s e w h e r e ( P . S . A s h t o n , p e r s comm.)

w h e re i t w as f o u n d g ro v / ing on t h e b a n k s o f l a r g e r i v e r s i n

t h e Tawau an d Sem po rn a r e g i o n s .

M o r p h o lo g y .

T h i s t r e e w h ic h i s s i m i l a r t o D u ab ang a g r a n d l f l o r a

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

d e s c r i b e d b y B a c k e r an d van S t e e n i s ( 1 9 5 k ) •

P o l l e n .

I t i s s i m i l a r t o t h a t o f D. g r a n d i f l o r a b u t c an

b e d i s t i n g u i s h e d from t h a t s p e c i e s ( T h a n ik a im o n i an d J a y a w e e r a ,

1 9 6 6 ) . H ow ever , s i n c e t h e s e two s p e c i e s a r e a l l o p a t r i c and

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

g e n u s , i t p r e s e n t e d no p r o b l e m s o f i d e n t i f i c a t i o n .

F l o w e r i n g b i o l o g y .

No d a t a was o b t a i n e d b u t t h e p r e s e n c e o f n e a r l y

open b u d s an d no f u l l y open f l o w e r s , c o u p l e d w i t h t h e p r e s e n c e

o f f a l l e n a n t h e r s and p e t a l s b e lo w t h e t r e e s a t m id day s u g g e s t

t h a t a n t h e s i s i s n o c t u r n a l . One s a p l i n g , o n l y 3 m h i g h , h a d

many b u d s and f r u i t s on i t . D. g r a n d i f l o r a n e v e r f l o w e r e d

u n t i l i t was c o n s i d e r a b l y l a r g e r t h a n t h i s .

P h e n o lo g y .

No d a t a . T r e e s w ere f l o w e r i n g n e a r S e m p o rn a ,

S ab ah on 25 D ecem ber 1 9 7 2 .

9 . M o r a c e a e .

Ciig g e n u o , A i* tocsypus » contcXj-ns soiuG clixjrop"bG3ro*“

p h i l o u s s p e c i e s . The s p e c i e s j c o n c e r n e d w ere n o t i d e n t i f i e d .

C o rn e r (1 9 5 2 ) r e c o r d e d "abou l t 20 s p e c i e s i n M a la y a , i n lo w la n dI

f o r e s t o r c u l t i v a t e d ” » He d e s c r i b e s 14 s p e c i e s i n c l u d i n g a l l

t h e commoner o n es an d e m p h a s i s e s t h e a b u n d a n c e o f p o l l e n

r e l e a s e d by some o f them ( p . 650) s u g g e s t i n g t h a t t h e y m ig h t

b e w ind p o l l i n a t e d . A b u nd an t p o l l e n w o u ld a l s o b e e x p e c t e d o f

a c h i r o p t e r o p h i l o u s s p e c i e s , e s p e c i a l l y , w h e re a s i n A r t o c a r p u s ,

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

Fig. 3 6 . P o l l e n g r a i n s o f some c h i r o p t e r o p h i l o u s p l a n t s .

Photographs marked (g) are of grains in guano samples from Eonycteris srelaea; those marked (f) are from flowers.

Musa truncata(f). Note the substance on the outer wall of the exine. This is sticky

Arenga spp.(g). Note the swollen bases to the spines.

Cocos nucifera(f)

Mangifera spp.(g).Eugenia malaccensis(f)

Oroxylum indicum(f)

3 7 . P o l l e n g r a i n s o f some c h i r o p t e r o p h i l o u s p l a n t s .

P h o to g rap h s marked (g) a r e o f g r a i n s i n guano

samples from E o n y c te r i s s p e l a e a ; th o s e marked ( f )

a r e from f l o w e r s .

Ceiba p e n t a n d r a ( g ) Durio g r a v e o l e n s ( f )

Bombax v a l e t o n i i ( f ) Durio z i b e t h i n u s ( f )

P a r k i a s p e c i o s a ( f ) T3(g)

38 . P o l l e n g r a i n s o f some c h i r o p t e r o p h i l o u s p l a n t s and

a n t h e r s o f A r to c a rp n s s p p . P h o to g rap h s marked (g)

a r e o f i t e m s i n th e guano o f E o n y c te r i s s n e l a e a ;

t h o s e marked ( f ) a r e from f lo w e r s .

B a r r i n g t o n i a r acem o sa ( f ) S o n n e r a t i a a l b a ( g )

A r to c a rp u s s p p . ( g )

A nther o f A r to ca rp u s A nther o f A r to c a rn u s spp .h e t e r o p h y l l u s ( f ) ( g ) . Note th e p o l l e n g r a i n s . )

F i g . 3 9 . D iagram t o show t h e p o s i t i o n o f f l o w e r s on some

M a l a y s i a n c h i r o p t e r o p h i l o u s t r e e s .

I = Musa ( s e c t i o n Eumusa) s p p .

^ - Cocos n u c i f e r a

3 . = A re n g a s p p . c f . A. p i n n a t a

A = M a n g i f e r a s p p . c f M, i n d i e a

3 = P ro x y lu m in d ic u m

6 = P a . j a n e l i a l o n g i f o l i a

7 = Bombax v a l e t o n i i

® = C e ib a p e n t a n d r a

9 a - D u r io s p p . c f . D. z i b e t h i n u s

9b = " c f . D. g r a v e o l e n s

10 = B a r r i n g t o n i a s p p . c f . B . r a c e m o s a

I I = P a r k i a s p p .

1 2 = E u g e n ia m a l a c c e n s l s

1 3 = D uabanga g r a n d i f l o r a

1A = S o n n e r a t i a a l b a o r S . o v a t a

15 = S o n n e r a t i a c a s e o l a r i s

\

89

MACROGLOSSUS MINIMUS.

1• E c o l o g i c a l d i s t r i b u t i o n .

1 , i . P e n i n s u l a r M a l a y s i a .

T a b le 4 and F i g . 40 show a l l t h e l o c a l i t i e s i n

P e n i n s u l a r M a l a y s i a from which, s p e c im e n s o f M. niininius h a v e ■

b e e n e x a m in e d , i n c l u d i n g t h o s e c a u g h t d u r i n g t h e s t u d y and

t h o s e i n t h e f o l l o w i n g c o l l e c t i o n s ; B r i t i s h M useum (N at. H i s t . )

(BM(NH)), L ondon; I n s t i t u t e f o r M e d i c a l R e s e a r c h ( IM R ), K u a la

Lumpur; R o y a l O n t a r i o Museum (ROM), C an ada ; S i n g a p o r e N a t i o n a l

C o l l e c t i o n (S N C ); U n i v e r s i t y o f M alaya c o l l e c t i o n (UMC). A i l

t h e m a i n l a n d r e c o r d s a r e from c o a s t a l d i s t r i c t s i n o r n e a r to

m an g ro v es an d t h e r e s t a r e f rom o f f s h o r e i s l a n d s . (T he c o n c e n t r a t i o n

o f r e c o r d s from t h e S e l a n g o r c o a s t ( T a b l e 4 ) r e f l e c t s t h e l a r g e

num ber o f c o l l e c t i o n s t h a t h a v e b e e n made c l o s e t o t h e c a p i t a l ,

K u a la L u m p u r .) T h i s s p e c i e s i s p r o b a b l y p r e s e n t a l o n g t h e c o a s t

o f P e n i n s u l a r M a l a y s i a w h e r e v e r m a n g ro v e s g ro w .

Many M a c r o g lo s s u s s o b r l n u s s p e c i m e n s i n t h e IMR

c o l l e c t i o n , i n c l u d i n g some from 6000 f t ( 1 8 0 0 m) a . s . l . on

Gunong B r i n c h a n g , P a h a n g , h a v e b e e n i n c o r r e c t l y i d e n t i f i e d a s

nM. l a g o c h i l u s " (= M. m in im us h e r e ) and r e c o r d e d a s s u c h by

Lim (1 9 6 6 , 1 9 7 3 ) .

1 . i i . B o r n e o .

M. m in im u s was common i n t h e m a n g ro v e s a t Bako

N a t i o n a l P a r k , S a ra w a k and was c a u g h t i n a n a r r o w to n g u e o f

m an g ro v es f r i n g i n g a c r e e k a t Tawau, S a b a h , H o w ever , one was

c a u g h t on t h e ed g e o f l o w la n d f o r e s t a t S em engkok , n e a r K u c h in g ,

Table i+.List of localities in Peninsular Malaysia from which

specimens of Macroglossus minimus were examined.

Locality State Coordinates Source

Penang Hill Pulau Pinang 5°25'N 1 0 0 ° 1 5'E ROMKuala Gula Perak 4°J+5'N 100°28'E UMC,

IMR,ROM,EM(NH)

Kuala Kulat Perak ? IMRRantau Panjang Selangor 3°02‘N 101°25'E This

UMC,study, ROM, IMR

P. Pintu Gedong Selangor 2°55'N 101°15’E SNCP. Tengah,Klang Selangor 3°00'N 101°16'E UMCP. Ketam, Klang Selangor 3°03'N 101 °1 6'E UMCCarey Island Selangor 2°/f8'N 1 0 1 °21'E This study, IMRKlang Selangor 3°02'N 101 °27'E BM(NH), IMR,

This studyEukit Laut Selangor 2°2|2'N 101 °31 'E IMRKuala Selangor Selangor 3°26'N 101 °1 V E This studyP. Tulai Pahang 2°5VN 10/f°07»E UMCSri Buat Island Pahang 2°Z+2'N 103°55'E SNCSelensing Pahang ? SNCSingapore Repb. of

Singapore1°22«N 103°48»E SNC

Note P = Pulau = Island.IMR = Institute of Medical Research, Kuala Lumpur.ROM = Royal Ontario Museum, Canada.SNC = Singapore National Collection.UMC = University of Malaya Collection.EM(NH)= Eritish Museum (Natural History).

F i g . 40. L o c a l i t y r e c o r d s of M a c r o g l o s s u s s p p . i n P e n i n s u l a r M a l a y s i a .

k m

© M . m i n i m u s

❖ M . s o b r i n u s

90

S a ra w a k an d s p e c i m e n s from many i n l a n d l o c a l i t i e s h a v e b e e n

ex am in ed ( F i g . i+1 a nd T a b le 5) •

2 . R o o s t i n g .

• No M. m in im u s w e re fo u n d r o o s t i n g d u r i n g t h e d ay

ci. 2 cums f suvt jLs.X e v i d e n c e sud. iDsiiSLVionx’1 c f csLjptxvc ^ x v s

some i n f o r m a t i o n on r o o s t i n g h a b i t s . A t R a n ta u P a n j a n g , some

b a t s w ere c a p t u r e d j u s t a f t e r d a r k a t t h e same n e t s i t e on

s e v e r a l n i g h t s , e . g .

Ad. S No. 20/1+9012.

N e t s i t e D a te Time

11+ : 5 J u l y 1972 2 0 . 1 0 h r s

• 1‘1+ 30 A u g u s t 1972 1 9 . 1 0 h r s

11+ 29 S e p te m b e r 1972 1 9 .1 0 h r s

11+ 10 November 1972 1 9 .3 5 h r s

, 1 1 + 28 J a n u a r y 1973 1 9 . 3 5 h r s

11+ 2 5 J u l y 1973 2 0 . 3 0 h r s

(D usk was b e tw e e n 1 9 .0 0 and 1 9 .1 0 h r s . )

T h i s show s t h a t some o f them r o o s t l o c a l l y and

i n t h e same a r e a f o r some t i m e .

M ost o f t h e v i l l a g e r s a t R a n ta u P a n j a n g knew

t h e w h e r e a b o u t s o f r o o s t i n g c o l o n i e s o f C y n o p te r u s b r a c h y o t i s

( P t e r o p o d i d a e ) and S c o t o p h i l u s t e m m i n c k i i L each ( V e s p e r t i l i o n i d a e )

i n c o c o n u t p a lm s , f r u i t t r e e s o r t h e i r h o u s e s . They w e re a l s o

aw are o f M y o t i s m y s t a c i n u s (K u h l) ( V e s p e r t i l i o n i d a e ) r o o s t i n g

>z

F I G . 4 1 . MAP OF B O R N E O S H O W I N G LOCALITY R ECORDS FORM A C R O G L O S S U S MINI MUS

110 E 115 E

300j 1--------1s c a l e km

110*E___

115°E

Table 5 .

Table showing localities in Eorneo from which specimens of Macroglossus minimus have been examined.

Locality State Coordinates Source

Niah Sarawak 3°48'N 113°47'E IMR, EM(NH)Long Dati, nr Long Lelang

Sarawak 3°27'N 115°10'E EM(NH)

Pa Mada Sarawak 3°35’N 115°331E UMCSemengkok F.R. Sarawak 1°26'N 1 10°20»E This studyBako N.P. Sarawak 1°/f9'N 110°27'E ti

Kg. Pang. Kuap Sarawak 7 IMRKiau Sabah 6°06'N 1 1 6°31’E SNCPoring, nr. Ranau

Sabah 6°06'N 116°32»E IMR

Tenom Sabah 5°13’N 115°57’E BM(NH)Tawau Sabah 4O20'N 118°0 1•E This studySebulu Kalimantan 0°1 6'S 117°00'E NAMRU-2Teleng Kalimantan 2°30'S 115°2VE NAMRU-2

Note Kg. Pang. = Kampong Pangkalan (village landing place). N.P. = National Park.F.R. = Forest Reserve.IMR = Institute of Medical Research, Kuala Lumpur. SNC = Singapore National Collection.UMC = University of Malaya Collection.

EM(NH) = British Museum (Natural History).NAMRU-2 = United States Naval Medical Research Unit,

No. 2.

9T

i n t h e u n f u r l e d new l e a v e s o f b a n a n a p l a n t s . H o w ev er , t h e y had

no k n o w le d g e o f t h e r o o s t i n g h a b i t s o f M a c r o g lo s s u s m in im u s .

T h i s s u g g e s t t h a t M. min im us r o O s t s o l i t a r i l y o r i n s m a l l g r o u p s ,

p o s s i b l y i n c o c o n u t p a lm s o r t h e c row ns o f f r u i t t r e e s w h e re

t h e y w ou ld n o t b e n o t i c e d .

Oil S6'v6jf8.x o c c a s i o n s , b a ts 'LllOUgut to M* minimus

w e re o b s e r v e d h a n g i n g from t h e f r o n d s o f c o c o n u t p a lm s f o r up

t o one h o u r d u r i n g t h e n i g h t . None w ere s e e n b e f o r e 2 2 , 0 0 h r s .

A l l w e re h a n g i n g a l o n e .

I n c a p t i v i t y , M. m in im us r o o s t e d f rom t h e f r o n d s

o f a s m a l l p a lm t r e e p l a c e d i n t h e c a g e o r from t h e r o o f o f t h e

c a g e . F i g . 42 show s t h e c h a r a c t e r i s t i c way t h e y p l a c e d one f o o t

e a c h s i d e o f t h e s t r u c t u r e t h e y hu ng f ro m . A f t e r 24 h o u r s i n

c a p t i v i t y , m o s t b a t s h ad s e l e c t e d a p r e f e r r e d r o o s t s i t e w h ich

t h e y u s e d t h e r e a f t e r . They r o o s t e d o u t o f r e a c h o f one a n o t h e r

b u t no a g g r e s s i o n was n o t e d b e tw e e n th em . D u r in g t h e d a y , t h e

w in g s e n v e lo p e d t h e h e a d an d abdomen o f t h e b a t s , i f d i s t u r b e d ,

t h e b a t s w i th d r e w t h e i r h e a d s from u n d e r t h e i r w in g s t o O b se rv e

t h e i r s u r r o u n d i n g s . I f a l a r m e d , th e y f o l d e d t h e i r w in g s by t h e i r

s i d e s i n r e a d i n e s s f o r f l i g h t .

A t d u s k , t h e y becam e a c t i v e , f l y i n g a b o u t t h e i r

c a g e . H ow ever , a s s o o n a s t h e y ha d f e d ( e a c h h w o u ld d r i n k a b o u t

2 ml o f t h e fo o d s o l u t i o n ) , t h e y a g a i n s l e p t f o r s e v e r a l h o u r s .

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

t h e i r thum bs ' ( F ig , 42 ) . )t

An a s t o n i s h i n g i n c i d e n t e m p h a s i s e d t h e i r p r o p e n s i t y

f o r r e t u r n i n g t o t h e i r a c c u s to m e d r o o s t s i t e ( a n d ’’fo o d s o u r c e " ) .

F ig . 4 2 . M acroglossus minimus.

a. Posture adopted for b. Typical roosting posture,urination and defaecation.

Fig. ^3 . Longitudinal sections of the tongue of M. minimus.

a. Filiform papillae near tip of tongue.I-------------------- I

0-5 mm

b. Conical papillae in mid-region of tongue.1_______________________________________ . 1

0*5 mm

92

F i v e a d u l t b a t s w e re c a u g h t a t R a n ta u P a n j a n g an d c a g e d i n

t h e f o y e r o f t h e G r e a t H a l l , U n i v e r s i t y o f M a la y a , f o r f i v e

d a y s d u r i n g an e x h i b i t i o n . They v/ere g i v e n a p a lm t r e e i n t h e i r

c a g e from w h ic h t o r o o s t . E ach b a t s e l e c t e d a r e g u l a r r o o s t

s i t e . On t h e l a s t e v e n i n g , t h e b a t s w ere r e l e a s e d , t h e c a g e

d i s m a n t l e d an d t h e p a lm t r e e rem ov ed 200 m t o t h e c a r p a r k o f

t h e S c h o o l o f B i o l o g i c a l S c i & n c e s . The f o l l o w i n g m o r n in g , one

o f t h e ' b a t s (w h ic h h a d b e e n r i n g e d p r i o r t o r e l e a s e ) was

r o o s t i n g from t h e same p l a c e on t h e same f r o n d o f t h e pa lm t r e e

t h a t i t h a d o c c u p i e d i n t h e c a g e . T h is ! b a t h a d become v e r y tam e

an d r e t u r n e d on f o u r o f t h e f o l l o w i n g t e n n i g h t s , e n t e r i n g my

room b y t h e w indow . I t s e t t l e d on my o u t s t r e t c h e d h an d a n d ,

a f t e r b e i n g f e d , s l e p t u n d e r my d e s k f o r h a l f t o one h o u r b e f o r e

f l y i n g aw ay .

3 . Food s o u r c e s o f M a c r o g lo s s u s m in im u s .

T a b le 6 l i s t s t h e p l a n t s from w h ic h M. m in im us

w e re shown t o f e e d . They w e re d e t e r m i n e d by o b s e r v i n g w h ic h

f l o w e r s t h e b a t s v i s i t e d an d b y i d e n t i f y i n g t h e p o l l e n fo u n d

on t h e i r b o d i e s a n d / o r i n t h e i r f a e c e s . O n ly c u l t i v a t e d b a n a n a s ,

Musa s p p . , w e re n o t d e t e c t a b l e by p o l l e n a n a l y s i s b e c a u s e t h e y

r e l e a s e no p o l l e n . The m o s t i m p o r t a n t fo o d s o u r c e s a t R a n ta u

P a n j a n g w e re S o n n e r a t i a a l b a , S . c a s e o l a r i s , Musa s p p , ( c u l t i v a t e d )

an d Cocos n u c i f e r a .

3 » i« S o n n e r a t i a ' a l b a a nd S . c a s e o l a r i s .

B o th t h e s e m angrove s p e c i e s a r e f a v o u r i t e fo o d

s o u r c e s o f ,M. m in im us a t R a n ta u P a n ja n g w h e re t h e f l o w e r s n e v e r

Table 6.The table records the plant species from which Macroglossus minimus fed at Rantau Panjang, Selangor.

Species Importance Abundance Flowering Origin Foodprovided

Bats seen feeding

Sonneratia alba +++ +++ Flushes wild p+n yesS. caseolaris +++ +++ continuous wild p+n yesS. ovata + + flushes wild p+n yesCocos nucifera +++ +++ continuous cult. p+?n noMusa spp. cult. +++ +++ continuous cult. n yesEugenia malaccensis ++ ++ seasonal cult. p+n yesCeiba pentandra + + seasonal cult. p+n * yesDurio zibethinus + + seasonal cult. p+n yesMangifera spp. ++ ++ seasonal cult. p=?n noRhizophora spp. + +++ wild p+?n no

Note. +++ = very important/abundant ++ = less important/abundant + = unimportant or rare

cult. = cultivated p = pollen n = nectar

9.3

fail to attract bats. Only S. alba grows at Telok Asam, Bako

National Park, where it was the most important food source for

Minimus itt January and Kay 1973 but, as there were fewer

bats there, not all flowers were visited (see p. 109).

Me mitt im us a p p r o a c h S o n n e r a t i a f l o w e r s from i n

4> n r : v t A - W U ^ - t o 1 *1 n r r7A W + r , l V n f m r \ v» rvJ . X W X U C X U V W J_ W ^ / A w a v / u U.<-<-A~ - t . J u l / X X U ^ U k> W V, W V i U O -

t i m e s t h e y f l y up t o a f l o w e r from b e lo w ( p a r t i c u l a r l y , when

a p p ro a c h in g . S . c a s e o l a r i s f l o w e r s w hich t e n d t o be s e m i - p e n d u l o u s

( F i g , 3 2 ) « They w ere f r e q u e n t l y s e e n t o h o v e r m o m e n t a r i l y b e f o r e

l a n d i n g on a f l o w e r b u t t h e y w ere n e v e r s e e n t o f e e d from a

f l o w e r b e f o r e l a n d i n g . When t h e y l a n d t h e y g r a s p t h e s e p a l s

o r t h e b a s e o f t h e c a l y x w i t h . t h e i r f e e t and a n c h o r t h e thumb

c law o n t o - t h e c a l y x . I n t h i s p o s i t i o n th e s t i g m a i s b r o u g h t

i n t o c o n t a c t w i t h t h e b a t s ' h e a d s o r abdomens and many o f th e

a n t h e r s a r e p r e s s e d a g a i n s t t h e b a t s ' abdomens o r t h e v e n t r a l

s u r f a c e s o f t h e i r w in g s ( F i g , V i) „• I n o r d e r t o o b t a i n n e c t a r ,

t h e y h av e t o b u r y t h e i r h e a d s d e e p among th e s t a m e n s so- t h a t

t h e h e a d s , t h r o a t s , abdomens a n a t h e v e n t r a l s u r f a c e s o f t h e i r

w in g s , e s p e c i a l l y t h e p l a g i o p a t a g i u m , and t h e w in g t i p s o f

f e e d i n g b a t s become l i b e r a l l y d u s t e d w i t h p o l l e n . Many o f t h e

ii* Minimus t h a t w ere c a u g h t a t R a n ta u P a n j a n g , Bako N a t i o n a l

P a r k and K u a la S e l a n g o r h a d S o n n e r a t i a p o l l e n on t h e s e p a r t s

o f t h e i r b o d i e s .

The b a t s u s u a l l y r e m a in on t h e f l o w e r f o r a v e r y

s h o r t t ime,; s e ld o m more th a n t h r e e s e c o n d s , f r e q u e n t l y l e s s

th a n tw o . When d e p a r t i n g , t h e y k e e p h o l d o f t h e c a l y x w i t h

t h e i r f e e t u n t i l t h e y h a v e t u r n e d a ro u n d and o p ened t h e i r w in g s .

F ig . 4 4 . M acroglossus minimus v i s i t i n g Sonneratia f lov /ers .

a. Approaching a Sonneratia ovata flower. Note the horizontal flight path.

b. Landed on a Sonneratia alba flower.

9k

They c a n r e l e a s e t h e h o l d o f t h e i r f e e t on t h e f i r s t downward

s t r o k e o f t h e w i n g s . I n t h i s w ay , t h e y can l e a v e t h e f l o w e r

w i t h o u t l o s i n g a ny h e i g h t .

Two m e th o d s o f i n g e s t i n g S o n n e r a t i a s p p . p o l l e n

w e re o b s e r v e d i n c a p t i v e M. m i n im u s .

1 • Aittix* f r O m u 1 xOWu'i , uliO u8.ts Xick.Ou

o f f a l l t h e p o l l e n w h ic h h ad b e e n d e p o s i t e d on th e m . T h e i r

l o n g to n g u e s w e re a b l e t o r e a c h a l m o s t a l l p a r t s o f t h e i r .

b o d i e s , i n c l u d i n g t h e i r h e a d s a s f a r b a c k a s t h e a r e a b e tw e e n

th e e a r s . W ith one o r two l i c k s t h e y w ere a b l e t o c l e a n a l m o s t

a l l t h e p o l l e n from t h e a r e a c o v e r e d b y t h e sw eep o f t h e t o n g u e .

Even w h i l e l y i n g e n t a n g l e d i n a m i s t n e t , t h e y w e re fo u n d

l i c k i n g p o l l e n o f f t h e i r b o d i e s . B a t s t h a t h ad b e e n i n t h e n e t s

f o r a s much a s f i v e m i n u t e s a l m o s t n e v e r h ad much p o l l e n on

th e m .

The to n g u e s o f M a c r o g lo s s u s s p p . , l i k e t h o s e o f

a l l p t e r o p i d s , a r e c o v e r e d i n p a p i l l a e . T h ese h a v e b e e n d e s c r i b e d

an d f i g u r e d i n A n d e r s e n ( 1 9 1 2 ) . W h ile t h e lo n g f i l i f o r m p a p i l l a e

fo u n d on t h e t i p s and m a r g in s o f t h e a n t e r i o r p a r t o f t h e to n g u e

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

o f n e c t a r ( s e e b e l o w ) , t h e r e m a i n i n g p a r t p r o d u c e s a s e r r a t e d

s u r f a c e w h ich a c t s a s a f i n e l y t o o t h e d comb. When t h e b a t s l i c k

p o l l e n from t h e i r b o d i e s , t h e to n g u e i s s l i g h t l y d e c u r v e d ,

c a u s i n g t h e p a p i l l a e t o s t a n d away f rom i t s s u r f a c e , t h u s i n c r e a s i n g

t h e i r e f f e c t i v e n e s s a s a comb ( F i g . kJ>£),

2 . O c c a s i o n a l l y , h a n g in g from t h e b a s e o f t h e c a l y x

b y i t s h i n d f e e t , a b a t g a t h e r e d t h e s t a m e n s b e tw e e n i t s f o r e a r m s

95

and l i c k e d t h e s u r f a c e s o f t h e c o m p a c te d a n t h e r s . No h a t s w ere

s e e n t o ea t . an a n t h e r n o r damage a f l o w e r i n a n y w ay .

B o th m e th o d s o f i n g e s t i o n w o u ld h a v e b e e n f e a s i b l e

i n t h e w i l d .

I n g e s t i o n o f n e c t a r .

N e c t a r i s h e l d i n t h e c u p - s h a p e d c a l y x o f S o n n e r a t i a

s p p . f l o w e r s w h ic h a r e t e r m i n a l on f l e x i b l e t w i g s t h a t b e n d

u n d e r t h e w e i g h t o f a v i s i t i n g b a t t h u s t i l t i n g t h e f l o w e r down­

w a rd s an d c a u s i n g t h e n e c t a r t o r u n f o r w a r d s . N e c t a r i s p r e v e n t e d

f rom e s c a p i n g by. t h e c l o s e l y p a c k e d f i l a m e n t s o f t h e a n t h e r s

w h ich r e t a i n i t b y c a p i l l a r i t y b u t i t can e a s i l y r u n down t h e

m o i s t t o n g u e o f t h e b a t .

I n c a p t i v i t y , b a t s f e d on a l i q u i d d i e t a lw a y s

b e n t t h e t i p o f t h e t o n g u e t h r o u g h 9 0 ° when i n s e r t i n g i t i n t o

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

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

t h e fo o d s o l u t i o n when t h e t o n g u e was w i th d r a w n . T h i s a c t i o n

i s o f o b v i o u s a d v a n t a g e w h e re a b a t m ig h t h a v e t o o b t a i n n e c t a r

f rom a f l o w e r w i t h a d eep c o r o l l a when c o n v e n t i o n a l l a p p i n g

w ou ld b e d i f f i c u l t i n t h e c o n f i n e d s p a c e .

3.ii. Musa s p p .

C u l t i v a t e d b a n a n a s w ere grown i n a l m o s t a l l o f

t h e c u l t i v a t e d a r e a a t R a n ta u P a n ja n g and w e re h e a v i l y u t i l i s e d

M* M inim us w h ich d r i n k t h e n e c t a r . E ach o f f o u r t e e n

i n f l o r e s c e n c e s t h a t w ere w a tc h e d c o n t i n o u s l y from 1 9 . 3 0 t o

2 0 , 3 0 h r s on v a r i o u s d a t e s i n 1 9 7 2 was v i s i t e d a t l e a s t t h r e e

96

t i m e s a nd one w as v i s i t e d n i n e t i m e s i n t h e two h o u r s o f

o b s e r v a t i o n s (mean 2 . 8 + 1 . 6 v i s i t s p e r i n f l o r e s c e n c e p e r h o u r )

K F ig , ^ 6 ) .

B e tw een 19*30 and 2 0 . 3 0 h r s a s many a s 22

M. m in im us w e re c a u g h t i n f o u r m i s t n e t s s e t c l o s e t o a g ro v e

o f f l o w e r i n g b a n a n a s . The u s u a l c a p t u r e r a t e f l u c t u a t e d b e tw e e n

t h a t e x t r e m e a n d z e r o , v e r y few b e i n g c a u g h t when t h e r e was a

b r i g h t moon a s t h e y w e re a b l e t o s e e t h e n e t s . On m o o n le s s

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

B an an a f l o w e r s c a n p r o d u c e a s much a s 2 . if ml o f

n e c t a r p e r i n f l o r e s c e n c e p e r n i g h t b u t t h e y r e l e a s e no p o l l e n .

N e c t a r i s s e c r e t e d b y n e c t a r i e s i n t h e r e c e p t a c l e o f t h e b a n a n a

f l o w e r an d h e l d i n t h e b a s e o f t h e f l o w e r (w h ic h f a c e s downwards)

b y s u r f a c e t e n s i o n ( F i g . 1 if ) . When t h e compound t e p a l and t h e

f r e e t e p a l a r e f o r c e d a p a r t , e . g . b y a b a t i n s e r t i n g i t s m u z z le

i n t o a f l o w e r - t h e ■ n e c t a r r u n s down t o form, a s m a l l p o o l a t t h e

a p e x o f t h e f r e e t e p a l ( c o m p a re F i g . 1 if an d if5«0 • The b a t t h e r e ­

f o r e , o n ly h a s t o t o u c h t h i s d ro p w i t h i t s m o i s t t o n g u e f o r t h e

n e c t a r t o r u n down i n t o i t s m o u th .

When f e e d i n g i n b a n a n a g r o v e s , M. m in im us f l e w

j u s t a b o v e t h e g r a s s an d f e r n s , w h ich fo rm ed a g ro u n d c o v e r

a b o u t 0 . 5 m h i g h , s e a r c h i n g f o r b a n a n a i n f l o r e s c e n c e s from b e lo w .

They w e re u s u a l l y c a u g h t i n t h e s e c o n d o r t h i r d s h e l v e s o f if

s h e l f m i s t n e t s s e t w i t h t h e l o w e s t s h e l f j u s t a b o v e g ro u n d

l e v e l , i . e . b e tw e e n 1 m an d 2 m a b o v e g ro u n d l e v e l . (M ost b a n a n a

i n f l o r e s c e n c e s w h ich h a n g b e lo w t h e l e a v e s , w e re b e tw e e n 3 m

an d 5 m a b o v e t h e g r o u n d .

I

I/

F ig . L Macroglossus minimus v i s i t i n g c u l t i v a t e d banana

b. Scratch marks in the"bloom" on the bract made by the bats. (Photographed the morning after anthesis when the flowers had fallen.

c. Enlargement of a bat feeding. Note: l. The anther (A) rubbing

over the bat's face.2. The use of the thumb

claw for anchorage.3. The bat has been ringed.

inflorescences,

a. Settled on inflorescence and about to feed.

N°.

of

infl

ore

scen

ces

F ig . if6 .

Histogram showing the number of visits to each of cultivated banana inflorescences made by Macroglossus minimus between 19.30 and 21.30 hrs on various dates in 1972. Data obtained at Rantau Panjang9 Selangor.

5-r

4 -

3 -

2 -

1 -

N ° . o f v i s i t s b y M . m i n i m u s

97

I n o r d e r t o f e e d ' f ro m t h e b a n a n a f l o w e r s t h e b a t s

m u s t swoop up t o t h e i n f l o r e s c e n c e and l a n d on t h e a b a x i a l

s u r f a c e o f t h e b r a c t t h a t l i e s b e lo w t h e e x p o s e d h an d o f f l o w e r s

( F i g . 4 5 a ) . The waxy bloom on t h e b r a c t g i v e s i t a " n o n - s l i p "

s u r f a c e t h a t h e l p s t h e b a t s t o o b t a i n p u r c h a s e b u t t h e y a r e

a l s o s u p p o r t e d b y t h e i r thumb , c la w s w h ich t h e y a n c h o r i n t o a

f l o w e r o r a b r a c t C F ig . 45 ) • From t h i s p o s i t i o n , t h e y can

i n s e r t t h e i r m u z z l e s i n t o t h e i n d i v i d u a l f l o w e r s on t h e i n f l o r e ­

s c e n c e i n s e a r c h o f n e c t a r . E ach v i s i t t o an i n f l o r e s c e n c e l a s t s

one t o f i v e s e c o n d s .

A l l b a n a n a i n f l o r e s c e n c e s t h a t w ere ex am in ed a f t e r

2 2 . 0 0 h r s a t R a n ta u P a n ja n g h a d c h a r a c t e r i s i t i c m a rk s w h ere t h e

w r i s t s an d f e e t o f v i s i t i n g b a t s h a d ru b b e d away t h e waxy b loom

t h a t c o v e r s t h e a b a x i a l s u r f a c e o f t h e b r a c t t h e y l a n d on ( F i g . 4 5 )

( e x c e p t i n f l o r e s c e n c e s t h a t w e re i n a c c e s s i b l e t o t h e b a t s e i t h e r

b e c a u s e t h e y w e re o v e rg ro w n b y c r e e p e r s o r b e c a u s e m i s t n e t s

ha d b e e n s e t a d j a c e n t t o t h e m ) . (E . s p e l a e a a l s o v i s i t e d t h e

f l o w e r s o f c u l t i v a t e d ,b an a n as b u t a t R a n ta u P a n j a n g i n much

lo w e r nu m b e rs t h a n M. m in im u s . ) I t was n o t p o s s i b l e t o i d e n t i f y

w h ic h s p e c i e s o f b a t h a d v i s i t e d a b a n a n a i n f l o r e s c e n c e from

t h e m ark s t h a t i t l e f t . The s i t u a t i o n was f u r t h e r c o m p l i c a t e d

when s e v e r a l b a t s h a d v i s i t e d t h e i n f l o r e s c e n c e . Some o f t h e

m ark s on t h e b r a c t s may h a v e b e e n made by E . s p e l a e a .b u t m o s t

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

M. m in im us a s w e l l . C o n t r a r y t o t h e f i n d i n g s o f F a e g r i an d v an

d e r P i j l ( 1 9 6 6 ) , I was u n a b l e t o c o u n t t h e num ber o f b a t s t h a t

h a d v i s i t e d a b a n a n a i n f l o r e s c e n c e from t h e m a rk s t h e y l e f t on

t h e f l o w e r s o r b r a c t s .

98

Many o f t h e b a t s t h a t w ere c a u g h t h a d w h i t e

p a t c h e s o f t h e waxy b loom on t h e i r w r i s t s an d f e e t sh o w in g

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

number o f b a t s w i t h s u c h w h i t e p a t c h e s c o u ld n o t b e made b e c a u s e

i t was so e a s i l y r u b b e d o f f when t h e y s t r u g g l e d i n t h e m i s t n e t s

O P C j l O h i O c a g S *

3 . i i i . Co c o s n u c i f e r a ( c o c o n u t s )

E v id e n c e t h a t M. m in im u s f e d from c o c o n u t f l o w e r s

came from a n a l y s i s o f p o l l e n i n t h e i r f a e c e s . The p o l l e n was

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

so no o b s e r v a t i o n s w e re made oh b a t s v i s i t i n g t h e f l o w e r s .

3.iv„ Other souces of pollen and nectar included Celba pentandra,

Durio zj.bethinus, Eugenia malaccensis, Manglf era spp„, Rhizophora

spp. and Sonneratta ovata..

Ceiba pentandra and Durio zibethinus.

T h e re w ere s e v e n C, p e n t a n d r a and one I), z i b e t h i n u s

t r e e s o l d enough t o f l o w e r i n t h e s t u d y a r e a a t R a n ta u P a n j a n g .

The f l o w e r i n g s e a s o n s o f b o t h w e re s h o r t and n e i t h e r s p e c i e s

w ere i m p o r t a n t t o M. m in im us t h e r e a l t h o u g h t h e b a t s f e d from

them when t h e y w e re a v a i l a b l e .

Eugenia-malaccensis.

When f e e d i n g from E, mal a c c e n s i s , t h e b a t s f l e w

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

b e f o r e l a n d i n g on t h e f l o w e r s ( s e e p . 7 2 and Fig* 39 ) • o r

sw o o p in g up t o them from b e lo w . S o m et im es a s many a s f i v e o r

99

s i x b a t s w e re f e e d i n g s i m u l t a n e o u s l y i n s i d e t h e c a n o p y o f one

t r e e . H ow ev er , t h i s s p e c i e s had v e r y s h o r t f l o w e r i n g s e a s o n s

( s e e p . 7 3 ) - a n d ■t h e r e w e re o n ly e l e v e n t r e e s r e c o r d e d i n t h e

s t u d y a r e a . T h u s , a l t h o u g h i t was h e a v i l y u s e d when a v a i l a b l e ,

i t s low an d i n t e r m i t t e n t a v a i l a b i l i t y l i m i t e d i t s im p o r t a n c e

a s a fo o d s o u r c e f o r M. m i n im u s .

M a n g i f e r a s p p .

E v id e n c e f o r M. m in im u s f e e d i n g f rom M a n g i f e r a

s p p . comes from t h e p r e s e n c e o f p o l l e n i n t h e f a e c e s o f some

b a t s . No b a t s w e re s e e n f e e d i n g on t h e f l o w e r s b u t no f l o w e r i n g

t r e e s w ere k e p t u n d e r w a tc h s i n c e t h i s p o l l e n s p e c i e s w as n o t

i d e n t i f i e d u n t i l a f t e r t h e c o m p l e t i o n o f f i e l d s t u d i e s .

R h iz o p h o r a s p p .

A l th o u g h two R h iz o p h o r a s p p . g rew v e r y commonly

i n t h e m a n g ro v e s a t R a n ta u P a n j a n g , M. m in im us w ere n e v e r

o b s e r v e d v i s i t i n g them an d t h e o n ly e v i d e n c e t h a t t h e y d i d so

comes from t h e p r e s e n c e o f p o l l e n i n t h e f a e c a l p e l l e t s

S o n n e r a t i a o v a t a .

A l l t h e r e m a r k s r e l a t i n g t o M. m in im us v i s i t i n g

S . a l b a a p p l y e q u a l l y w e l l t o S . o v a t a w h ic h a l s o f l o w e r e d i n

f l u s h e s . I t v/as a r a r e t r e e i n t h e s t u d y a r e a a t R a n ta u P a n j a n g ;

o n l y f i v e w e re fo u n d on t h e n o r t h b a n k and e i g h t on t h e s o u t h

bank, o f t h e r i v e r . I t s s c a r c i t y l i m i t e d i t s i m p o r t a n c e t o

M. m in im us a s a fo o d s o u r c e .

100

4 . A n a l y s i s o f p o l l e n c o l l e c t e d from M. m i n im u s ♦

4 . i . Faecal samples from Rantau Panjang.

T a b le 7 , p a r t c , r e c o r d s t h e n u m e r i c a l p r o p o r t i o n

o f e a c h d i f f e r e n t p o l l e n t y p e i d e n t i f i e d i n 1 6 2 f a e c a l s a m p le s

from M. m in im us c a u g h t i n t h i s s t u d y a r e a . The p o l l e n o f t h e

m angrove t r e e s , S o n n e r a t i a a l b a ( i n c l u d i n g S . o v a t a ) an d S .

c a s e o l a r i s , w e re t h e m o s t n um e ro u s t y p e s , b e i n g p r e s e n t i n

1 3 0 ( 8 0 , 2 % ), an d t h e o n l y p o l l e n s p e c i e s i n 8 0 ( 4 9 . 4 %) o f t h e

s a m p l e s .

M ost S. a l b a t r e e s f l o w e r e d t o g d t h e b i n p r o l o n g e d

f l u s h e s w h i l e S . c a s e o l a r i s f l o w e r e d c o n t i n o u s l y ( p . 8 l ) .

T a b le 7 , p a r t s a and b show s t h e p r o p o r t i o n s o f e a c h p o l l e n

t y p e d u r i n g ( a ) and b e tw e e n (b ) S , a l b a f l o w e r i n g f l u s h e s . The *

f r e q u e n c y w i t h w h ic h S . a l b a p o l l e n O c c u r r e d i n t h e s a m p le s

f e l l f rom 6 9 »2 % d u r i n g f l u s h e s t o 17,2% b e tw e e n f l u s h e s ( t a b l e

7 , a an d b , l a s t c o lu m n ) . B e tw een f l u s h e s t h e r e was a l s o an

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

s a m p le s i n w h ic h i t was p r e s e n t ( T a b l e 8 ) .

T h e re was no c o r r e s p o n d i n g d i f f e r e n c e i n t h e

f r e q u e n c y w i t h v /h ich S . c a s e o l a r i s p o l l e n o c c u r r e d i n t h e

s a m p le s b e tw e e n p e r i o d s o f f l o w e r i n g and n o n - f l o w e r i n g o f S . a l b a

( T a b le 7, a an d b , l a s t co lum n) a l t h o u g h t h e p r o p o r t i o n o f

5 , c a s e o l a r i s p o l l e n i n t h o s e s a m p le s i n w h ic h i t o c c u r r e d v/as

*F o o t n o t e The f r e q u e n c y v / i th w h ich a g i v e n p o l l e n s p e c i e s

o c c u r r e d i n n s a m p le s i s d e f i n e d h e r e a s t h e p e r c e n t a g e o f n

s a m p le s i n v /h ich t h a t p o l l e n s p e c i e s v/as p r e s e n t .

T a b le 7*

The t a b l e r e c o r d s t h e a b u n d a n c e o f e a c h p o l l e n ty p e

fo u n d i n f a e c a l s a m p le s c o l l e c t e d from 162 M a c r o g lo s s u s

m in im us a t R a n ta u P a n j a n g , S e l a n g o r .

a = s a m p le s c o l l e c t e d d u r i n g S . a l b a f l o w e r i n g f l u s h e s (n = 10if)

b = '< ■» b e tw e e n » » » (n- = 5 8 )

c = t o t a l ( i . e . a + b c om bin ed ) (n = 1 6 2 )

The f i g u r e s show t h e num ber o f s a m p le s c o n t a i n i n g 100%, 75 to

99%j 5 0 t o 7^%> 2 5 t o k9% o r l e s s t h a n 2 5 % ( b u t a t l e a s t one

p o l l e n g r a i n ) r e s p e c t i v e l y o f e a c h p o l l e n s p e c i e s . The f i g u r e s

i n b r a c k e t s a r e p e r c e n t a g e s o f t h e t o t a l num ber o f s a m p l e s .

N o te S . a l b a = S o n n e r a t l a a l b a ( m a n g r o v e s ) ,

S . c a s . = S o n n e r a t i a c a s e o l a r i s (m a n g ro v e s )

Cocos = Cocos n u c i f e r a ( c o c o n u t )

C e ib a = C e ib a p e n t a n d r a (k a p o k )

E u g . = ■*E u g e n ia m a l a c c e n s i s (M alay a p p l e )

Mang. = M a n g i f e r a s p p . (mango)

D u r io = D u r io z i b e t h i n u s ( d u r i a n )

R h i z . = R h iz o p h o r a s p p . (m an g ro v e )

«T h ese two p o l l e n t y p e s w e re i d e n t i f i e d t o g e n u s l e v e l o n ly #

B a t s p r o b a b l y f e e d on o t h e r s p e c i e s i n e a c h g e n u s b u t t h e s e

s p e c i e s w e re t h e o n l y o n es a t R a n ta u P a n j a n g w h ic h b a t s w e re

r e c o r d e d v i s i t i n g .

Species 100% • 75-99% 50-74% 25-49% < 2 5 % total

S.alba a 5'+(51 .9) 8 ( 7 .7 ) 7 ( 6 .7 ) 7( 6 .7 ) 1 1 ( 1 0 . 6 ) 8 7 (8 3 .7 )+ b 2 6 (4 4 .8 ) 7 (1 2 .1 ) 5 ( 8 . 6 ) 0 5 ( 8 . 6 ) 4 3 (7 4 .1 )

S .cas. c 8 0 (4 9 .4 ) 1 5( 9 .3 ) 1 2 (7 .4 ) 7( 4 .3 ) 1 6 ( 9 .9 ) 1 3 0 ( 8 0 . 2 )

S. alba a 55 (3 5 .7 ) 14 (13 .5 ) 7 ( 6 .7 ) 5( 4 .8 ) 11 ( 1 0 . 6 ) 7 2 (69 .2 )b 2 ( 3 .4 ) 1 ( 1 .7) 2 ( 3 .5 ) 1 ( 1 .7) 4( 6 .9 ) 10(1 7 . 2 )c 3 7 (2 2 .8 ) 1 5( 9 .3 ) 9 ( 5 .6 ) 6 ( 3 .7 ) 1 5( 9 .3 ) 8 2 ( 5 0 . 6 )

S . c as « a 1 4 (1 3 .5 ) 9( 8 . 6 ) 6 ( 5 .8 ) 11 ( 1 0 . 6 ) 9( 8 . 6 ) 49(47.1 )b 1 5 (2 5 .9 ) 7 (1 2 .1 ) 2 ( 3 .4 ) 1 ( 1 .7) 2( 3 .4 ) 2 7 (4 6 .6 )c 2 9 (1 7 .4 ) 16 ( 9 .9 ) 8 ( 4 .9 ) 12 ( 7 .4 ) 11( 6 .5 ) 76 (4 6 .9 )

Cocos a 6 ( 5 .8 ) 0 6 ( 5 .8 ) 2 ( 1 . 9 ) 7( 6 .7 ) 2 1 ( 2 0 . 2 )b 11 (1 9 .0 ) 1( 1 .7) 1 ( 1 .7 ) 0 3( 5 . 2 ) 1 6 ( 2 7 . 6 )c 1 7 (10 .5 ) 1 ( 0 . 6 ) 7 ( 4 .3 ) 2 ( 1 . 2 ) 1 0 ( 6 . 2 ) 3 7 (2 2 .8 )

Ceiba a 0 2 ( 1 . 9 ) 1 (1 . 0 ) 0 4( 3 .8 ) 7( 6 .7 )b 0 0 0 0 0 0c 0 2 ( 1 . 2 ) 1 ( 0 . 6 ) 0 4( 2 .5 ) 7( 4 .3 )

Eug. a 3( 2 .9 ) 6 ( 5 .8 ) 1 (1 . 0 ) 0 7( 6 .7 ) 17 (16 .3 )b 3( 5 . 2 ) 0 0 2( 3 .4 ) 2( 3 .4 ) 7 (1 2 .1 )c 6 ( 3 . 7 ) 6 ( 3 . 7 ) 1 ( 0 . 6 ) 2 ( 1 . 2 ) 9( 5 . 6 ) 2 4 (1 4 .8 )

Hang. a 2( 1 .9) 2 ( 1 . 9 ) 0 0 1 ( 1 . 0 ) 5( 4 . 8 )b 2 ( 3 . 5 ) 0 0 0 1 ( 1 .7) 3( 5 . 2 )c 4( 2 .5 ) 2 ( 1 . 2 ) 0 0 2 ( 1 . 2 ) 8 ( 4 .9 )

Durio a 0 0 0 0 2( 1 .9) 2 ( 1 . 9 )b 1 ( 1 .7) 0 0 0 1 ( 1 .7) 2( 3 .4 )c 1 ( 0 . 6 ) 0 0 0 3( 1 .9) 4( 2 .5 )

Rhiz. a 0 0 0 1 ( 1 . 0 ) 1 ( 1 . 0 ) 2 ( 2 . 0 )b 0 0 0 0 0 0c 0 0 0 1 ( 0 . 6 ) 1 ( 0 . 6 ) 2 ( 1 . 2 )

T a b le 8 .

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

p r o p o r t i o n s o f t h r e e p o l l e n s p e c i e s m o s t commonly c o n t a i n e d

i n p o l l e n s a m p le s c o l l e c t e d from t h e b o d i e s o r f a e c e s o f

M a c r o g lo s s u s m in im us c a u g h t a t R a n ta u P a n j a n g d u r i n g and

u 6 *tVv 6 Six f l o w e r i n g x lu s l i e S Ox 30nuej/8.t.lfci Bluy.#

P a r t A.

Each f i g u r e i s t h e p e r c e n t a g e o f t h o s e s a m p le s

c o n t a i n i n g t h e g i v e n p o l l e n t y p e i n ' w h i c h t h a t p o l l e n ty p e

made up 75% o r more o f t h e p o l l e n p r e s e n t ( e . g . d u r i n g

S . a l b a f l o w e r i n g f l u s h e s Jj-6.9% o f t h e s a m p le s c o n t a i n i n g

an y S . c a s e o l a r i s p o l l e n c o n t a i n e d 75% o r more S . c a s e o l a r i s

p o l l e n .

The f i g u r e s i n b r a c k e t s ' a r e t h e d i f f e r e n c e s b e tw e e n

t h e p e r c e n t a g e s r e c o r d e d d u r i n g an d b e tw e e n S . a l b a f l o w e r i n g

f l u s h e s ; i f + v e , t h e y i n d i c a t e t h a t more s a m p le s c o n t a i n e d

75% o r more o f t h a t p o l l e n t y p e d u r i n g S . a l b a f l o w e r i n g

f l u s h e s t h a n b e tw e e n them ( i . e . an i n c r e a s e i n t h e a b u n d a n c e

o f t h a t p o l l e n i n t h e s a m p le s i n v /h ich i t o c c u r r e d ) and v i c e

v e r s a .

P a r t B .

As i n P a r t A b u t e a c h f i g u r e i s t h e p e r c e n t a g e o f t h o s e

s a m p le s c o n t a i n i n g t h e g i v e n p o l l e n t y p e i n w h ic h t h a t p o l l e n

t y p e made up 2 5 % o r l e s s o f t h e p o l l e n p r e s e n t .

I f t h e d i f f e r e n c e s a r e +ve t h e y i n d i c a t e more s a m p le s

c o n t a i n e d 25% o r l e s s o f t h a t p o l l e n t y p e d u r i n g S . a l b a

f l o w e r i n g f l u s h e s t h a n b e tw e e n them ( i . e . a d e c r e a s e i n t h e

ab u n d a n c e o f t h a t p o l l e n i n t h e s a m p le s i n v /h ich i t o c c u r r e d )

and v i c e v e r s a .

Part A.

Samplefrom

S . alba During Eetween

S. caseolaris During Eetween

C. nucifera During Eetween

Faeces£R no/ 70 no/OU • V-y/C/ yu 9\J /<J

(+38 .0 )

». £ ro/ Oi co/ • y/u O i • yfv

( - 3 4 .6 )

O 0 /Trt/ nc r>o/ *-u • u/o r > »u/o

( - 4 6 . 4 )

Bodies71.25% 52.0%

(+19.25)

83.3% 75.7%

(+7.6)

0 0

(0)

Part E.

Samplefrom

S . alba S. caseolaris C. nuciferaDuring Between During Between During Between

Faeces15.3% 40.0%

( - 2 4 . 7 )

18.3% 7.4%

(+10.9)

33.3% 18.7%

(+14 .6)

Bodies10.0% 28.0%

( - 1 8 . 0 )

8.1% 6.1%

(+2.0

100.0% 60.0%

(+40 .0 )

101

h i g h e r b e tw e e n S . a l b a f l u s h e s t h a n d u r i n g them ( T a b l e 8 ) ,

B e s i d e s S o n n e r a t l a , t h e o n l y p o l l e n t h a t was r e g u l a r l y p r e s e n t

i n f a e c a l s a m p le s i n a s i g n i f i c a n t q u a n t i t y was t h a t o f t h e

c o c o n u t , Cocos n u c i f e r a , (C o c o n u ts w ere u b i q u i t o u s i n t h e

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

c o n t i n o u s i y , ) T h e re was o n ly a s l i g h t c h a n g e i n t h e f r e q u e n c y

o f o c c u r r e n c e o f c o c o n u t p o l l e n , i n t h e s a m p l e s , f rom 2 0 . 2 %

d u r i n g f l o w e r i n g f l u s h e s o f S . a l b a t o 27 .6% b e tw e e n them

( T a b l e 7 > a an d b , l a s t c o lu m n ) . H ow ever , b e tw e e n S . a l b a

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

o f c o c o n u t p o l l e n i n t h e s a m p le s i n w h ic h i t o c c u r r e d ( T a b le 8 ) .

A t R a n ta u P a n j a n g , M. m in im us f r e e l y f e d from a l l

t h e o t h e r p l a n t s l i s t e d i n T a b l e 6 when t h e y w ere a v a i l a b l e ,

e x c e p t R h iz o p h o r a s p p . C u l t i v a t e d b a n a n a s w e re an i m p o r t a n t

f o o d s o u r c e o f n e c t a r b u t t h e y p r o d u c e no p o l l e n a n d , t h e r e f o r e ,

do n o t f e a t u r e i n t h o s e s a m p l e s . The r e m a i n d e r a r e n o t i m p o r t a n t

fo o d s o u r c e s e i t h e r b e c a u s e t h e t r e e s a r e r a r e o r b e c a u s e t h e i r

s h o r t f l o w e r i n g s e a s o n s l i m i t t h e i r a v a i l a b i l i t y , o r b o t h . The

p o l l e n o f a l l t h e s e s p e c i e s c o n s e q u e n t l y fo rm ed a s m a l l p a r t

o f a l l t h e p o l l e n c o n t a i n e d i n t h e f a e c a l s a m p le s c o l l e c t e d

from M. m in im u s .

i f . i i . Body s a m p le s from R a n ta u P a n j a n g , S e l a n g o r .

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

from t h e b o d i e s an d m em branes o f M. m in im us a t R a n ta u P a n ja n g

a r e g i v e n i n T a b le 9 . P a r t s a an d b g i v e t h e p r o p o r t i o n s o f

e a c h p o l l e n t y p e d u r i n g ( a ) an d b e tw e e n (b ) f l o w e r i n g f l u s h e s

o f S . a l b a w h i l e p a r t c g i v e s t h e com bined r e s u l t . S o n n e r a t i a

T a b le 9*

The t a b l e r e c o r d s t h e a b u n d a n c e o f e a c h p o l l e n t y p e

i n p o l l e n s a m p le s c o l l e c t e d from t h e b o d i e s o f 159 M a c r o g lo s s u s

min im us a t R a n ta u P a n j a n g , S e l a n g o r .

a = s a m p le s c o l l e c t e d d u r i n g S . a l b a f l o w e r i n g f l u s h e s ( n = 1 1 0 )

b = ” ” b e tw e e n H ” (n = 4 9 )

c = t o t a l ( i . e . a + b co m b in ed ) (n = 1 5 9 )

The f i g u r e s show t h e num ber o f s a m p le s c o n t a i n i n g 100%, 75 t o

9 9 %) 50 t o 7 4 %) 2 5 t o 4 9 % o r l e s s t h a n 2 5 % ( b u t a t l e a s t one

p o l l e n g r a i n ) r e s p e c t i v e l y o f e a c h p o l l e n s p e c i e s . The f i g u r e s

i n b r a c k e t s a r e p e r c e n t a g e s o f t h e t o t a l num ber o f s a m p l e s .

N o te S . a l b a = S o n n e r a t i a a l b a (m a n g ro v e s )

S . c a s . = S o n n e r a t i a c a s e o l a r i s (m a n g ro v e s )

Cocos = Cocos n u c i f e r a ( c o c o n u t )

C e ib a = C e ib a p e n t a n d r a (k a p o k )*

E u g . = E u g e n ia m a l a c c e n s i s (M alay a p p l e )

Mang. = M a n g i f e r a s p p . (mango)

*T h i s p o l l e n t y p e v/as i d e n t i f i e d t o g e n u s l e v e l o n l y . B a t s

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

v/as t h e o n l y one r e c o r d e d a t R a n ta u P a n j a n g w h ic h b a t s w e re

v i s i t i n g .

Species 100% 75-99% 5 0 - 74% 2 5 - 4 9% < 25% Total

S.alba a 7 7 (7 0 .0 ) 1 2 (1 0 .9 ) 3( 2 .7 ) 0 3 ( 2 .7 ) 9 5 (8 6 .4 )+ b 3 4 (6 9 .4 ) 6 ( 1 2 .2 ) 1( 2 .0 ) 0 5 (1 0 .2 ) 4 6 (9 3 .9 )

S.cas. c 1l i ( 6 9 .8 ) 1 8 ( 1 1 . 3 ) 4( 2 . 5 ) 0 8( 5 , 0 ) 1 4 1 (8 8 ,7 )

S .alba a **3(39.1) 14 (1 2 .7 ) 1 5 (1 3 .6 ) 0 8( 7 .3 ) 8 0 (7 2 .7 )b 9 ( 1 8 .**) 4( 8 .2 ) 0 5 (1 0 .2 ) 7 (1 4 .3 ) 25(51 .0)c 52 (32 .7 ) 18(11 .3) 1 5( 9 .4 ) 5( 3 .1 ) 15( 9 .4 ) 105(66 .0 )

S.cas. a 2 4 ( 2 1 .8) 7( 6 .4 ) 0 3( 2 .7 ) 3 ( 2 .7 ) 3 7 (3 3 .6 )b 18 (36 .7 ) 7 (1 4 .3 ) 6 (1 2 .2 ) 0 2( 4 .1 ) 3 3 (6 7 .3 )c 4 2 (2 6 .7 ) 14( 8 .8 ) 6( 3 .8 ) 3( 1 .9 ) 5( 3 .1 ) 7 0 (44 .0 )

Cocos, a 0 0 0 0 7( 6 .4 ) 7( 6 .4 )b 0 0 0 2( 4 .1 ) 3( 6 .1 ) 5 (1 0 .2 )c 0 0 0 2( 1 .3) 10( 6 .3 ) 12( 7 .5 )

Ceiba a 0 4( 3 .6 ) 3( 2 .7 ) 0 10( 9*1) 1 7 (1 5 .5 )b K 2 .0 ) 0 2( 4 .1 ) 2( 4 .1 ) 6 (1 2 .2 ) 11 (2 2 .4 )c 1( 0 .6 ) 4( 2 .5 ) 5( 3 .1 ) 2( 1 .3) 1 6 (1 0 .1 ) 2 8 (1 7 .6 )

Eug. a 0 0 0 4( 3 .6 ) 13(11 .8) 1 7 (1 5 .5 )b 2( 4 .1 ) 0 0 2( 4 .1 ) 5 (1 0 .2 ) 9 (1 8 .4 )c 2( 1 .3 ) 0 0 6( 3 .8 ) 18(11 .3) 2 6 (1 6 .3 )

Mang. a 0 0 0 0 8( 7 .3 ) 8( 7 .3 )b 0 0 0 0 3( 6 .1 ) 3( 6 .1 )c 0 0 0 0 11( 6 .9 ) 11( 6 .9 )

102

was c l e a r l y t h e m o s t i m p o r t a n t p o l l e n g e n u s w i t h 1 1 1 ( 6 9 *8 %)

o f t h e 159 s a m p le s c o n t a i n i n g o n l y S o n n e r a t i a p o l l e n and 141

(88 .7% ) c o n t a i n i n g a t l e a s t some o f i t .

T a b le 9 > p a r t s a and b ( l a s t co lum n) show t h a t

t h e f r e q u e n c y o f o c c u r r e n c e o f S . a l b a p o l l e n i n t h e s a m p le s

w as r e d u c e d f ro m 72 .7% d u r i n g S . a l b a f l o w e r i n g f l u s h e s t o

5 1 • 0 % b e tw e e n th e m . T h ese t r e n d s a r e s i m i l a r t o b u t s m a l l e r

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

p r o p o r t i o n o f S . a l b a i n t h o s e s a m p le s i n v /h ich i t o c c u r r e d

was a l s o r e d u c e d b e tw e e n f l u s h e s ( T a b le 8 ) .

D i f f e r e n c e s i n t h e q u a n t i t i e s o f S . c a s e o l a r i s

p o l l e n i n t h e s e s a m p le s d i d n o t f o l l o w t h e t r e n d s o b s e r v e d i n

t h e f a e c a l s a m p l e s . Thus i n t h e s e s a m p l e s , t h e f r e q u e n c y v / i th

w h ich i t was p r e s e n t d o u b le d from 53 .6% d u r i n g S . a l b a f l o w e r i n g

f l u s h e s t o 67*3% b e tw e e n them ( T a b l e 9 , p a r t s a and b , l a s t

c o lu m n ) . H ow ev er , t h e . p r o p o r t i o n o f S . c a s e o l a r i s p o l l e n i n

t h o s e s a m p le s i n w h ic h i t o c c u r r e d r e m a in e d f a i r l y c o n s t a n t

( T a b l e 8 ) . T h i s i s t h e t r e n d t h a t was e x p e c t e d i n t h e f a e c a l

s a m p l e s .

The p o l l e n o f C e ib a p e n t a n d r a an d E u g e n ia m a l a c c e n s i s

v/as commonly r e c o v e r e d f rom t h e h e a d s an d abdom ens o f M. m in im us

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

t y p e s w ere r a r e l y e n c o u n t e r e d . P o l l e n o f D u r io an d R h iz o p h o r a

w ere n e v e r f o u n d on t h e b o d i e s o f t h i s s p e c i e s o f b a t a t R a n ta u

P a n j a n g .

4 * i i i . F a e c a l and bo d y s a m p le s from Bako N a t i o n a l P a r k .

T a b le 10 shows t h e n u m e r i c a l p r o p o r t i o n s o f p o l l e n

Table 10.

The table records the abundance of each pollen type

found in 23 faecal samples and 10 samples from the bodies of

Macroglossus minimus caught at Telok Asam, Eako National Park,

Sarawak.

The figures show the number of samples containing

100%, 75 to 99%, 50 to 74%, 25 to 49% or less than 25% (but

at least one pollen grain) respectively of each pollen species.

The figures in brackets are percentages of the total number of

samples.

From the faeces of the bats (n = 23).

100% 75-99% 50-74% 25-49% < 25% total

S.alba

Bug.

Durio

?

19(82.6)

0

0

0

1( 4.3)

2( 8.7)

0

0

2( 8.7)

0

0

0

0

2( 8.7)

0

0

1( 4.3)

2( 8.7)

1( 4.3)

1( 4.3)

2 3 (100 .0 )

6( 26.1)

K *f.3)

1( 4.3)

From the bodies of the bats (n =10)

S.alba 7 (7 0 .0 ) 2 ( 2 0 .0 ) 1 (1 0 .0 ) 0 0 10(100 .0 )

Bug. 0 0 0 1 (1 0 .0 ) 2 ( 2 0 .0 ) 3( 30 .0 )

Note S.alba = Sonneratia alba.

Eug. = Eugenia spp (species unknown) .

Durio = Durio spp. (possibly D. carinatus).

103

s p e c i e s p r e s e n t i n 2 3 f a e c a l s a m p le s an d 10 s a m p le s from t h e

b o d i e s o f M„ m in im u s c a u g h t i n Bako N a t i o n a l P a r k in. J a n u a r y

and May 1 9 7 3 . E x c e p t f o r t h r e e f a e c a l s a m p le s fro m b a t s c a u g h t

i n t h e s c r u b b e h i n d t h e p a r k b u i l d i n g s i n May 19 7 3 j a l l t h e

s a m p le s w e re c o l l e c t e d from b a t s m i s t n e t t e d a l o n g t h e b r i d g e

t h r o u g h t h e S . a l b a g r o v e ( F i g s . 7 and 8 ) .

S . a l b a was f l o w e r i n g d u r i n g b o t h v i s i t s a l t h o u g h

t h e r e w ere fe w e r f l o w e r s i n May when t h e t r e e s w e re n e a r i n g

t h e end o f a f l o w e r i n g f l u s h .

S . a l b a p o l l e n o c c u r r e d i n e v e r y s a m p le and i t

c o n t r i b u t e d more t h a n 50% o f t h e p o l l e r f p r e s e n t i n a l l e x c e p t

two o f th e m . An unknown s p e c i e s o f E u g e n ia was t h e rimly o t h e r

p o l l e n t y p e t h a t o c c u r r e d f r e q u e n t l y . I t was p r e s e n t i n J a n u a r y

a n d May. I n J a n u a r y , one s m o o th , t r i - p o r a t e D u r io s p p . p o l l e n

g r a i n v/as fo u n d i n a f a e c a l s a m p l e . The s p e c i e s i s unknown b u t

i t may h a v e b e e n D. c a r i n a t u s v /h ich i s common a t T e lo k Asam.

(Many t r e e s h a d young f r u i t s i n d i c a t i n g t h a t t h e y h a d f l o w e r e d

r e c e n t l y and one o r two may s t i l l h a v e b e e n i n f l o w e r . ) T h e re

was one o t h e r u n i d e n t i f i e d p o l l e n g r a i n i n one f a e c a l s a m p le

i n J a n u a r y .

4 » i v . Samples obtained from elsewhere.

A few p o l l e n s a m p le s from M. m in im us v/ere o b t a i n e d

e l s e w h e r e . A t .K u a la S e l a n g o r e l e v e n f a e c a l s a m p le s an d f o u r

s a m p le s from t h e b o d i e s o f b a t s w ere c o l l e c t e d from M. m in im u s ..

m i s t n e t t e d a d j a c e n t t o f l o w e r i n g S o n n e r a t i a a l b a t r e e s on

12 F e b r u a r y 1 9 7 2 . T h e re w ere many k a p o k ( C e ib a p e n t a n d r a ) t r e e s

104

g ro w in g n e a r b y . P o l l e n r e c o v e r e d from t h e i r b o d i e s was a l l

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

p r o p o r t i o n o f C. p e n t a n d r a ( T a b l e 11 ) .

On 30 December '1972 a t Tawau, S a b a h , two p o l l e n

s a m p le s w e re c o l l e c t e d from t h e h e a d s o f M. m in im us c a u g h t i n

m i s t n e t s s e t among m a n g ro v e s f r i n g i n g a t i d a l s t r e a m t h a t r a n

th r o u g h a r u b b e r e s t a t e . A l th o u g h S o n n e r a t i a a l b a grew n e a r b y ,

no t r e e s w e re fo u n d i n f l o w e r t h e n . An unknown s p e c i e s o f

E u g e n ia fo rm e d 89.2% and 99.9% r e s p e c t i v e l y o f t h e p o l l e n

g r a i n s i n t h e s e s a m p l e s . A l l t h e r e m a i n d e r w e re C e ib a p e n t a n d r a

T h e re was no S . a l b a p o l l e n .

S a m p le s w e re o b t a i n e d from t h e a l i m e n t a r y c a n a l s

o f p r e s e r v e d s p e c i m e n s i n t h e U n i v e r s i t y o f M a lay a c o l l e c t i o n ,

t h r e e from K u a la G u la and f i v e f rom P u la u K e tam . A l l c o n t a i n e d

t h e p o l l e n o f S o n n e r a t i a a l b a b u t one a l s o c o n t a i n e d a l i t t l e

( l e s s t h a n 1%) R h iz o p h o r a s p p . p o l l e n .

5 . I t e m s o t h e r t h a n p o l l e n i n t h e f a e c e s o f M. m in im u s .

M ost f a e c a l p e l l e t s from M. m in im us v a r i e d i n

c o l o u r from b r i g h t y e l l o w t o w h i t e . They w e re s u f f i c i e n t l y

f i r m t o r e t a i n t h e i r c y l i n d e r i c a l s h a p e ev en when h a n d l e d w i t h

f o r c e p s . They w ere made up a l m o s t e n t i r e l y o f p o l l e n e x i n e s ,

b u t from t i m e t o t i m e a w ide v a r i e t y o f o t h e r i t e m s w e re some­

t i m e s p r e s e n t . I t e m s b e s i d e s p o l l e n fo u n d i n f a e c a l s a m p le s

f rom R a n ta u P a n j a n g may b e d i v i d e d i n t o t h r e e g r o u p s

I t e m s o f unknbwn o r i g i n .

^ • I t e m s o f p l a n t o r i g i n ,

i i i . I t e m s o f a n i m a l o r i g i n .

Table 11 .

Table recording the proportions of Sonneratia alba and Ceiba pentandra pollen in faecal samples collected from 11 Macroglossus minimus at Kuala Selangor on 12 February 1972. No other pollen types were present in the samples.

No. of samples % S . alba % C. pentandra

6 100.0 -1 99.9 0.1

1 97.3 2.7

1 88.9 11 .1

1 5.3 94.7

1 5.1 94.9

II

/

105

i . Items o f unknown o r i g i n .

Amorphous m a t t e r , w h ich a p p e a r e d g r a n u l a r u n d e r

t h e m i c r o s c o p e , was p r e s e n t i n 45 ( 2 7 . 8 %) o f t h e f a e c a l s a m p le s

from R a n ta u P a n j a n g . I t was n o t s t a i n e d b y b a s i c f u s c h i n . When­

e v e r i t was common i n f a e c a l p e l l e t s , t h e y t e n d e d t o b e d a r k e r

i n c o l o u r ( o f t e n d a r k g r e e n o r b row n) and s e m i—l i c j u i d i n

c o n s i s t e n c y . I n e i g h t s a m p le s i t was a c c o m p a n ie d b y s m a l l

d r o p l e t s o f o i l w h ic h s t a i n e d r e d i n Sudan IV .

ii. Items of plant origin.

F r a g m e n t s o f e p i d e r m a l t i s s u e w e re p r e s e n t i n

9 (5*6%) o f t h e s a m p le s w i t h s t o m a t a p r e s e n t i n 3 ( F i g . 4 7 ) . The

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

o f v a s c u l a r c e l l s w i t h h e l i c a l t h i c k e n i n g s i n t h e i r w a l l s w ere

r e c o r d e d f i v e t i m e s , i n t h r e e t h e y w ere a s s o c i a t e d w i t h e p i d e r m a l

t i s s u e . T h ese p l a n t p a r t i c l e s w e re a c c o m p a n ie d b y am orphous

m a t t e r i n 7 (64%) o f t h e 11 s a m p le s i n w h ic h t h e y w ere f o u n d .

F u n g a l s p o r e s w e re p r e s e n t i n 11 ( 6 .8 % ) . They w e re a c c o m p a n ie d

'b y f u n g a l h y p h a e i n f o u r and s p o r a n g i a w e re a l s o fo u n d i n one

o f th e m . They w e re a p p a r e n t l y a l l p h y c o m y c e te s ( D r . Nawawi,

p e r s , comm.)

iii. Items of animal origin.

These were of three types

a . Arthropod remains. The following arthropod - remains were

found in the faecal samples from Rantau Panjang.

ARACHNIDA. A c a r i n a ( m i t e s ) . M i t e s w ere fo u n d i n 15 (9 .1% ) s a m p le s

( F i g . 47 ) •. One s a m p le , o n ly a s m a l l p o r t i o n ( a b o u t

10%) o f t h e w h o le f a e c a l p e l l e t , c o n t a i n e d 36 m i t e s

e . Fungal s p o r e s .

c . Nematodes( n o te a l s o one mammalian h a i r (H) .)

d .E p id e rm a l t i s s u e i n c l u d i n g s to m a ta

F ig . if7. I te m s found i n th e f a e c e s of M acroglossus s o b r i n u s( a , b , c , d ) and i n th e s tomachs o f E o n v c t e r i s s p e l a e a (e)

l'< f e (A car ina ) .

106

o f a t l e a s t two s p e c i e s .

A r a n e i d a ( s p i d e r s ) . A s m a l l s p i d e r w as fo u n d i n one

s a m p le .

INSECTA. T h y s a n o p t e r a ( t h r i p s ) . T h r i p s w ere fo u n d i n e i g h t

s a m p l e s .

D i p t e r a ( f l i e s ) . One s m a l l n e m a t o c e r a n was fo u n d

i n e a c h o f two s a m p l e s .

H y m e n o p te r a ( a n t s , b e e s and w a s p s ) . S e v e r a l s m a l l

a n t s w e re fo u n d i n t h r e e s a m p le s a nd u n i d e n t i f i e d ,

s m a l l , w in g e d h y m e n o p te r a n s i n tw o .

Most o f t h e s e a n i m a l s w e re v e r y s m a l l ( i . e . n o t more t h a n 1 mm

l o n g ) . T h e i r b o d i e s w ere se ld o m damaged e x t e n s i v e l y an d t h e y

f r e q u e n t l y r e t a i n e d a l l t h e i r a p p e n d a g e s , i n c l u d i n g w in g s i n

w in g e d i n s e c t s . D e t a i l s o f i n t e r n a l ana tom y w ere s t i l l v i s i b l e

i n some o f th e m . They w e re p r o b a b l y i n g e s t e d a c c i d e n t a l l y ;

w h i l e t h e b a t s w e re f e e d i n g from f l o w e r s .

R em a in s o f l a r g e r i n s e c t s w ere a l s o p r e s e n t . T h e re

w e re i n s e c t s c a l e s , p o s s i b l y from m o th s , i n 2^ (1 i f .8%) o f t h e

s a m p l e s . T h ese w ere a c c o m p a n ie d b y i n s e c t t a r s i , p r o b a b l y a l s o

from m o th s an d odd f r a g m e n t s o f b row n c h i t i n i n 7 o f t h e s a m p l e s .

The f a c e o f one a d u l t f e m a le c a u g h t on 25 J u l y 1973 was c o v e r e d

b y i n s e c t s c a l e s . The b a t was d i s s e c t e d an d i t s s to m a c h c o n t e n t s

ex am in ed i n t h e l a b o r a t o r y . R em ain s o f two i n s e c t l e g s , a t a r s u s

an d a t a r s u s w i t h p a r t o f t h e a t t a c h e d t i b i a w e re p r e s e n t . Each

t a r s u s was a p p r o x i m a t e l y 2 mm lo n g and t h i c k l y c o v e r e d w i t h

h a i r - l i k e s c a l e s . T h e re w e re a l s o many t i n y f r a g m e n t s o f brown

c h i t i n . U nder t h e m i c r o s c o p e , t h e r e m a i n d e r o f t h e g r e y s to m a ch

/

/II

107

c o n t e n t s w e re com posed e n t i r e l y o f am o rp ho us m a t t e r , i n s e c t

s c a l e s , f a t d r o p l e t s an d t h e o c c a s i o n a l e x i n e s o f c o c o n u t

p o l l e n g r a i n s .

b . Mammalian hair, indistinguishable from that of M. minimus

itself, was present in 90 (55 .6% ) of the Rantau Panjang samples.

This was undoubtedly ingested by bats cleaning pollen from their

fur or when grooming.

c . P a r a s i t e s o f t h e a l i m e n t a r y c a n a l . V ery l a r g e nu m b ers o f

s m a l l n e m a to d e s w e re fo u n d i n t h r e e f a e c a l s a m p le s ( F i g . 4 7 ) .

C e s to d e o n c h o s p h e r e s , c h a r a c t e r i s e d b y t h e s i x h o o k s o f t h e

h e x a c a n t h em b ry o , w e re p r e s e n t i n tw o , one o f w h ic h a l s o c o n t a i n e d

s e v e r a l , d e t a c h e d , c e s t o d e p r o g l o t t i d s . ( T h r e e c e s t o d e s w e re

fo u n d i n t h e i l i u m o f one s p e c im e n c a u g h t a t R a n ta u P a n j a n g . )

6 . F l i g h t .

The b o d i e s o f M. m in im us a r e n o t u n d u ^ l y com p ac t

o r s t r e a m l i n e d . The f u r i s t h i c k an d l o n g . I t d o e s n o t l i e

c l o s e l y a g a i n s t t h e b o d y .

D a ta on w ing m e a s u re m e n ts a r e g i v e n i n T a b l e s 12

an d 1 3 . T h e re v/as very- l i t t l e d i f f e r e n c e i n t h e mean l e n g t h o f

t h e f o r e a r m , 3 r d d i g i t o r 5 ^ d i g i t o f a d u l t m a le s a nd a d u l t

f e m a l e s b u t n o n - p a r o u s f e m a l e s w e ig h e d l e s s t h a n a d u l t m a l e s .

The i n d i c e s r e c o r d e d i n T a b le 12 a r e i n d e p e n d e n t o f w e i g h t an d

t h e d a t a f o r a l l a d u l t s a r e co m b ined b u t t h o s e r e c o r d e d i n

T a b le 13 a r e r e l a t e d t o w e i g h t and s o a d u l t m a le s an d n o n - p a r o u s

a d u l t f e m a l e s a r e t r e a t e d s e p a r a t e l y . Compared t o o t h e r p t e r o p i d s

( F i n d l e y , S t u d i e r an d W i l s o n , 1972) M. m in im u s h a s l o n g , b r o a d

w in g s w i t h a l o n g w in g t i p t o f o r e a r m r a t i o a nd t h u s a .t

Table 12.

The t a b l e shows some wing c h a r a c t e r i s t i c s o f

M acroglossus minimus , M. s o b r i n u s and E o n y c te r i s s p e l a e a .

(These c h a r a c t e r i s t i c s a r e in d e p e n d e n t o f w e ig h t and so

d a t a f o r a d u l t s o f b o t h s s ex e s a r e combined.)

F.A. Tip in d e x A.R. in d e x Area in d ex

M.s .

(1)

Mean

n

Range

44.1 7+0.96

19

42 . 2 - 4 5 . 5

1 .92+0 .03

18

1 . 8 6 - 1 .9 7

1 .98+0.06

18

1 .7 4 - 2 .0 3

1 68.63+1 0 .06

18

149.2-1 90 .9

M.m.(2)

Mean

n

Range

42.40+1 .39

34

4 0 . 0 - 4 5 . 0

1 .91+0.05

34

1 . 8 1 - 2 . 0 2

1 .98+0 .04

34

1 .9 0 -2 .0 7

153 .73+8.64

34

133.4-1 71 .6

E . s .

(3)

Mean

n

Range

65 .6 2 + 2 .3 ?

62

61 . 0 - 7 0 .0

1 .73+0 .05

62

1 .61 - 1 .8 3

2 .3 3 + 0 .0 5

62

2 . 2 5 - 2 . 4 3

275.63+21 .77

62

2 2 6 .8 -3 2 6 .4

p = 0.01 1 / 2 / 3 0 = 2 ) / 3 ( l = 2 ) / 3 1 / 2 / 3

N . B . j - M .s . = M acrog lossus s o b r i n u s ; M.m. = M. min imus;

E . s . = E o n y c te r i s s p e l a e a .

F.A. = fo rearm and A.R. = a s p e c t r a t i o .

Table 13.

The t a b l e shows th e w e ig h t and two w e ig h t dependen t

c h a r a c t e r i s t i c s o f M acrog lossus s o b r i n u s , M. minimus and

E o n y c te r i s s p e l a e a . (A du l t males and n on-pa rous fem ales a r e

t r e a t e d s e p a r a t e l y . )

M . s . O ) M.m.(2) E . s . (3) p = 0 .0 1

Weight

c?

Mean

n

Range

21 .67+1 .03

9

2 0 . 0 - 2 3 . 0

1 8 .6 7 + 1 .80

9

1 6 . 0 - 2 1 . 0

5 9 .03+4 .09

26

5 2 .5 - 7 0 .0

( 1 =2 ) #

(gins)

?

Mean

n

Range

2 1 .58+1.63

6

2 0 . 0 - 2 4 . 0

1 7.6+1 .39

22

1 5 . 0 - 1 9 . 0

4 8 .6 7 + 5 .0 8

27

41 . 0 - 5 7 . 5

( 1 =2 ) #

Loading

Mean

n

Range

0 .1 3 +0 .01

8

0 .11 - 0 .1 4

0 .12+0 .009

9

0 . 1 0 - 0 . 1 3

0 . 2 0 +0 .01

25

0 . 1 8 - 0 . 2 4

( 1 =2 ) #

in d ex

?

Mean

n

Range

0 . 13+0 . 0 0 8

6

0 . 1 2 - 0 . 1 4

0 .11+0 .009

13

0 . 1 0 - 0 . 1 3

0 .1 8 +0 .01 8

27

0 . 1 4 - 0 . 2 2

( 1 =2 ) #

Length

Mean

n

Range

5 .95+0 .26

8

5 . 7 0 - 6 . 4 5

6 .70+0 .58

9

6 .1 0 - 7 .7 5

3 . 1 2 +0 . 1 8

25

2 . 8 6 - 3 . 5 0

1/ 2#

in d ex

?

Mean

n

Range

6 .05+0 .37

6

5 .7 4 - 6 .5 0

7 .15+0 .52

13

6 . 2 0 - 8 .0 7

3 .6 5 J P .3 6

27

3 . 1 3 - 4 . 2 2

1 / 2 #

108 .

low w in g l o a d i n g i n d e x . T h e se c h a r a c t e r s s u g g e s t s lo w f l i g h t

an d g r e a t m a n o e u v r a b i l i t y .

6 . i i . F l i g h t .

•M. m in im us f l i e s a t a l l h e i g h t s , f rom a b o u t 10 cm

a b o v e g ro u n d l e v e l ( o v e r t h e dyke a l o n g s i d e jS. a l b a t r e e s ) t o

t h e t o p s o f t h e t r e e s , H ow ev er , when f e e d i n g from b a n a n a f l o w e r s

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

f rom b e lo w .

They do n o t f l y f a s t b u t a r e v e r y a g i l e an d c an

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

o f S o n n e r a t i a an d E u g e n ia t r e e s w i t h r e m a r k a b l e d e x t e r i t y . I n

c a p t i v i t y t h e y c o u l d f l y b a c k an d f o r t h a c r o s s t h e i r c a g e (1 m

x 1 m x 1 .3 m h i g h ) h o v e r i n g b r i e f l y i n f r o n t o f t h e g a u z e

n e t t i n g on t h e s i d e s b e f o r e t u r n i n g a ro u n d ( F i g . k°l ) . When

f o r a g i n g , t h e i r f l i g h t p a t h s w e re e r r a t i c . H a v in g f e d from ’ one

f l o w e r t h e y w o u ld c i r c l e t h e t r e e o r f l y s e v e r a l y a r d s away

b e f o r e r e t u r n i n g t o f e e d on a n o t h e r f l o w e r ( e x c e p t when f e e d i n g

from 13. m a l a c c e n s i s when t h e y u s u a l l y r e m a in e d w i t h i n t h e can o p y

b e tw e e n s u c c e s s i v e v i s i t s t o . f l o w e r s ) . A t y p i c a l f l i g h t p a t h

f o r one b a t f e e d in g jb n S . a l b a f l o w e r s a t s i t e 1 a t E a n ta u

P a n j a n g i s i l l u s t r a t e d i n Fig-. 5 0 .

6 . i i i . A c t i v i t y t i m e .

T h e re was a p e a k o f a c t i v i t y i n t h e f i r s t two

h o u r s a f t e r d a r k , i . e . f rom a p p r o x i m a t e l y 1 9 . 1 5 t o 2 1 . 1 5 h r s .

T h i s f e l l o f f b e tw e e n 2 1 .1 5 and 2 2 .1 5 h r s an d r e m a in e d a t a

low l e v e l f o r t h e r e s t o f t h e n i g h t . No p red aw n p e a k i n a c t i v i t y

w as r e c o r d e d . T h i s i s r e f l e c t e d i n t h e num ber o f b a t s c a u g h t

p e r h o u r a t R a n ta u P a n j a n g . The n um bers a r e e x p r e s s e d a s

F i g . ^ 8 . M acrog lossus s o b r i n u s - showing th e upward d i r e c t e d eyes s u i t a b l e f o r s e a r c h i n g f o r Musa i n f l o r e s c e n c e s from be low .

Fig* 4 9 . M acrog lossus minimus h o v e r in g i n one c o rn e r o f i t s cage .N o t e : - 1 . The h in d l imbs a r e h e l d t o g e t h e r to

g iv e a maximum wing a r e a .2 . The body i s h e ld a t an a c u t e a n g le

to th e h o r i z o n t a l .

F i g . 5 0 . D iagram sh o w in g t y p i c a l f l i g h t p a t h s f o r E o n y c t e r i s

s p e l a e a ( —— ) and. M a c r o g lo s s u s mi n i mu s (•— )

v i s i t i n g S o n n e r a t i a a l b a f l o w e r s . (The p a t h f o r

m in im us i s b a s e d on a p a t h r e c o r d e d a t S i t e # 1 ,

R a n ta u P a n j a n g , S e l a n g o r . T h a t f o r E . s p e l a e a i s

e n t i r e l y d i a g r a m m a t i c . )

DYKEM A N G R O V E FOREST

S O N N E R A T I A

A L B A

S O N N E R A T I AALBA

S O N N E R A T I AA L B A

109

W i l l i a m s ' mean ( T a b l e li+, F i g . 51 ) « ( W i l l i a m s ' mean i s . c a l c u l a t e d

from : -

w here M .= W i l l i a m s ' m ean, wn = t h e num ber o f b a t s c a u g h t p e r h o u r an d

N = t h e num ber o f h o u r s f o r w h ic h v a l u e s o f n a r e a v a i l a b l e .

T h i s m o d i f i c a t i o n o f t h e f o r m u l a f o r a g e o m e t r i c mean a l l o w s

v a l u e s o f n = 0 t o b e i n c l u d e d (Haddow, 1 95A-> 1 9 6 0 ) . I t h a s

t h e a d v a n t a g e o v e r t h e n o rm a l mean o f r e d u c i n g t h e d i s t o r t i o n s

p r o d u c e d b y a b n o r m a l l y l a r g e o r s m a l l v a l u e s o f n . )

t h e l o c a t i o n o f t h e n e t s was ch a n g e d d u r i n g t h e n i g h t ( F i g . 5 2 ) .

T h e r e f o r e , t h e d r o p i n n um b ers o f b a t s c a u g h t p e r h o u r a f t e r

2 1 . 1 5 h r s was g e n u i n e a nd n o t an a r t i f a c t c a u s e d b y t h e b a t s

l e a r n i n g t h e p o s i t i o n s o f t h e n e t s and s u b s e q u e n t l y a v o i d i n g

th em .

7 . A bundance a t Bako N a t i o n a l P a r k com pared t o R a n ta u P a n j a n g .

I n c o m p a r i s o n t o R a n ta u P a n j a n g , v e r y few M. m in im us

w ere s e e n i n t h e m a n g ro v e s a t B a k o . T h i s s u b j e c t i v e v iew was

s u p p o r t e d by c o m p a r in g t h e c a p t u r e r a t e . o f b a t s a t e a c h l o c a l i t y .

A mean o f 1 .0 6 M. m in im us w e re c a u g h t p e r n e t p e r n i g h t a t Bako

com pared t o a n e s t i m a t e o f 7 »kk p e r n e t p e r n i g h t a t . R a n t a u

P a n j a n g . (An e s t i m a t e o f t h e number o f b a t s c a u g h t p e r n e t p e r

n i g h t a t R a n ta u P a n ja n g was o b t a i n e d by summing W i l l i a m s * mean

Log (Mw + 1 )lo g - (n + 1 )

The a c t i v i t y p a t t e r n r e m a in e d c o n s t a n t e v en when

T a b le 11f• The t a b l e shows t h e t o t a l , mean and W i l l i a m s 1 mean

f o r t h e num ber o f M a c r o g lo s s u s m in im us c a u g h t p e r

h o u r d u r i n g e a c h h o u r o f t h e n i g h t a t R a n ta u

P a n j a n g , S e l a n g o r d u r i n g 1972 and 1973*

F i g . 51• The f i g u r e i l l u s t r a t e s g r a p h i c a l l y t h e d a t a g i v e n

i n T a b le 1if .

W i l l i a m s ’ m e a n

1 9 - 0 0 - 1 9 - 1 5 1

------- 1-------- 1--------1_____ ■ ■ ■

19-15 - 20-15 ||1 1

20-15 - 21-15 i l l ! !

21-15 - 22-15 i l l

22-15 - 23-15 ! ! ! ! !------- ,—

:: i

00-15 - 01-15 1 1 1 !

01-15 - 02-15 I I

02 - 15 - 03-15 1 1 1

0 3-15 - 04-15 I I I !

04-15 - 05-15 I I I !

05-15 - 06-15 I I I !

o o o o O O ro rv ro roVn -P- vn rv —* O VN ro —* o VO VO• • • • • • « • • • • •—* —» —» —* —- --» —* —» —» —» —» O i-3Vn VJ1 VJ1 vn vn vn vn vr vn vn vn o H-1 i i i i i i i i l i 1 ao o o o o o o rv rv ro rv —- CDCTs Vn -p- VN rv —* o Vn rv —* o vO• • • • • • • • • • • •—* —* —- —» —» —■ —» —» —A —* —» —»vn vn vn Vn vn VJl vn vr vn vn vn vn

p sH- O

rv rv -P- VJ1 vo —* —* Vn -P- -p- vn vn cq »—* 00 —• ro oo O o S 'cr- OC0 Mj

a ^rv VN <Ts CO -p- ro VN vn —• ro rv VN cd o

ro OV -VI _* O vn (-■ •-P- cr\ - a co

CO o

m, s ;CD o

VN ro CTv vn ~s3 ro -P- -P- —» rv rv rv 3 •—- —* O) -p- rv -p- cd—- — * VN 1—* oCD MjCO

v-3vn vn —- —- —* -p- -VI —' rv -p- -P- vn O

ro vn —* VN -VJ o vn rv OO c+

vn vn -VJ o CD1—1

t-3o

rv rv vn ro _» V* -P- VN cr\ OO vO o c-r• • • • • • • • • • • • 35vn. vn o CT\ ro vO VN VM —» vO ON (—1

S'P

3 i—1ro r\j rv —* o VN VN rv -P- crv o CD M• • • • • • • • • • • • CD H-vn vn o vO -O av ON -P- rv -o •p- 3 CD£3

03

F i g , 52. H is tog ram to show th e number o f M acrog lossus minimus

c a u g h t p e r hour on t h r e e n i g h t s on which th e m is t

n e t s were moved from one n e t s i t e t o a n o t h e r a t

ab o u t m id n ig h t . Data o b t a in e d a t Rantau P an ja n g ,

S e l a n g o r .

5

3

2

1

0

in in in in in in in in in mrH rH rH »—1 1—1 rH rH rH rH rH

•

O

•

rH•

CM

•

CO

•

0•

rH

•

CM

•

CO

• •

inCM

|

CM

|CM

ICM

101

0 0 0 O O

in in in in inI

in1

in

1

in1

in1

mH rH rH rH rH rH rH rH r*H rH• «

O

•

rH

•

CM

•

CO

«

0•

rH CM

•

CO

•

rH CM CN CM CM 0 O O O 0

Time

05

.15

-06

.15

■ II ' '

11°

f o r t h e num ber o f b a t s . c a u g h t p e r h o u r ( T a b le 1 4 ) an d d i v i d i n g

t h e sum b y t h e u s u a l num ber o f n e t s s e t p e r n i g h t ( i . e . 5 ) .

T h i s t a k e s i n t o a c c o u n t t h e ch an g e i n t h e num ber o f b a t s

c a u g h t p e r h o u r d u r i n g t h e n i g h t . S i n c e t h e n e t s w ere s e t a l l

n i g h t a t Bako t h e num ber o f b a t s c a u g h t t h e r e p e r n e t n i g h t i s

SS..GIL a-jr Ccll.Cu.lS.o6 Cl 9 S ilu o in C S oil 6 SSiuC XictS W65T6 U 56u So uOoli

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

The s u b j e c t i v e v iew i s a l s o s u p p o r t e d b y t h e -

num ber o f b a t s v i s i t i n g S o n n e r a t i a a l b a f l o w e r s a t e a c h l o c a l i t y .

Twice a t R a n ta u P a n j a n g t h r e e jS . a l b a f l o w e r s w e re o b s e r v e d f o r

15 m i n u t e s i n e a c h h o u r b e tw e e n 1 9 .3 0 and 0 0 . 4 5 h r s an d once

t h r o u g h o u t t h e n i g h t . T h i s was r e p e a t e d i n Bako N a t i o n a l P a r k

on two c o m p l e t e n i g h t s i n J a n u a r y 1 9 7 3 . The mean num ber o f b a t s

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

i s g i v e n i n F i g , 53 * ( I t i s p o s s i b l e t h a t some o f t h e b a t s

r e c o r d e d a f t e r 2 0 .3 0 ' h r s w e re E o n y c t e r i s s p e l a e a b u t t h a t s p e c i e s

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

and i t s l a r g e r s i z e . ) At R a n ta u p a n j a n g , t h e r e s u l t s show t h e

same a c t i v i t y p a t t e r n d u r i n g t h e n i g h t a s r e s u l t s from c a p t u r e

r a t e s do I F i g . 51 ) . At B ak o , w h e re b a t s w ere o n l y s e e n t o l a n d

on two o f t h e s i x f l o w e r s , t h e r e s u l t s v/ere t o o s m a l l t o g i v e

a n y i n d i c a t i o n o f a c t i v i t y p a t t e r n .

8 . R in g i n g results. *

Population structure. At Rantau Panjang, 1063 M. minimus were

captured, examined, ringed and released. 702 (66%) were adults

with a sex ratio of 286 (40.7%) males to 416 (59.3%) females.

The d i f f e r e n c e i s s i g n i f i c a n t (p ^ 0 . 0 0 1 ) . 361 (34 .0% ) w ere

Fig* 53. H is tog ram s showing th e number o f t im es M acrog lossus

minimus landed on S o n n e r a t i a a lb a f lo w e r s a t Rantau

P a n ja n g , S e la n g o r and a t Telok Asara, Bako N a t i o n a l

P a r k , Sarawak.

Mean of: 2 n i g h t s o b s e r v a t i o n one n i g h t ' s o b s e r v a t i o n

toa>

££•HEITs

£<D

u<1)5o

uCDP<(0£E

E

5E|

.Oto4->■H00

t I>

2

1 A

Mean o f 2 n i g h t s o b s e r v a t i o n

________________ A ____________________

m - r mir\ in in in-d- -S' -4" -S'• . • .O' O CM«— C\J OJ CM1 1 1 1O O O Om m m ■ tn• • . •cn o CMc— CM <M CM

in in in in in-3 -S- -S' ~r -S'• *• • • .m o' CM mCM1 01 01 o1 oio o o o om m m tn m• • • • •o r— CM mCM o o O o

ITS

-d"01

oK\

o

i n-S '

•i no

iom

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Time

111

s u b a c l u l t s w i t h a s e x r a t i o o f 189 ( 5 2 .^%) m a le s t o 172 (1)7.6%)

f e m a l e s . T h i s d i f f e r e n c e i s n o t s i g n i f i c a n t (p = < 0 . 5 > 0 . 3 ) »

The r a t i o o f a d u l t s t o s u b a d u l t s was f a i r l y c o n s t a n t t h r o u g h o u t

t h e s t u d y ,

9 . M ovem ents ,

E v id e n c e f o r movem ents o f M. m in im us a t R a n ta u

P a n j a n g comes from two s o u r c e s , i ) r e c a p t u r e o f r i n g e d b a t s and

i i ) a n a l y s i s o f p o l l e n on t h e i r b o d i e s and i n t h e i r f a e c e s .

9 . 1 , E v id e n c e from R e t r a p d a t a .

r e c a p t u r e d one o r more n i g h t ( s ) a f t e r t h e y w ere r i n g e d ( T a b le 15)

g i v i n g 332 r e c a p t u r e r e c o r d s . ( T h i s e x c l u d e s 164 r e c o r d s f o r

t h e s e c o n d o r s u b s e q u e n t t i m e s t h a t a b a t was r e c a p t u r e d d u r i n g

t h e same n i g h t . ) 191 (57»5%) o f t h e s e r e c a p t u r e d b a t s w ere

c a u g h t a t t h e s i t e s a t w h ich t h e y w ere r i n g e d o r h a d p r e v i o u s l y

b e e n r e c a p t u r e d .

r e m a in a t o r r e t u r n r e p e a t e d l y t o t h e same p l a c e s . T h i s a p p l i e d

t o a l l ag e an d s e x c l a s s e s . I t c a n b e f u r t h e r e m p h a s i s e d by

i n d i v i d u a l r e c o r d s , e . g . 1) S u b a d u l t <? R in g No. 2 0 /4 8 7 4 3

R in g e d a t s i t e 7 on 14 M arch 1 9 7 3 .

222 (20 .9% ) o f t h e 1063 r i n g e d M. m in im us w ere

T h e re i s t h u s a s t r o n g t e n d e n c y f o r some b a t s t o

R e c a p t u r e s i t e D a te Time

7

7

7

7 v

14 ' M arch 1972 ' 2 1 . 3 0 h r s

3 A u g u s t 1972 2 1 .0 0 h r s ( n e a r l y m a t u r e )

7 November 1 972 2 1 . 3 0 h r s ( m a tu r e )

21 March 1973 1 9 .3 5 h r s , 2 0 . 3 0 h r s

Table 1 5 .

Table recording the number of Macroglossus minimus captured and the number of nights on which they were recaptured at Rantau Panjang, Selangor. (The data excludes records of the second or subsequent times any bat was recaptured in one night. )

<? ad. ? ad . (f sad. ? sad. Total

Captured once 21 8 338 144 144 844only 7 6 . 2% 81 . 2% 74.9% 8 3 . 2% 79.2%

Recaptured once 49 54 3 k 18 15517.1% 13.0% 1 8 , 1% 1 0 . k% 14.5%

" twice 7 14 10 7 382.4% 3.4% 3 . 0% k .0 % 3.6%

" three 5 7 k 2 18times 1 .7% 1 .7% 2 .0 % 1 .2% 1 .7%

" four 5 3 - 1 9times 1 .7% 0.7% 0 . 6% 0 . 8%

" five 1 - - - 1times 0.3% 0 . 1%

11 six 1 - - — 1times 0.3% 0 . 1%

Total 286 41 6 192 172 1 066

68 78 k S 28 222Total recaptured

2 3 . 8% 1 8 . 7% 2 5 . 0% 1 6 . 3% 2 0 .8 %

112

e . g . 2 ) A d u l t ? R in g No. 2 0 /4 8 8 5 9 *

R in g e d a t s i t e 9 a t 19*45 h r s on 6 A p r i l 1 9 7 2 .

Recapture s i te Date Time

9 6 A p r i l 1972 2 0 .3 0 h r s , 2 2 . 4 5 h r s , 0 6 .1 5 h r s .

9 22 May 1972 19*20 h r s , 2 0 .2 0 h r s , 2 1 . 0 0 h r s ,

21.40 h rs .

9 19 J u l y 1972 19*40 h r s , 2 0 . 1 5 h r s .

9 6 S e p t . 1972 19*25 h r s , 19*45 h r s , 2 0 .3 0 h r s .

9 3 March 1973 19*35 h r s , 2 0 . 3 5 h r s .

14-1 ( 4 2 . 5 %) o f t h e r e c a p t u r e d b a t s w e re r e c a u g h t

a t s i t e s w h e re t h e y h ad n o t p r e v i o u s l y b e e n r e c o r d e d ( T a b l e 1 6) .

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

d i f f e r e n t num bers o f b a t s w ere c a u g h t a t d i f f e r e n t s i t e s , e . g .

t h e r e w e re t h r e e r e c o r d s e a c h o f movements from s i t e s 1 t o 14

and 7 t o 14 a l t h o u g h 150 b a t s w ere c a u g h t a t s i t e 1 and 387

at s i te 7 . Despite th is , the movements are not random. Thus

45*4% o f a l l t h e movem ents w e re b e tw e e n t h e r i v e r and t h e 1

c u l t i v a t e d a r e a a l t h o u g h o n l y 2 4 . 8 % o f t h e b a t s w ere c a u g h t

a l o n g t h e r i v e r ( i . e . a t s i t e s 1 , 2 , 1 0 , 12 and 1 3 ) .

Movem ents w ere r e c o r d e d b e tw e e n t h e r i v e r and

a l l o t h e r s i t e s w i t h i n t h e s t u d y a r e a e x c e p t s i t e 13 w h e re

v e r y few b a t s w e re c a p t u r e d anyway ( T a b le 1 6 ) , e . g . F i g . 54

shows t h e m ovem ents r e c o r d e d b e tw e e n s i t e 1 and a l l o t h e r s i t e s

b u t m o s t m ovem ents from s i t e s w i t h i n t h e c u l t i v a t e d a r e a w ere

e i t h e r to w a r d s o r away from t h e r i v e r ( e . g . F i g . 5 5 ) .

Table 16 .

The t a b l e shows a l l r e c o rd e d movements o f M acroglossus

minimus between n e t s i t e s a t P an ta u P a n jan g , S e l a n g o r . (For

th e l o c a t i o n o f n e t s i t e s , see F i g . 3*)

2 13

3 1

4 6 1

5 4 5 3

6 9 2 2 8

7 1 3 1 2 2

8 4 1 2 2

9 5 5 1 1 1

10 1 1

11

12 2 2 8 3 1 2

13

14 3 9 2 3 9 • 4

15 3 3Natsite 1 2 3 4 5 6 7 8 9 10 11 12 13 14 1 5

I

F i g . 5 ^ . Map o f t h e s t u d y a r e a , R a n ta u P a n ja n g s h o w in g move­

m e n ts o f M a c r o g lo s s u s m in im u s b e tw e e n Net#1 and a l l

o t h e r n e t s i t e s . F i g u r e s r e c o r d t h e num ber o f move­

m e n t s . F i g u r e s i n b r a c k e t s a r e p e r c e n t a g e b a t s from

e a c h n e t s i t e a l s o c a u g h t a t N e t # 1 .

F i g . 55* Map o f t h e s t u d y a r e a a t R a n ta u P a n ja n g sh o w in g

m ovem ents o f M a c r o g lo s s u s m in im us b e tw e e n N e t # 1 4

an d a i l o t h e r n e t s i t e s . F i g u r e s r e c o r d t h e num ber

o f m o v em en ts . F i g u r e s i n b r a c k e t s a r e % b a t s from

e a c h n e t s i t e a l s o c a u g h t a t S i t e # 1 4 .

S i t e 14

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J-XAjs*j. )i"v m a n g ro v e

113

The overall pattern of movement within the study

area was of hats flying along the river hank (and at full raoon

they could clearly be seen flying along the dyke or the water

front) when feeding on Sonneratia spp., or at right angles to

the river when feeding in the cultivated area. Bats roosting

in the cultivated area probably travelled to the river, feeding

en route, and then hack and forth along the river before

returning to the place from which they started. Bats roosting

along the river probably fed along the river, occasionally

moving into.the cultivated area too.

9.ii. Evidence from pollen analysis.No differences were found in the pollen content

of faecal samples collected at different net sites but bats

with poilen on their bodies caught near a flowering tree on

which they fed usually (but not invariably) had pollen of that

tree species on their bodies.

Thus bats were caught along the river with Cocos,

Eugenia and Ceiba as well as Sonneratia spp. pollen in their

faeces (and with white markings from the bracts of banana

inflorescences on their wrists and feet). Others were caught

in the cultivated area with Sonneratia caseolaris and S. alba

pollen in their faeces and frequently on their bodies too.

Three bats that were caught in the cultivated

area more than once in one night had Sonneratia pollen on them

when they were recaught.

m

They w e r e - : ~

1 . & s u b a d u l t ; N e t 9 ; 6 S e p te m b e r 1 9 7 2 . C a u g h t a t : -

1 9 . 1 5 h r s No p o l l e n on b o d y .

2 2 . 0 0 h r s S o n n e r a t i a p o l l e n on w in g .

2 . & a d u l t ; N e t 1 4 ; 10 November 1 9 7 2 . C au g h t a t

1 9 .3 0 h r s No p o l l e n on b o d y ,

2 2 . 2 0 h r s S o n n e r a t i a p o l l e n on w in g .

3 . ? a d u l t ; N e t 4 ; 5 J u l y 1973* C au g h t a t

1 9 .3 0 h r s No p o l l e n on b o d y .

2 0 . 3 5 h r s »

2 2 .0 0 h r s S o n n e r a t i a p o l l e n on b o d y .

T h e se b a t s m u s t , t h e r e f o r e , h a v e t r a v e l l e d from

t h e n e t s i t e ( F i g . ) t o t h e r i v e r , f e d t h e r e f ro m S o n n e r a t i a

f l o w e r s and r e t u r n e d t o t h e same n e t s i t e .

1 0 . R e p r o d u c t i o n .

D a ta on r e p r o d u c t i o n w e re o b t a i n e d a t R a n ta u

P a n ja n g and a t Bako d u r i n g t h e c o u r s e o f o t h e r s t u d i e s .

1 0 . i . R a n ta u P a n j a n g .

The t e s t e s o f a l l a d u l t m a le s w ere c o n t a i n e d i n

a s c r o t u m . M e a s u re d th r o u g h t h e s k i n o f t h e s c r o tu m on l i v e

b a t s t h e y w ere 8 . 0 t o 11 .0 mm i n l e n g t h x 7 . 0 t o 9 . 0 mm i n

w i d t h . D i s s e c t e d o u t from t h e s c r o tu m t h e y m e a s u re d 7 . 0 t o

9 . 0 mm i n l e n g t h x 6 . 0 t o 8 . 0 mm i n w i d t h . T e s t e s w ere d i s s e c t e d

from a d u l t m a l e s i n e v e r y month from F e b r u a r y 1 972 t o J a n u a r y

/

/

115

1973« The e p i d i d y m i s o f a l l s p e c im e n s c o n t a i n e d a w h i t e l i q u i d

p re su m ed t o b e s p e r m a t o z o a . Once s p e r m a t o g e n e s i s was e s t a b l i s h e d

i t t o o k p l a c e c o n t i n o u s l y .

A l l m a le s had a d e r m a l g l a n d on t h e lo w e r t h r o a t

w h ic h , i n M. m in im u s i n P e n i n s u l a r M a l a y s i a , was a lw a y s a b s e n t

f rom a d u l t f e m a l e s . From b i r t h t o f i r s t f l i g h t , i t was u n p ig m e n te d

and i n c o n s p i c u o u s ; i n f r e e f l y i n g s u b a d u l t s i t becam e g r a d u a l l y

more p e r m a n e n t and t u r n e d b r i g h t r e d and a t s e x u a l m a t u r i t y

i t was up t o 2 mm w id e , d i s t i n c t l y r a i s e d i n t o a r i d g e and b r i g h t

r e d t o c h o c o l a t e b row n i n c o l o u r . I t p ro d u c e d no d i s c h a r g e o r

d e t e c t a b l e s c e n t . I t s f u n c t i o n i s n o t known.

The r e p r o d u c t i v e s t a t u s o f 473 a d u l t f e m a le s

c a u g h t i n 1972 i s su m m a r i s e d i n F i g . 5 6 . The f i g u r e i n c l u d e s

a l l r e t r a p r e c o r d s e x c e p t w h e re a f e m a le was c a u g h t t w i c e i n one

c a l e n d e r m o n th , w h e re o n l y t h e f i r s t r e c o r d h a s b e e n i n c l u d e d .

P r o f i c i e n c y i n d e t e c t i n g t h e e a r l y s t a g e s o f p r e g n a n c y b y p a l p a t i o n

im p ro v e d v / i th e x p e r i e n c e , c o n s e q u e n t l y t h e r e c o r d s from F e b r u a r y

t o A p r i l 1 972 a t t h e b e g i n n i n g o f t h e s t u d y show a b n o r m a l l y low

p r e g n a n c y and h i g h " n o t p r e g n a n t " r a t e s b e c a u s e s e v e r a l e a r l y

p r e g n a n c i e s w e n t u n d e t e c t e d . V ery few d a t a w e re o b t a i n e d a f t e r

O c to b e r 1 9 7 2 .

N e v e r t h e l e s s , F i g . 56 shows t h a t p r e g n a n c y

o c c u r r e d i n e v e r y m onth f o r w h ich d a t a a r e a v a i l a b l e ( i . e . i n

a l l m on th s e x c e p t J a n u a r y : Some b a t s w h ich w e re n e a r i n g f u l l

t e r m p r e g n a n c y i n F e b r u a r y m u s t h a v e b e e n p r e g n a n t i n J a n u a r y . )

The a p p a r e n t p e a k f rom May t o S e p te m b e r i s ari a r t i f a c t b e c a u s e

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

FIG. 55 .Histograms showing the reproductive status of 473

adult female Macroglossus minimus caught at Rantau Panjang, Selangor.N = 37 80 43 38 60 48 97 41 13 8 8

100

tr>C

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rorH

60

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2 0 -

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80-

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Months

116

sa m p le s i z e a f t e r S e p te m b e r . The d a t a f o r t h o s e m on ths p r o b a b l y

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

more i n f o r m a t i o n w ou ld b e r e q u i r e d t o v e r i f y t h a t . The d a t a on

l a c t a t i o n s u p p o r t s t h i s v i e w .

L a c t a t i o n was more e a s i l y and a c c u r a t e l y d e t e c t a b l e

th a n p r e g n a n c y a n u i t w a s o f c o n s i d e r a b l y s h o r t e r d u r a t i o n th a n

g e s t a t i o n (w h ic h l a s t e d o v e r t h r e e m o n t h s ) . T h e r e f o r e , m o n th ly

r e c o r d s o f t h e p e r c e n t a g e o f a d u l t f e m a le s i n l a c t a t i o n may b e

e x p e c t e d t o p o r t r a y more c l e a r l y a n y s e a s o n a l v a r i a t i o n i n t h e

r e p r o d u c t i v e r a t e t h a n p r e g n a n c y r e c o r d s d o . From F e b r u a r y t o

O c to b e r 1972 t h e p r o p o r t i o n o f a d u l t f e m a le s i n l a c t a t i o n r e m a in e d

f a i r l y c o n s t a n t , i . e . b e tw e e n 4 3 .2% i n F e b r u a r y and 20 .8% i n

J u l y . The d i f f e r e n c e i s n o t s i g n i f i c a n t (p < 0 . 1 > 0 . 0 5 ) . I t

d o u b le d i n November and D ecem ber b u t t h e s a m p le s i z e was v e r y

s m a l l i n t h e s e m onths and so t h e f i g u r e s a r e n o t m e a n i n g f u l .

The r e s u l t s t o d a t e a r e c o n s i s t e n t w i t h t h e h y p o t h e s i s t h a t

t h e r e i s l i t t l e s e a s o n a l v a r i a t i o n i n t h e r e p r o d u c t i v e r a t e .

1 0 . i i . G e s t a t i o n .

A minimum f i g u r e f o r t h e d u r a t i o n o f g e s t a t i o n

was o b t a i n e d from r e t r a p d a t a o f b a t s c a u g h t i n e a r l y p r e g n a n c y

an d a g a i n j u s t p r i o r t o p a r t u r i t i o n . T a b le 17 shows t h e s i x

l o n g e s t i n t e r v a l s r e c o r d e d . ( P r e g n a n c y was s u b d i v i d e d i n t o f o u r

p h a s e s ; e a r l y p r e g n a n c y , 1 / 3 p r e g n a n t , 2 / 3 p r e g n a n t and 3 / 3

p r e g n a n t , a c c o r d i n g t o t h e s i z e and d e v e lo p m e n t o f t h e e m bryo .

(The c a t e g o r i e s a r e d e f i n e d on p . 2 2 . )

The l o n g e s t i n t e r v a l r e c o r d e d was 1 08 d a y s . The

b a t c o n c e r n e d h a d a v e r y s m a l l u t e r i n e s w e l l i n g , a b o u t 2 mm i n

Table 17.

R e c a p tu re d a t a o f 6 M acroglossus minimus cau g h t a t

Rantau P an jan g which i n d i c a t e th e l e n g t h o f g e s t a t i o n .

No Date o f c a p t u r e

Date o f r e c a p t u r e S t a t u s change

No. o f days

1 22 May 72 6 S e p t . 72 L, e a r l y p r to 3 / 3 p r , ME 108

2 6 A p r . 72 16 J u l y 72 L, e a r l y p r t o 3 / 3 p r , ME 101

3 5 J u l y 72 10 O c t . 72 e a r l y p r to 3 / 3 p r 97

4 3 Aug. 72 7 Nov. 72 e a r l y p r to 3 / 3 p r 96

5 9 May 72 9 Aug. 72 e a r l y p r to 3 / 3 p r 92

6 10 Nov 72 28 June 73 1 / 3 p r to 3 /3 p r , ME 79

N.B. L = l a c t a t i n g ; p r = p r e g n a n t ; ME = mammary g la n d s e n l a r g e d

Table 1 8 .

E s t im a te d d u r a t i o n o f th e fou r phases o f p regnancy i n

M acroglossus minimus c a p tu r e d a t Rantau P a n ja n g , S e l a n g o r ,

assuming th e l e n g t h o f g e s t a t i o n i s 120 d a y s .

Phase No. of bats % estimated no. of days

early pregnancy 97 3 5 .8 43

1 /3 pregnant 65 24 .0 29

2 /3 pregnant 53 1 9 .6 23

3 /3 pregnant 56 2 0 .7 25

Total 271 100.1 120

117

d i a m e t e r , o n l y j u s t d e t e c t a b l e b y p a l p a t i o n on 22 May 1 9 7 2 .

I t was r e c a p t u r e d on 6 S e p te m b e r 1972 j u s t p r i o r t o p a r t u r i t i o n

when i t s mammary g l a n d s w ere e n l a r g e d and c o l o s t r u m was e x t r a c t e d

f ro m t h e n i p p l e s b y g e n t l e p r e s s u r e . G e s t a t i o n m u s t h a v e l a s t e d

a few d a y s l o n g e r , i . e . p r o b a b l y a b o u t 120 d a y s ( s e e a l s o b e l o w ) .

I f t h e h y p o t h e s i s t h a t b r e e d i n g i s c o n t i n o u s i s

c o r r e c t , t h e n t h e p e r c e n t a g e o f p r e g n a n t f e m a le s i n e a c h p h a s e

o f _ p r e g n a n c y w i l l b e e q u a l t o t h e p e r c e n t a g e o f t h e g e s t a t i o n

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

a c t u a l g e s t a t i o n t im e i s known t h e a c t u a l d u r a t i o n f o r e a c h

p h a s e o f p r e g n a n c y c an b e c a l c u l a t e d ( T a b l e 18 ) .

M O . i i i . L a c t a t i o n .

D a ta i n d i c a t i n g t h e d u r a t i o n o f l a c t a t i o n was

' o b t a i n e d fro m r e c a p t u r e d b a t s ( T a b l e 1 9 ) . The l o n g e s t p e r i o d s

o f l a c t a t i o n r e c o r d e d w e re 75 d a y s and 69 d a y s . The f o r m e r

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

g i v e n b i r t h - when f i r s t c a u g h t . She was a g a i n i n e a r l y p r e g n a n c y

and h a d n e a r l y c e a s e d l a c t a t i n g when r e c a u g h t . The l a t t e r r e c o r d

Was o f a f e m a le v e r y n e a r t o p a r t u r i t i o n when f i r s t c a u g h t

( c o l o s t r u m was m a n u a l ly e x t r a c t e d from t h e n i p p l e s ) . She was

1 / 3 p r e g n a n t a nd h a d a l m o s t c e a s e d l a c t a t i n g when r e c a u g h t .

L a c t a t i o n and e a r l y p r e g n a n c y f r e q u e n t l y o v e r l a p p e d

( F i g . 5 6 ) an(i one f e m a le was r e c o r d e d a s 1 / 3 p r e g n a n t and

l a c t a t i n g . The o v e r l a p o f p r e g n a n c y and l a c t a t i o n was c o n f i r m e d

i n s e v e r a l i n s t a n c e s b y d i s s e c t i o n . T h i s i s i n d i c a t i v e o f a

p o s t - p a r t u m e s t r o u s b u t more w ork i s n e e d e d t o v e r i f y t h i s .

Table 19*

R ec ap tu re d a t a o f 5 M acroglossus minimus cau g h t a t

Rantau P an jan g which i n d i c a t e th e l e n g t h o f j t f J S * .

No.Date o f c s p t u r s

Date o fr* Q n f iit*p- —-~.r ~ ~

S t a t u s change No. o frl n y o

1 27 Mar. 72 10 June 72 L(p r? ) to L, e a r l y p r 75

2 30 Aug. 72 8 Dec. 72 3 /3 ME to 1 /3 p r , L 69

3 10 O c t . 72 1 Dec. 72 L, n o t p r to L, e a r l y p r 52

4 6 Apr. 72 22 May 72 L( c yng) to L, e a r l y p r 46

5 29 S e p t . 72 10 Nov. 72 L, n o t p r to L, e a r l y p r 42

N.B. L = l a c t a t i n g ; p r = p r e g n a n t ; c yng = c a r r y i n g young.

>

Table 20 .

R ec a p tu re d a t a o f 6 M acrog lossus minimus cau g h t a t

Rantau P an jan g which i n d i c a t e th e i n t e r v a l be tween s u c c e s s i v e

p r e g n a n c i e s .

No.Date o f c a p t u r e

Date of r e c a p t u r e S t a t u s change

No. o fdays

1 1 May 72 10 O ct . 72 L, n o t p r to L, n o t p r 163

2 6 A pr . 72 6 S e p t . 72 c yng to 3 /3 p r , ME 153

3 14 Mar. 72 12 Aug. 72 L (p r? ) to L, e a r l y p r 151

4 30 Aug. 72 28 J a n . 73 3 /3 p r , ME to 3 / 3 p r , ME 1 50

5 14 Mar. 72 4 Aug. 72 L (p r? ) to L, e a r l y p r 143

6 14 Mar. 72 3 Aug. 72 L, n o t p r to e a r l y p r 142

N.B. L = l a c t a t i n g ; p r = p r e g n a n t ; c yng = c a r r y i n g young; HE = Mammary g la n d s e n l a r g e d .

118

1 0 . i v . R e p r o d u c t i v e r a t e . '

R e c a p t u r e o f f e m a le s i n t h e same p h a s e o f

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

o f s u c c e s s i v e y o u n g . T a b le 20 p r e s e n t s d a t a sh o w in g t h e

a p p r o x i m a t e i n t e r v a l b e tw e e n p r e g n a n c i e s f o r s i x b a t s . The

t im e v a r i e d from a l i t t l e o v e r 140 d a y s t o a b o u t 160 d a y s .

The i n t e r v a l s a l s o e s t a b l i s h a maximum p o s s i b l e g e s t a t i o n t i m e ,

1 0 .V . The r e p r o d u c t i v e c y c l e .

M a c r o g lo s s u s m in im us a t R a n ta u P a n j a n g shows a

s e a s o n a l p o l y e s t r y ( a s d e f i n e d b y W i l s o n , 1 9 7 3 ) . The n o rm a l

c y c l e e n t a i l s t h e b i r t h o f one young a b o u t e v e r y f i v e m o n th s .

G e s t a t i o n l a s t s a b o u t 120 d a y s (minimum 108 d a y s ; maximum 140

d a y s ) f o l l o w e d b y l a c t a t i o n w h ic h l a s t s 60 t o 70 ' d a y s . The

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

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

p a r tu m e s t r o u s ,

l O . v i . C a r r y i n g o f y o u n g .

F e m a le s do n o t n o r m a l l y c a r r y t h e i r young w i t h

them when f o r a g i n g . Of 198 l a c t a t i n g b a t s c a u g h t a t R a n ta u

P a n ja n g and K u a la S e l a n g o r 5 (2 .5% ) c a r r i e d y o u n g . No f e m a le s

w e re c a u g h t e l s e w h e r e c a r r y i n g y o u n g . The f i v e j u v e n i l e s t h a t

w e re fo u n d a t t a c h e d t o l a c t a t i n g f e m a le s w e re a l l v e r y y o u ng :

T h re e w e re n e o n a t e s ( w e i g h t 4 g> n = 3 ; f o r e a r m l e n g t h 20 mm,

n = 2 ) . One w e ig h e d 5 g w i t h a f o r e a r m l e n g t h o f 24 mm and

t h e l a s t , k n o w n t o b e 10 d a y s o l d , w e i g h e d 5 g w i t h a f o r e a r m

l e n g t h o f 2 7 mm.

119

1 0 . v i i . S p e c i f i c f o e t a l g ro w th v e l o c i t y .

H u g g e t t and. W iddas (1 9 5 0 ) d e v i s e d a f o r m u la

w h e re b y an e s t i m a t e o f t h e s p e c i f i c f o e t a l g ro w th o f mammals

c a n b e c a l c u l a t e d from t h e g e s t a t i o n p e r i o d i n d a y s and t h e

b i r t h w e i g h t b y : -

a • V 7 "

(* - y

w h ere a = s p e c i f i c f o e t a l g ro w th v e l o c i t y ,

W = b i r t h w e i g h t ,

t = g e s t a t i o n , an d

' t = i n t e r c e p t o f b o s s - r u m p l e n g t h on t h e t i m e a x i s .

No d a t a was o b t a i n e d by w h ic h t c a n b e c a l c u l a t e d and an

e s t i m a t e o f 0 . 2 x g e s t a t i o n t im e s u g g e s t e d by H u g g e t t and

W iddas ( o p p . c i t . ) f o r mammals w i t h a g e s t a t i o n o f 100 - A-00

d a y s h a s b e e n e m p lo y e d . U s in g t h i s e s t i m a t e , ja = 0 . 0 1 7 .

l O . v i i i . P o s t n a t a l g ro w th r a t e .

One a d u l t f e m a le was c a u g h t t w i c e i n t e n d a y s

c a r r y i n g an i n f a n t . When f i r s t c a u g h t on 10 M arch 1 9 7 2 , t h e

y oung was a n e o n a t e s t i l l w e t w i t h a m n i o t i c f l u i d . I t s f o r e a r m

l e n g t h was 20 mm a nd w e i g h t ^ g . I t s e y e s w e re c l o s e d . By 20

M arch t h e f o r e a r m h ad i n c r e a s e d t o 27 mm and t h e w e i g h t t o 5 g .

The e y e s w e re o p e n . R e t r a p d a t a from young c a p a b l e o f f l i g h t

i n d i c a t e t h a t t h e i r i n c r e a s e i n w e i g h t was a p p r o x i m a t e l y one

g ra m /m o n th

120

R e l a t i v e g ro w th r a t e s o f t h e f o r e a r m and w e i g h t r , .

I n F i g . 5 7 t h e f o r e a r m l e n g t h h a s b e e n p l o t t e d

a g a i n s t w e i g h t f o r s u b a d u l t b a t s . S in c e no f r e e f l y i n g young

w i t h f o r e a r m s l e s s t h a n 3 6 . 5 mm w ere c a u g h t i t h a s b e e n a ssum ed

t h a t f i r s t f l i g h t o c c u r r e d a t a b o u t t h a t s i z e .

F i g , 57 shows t h a t p r i o r t o f i r s t f l i g h t t h e r e '

v/as a v e r y r a p i d g ro w th r a t e o f t h e f o r e a r m com p ared t o t h e

• w e i g h t . H ow ev er , a f t e r f i r s t f l i g h t , t h e r e l a t i v e g ro w th r a t e s

c h a n g e s o t h a t b o t h r e a c h a d u l t d im e n s i o n s a t a b o u t t h e same

t i m e .

A b s o l u t e g r o w th r a t e s .

S i n c e l a c t a t i o n l a s t s a b o u t 70 d a y s an d s i n c e

t h e y oung a r e c a p a b l e o f f e e d i n g on t h e i r own p r i o r t o w e a n in g ,

n e o n a t e s m u s t a t t a i n f r e e f l i g h t i n l e s s t h a n 7 0 d a y s , i . e . t h e i r

minimum mean g ro w th r a t e i s a b o u t 0 . 0 5 7 g p e r d a y . A f t e r f r e e

f l i g h t , t h e g ro w th r a t e d r o p s t o a b o u t 0 . 0 3 3 g p e r d ay ( i . e . -

1 g p e r m o n t h ) .

They t h u s r e a c h a d u l t w e i g h t b e tw e e n s i x and

e i g h t m o n th s f rom b i r t h ' .

1 0 . i x . Bako N a t i o n a l P a r k .

T a b le 21 show s t h e r e p r o d u c t i v e s t a t u s o f a d u l t

f e m a le M. m in im us c a u g h t a t T e lo k Asara, Bako N a t i o n a l P a r k .

I n J a n u a r y 1973 8 (50%) o f t h e 16 a d u l t f e m a l e s t h a t w e re c a u g h t

w e re 1 / 3 p r e g n a n t . ( A t R a n ta u P a n j a n g t h i s p h a s e w as c a l c u l a t e d

t o l a s t j u s t u n d e r 2 0 % o f e a c h r e p r o d u c t i v e c y c l e a n d s o , i f

b r e e d i n g w as a l s o c o n t i n o u s a t T e lo k Asam, t h e e x p e c t e d num ber

o f f e m a le s i n t h i s p h a s e o f p r e g n a n c y w o u ld b e 2 0 % o f 1 6 , i . e . 3 »

F i g . 5 7 . The f o r e a r m s o f young M a c r o g lo s s u s m in im us p l o t t e d

a g a i n s t t h e i r w e i g h t . S u b a d u l t s ( c a p a b l e o f f r e e

f l i g h t ) an d j u v e n i l e s ( i n c a p a b l e o f f l i g h t ) a r e

t r e a t e d s e p a r a t e l y and a r e g r e s s i o n l i n e h a s b e e n

f i t t e d t o t h e d a t a f o r • s u b a d u l t s . D a ta f o r one

j u v e n i l e c a u g h t t w i c e i n 10 d a y s a r e j o i n e d b y a

d o t t e d l i n e . A l l d a t a from R a n ta u P a n j a n g , S e l a n g o r .

FOR

EA

RM

[M

Mj

45

40

A

A AA A A

A A A

A A

9

35

30

25

20i

A x 2

A SUBADULT, CAPABLE OF FLIGHT.

A J U V E N I L E , N O T ” ” »

O ADULT, M E A N .

158 12

W E I G H T ( G R A A \ S j16 20

Table 21 .

Table showing th e r e p r o d u c t i v e s t a t u s o f a d u l t female

M acrog lossus minimus caugh t i n Bako N a t i o n a l P a rk , Sarawak.

T o t a l L n o fixu v j y

T V . Y»J b '1

a o 1 v« vi i / ~ Z ~ ~ 1 / -) / 7 ---------<-/ .vr-*- •7 /7 . —

J a n u a ry

197316 1 1 8 2 h

May1973

7 2 1 b

N.B. L = l a c t a t i n g ; p r = p r e g n a n t .

Table 22 .

Tab le showing th e r a t i o of a d u l t to s u b a d u l t

M acrog lossus minimus c augh t i n Bako N a t i o n a l P a r k , Sarawak.

T o t a l Adul ts S u b a d u l t s

J a n u a ry1973

22 20(90.9%) 2( 9.1%)

May

197332 H (13 .75% ) 18(56.25%)

121

The o b s e r v e d num ber d i f f e r s s i g n i f i c a n t l y from t h e e x p e c t e d ’

(p = 0 . 0 1 ) . I n May 1973 o n l y s e v e n a d u l t f e m a l e s v/ere c a u g h t .

H ow ever , k (57%) w e re 3 / 3 p r e g n a n t and two w ere h e a v i l y l a c t a t i n g

sh o w in g t h a t t h e y ha d r e c e n t l y g i v e n b i r t h .

A l th o u g h t h e s a m p le s i z e s a r e s m a l l t h e y s t r o n g l y

i n d i c a t e t h a t t h e r e Was s y n c h r o n y i n t h e b r e e d i n g c y c l e s o f

some members o f t h e p o p u l a t i o n i n b o t h m o n th s . S i n c e t h e r e p r o d u c t i v e

c y c l e l a s t s f i v e m o n ths and t h e v i s i t s t o Bako w e re f o u r m on ths

a p a r t , t h e g r o u p o f 3 / 3 p r e g n a n t f e m a le s r e c o r d e d i n May c o u l d

n o t h a v e b e e n p a r t o f t h e 1 / 3 p r e g n a n t f e m a le s r e c o r d e d i n

J a n u a r y .

The r a t i o s o f s u b a d u l t s t o a d u l t s i n e a c h m onth

( T a b le 22 ) s u p p o r t t h e h y p o t h e s i s t h a t p r e g n a n c y w as s y n c h r o n o u s

among some f e m a l e s i n J a n u a r y 1 9 7 3 . I n J a n u a r y 22 b a t s w e re

c a u g h t b u t o n l y 2 (9 .1% ) w e re s u b a d u l t s . I n May 32 b a t s w e re

c a u g h t a n d 18 (5 6 .2% ) w e re s u b ' a d u l t s . I n b o t h m o n th s t h e p r o p o r t i o n

o f s u b a d u l t s i n t h e p o p u l a t i o n d i f f e r e d from t h e mean p r o p o r t i o n

£3A%) r e c o r d e d a t R a n ta u P a n j a n g w h e re b r e e d i n g w as c o n t i n o u s ,

(p = < 0 . 0 5 ^ 0 . 0 2 ) . They a l s o d i f f e r e d s i g n i f i c a n t l y from

e a c h o t h e r (p = < 0 . 0 1 > 0 . 0 0 1 ) .

15 (83%) o f t h e 18 s u b a d u l t s c a u g h t i n May w e ig h e d

b e tw e e n 11 . 5 g an d 1 2 . 5 g (mean 1 1 . 7 g ) , i . e . a b o u t k g l e s s

t h a n t h e mean w e i g h t o f a d u l t m a le s c a u g h t a t T e lo k Asam. T hese

s u b a d u l t s w e re p r o b a b l y a l l o f s i m i l a r a g e .

U s in g t h e e s t i m a t e s o b t a i n e d a t R a n ta u P a n j a n g

f o r t h e d u r a t i o n o f e a c h p h a s e o f p r e g n a n c y and f o r t h e g ro w th

r a t e s o f y o u n g , t h e young o f f e m a le s t h a t w e re 1 / 3 p r e g n a n t

i n J a n u a r y w o u ld b e e x p e c t e d t o w e ig h a b o u t 11 g b y May. T h i s

122

i s v e r y c l o s e t o t h e mean w e i g h t o f t h e l a r g e g r o u p o f young

r e c o r d e d a t T e lo k Asara i n May and s u p p o r t s t h e h y p o t h e s i s t h a t

t h e y w e re t h e o f f s p r i n g o f a g r o u p o f f e m a le s t h a t w ou ld h a v e

b e e n 1 / 3 p r e g n a n t i n J a n u a r y an d t h e r e f o r e t h a t b r e e d i n g was

s y n c h r o n i s e d among some members o f t h e p o p u l a t i o n a t T e lo k Asam

i n J a n u a r y 1973*

More w o rk i s n e e d e d t o v e r i f y s y n c h r o n o u s b r e e d i n g

a t T e lo k Asam b u t t h i s w o u ld b e a v e r y i n t e r e s t i n g t o p i c f o r

i n v e s t i g a t i o n .

123

MACROGLOSSUS SOBRINUS.

1 . E c o l o g i c a l d i s t r i b u t i o n .

M a c r o g l o s s u s s o b r i n u s was c a u g h t o v e r a w ide

a l t i t u d i n a l r a n g e , from 10 m a . s . l . on t h e ed g e o f t h e T e lo k

F o r e s t R e s e r v e , S e l a n g o r t o 1800 m a . s . l . on Gunong B r i n c h a n g ,

P a h a n g . H ow ev er , i t was n e v e r t a k e n f a r from Musa t r u n c a t a o r

M. m a l a c c e n s i s , two common s p e c i e s o f w i l d b a n a n a s . Thus i t

was n e v e r t a k e n i n p r i m a r y f o r e s t n o r i n m a n g ro v e s a l t h o u g h

i t was t a k e n on t h e e d g e s o f t h e Meru F o r e s t R e s e r v e an d t h e

T e lo k F o r e s t R e s e r v e w h ic h a r e 11 km an d 3 km r e s p e c t i v e l y

from m angrove swamps. (B o th c a p t u r e s i t e s w e re s e c o n d a r y f o r e s t s . )

The l o c a l i t i e s a t w h ich i t was c a u g h t a r e l i s t e d

i n T a b le 23 . T h e se and o t h e r l o c a l i t i e s i n M a l a y s i a from w h ic h

s p e c i m e n s h a v e b e e n ex am in ed a r e l o c a t e d on F i g . 40 an d l i s t e d

i n S a b le 24 t o g e t h e r w i t h t h e c o l l e c t i o n s i n w h ic h t h e s p e c im e n s

a r e d e p o s i t e d . Lira (1 9 6 6 ) r e c o r d e d t h i s s p e c i e s ( a s M. m in im us ,

s e n s u A n d e r s e n , 1912.) from t h e m angrove f o r e s t s b u t r e e x a m i n a t i o n

o f t h e s p e c im e n s (now d e p o s i t e d i n t h e IMR c o l l e c t i o n ) on w h ich

he b a s e d h i s r e c o r d s showed them t o be m i s i d e n t i f i e d s p e c im e n s

o f M. m i n im u s .

2 . R o o s t i n g .

No w i l d s o b r i n u s w ere fo un d r o o s t i n g e i t h e r

by day o r by n i g h t . A n o t e on t h e l a b e l o f o n e , No. 84749 i n

t h e IMR c o l l e c t i o n s t a t e s " c a u g h t i n l i v e bamboo t r e e 15 f t

from kampong b y h a n d " . ("Kampong" = Kg. B a tu E n a m b la s , i . e .

v i l l a g e a t t h e 1 6 ^ m i l e p o s t on t h e T a p a h - R i n g l e t r o a d . )

Table 2 3 .

The table lists the localities in Peninsular Malaysia

at which Macroglossus sobrinus was caught and records the

number of pollen samples collected at each locality.

LocalityNo. No. Total Pollen from

Ringed Killed caught Gut and faeces Fur

Kajang 17 3 20 13 7

F.S.C. 21 1 22 10 5

Telok F.R. - 2 2 2 -

Meru F.R. - 3 3 3 1

li+^mile, Tape'n - 5 5 5 2to Ringlet rd.

17th mile, " 7 7 7 3

G. Erinchang - 6 6 6 1

Total 00 27 65 46 19

Note F.S.C. = Field,Studies Centre, Ulu Gorabak.

F.R. = Forest Reserve.

G. = Gunong = Mountain.

Ir

I

Table 2 l { .

The t a b l e r e c o r d s th e l o c a l i t i e s i n P e n i n s u l a r

M alays ia from which specimens o f M acroglossus s o b r in u s were

examined ( e x c l u d i n g th o se l i s t e d i n Table 2 3 ) .

L o c a l i t y S t a t e C o o rd in a te s C o l l e c t i o n

Segamat Lab is F .R . J ohore approx 3°30 'N 103°00E IMR

G. Ledang ti 3°28 'N 102°39 'E IMR

G. J e r a i Kedah 5°*tS'N 100°26 'E IMR

F o r t Erook K e la n tan ? UMC

Kota Bahru ii 6°05 'N 102°19 'E IMR

Eentong Pahang 3°3^ 'N 101°5 1 'E IMR

G. Eenom it 3 ° 5 T N 102°05 'E IMR

F r a s e r ' s H i l l ii 3°Ml'N 1 01 °^f21E UMC

Gent ing Sempah ii 3°22 'N 101° 5 0 ' E UMC

G. Gajah P erak Z*°25'N 101 °1 1 'E IMR

Maxwell H i l l it i f °5 l 'N 100°48 'E SNC, UMC

E u k i t Lanjang S e la n g o r 3 U12'N 1 01 °3 9 'E IMR

E u k i t Legong it 3°14 'N 101°if1 'E IMR

Kuala Lumpur ii 3°09 'N 101° 4 8 'E IMR

Ulu Langat ii 3 O06'N 101°52 'E IMR

Note G. = Gunong = M ounta in .F .R . = F o r e s t R e s e rv e .IMR = I n s t i t u t e o f M edica l R e se a rc h , Kuala Lumpur. SNC = S in g a p o re N a t i o n a l C o l l e c t i o n .UMC = U n i v e r s i t y o f Malaya C o l l e c t i o n .

//

122|

C a r e f u l s e a r c h e s f o r r o o s t i n g b a t s w ere made ( p a r t i c u l a r l y

a t K a ja n g ) a m o n g s t t h e b u n c h e s o f d e ad l e a v e s t h a t o f t e n

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

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

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

new l e a v e s (w h e re M y o t l s m y s t a c i n u s was commonly fo u n d r o o s t i n g )

and among t h e h a n d s o f f r u i t on f r u i t i n g b a n a n a s .

I n c a p t i v i t y t h e y s u s p e n d e d t h e m s e l v e s from t h e

m i d r i b s o r f r o n d s o f a s m a l l p a lm t r e e p l a c e d i n t h e i r c a g e

o r from t h e g a u z e n e t t i n g on t h e r o o f o f t h e i r c a g e . T h e i r

mode o f g r a s p i n g t h e s u p p o r t was e x a c t l y a s d e s c r i b e d f o r

M. m in im u s t i . e . w i t h one f o o t p l a c e d e a c h s i d e o f t h e s u p p o r t

(S e e F i g . 2 f o r M. m in im u s ) a n d w i t h t h e b o d y e n v e lo p e d by

t h e w i n g s . The h e a d was a l s o c o v e r e d b y t h e w in g s u n l e s s t h e

b a t was aw akened an d a l e r t . They s u s p e n d e d t h e m s e l v e s by t h e i r

thum bs t o u r i n a t e an d d e f a e c a t e .

3 . Food s o u r c e s o f M a c r o g lo s s u s s o b r i n u s . -

A l th o u g h o n ly a p r e l i m i n a r y s t u d y o f M a c r o g lo s s u s

s o b r i n u s was u n d e r t a k e n , i t was fo u n d t h a t a v e r y c l o s e r e l a t i o n ­

s h i p e x i s t e d b e tw e e n i t an d two s p e c i e s o f w i l d b a n a n a s (Musa

m a l a c c e n s l s a n d M. t r u n c a t a , i n c l u d i n g t h e i r h y b r i d p r o g e n y )

on w h ich t h e y f e d . U nder c e r t a i n c o n d i t i o n s t h e y a l s o f e e d on

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

f e e d i n g from Musa v l o l a s c e n s

Musa m a l a c c e n s i s , M. t r u n c a t a an d t h e i r n a t u r a l h y b r i d s

Musa t r u n c a t a r e p l a c e s M. m a l a c c e n s i s a t h i g h

/

j

125

a l t i t u d e s b u t t h e r e i s . a b r o a d r e g i o n o f o v e r l a p i n w h ich

n a t u r a l h y b r i d s b e tw e e n them a r e common (C h en , 1 9 7 2 ; Simmonds,

19 55 ; p e r s . o b s . ) . On m o o n l i g h t n i g h t s M a c r o g l o s s u s s o b r i n u s

w as s e e n l a n d i n g on t h e i n f l o r e s c e n c e s o f b o t h t h e s e s p e c i e s

o f Musa an d t h e i r h y b r i d s a t a num ber o f l o c a l i t i e s . R e g u l a r

o b s e r v a t i o n s w e re made a t f i v e o f them ( T a b le 2 5 ) .

I n t h e b o t a n i c a l l y s t e r i l e c o n d i t i o n s p r e v a i l i n g

i n ' t h e r u b b e r e s t a t e i n K a ja n g , Musa m a l a c c e n s i s was t h e o n ly

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

e x c e p t f o r f o u r f l o w e r i n g c u l t i v a t e d b a n a n a p l a n t s , an d a t

h i g h e l e v a t i o n s on Gunong B r i n c h a n g ( w h e r e M. s o b r i n u s was

o b s e r v e d l a n d i n g on t h e i n f l o r e s c e n c e s o f M. t r u n c a t a ) no o t h e r

known c h i r o p t e r o p h i l o u s p l a n t s w ere f o u n d . T h e r e f o r e , a t K a ja n g

an d p r o b a b l y a t h i g h a l t i t u d e s on Gunong B r i n c h a n g , M. s o b r i n u s

was l a r g e l y d e p e n d e n t on Musa s p p . f o r (food.

3 . i i . C u l t i v a t e d b a n a n a s .

C u l t i v a t e d b a n a n a s a r e p a r t vh e n o c a r p i c d e r i v a t i v e s

0:f ^ u s a a c u m i n a t a ( s e n s u l a t o ) o r h y b r i d s b e tw e e n t h a t s p e c i e s

an d Musa b a l b i s i a n a (S im m onds, 1 9 6 2 ) . T h e i r a n t h e r s r e l e a s e

no p o l l e n b u t t h e y do p r o d u c e up t o 2.1+ ml o f n e c t a r p e r

i n f l o r e s c e n c e p e r n i g h t f o r o v e r s i x m o n th s .

Where c u l t i v a t e d b a n a n a s w e re grow n n e a r Musa

m a l a c c e n s i s o r M. t r u n c a t a ( o r t h e i r h y b r i d s ) ( e . g . on t h e edge

o f t h e Meru F o r e s t R e s e r v e , a t t h e F . S . C . U lu Gombak an d a t

K a j a n g ) , M a c r o g lo s s u s s o b r i n u s was s e e n t o l a n d on them t o f e e d .

H ow ever , w h e re no w i l d b a n a n a s grew n e a r b y , M, s o b r i n u s was n o t

fo u n d among c u l t i v a t e d b a n a n a s . E . g . f o u r m i s t n e t s w e re s e t

The t a b l e show s t h e num ber o f v i s i t s t o c lu m ps o f

f l o w e r i n g b a n a n a s p e r u n i t t im e b y M a c r o g lo s s u s s o b r i n u s .

N o te 1

N o te 2

N o te 3

T-R r d . = T a p a h - R i n g l e t r o a d ,

F . S . C . = F i e l d S t u d i e s C e n t r e , U lu Gombak,

G. = Gunong = M o u n t a in ,

i n f l o r . = i n f l o r e s c e n c e .

A v i s i t r e f e r s t o a n y o c c a s i o n on w h ic h a b a t

l a n d e d on an y o f t h e i n f l o r e s c e n c e s i n t h e c lum p

b e i n g w a tc h e d i r r e s p e c t i v e o f how many i t l a n d e d

on o r how many t i m e s i t l a n d e d on them s o l o n g a s

i t r e m a in e d i n t h e i r im m e d ia te v i c i n i t y .

O b s e r v a t i o n s w e re c o n t i n o u s f o r p a r t o f e a c h h o u r .

’•Each" r e f e r s t o t h e d u r a t i o n i n m i n u t e s o f e a c h

p e r i o d o f o b s e r v a t i o n and ’• T o t a l ’' i s t h e t o t a l

t im e i n h o u r s t h a t t h e i n f l o r e s c e n c e s w e re w a t c h e d .

No. of Observations No. ofLocality Musa spp. inflor. Time Each Total visits

G. Erinchang M. truncata 2 1 9 . 25-21 .1 0 45 /60 I i 211 2 2 2 . 3 0 - 0 0 .1 5 11 1 i 1

I7thmile, T-R rd.

M. truncata 2 2 0 . 0 0 - 0 0 .3 0 3 0 /6 0 2 i 3

1T-R rd.

M. malaccensis x M. truncata

3 19 .3 0 -2 2 .0 0 3 0 /6 0 i i 1i

ii 3 01+. 00 -05 .30 25/30 i i 0

F.S.C. Cultivated 4 1 9.30-23.1+5 15/30 n 3ii 3 11 it 2± 3

Kajang M. malaccensis 2 1 9 .3 0 -2 3 .4 5 1 5 /6 0 i £ 311 2 ti ti 1* 2n 2 11 11 1* 1ii 2 n n 1* 2it 2 11 11 1 i 2

Total 19 23

I4 tnmile, T-R rd.

M. violascens 1 1 9 .3 0 -2 2 .0 0 3 0 / 6 0 1 i 0

ti 1 0 4 .0 0 -0 5 .3 0 25/30 1 i 0

Kajang M. violascens 2 1 9 .3 0 -2 3 .4 5 1 5 /6 0 u 02 11 n 1* 02 11 11 u 02 ii 11 1 £ 02 11 n 1* 0

Total 9 0

126

from d u sk u n t i l 2 3 . 0 0 h r s on e a c h o f two n i g h t s i n a l a r g e

g r o v e o f c u l t i v a t e d b a n a n a s w h ic h grew a r o u n d a v i l l a g e i n

t h e m id d le o f a r u b b e r p l a n t a t i o n n e a r Meru v i l l a g e , 5 km from

t h e Meru- F o r e s t R e s e r v e w h e re M. s o b r i n u s was c a u g h t and 7 km

from R a n ta u P a n j a n g w here M. m in im us was so common. No M a c r o g lo s s u s

o f s i fix git s p e c i g s y/GjTG c au g h 4c or* so o n t-h.oi' s (s . l t i io i ig i i E# spo l aG a

was c a u g h t and s e e n ) .

S i m i l a r l y i n t h e U n i v e r s i t y o f M a lay a campus w h ere

c u l t i v a t e d b a n a n a s w ere n u m e r o u s , a l t h o u g h s c a t t e r e d i n s m a l l

p a t c h e s , a n d w h e re D u r io z i b e t h i n u s and O ro xy lu m in d i c u m ( b o t h

c h i r o p t e r o p h i l o u s t r e e s ) w e re g row n , no M. s o b r i n u s w ere e i t h e r

c a u g h t o r s e e n ( f o r d a t a on o b s e r v a t i o n s on t h e 0 . i n d i c u m t r e e s

a t n i g h t s e e u n d e r E . s p e l a e a , p . 1 8 0 ) . At R a n ta u P a n ja n g w h e re

t h e r e w ere no w i l d Musa s p p . , o n l y M a c r o g lo s s u s m in im us v/as

c a u g h t among t h e c u l t i v a t e d b a n a n a s .

3 . i i i . Musa v i o l a s c e n s .

Musa v i o l a s c e n s i s t h e t h i r d common, w i l d s p e c i e s

o f Musa i n P e n i n s u l a r M a l a y s i a . I t s i n f l o r e s c e n c e d i f f e r s f rom

t h e s p e c i e s m e n t io n e d a b o v e i n many m o r p h o l o g i c a l c h a r a c t e r s ,

t h e m o s t i m p o r t a n t o f w h ic h i s i t s u p r i g h t p o s t u r e ( F i g . 11) .

M a c r o g l o s s u s s o b r i n u s was n e v e r s e e n t o v i s i t

Musa v i o l a s c e n s . I n n i n e h o u r s d u r i n g w h ic h i n f l o r e s c e n c e s o f

t h i s s p e c i e s w e re k e p t u n d e r o b s e r v a t i o n , no v i s i t b y an y s p e c i e s

o f b a t was n o t e d a l t h o u g h M. s o b r i n u s v i s i t e d Musa m a l a c c e n s i s

an d M. m a l a c c e n s i s x t r u n c a t a t h a t g rew a l o n g s i d e a n d w h ich

w ere w a tc h e d s i m u l t a n e o u s l y ^ e l e v e n t i m e s ( T a b l e 2 3 ) .

A t K a j a n g , w h e re Musa v i o l a s c e n s was v e r y common .

127/

two 10 m n e t s w ere s e t on t h r e e n i g h t s from d u s k t o m i d n i g h t

i n t h e h e a d o f a s m a l l v a l l e y ( s i t e 3 , F i g . 3 ) w h e re 19

M. v i o l a s c e n s an d 2 M. m a l a c c e n s i s w ere f l o w e r i n g , b u t no

M a c r o g l o s s u s s o b r i n u s was c a u g h t i n th e m . I f M. s o b r i n u s d i d

f e e d from M. v i o l a s c e n s f l o w e r s , some w ould p r o b a b l y h a v e b e e n

C & U gh 't u 6 r c W h6 i~6 JCi i 6 1 '6 WS T B iu01“6 f l 0 w 6 x “i n g K • V l. 0 .1 <5. S C 6 i i5 t h & ii

t h e r e w e re f l o w e r i n g p l a n t s o f e i t h e r Musa s p p . a t an y o t h e r

s i t e i n t h e s t u d y a r e a .

The o n l y o t h e r K a ja n g s i t e s a t w h ic h no M. s o b r i n u s

w ere c a u g h t w e re s i t e s 6 a n d 7 . H ow ever , o n ly one 10 m n e t was

s e t on two n i g h t s from d u sk u n t i l m i d n i g h t a t s i t e 6 and one

10 m n e t f o r one n i g h t a t s i t e . 7 o v e r t h e same t i m e . F u r t h e r m o r e ,

b o t h t h e s e s i t e s w e re v e r y e x p o s e d an d t h e few Musa s p p .

i n f l o r e s c e n c e s i n t h e v i c i n i t y w ere w i d e l y s c a t t e r e d .

3 . i v . V i s i t i n g b a n a n a f l o w e r s .

When f o r a g i n g among b a n a n a p l a n t s y M a c r o g lo s s u s

s o b r i n u s f l y c l o s e t o t h e g ro u n d s e a r c h i n g f o r i n f l o r e s c e n c e s

f rom b e lo w , sw o o p in g up t o a l i g h t when one i s l o c a t e d . ( I t

w ou ld h a v e b e e n i m p o s s i b l e f o r them t o f o r a g e a m o n g s t t h e d e n s e

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

b e lo w t h e l e a v e s ( F i g . 11 ) p r e v e n t e d t h e s e a r c h i n g from a b o v e .

No p h o t o g r a p h s w e re o b t a i n e d o f t h i s s p e c i e s on b a n a n a i n f l o r e s c e n c e s

b u t a s f a r a s c o u l d b e s e e n . t h e y l a n d and f e e d i n t h e same way

a s do M. m in im u s on c u l t i v a t e d b a n a n a s a t R a n ta u P a n j a n g . 18

M. s o b r i n u s w e re c a u g h t w i t h Musa s p p . p o l l e n a d h e r i n g t o t h e i r

m u z z l e s .

F i g . show ed M. m in im us f e e d i n g from a c u l t i v a t e d

128

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

i n s e t t s i t s m u z z le i n t o a f e r t i l e m ale f l o w e r , t h e a n t h e r s b r u s h

o v e r t h e m u z z le d e p o s i t i n g p o l l e n a l o n g i t . I t a l s o d e m o n s t r a t e s

how t h e f l o w e r i s f o r c e d open c a u s i n g t h e n e c t a r t o r u n down

from t h e b a s e t o t h e t i p o f t h e f r e e t e p a l , an d i n t o t h e b a t ' s

m outh a s d e s c r i b e d b e f o r e .

Where s e v e r a l b a n a n a s w ere f l o w e r i n g i n one g r o v e ,

v i s i t i n g M. s o b r i n u s move r a p i d l y from one i n f l o r e s c e n c e t o

a n o t h e r , s o m e t im e s r e t u r n i n g two o r t h r e e t i m e s t o some o f th em .

E ach t im e t h e b a t s e t t l e s f o r o n l y two o r t h r e e s e c o n d s .

A l l o b s e r v a t i o n s o f M. s o b r i n u s l a n d i n g on b a n a n a

i n f l o r e s c e n c e s w e re made by m o o n l i g h t . ( T o r c h l i g h t was u s e d t o

i l l u m i n a t e t h r e e i n f l o r e s c e n c e s b e tw e e n 0 4 . 0 0 an d 0 5 . 3 0 h r s a t

t h e 1 4 ^ m i l e , T a p a h - R i n g l e t r o a d b u t no b a t s v i s i t e d them d u r i n g

t h i s p e r i o d . ) Even when t h e r e i s a f u l l moon t h e d e e p s h a d e

i n s i d e a b a n a n a g r o v e an d t h e r a p i d i t y w i t h w h ic h M. s o b r i n u s

move from one p l a n t t o a n o t h e r made i t v e r y d i f f i c u l t t o r e c o r d

e a c h l a n d i n g on e a c h i n f l o r e s c e n c e . T h e r e f o r e , " a v i s i t ” a s

r e c o r d e d i n T a b le 25 r e f e r s t o an y o c c a s i o n on w h ic h a b a t l a n d e d

on one o r more o f t h e i n f l o r e s c e n c e s b e i n g w a tc h e d i r r e s p e c t i v e

o f hov/ many o f t h e i n f l o r e s c e n c e s i n t h a t g r o u p o f p l a n t s i t

l a n d e d on o r how many t i m e s i t l a n d e d on a n y o f them so l o n g

a s i t r e m a in e d i n t h e im m e d ia te v i c i n i t y . S i n c e i n t e r v a l s b e tw e e n

v i s i t s w e re n o r m a l l y f a i r l y l o n g ( e . g . t h e s h o r t e s t r e c o r d e d

i n t e r v a l was s e v e n m i n u t e s ) an d t h e d u r a t i o n o f ,v i s i t s , a s

d e f i n e d h e r e , was u s u a l l y b e tw e e n two and t h i r t y s e c o n d s ,

l i t t l e p r a c t i c a l d i f f i c u l t y was e x p e r i e n c e d i n d e c i d i n g w h a t

t i lc o n s t i t u t e d a v i s i t . At K a ja n g and a t . t h e 14 m i l e , T a p a h -

R i n g l e t r o a d , w h e re M. m a l a c c e n s i s ( o r M. m a l a c c e n s i s x t r u n c a t a )

an d _M. v i o l a s c e n s w e re w a tc h e d s i m u l t a n e o u s l y , p a r t i c u l a r

a t t e n t i o n was p a i d t o w h ic h i n f l o r e s c e n c e s t h e b a t s l a n d e d o n .

3 . v . P l a n t s b e s i d e s Musa s p p .

M a c r o g l o s s u s s o b r i n u s was n o t s e e n t o v i s i t t h e

f l o w e r s o f an y o t h e r p l a n t s p e c i e s a l t h o u g h t h e r e i s e v i d e n c e

f rom a n a l y s i s o f p o l l e n on t h e i r b o d i e s and i n t h e i r f a e c e s

t h a t t h e y v i s i t D uabanga g r a n d i f l o r a and p o s s i b l y D u r io z i b e t h i n u s .

The o b s e r v a t i o n s made on v i s i t s b y b a t s t o o t h e r c h i r o p t e r o p h i l o u s

p l a n t s a r e d e s c r i b e d u n d e r M a c r o g lo s s u s m in im u s an d E o n y c t e r i s

s p e l a e a .

k* A n a l y s i s o f p o l l e n s a m p le s from M a c r o g lo s s u s s o b r i n u s .

The n u m e r i c a l c o m p o s i t i o n o f p o l l e n i n t h e f a e c e s

an d g u t s a m p le s from ^6 M. s o b r i n u s a r e g i v e n i n T a b le 26 .

I n a d d i t i o n t o t h e s e , p o l l e n s a m p le s w ere o b t a i n e d from t h e

f u r on t h e h e a d s o f 19 b a t s .

i * . i . P o l l e n s a m p le s o b t a i n e d from b a t s c a u g h t a t K a ja n g .

Ten o f t h e g u t a nd f a e c a l s a m p le s an d s e v e n o f

t h e s a m p le s f rom t h e h e a d s o f b a t s w ere o b t a i n e d a t K a ja n g w h ere

t h e r e w e re no known c h i r o p t e r o p h i l o u s p l a n t s p e c i e s b e s i d e s

Musa m a l a c c e n s i s ( a n d c u l t i v a t e d b a n a n a s ) . T h e se s a m p le s

c o n t a i n e d o n l y Musa s p p . p o l l e n .

4 » i i . P o l l e n s a m p le s o b t a i n e d from b a t s c a u g h t u n d e r c h i r o p t e r o p h i l o u s

t r e e s b e s i d e s Musa s p p .

a ) U nder P a r k i a s p e c i o s a . S am p le s o f t h e g u t c o n t e n t s w ere

Table 26.

The table records the ebundance of each pollen type

found in gut or faecal samples collected from 46 Macroglossus

sobrinus in Peninsular Malaysia.

The f i g u r e s show th e number of s am r le s c o n t a i n i n g

100%, 75 to 99%, 50 to 74%, 25 to 49% or l e s s th an 25% (b u t

a t l e a s t one p o l l e n g r a i n ) r e s p e c t i v e l y o f each p o l l e n s p e c i e s .

The f i g u r e s i n b r a c k e t s a r e th e p e r c e n t a g e s of th e t o t a l

number o f s am p le s .

100% 75-99% 50-74% 25-49% < 25% T o t a l

Musa s p p .

Duabanga

Durio sp p .

36(78.3)

1( 2.2)

N i l

4(8.7)

4(8.7)

N i l

1(2.2)

N i l

N i l

N i l

1(2.2)

N i l

4(8.7)

1(2.2)

2(4.3)

45(97.8)

7(15.2)

2( 4.3)

Note buabanga = Duabanga g r a n d ! f l o r a .

■ Ij

130

o b t a i n e d from f i v e M. s o b r i n u s c a u g h t i n J u n e 1973 u n d e r at ilf l o w e r i n g P . s p e c i o s a t r e e i n s e c o n d a r y f o r e s t a t t h e 14

m i l e , T a p a h - R i n g l e t r o a d . Musa s p p . p o l l e n was t h e o n l y p o l l e n

s p e c i e s p r e s e n t i n t h e s e s a m p le s a nd i n two o t h e r s c o l l e c t e d

f r o m ' t h e m u z z l e s o f t h e s e b a t s .

b ) U nder D u r io z l b e t h i n u s . F a e c a l s a m p le s w e re c o l l e c t e d from

s i x b a t s c a u g h t u n d e r a f l o w e r i n g D. z i b e t h i n u s t r e e a t t h e

F . S . C . , U lu Gombak b e tw e e n 30 March and 5 A p r i l 1 9 7 3 . F o u r o f

them c o n t a i n e d o n l y Musa s p p . p o l l e n . One c o n t a i n e d 19 (4%)

D u r io s p p . g r a i n s among 500 p o l l e n g r a i n s t h a t w e re c o u n te d

an d one c o n t a i n e d one D u r io s p p . g r a i n an d one D uabanga g r a n d i f l o r a

g r a i n o u t o f 231 p o l l e n g r a i n s i n t h e s a m p le . Musa s p p . made

up t h e r e s t o f t h e s e two s a m p l e s . ( C o n t a m i n a t i o n may h a v e b e e n

r e s p o n s i b l e f o r some o f t h e D u r io s p p . p o l l e n an d t h e one D.

g r a h d i f l o r a p o l l e n g r a i n i n t h e s e s a m p l e s . ) Two s a m p le s from

t h e m u z z le s o f t h e s e b a t s c o n t a i n e d o n ly Musa s p p . p o l l e n .

c ) U nder D uabanga g r a n d i f l o r a . G ut s a m p le s w ere c o l l e c t e d from

s e v e n b a t s c a u g h t u n d e r a s t a n d o f f l o w e r i n g D. g r a n d i f l o r a

t r e e s i n a c l e a r i n g , i n s e c o n d a r y f o r e s t a t t h e 1 7 ^ m i l e , T a p a h -

R i n g l e t r o a d on 14 M arch 1 9 7 3 . Musa t r u n c a t a was common a ro u n d

t h e m a r g in s o f t h e c l e a r i n g . One s a m p le c o n t a i n e d 100% D.

g r a n d i f l o r a p o l l e n . T h re e c o n t a i n e d from 75% t o 99% D. g r a n d i f l o r a

p o l l e n an d one c o n t a i n e d 37% D. g r a n d i f l o r a p o l l e n . A l l t h e

r e m a i n i n g p o l l e n was Musa s p p . i n c l u d i n g two t h a t c o n t a i n e d

o n l y Musa s p p . p o l l e n .

P o l l e n s a m p le s w e re c o l l e c t e d from t h e h e a d s o f

t h r e e o f t h e s e b a t s . One c o n t a i n e d 100% Musa s p p . p o l l e n ( a l l

131

on t h e m u z z l e ) , one 100% D. g r a n d i f l o r a p o l l e n ( a l l o v e r h e a d

an d c h e s t ) and t h e t h i r d c o n t a i n e d a m i x t u r e o f a p p r o x i m a t e l y

k0% Musa s p p . p o l l e n an d 60% D. g r a n d i f l o r a p o l l e n on t h e

m u z z l e , b u t o n l y D. g r a n d i f l o r a e l s e w h e r e on i t s h e a d .

4 . i i i . O th e r p o l l e n s a m p le s o b t a i n e d from M a c r o g l o s s u s s o b r i n u s .

bo d y s a m p le s w e re c o l l e c t e d from b a t s c a u g h t n e a r t o f l o w e r i n g

Musa s p p . i n o r on t h e edge o f s e c o n d a r y f o r e s t .

c h i r o p t e r o p h i l o u s t r e e s , b e s i d e s v a r i u o s Musa s p p . , e x c e p t a s

d e s c r i b e d a b o v e . H o w ever , o t h e r c h i r o p t e r o p h i l o u s s p e c i e s o f t e n

g rew i n t h e v i c i n i t y o f t h e n e t l o c a t i o n s ( e x c e p t on Gunong ,

B r i n c h a n g w h e re no o t h e r known c h i r o p t e r o p h i l o u s t r e e s w e re

s e e n ) : G ut s a m p le s w e re c o l l e c t e d from f o u r b a t s c a u g h t on t h e

m o u n ta in t o g e t h e r w i t h a s a m p le from t h e m u z z le o f one o f

them ; t h e y a l l c o n t a i n e d o n l y Musa s p p ; p o l l e n . E . g . t h e s i t e

a t t h e 1 4 ^ m i l e , T a p a h - R i n g l e t r o a d , was w i t h i n 200 m o f a

g r o v e o f P roxy lum i n d i c u m t r e e s i n w h ic h t h e r e w e re s i x t r e e s

c a r r y i n g a t o t a l o f t h i r t e e n i n f l o r e s c e n c e s , a t l e a s t two

f l o w e r i n g D uabanga g r a n d i f l o r a t r e e s and t h r e e o t h e r P a r k i a

s p e c i o s a t r e e s .

p r e s e n t i n t h e s e s a m p le s e x c e p t i n one f a e c a l s a m p le from t h e

F . S . C . , U lu Gombak i n w h ic h 97% o f t h e 500 p o l l e n g r a i n s t h a t

w e re c o u n t e d was D. g r a n d i f l o r a . The r e m a i n i n g 3% was Musa s p p .

The r e m a i n i n g 18 g u t and f a e c a l s a m p le s an d 13

No a t t e m p t was made t o c a t c h M. s o b r i n u s n e a r t o

P o l l e n o f Musa s p p . was t h e o n ly £ o l l e n v t y p e t

132

5« I t e m s b e s i d e s p o l l e n i n t h e g u t and f a e c a l s a m p le s o f

M a c r o g l o s s u s s o b r l n u s .

5 . i . I t e m s o f unknown o r i g i n .

A morphous m a t t e r s i m i l a r t o t h a t f o u n d i n M a c r o g lo s s u s

m in im us f a e c a l s a m p le s was o n l y r e c o r d e d i n two (2+ .3%) o f t h e

g u t and f a e c a l s a m p le s from M. s o b r l n u s . I n b o t h i t was a s s o c i a t e d

w i t h Musa s p p . p o l l e n . I t was n o t a b u n d a n t i n e i t h e r .

5 . i i . I t e m s o f p l a n t o r i g i n .

S m a l l c lu m p s o f p l a n t c e l l s w e re p r e s e n t i n one

sa m p le from U lu Gombak (w h ic h a l s o c o n t a i n e d a m o rph ou s m a t t e r ) .

No e p i d e r m a l , v a s c u l a r o r o t h e r c h a r a c t e r i s t i c c e l l t y p e s w ere

p r e s e n t an d t h e t i s s u e from w h ich t h e y w e re d e r i v e d i s n o t

know n. A s t r i p o f f i b r o u s t i s s u e was p r e s e n t i n one s a m p le

f rom K a j a n g . B o th o f t h e a b o v e s a m p le s a l s o c o n t a i n e d Musa s p p .

p o l l e n g r a i n s . No f u n g a l r e m a i n s w e re r e c o g n i s e d . .

5 . i i i . I t e m s o f a n i m a l o r i g i n .

M i t e s ( A c a r i n a ) w ere p r e s e n t i n 8 ( 1 7 . A-%) s a m p le s

i n c l u d i n g two fro m 1800 m on Gunong B r i n c h a n g a n d f o u r from

K a j a n g . T h ese w e re p r o b a b l y i n g e s t e d a c c i d e n t a l l y w h i l e f e e d i n g

on b a n a n a f l o w e r s a s t h e r e w e re no o t h e r known a l t e r n a t i v e fo o d

s o u r c e s from w h ic h t h e y c o u l d h a v e come a t e i t h e r o f t h e s e

l o c a l i t i e s . The r e m a i n i n g two w e re from U lu Gombak and 1 7 ^

m i l e , T a p a h - R i n g l e t r o a d r e s p e c t i v e l y . The l a s t s a m p le a l s o

c o n t a i n e d one s m a l l n e m a t o c e r a n ( D i p t e r a ) . S m a l l f r a g m e n t s o f

c h i t i n and i n s e c t s c a l e s w e re fo u n d i n one s a m p le f rom U luf Jn

Gombak an d i n s e c t s c a l e s w e re p r e s e n t i n one from 14 m i l e ,

T a p a h - R i n g l e t r o a d .

F u r w as a b s e n t e x c e p t i n two s a m p le s from 1 7 ^

m i l e , T a p a h - R i n g l e t r o a d (w h ic h a l s o c o n t a i n e d a h i g h p r o p o r t i o n

o f D. g r a n d i f l o r a p o l l e n ) .

No p a r a s i t e s w e re r e c o v e r e d from t h e a l i m e n t a r y

c a n a l s o r t h e f a .e c e s o f M. s o b r i n u s .

6 . F l i g h t .

6 . i . M o r p h o l o g i c a l c h a r a c t e r i s t i c s r e l a t i n g t o f l i g h t .

The w in g c h a r a c t e r i s t i c s o f M a c r o g l o s s u s s o b r i n u s

a r e v e r y s i m i l a r t o t h o s e o f M. m in im us ( T a b l e s 12 an d 13 ) -and

th e comments g i v e n f o r t h e l a t t e r s p e c i e s ( p . 107) a p p l y e q u a l l y .

L ik e M. m i n im u s , t h e b o d i e s o f M. s o b r i n u s a r e

n o t u n d u e ly c o m p ac t o r s t r e a m l i n e d . The f u r i s t h i c k an d l o n g .

I t d o e s n o t l i e c l o s e l y a g a i n s t t h e s k i n a s i t d o e s i n E o n y c t e r i s

s p e l a e a .

6 . i i . F l i g h t .

M a c r o g lo s s u s s o b r i n u s was c a p a b l e o f f l y i n g

f r e e l y a nd w i t h g r e a t a g i l i t y b e tw e e n t h e p s e u d o s t e m s o f

b a n a n a s g ro w in g i n d e n s e g r o v e s ( e . g . F i g . 11 ) . ( C a p t i v e b a t s

w e re a b l e t o f l y b a c k and f o r t h i n a c a g e m e a s u r i n g \ m x 1 m

x 1 . 3 m h i g h , b u t w i t h l e s s f a c i l i t y t h a n M. m in im u s ) . I n more

open s i t u a t i o n s t h e i r f l i g h t was f a s t and d i r e c t and i t was

t h e n d i f f i c u l t t o d i s t i n g u i s h them by s i g h t f rom E o n y c t e r i s

s p e l a e a ( e s p e c i a l l y v e r y young E . s p e l a e a w h ic h t e n d e d t o f l y

more s l o w l y an d w h ich w ere a p p r e c i a b l y s m a l l e r t h a n t h e a d u l t s ) .

' s o b r i n u s f o r a g e d a l o n e n o t i n f l o c k s . (T h is

m

a f f o r d e d a n o t h e r m eans o f d i s t i n g u i s h i n g i t b y s i g h t a t n i g h t

from E . s p e l a e a w h ic h u s u a l l y t r a v e l l e d i n f l o c k s ) .

7 . A c t i v i t y p a t t e r n . .

F i g . 58 and T a b le 27 show W i l l i a m s ' mean f o r t h e

iiuiul)G!T o f 1' iaCro^ 1 o s 5 u s s coi~iLiiu3 CcLugliu p e r i io u r dujriLng 6s.ch.

h o u r o f t h e n i g h t f rom 19*15 t o 0 6 .1 5 h r s . D a ta fro m 53 c a p t u r e

r e c o r d s h a v e b e e n i n c l u d e d . I n s u f f i c i e n t d a t a i s a v a i l a b l e on t h e

t im e o f c a p t u r e o f b a t s a t an y one p l a c e t o d raw a h i s t o g r a m

f o r a s i n g l e l o c a l i t y . T h e r e f o r e d a t a h a s b e e n i n c l u d e d from

a l l l o c a l i t i e s a t w h ic h t h e s p e c i e s was c a u g h t e x c e p t a s s p e c i f i e d

b e lo w .

R e c o r d s w ere o m i t t e d w h e r e , i n h i n d s i g h t , i t

was r e a l i s e d t h a t t h e m i s t n e t s h a d b e e n s e t i n i n a p p r o p r i a t e

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

b a n a n a s . T h i s i n c l u d e s s e v e r a l r e c o r d s from U lu Gombak and some

from e l s e w h e r e . D a ta was u n a v a i l a b l e f o r t h r e e n i g h t s from

1 9 .1 5 t o 0 0 . 1 5 h r s an d one from 19*15 t o 2 1 . 1 5 h r s a t K a j a n g .

On e a c h o c c a s i o n t h a t h a s b e e n i n c l u d e d , f i v e 10 m

n e t s w ere u s e d e x c e p t t w i c e a t K a ja n g when s i x n e t s w ere s e t

an d t h r e e t i m e s a t U lu Gombak when f o u r n e t s w e re s e t . The

f i g u r e s i n T a b le 27 a r e t h e r e f o r e n e a r l y p r o p o r t i o n a l t o t h e

num ber o f b a t s c a u g h t , p e r h o u r p e r n e t .

The i n c l u s i o n o f d a t a from d i f f e r e n t l o c a l i t i e s

i n t r o d u c e s two f a c t o r s w h ic h c o m p l i c a t e an y i n t e r p r e t a t i o n o f

t h e a c t i v i t y p a t t e r n o f M. s o b r l n u s d u r i n g t h e n i g h t . T h ese

a r e

1 . P o s s i b l e v a r i a t i o n i n p o p u l a t i o n d e n s i t y b e tw e e n

Table 27 .The t a b l e shows th e t o t a l mean and W i l l i a m s ' mean

for the number of Macroglossus sobrlnus caught per hour in Peninsular Malaysia.

Time No. of n i g h t s No. o f b a t s No. o f b a t s > p e r hour

W il l iams 'mean

1 9.00-1 9.1 5 1 5 0 0 0

1 9 .1 5 -2 0 .1 5 15 13 0 .8 7 0 .6 1

20.1 5-21 .1 5 15 10 0 .6 7 0 .4 721 .1 5-22.1 5 14 9 0 .6 4 0 .322 2 .1 5 -2 5 .1 5 14 6 0 .43 0 .4 02 3 . 1 5-CO. 15 1 4 • 5 0 .3 6 0 .2 500.1 5-01 .1 5 10 1 0 .1 0 0 . 1 ?01 .1 5-02.1 5 8 2 0 .2 5 . , 0 .1 902.1 5-03.1 5 8 2 0 .2 5 0.1 50 3 .1 5—Oh. 1 5 8 3 0 .3 7 0 .2 5Oh.15-05.1 5 8 2 0 .2 5 0.1 905.1 5-06.1 5 8 0 0 0

F ig . 58.The figure illustrates graphically the data for Williams'

mean recorded in Table 2?»

o.6_

0 . 5-

c(0| O.lH

a 0 -3 -a■HHrH

■H

o.r

IAIAir \ m lA lA ITS inin

inin in in in

Time

l o c a l i t i e s , e . g . i f t h e p o p u l a t i o n was u n u s u a l l y d e n s e a t

l o c a l i t i e s w h e re r e c o r d s w ere o n ly a v a i l a b l e up t o m i d n i g h t

b u t u n u s u a l l y low w h e re r e c o r d s w ere a v a i l a b l e f o r t h e w h o le

n i g h t , an a b n o r m a l l y h i g h e s t i m a t e o f r e l a t i v e a c t i v i t y w ou ld

b e made f o r t h e p e r i o d t o m i d n i g h t . H ow ever, no e v i d e n c e s u g g e s t

t h a t t h i s ws.s t h s c s s s a t s n v oif t h e l o c a l i . t i . s s v i s i . t s c lc

2 . A t K a ja n g an d a t t h e F . S . C . , U lu Gombak, b a t s

w ere r i n g e d an d r e l e a s e d . A t o t h e r l o c a l i t i e s , b a t s w ere u s u a l l y

n o t r e l e a s e d an d n e v e r r i n g e d . S in c e r e c a p t u r e r e c o r d s h av e

b e e n i n c l u d e d w h e re a p p r o p r i a t e , t h i s may h a v e a f f e c t e d r e s u l t s .

H ow ever , an y e f f e c t s a r e n o t l i k e l y t o b e l a r g e b e c a u s e , b e s i d e s

K a ja n g an d t h e F . S . C . , n e t s w e re n o t s e t a t p r e c i s e l y t h e same

l o c a l i t y f o r more t h a n one n i g h t a nd o n ly two o f t h e 37 r i n g i n g

r e c o r d s t h a t a r e i n c l u d e d w ere o f b a t s c a u g h t t w i c e i n t h e same

n i g h t .

T h e r e f o r e , t h e h i s t o g r a m p r o b a b l y d o e s show t h e

. a c t i v i t y p a t t e r n f o r M. s o b r i n u s d u r i n g t h e n i g h t .

8 . R i n g i n g r e s u l t s .

8 . i . P o p u l a t i o n s t r u c t u r e .

B e s i d e s t h e 65 M a c r o g lo s s u s s o b r i n u s c a u g h t d u r i n g

t h e s t u d y , 1+7 s p e c i m e n s from P e n i n s u l a r M a l a y s i a ( m o s t l y from

P a h a n g , P e r a k an d S e l a n g o r ) d e p o s i t e d i n t h e IMR c o l l e c t i o n

w e re e x a m in e d . The s e x r a t i o o f a d u l t s t o s u b a d u l t s a r e g iv e n

i n T a b le 28 . I n b o t h g r o u p s t h e r e i s an e x c e s s o f f e m a le s w i t h

a mean r a t i o o f hk*7% m a le t o 55*3% f e m a l e . The d i f f e r e n c e i s

n o t s i g n i f i c a n t (p = 0 . 5 ) and b o t h show an a d u l t : s u b a d u l t r a t i o

o f a p p r o x i m a t e l y 5:1 •

Table 28 .

The t a b l e r e c o r d s t h e sex r a t i o and th e r a t i o of

a d u l t to s u b a d u l t M acrog lossus s o b r in u s caugh t i n P e n i n s u l a r

M alays ia

Total no. specimens

Source c f ad. ? ad. Total ad.

Totalsad.

65 This studyn %

25(4 6 .3 )

29(5 3 .7 )

54(8 3 .1 )

1 V(1 6 .9 )

47 IMR n

collection %17

( 4 2 .3)23

(57 .5 )40

(8 5 .1 )7

(14 .9 )

112 Total 0 %

42

(4 4 .7 )52

(55 .3 )94

(8 3 .9 )18

( 1 6 . 1 )

Note ad. = adult,sad. = subadult,IMR = Institute of Medical Research, Kuala Lumpur.

136

8 . i ± . M ovem ents .

F . S . C . , U lu Gombak. 21 M a c r o g l o s s u s s o b r i n u s w e re r i n g e d a t

t h e F . .S .C . , U lu Gombak. None o f t h e s e v e n a d u l t o r two s u b a d u l t

m a le s w ere r e c a p t u r e d . H o w ever , f i v e o f t h e n i n e a d u l t f e m a le s

w ere r e c a p t u r e d ( t h r e e w ere c a p t u r e d tw ic e an d .two t h r e e t i m e s ) .

The l o n g e s t i n t e r v a l b e tw e e n c a p t u r e and r e c a p t u r e o f e a c h o f

t h e s e f i v e b a t s was 6 , 6 1 , 7 8 , 96 an d 356 d a y s r e s p e c t i v e l y .

The o n l y s u b a d u l t f e m a le c a u g h t t h e r e was c a p t u r e d t h r e e t i m e s

i n t e n d a y s .

K a j a n g . 20 M. s o b r i n u s w ere r i n g e d a t K a ja n g b e tw e e n 16 O c to b e r

and 20 December 1 9 7 3 . T h re e o f t h e e i g h t a d u l t m a le s w ere n o t

r e c a p t u r e d , two w e re c a p t u r e d t w i c e , one t h r e e t i m e s an d two

f i v e t i m e s . Two s u b a d u l t m a l e s , one s u b a d u l t f e m a le an d f i v e

o u t o f s i x a d u l t f e m a le s w e re n o t r e c a p t u r e d b u t one a d u l t

f e m a le was c a p t u r e d t h r e e t i m e s . I n t e r v a l s b e tw e e n c a p t u r e and

r e c a p t u r e v a r i e d from t w i c e i n a n i g h t t o 65 d a y s . (T he l a t t e r ,

a n a d u l t m a le No. 20A -9660 was c a p t u r e d on t h e f i r s t and

r e c a p t u r e d on t h e l a s t n i g h t o f r i n g i n g a t K a j a n g . I t was a l s o

c a u g h t on two i n t e r v e n i n g n i g h t s . ) R e c a p t u r e d a t a i s su m m ar ised

i n T a b le .29-

D e s p i t e t h e a p p a r e n t d i f f e r e n c e s i n t h e r e c a p t u r e

r a t e o f m a le s an d f e m a l e s (w h ic h may b e an a r t i f a c t due t o t h e

s m a l l sa m p le s i z e s ) a t e a c h l o c a l i t y , i t . i s c l e a r t h a t some

s o b r i n u s o f b o t h s e x e s r e m a in i n o r r e t u r n t o t h e same s i t e s

o v e r e x t e n s i v e p e r i o d s o f t i m e .

Some e v i d e n c e o f movement w i t h i n t h e f e e d i n g

r a n g e was o b t a i n e d a t K a j a n g . The l o c a t i o n o f n e t s i t e s i n

r e l a t i o n t o t h e d i s t r i b u t i o n o f b a n a n a f l o w e r s t h e r e i s

Table 2 9 #

The t a b l e shows d e t a i l s o f the r e c a p t u r e r e c o r d s

f o r M acroglossus s o b r in u s cau g h t a t Kajang , S e l a n g o r . A l l

b a t s were a d u l t s . For th e l o c a t i o n o f n e t s i t e s see F i g . 59#

No. Sex Dates on which captured

Net-No. Time

20/49680 female 212720

Nov.- Nov. Dec.

197319731973

59

10

20/49658 male 16 Oct. 1973 •121 Oct. 1973 1

20/49660 male 16 Oct. 1973 2 2 0 .00 hrsit 2 20 .30 hrs

21 Nov. 1973 427 Nov. 1973 920 Dec. 1973 10

20/49685 male 21 Nov. 1973 527 Nov. 1973 8 2 0 .30 hrs

ii 8 2 2 .30 hrs20 Dec. 1973 5 19.30 hrs

n 8 21 .30 hrs20/49686 male 21

2720

Nov. Nov. Dec.

197319731973

585

30/01 765 male 2120

Nov. Dec.

19731973

55

137

i l l u s t r a t e d i n F i g . 59 w here a r e a s i n w h i c h ^ i n d i v i d u a l b a t s

w ere r e c a p t u r e d h a v e b e e n s u p e r im p o s e d on t h e t o p o g r a p h i c a l

map. Mi s o b r i n u s moved e x t e n s i v e l y a l o n g t h e s t r e a m s from one

c lum p o f f l o w e r i n g b a n a n a s t o a n o t h e r . They p r o b a b l y v i s i t e d

a l l p a r t s o f t h e a r e a a t w h ic h Musa m a l a c c e n s i s g rew ( a l t h o u g h

d a t a i s n o t s u f f i c i e n t t o show t h i s ) , . The Granges o f a d u l t m a le s

o v e r l a p e a c h o t h e r e x t e n s i v e l y an d t h e t h r e e f e m a l e s t h a t w ere

k i l l e d w e re a l l c a u g h t w i th in one h o u r o f e a c h o t h e r a t n e t

s i t e 1 .

9 . A bundance o f M a c r o g l o s s u s s o b r i n u s a t K a ja n g i n r e l a t i o n t o

t h e a b u n d a n c e o f b a n a n a ' f l o w e r s .

A t K a j a n g , e i g h t a d u l t m a l e s , n i n e a d u l t f e m a le s

( i n c l u d i n g t h r e e k i l l e d on 5 O c to b e r 1973) and t h r e e s u b a d u l t s

w e re c a u g h t . The a d u l t s e x r a t i o i s k7% m ale t o 53% fe m a le and

85% o f t h e b a t s w ere a d u l t . D e s p i t e t h e s m a l l s a m p le s i z e s ,

t h e s e f i g u r e s a p p r o x i m a t e v e r y c l o s e l y t o t h o s e g i v e n i n T a b le 28

f 6 r 112 M. s o b r i n u s ex am in ed a l t o g e t h e r i n P e n i n s u l a r M a l a y s i a .

On 20 December 1 9 7 3 , t h e l a s t d a y on w h ich b a t s

w e re c a u g h t a t K a j a n g , s e v e n M. s o b r i n u s w ere c a u g h t . F i v e (71%)

w e re r e c a p t u r e s , i n c l u d i n g a l l f o u r a d u l t m a le s c a p t u r e d t h a t

n i g h t . On t h e p e n u l t i m a t e o c c a s i o n , f o u r (50%) o f t h e e i g h t

b a t s c a p t u r e d w e re r e c a p t u r e s ( i n c l u d i n g t h r e e o u t o f f o u r

a d u l t m a l e s ) .

The h i g h r e c a p t u r e r a t e s e s p e c i a l l y f o r a d u l t

m a le s c o u p l e d w i t h t h e c l o s e a g r e e m e n t o f a g e an d s e x r a t i o s

a t K a ja n g w i t h t h o s e fo u n d f o r t h e s p e c i e s a s a w h o le s u g g e s t

t h a t t h e m a j o r i t y o f t h e p o p u l a t i o n i n t h e s t u d y a r e a had b e e n

r i n g e d . T h e r e f o r e , t h e r e w e re p r o b a b l y b e tw e e n tw e n ty and

FIG 59 THE K N O W N R A N G E S O F T WO M A C R O G L O S S U S SOBRI NUS AT K A J A N G BASED O N RECAPTURE RECORDS

W \ ' 2 v

\ 2 m ? 1 0 v

0-5 ki

N o t s i t e sf l o w e r i n g M u s a m a l a c c e n s i s

» M . v i o l a s c e n s" c u l t i v a t e d b a n a n a s

m

138

t h i r t y i n d i v i d u a l s ; - i t m u s t b e e m p h a s i s e d t h a t t h i s i s a

t e n t a t i v e f i g u r e .

The t o t a l num ber o f Musa m a l a c c e n s i s i n f l o r e s c e n c e s

i n t h e s t u d y a r e a on 21 November 1973 was 5 2 . T h e re w e re a l s o

f o u r c u l t i v a t e d b a n a n a i n f l o r e s c e n c e s and a p p r o x i m a t e l y 49

Musa v i o i a s c e n s i n f l o r e s c e n c e s . Beyond t h e s t u d y a r e a t h e o w ners

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

t h e r e w ere no o t h e r M. m a l a c c e n s i s f l o w e r s w i t h i n 1 km ( e x c e p t

two a b o u t 0 . 5 km n o r t h w e s t o f n e t 5 on a s e p a r a t e d r a i n a g e

s y s te m ) an d p r o b a b l y no ne f o r a c o n s i d e r a b l e d i s t a n c e b e y o n d .

t h a t . T h e re w e re b e tw e e n 15 an d 20 c u l t i v a t e d b a n a n a i n f l o r e s c e n c e s

w i t h i n 1- km o f t h e s t u d y a r e a , m o s t l y c o n c e n t r a t e d a r o u n d a

v i l l a g e , a b o u t 0 . 5 km t o t h e s o u t h e a s t .

S i n c e c u l t i v a t e d b a n a n a s r e l e a s e no p o l l e n and

t h e b a t s w e re a p p a r e n t l y n o t i n g e s t i n g p o l l e n from any s o u r c e

o t h e r t h a n M. m a l a c c e n s i s , 52 M. m a l a c c e n s i s f l o w e r s w ere

p r o v i d i n g a l l t h e p o l l e n r e q u i r e d b y 20 t o 30 M a c r o g l o s s u s -

s o b r i n u s (= 1 . 7 t o 2 . 6 i n f l o r e s c e n c e s p e r b a t ) . The c a r b o h y d r a t e

r e q u i r e m e n t s o f t h e b a t s w ore b e i n g s u p p l e m e n t e d b y n e c t a r from

c u l t i v a t e d b a n a n a s and a d d i t i o n a l p r o t e i n may h a v e b e e n o b t a i n e d

from i n s e c t s .

(N .B . i t h a s b e e n a s s u m e d , p o s s i b l y i n c o r r e c t l y , t h a t ( a ) t h e s e

b a t s ' w e r e n o t t r a v e l l i n g b e y o n d t h e s t u d y a r e a t o f e e d e x c e p t

t o some c u l t i v a t e d b a n a n a s w i t h i n 1 km, (b) t h e p o p u l a t i o n i n

t h e s t u d y a r e a n u m b ered b e tw e e n tw e n ty an d t h i r t y i n d i v i d u a l s ,

( c ) the bats did not feed on the flowers of Musa vioiascens. I t

should also be noted that a few Eonycteris spelaea also fed from

the banana flowers in the study area.

139

1 0 . R e p r o d u c t i o n .

Very l i t t l e data is available on the reproduction

of Macroglossus sobrinus. Adult males were caught in every

month of the' year except February, May and August. They a l l had

scrotal testes measuring approximately 9 mm in .length x 7 mm

in width and a white substance, presumably Spermatozoa, was

present in the epididymis of a l l that were dissected. Adult

males in the IMR collection collected in May and August had

testes of similar dimensions. Spermatogenesis i s probably

continous in adult males.

Pregnancy was recorded in March, April, September,

November and December and lactation was recorded in March, April,

June, July, September, October and November

In Peninsular Malaysia, M. sobrinus i s probably

an aseasonal, polyestrus species (see Wilson, 1 97 3 )*

thOne f e m a le c a u g h t a t t h e 17 m i l e , T a p a h - R i n g l e t

r o a d on 1 /+ March 1 973» w&s l a c t a t i n g h e a v i l y . The r i g h t u t e r i n e

h o rn was s t i l l c o n s i d e r a b l y d i s t e n d e d from h a v i n g c o n t a i n e d an

embryo and t h e r e was a s m a l l b u t d i s t i n c t s w e l l i n g a b o u t 1 . 5 mm

i n d i a m e t e r i n t h e l e f t u t e r i n e h o r n i n d i c a t i n g e a r l y p r e g n a n c y .

T h i s b a t showed t h a t e a r l y p r e g n a n c y can o v e r l a p l a c t a t i o n ,

t h a t t h e r e i s a p o s t - p a r t u m e s t r o u s an d t h a t e m b r y o n ic d e v e l o p ­

m en t may o c c u r i n e i t h e r u t e r i n e h o r n .

140

EONYCTERIS SPELAEA

! . E c o l o g i c a l d i s t r i b u t i o n i n P e n i n s u l a r M a l a y s i a .

E o n y c t e r i s s p e l a e a h a s b e e n c o l l e c t e d from a l l

s t a t e s i n P e n i n s u l a r M a l a y s i a ( F i g . 60 ) . D u r in g t h e p r e s e n t s t u d y

. i t was c a u g h t i n a num ber o f l o c a l i t i e s c o v e r i n g m o s t m a jo r

v e g e t a t i o n t y p e s i n t h e c o u n t r y (m a n g ro v e s , v a r i o u s c u l t i v a t i o n

t y p e s i n c l u d i n g m o n o c u l t u r e s and m ixed c u l t i v a t i o n , p r i m a r y and

s e c o n d a r y lo w la n d f o r e s t and s e c o n d a r y m o n tan e f o r e s t ) and r a n g i n g

i n a l t i t u d e from s e a l e v e l t o 1 800 m a . s . l . ( T a b l e 3 0 ) . A t many

o f t h e s e l o c a l i t i e s , o b s e r v a t i o n s w ere a l s o made on b a t s f e e d i n g

from f l o w e r s .

2 . R o o s t s .

Enorm ous c o l o n i e s o f E o n y c t e r i s s p e l a e a r o o s t

i n c a v e s . T h re e c a v e s c o n t a i n i n g r o o s t s o f many th o u s a n d E . s p e l a e a ,

a s w e l l a s o t h e r b a t s p e c i e s , w e re v i s i t e d i n P e n i n s u l a r M a l a y s i a .

They w e re Gua P o n d o k , n e a r K u a la K a n g s a r , P e r a k ; Gua S a n d in g ,

n e a r K am par, P e r a k an d B a tu C a v e s , n e a r K u a la L um pur. S e l a n g o r

( F i g . 2 ) . A s i n g l e b a t was c a u g h t r o o s t i n g i n a h o u s e a t R a n ta u

P a n j a n g b u t v i l l a g e r s c l a i m e d i t was u n u s u a l t o f i n d t h i s s p e c i e s

i n t h e i r d w e l l i n g s . No o t h e r e x t r a - c a v e r n i c o l o u s r o o s t s i t e s w e re

l o c a t e d d u r i n g t h e s t u d y .

The t h r e e c a v e s i n w h ich E . s u e l a e a w e re p r e s e n t

s h a r e d c e r t a i n c h a r a c t e r i s t i c s :

1 . They w ere a l l i n l i m e s t o n e o u t c r o p s ( b u t no l a r g e c a v e s h av e

b e e n r e p o r t e d i n P e n i n s u l a r M a l a y s i a e x c e p t i n l i m e s t o n e ) ,

2 . The c a v e s t h e m s e l v e s v/ere v e r y l a r g e . E . s p e l a e a se ld o m

F i g . 60. L o c a l i t y r e c o r d s of E o n y c t e r i s s p e l c e a in P e n i n s u l a r M a l a y s i a .

♦ ♦

k m

$ E . s p e l a e a

Table 30 .The t a b l e chows th e l o c a l i t i e s a t which E o n y c te r i s

s p e l a e a was c a u g h t , t h e number t h a t were r in g e d and the h a b i t a t s i n which they were caugh t d u r in g th e s t u d y . ( A l l l o c a l i t i e s a r e marked on F i g s . 1 o r 2 . )

L o c a l i t y S t a t e No.caugh t

No.r in g e d H a b i t a t

Batu Caves S e la n g o r 104 16

Gua Pondok Perak n i l * - ► Limestone cavesGua Sanding tt 68 42I 5 t h m i l e , U.G. S e la n g o r 53 52U t h m i l e , T-R F erak 2 - 2° lowland f o r e s tI 7 t h m i l e , T-R ii 1 -2 l s t m i l e , U.G. S e la n g o r 2 - 1 0 lowland f o r e s tG. B r inchang Pahang 1 - 2° montane f o r e s tKuala S e lan g o r S e la n g o r 4 - l>■ mangroves n e x t toRantau Pan jang ii 206 192 J mixed v e g e t a t i o nBako N.P. Sarawak 2 2 mangroves n e x t to

p r im ary v e g e t a t i o nMeru F.R. S e la n g o r 3 3 mixed c u l t i v a t i o n

n e x t to 2° f o r e s tMeru ( v i l l a g e ) ti 4 4 1 bananas i n ru b b e rKajang n 14 11 " e s t a t e

Kuala Lumpur ii 2 2 suburban garden

T o t a l 466 324

Note 1° = p r im a r y ,2° = s e c o n d a ry ,F .R . = f o r e s t r e s e r v e ,G. = Gunong = Mountain ,N.P . = N a t i o n a l P a rk ,T-R = Tapah R i n g l e t r o a d ,U.G. = Ulu Gombak.*

Gua Pondok - No l i v e b a t s caugh t a l th o u g h many were s e e n . E o d ie s were r e c o v e re d from th e cave f l o o r .

1t-1

p e n e t r a t e d p o r t i o n s w h e re t h e r o o f was l e s s t h a n 7 m a b o v e t h e

f l o o r an d t h e p a s s a g e w a y was e q u a l l y w id e . ( I n B a tu C aves a few

b a t s p a s s e d t h r o u g h t h e r e l a t i v e l y n a r r o w p a s s a g e b e tw e e n C av ern

B a nd t h e G r e a t Boom ( F i g . 6 ) b u t m o s t b a t s i n C a v e rn A an d B

l e f t v i a t h e " e n t r a n c e " w h i l e t h o s e i n C a v e rn C a n d t h e G r e a t

Boom l e f t v i a t h e h o l e i n t h e r o o f o f t h e G r e a t Room.) The b a t s

u s u a l l y r o o s t e d 10 n o r more a b o v e t h e f l o o r . The c a v e s w e re n o t

n e c e s s a r i l y d e e p : Gua S a n d in g i s h o r s e - s h o e s h a p e d w i t h a l a r g e

e n t r a n c e a t e a c h e n d an d no p o i n t i n t h e c a v e i s more t h a n 5 0 m

• f ro m one e n t r a n c e o r t h e o t h e r . I n c o n t r a s t , i n C a v e rn C, B a tu

C a v e s , some b a t s r o o s t i n t o t a l d a r k n e s s a b o u t kOO m from t h e i r

e x i t i n t h e G r e a t Room an d 300 m from t h e n e a r e s t l i g h t s o u r c e ,

a s m a l l c h im ney n e a r t h e mouth o f C av e rn C

3 . The e n t r a n c e s t o a l l t h e c a v e s a r e l a r g e , 7 m o r more h i g h

an d a s w i d e . They a l l open a b o v e g ro u n d l e v e l an d a r e u n o b s t r u c t e d

b y t r e e s o r o t h e r o b s t a c l e s . The h o l e i n t h e r o o f o f t h e G r e a t

Room i s e x c e p t i o n a l i n t h a t t h e b a t s h a v e t o f l y u p w a rd s t o e x i t .

No m e a s u re m e n ts o f t h e h u m i d i t y o r t e m p e r a t u r e

o f t h e a i r a d j a c e n t t o t h o s e p o r t i o n s o f t h e c a v e r o o f s on w h ich

E . s p e l a e a r o o s t e d a r e a v a i l a b l e b u t t h e b a t s a p p e a r e d t o t o l e r a t e

w id e p a r a m e t e r s f o r b o t h f a c t o r s . Deep i n s i d e a c a v e , t e m p e r a t u r e

an d h u m i d i t y a r e f a i r l y c o n s t a n t ( B u l l o c k , 1 9 6 5 ) . T h i s i s t r u e

o f C a v e rn C o f B a tu C av es (M c C lu re , Lira and W inn, 1 9 6 ? ) w h ere

t h e r e l a t i v e h u m i d i t y a p p r o a c h e s 100% an d a l i g h t m i s t i s p r e s e n t

i n t h e r a i n y m o n t h s . I n c o n t r a s t , t h e su n s t r e a m s t h r o u g h t h e

h o l e i n t h e c e i l i n g o f t h e G r e a t Room r a p i d l y d r y i n g an y m o i s t u r e

and w arm ing t h e a i r , p a r t i c u l a r l y a t t h e end w h e re t h e ch im n ey

F i g . 6 1 . H i s t o g r a m s sh o w in g i n d i c e s o f t h e r e l a t i v e a b u n ­

d a n c e o f E o n y c t e r i s s p e l a e a i n B a tu C av es and

t h e q u a n t i t y o f w a t e r p e r c o l a t i n g i n t o t h e c a v e s

e a c h m o n th „

The i n d i c e s a r e c a l c u l a t e d a s f o l l o w s :

A bundance i n d e x . Each w eek t h e b a t s w e re r e c o r d e d

a s " v e r y a b u n d a n t " , " a b u n d a n t " o r " l e s s a b u n d a n t "

( f o r d e f i n i t i o n s s e e p . 27 ) • T h e se c a t e g o r i e s

? /e re a l l o c a t e d n u m e r i c a l v a l u e s , 3 , 2 a n d 1 r e s - .

p e c t i v e l y . The sum o f t h e w e e k ly v a l u e s i n e a c h

c a l e n d e r month (A p p e n d ix 3) was d i v i d e d b y t h e

num ber o f w eeks i n t h e m onth t o g i v e a mean v a l u e ,

t h e a b u n d a n c e i n d e x .

W a te r i n d e x . T h i s w as c a l c u l a t e d t h e same way

b u t t h e f i g u r e s 1 , 2 an d 3 w e re a l l o c a t e d t o

c a t e g o r i e s " d r y " , " w e t" an d " v e r y w e t" r e s p e c t ­

i v e l y ( p . 2 ? and a p p e n d ix 3 ) .

N .B . Weeks f a l l i n g b e tw e e n two c a l e n d e r m o n th s

w e re i n c l u d e d i n b o t h m o n th s u n l e s s o n l y one o r

two d a y s w e re i n one o f t h e m o n th s and f i v e o r

s i x d a y s i n t h e o t h e r when t h e d a t a f o r t h a t

week was i n c l u d e d i n t h e l a t t e r m o n th .

A b u n d a n c e I n d e xo _* to tsj'Jn o tn O u i

usOJ

W a t e r I n d e xO _fc fo N>

o cn o u i o cn

142

i s s i t u a t e d . E . s p e l a e a r o o s t e d i n b o t h s i t u a t i o n s .

2 . i . A bundance i n t h e r o o s t s .

No a t t e m p t was made t o c o u n t t h e num ber o f

E . s p e l a e a p r e s e n t i n B a tu C aves b u t e a c h week a s u b j e c t i v e

a s s e s s m e n t o f t h e i r a b u n d a n c e was r e c o r d e d . The f u l l r e s u l t s

a r e i n c l u d e d i n A p p en d ix 3* They a r e a l s o su m m a r i s e d i n F i g . 61 .

T h e re was no d i s c e r n a b l e r e l a t i o n s h i p b e tw e e n t h e a b u n d a n c e o f

b a t s and e i t h e r . t h e r a i n f a l l o r t h e t im e o f y e a r ( F i g s . 4 an d 61 ) .

2 . i i . B e h a v io u r i n t h e r o o s t .

s p e l a e a hung from t h e c a v e c e i l i n g s i n g r o u p s

o f two t o s e v e r a l h u n d r e d i n d i v i d u a l s ; s i n g l e b a t s w e re o c c a s i o n a l l y

s e e n . The same p l a c e s on t h e ,r o o f w e re u s e d r e p e a t e d l y so t h a t

s m a l l h o l l o w s w e re w orn i n t o t h e l i m e s t o n e . T h e se h o l l o w s showed

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

o f t h e r o c k ’ s d i r t y s u r f a c e . I n B a tu C aves w h e re l a r g e a r e a s o f

c e i l i n g w e re u n u s e d o r o n l y i n t e r m i t t e n t l y u s e d ( i n d i c a t i n g t h a t

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

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

' l o c a t e d i n n a t u r a l h o l l o w s i n t h e c a v e r o o f . I n C a v e rn C p r e c i s e l y

t h e same s i t e s w e re a l l u s e d t h r o u g h o u t t h e 22 m o n th s t h e y w ere

o b s e r v e d a l t h o u g h t h e num ber o f b a t s a t e a c h s i t e f l u c t u a t e d .

When n um b ers w e re v e r y h i g h , two a d d i t i o n a l s i t e s i n C a v e rn B

w ere u s e d b u t t h e s e w e re a b a n d o n e d when nu m b ers d i m i n i s h e d a g a i n .

I n Gua S a n d in g w h e re l e s s r o o f s p a c e was a v a i l a b l e

t h e r e was a v e r y l a r g e g ro u p i n t h e c e n t r e o f t h e c a v e and many

s m a l l e r g r o u p s a r o u n d i t s p e r i p h e r y .

2 . i i i . A c t i v i t y i n t h e r o o s t .

D u r in g t h e day t h e r e was a c o n t i n o u s n o i s e i n t h e

c a v e s a s t h e b a t s c h a t t e r e d and s q u e a l e d w h i l e j o s t l i n g f o r s p a c e .

B a t s f r e q u e n t l y f l e w fro m one g ro u p t o a n o t h e r b u t w h e t h e r t h e y

moved v o l u n t a r i l y o r h ad b e e n d i s l o d g e d b y members o f t h e i r

o r i g i n a l g ro u p i s n o t know n. I n Gua S a n d in g (w h e re i t was p o s s i b l e

t o o b s e r v e them w i t h b i n o c u l a r s ) t h e a r r i v a l o f a new i n d i v i d u a l

i n a s m a l l g ro u p c a u s e d s e v e r a l s e c o n d s o f j o s t l i n g w h ic h u s u a l l y

r e s u l t e d i n t h e same o r a n o t h e r b a t b e i n g d i s l o d g e d a nd f l y i n g

. e l s e w h e r e . A ls o b a t s f r e q u e n t l y awoke an d b e g a n t o groom an d

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

s q u a b b l i n g o f t e n d i s l o d g e d one o r two o f th em .

2 . i v . R o o s t e x i t a n d e n t r y .

E o n y c t e r i s s p e l a e a b e g a n t o l e a v e t h e r o o s t j u s t

b e f o r e d u s k , i . e . b e tw e e n 1 8 .5 0 an d 19*00 h r s e a c h e v e n i n g . L i g h t

r a i n d i d n o t p r e v e n t them l e a v i n g . F o r a b o u t 90 m i n u t e s p r i o r

t o d e p a r t u r e t h e y becam e p r o g r e s s i v e l y more n o i s y an d f l e w b a c k

and f o r t h w i t h i n t h e c a v e s . The m a j o r i t y d e p a r t e d b y 2 0 . 0 0 h r s

b u t some l e f t a s l a t e a s 21 .0 0 h r s a f t e r w h ich t h e few t h a t

r e m a in e d i n t h e c a v e a p p a r e n t l y s t a y e d a l l n i g h t .

A t Gua S a n d in g a n e t was s e t a c r o s s one e n t r a n c e

a l l n i g h t on 16 N ovember 1 9 7 2 . From 1 8 .5 0 h r s a d u l t s a n d . s u b a d u l t s

o f b o t h s e x e s w e re c a u g h t l e a v i n g t h e c a v e . By 1 9 .2 0 h r s few b a t s

w e re b e i n g c a u g h t a n d t h e s e w e re m o s t l y s u b a d u l t s . A f t e r 1 9 .2 0

h r s o n ly s u b a d u l t s w e re c a u g h t ( e n t e r i n g and l e a v i n g t h e c a v e )

u n t i l 01 .30 h r s when a d u l t s b e g a n t o r e t u r n , some w i t h p o l l e n

on t h e i r b o d i e s . Two had so much p o l l e n ( a l l C e ib a p e n t a n d r a )

12+4

on them t h a t t h e y a p p e a r e d y e l l o w , n o t brown! The b a t s

c o n t i m i e d r e t u r n i n g u n t i l a f t e r 0 6 . 0 0 h r s . Many o f t h o s e t h a t

r e t u r n e d e a r l i e r r o o s t e d i n t h e c a v e b u t l a r g e n u m b ers m i l l e d

a ro u n d t h e e n t r a n c e from 0 1 . 3 0 h r s t o dawn.

2 . v . P o p u l a t i o n s t r u c t u r e i n B a tu C a v e s .

102+ E o n y c t e r i s s p e l a e a w ere c a u g h t i n B a tu C aves

d u r i n g t h i s s t u d y . 83 o f them w ere k i l l e d b e tw e e n 21 and 26

F e b r u a r y 1973 f o r a n a l y s i s o f p o l l e n i n t h e a l i m e n t a r y c a n a l .

D u r in g t h i s o p e r a t i o n many s u b a d u l t s w e re r e l e a s e d u n r e c o r d e d

b u t a l l a d u l t s w e re r e c o r d e d . D e s p i t e t h i s 52 (50%) o f t h e

r e c o r d e d b a t s w e re s u b a d u l t s . Of t h e 52 a d u l t s , 23 (2+2+.2%)

w e re m a le and 29 (55*3%) w e re f e m a l e . The d i f f e r e n c e was n o t

s i g n i f i c a n t ( p > 0 . 0 5 ) . T hus s u b a d u l t s o u tn u m b e re d a d u l t

f e m a le s b y n e a r l y 2 : 1 .

B eck a n d Lim (1 9 ? 3 ) r e p o r t e d c a t c h i n g 302+1

E* s p e l a e a i n B a tu C aves b e tw e e n March 1966 and F e b r u a r y 1 9 6 8 .

T h e se i n c l u d e d 973 a d u l t m a l e s , 1060 a d u l t f e m a l e s an d 1008

’• im m atu re b a t s o f b o t h s e x e s ” . ( T h e i r d e f i n i t i o n o f " i m m a tu r e ”

i n c l u d e d b a t s o f a l l a g e s from b i r t h t o t h e o n s e t o f s e x u a l

m a t u r i t y . ) A n a l y s i s o f t h e i r d a t a g i v e r a t i o s o f 2+7 , 9 % a d u l t

m a l e s t o 52 .1% a d u l t f e m a l e s . The d i f f e r e n c e w as n o t s t a t i s t i c ­

a l l y s i g n i f i c a n t (p > 0 . 0 5 *C0.06) and 76.1% a d u l t s t o 23.9%

s u b a d u l t s ( e x c l u d i n g 371 s u c k l i n g young a t t a c h e d t o t h e i r

m o t h e r s : I f t h e s e a r e i n c l u d e d t h e r a t i o i s 66 .9% a d u l t t o

33 .1% s u b a d u l t . ) T h e re w e re a p p r o x i m a t e l y e q u a l n u m b ers o f a

a d u l t f e m a l e s an d y o u n g .

2 . v i . S e g r e g a t i o n o f s e x and a g e c l a s s e s i n t h e r o o s t .

I t i s p r o b a b l e t h a t t h e r e i s some d e g r e e o f

s e g r e g a t i o n o f a g e and s e x c l a s s e s i n t h e r o o s t . I n Gua S a n d in g

U 5

one ch im n ey n e a r t h e e n t r a n c e a lw a y s c o n t a i n e d a g r o u p o f 100

t o 200 s u b a d u l t s . The ch im n ey was n o t d a r k and t h e b a t s c o u l d b e

o b s e r v e d t h r o u g h b i n o c u l a r s d u r i n g t h e d a y . They w e re e a s i l y

i d e n t i f i e d a s s u b a d u l t s by t h e i r s m a l l s i z e (when f i r s t n o t i c e d

t h e y w ere t h o u g h t t o b e P e n t h e t o r l u c a s i ( D o b s o n ) , a much s m a l l e r

o» •»*«./•*✓*» *100 TP fin rw> o a/* r» n c H Art ooirovi uror>o a on rrVi 4- 4 y>

a m i s t n e t s e t a t t h e b a s e o f t h e ch im n ey a s t h e y l e f t a t d u s k .

A l l w ere s u b a d u l t s ( w e i g h t s b e tw e e n 25 and 37 g) • I n B a tu C a v e s ,

e v id e n c e from g uan o a n a l y s i s s u g g e s t s t h a t one g r o u p i n C av e rn

C from w h ic h guan o was c o l l e c t e d c o n t a i n e d many s u b a d u l t s ^

A d u l t s w ere n o t s h o t f rom t h e r o o f o f B a tu C aves

b e c a u s e o f t h e p o s s i b l e e f f e c t s d i s t u r b a n c e o f t h i s k i n d m ig h t

h av e h ad on t h e c o m p o s i t i o n o f g r o u p s r o o s t i n g a b o v e t h e guano!

c o l l e c t i o n t r a y s an d c o n s e q u e n t e f f e c t s on t h e gu an o c o l l e c t e d

b u t t h e IMR r e g i s t e r r e c o r d s f o u r s e r i e s o f E . s p e l a e a t o t a l l i n g

235 b a t s t h a t w e re s h o t n e a r t h e e n t r a n c e i n s i d e B a t u C aves

b e tw e e n 1951 an d 1955* They w e re ex am in ed by D r . J . L . H a r r i s o n .

The d a t a i n T a b le 31 w e re d e r i v e d from t h e s e e n t r i e s i n t h e r e g i s t e r .

They show two p o i n t s o f i n t e r e s t : F i r s t l y , i n t h e two l a r g e r

s e r i e s (3 an d /+) t h e a d u l t m a le : t o a d u l t f e m a le r a t i o d i f f e r e d

s i g n i f i c a n t l y from t h e e x p e c t e d 1:1 r a t i o ; i n s e r i e s 3 t h e r e w ere

more f e m a le s t h a n m a le s an d i n s e r i e s ^ v i c e v e r s a ( p < 0 .00 1

f o r b o t h ) . S e c o n d l y , t h e r e w e re v e r y few s u b a d u l t s i n t h e s e

s a m p l e s .

T h e re i s s t r o n g e v i d e n c e t h a t many s u b a d u l t s

w e re r e c o r d e d a s a d u l t : e . g . Many " a d u l t m a l e s " w e re r e c o r d e d

a s h a v in g t e s t e s " n o t d e s c e n d e d " . T h e se w ere p r e s u m a b ly s u b a d u l t s

w i t h v e r y s m a l l t e s t e s b e c a u s e t h e t e s t e s o f a d u l t an d s u b a d u l t

Table 31 .The t a b l e show the r a t i o o f a d u l t males t o a d u l t

females and th e p e r c e n t a g e o f s u b a d u l t s i n fo u r samples o f E o n y c te r i s s r e l a e a s h o t from th e r o o f o f Batu Caves between 1951 and 1955* The b a s i c d a t a was e x t r a c t e d from th e r e g i s t e r o f th e I n s t i t u t e o f M edica l R e s e a rc h , Kuala Lumpur. F ig u re s i n p a r e n t h e s i s a r e p e r c e n t a g e s .

Group Date Totalshot

Totaladult

Adultmales

Adultfemales P

Totalyoung

1 1 3 Nov. 1 51 29 2 8 (9 6 .6 ) 1 0 (35 .7 ) 1 8 (6 4 .3 ) >0.1 1 ( 3 .4 )2 27 Feb. »52 30 2 7 (9 0 .0 ) 12(44./+) 1 5 (5 5 .6 ) >0.1 3 ( 1 0 .0 )3 25 May ’ 53 115 108(93 .9 ) 3 2 (7 9 .6 ) 7 6 (7 0 .4 ) <0.001 7 ( 6 .1 )4 19 J u l y *55 61 5 9 (9 6 .7 ) 4 6 (7 8 .0 ) 1 3 (2 2 .0 ) <0.001 2 ( 3 .3 )

Total 235 222(9/1.5) 100(45 .0 ) 1 1 2(55 .0 ) >0.1 1 3 (5 .5 )

Note p i s t h e p r o b a b i l i t y t h a t each sample was s h o t from groups composed o f e q u a l numbers o f a d u l t males and f e m a le s .

Table 32 .The t a b l e r e c o r d s th e same d a t a as Table 31 f o r groups

3 and 4 i n t h a t t a b l e . However, males i n which th e t e s t e s were n o t descended and fem ales w eigh ing l e s s than 4 0 .0 g which we r e r e c o rd e d as a d u l t i n th e I .M .R . r e g i s t e r and i n Tab le 29 have been r e d e s i g n a t e d s u b a d u l t i n t h i s t a b l e .

Group Total Total Adult Adult Totalshot Adult males females P young3 115 91 (7 9 .1 ) 2 5 (2 7 .9 ) 6 6 (72 .5 ) <0 .0 0 1 2 4 (2 0 .9 )4 61 3 9 (6 3 .9 ) 2 7 (6 9 .2 ) 1 2 (30 .8 ) < 0 .02 2 2 (3 6 .1 )

146

E o n y c t e r i s s p e l a e a a r e c o n t a i n e d i n a s c r o tu m an d t h e r e f o r e a r e

d e s c e n d e d . ( I n t h i s s t u d y a d u l t s a r e d e f i n e d a s b e i n g s e x u a l l y

m a tu r e an d a d u l t m a l e s a r e d i s t i n g u i s h e d from s u b a d u l t s by t h e i r

l a r g e t e s t e s , p . 21 ) . E . g . 2 , " A d u l t f e m a le " No. 4 1 58 9 w e ig h e d

1 8 g b u t t h e mean w e i g h t o f n o n - p a r o u s a d u l t f e m a l e s r e c o r d e d

d u r i n g t h i s s tud .y w as 4 9*97 S> t h e minimum w e i g h t was 4 0 . 0 g .

I n T a b le 32 t h e d a t a f o r g r o u p s 3 and 4 o f

T a b le 31 h a v e b e e n - r e v i s e d so a s t o i n c l u d e a s a d u l t s o n ly t h o s e

m a le s w i t h " d e s c e n d e d t e s t e s " an d t h o s e f e m a le s w e i g h i n g o v e r

4-0.0 g . The r e m a i n d e r h a v e b e e n r e d e s i g n a t e d a s y o u n g . ( I n s u f f i c i e n t

d a t a w e re a v a i l a b l e t o p e r m i t r e v i s i o n s o f g r o u p s 1 an d 2 . ) The

d i f f e r e n c e s i n t h e r a t i o s o f a d u l t m a l e s t o a d u l t f e m a l e s a r e

s t i l l s i g n i f i c a n t , i . e . t h e r e a r e s i g n i f i c a n t l y more f e m a l e s ' t h a n

m a le s i n g ro u p 3 (p < 0 .001 ) an d v i c e v e r s a i n g ro u p 4 (p < 0 ,02 .)

b u t t h e r a t i o o f s u b a d u l t s t o a d u l t s a p p r o a c h e s t h e e x p e c t e d r a t i o

o f 3 3 .1 % : 66.9% r e c o r d e d e a r l i e r .

The m o s t p l a u s i b l e e x p l a n a t i o n o f t h i s d a t a i s

t h a t a d u l t m a le s an d a d u l t f e m a l e s w e re s e g r e g a t e d i n t o s e p a r a t e

r o o s t i n g g r o u p s an d t h a t s e r i e s 3 was m o s t l y c o l l e c t e d from f e m a le

g r o u p s w h i l e s e r i e s 4 was m o s t l y c o l l e c t e d from m ale g r o u p s .

3 . Guano c o l l e c t e d from B a tu C a v e s .

3 - i . Guano c o l l e c t i o n

Guano was c o l l e c t e d from t h e f o u r t r a y s i n C av ern

C, B a tu C a v e s , e a c h week from 19 F e b r u a r y 1972 t o 28 J a n u a r y 1974

e x c e p t s i x t i m e s when t h e i n t e r v a l s b e tw e e n c o l l e c t i o n s w ere ;

f i v e d a y s ( 1 ) , s i x d a y s ( 1 ) , t e n d a y s (2 ) and two w eeks ( 2 ) .

99 c o l l e c t i o n s w e re made from e a c h t r a y b u t some w e re d i s c a r d e d

b e c a u s e t h e y w e re c o n t a m i n a t e d ( p . 2 7 ) o r b e c a u s e t h e r e was so

l i t t l e gu ano on them t h a t r e a l i s t i c p o l l e n c o u n t s c o u l d n o t b e

m ade. The num ber o f p o l l e n c o u n t s made from e a c h t r a y w e re

T ra y I

I I

I I I

IV

90

85

96

The f r e s h w e i g h t o f e a c h s a m p le , t h e p e r c e n t a g e

. d r y w e i g h t o f a s u b s a m p le and t h e e s t i m a t e d t o t a l d r y w e i g h t o f

e a c h s a m p le a r e g i v e n i n A p p en d ix 3 .

3 « i i . W e ig h t o f g uan o c o l l e c t e d .

The f r e s h w e i g h t o f gu ano on e a c h t r a y f l u c t u a t e d

e n o r m o u s ly , s o m e t im e s from week t o week a s a r e s u l t o f w a t e r

d r i p p i n g from t h e r o o f o n to t h e p o l y t h e n e s h e e t s an d d i l u t i n g

i t . T h e re w e re c o n t i n o u s d r i p s f a l l i n g o n to T ra y 1 , 2 and 4 . The

volum e o f w a t e r d r i p p i n g o n to e a c h was r e l a t e d t o t h e r a i n f a l l

o v e r t h e B a tu C aves h i l l . A t t i m e s o f h e a v y and f r e q u e n t s t o r m s ,

w a t e r a l s o d r i p p e d o n to T ra y 3* The e f f e c t s o f w a t e r d r i p on t h e

f r e s h w e i g h t i s i l l u s t r a t e d b y t h e f o l l o w i n g f i g u r e s from T ra y 1 .

Sam ple F r e s h y e i g h t % d r y w e i g h t

number. .. i n gms. o f s u b s a m p le

117

121

125

345

1146

1672

14.9%

4.4%

3.0%

E s t i m a t e d d r y R a i n f a l l

w t . i n gms. f o r w eek .

51 .4

5 0 ,4

5 0 .2

0 . 4 3 ”

1 .2 9 "

1 .3 3 "

N o te t h a t t h e r e was c o n s i d e r a b l y more w a t e r i n s a m p le 1 25 t h a n

121 a l t h o u g h t h e r a i n f a l l was a b o u t e q u a l f o r t h e two w e e k s .

T h i s phenom enon was f r e q u e n t l y o b s e r v e d . The p r o b a b l e e x p l a n a t i o n

i s t h a t t h e h i l l a c t s a s a r e s e r v o i r and when r a i n f a l l s a f t e r

a d r y s p e l l some w a t e r i s i n i t i a l l y a b s o r b e d by t h e s o i l o r ,

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

m o s t o f t h e w a t e r d r i p d i r e c t l y i n t o t h e c a v e . Thus a f t e r a d r y

s p e l l t h e r e i s a t i m e l a g b e tw e e n h e a v y r a i n a n d maximum flo w

o f w a t e r i n t o t h e c a v e .

D u r in g d r y p e r i o d s t h e f r e s h w e i g h t o f g uano on

T ra y 3 was a f f e c t e d by e v a p o r a t i o n . The p e r c e n t a g e d r y w e i g h t

r o s e t o a s much a s 21 .1% b e tw e e n 26 and 31 J u l y 1 9 7 2 . (The % d r y

w e i g h t o f f r e s h l y d e f a e c a t e d guan o was b e t ween 1 1 . 0 and 14.0%

i n Gua S a n d in g w h e re f r e s h f a e c a l p e l l e t s w e re c o l l e c t e d . T h i s

r a n g e i s p r o b a b l y t h e same a s t h a t o f guano from B a tu C a v e s . )

N e v e r t h e l e s s , t h e r e w ere f l u c t u a t i o n s i n t h e

e s t i m a t e d d r y w e i g h t o f s a m p l e s . I n a c c u r a c y i n h e r e n t i n t h e .

m e th o d s o f c o l l e c t i n g t h e g uano a n d e s t i m a t i n g t h e d r y w e i g h t

u n d o u b t e d l y a c c o u n t f o r some o f t h i s . The e f f e c t s w o u ld be

g r e a t e s t w h e re t h e % d r y w e i g h t was h i g h e s t b u t e v e n h e r e e r r o r s

w e re t h o u g h t u n l i k e l y t o e x c e e d 10% and w e re p r o b a b l y b e lo w 5%

u s u a l l y .

The m a j o r c a u s e o f t h e f l u c t u a t i o n s was t h o u g h t

t o b e v a r i a t i o n i n t h e num ber o f b a t s r o o s t i n g a b o v e t h e t r a y s .

H o w ever , d r y w e i g h t c a n n o t b e u s e d a s a m e a s u re o f t h e num ber

o f b a t s i n t h e g ro u p ab o v e t h e t r a y o r i n t h e c a v e b e c a u s e once

t h e r o o s t i n g s i t e s on t h e r o o f a r e a i m m e d i a t e l y ab o v e t h e t r a y

a r e s a t u r a t e d t h e r e c an b e no f u r t h e r i n c r e a s e i n t h e num ber o f

b a t s o v e r t h e t r a y an d c o n s e q u e n t l y i n t h e g uan o f a l l i n g on t h e

149

t r a y b u t more b a t s c o u l d j o i n t h e g r o u p , r o o s t i n g on i t s p e r i p h e r y .

C o n v e r s e l y , b a t s r o o s t i n g on t h e p e r i p h e r y o f t h e g r o u p c o u ld

move e l s e w h e r e w i t h o u t a f f e c t i n g t h e q u a n t i t y o f g uan o c o l l e c t e d .

T h e re w ere many l o c a l movem ents w i t h i n t h e c a v e made b y b a t s

l e a v i n g one g r o u p t o j o i n a n o t h e r T h e r e a s o n s f o r t h e s e m ovem ents

a r e n o t known b u t t h e y may h a v e b e e n a s s o c i a t e d w i t h t h e q u a n t i t y

o f w a t e r d r i p p i n g f ro m t h e r o o f , e . g . e x c e s s i v e w a t e r d r i p p i n g

from a r o o s t o r ( a s was s e e n i n C a v e rn A) e v en r u n n i n g o v e r t h e

r o c k on w h ic h t h e b a t s ha d b e e n h a n g in g f ro m , m i g h t f o r c e some

o r a l l t h e b a t s i n t h a t g ro u p t o e m i g r a t e . T h e se movem ents w i t h i n

t h e c a v e c o u l d h a v e a f f e c t e d t h e q u a n t i t y o f g u an o f a l l i n g on t h e

t r a y s b u t i m m i g r a t i o n t o and e m i g r a t i o n from t h e c a v e may h a v e

b e e n i m p o r t a n t to o .

T h e re was so m e t im e s a n i n c r e a s e i n t h e d r y w e i g h t

o f g uano f o l l o w i n g a p e r i o d o f h e a v y r a i n . T h i s may h av e b e e n

c a u s e d b y an i n f l u x o f b a t s , t h a t h a d b e e n r o o s t i n g o u t s i d e t h e

c a v e s , s e e k i n g s h e l t e r .

4 . Food s o u r c e s o f E . s p e l a e a i n P e n i n s u l a r M a l a y s i a .

A l l t h e p l a n t s t h a t E . s p e l a e a was known t o f e e d

from i n P e n i n s u l a r M a l a y s i a a r e l i s t e d i n T a b le 33 • They i n c l u d e

tfro f a m i l i e s o f m o n o c o ty le d o n s an d e l e v e n o f d i c o t y l e d o n s . I n

a d d i t i o n , many s p e c i e s o f e x o t i c c h i r o p t e r o p h i l o u s p l a n t s a r e

so m e t im e s grown i n P e n i n s u l a r M a l a y s i a a s o r n a m e n t a l p l a n t s ,

e . g . Agave s p p . ( A m a r y l l i d a c e a e ) , A d a n s o n ia d l g l t a t a L. (B om b acaceae )

an d C r e s c e n t i a c u j e t e L . ( B i g n o n i a c e a e ) . T h i s l i s t i s n o t e x h a u s t i v e .

E . s p e l a e a may f e e d from them when a v a i l a b l e , e . g . van d e r P i j l

Table 33 .The table records all the plants that Eonycteris

spelaea was shown to feed from in Peninsular Malaysia.

Family Species Bats seen on flower

Pollen identi fied

Musaceae Musa malaccensis + 1M. truncata + ►To genusM. spp. cultivated + No pollen

palmae Cocos nucifera To speciesArenga spp. To genus

Anacardiaceae Mangifera spp. To genusBignoniaceae Oroxylum indicum + To species

Pajanetia longifolia *Bombacaceae Bombax valetonii + To species

Ceiba pentandra + To speciesDurio zibethinus + •To genusD. graveolens +

Compositae ? To familyLecythidaceae Barringtonia spp. To genusLeguminosae Parkia .iavanica +

P. speciosa + ►To genusP. singularis

Moraceae Artocarpus spp. To genusMyrtaceae Eugenia malaccensis + To genusRhizophoraceae Rhizophora spp. To genusSapotaceae ? To familySonneratiaceae Duabanga grandiflora + To species

Sonneratia alba +S. caseolaris + ► To speciesS . ovata +

Claw marks on fallen corollas similar to those of E. srelaea on Oroxylum indicum but possibly made by Macroglossus sobrinus.

Ii

• 1 50

(1 9 3 6 ) r e c o r d e d i t f e e d i n g from Agave a n g n s t i f o l i a Haw. and

A d a n s o n ia d i g i t a t a i n I n d o n e s i a .

5 . P o l l e n t y p e s fo u n d i n g uano from B a tu C a v e s .

Not a l l the plants lis ted in Table 33 have pollen

which is universally id e n tif iab le . Those that do are the three

species of Sonneratia. Duabanga grandiflora (and D. moluccana

in Borneo) and Cocos nucifera .

H ow ev er , Bombax v a l e t o n i i and Cei b a p e n t a n d r a

(w h ic h h a v e p o l l e n i d e n t i f i a b l e t o t h e g e n u s l e v e l ) w e re t h e

o n l y r e p r e s e n t a t i v e s o f t h e i r g e n e r a i n S e l a n g o r an d Oro x y lu m

in d i c u m was t h e o n l y known c h i r o p t e r o p h i l o u s s p e c i e s o f B i g n o n i a c e a e

r e c o r d e d i n S e l a n g o r . The p o l l e n o f t h e s e s p e c i e s f o u n d i n g uano

from B a tu C aves c o u l d t h e r e f o r e b e i d e n t i f i e d t o s p e c i e s l e v e l .

The p o l l e n o f P a r k i a j a v a n i c a , P . s i n g u l a r i s and

£♦ s p e c i o s a c o u l d n o t b e d i s t i n g u i s h e d from one a n o t h e r . The p o l l e n

£ • s u m a t r a n a was n o t e x a m in e d . H ow ever , P . s i n g u l a r i s and P .

s u m a t r a n a a r e r a r e a nd P . . j a v a n i c a an d P . s p e c i o s a d i d n o t f l o w e r

s i m u l t a n e o u s l y . so m o s t o f t h e P a r k i a s p p . p o l l e n i n t h e guano

ca n b e a t t r i b u t e d t o e i t h e r P . j a v a n i c a o r P . s p e c i o s a d e p e n d in g

on w h ic h was f l o w e r i n g when t h e gu ano was d e p o s i t e d . N e v e r t h e l e s s ,

P a r k i a s p p . p o l l e n h a s b e e n t r e a t e d a s a s i n g l e p o l l e n t y p e i n

t h e f o l l o w i n g a n a l y s i s ,

The p o l l e n o f a l l o t h e r p l a n t s l i s t e d i n T a b le 33

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

C o m p o s i t a e , M o ra c e a e an d S a p o t a c e a e . M o rac eae p o l l e n was f r e q u e n t l y

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

s p p ; and h a s a l l b e e n a t t r i b u t e d t o A r t o c a r p u s .

1 51

The p o l l e n o f Musa s p p . d o e s n o t p r e s e r v e i n

guano a l t h o u g h t h e e x i n e p a s s e s t h r o u g h t h e g u t o f E . s p e l a e a

a p p a r e n t l y undam aged a s shown b y t h e p r e s e n c e o f w h o le g r a i n s

i n t h e f r e s h f a e c e s o f b a t s c a u g h t w h i l e f o r a g i n g . I t was v e r y

se ld o m fo u n d i n t h e B a tu C aves guano s a m p le s b u t i t was p r e s e n t

i n t h e a l i m e n t a r y c a n a l s o f some E . s p e l a e a c a u g h t i n B a tu C a v e s .

H ow ever , i n t h e gu ano t h e r e w e re f r e q u e n t l y many s m a l l f r a g m e n t s

o f t i s s u e w h ic h s t a i n e d i n b a s i c f u s c h i n t o a s i m i l a r e x t e n t a s

t h e e x i n e s o f Musa s p p . p o l l e n g r a i n s and w h ich w e re t h o u g h t t o

b e t h e r e m a i n s o f r u p t u r e d Musa s p p . p o l l e n g r a i n s . T h ese f r a g m e n t s

w ere e x c lu d e d from a l l c o u n t s o f p o l l e n i n t h e g u a n o .

T h e re w e re a n o t h e r s i x p o l l e n t y p e s i n t h e g uano

w h ic h w e re n o t i d e n t i f i e d e v en t o f a m i l y l e v e l . T h e se w e re c o d e -

named T3, T/f, T 6 , T 10 , T12 and T13» T3 ( F i g . 37 ) was v e r y s i m i l a r

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

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

c o n s i d e r a b l y l a r g e r t h a n an y known n o r m a l o r m u t a n t fo rm s o f

S o n n e r a t i a . D r . J . M u l l e r ( p a l y n o l o g i s t a t t h e R e j k s h e r b a r i u m ,

L e id e n ) who h a s h a d c o n s i d e r a b l e e x p e r i e n c e w i t h t h e p o l l e n o f

S o n n e r a t i a ( M u l l e r , 19 6 9 ; M u l l e r an d H o u -L iu , 1966) e x p r e s s e d

t h e v iew ( i n l i t . ) t h a t i t was " a S o n n e r a t i a p o l l e n t y p e w h ic h

I h a v e n e v e r s e e n b e f o r e . I am n o t s u r e w h ich s p e c i e s p r o d u c e d

i t . I t may h a v e b e e n S . a l b a , S . o v a t a o r some h y b r i d . " The

f o l l o w i n g ' f a c t s . c o n t r a d i c t t h i s v ie w . No g r a i n s o f T3 w ere fo u n d

on t h e b o d i e s o r i n t h e f a e c e s o f M a c r o g lo s s u s m in im u s w h ic h

r e l i e d v e r y h e a v i l y on S o n n e r a t i a s p p . f o r f o o d . O n ly one E o n y c t e r i s

s p e l a e a was c a u g h t w i t h p o l l e n o f t y p e T3 on i t s b o d y ( i n d i c a t i n g

152

r e c e n t f e e d i n g from t h e p l a n t from w h ich i t c a m e ) . T h a t was a t

t h e F . S . C . , U lu Gombak, i . e . i n s e c o n d a r y f o r e s t 40 km i n l a n d .

P e a k s i n t h e q u a n t i t y o f T3 p o l l e n i n t h e guano o f B a tu Gaves

d i d n o t c o i n c i d e w i t h p e a k s i n t h e p o l l e n o f a n y s p e c i e s o f

S o n n e r a t i a . •

I t s i d e n t i t y r e m a i n s p r o b l e m a t i c a l b u t t h e

p o s s i b i l i t y o f i t b e i n g a m u t a n t fo rm o f D uabanga g r a n d i f l o r a

( a l s o S o n n e r a t i a c e a e ) s h o u l d be c o n s i d e r e d .

Amongst t h e r e m a i n i n g u n i d e n t i f i e d t y p e s o n ly

T10 was o f an y c o n s e q u e n c e , a c c o u n t i n g f o r b e tw e e n 0 .24% and

0 .38% o f a l l t h e p o l l e n on e a c h gu an o t r a y . The r e s t e a c h a c c o u n t e d

f o r 0 .03% o r l e s s o f a l l t h e p o l l e n on e a c h t r a y . The p o s s i b i l i t y

t h a t t h e y w e re i n t r o d u c e d t o t h e s a m p le s by c o n t a m i n a t i o n c a n n o t

b e r u l e d o u t .

6 . A n a l y s i s o f p o l l e n i n t h e guano from B a tu C a v e s .

6 i . P r e s e n t a t i o n a n d l i m i t a t i o n o f t h e d a t a .

The d a t a i s p r e s e n t e d i n t e r m s o f t h e n u m e r i c a l

p r o p o r t i o n o f e a c h p o l l e n t y p e . S i n c e t h e p o l l e n g r a i n s o f '

d i f f e r e n t s p e c i e s v a r y c o n s i d e r a b l y i n s i z e , t h e d a t a do n o t

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

n o t h i n g i s known o f t h e i r n u t r i t i v e v a l u e ( e i t h e r o f t h e i r

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

t h e i r n u t r i v t i v e i m p o r t a n c e t o t h e b a t s .

The p u r p o s e o f t h e a n a l y s i s was t o i d e n t i f y t h e

s o u r c e s o f p o l l e n an d t h e u s e made o f them by th& b a t s . T h i s

i n f o r m a t i o n can b e d e r i v e d from i d e n t i f y i n g t h e p o l l e n p r e s e n t

i n t h e guano a nd from c h a n g e s i n t h e n u m e r i c a l p r o p o r t i o n s o f

153

the pollen types from week to week. A study of the nutritive

value of the different pollen types must await a study of the

nutrition of the bats concerned. That was not the purpose of

the study. However, in order that the difference in the appearance

of the data had i t been presented in terms of the volumetric

proportions of the pollen con be appreciated volume indices

of all the more common pollen types are given in Table .The

volume indices relate the volume of each pollen type to the

volume of Artocarpus spp. which was arbitrarily allocated a

volume index of one as i t was the smallest common pollen type.

Clearly Artocarpus spp. and Duabanga grandiflora

would account for a very much smaller proportion of the volume

of pollen than they do of the numerical proportion, and Durio

spp. and Parkia spp. when present would account for a very much

larger proportion of the volume of the pollen present than they

do of the numerical proportion. (Parkia spp. polyads were counted

as representing one unit not as 16 or 32 individual grains.) The

difference between the numerical and volumetric proportions would

be more pronounced when the large grains formed a small numerical

proportion of a sample than when they formed a large numerical

proportion of the grains present. The following hypothetical

example illustrates this point.

Volumetric

index no.

Numerical

proportion

Volumetric

proportion

Parkia Artocarpus P a r k i a Artocarpus P a r k i a Artocarpus

138.25 1 10% 90% 93.9 6.1

138.25 1 90% 10% 99.9 0.1

T a b le 3k •

The t a b l e show s t h e d i m e n s i o n s o f some p o l l e n t y p e s '

t h a t o c c u r i n t h e 'guano o f E o n y c t e r i s s p e l a e a , a s w e l l a s

t h e a p p r o x i m a t e vo lum e o f e a c h r e l a t i v e t o A r t o c a r p u s s p p ,

p o l l e n ( i . e . t h e i r volum e i n d e x ) .

N o te 1 b - a = l e n g t h from t h e b a s e t o t h e o p p o s i t e

a n g l e ( o f t r i a n g u l a r g r a i n s ) ,

d = d i a m e t e r ( o f s p h e r i c a l o r n e a r l y s p h e r i c a l

g r a i n s ) .

p x eq = p o l a r l e n g t h x e q u a t o r i a l d i a m e t e r ( f o r

a l l o t h e r g r a i n s ) .

N o te 2 M e a s u re m e n ts w e re made from f r e s h p o l l e n c o l l e c t e d

from f l o w e r s and m oun ted i n an a q u e o u s s o l u t i o n

o f p o l y v i n a l a l c o h o l (PVA) e x c e p t ' f o r s p e c i e s

m arked * ; m e a s u re m e n ts f o r t h e s e s p e c i e s w ere

from g r a i n s fo u n d i n g uan o s a m p l e s .

N o te 3 C a l c u l a t i o n o f t h e volum e i n d e x , p l a s t i c e n e m o d e ls

o f t h e p o l l e n g r a i n s w e re made t o s c a l e a c c o r d i n g

t o t h e d im e n s i o n s r e c o r d e d h e r e . (No m e a s u re m e n ts

o f t h e p o l a r l e n g t h o f t r i a n g u l a r g r a i n s w e re

r e c o r d e d . The p o l a r l e n g t h s o f t h e m o d e ls w ere

t h u s a p p r o x i m a t i o n s b a s e d on t h e k n o w le d g e o f t h e

s h a p e o f t h e g r a i n s . ) The m o d e ls w ere v /e ig hed on a

beam b a l a n c e t o t h e n e a r e s t 0 .1 g . The volum e

i n d e x r a t i o s w e re c a l c u l a t e d from t h e w e i g h t r a t i o s

o f t h e m o d e l s .

N o te k : - P a r k i a s p p . p o l y a d s w e re c o u n t e d a s s i n g l e u n i t s .

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

n o t s i n g l e p o l l e n g r a i n s .

Species 25

Measurem

entstak

en

Mean

size

(u)

Wt.

of m

odel (gram

s)

Volum

e

index

Musa truncata 201 d 73.0 7.9 19.75

Cocos nucifera 20 p x eq 97.8 X 76.1 13.6 34.0

Arenga spp.*

20 p x eq 59.8 X 46.2 2.3 5.75

Mangifera spp.«

10 d 38.0 1 .4 3.5

Oroxylum indicum 20 p x eq 130.0 X 125.0 44.0 110.0

Bombax valetonii 20 b - a 97.8 11 .0 27.5

Ceiba pentandra 20 p x eq 42.3 X 97.8 15.5 38.75

Durio zibethinus 2 0 d 130.4 44.4 111 . 0

Parkia speciosa 20 polyad 1 * 2 * 8 X 1 52.855.3 138.25

X 108.7

Eugenia malaccensis 20 b - a 27 .2 0.3 0.75

Duabanga grandiflora 20 p x eq 32.6 X 24.4 0.4 1 . 0

Sonneratia alba 20 p x eq 87.0 X 54.3 7.2 18 .0

S. ovata 20 ip x eq 77.4 X 52 .0 5.5 13.75

S. caseolaris 20 p x eq 76.1 X 54.3 5.4 13.5

Artocarpus elasticus 2 0 j p x eq 2 0 . 0 X 27 .2 0.4 1 . 0

T3*

2 0 p x eq 141 .3 X 11 9.6 36.8 92 .0

T10*

3 b - a 59.8 3.5 7.75

1 5k

The d a t a d o e s n o t r e f l e c t t h e i m p o r t a n c e o f t h e

p l a n t s a s a s o u r c e o f n e c t a r . , e , g . Oroxylum i n d i c u m p r o d u c e s

r e l a t i v e l y l i t t l e p o l l e n w h ic h i s d e p o s i t e d a c c u r a t e l y on t h e

h e a d o f a b a t a s i t p r o b e s f o r n e c t a r b u t P a r k i a s p p . o r S o n n e r a t i a

s p p . d e p o s i t a b u n d a n t p o l l e n on a b a t * s h e a d and b o d y w h i l e

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

b u t l i t t l e o r rio n e c t a r . Musa s p p . a r e i m p o r t a n t s o u r c e s o f n e c t a r

b u t a r e h a r d l y r e p r e s e n t e d by p o l l e n i n t h e gu ano s a m p le s b e c a u s e

i t s p o l l e n b r e a k s down e a s i l y .

The d a t a d o e s show t h e s p e c i e s ( e x c e p t Musa s p p . )

from w h ich E . s p e l a e a o b t a i n e d p o l l e n , t h e d a t e s on w h ic h t h e y

o b t a i n e d p o l l e n from th e m , t h e i m p o r t a n t p o l l e n p r o d u c i n g p l a n t s

( b u t , a s e x p l a i n e d a b o v e , n o t t h e r e l a t i v e n u t r i t i v e i m p o r t a n c e

o f t h o s e s p e c i e s ) and i t i n d i c a t e s i n some i n s t a n c e s p r e f e r e n c e s

shown by t h e b a t s f o r d i f f e r e n t s p e c i e s w h ic h w ere a v a i l a b l e

s i m u l t a n e o u s l y I t a l s o shows an y d i f f e r e n c e s i n t h e p o l l e n d i e t s

o f t h e f o u r r o o s t i n g g r o u p s from w h ic h g uano was c o l l e c t e d .

The d a t a h a s b e e n p r e s e n t e d i n t h r e e s e c t i o n s .

1 . T a b le 35 g i v e s t h e mean n u m e r i c a l p e r c e n t a g e o f e a c h p o l l e n

t y p e on e a c h t a b l e .

2 . The w e e k ly v a l u e s o f t h e n u m e r i c a l p r o p o r t i o n o f e a c h o f t h e

f o l l o w i n g p o l l e n t y p e s i s p l o t t e d o n to k i t e d i a g r a m s ( F i g s . 62

t o 65 ) ’• S o n n e r a t i a a l b a ( i n c l u d i n g S . o v a t a ) , S . c a s e o l a r i s ,

D uabanga g r a n d i f l o r a , P a r k i a ( s p e c i e s c o m b i n e d ) , A r t o c a r n u s

s PP*j E u g e n ia s p p . , D u r io s p p . , Bombax v a l e t o n i i , C e ib a

p e n t a n d r a , T3, T8 ( S a p o t a c e a e ) and Cocos n u c i f e r a . O th e r

p o l l e n t y p e s v/ere r a r e i n t h e s a m p l e s . The f l o w e r i n g s e a s o n

Table 35 .The t a b l e shows th e mean p e r c e n ta g e o f each p o l l e n

type found i n each o f fou r p o l l e n c o l l e c t i n g t r a y s i n Cavern C o f Batu Caves, S e l a n g o r . The guano was from E o n y c te r i s s p e l a e a

Family SpeciesGuano Tray No.

1 2 3 4

Musaceae Musa spp. - - <0.01 0.01 <£>•01

Palmae Cocos nucifera 0.48 1 .50 1 -95 1 .54 f .S?Arenga spp. <0.01 <0.01 <0 .01 <0.01 <"001

Anacardiaceae Mangifera spp. 0.13 0 .3 0 0.10 0.14 0 -1?

Bignoniaceae Oroxylum indicum 0.06 0.05 0.12 0.10 0 -0?

Bombacaceae Bombax valetonii 0.46 0.94 0.52 0.86 C? 6

Ceiba rentandra 0.65 1 .49 1 .61 2.70 I-Durio spp. (1) 2.67 3.07 3.46 4.41 3-40

(2) - <0.01 <0.01 - < .0 0 1

Compositae ? 0.02 0.03 0.02 0.01 0 olLecyt-hidaceae Earringtonia spp. <0.01 0.01 <0.01 <0.01 Cfc.QILeguminosae Parkia spp. 11 .38 12.96 10.86 9.25 tl-uMoraceae Artocarnus spp. 1 7.80 29.73 28.93 34.77Myrtaceae Eugenia spp. 5.47 3.73 3-69 4.32 A *Rhizophoraceae Rhizophora spp. 0.01 0.01 0.01 0.01 0 *Sapotaceae T8 1 .84 1 .76 0.66 1 .61 I - 4 VSonneratiaceae Duabanga grandiflora 50.13 26.84 24.91 18.36 ’ t>6

Sonneratia alba (incl. S. ovata)

6.65 14.04 13.75 12.42 11- 71

S. caseolaris 1 .40 3.84 9.26 8.11? T3 0.65 0.89 0.77 0.74 0'%? T4 - 0.02 0.01 0.02 o-ot

? T6 - - <0.01 - ■cc-oi? T10 0.33 0.24 0.38 0.24 O'lo

! ? T12 <0.01 - <0.01 0.03 0 • £> 1

? T13 - - <0.01 - < o-M? ? 0.01 0.03 0.01 0.04

(00 • 5 ?

155

of each s p e c i e s has been superimposed on th e d ia g ra m s . Each

guano t r a y i s t r e a t e d s e p a r a t e l y .

3 . The n u m e r ic a l p r o p o r t i o n o f a l l e x c ep t th e r a r e s t p o l l e n ty p e s

a r e g iv en i n Appendix 4»

6 . i i . Major d i f f e r e n c e s i n th e p r o p o r t i o n s of p o l l e n on d i f f e r e n t

t r a y s .

A l t o g e t h e r , 188,000 p o l l e n g r a i n s from th e guano

c o l l e c t e d from E a tu Caves were i d e n t i f i e d as f a r a s p o s s i b l e and

counted ( e x c l u d i n g r e p e a t c o u n t s o f some samples t o de te rm in e

th e a c c u ra c y o f th e c o u n t in g method, Appendix 2 ) . Normally

500 g r a i n s were coun ted p e r sample b u t sometimes 1000 g r a i n s

were c o u n te d . The t o t a l number o f c o u n ts p e r t r a y a r e a s f o l l o w s ; -

Counts o f 500 g r a i n s each 89 83 84 86 342

Counts o f 1000 g r a i n s each 1 2 12 2 17

T o t a l number o f samples 90 85 96 88 359

T o t a l number o f g r a i n s . 45500 43500 54000 45000 188000

T able 35 r e c o r d s th e mean p e r c e n t a g e o f each

p o l l e n type on each o f th e t r a y s . The f i g u r e s were o b t a in e d by

c a l c u l a t i n g th e a r i t h m e t i c mean o f th e p e r c e n t a g e s o f each p o l l e n

type from each weekly coun t th ro u g h o u t th e s t u d y .

Two i m p o r t a n t f a c t s emerge from t h i s t a b l e :

F i r s t l y , w i th th e e x c e p t io n o f some p o l l e n ty p e s t h a t a c co u n t

f o r 0 .0 3 % o r l e s s o f t h e p o l l e n from any t r a y , a l l p o l l e n ty p e s

I■ j ■

, 156/

o c c u r r e d on e a c h t r a y ; S e c o n d l y , T ra y I d i f f e r e d . i n some r e s p e c t s

from t h e o t h e r t h r e e . T h u s , S o n n e r a t i a a l b a , S . c a s e o l a r i s and

A r t o c a r p u s s p p . p o l l e n w e re m a r k e d ly l e s s nu m ero u s on T ra y I

t h a n on an y o t h e r t r a y e x c e p t t h a t t h e r e was a l s o a v e r y l i t t l e

S . c a s e o l a r i s p o l l e n on T ra y I I . A l s o , C ocos n u c i f e r a and C e ib a

p e n t a n d r a , a l t h o u g h n o t v e r y common on an y o f t h e . t r a c t s w ere

two o r t h r e e t i m e s more nu m ero u s on T r a y s I I , I I I a n d IV t h a n

on T ra y I . The d i f f e r e n c e s a r e s i g n i f i c a n t t o p „= 0 .0 5 *

I n c o n t r a s t , D uabanga g r a n d i f l o r a was t w i c e a s

n u m erou s on T ra y I ( c o n t r i b u t i n g 50.13% o f a l l t h e p o l l e n c o u n t e d

from t h a t s i t e ) a s on T r a y s I I , I I I an d IV . O roxylum I n d i c u m ,

w h i l e n o t common on an y o f t h e t r a y s was f i v e t o t w e lv e t i m e s

a s n um ero u s on T ra y I a s on T r a y s I I , I I I an d IV . The d i f f e r e n c e s

a r e s i g n i f i c a n t t o p = 0 . 0 5 *

6 . i i i . A n a l y s i s o f t h e p o l l e n on t h e t r a y s .

S p e c i e s w i t h one A n n u a l f l o w e r i n g s e a s o n .

P a r k i a s p p . ( L e g u m i n o s a e ) .

The r e s u l t s f o r P a r k i a s p p . w e re more o r l e s s

u n i f o r m on a l l f o u r t r a y s ( F i g s . 62 t o 65 ) • T h e re w e re m a jo r

p e a k s o f P a r k i a s p p , p o l l e n from May t o J u l y 1972 an d May t o

J u n e 1973 w h ic h c o i n c i d e d w i t h t h e f l o w e r i n g s e a s o n s o f P .

s p e c i o s a e a c h y e a r . T h e re w e re s m a l l e r p e a k s i n O c t o b e r 1972

an d O c to b e r t o December 1973 w h ich c o i n c i d e d w i t h t h e f l o w e r i n g

s e a s o n s o f P . j a v a n i c a e a c h y e a r . e x c e p t December 1 9 7 3 . The p o l l e n

fo u n d i n D ecem ber 1 973 May h a v e come from P . s i n g u l a r i s a s

h e r b a r i u m s p e c i m e n s s u g g e s t s i t f l o w e r s b e tw e e n D ecem ber and

M a rc h . B o th p e a k s i n 1973 w e re n o t i c e a b l y s m a l l e r t h a n i n 1972

r e f l e c t i n g t h e p o o r f l o w e r i n g t h a t was r e c o r d e d f o r e a c h s p e c i e s

J

0

N

O

s

J

J

M

A

F

J

D

N

O

s

J

J

M

DurioP ark ia C aiba T* C ocot

The figure shows the weekly % of

the 12 most numerous pollen types

in the guano of Eonycteris spelaea

collected from Tray I , Cavern C,

Batu Caves, Selangor.

C sibaS. a lb a Artocarpus Durio

Fig. 63. The figure shows the weekly % of 'the

12 most numerous pollen types in the

guano of Eonycteris spelaea collected

from Tray I I , Cavern C, Batu Caves,

Selangor.

? , \ ( . / \ ... 1)1A 1 / Y 1 ;

❖ J L ) < p L ii1 1

O)

Y ’ s X1 i »

i 1---- ?-----

Y 1 <>f

1Y Y . ” 1

\- O . n

T i------- —

Y / /\ It

/? s i i i I\ /

------- --- 1 i 1. . . j-

Y— i—

Y| A 1 19 6 1

i sii i ii i1

-----1-----A

Cy ) \---------- ----------

i ip- •

) \1 i | 1

\ I l ii

....X <S

♦ ii p §§ 1

lI \ )

ii i. ft i. J H

J\ X ? I i <V>i i

\ ■ o »I1

i

} I b T7 1 1 ii 0

|-- a >< 1 1

I___i

i .... 0 ": 1 V i A l i

?_ - |---- ^

? Yi A 1 Y i 1 T— i

( I > )< i| V 0 J 1■ I . . .

i— P—

A hS .a lba S.c6ta& larU Dwabanga A rtocarpus Eugania

Fig. 6 k . The figure shows the weekly % of the

12 most numerous pollen types in the

guano of Eonycteris spelaea collected

from Tray I I I , Cavern C, Batu Caves,

Selangor.

J

D

N

O

s

A

J

J

M

A

M

f

J

D

N

O

t

A

M

A

M

S. a lb a V caM oiarii Ouabarvga Park ia A rto ca rp a i Eu«*nio Dario Bombax C oiba C « «

Fig.$5-'. The figure shows the weekly % of the 12

most numerous pollen types in the guano

of Eonycteris spelaea collected from Tray

IV, Cavern C, Batu Caves, Selangor.

/

/

: ■ 157

i n 1 9 7 3 . No d a t a was o b t a i n e d on t h e r e l a t i v e a b u n d a n c e o f t h e

t r e e s b u t s i n c e P . s p e c i o s a i s o f t e n l e f t s t a n d i n g d u r i n g f o r e s t

c l e a r a n c e , o r ev en p l a n t e d i n some v i l l a g e s f o r i t s e d i b l e f r u i t ,

i t may b e commoner t h a n P . . j a v a n i c a , t h u s e x p l a i n i n g t h e l a r g e r

p e a k s w h ich c o i n c i d e w i t h t h e f l o w e r i n g o f P . s p e c i o s a .

P a r k i a s p p . p n ly a d g w a re p r e s e n t o u t s i d e t h e s e

t i m e s i n v e r y s m a l l q u a n t i t i e s i n a l m o s t a l l s a m p l e s . T h i s i s

n o t s u r p r i s i n g b e c a u s e a l o n g t h e T a p a h - R i n g l e t r o a d o c c a s i o n a l

P . s p e c i o s a t r e e s w e re fo u n d i n f l o w e r e v e r y m onth from May 1972

(when r e c o r d s b e g a n ) t o J a n u a r y 197A-.

Bombax v a l e t o n i i and C e ib a p e n t a n d r a (B om b acaceae )

M ino r p e a k s a p p e a r e d f o r b o t h s p e c i e s on T ra y s

I I , I I I and e s p e c i a l l y IV , a l l c o i n c i d i n g w i t h t h e f l o w e r i n g

s e a s o n s o f t h e t r e e s . V ery l i t t l e p o l l e n o f e i t h e r s p e c i e s was

r e c o v e r e d on T ra y I . The o c c u r r e n c e o f B . v a l e t o n i i p o l l e n was

l a r g e l y c o n f i n e d t o t h e f l o w e r i n g s e a s o n b u t C. p e n t a n d r a was

f r e q u e n t l y p r e s e n t i n v e r y s m a l l q u a n t i t i e s t h r o u g h o u t t h e y e a r .

T h i s i s c o n s i s t e n t w i t h t h e k n o w le d g e t h a t f l o w e r i n g o f B . v a l e t o n i i

i s m o s t l y r e s t r i c t e d t o t h e f l o w e r i n g s e a s o n b u t C. p e n t a n d r a

t r e e s may o c c a s i o n a l l y f l o w e r a t a n y t i m e .

S p e c i e s w i t h two a n n u a l f l o w e r i n g s e a s o n s .

D u r lo s p p . (B o m b acaceae )

P e a k s o c c u r r e d on a l l f o u r t r a y s i n M arch , May

an d S e p te m b e r t o O c to b e r 1972 a nd i n A p r i l a n d S e p te m b e r 1973

( F i g s . 6 2 t o 6 5 ) . T h e re was c o n s i d e r a b l e v a r i a t i o n i n t h e

s i z e o f p e a k s on d i f f e r e n t t r a y s b u t t h e y w ere a l l c o n s i s t e n t l y

I\

■ I

. 1 5 8

s m a l l e r on T r a y I t h a n o n ' a n y o t h e r t r a y . The p e a k s i n 1973 w e re

g e n e r a l l y h i g h e r t h a n i n 1972 an d on T r a y s I , I I a n d IV t h e

h i g h e s t p e a k was i n A p r i l 1 973 h u t on T ra y I I I t h e h i g h e s t was

i n S e p te m b e r 1973• A l l t h e p e a k s c o i n c i d e d w i t h f l o w e r i n g o f

D. z i b e t h l n u s .

The h e a v i e s t f l o w e r i n g s e a s o n r e c o r d e d f o r D.

z i b e t h i n u s d u r i n g t h e s t u d y was i n A p r i l 1973 an d t h e p o o r e s t

was i n S e p te m b e r 1973 when many D. z i b e t h i n u s t r e e s f a i l e d t o

f l o w e r an d o t h e r s p r o d u c e d o n l y a few f l o w e r s . T h i s a g r e e s w i t h

t h e h i g h e s t p e a k o f p o l l e n i n t h e gu ano on t h r e e o f t h e f o u r

t r a y s o c c u r r i n g i n A p r i l 1973 b u t t h e r e a s o n f o r t h e r e l a t i v e l y

h i g h p e a k r e c o r d e d i n S p e te m b e r t h a t y e a r i s n o t c l e a r . L i t t l e

i s known o f t h e w i l d s p e c i e s o f D u r io and t h e y may h av e f l o w e r e d

w e l l i n S e p te m b e r 1973*

V e r u c u l a t e D u r io s p p . p o l l e n was v e r y o c c a s i o n a l l y

fo u n d on T r a y s I I a n d I I I ( D u r lo s p p . t y p e 2 , T a b le 35 ) • (F u c h s

(1967)) exam in ed t h e p o l l e n o f 20 s p e c i e s o f D u r i o . He fo u n d s i m i l a r

g r a i n s i n D. o b lo n g u s an d D. t e s t u d i n a r u m B e c c a r i and p u b l i s h e d

a p h o t o g r a p h o f t h e l a t t e r . )

T8 ( S a p o t a c e a e )

A l l f o u r t r a y s h ad one m a jo r p e a k i n May 1 9 7 2 .

T r a y s I I , I I I an d IV showed a m in o r p e a k a l i t t l e e a r l i e r , i . e .

i n March 1 9 7 2 . C o r r e s p o n d i n g b u t v e r y s m a l l p e a k s w ere o b s e r v e d

from F e b r u a r y t o March an d i n May 1973 on T ra y I I a nd I I I ( F i g s .

63 and 6^ ) b u t o n l y t h e f o r m e r was r e c o r d e d on T r a y s I an d IV

( F i g s . 62 and 6 5 ) . B e s i d e s t h e s e p e a k s i t o c c u r r e d o c c a s i o n a l l y

t h r o u g h o u t t h e y e a r . I t s o r i g i n i s unknown a n d more t h a n one s p e c i e s

i ' ' -

159

•* . . . . ‘

may b e i n v o l v e d . -

S p e c i e s f l o w e r i n g s e a s o n a l l y o r i n f l u s h e s more t h a n - t w i c e p e r annum .

S o n n e r a t i a a l b a ( i n c l u d i n g S . o v a t a ) ( S o n n e r a t i a c e a e )

W ith one e x c e p t i o n ( s e e b e lo w ) S . a l b a p o l l e n

beqame a b u n d a n t i n t h e gu ano i n B a tu Caves d u r i n g e a c h f l o w e r i n g

f l u s h . However i t s a b u n d a n c e v a r i e d from one t r a y t o a n o t h e r .

■Tray^I c o n s i s t e n t l y h a d l e s s t h a n d i d t h e o t h e r t h r e e t r a y s ( T a b l e

' 35 ) anc* T ra y IV h a d more t h a n t h e o t h e r s i n 1 973 ( F i g s 62 t o 65) •

N orm al f l o w e r in g - f l u s h e s o f S . a l b a l a s t e d a b o u t

one m onth b u t two f l u s h e s e a c h l a s t e d a b o u t 2 . 5 m o n th s . D u r in g

t h e f i r s t l o n g f l u s h from m i d - A p r i l t o e a r l y J u l y 1972 f l o w e r i n g

w as som ewhat r e t a r d e d an d n e v e r p a r t i c u l a r l y h e a v y a l t h o u g h

f l o w e r s w ere a lw a y s p r e s e n t . V ery l i t t l e S . a l b a p o l l e n o c c u r r e d

i n t h e gu ano c o l l e c t - e d i n B a tu C aves t h e n . I n s t e a d t h e m a j o r i t y / N

o f g r a i n s w e re T8 ( S a p o t a c e a e ) e a r l i e r and P a r k i a s p p . ( p r o b a b l y

P . s p e c i o s a ) l a t e r . T8 h a s y e t - t o b e i d e n t i f i e d b u t P . s p e c i o s a

f l o w e r e d h e a v i l y a t t h a t t i m e .N S i n c q T8 p o l l e n was so a b u n d a n t

on a l l t a b l e s t h e s p e c i e s fpom w h ich i t came,.can r e a s o n a b l y be

i n f e r r e d t o h a v e f l o w e r e d w e l l . The b a t s a p p a r e n t l y f e d f rom

P a r k i a s p p . a nd T8 i n p r e f e r e n c e t o S . a l b a i n t h a t i n s t a n c e

A t t h e c o r r e s p o n d i n g t im e i n 1973 t h e r e was v e r y

l i t t l e T8 p o l l e n i n t h e s a m p le s an d P . s p e c i o s a h a d a v e r y p o o r

f l o w e r i n g s e a s o n w h i l e t h e £>, a l b a f l u s h was n o r m a l . A l th o u g h

some P a r k i a s p p . p o l l e n was p r e s e n t t h e r e was a l s o a p e a k i n t h e

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

D u r in g t h e s e c o n d e x t e n d e d f l o w e r i n g f l u s h o f

S . h l b a from l a t e S e p te m b e r t o e a r l y December 1 9 7 3 , S . a l b a p o l l e n

i ■ ■ ■

" , ■ . 160

was a b u n d a n t i n t h e guan o from a l l f o u r t r a y s , Howex*er t h e r e was

a r e d u c t i o n i n t h e q u a n t i t y o f S . a l b a p o l l e n on a l l t r a y s f o r

a f o r t n i g h t f rom l a t e O c to b e r t o e a r l y November t h a t y e a r . The

r e a s o n i s n o t known b n t i t may r e f l e c t a r e d u c t i o n i n t h e a b u n d a n c e

o f f l o w e r s t h a t wais n o t r e c o r d e d i n t h e f i e l d ,

- Many s a m p le s c o l l e c t e d o u t s i d e t h e f l o w e r i n g

f l u s h e s c o n t a i n e d p o l l e n r e c o r d e d a s S . a l b a . - O c c a s i o n a l S . a l b a

t r e e s c o u l d a lw a y s b e fo u n d i n f l o w e r b u t some o f t h i s p o l l e n

was u n d o u b t e d l y from S . o v a t a .1 *

T r 'ees t h a t f l o w e r c o n t i n o u s l y .

S o n n e r a t i a c a s e o l a r i s ( S o n n e r a t i a c e a e ) .

- c a s e o l a r i s p o l l e n was p r e s e n t i n m o s t s a m p le s

from a l l t r a y s b u t i t s a b u n d a n c e f l u c t u a t e d : . On T ra y I i t was

n e v e r n u m erou s (>27.2% once., o t h e r w i s e <9%) b u t on T ra y I I and

e s p e c i a l l y T r a y s I I I and IV i t was a t t i m e s a m a j o r c o n s t i t u e n t

o f t h e g uano ( e . g . fo r m in g up t o k0% on T ra y I I I i n s a m p le 91)

( .A p p e n d ix ^ ) . I t was g e n e r a l l y more a b u n d a n t i n 1973 t h a n i n\

1 9 7 2 . The r e a s o n s f o r t h e f l u c t u a t i o n s a r e n o t c l e a r . They do

n o t a lw a y s c o i n c i d e w i t h p e a k s i n S . a l b a p o l l e n (when more o f

t h e b a t s m i g h t b e - e x p e c t e d t o v i s i t c o a s t a l d i s t r i c t s ) b u t t h e r e

i s a s l i g h t t e n d e n c y f o r p e a k s ' t o p r e c e d e o r s u c c e e d p e a k s i n

S . a l b a p o l l e n ./

D uabanga g r a n d i f l o r a ( S o n n e r a t i a c e a e ) .

/ D. g r a n d i f l o r a p o l l e n was p r e s e n t i n a l m o s t e v e r y

s a m p le b u t i t s a b u n d a n c e on a l l t r a y s f l u c t u a t e d w i d e l y . I t wasv

v e r y common t h r o u g h o u t t h e s t u d y on T ra y I w h ere i t c o n t r i b u t e d

1 61

50,.13% o f a l l t h e p o l l e n ' g r a i n s ' c o u n t e d . On T r a y s I I , I I I and

IV i t c o n t r i b u t e d o n l y h a l f t h a t q u a n t i t y ( T a b le 3 5 ) . On T ra y I

m o s t r e d u c t i o n s i n t h e p r o p o r t i o n o f D. g r a n d i f l o r a p o l l e n

c o i n c i d e d w i t h p e a k s i n t h e p r o p o r t i o n o f p o l l e n fro m s e a s o n a l l y

f l o w e r i n g s p e c i e s . E . g . f rom F e b r u a r y ’1 972 t o F e b r u a r y 1 9 7 3 '

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

s u c c e s s i v e l y w i t h p e a k s i n p o l l e n o f ’A r t o c a r p u s s p p . , T8 ( S a p o t a c e a e ) ,

P a r k l a s p p . , E u g e n ia s p p . , S o n n e r a t l a a l b a , E u g e n ia ' s p p . , P a r k l a

SPP*> S . a l b a a nd f i n a l l y A r t o c a r p u s s p p . ( F i g . 62 ) .

On T r a y s ' l l , I I I an d IV D. g r a n d i f l o r a p o l l e n

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

A p r i l b u t - f r o m A p r i l t o A u g u s t i t was much l e s s n u m e r o u s . From

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

t o t h e p r e s e n c e o f p o l l e n from s e a s o n a l l y f l o w e r i n g s p e c i e s , b u t

from A p r i l -to A u g u s t when i t was r e l a t i v e l y s c a r c e i t was u s u a l l y

r e p l a . c e d by A r t o c a r p u s s p p . p o l l e n an d t h e f l u c t u a t i o n s i n r e s p o n s e

t o p e a k s i n s e a s o n a l l y f l o w e r i n g s p e c i e s w e re r e f l e c t e d i n t h e

l a t t e r g e n u s r a t h e r t h a n i n D . / g r a n d l f l o r a .

The d a t a s u g g e s t s t h a t D, g r a n d i f l o r a was an

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

o v e r T r a y - I an d f o r t h o s e r o o s t i n g o v e r T r a y s I I , I I I an d IV

when Ar t o c a r p u s s p p . p o l l e n was l e s s a v a i l a b l e t o th e m . H ow ever,

many o f t h e b a t s r o o s t i n g a b o v e a l l t h e t r a y s s w i t c h e d l a r g e l y

t o s e a s o n a l l y f l o w e r i n g t r e e s f o r fo o d when t h e y becam e a v a i l a b l e .

The u s e o f J ) . g r a n d i f l o r a t h r o u g h o u t t h e y e a r by b a t s r o o s t i n g

o v e r T ra y I a c c o u n t s a t l e a s t i n p a r t f o r t h e h i g h e r p r o p o r t i o n

Sran (^ i f l o r a p o l l e n on t h a t t r a y r e c o r d e d i n T a b le 35 •

Ar t o c a r p u s sp i j . (M o ra c e a e ) 'N

The species of Artocarpus on which the bats

fed have not been identified and so their phenology is unknown.

T h i s makes i n t e r p r e t a t i o n o f t h e d a t a t e n t a t i v e . A r t o c a r p u s s p p .

p o l l e n was p r e s e n t i n a l m o s t a l l s a m p le s from a l l f o u r t r a y s . .

I t was se ld o m v e r y common on T r a y I b u t on T r a y s I I , I I I and IV

i t l a r g e l y r e p l a c e d Dugtbanga g r a n d ! f l o r a p o l l e n from A p r i l t o

A u g u s t i n b o t h y e a r s and was common a t o t h e r t i m e s to o ( F i g s . 62

t o 6 5 ) •

P e a k s _ i n t h e p r o p o r t i o n o f p o l l e n o f s e a s o n a l l y

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

s p p . p o l l e n on T r a y s I I , I I I an d IV e x c e p t when i t was r e p l a c e d

ky £,• g r a n d i f l o r a any w ay . T h i s i s c l e a r l y d e m o n s t r a t e d on T r a y s

I I I and IV ( F i g s . 6^ t o 65 ) and t o a l e s s e r e x t e n t on T ra y II

( F i g . 63 )• w h e n i n A p r i l 197?- A r t o c a r p u s s p p . r e p l a c e d D. g r a n d i f l o r a

an d i n s u b s e q u e n t m o n th s r e f l e c t e d p e a k s i n T8 ( S a p o t a c e a e ) ,

P a r k ! a s p p . ( p r o b a b l y £ . s p e c i o s a ) , E u g e n ia s p p . and S o n n e r a t i a

a l b a p o l l e n r e s p e c t i v e l y . T h e re w ere t i m e s when A r t o c a r p u s s p p .

a n d D. g r a n d i f l o r a w e re b o t h common a n d r e f l e c t e d m in o r f l u c t u a t i o n s

i n e a c h o t h e r , e . g . on T r a y s II and IV ( F i g s . 63 and 65 ) from

J a n u a r y t o M arch 1973* A f u l l u n d e r s t a n d i n g o f t h e r e l a t i o n s h i p s

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

gu ano m u s t a w a i t f u r t h e r k n o w le d g e o f t h e p h e n o lo g y and d i s t r i b u t i o n

o f t h e A r t o c a r p u s ; : s p p . an d a l s o o f w h ich a g e o r s e x c l a s s e s o f

b a t f e e d from i t m o s t l y .

Cocos nucifera.

C o c o n u t p o l l e n was f r e q u e n t l y p r e s e n t t h r o u g h o u t

1972 and 1973* However, i t was less abundant on Tray I than on

Trays I I , I I I and IV and was never abundant on any of them. The

re la tive scarcity on Ti'ay I may be correlated with the re la tive

scarcity of Sonneratia alba pollen on Tray I which indicated

that bats roosting over th is tray did not go to the coast as

often as diu those roosting over the other- tray s.,. Since coconut

phlms. are particu larly abundant along the coast bats that went

there would have more opportunity to feed from them than those

that did not' go there so often.

Other species, _

Eugenia spp. (Myrtaceae)

Eugenia spp. pollen was present in small amounts

i n most samples but in August 1972 and 1973 i t became veryt

abundant in the guano. These peaks coincided with flowering of

E. malaccensis in Selangor, which probably accounted, a t least

in part, for them. However, flowering of E. malaccensis in March

each year either did not coincide with peaks or, sometimes,

coincided with very small peaks in 'the proportion bf Eugenia spp.

pollen in the guano. The reason for the absence of larger peaks-

in the proportion of Eugenia spp..pollen in Mhrch each year is

not known

'Another peak in Eugenia spp. pollen occurred in

April each year except’ on Tray IV (Fig 65 ) and in 1972 on Tray

I I (Fig. 5 3 ) . Other small peaks occurred from time to time but

they were not consistent over the two years and were not present

on a l l the trays. Undoubtedly, Eugenia spp. besides E. malaccensis

1 Qi

p r o v i d e d fo o d f o r E . s p e l a e a .

(N .B . E u g e n ia s p p . p o l l e n g r a i n s a r e v e r y s m a l l ( T a b l e 3 ) and

do n o t s t a i n d e e p l y i n b a s i c . f u s c h i h ; T h e i r a b u n d a n c e i n some

s a m p le s may h a v e b e e n u n d e r e s t i m a t e d or, when n o t a b u n d a n t f t h e i r

p r e s e n c e o v e r l o o k e d . ) ' .

- ^ 3 .

' T3 p o l l e n w as v e r y o f t e n p r e s e n t i n v e r y s m a l l

nu m b ers on a l l t r a y s . T h e re was g e n e r a l l y l e s s on T ra y I t h a n

on T r a y s I I , I I I an d IV ( T a b l e 3 5 ,' an d F i g s 62 t o 65 ) • T h e re

w e re . s m a l l b u t e x t e n d e d p e a k s from- December 1 972 t o J a n u a r y 1973

a n d from N ovember 1973 t o J a n u a r y 1974» T h i s d o e s n o t c o i n c i d e

w i t h p e a k s i n t h e p r o p o r t i o n o f S o n n e r a t i a a l b a p o l l e n i n t h e

g u a n o , s u g g e s t i n g t h a t T3 i s n o t a n a b e r r a n t fo rm o f S . a l b a

p o l l e n ( s e e p . 1 5 1 ) •

The r e m a i n i n g p o l l e n s p e c i e s , i . e . A re n g a s p p . ,

B a r r i n g t o n i a s p p . , M a n g i f e r a s p p . , O roxylum i n d i c u m , B h iz o p h o r a

s p p . , Tif, T6 , T 1 0 , T12 and T13 w ere a l l r a r e , s e ld o m f o r m in g

m ore t h a n 1 t o 2 % o f t h e p o l l e n g r a i n s i n a n y s a m p le an d ev en

when com bin ed t h e y fo rm ed l e s s t h a n 0 . 6 % o f a l l t h e p o l l e n c o u n te d

d u r i n g t h e s t u d y ( T a b l e 35 ) •

I t e m s b e s i d e s p o l l e n fo u n d i n t h e g u a n o .

I n s e c t s w e re o f t e n drow ned i n t h e gu ano on t h e

guano t r a y s a n d d i p t e r a n s o f t e n l a i d e g g s on T ra y I . Fungus grew

on a l l o f th e m . T h e r e f o r e , i n s e c t r e m a i n s and f u n g a l s p o r e s fo un d

i n t h e s a m p le s c o u l d n o t b e a t t r i b u t e d w i t h c e r t a i n t y t o t h e b a t s .

H ow ever , some i t e m s o f p l a n t o r i g i n m u s t h a v e come from t h e b a t s .

165

These included anthers of Parkia spp. and Artocarpus spp.,

frequently with pollen grains inside them ( Fig. 38 ), and

small fragments of plant tissue which had usually decomposed.

Spiral structures, presumed to be the helical thickening structures

from plant vascular tissue were also- present,in most samples.

The plant tissues, besides pollen and anthers, were never common

in the samples and when chiropterophilous flowers were abundant,

e.g. during the flowering season of Parkia speciosa or during

flowering flushes of Sonneratia alba they were v ir tua lly absent.

Their presence a t other times shows that the bats did feed on

plant products (possibly ripe fru it?) besides pollen and nectar

but thatv when available pollen and nectar were preferred.

7 . F r e s h gu ano from Gua S a n d i n g .

7 . i . Collection of samples . .

19 gu ano s a m p le s c o m p r i s i n g 2 0 0 f r e s h f a e c a l p e l l e t s

e a c h w e re c o l l e c t e d from Gua S a n d in g on one d ay e a c h m on th from

May 1 972 t o D ecem ber 1 973 e x c e p t i n May 1 973-. 500 g r a i n s w e re

c o u n t e d f rom e a c h s a m p l e . A t o t a l o f 9 5 0 0 g r a i n s w e re c o u n t e d .

7 . i i . P r e s e n t a t i o n an d l i m i t a t i o n o f d a t a .

Since the samples were derived from fresh guano

collected on one day each month they are no more than "spot checks"

on the pollen present in the. guano on the date of collection.

They therefore d iffer from the continous collections made in

Batu Caves and for th is reason k ite diagrams, such as were drawnt

for the Batu Caves samples, cannot be drawn for the Gua Sanding

samples. Another difference i s that the samples from Gua Sanding

166

w ere c o l l e c t e d ra n d o m ly from a w id e a r e a on t h e c a v e f l o o r . They

d i d n o t a l l o r i g i n a t e from a s i n g l e r o o s t i n g g r o u p o f b a t s - a s

d i d t h e f o u r s e r i e s o f . s a m p l e s from B a tu C a v e s .

The d a t a i s p r e s e n t e d i n T a b le 36 a s t h e n u m e r i c a l

p e r c e n t a g e o f e a c h p o l l e n t y p e p r e s e n t i n e a c h m o n th ly s a m p le .

The r e m a r k s c o n c e r n i n g t h e v a l u e o f n u m e r i c a l d a t a a s com pared

w i t h t h e v o l u m e t r i c d a t a on p.. 1 52 a p p l y e q u a l l y h e r e .

7 . i i i . V a r i e t y o f p o l - le n t y p e s .

1 8 p o l l e n t y p e s w e re fo u n d i n t h e g uan o from

G u a -S a n d in g ; 7 f e w e r t h a n i n gu ano from B a tu C a v e s . A l l p o l l e n*

t y p e s fo u n d i n Gua S a n d in g w e re a l s o fo u n d i n B a tu C a v e s . The

s e v e n t y p e s fo u n d i n B a tu C av es b u t n o t Gua S a n d in g w e re r a r e

i n B a tu C a v e s . They w e re : - D u r lo s p p . w i t h a v e r u c u l a t e e x i n e

(= P u r i o s p p . t y p e 2 , T a b le 35 )» B a r r l n g t o n i a s p p . j R h iz o p h o r a

s p p . , T ^ , T6, T12 an d T13«

7 . i v . N u m e r i c a l p r o p o r t i o n s o f e a c h p o l l e n t y p e .

The s a m p le s showed s e v e r a l v e r y m ark ed d i f f e r e n c e s

t o t h o s e from B a tu C a v e s . Thus S o n n e r a t i a a l b a was r a r e i n Gua

S a n d i n g . I t fo rm ed k,2.% o f t h e p o l l e n i n J u n e 1973> 1.8% i n

O c to b e r 1973 a nd l e s s t h a n 1% i n t h e r e m a i n i n g s a m p l e s . I t was

a b s e n t f rom 10 (53%) o f t h e s a m p l e s . O n ly 0 .^5% o f a l l t h e p o l l e n

c o u n te d was S . a l b a ( c f . B a tu C a v e s , T a b le 35 , w h e re S . a l b a

c o n t r i b u t e d 6 .5% t o J\k»k% o f a l l t h e p o l l e n c o u n t e d on e a c h t a b l e . )

I n c o n t r a s t , S . c a s e o l a r i s was t h e m o s t n u m erou s s i n g l e p o l l e n

s p e c i e s i n s a m p le s from Gua S a n d i n g , m ak ing up 25.2% o f a l l t h e

p o l l e n p r e s e n t ( c f . 1 .**% t o 9.3% i n B a tu C a v e s , T a b le 35 ) • I t

o c c u r r e d i n e v e r y sa m p le a nd c o n t r i b u t e d 65.2% o f t h e p o l l e n i n

T a b l e 3 6 .

The t a b l e r e c o r d s t h e ° / o f e a c h p o l l e n t y p e i n t h e g u a n o o f E o n y c t e r i s s p e l a e a c o l l e c t e d

m o n t h l y f r o m Gua S a n d i n g , P e r a k b e t w e e n May 19 7 2 a n d D e c e m b e r 1 9 7 3 .

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May 0 . 6 3 . 9 1 . 0 1 . 2 1 6 . 5 1 . 2 1 . 2 1 . 3 6 . 9 0 . 4 5 5 . 2 1 . 2 2 . 5

J u n 0 . 2 1 8 . 8 0 . 6 0 . 8 0 . 8 6 5 . 5 0 . 8 8 . 1 3 . 6 0 . 9

J u l 1 6 . 0 0 . 2 0 . 5 2 . 3 0 . 4 5 8 . 5 0 . 4 2 . 3 0 . 8 1 5 . 7 2 . 4 0 . 5

Aug 2 0 . 8 0 . 6 5 . 2 5 2 . 4 4 . 6 1 4 . 6 2 . 0

S e p 0 . 2 2 0 . 9 1 . 0 2 3 . 3 9 . 1 4 . 6 8 . 7 0 . 2 2 9 . 4 2 . 6

O c t 1 0 . 2 3 . 8 4 . 4 4 . 4 2 0 . 8 2 . 4 2 8 . 0 2 2 . 0 4 . 0

Nov 1 9 . 1 1 . 8 1 . 8 0 . 4 4 . 3 0 . 2 0 . 2 7 . 9 0 . 4 6 2 . 8 0 . 8 0 . 4

Dec 6 . 0 2 . 6 2 0 . 0 4 . 4 0 . 4 0 . 2 4 . 4 4 1 . 4 1 5 . 2 . 3 . 6 1 . 8

J a n 3 7 . 6 0 . 2 5 . 6 1 7 . 4 0 . 2 5 . 8 3 . 4 2 9 . 4 0 . 4

F e b 7 . 0 0 . 2 3 . 6 0 . 2 0 . 4 1 2 . 4 4 . 8 6 4 . 4 0 . 2 5 . 8 1 . 2 0 . 2 0 . 2

Mar 0 . 2 6 . 8 6 . 8 0 . 2 1 2 . 4 6 9 . 0 2 . 2 2 . 2 0 . 2

A p r 1 7 . 8 0 . 4 1 . 2 9 . 6 1 0 . 6 3 7 . 2 2 3 . 0 0 . 2

May 7 ? 7 ? ? 7 ? 7 7 7 7 7 ? 7 7 7 ? 7

J u n 0 . 6 2 1 . 0 1 . 2 0 . 4 4 8 . 6 3 . 8 4 . 2 1 7 . 6 2 . 6

J u l 0 . 2 1 4 . 4 5 . 0 0 . 8 0 . 2 2 . 0 9 . 5 0 . 4 6 5 . 2 2 . 0

Aug 1 . 8 3 . 6 0 . 2 8 . 8 2 0 . 6 7 . 2 4 . 0 2 6 . 4 0 . 4 2 3 . 8 3 . 0 0 . 4

S e p 1 6 . 2 0 . 2 1 2 . 2 3 . 2 1 . 6 5 1 . 2 0 . 2 1 0 . 6 4 . 6

O c t 1 6 . 0 0 . 4 7 . 4 0 . 4 0 . 2 9 . 0 2 . 2 1 2 . 8 1 . 8 4 2 . 6 7 . 0

Nov 1 9 . 2 0 . 4 3 3 . 8 0 . 2 3 . 6 0 . 2 2 . 4 3 9 . 4 0 . 8

Dec 0 . 6 1 5 . 0 0 . A 2 4 . 8 5 . 4 0 . 2 0 . 2 2 1 . 8 2 0 . 4 7 . 8 3 . 4

Mean 0 . 1 0 . 1 1 4 , 8 0 . 5 o . e 2 . 8 5 . 3 5 . 2 0 . 1 1 4 . 6 8 . 3 I .1 1 9 . 2 0 . 5 2 5 . 2 2 . 4 0 . 1 0 . 3

A

1 6 7

J u ly 1973.

A r to c a rp u s spp . and Duabanga g r a n d i f l o r a were

l e s s abundan t i n Gua Sanding,- e s p e c i a l l y th e former which

c o n t r i b u t e d o n ly 2.4% of a l l t h e p o l l e n counted (cf. 17.8% to

34.7% i n E a tu Caves, Table 35 ) •

Arenga spp . which was r a r e ±n Batu Caves ( < 0.01%

of a l l t h e p o l l e n coun ted) was p r e s e n t i n ev e ry sample from Gua

Sand ing . I t c o n t r i b u t e d 1 4.84% of a l l th e p o l l e n p r e s e n t and th e

monthly p r o p o r t i o n s ranged from 0.2% to 37.6%. No p a t t e r n to th e ' /

f l u c t u a t i o n s w as ' o b s e r v e d . I t d id n o t r e f l e c t peaks i n th e p o l l e n

of s e a s o n a l l y f l o w e r i n g s p e c i e s , e . g .

Year Month % P a r k i a s p p . % Arenga s p p .

1972 , May 1 .2 3 . 9• June 65 .2 1 8 . 8

J u l y 5 8 .5 1 6 .0

August 5 .2 20 .8

1973 June 4 8 .6 21 .0

J u l y 2 .0 14 .4

August 7 .2 1 .8

Thus i t d id n o t behave as d id Duabanga g r a n d i f l o r a

and A r to c a rp u s spp . i n E a tu Caves.

O ther s p e c i e s behaved i n a s i m i l a r way i n Gua

Sanding and Batu Caves. Thus t h e r e were major peaks i n th e

p r o p o r t i o n o f P a r k i a s p p . p o l l e n ( p ro b a b ly P . s p e c i o s a ) i n June

168

/.to July 1972 and June to August 1 9 7 3 . Minor peaks (probably of

£• .lavaflica) occurred in October each year. There were peaks in

the proportion of Durio spp.pollen in May and September 1972

and April (and May?) and August to September 1973*

There were higher than normal proportions of

Eugenia spp. pollen in May, August and December 1972 and February,

July and December 1973 (no records for May 1 9 7 3 ) . The peaks in

August 1 9 7 2 , February 1973 and July 1973 probably coincide with

the |>eaks observed in March and August in Batu Caves ( i . e . when

malaccensis flowered) but the 1973 peaks were one month earl ie r

in Perak than in Selangor. Peaks in the proportion of Eugenia

spp. pollen occurred in some Batp Caves samples in December 1972

and January 1 9 7 3 . They may have been of the same species as that

which caused peaks in Gua Sanding in December each year, the

season again being advanced one month in Perak l* J**73.

C e ib a p e n t a n d r a was p r e s e n t a l m o s t e v e r y month

b u t w i t h a p e a k from O c to b e r 1972 t o March 1973 ( e s p e c i a l l y

J a n u a r y 1973) a n d O c to b e r t o D ecem ber 1973 ( t h e l a s t sam p le

c o l l e c t e d from Gua S a n d i n g ) . Bombax v a l e t o n i i was p r e s e n t f rom

November 1972 t o J a n u a r y 1973 an d i n December 1 9 7 3 . Odd g r a i n s

o c c u r r e d i n o t h e r m o n th s . T h e se s e a s o n s a r e s i m i l a r t o t h o s e f o r

p o l l e n i n B a tu C a v e s , t h e o c c u r r e n c e o f C. p e n t a n d r a b e i n g l e s s

d e f i n e d i n gu an o f rom b o t h c a v e s .

The s m a l l p e a k o f T8 ( S a p o t a c e a e ) p o l l e n i n guano

f rom Gua S a n d in g i n O c to b e r 1972 was n o t o b s e r v e d i n t h e B a tu

C av es s a m p le s b u t t h e p e a k i n F e b r u a r y and M arch 1973 was o b s e r v e d

i n guano f rom b o t h l o c a l i t i e s .

169

I n 1 972 O roxylum in d i c u m p o l l e n i n t h e s a m p le s

from Gua S a n d in g d i d n o t c o i n c i d e w i t h t h e f l o w e r i n g p e a k

o b s e r v e d on t h e n e a r b y T a p a h - R i n g l e t r o a d . I t d i d s o i n 1973

( s e e p . 5 3 ) .

The r e m a i n i n g p o l l e n s p e c i e s i . e . Musa s p p . ,

C o c o s •n u c i r e r a , M a n g ix e ra s p p . , ? ( C o m p o s i t a e ) , T3 an d T1 0 w e re '

r a r e i n Gua S a n d i n g .

8 . S u b f o s s i l d e p o s i t s .

8 . i . E a t u C a v e s . . x

The s o i l p r o f i l e e x p o s e d i n t h e s i d e o f a w a t e r

c h a n n e l i n t h e P en ny room show ed f i v e d i s t i n c t l a y e r s . A l a y e r

o f w h i t e c l a y a b o u t 5 cm t h i c k c o v e r e d t h e b e d r o c k and was

c o v e r e d by a t h i n l a y ’b r o f b l a c k s o i l 3 cm t h i c k . E e tw e e n 8 cm

an d i|-l cm ab o v e b e d r o c k was a l a y e r o f f i n e g r a i n e d s o i l c o n t a i n i n g\

many p a r t i c l e s o f l i m e s t o n e . I t s c o l o u r ch a n g e d g r a d u a l l y from

p a l e o c h r e a t I t s b a s e t o r e d a t t h e t o p . B e tw ee n if1 cm and 7 6 cm

from b e d r o c k t h e r e was a l a y e r o f v e r y d a r k s o i l , a p p a r e n t l y

r i c h i n o r g a n i c m a t t e r . The t o p l a y e r from ? 6 cm t o 107 cm was

brow n e a r t h . ' .A l a y e r o f c h a r c o a l was fo u n d b e tw e e n 91 cm an d 9 7

cm from b e d r o c k . .

C o re s w e re d i f f i c u l t t o c u t due t o t h e co m p a c t

n a t u r e o f t h e s o i l . The s a m p le s t h a t w ere o b t a i n e d s u g g e s t e d a

p r o f i l e s i m i l a r ' t o t h a t d e s c r i b e d a b o v e .

No p o l l e n was fo u n d i n an y s a m p le s e x c e p t i n t h e

t o p 2 cm and t h a t was v e r y r e c e n t b u t m o s t g r a i n s w e re more o r

l e s s d e c a y e d .

8 » i i . Gua S a n d i n g .

S e v e r 'a l c o r e s w e re o b t a i n e d from t h e s o i l s u r f a c e

t o b e d r o c k . They v a r i e d from ^0 t o 70 cm l o n g . A l l w e re composed

o f w h i t i s h , g r a n u l a r p a r t i c l e s o f l i m e s t o h e . No p o l l e n was

d i s c o v e r e d i n an y o f them bu.t d i s i n t e g r a t i n g g r a i n s w ere c o l l e c t e d

from a. t h i n l a y e r o f s l u d g e on t h e s u r f a c e o f t h e d e p o s i t . At

t i m e s a r i v e r ' o u t s i d e t h e c a v e f l o o d e d an d w a t e r f lo w e d th r o u g h

t h e c a v e . -T h is may h a v e c a r r i e d away t h e decomposed, o r g a n i c m a t t e r

l e a v i n g t h e h e a v i e r l i m e s t o n e p a r t i c l e s t o fo rm a b e d o f a l m o s t

p u r e l i m e s t o n e , s a n d .

9 . P o l l e n s a m p le s c o l l e c t e d from E o n y c t e r i s s p e l a e a .

P o l l e n s a m p le s w e re c o l l e c t e d f rom t h e b o d i e s)

o f 2i+’b a t s an d f a e c a l s a m p le s w ere c o l l e c t e d from 37 b a t s . I nv '\

m o s t i n s t a n c e s t h e p o l l e n on t h e b o d i e s o f t h e b a t s was o f p l a n t s

g ro w in g i n t h e v i c i n i t y o f t h e c a p t u r e s i t e , e . g . p o l l e n from

t h e b o d i e s o f b a t s c a u g h t u n d e r a f l o w e r i n g D u r io z i b e t h i n u s

t r e e a t t h e F . S . C . , U lu Gombak was u s u a l l y p u r i o s p p . p o l l e n ,

an d p o l l e n c o l l e c t e d from b a t ^ c a u g h t a t R a n ta u P a n j a n g was

u s u a l l y G o n n e r a t i a s p p . p o l l e n .

The p o l l e n i n f a e c a l s a m p le s show ed a l i t t l e

more v a r i a t i o n , e . g . s a m p le s from b a t s c a u g h t u n d e r a f l o w e r i n g

D u r io z i b e t h i n u s t r e e a t t h e F . G . C . , U lu Gombak m o s t l y c o n t a i n e d

D u r io s p p . p o l l e n b u t D uabanga g r a n d i f l o r a p o l l e n an d Musa s p p .

p o l l e n w ere a l s o p r e s e n t i n some o f th em , p o l l e n i n s a m p le s

c o l l e c t e d a t R a n ta u P a n j a n g commonly c o n t a i n e d G o n n e r a t i a s p p .

p o l l e n b u t A r t o c a r p u s s p p . , Bombax v a l e t o n l l , C e ib a p e n t a n d r a ,

171

D u r io s p p . , E u g e n ia s p p . a n d P a r k i a s p p . p o l l e n w e re a l s o fo u n d

i n som e. .

A d e t a i l e d a n a l y s i s o f t h e p o l l e n p r e s e n t i n

t h e s e S am p les w o u ld o n l y r e f l e c t t h e h a b i t a t s i n w h ic h t h e b a t s

w e re c a u g h t . I t w o u ld n o t show t h e ' r e l a t i v e i m p o r t a n c e t o t h e

b a t s o f an y o f t h e p o l l e n t y p e s e n c o u n t e r e d . F o r t h i s r e a s o n

a d e t a i l e d a n a l y s i s i s o m i t t e d an d a n y s a m p le s t h a t r e v e a l

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

m e n t io n e d i n o t h e r r e l e v a n t s e c t i o n s o f t h e t e x t .

I t e m s b e s i d . e s p o l l e n fo u n d i n f a e c a l s a m p le s

a r e d e s c r i b e d h e r e . They a r e d i v i d e d i n t o t h r e e g r o u p s

i . I t e m s o f unknown o r i g i n .

i i . I t e m s o f p l a n t o r i g i n . ,

i i i . I t e m s o f a n i m a l o r i g i n .

i . I t e m s o f unknown o r i g i n .✓

A morphous m a t t e r was p r e s e n t i n 17 (^5*9%) o f

t h e s a m p le s b u t o i l d r o p l e t s w ere o n l y fo u n d i n tw o , b o t h s u b a d u l t s .

i i . I t e m s o f p l a n t o r i g i n .

F r a g m e n t s o f p l a n t t i s s u e , i n c l u d i n g s m a l l g r o u p s

o f c e l l s an d v a s c u l a r t i s s u e w e re fo u n d i n 11 ( 2 9 . 7 %) o f t h e

s a m p le s ; t h r e e o f them a l s o ^ c o n t a i n e d e p i d e r m a l t i s s u e w i t h

n u m ero us s t o m a t a i n e a c h p i e c e .

A few f u n g a l s p o r e s w e re fo u n d i n two s a m p l e s .

The s to m a c h c o n t e n t s o f one a d u l t f e m a le c a u g h t on 1 i|- March 1972

a t R a n ta u P a n j a n g i n c l u d e d a few D u r io S pp . g r a i n s b u t t h e m a j o r i t y

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

172

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

f r u i t i n g b o d y ( F i g . /+? ) .

i i i . I t e m s o f a n i m a l o r i g i n .

S m a l l a r t h r o p o d s ( m i t e s , A c a r i n a an d t h r i p s ,

T h y s a n o p t e r a ) w e re fo u n d i n f o u r (10 .8% ) s a m p le s an d i n s e c t s c a l e s

i n two s a m p le s (5 * ^ % ). No o t h e r e v i d e n c e t h a t t h e b a t s h a d i n g e s t e d

a n y a n i m a l s was f o u n d .

Mammalian h a i r , a l m o s t c e r t a i n l y t h a t o f E . s p e l a e a

i t s e l f , w a s , f o u n d i n 21 (56 .7% ) o f t h e s a m p l e s . T h i s was p r o b a b l y

i n g e s t e d by t h e b a t s e i t h e r w h i l e g ro o m in g o r c l e a n i n g p o l l e n

from t h e i r f u r .

No e v i d e n c e o f an y n e m a t o d e s , c e S t o d e s o r o t h e r

p a r a s i t e s o f t h e a l i m e n t a r y c a n a l w e re fo u nd i n t h e f a e c a l s a m p l e s .

1 0 . F l i g h t .

1 0 . i . M o r p h o l o g i c a l p r o p e r t i e s r e l a t e d t o f l i g h t .

The b o d i e s o f E o n y c t e r i s s p e l a e a a r e s t r e a m l i n e d

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

t h e s k i n .

D a ta on w ing d i m e n s i o n s a r e g i v e n i n T a b l e s 12

an d 13 • They a r e b a s e d on m e a s u re m e n ts from 25 a d u l t m a le s and

27 a d u l t n o n - p a r o u s f e m a l e s . No d i f f e r e n c e s b e tw e e n m a le s and

f e m a le s w ere fo u n d i n t h e l i n e a r m e a s u re m e n ts u s e d t o c a l c u l a t e

t h e i n d i c e s r e c o r d e d i n t h e t a b l e s b u t n o n - p a r o u s a d u l t f e m a le s

w e ig h l e s s t h a n a d u l t m a l e s . T h e r e f o r e , d a t a from a l l t h e b a t s

h a v e b e e n co m b in ed t o c a l c u l a t e t h e i n d i c e s g i v e n on T a b le 12

w h ic h a r e i n d e p e n d e n t o f w e i g h t b u t m a le s and f e m a l e s a r e t r e a t e d

1 73

s e p a r a t e l y on T a b le 13 w h e re t h e I n d i c e s r e c o r d e d t h e r e a r e

r e l a t e d t o w e i g h t .

Compared t o o t h e r b a t s , i n c l u d i n g some p t e r o p i d s ,

s t u d i e d by F i n d l e y , S t u d i e r and W il so n ( 1 9 7 2 ) , E . s p e l a e a h a s

s h o r t , n a r r o w w in g s w i t h a a v e r a g e w ing t i p t o f o r e a r m r a t i o .

C o n s e q u e n t l y i t h a s a h i g h w i n g l e a d i n g i n d e x a nd a low w in g

l e n g t h i n d e x . T h e se a r e c h a r a c t e r i s t i c o f f a s t f l i g h t . ( F i n d l e y ,

S t u d i e r an d W i l s o n , 1 9 7 2 ) .

B e c a u s e n o n - p a r o u s a d u l t f e m a le s w e ig h l e s s t h a n

a d u l t m a le s t h e y h a v e a lo w e r w ing l o a d i n g i n d e x an d a h i g h e r

w in g l e n g t h i n d e x t h a n a d u l t m a l e s b u t a s w e i g h t i n c r e a s e s d u r i n g

p r e g n a n c y u n t i l f e m a l e s n e a r i n g p a r t u r i t i o n w e ig h a b o u t t h e same

as" a d u l t m a le s t h e , v a l u e s o f t h e s e two i n d i c e s c h a n g e d u r i n g

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

t o t h e v a l u e s f o r a d u l t m a l e s .

D e t a i l s o f t h e w'ing m e a s u re m e n ts o f y oung b a t s

a r e g i v e n on p . 1 8 8 e t . s e q .

1 0 . i i . F l i g h t .

The f l i g h t o f E . s p e l a e a i s f a s t an d d i r e c t .

T h i s b a t i s n o t c a p a b l e o f h o v e r i n g an d d o e s n o t p e n e t r a t e t h i c k

v e g e t a t i o n s u c h a s d e n s e b a n a n a g r o v e s .

^ P h o t o g r a p h s t a k e n on t h e U n i v e r s i t y o f M a lay a

campus o f E . s p e l a e a f e e d i n g from f l o w e r s o f P ro x y lu m in d i c u m

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

t h e f l o w e r s , ( o . i n d i c u m i s a s m a l l t r e e . One o r two l a r g e " t r u m p e t ! '

s h a p e d m aroon an d w h i t e f l o w e r s op en e a c h n i g h t on a r a c h i s t h a t

h o l d s t h e i n f l o r e s c e n c e w e l l away from t h e v e g e t a t i o n ( F i g s . 1 6

17b

and 1 7 .

B a t s a p p r o a c h e d a f l o w e r h o r i z o n t a l l y b u t 0 , 3 u

t o 0®5 in b e lo w i t u n t i l t h e y w ere a b o u t 0 . 5 w from i t ( F i g . 6 6 a

when t h e y sw ooped u p w a r d s , r a i s i n g t h e i r w in g s o v e r t h e i r b a c k s

( F i g . 6 6 b ) . When " s t a n d i n g " n e a r l y v e r t i c a l l y i n t h e a i r and

l e v e l w i t h t h e c o r o l l a m o u th , a p o w e r f u l " d o w n s t r o k e " o f t h e

w in g s b r a k e d t h e f o r w a r d m o t io n ( F i g . 6 6 c ) e n a b l i n g t h e b a t s

t o make a s o f t l a n d i n g i n a p o s t u r e s u i t a b l e f o r h o l d i n g o n to

t h e f l o w e r ( w i t h t h e thumb c la w s a n c h o r e d i n t h e c o r o l l a t u b e

and t h e f e e t g r i p p i n g t h e w r i n k l e d p e t a l s ) an d f e e d i n g ( F i g . 6 6 d ) .

When d e p a r t i n g t h e y r o l l e d b a c k w a r d s o f f t h e

f l o w e r ( F i g . 6 6 e ) i n i t i a l l y f a l l i n g h e a d f i r s t an d t h e n l e v e l l i n g

o u t an d f l y i n g away ( F i g . 6 6 f ) .

O th e r f l o w e r s . The f l o w e r s p e c i e s t h a t E . s p e l a e a

w e re s e e n v i s i t i n g a r e l i s t e d i n T a b le 3 3 , The b a t s a p p r o a c h e d

' a l l t h e s e f l o w e r s f rom b e lo w , sw o o p in g up a t . t h e l a s t m i n u te a n d ,

on d e p a r t u r e , t h e y d r o p p e d a l i t t l e b e f o r e f l y i n g aw ay . The ■

p o s i t i o n and t h e m o r p h o lo g y o f t h e f l o w e r s o f some o f t h e s e p l a n t s

e f f e c t i v e l y p r e v e n t s an y o t h e r mode o f a p p r o a c h , e . g . Musa s p p .

i n w h ic h t h e l a r g e s u b t e n d i n g b r a c t w ou ld p r o h i b i t a h o r i z o n t a l

a p p r o a c h o r D u r io z i b e t h i n u s w h e re t h e p o s i t i o n o f t h e f l o w e r s

u n d e r t h e b r a n c h e s ( F i g . 69 ) w ou ld h a v e t h e same e f f e c t .

1 1 . D i s t a n c e s t r a v e l l e d by E . s p e l a e a f o r f o o d .

1 1 . i . E v id e n c e from r i n g i n g r e c o v e r i e s .

No r e c o v e r i e s o f b a t s r i n g e d d u r i n g t h e p r e s e n t

s t u d y g i v e an y e v i d e n c e a s t o t h e f e e d i n g r a n g e o f t h i s s p e c i e s .

Fig. 66* A series of photographs to show the approach by Bonycteris srelaea to Oroxylum indicum flowers. In a,d and e the camera was level with the inflorescence. In b,c and f it was underneath the inflorescence.(See also following page.)

a. Swooping up to the inflorescence.

Raising its wings over its back.

Braking stroke prior to landing.

F ig . 66 continued

Feeding from the flower. Note position of thumb claws and feet.

Throwing itself backwards from the flower.

Falling after leaving the flower (so as to gather speed?) prior to flying away.

175

H ow ever, t h r e e h a t s r i n g e d by Dr-. A . J .B e c k and Lim Boo L i a t

- in B a tu C aves w e re r e c a p t u r e d d u r i n g t h e p r e s e n t s t u d y ; two .58

km away a t R a n ta u P a n j a n g and one 23 km away a t S e r d a n g . Two

o t h e r r e c o v e r i e s h a v e been- r e p o r t e d , one from K u a la Lumpur and

one from B a tu C av es ( T a b l e 3 7 , ) . B eck and Lim (1973) r e p o r t e d

n o r e c a p l u r e s a w a y from- B a l u C a v e s . T h e i n t e r v a l s b e t w e e n r i n g i n g

and r e c o v e r y a r e l o n g an d t h e r e i s no e v i d e n c e t h a t t h e b a t s

w e re s t i l l r o o s t i n g i n B a tu C aves a t t h e t im e o f r e c a p t u r e

( e x c e p t f o r one r e c a p t u r e d t h e r e ) .

11 . i i . E v id e n c e from p o l l e n i n t h e g uan o from B a t u C a v e s .

The s to m a c h s an d a l i m e n t a r y c a n a l s o f E . s p e l a e a

v c a u g h t w h i l e f o r a g i n g a t n i g h t a nd t h e r e f o r e w i t h fo o d i n t h e

s to m a c h w ere em pty w i t h i n 2i+ h o u r s i f t h e b a t s w e re k e p t i n

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

l e a v i n g B a tu C aves i n t h e e v e n i n g h a d em pty s t o m a c h s b u t u s u a l l y

h a d some fo o d i n t h e lo w e r g u t . T h i s w ou ld u n d o u b t e d l y b e v o id e d

d u r i n g t h e n i g h t . T h e r e f o r e , i t c a n b e a ssu m ed t h a t m o s t i f n o t

a l l i n d i g e s t a b l e m a t t e r w o u ld h a v e b e e n e x p e l l e d a s f a e c e s w i t h i n

24 h o u r s o f i t s i n g e s t i o n . ( T h i s i s e n t i r e l y i n k e e p i n g w i t h w h a t

i s known f o r o t h e r m e g a c h i r o p t e r a n s w h ic h a r e w e l l known f o r th e , t

r a p i d i t y w i t h w h ic h fo o d p a s s e s t h r o u g h t h e a l i m e n t a r y c a n a l

( e . g . W a lk e r , 1 9 6 4 ) . T h e r e f o r e a l m o s t a l l t h e p o l l e n i n t h e

g uan o c o l l e c t e d on t h e t r a y s i n B a tu C aves w ou ld h a v e b e e n

i n g e s t e d w i t h i n 2 4 h o u r s o f i t s d e p o s i t i o n , i . e . t h e p r e v i o u s

n i g h t .

I f . t h e o r i g i n o f t h e p o l l e n on t h e t r a y s i s known

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

Table 37.

The table shows details of recovery records for Eonycteris snelaea ringed by A.J.Beck and lim Eoo Liat in Batu Caves, Selangor between 1965 and 1968.

Ring No.Time between capture and recapture

RecaptureLocality

Distance in km Source

050-13297 6 months Kuala Lumpur 1 9 km MAPS

060-1033*f 60 months Rantau Panjang 38 km ANS

060-11171+ 12 months Batu Caves - MAPS

060-11 6i*7 57 months Serdang 23 km ANS

060-25809 52 months * Rantau Panjang 38 km ANS

Note :- ANS = recaptured during this study,MAPS = recapture reported by the Migratory Animals

Pathological Survey Head office, Bangkok, Thailand by courtesy of Dr. H. McClure.

176

r e t u r n e d , f rom t h e r e t o B a tu C aves t h e same n i g h t . I t i s

. p a r t i c u l a r l y f o r t u n a t e t h a t S o n n e r a t i a p o l l e n c a n "be i d e n t i f i e d

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

b e a c c u r a t e l y m apped . S . a l b a p o l l e n , i n , p a r t i c u l a r , was some­

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

c o l l e c t e d one week i t made up 42 .6% o f t h e p o m s u 011 r r a y x ,

96.8% on T ra y I I , 8 0 . 4 % on T ra y I I I an d 87.0% on T ra y IV ( s a m p l e s

333 - 3 3 6 , A p p en d ix if ) . The d i s t r i b u t i o n o f S . a l b a and S .

c a s e o l a r i s i n S e l a n g o r and t h e r e l a t i v e p o s i t i o n o f B a tu C aves

a r e shown i n F i g . 67 . The c l o s e s t p o i n t t o B a tu C av es a t w h ich

S. a l b a g ro w s i s a t R a n ta u P a n j a n g , 38 km aw ay . Thus E o n y c t e r i s

s p e l a e a a f t e r f e e d i n g on S . a l b a m u s t f l y a minimum o f 38 km

b a c k t o B a tu C aves t h e same n i g h t . H ow ever , S . a l b a a l s o o c c u r s

a l o n g 53 km o f c o a s t l i n e from C a re y I s l a n d t o K u a la S e l a n g o r .

T h i s i s a l l w i t h i n 50 km o f B a tu C aves and p r o b a b l y w i t h i n t h e

f e e d i n g r a n g e o f E . s p e l a e a from B a tu C a v e s .

12. Activity times at Rantau Panjang and elsewhere.

The majority of Eonycteris spelaea le f t Batu

Caves between 19.00 and 1 9 .4 5 hrs each night. At the F . S . C . ,

U lu Gombak (9 km from B a tu C av es) t h e y w ere f i r s t s e e n f e e d i n g

on F l o w e r i n g D u r io z i b e t h i n u s t r e e s a t a b o u t 2 0 . 0 0 h r s and w e re

c a u g h t t h e r e i n m i s t n e t s from 2 0 . 1 5 h r s . A t R a n ta u P a n j a n g

(3 8 km from B a tu C av es) o n l y two o u t o f 206 c a p t u r e d b a t s w e re

c a u g h t b e f o r e 2 0 . 1 5 h r s (o n e a t 1 9 .3 0 h r s an d t h e o t h e r a t 19«35

h r s , s u g g e s t i n g t h a t t h e y h ad r o o s t e d l o c a l l y ) . T h i r t e e n w ere

c a u g h t b e tw e e n 2 0 . 1 5 a r d 2 0 . 4 5 h r s b u t from 2 0 . 4 5 h r s t h e y w ere

F i g . 67 M a p of S e l a n g o r s h o w i n g t h e d i s t r i b u t i o n of

S o n n e r a t i a s p p . in r e l a t i o n to B a t u C a v e s .

38 km f r o m >

Bat u / C av ey

5 0 km f rom >

Batu / C aves/

K u a l a S e l a n g o r

G o m b a k

B a t uC a v e s

B u k i tL a n j a n g

K u a l a ^ L u m p u r

R a n t a uP a n j a n g

&&& J Batu Tig a 1

S e r d a n g

a r e ys t a n d

S. a l b a

S. c a s e o l a r i sk m

177

c a u g h t r e g u l a r l y \ w i t h a p e a k i n t h e e a r l y e v e n i n g b e tw e e n 2 1 . 1 5

-and 2 2 , 1 5 h r s u n t i l 0 1 , 1 5 h r s when num bers d ro p p e d o f f . None

w e re c a u g h t a f t e r 0 3* 25 h r s ( T a b l e 3 8 and F i g . 68 ) . T h e re i s

a t im e l a g o f a b o u t one h o u r b e tw e e n t h e t im e b a t s b e g a n t o

em erge from B a tu C aves ( i 8 » 5 0 t o 19*00 h r s ) a n d t h e t i m e t h e y

w ere f i r s t c a u g h t a t U lu Gombak and o f a b o u t 1-j h o u r s b e f o r e

t h e y w e re f i r s t c a u g h t a t R a n ta u P a n j a n g , The f l i g h t s p e e d o f

s p e l a e a i s n o t known b u t i t c e r t a i n l y e x c e e d s 26 km p e r h o u r

w h ic h -w o u ld b e r e q u i r e d o f b a t s i f t h e y w e re t r a v e l f ro m B a tu

C a v e s - t o R a n ta u P a n j a n g i n 1 h o u r s .

I f t h e b a t s a t t h e F . S . C . , U lu Gombak and a t

R a n ta u P a n j a n g d o ' come from B a tu C aves (n o o t h e r r o o s t s w e re

l o c a t e d , c e r t a i n l y no o t h e r l a b g e r o o s t s o c c u r i n S e l a n g o r ) i t

m ig h t b e e x p e c t e d t h a t t h e y s o m e t im e s f e e d en r o u t e .

E v id e n c e t h a t some E . s p e l a e a c a u g h t a t R a n ta u

P a n j a n g h a d t r a v e l l e d t h e r e from some d i s t a n c e i n l a n d a n d f e d

en r o u t e came from s i x a d u l t s t h a t w ere c a u g h t a t R a n ta u P a n j a n g

w i t h a h e a v y d u s t i n g o f P a r k i a s p p . . p o l l e n on t h e i r b o d i e s and

m em bran es . T h re e o f t h o s e w ere c a u g h t t o g e t h e r a t 2 0 .2 0 h r s on

8 December 1 9 7 2 ; one o f them a l s o had a P a r k i a s p e c i o s a l e a f l e t

a d h e r i n g t o a s t i c k y p a t c h on t h e t i p o f one w in g . (The s t i c k y

p a t c h was p r o b a b l y r e s i n from a t r e e . ) T h e re was no P . s p e c i o s a

g ro w in g a t R a n ta u P a n j a n g n o r a l o n g t h i s p o r t i o n o f t h e c o a s t .

T hese b a t s m u s t , t h e r e f o r e , h av e come to R a n ta u P a n j a n g from

i n l a n d . The c l o s e s t p o i n t a t w h ich i t i s known t o grow i s i n

t h e Meru F o r e s t R e s e r v e ( F i g . 2 ) b u t i t a l s o p l e n t i f u l i n

m o s t o t h e r f o r e s t r e s e r v e s i n S e l a n g o r .

T a b le 3 8 . The t a b l e show s t h e t o t a l , mean an d W i l l i a m s 1 mean

f o r t h e num ber o f E o n y c t e r i s s p e l a e a c a u g h t p e r

h o u r ( b u t e x c l u d i n g t h o s e c a u g h t i n o r n e a r c a v e

r o o s t s ) i n P s n i n s u l s r Ms.ls.ysia.*

F i g . 6 8 . The f i g u r e i l l u s t r a t e s g r a p h i c a l l y t h e d a t a f o r

W i l l i a m s 1 mean r e c o r d e d i n T a b l e 3 8 .

Time No. o f n i g h t s

No. o f males

No. o f females T o ta l T o t a l / h r V7 i 1 l i ams1

mean

19.00-19 .1 5 50 0 0 0 0 0

1 9 .1 5 -2 0 .1 5 50 0 2 2 0 .0 4 0 .120.15-21 .1 48 15 21 54 0.71 0 .421 .1 5 -2 2 .1 5 42 14 51 65 1 .54 1 .12 2 . 1 5 - 2 5 . 1 5 51 1 2 25 55 1 .15 0 . 72 3 .1 5 -0 0 .1 5 18 9 15 22 1 . 2 2 0 . 800.15-01 .1 5 11 4 9 15 1 . 1 8 0 . 801 .1 5 - 02 .1 5 9 5 2 5 0 .5 5 0 .40 2 .1 5 -0 3 .1 5 5 0 4 4 0 . 8 0 0 . 6

0 3 . 1 5 - 0 4 . 1 5 4 0 1 1 0 . 2 5 0 .20 4 .1 5 - 05 .1 5 2 0 0 0 0 005.1 5-06.1 5 2 0 0 0 0 0

0-7

S ° ' 6 "05<De

0-5-03ec6

•HrHiH■H& 0-3

Si:

LAi a la

O ' fA

ir\ l aLA LA LA

O ' LAOJ

Tine

178

13* R i n g i n g r e s u l t s . . '

R e t r a p , d a t a f o r E o n y c t e r i s s p e l a e a r i n g e d d u r i n g

t h e s t u d y i s l i m i t e d a s o n l y 1? i n d i v i d u a l s w e re r e t r a p p e d .

E i g h t o f them w e re c a u g h t t h r e e t i m e s i n one n i g h t an d t e n on

two o r more n i g h t s i n c l u d i n g one w h ic h was c a u g h t t w i c e i n one

-1- x - - -1 :_____x.1, _* j. 7 m _ 1- -1 _ -7 \ mi- _ -i . a. _ _-i~i a g i i i ttu u u a . cuxu untsx* u.Lg,nb v.j.ctuxt? ) * xnt? ucxi/cx

two t e n d e h c i e s ^ ^ ^ '

■ F i r s t l y , t h e r e c a p t u r e o f h a t s two o r t h r e e /

t i m e s . i n a n i g h t a t R a n ta u P a n j a n g and K a ja n g i n d i c a t e s t h a t

some E . s p e l a e a r e m a in i n t h e same a r e a f o r S e v e r a l h o u r s

d u r i n g t h e n i g h t , r e t u r n i n g t o t h e same p l a c e s f ro m t im e t o

t i m e . The t im e i n t e r v a l b e tw e e n s u c h r e c a p t u r e s v a r i e d , e . g .

one b a t was c a u g h t t h r e e t i m e s ' i n 55 m i n u te s w i t h i n t e r v a l s o f

1 5 and 40 m i n u t e s b e tw e e n e a c h c a p t u r e w h i l e a n o t h e r was

r e c a p t u r e d 3 h o u r s 2 5 m i n u t e s a f t e r b e i n g r i n g e d .

S e c o n d l y , t h e r e c a p t u r e o f b a t s a t t h e same

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

t o o r r e m a in i n t h e same a r e a s f o r some d a y s , s o m e t im e s o v e r

l o n g p e r i o d s , e . g . one b a t r e c a p t u r e d a t R a n ta u P a n j a n g 140 d ay s

" a f t e r i t was r i n g e d t h e r e and a n o t h e r was c a u g h t f o u r t i m e s o v e r

89 d a y s a t t h e F o S . C . , U lu Gombak. H ow ev er , t h e m o s t r e m a r k a b l e

f e a t u r e o f t h e s e r e c o r d s i s t h a t a t R a n ta u P a n j a n g (w h e re m i s t

n e t s w ere s e t a t 15 d i f f e r e n t s i t e s ) s i x o f t h e s e v e n b a t s c a u g h t

on. two d i f f e r e n t n i g h t s w e re r e c a p t u r e d i n t h e same n e t t h a t

t h e y h a d o r i g i n a l l y b e e n c a u g h t i n ( T a b l e 39 ) • T h i s s u g g e s t s

t h a t t h e b a t s h a d becom e f a m i l i a r w i t h t h e o f t e n f r e q u e n t e d

f e e d i n g a r e a s an d f o l l o w r e g u l a r r o u t e s i n t h o s e a r e a s .

Table 39 .The t a b l e r e c o r d s th e i n t e r v a l between c a p t u r e and

r e c a p t u r e and th e l o c a l i t i t e s a t which th e y were caugh t f o r a l l E o n v c te r i s s n e l a e a t h a t were r i n g e d and r e c a p t u r e d d u r in g th e s tu d y

Bat No. Sex/Age L o c a l i t i e s Date Time i n t e r v a l1 •? Ad. F .S .C . 22 J a n .

3 Feb. 1 2 F e b .

197219721972 V s days} * days

J

2 ? Ad. R .P .#2 10 Feb. 14 Feb.

19721972 | 4 Days

3 ? Ad. R . P . #6 10 Mar. 4 May

19721972 | 55 days

4 •? Ad. R . P . #6 10 Mar.it

2 June

1972

1972V 2 h r s .^|84 days .

5 ? Ad. R . P . #8 27 Mar. 9 May

19721972 | 43 days

6 ? Ad. R . P . #9 6 Apr. n

1972 j- 1 . 2 5 h r s .

7 ? Ad. R .P .#11 #12

6 Aug.ii

1972 | 0 .5 0 h r s . J8 ? Ad. R . P . #11 6 Aug.

n1972 | 2 .4 5 h r s .

9 ? Ad. R . P . #11 6 Aug. 9 Aug.

19721972

/J 3 days

10 ? Ad. R.P .#9 6 S e p t ••

. 1972 | 0 .3 0 h r s

11 ? Sad . R .P .#4 10 Oct . 1 Dec.

19721972 j 52 days

12 (? Ad. F .S .C . 17 J a n . 13 Feb.

9 Apr. 15 A pr .

1972197219721972

] 1 7 days')166 days > 89 days

6 days j

13 8* Ad. R .P .#8 27 Mar.Y!

1972 | 2 .3 5 h r s .

14 $ Ad. R .P .#9 6 Apr.ii

ii

1972/

j 1 .50 h r s .J

15 d* Ad. F .S .C . 9 Apr. 29 June

19721972 j 81 days

16 d* Ad. R . P . #10 11 J u l y 28 Nov.

19721972 |l 40 days

17 cf Ad. R . P . #15 1 S e p t ,itii

1972| 0 . 5 5 h r s .

N o t e : - R .p . = Rantau P a n jan g , F .S .C . = F i e l d S t u d i e s C e n t r e , Ulu Gorabak; # = Net s i t e number ( s e e F ig . 3 ) .

179

1 4 . ' F o r a g i n g . _ .

1 4 . i * F l o c k s . ■ . . .

Eonycteris spelaea- foraged in flocks. It. was not

p o s s i b l e to- a c c u r a t e l y c o u n t t h e number o f b a t s i n a n y f l o c k

b u t e s t i m a t e s o f t h e num bers v a r i e d from f i v e t o t w e n t y ,

o c c a s i o n a l l y t h e r e w e re t h o u g h t t o be a s many a s f i f t y b a t s i n

a flock. Pertinent observations were made at the F.S.C., Ulu

Gombak on 14 A p r i l s 973 when t h e moon was n e a r l y f u l l . A

f l o w e r i n g d u r i a n t r e e ( D u r io z i b e t h i n u s ) was k e p t u n d e r o b s e r v a t i o n

from d u sk u n t i l 2 3 .0 0 h r s . B i n o c u l a r s (x 8 m a g n i f i c a t i o n ) w e re

u s e d t o w a tc h t h e b a t s . To t h e s o u t h t h e r e was a c l e a r i n g g i v i n g

an open v ie w o f t h e s k y f o r s e v e r a l h u n d re d m e t r e s down t h e

Gombak v a l l e y . E l s e w h e r e t h e v iew was b l o c k e d b y t r e e s . T h e re

w e re o t h e r f l o w e r i n g d u r i a n t r e e s i n t h e n e i g h b o u r h o o d . ■

' The f i r s t flock of E. spelaea reached the tree

a t 2 0 . 0 5 h r s . A t l e a s t s i x d i f f e r e n t f l o c k s v i s i t e d i t d u r i n g/

t h e n e x t 55 m i n u t e s a n d f i v e b e tw e e n 21 .0 0 and 2 2 . 0 0 h r s b u t

t h e num ber d r o p p e d t o t h r e e f l o c k s b e tw e e n 2 2 . 0 0 a n d 2 3 . 0 0 h r s .

T h e re w e re u s u a l l y b e tw e e n t e n an d tw e n ty b a t s i n e a c h f l o c k .

They w ere c l e a r l y s i l h o u e t t e d a g a i n s t t h e sk y a s t h e y a p p r o a c h e d

t h e t r e e fro m t h e s o u t h . On r e a c h i n g t h e t r e e t h e co m p ac t u n i t

b r o k e up a s i n d i v i d u a l s c i r c l e d i t , a l i g h t i n g e v e r y now and t h e n

t o f e e d ( F i g . 6 9 ) . The b a t s r e m a in e d on one i n f l o r e s c e n c e o n l y

a s h o r t t im e ( f i v e t o t e n s e c o n d s ) b u t o c c a s i o n a l l y s c r a m b le d

a l o n g t h e b r a n c h fro m one i n f l o r e s c e n c e t o t h e n e x t . When t h i s

h a p p e n e d t h e y s o m e t im e s w e re s e t t l e d f o r up t o a m i n u te b e f o r e

f l y i n g aw ay . On e a c h i n f l o r e s c e n c e t h e y w e re s e e n r a p i d l y p r o b i n g

one f l o w e r a f t e r a n o t h e r .

F i g . 69* E o n y c t e r i s s p e l a e a f e e d in g from th e f lo w e r s o f Durio z i b e t h i n u s .Note t h e p o l l e n on th e b a t ’ s head and th e way th e s t ig m a (S ) and a n t h e r s come i n c o n t a c t w i th th e b a t ’s abdomen, wings and h ead .

1 So

The t im e t h a t e a c h f l o c k s t a y e d a t t h e t r e e

v a r i e d from f i v e t o tw e n ty m i n u t e s . No f l o c k s w e re o b s e r v e d

r e f o r m i n g o r d e p a r t i n g . A p p a r e n t l y t h e y d i d n o t i m m e d i a t e l y f l y

s o u t h a g a i n . O c c a s i o n a l l y two f l o c k s w ou ld o v e r l a p an d t h e r e

w o u ld h e a s many a s t h i r t y o r more h a t s a ro u n d t h e t r e e a t t h e

same tx m e . T h e se f l o c k s a p p a r e n t l y d i d n o t u n i t e f o r , i n v a r i a b l y ,

t h e r e was a su d d e n d r o p i n t h e num ber o f b a t s p r e s e n t i n d i c a t i n g

t h a t some h a d d e p a r t e d . I t i s n o t known i f t h e r e was a n y i n t e r ­

c h a n g e o f i n d i v i d u a l s b e tv /e e n t h e two f l o c k s when t h e y m et i n

t h i s ■w ay .

D e t a i l e d o b s e r v a t i o n s w ere a l s o made o f E . s p e l a e a

v i s i t i n g O roxylum in d i c u m f l o w e r s on t h e U n i v e r s i t y o f M a la y a

campus b e tw e e n 5 an d 19 A u g u s t 1973* T h e re w ere two t r e e s g ro w in g

c l o s e t o g e t h e r i n t h e cam p u s . D u r in g t h e p e r i o d o f o b s e r v a t i o n

o ne t r e e h a d two i n f l o r e s c e n c e s and t h e o t h e r f i v e u n t i l one o f

t h e f i v e b r o k e o f f on 1^ A u g u s t . E ach i n f l o r e scen .ce h a d two

f l o w e r s p e r n i g h t e x c e p t one w h ich h ad t h r e e f l o w e r s o n c e . R e c o rd s

w e re k e p t o f t h e num ber o f b a t s v i s i t n g e i t h e r f l o w e r on one

i n f l o r e s c e n c e , w h ic h ( u n l e s s t h e moon was b r i g h t ) was i l l u m i n a t e d

b y a t o r c h w i t h w eak b a t t e r i e s , p l a c e d on t h e g ro u n d w i t h t h e

beam s h i n i n g on t h e f l o w e r s . P h o t o g r a p h s w e re a l s o t a k e n ( F i g s . 66

a t o f ) . (A d e s c r i p t i o n o f t h e mode o f a p p r o a c h t o an d d e p a r t u r e

from t h e f l o w e r s was g i v e n on p . 174*)

U n l i k e t h e Gombak v a l l e y s i t u a t i o n d e s c r i b e d

a b o v e , d i s c r e e t f l o c k s w ere n o t o b s e r v e d a p p r o a c h i n g and c i r c l i n g

t h e t r e e s b u t v i s i t s t o t h e f l o w e r s came i n w aves a s members o f

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

( F i g . 7 0 ) ,

F ig . 7 0 .The f i g u r e shows th e number o f E o n y c te r i s soe loe o

and t h e t ime t h a t th e y landed on one Oroxylum indlcum i n f l o r e s c e n c e . The o b s e r v a t i o n s were made on th e U n i v e r s i t y o f Malaya campus between 5 and 19 August 1973.

5 August 1973 14 August 1973 19 Augus t 1 973

2 0 .OOhrs .10 .2 0 .30 .40 .50

2 1 .OOhrs . 10 .2 0 .30 .40 .50

22 .OOhrs .10 .20 .30 .40 .50

2 3 . OOhrs . 10 .2 0 .30 .40 .50

2 4 . OOhrs .10 .2 0 .30 .40 .50

0 1 .OOhrs .10 .2 0 .30 .40 .50

0 2 . OOhrs

8

::::: 9

11 43

55

iia 11

NOOBSERVATIONS

1

17

13

1 30

1

5 )NO

OBSERVATIONS

NOOBSERVATIONS

rJ)

] 8

15

J18

T o ta l h r s . T o ta l b a t s B a t s / h r .

59218.4

5499 .8

5448.8

1 81

S i m i l a r w aves w e re r e c o r d e d v i s i t i n g S o n n e r a t i a

a l b a a t R a n ta u P a n j a n g , e . g . on 15 J u n e 197.5 f o u r f l o w e r s on

one t r e e i n a g r o v e o f f l o w e r i n g _S_. a l b a w e re w a tc h e d from 1 9 .3 0

t o 2 2 .3 0 h r s . B e tw een 2 1 . 2 0 and 2 1 . ^ 0 h r s t h e f l o w e r s w e re v i s i t e d

29 t i m e s b y E . s p e l a e a ; t h e r e w e re no o t h e r v i s i t s b y b a t s

i d e n t i f i e d a s E . s p e l a e a d u r i n g t h e t im e o f o b s e r v a t i o n . S i m i l a r

w aves w e re f r e q u e n t l y s e e n w h i l e m i s t - n e t t i n g a l o n g t h e dyke

b y t h e m a n g ro v es a t R a n ta u P a n j a n g b u t p r e c i s e r e c o r d s o f t h e

number o f v i s i t s b y b a t s t o i n d i v i d u a l f l o w e r s w e re n o t k e p t .

Waves o f E . s p e l a e a w e re a l s o s e e n f e e d i n g from f l o w e r i n g P a r k i a

s p e c i o s a , Bombax v a l e t o n i i an d D uabanga g r a n d i f l o r a b u t d e t a i l e d

n o t e s o f t h e num ber o f b a t s and t i m e s w ere n o t k e p t .

1 ^ - . i i . C o m p o s i t i o n o f f l o c k s .

I t w as n o t p o s s i b l e t o c a t c h c o m p l e t e f e e d i n g

f l o c k s o f E o n y c t e r i s s p e l a e a from w h ic h t o d e t e r m i n e t h e i r age

an d s e x c l a s s c o m p o s i t i o n . H o w ever , some e v i d e n c e on t h i s t o p i c

v/as o b t a i n e d a t R a n ta u P a n j a n g .

A t R a n ta u P a n j a n g t h i s s p e c i e s was o c c a s i o n a l l y

c a u g h t one a t a t im e b u t u s u a l l y two o r m ore w ere n e t t e d a t t h e

same t i m e . T h ese w e re members o f f l o c k s f e e d i n g from e i t h e r

c u l t i v a t e d b a n a n a s o r S o n n e r a t i a s p p . f l o w e r s an d w h ic h w ere

t h e r e f o r e f l y i n g n e a r g ro u n d l e v e l . Among t h e s e s m a l l g r o u p s

t h e r e w ere e x a m p le s o f a s s o c i a t i o n s c o m p r i s i n g a l l a d u l t m a l e s ,

a l l a d u l t f e m a l e s , a d u l t m a le s w i t h a d u l t f e m a l e s a nd o f s u b ­

a d u l t s w i t h a l l o f t h e a b o v e .

182

P a r t i a l s e g r e g a t i o n o f a g e an d s e x c l a s s e s o f E . s p e l a e a

when f e e d i n g .

a ) E v id e n c e from m i s t n e t t i n g d a t a .

The p e r c e n t a g e o f a d u l t m a le , a d u l t f e m a le and

s u b a d u l t , E o n y c t e r i s s p e l a e a ( s e x e s co m bined b u t e x c l u d i n g

s u c k l i n g yo ung a t t a c h e d t o m o t h e r s ) f o r 2 ,6 7 0 E . s p e l a e a c a u g h t

i n B a tu C aves w e re 3 6.4%> 39 .7% and 23.8% r e s p e c t i v e l y ■( f i g u r e s

c a l c u l a t e d f ro m d a t a g i v e n b y B eck an d Lim, 1 9 7 3 ) . H o w ever , t h e

e q u i v a l e n t r a t i o s f o r E . s p e l a e a c a u g h t a t n i g h t o u t s i d e t h e

** c a v e s d i f f e r m a r k e d ly from t h e s e f i g u r e s ( F i g . 71 a nd T a b le kO)

I n t h e two c o a s t a l l o c a l i t i e s , R a n ta u P a n j a n g

an d K u a la L a n g a t , t h e r e w ere p r o p o r t i o n a l l y more a d u l t f e m a le s

and fe w e r a d u l t m a l e s a n d s u b a d u l t s t h a n i n B a tu C a v e s . The

d i f f e r e n c e s i n t h e a d u l t s e x r a t i o com pared t o B a tu C aves was

s i g n i f i c a n t a t R a n ta u P a n j a n g ( p < 0 , 0 0 1 ) b u t n o t a t K u a la L a n g a t

w h e re t h e s a m p le s i z e was v e r y s m a l l (p = 0 . 3 ) . The d i f f e r e n c e

. i n t h e a d u l t t o s u b a d u l t r a t i o co m pared t o B a tu C aves was n o t

s i g n i f i c a n t a t R a n ta u P a n j a n g (p < 0 . 2 > 0 . 1 ) b u t was so a t

K u a la L a n g a t ( p < 0 .01 > 0 . 0 0 1 ) .

A t t h e two i n l a n d l o c a l i t i e s , B u k i t L a n ja n g

and t h e F . S . C . , U lu Gombak s u b a d u l t s o u tn u m b e re d a d u l t s ( a t t h e

F . S . C . , U lu Gombak n e a r l y s i x t i m e s a s many s u b a d u l t s a s a d u l t

f e m a l e s w e re c a u g h t ) . The r a t i o o f s u b a d u l t s t o a d u l t s d i f f e r e d

s i g n i f i c a n t l y from t h e same r a t i o i n B a tu C aves ( F . S . C . , Ulu

Gombak p < 0 . 0 0 1 ; B u k i t L a n ja n g p < 0 . 0 0 1 ) .

A t t h e F . S . C . , U lu Gombak t w i c e a s i ^ n y a d u l t

m a le s a s a d u l t f e m a le s w ere c a u g h t . H ow ever , t h e s a m p le i s v e r y

71. M a p s h o w i n g % a d u l t m a l e , % a d u l t f e m a l e a n d % s u b a d u l t E. s p e l a e a c a u g h t a t 5 loca l i t i e s

3 U i u G o m b a k

B a t u -- Caves "

o 16 B u k i t ^ ) ? 33 L a n j a n g * ^ ; o v e r

Kuala^Lumpur ,

R a n t a uP a n j a n g S e r d a n

K u a l aL a n g a t

Table 4 0 .

The t a b l e shov/s th e p e r c e n ta g e of a d u l t male , a d u l t

female and s u b a d u l t E o n y c te r i s s p e l a e a caugh t i n Batu Caves

and a t fo u r l o c a l i t i e s o u t s i d e Batu Caves.

L o c a l i t y T o t a l Ad. d* Ad. ? T o t a l Ad. Sad .d* + ?

■ #Batu Caves

Ulu Gombak

RantauPanjang

KualaLangat

B u k i tLanjang

**2670

53

206

32

74

973(36 .4 )

13 (24 .5 )

46 (2 2 .3 )

1 2 (3 7 .5 )

1 2 ( 1 6 . 2 )

1060(39 .7)

6(11 .3)

120(58 .3)

19 (59 .4 )

24 (32 .4 )

2033(76 .1 )

19 (3 5 .8 )

1 6 6 ( 8 0 . 6 )

3 1 (9 6 .9 )

3 6 (4 8 .6 )

637(23 .8)

34 (6 4 .1 )

40 (1 9 .4 )

1 (3 .1 )

38(51 .4)

«d a t a from Beck and Lira (1973) ,

* *e x c lu d in g 371 s u c k l i n g young a t t a c h e d to t h e i r m o th e rs ,

• * *d a t a e x t r a c t e d from th e c o l l e c t i o n r e g i s t e r o f th e I n s t i t u t e o f M edica l R e se rac h , Kuala Lumpur.

183

s m a l l and t h e d i f f e r e n c e from t h e n e a r l y 1:1 r a t i o i n B a tu C aves

i s n o t s i g n i f i c a n t ( p < 0 .1 > 0 . 0 5 ) . A t B u k i t L a n ja n g (w h ich

l i e s on a l i n e from B a tu C aves t o t h e c o a s t i n t h e r e g i o n o f

R a n ta u P a n j a n g ) t h e r e w e re t w i c e a s many a d u l t f e m a l e s a s m a l e s .

A g a in , t h e s a m p le s i z e i s s m a l l and t h e s e x r a t i o d o e s n o t d i f f e r

-Li — j>.« i i . ^ o. -n _ j— ~ ~ f -« r \ v_ r \ r \ rz \O X 5 U 1 x x u c i u o - L , y i i u u i u n c x u \ j x u c i o u v a v c o ^ j j kj • i W t t / y y •

The d a t a shows t h a t more a d u l t f e m a le s t h a n

a d u l t m a le s o r s u b a d u l t s f e e d a l o n g t h e c o a s t an d t h a t m os t

s u b a d u l t s r e m a in i n t h e v i c i n i t y o f B a tu C a v e s . I t a l s o s u g g e s t s

t h a t a d u l t m a l e s f e e d i n t h e h i l l s more t h a n a d u l t f e m a le s d o .

The h i g h e r p r o p o r t i o n o f a d u l t f e m a le s s u g g e s t e d a t B u k i t '

L a n j a n g may r e s u l t f rom a d u l t f e m a l e s f e e d i n g i n t h e f o r e s t s

en r o u t e t o t h e c o a s t s .

( D a ta from K u a la L a n g a t and B u k i t L a n ja n g w ere

e x t r a c t e d fro m t h e IMR c o l l e c t i o n r e g i s t e r . A t K u a la L a n g a t t h ei

b a t s w e re n e t t e d c l o s e t o t h e g ro u n d b u t i n B u k i t L a n ja n g t h e

n e t s i n w h ic h Eh s p e l a e a w e re c a u g h t w e re s e t on a e r i a l w alkw ays

i n o r o v e r t h e c a n o p y o f t h e f o r e s t (Lira Boo L i a t , p e r s . comm.)

b . E v id e n c e from p o l l e n a n a l y s i s .

B e tw een 21 an d 26 F e b r u a r y 1973 ( d u r i n g a f l o w e r i n g

f l u s h o f S o n n e r a t i a a l b a a t R a n ta u P a n ja n g ) 83 E o n y c t e r i s s p e l a e a

v/ere c a u g h t an d e t h e r i s e d a s t h e y em erged from t h e mouth o f

C av e rn C, B a tu C a v e s , i n t h e e v e n i n g . They i n c l u d e d 21 a d u l t

m a l e s , 27 a d u l t f e m a le s an d 35 s u b a d u l t s ( s e x e s c o m b i n e d ) . Any

m a t t e r i n t h e a l i m e n t a r y c a n a l v/as ex am in ed f o r p o l l e n .

P o l l e n was fo u n d i n 19 a d u l t m a l e s , 22 a d u l t

f e m a le s and 22 s u b a d u l t s . The s to m a c h s o f some s u b a d u l t s a l s o

184

c o n t a i n e d a w h i t e s u b s t a n c e p re s u m e d t o b e m i l k ( s e v e n o f t h e

s m a l l e r o n e s c o n t a i n e d m i lk b u t no p o l l e n ) » . The a l i m e n t a r y c a n a l s

o f t h e r e m a i n i n g 2 a d u l t m a l e s , 5 a d u l t f e m a le s an d 6 s u b a d u l t s

w ere em p ty .

The r e s u l t s o f t h e p o l l e n a n a l y s i s a r e g i v e n

*1 n m - i CP no ~— - -* -o » f *— *

A d u l t m a l e s . 19 b a t s c o n t a i n e d p o l l e n , D uabanga g r a n d l f l o r a

was t h e m o s t a b u n d a n t p o l l e n t y p e p r e s e n t . I t was t h e o n ly s p e c i e s

i n 5 ( 2 6 .0 % ) o f t h e b a t s an d fo rm ed 75% o r more o f t h e p o l l e n i n

12 (63%) o f th e m . I t was a b s e n t from o n l y 2 (1 1 % ). S o n n e r a t i a

( a l l s p e c i e s co m b in e d ) w as n o t a b u n d a n t . I t was t h e o n ly p o l l e n

t y p e i n 1 (5%) o f t h e s a m p le s and fo rm e d 75% o r m ore o f t h e

p o l l e n i n 2 (11%) o f th e m . I t was a b s e n t from 8 (42%) and p r e s e n t

i n s m a l l q u a n t i t i e s i n t h e r e m a i n d e r .

A r t o c a r p u s s p p . made up 53% an d 86% r e s p e c t i v e l y

o f t h e p o l l e n i n two s a m p l e s . Musa s p p , made up 5«5% i n on e

s a m p l e . P a r k i a s p p . , E u g e n ia s p p . a n d C e ib a p e n t a n d r a w e re p r e s e n t

i n s m a l l q u a n t i t i e s ( < 1 . 0 % ) ' i n s e v e r a l s a m p l e s .

A d u l t f e m a l e s . 22 b a t s c o n t a i n e d p o l l e n . S o n n e r a t i a ( a l l s p e c i e s

co m b ined ) was t h e m o s t a b u n d a n t p o l l e n t y p e . I t was t h e o n ly

t y p e i n 8 (36%) o f t h e s a m p le s and fo rm ed 75% o r more o f t h e

p o l l e n i n 14 (64%) o f th e m . I t was a b s e n t f rom o n l y 2 (9 % ).

Duabanga g r a n d i f l o r a was t h e o n l y p o l l e n t y p e i n 2 (9%) o f t h e

s a m p le s and fo rm ed 7 5 % o r more o f t h e p o l l e n i n 5 ( 2 3 %) o f th em .

I t was a b s e n t f rom 11 (50 % ).

O th e r p o l l e n t y p e s p r e s e n t i n t h e s a m p le s i n c l u d e d

C e ib a p e n t a n d r a . Bombax v a l e t o n i i , E u g e n ia s p p . , P a r k i a s p p .

an d A r t o c a r p u s s p p . They w ere o n l y p r e s e n t i n v e r y s m a l l

/

F i g . ? 2 . H i s t o g r a m s s h o w in g t h e p r o p o r t i o n s o f p o l l e n o f

S o n n e r a t i a s p p . , D uabanga g r a n d l f l o r a and o t h e r

t y p e s i n t h e a l i m e n t a r y c a n a l s o f E o n y c t e r i s s p e l a e a

c a u g h t e m e r g in g from B a tu C a v e s , S e l a n g o r b e tw e e n

21 an d 26 F e b r u a r y 1973*

oI—

S’ S iN>

N u m b e r o f b a t s

3!L ptoro 05

_ _ L _oo

— Lroi 05

100

—I

1007 5 - 9 95 0 - 7 42 5 - 4 9

0 - 2 4 05

3 +0« >ro clro .

ro_ u

05_L

00II

mO33<D

</i-a-a

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7 5 - 9 95 0 - 7 42 5 - 4 9

0 - 2 4

oo:

1007 5 - 9 95 0 - 7 42 5 - 4 9

0 - 2 40 m

3"

(D

185

q u a n t i t i e s ( 3%) "but one s a m p le c o n t a i n e d 10 .5% Musa s p p .

p o l l e n .

S u b a d u l t s ( s e x e s c o m b in e d ) . 22 b a t s c o n t a i n e d p o l l e n . D uabanga

g r a n d i f l o r a was p r e s e n t i n a l l o f th em . I t made up 75% o r more

o f t h e p o l l e n p r e s e n t i n 21 (95%) o f t h e b a t s b u t fo rm ed 100%

i n o n l y 3 (14%) o f th em . I t was a b s e n t from 1 (5% )• S o n n e r a t i a

s p p . was a b s e n t f rom o n l y 6 (27 .3% ) o f t h e s a m p le s b u t p r e s e n t

i n 16 (72 ,7 % ) o f th em . I n two b a t s t h a t w ere n e a r l y f u l l grown

i t made up 1 5"»0% and 1 8 . 8 % r e s p e c t i v e l y o f t h e p o l l e n p r e s e n t .

Among t h e r e m a i n i n g 14 i t was p r e s e n t i n s m a l l q u a n t i t i e s m ak ing

up b e tw e e n < 0 . 0 1 % and 7*5% (mean 2 . 2 5 % ) ' o f t h e p o l l e n p r e s e n t

i n t h e b a t s . Thus m o s t s a m p le s from s u b a d u l t s c o n t a i n e d 95% o r

m ors D. g r a n d i f l o r a p o l l e n and 5% o r l e s s S o n n e r a t i a s p p , p o l l e n .

15* R e p r o d u c t i o n i n E . s p e l a e a .

15*3-* M a l e s .

The t e s t e s o f a l l m a t u r e m a le s w e re l a r g e ,

m e a s u r in g up t o 13 mm i n l e n g t h b y 10 mm i n w i d t h . They w ere

c o n t a i n e d i n a s c r o t u m . M a tu re m a le s w e re c a u g h t i n e v e r y month

o f t h e y e a r . S p e c im e n s w e re n o t c o l l e c t e d r e g u l a r l y f o r d i s s e c t i o n

b u t d u r i n g t h e c o u r s e o f o t h e r w ork t h e e p i d i d y m i s o f 1 6 w ere

exam ined.. A l l c o n t a i n e d s p e r m a t o z o a . Male E . s p e l a e a a p p a r e n t l y

r e m a in s e x u a l l y a c t i v e t h r o u g h o u t t h e i r a d u l t l i f e .

1 5 . i i » F e m a l e s .1

P r e g n a n t an d l a c t a t i n g a d u l t f e m a l e s w e re c a u g h t

t h r o u g h o u t t h e y e a r ( T a b l e 4 l ) b u t t h e d a ta ! i s j i n s u f f i c i e n t

t o show w h e th e r a n y p e a k s o c c u r i n e i t h e r c a t e g o r y d u r i n g t h e

Table 41 .

The t a b l e r e c o r d s th e r e p r o d u c t i v e s t a t u s o f a l l

t h e a d u l t female E o n y c te r i s s p e l a e a caugh t i n P e n i n s u l a r

M a la y s ia each month d u r in g th e s tu d y , i . e . from F eb ru a ry

1972 to J a n u a r y 1974.

J F M A M J J A S 0 N D T o t a l

e a r l y p r - - 1 2 - 1 2 2 - - 2 3 13

1 /3 p r 1 5 1 2 12 2 - 1 - - 2 1 27

2 /3 p r 1 6 2 1 7 1 1 1 1 1 2 2 26

3 / 3 p r 2 7 1 - 4 2 1 3 1 - 1 - 22

T o t a l p r 4 18 5 5 23 6 4 7 2 1 7 6 88

L 1 15 6 6 3 3 8 16 2 1 4 5 70

NLNP - 8 13 4 3 2 1 2 1 - 4 - 38

T o t a l 5 41 24 15 29 11 13 25 5 2 15 11 1 85*

* The t o t a l i s 11 l e s s th an th e sum o f t h e s u b t o t a l s becau se

e le v e n b a t s t h a t were b o th p r e g n a n t and l a c t a t i n g have

been r e c o rd e d tw ic e ; once i n each r e l e v a n t column.

Note p r = p r e g n a n t ,

L = l a c t a t i n g ,

NLNP = n o t l a c t a t i n g and p o s s i b l y n o t p r e g n a n t (no

embryo d e t e c t e d by p a l p a t i o n ) .

186

(

1y e a r . No d a t a was o b t a i n e d t o c o n f i r m t h e g e s t a t i o n o r l a c t a t i o n

t i m e s r e p o r t e d b y B eck and Lim (1 9 7 3 ) b u t c o n c u r r e n t s t a t e s o f

l a c t a t i o n and e a r l y p r e g n a n c y ( d e t e r m i n e d by p a l p a t i o n ) w ere

r e c o r d e d e l e v e n t i m e s i n r i n g e d b a t s a nd c o n f i r m e d b y d i s s e c t i o n

i n t h r e e m o r e . I n s e v e n d i s s e c t e d f e m a l e s , e m b r y o s w e re fo u n d i n

"til© 3? x "b w t © 2? i i i © « N o rx s wcx*© £ 0x2.1*. ci x n t l i c l o f t w c 37x11^ i i o r i i #

The IMR R e g i s t e r (B ooks 9 and 1 6 ) r e c o r d em bryos i n t h e r i g h t

u t e r i n e h o r n i n 3h f e m a le s b u t no n e i n t h e l e f t u t e r i n e h o r n , v ■

1 5 « i v . C a r r y i n g o f y o u n g . -

Of 21 lactating female Eonycteris spelaea caught

l e a v i n g Qua S a n d in g and B a tu C aves 17 (80 .9% ) w e re c a r r y i n g

j u v e n i l e s t h a t v a r i e d i n h e i g h t f rom 8 . 5 g ( a b o u t b i r t h w e i g h t )\

t o 2 0 . 0 g ( a p p r o x i m a t e l y t h e w e i g h t o f s u b a d u l t s a t f i r s t f l i g h t ) .k

H o w ever , of/ 51 l a c t a t i n g f e m a l e s c a u g h t away from t h e c a v e s a t

n i g h t , o n l y 5 (9*8%) c a r r i e d j u v e n i l e s . They w e ig h e d from 9 . 0

t o 1 0 . 5 g . The d i f f e r e n c e i n p r o p o r t i o n o f l a c t a t i n g f e m a le s

c a r r y i n g j u v e n i l e s a s t h e y l e f t t h e c a v e s an d c a r r y i n g j u v e n i l e s

i n t h e f e e d i n g a r e a s i s s i g n i f i c a n t (p <£ 0 . 0 0 1 ) . _

The f i v e j u v e n i l e s c a u g h t w i t h t h e i r m o t h e r s

i n t h e f e e d i n g a r e a s w e re a l l e s t i m a t e d t o b e l e s s t h a n a week

o l d . One was s t i l l w e t from a m n i o t i c f l u i d a t 2 ^ . 0 0 h r s an d m u s t

h a v e b e e n b o r n n e a r b y ; i . e . o u t s i d e t h e r o o s t . (When r e c a u g h t

a t 0 2 .0 0 h r s i t s f u r had d r i e d ) . The f u r o f a n o t h e r was d r y b u t

i t s u m b i l i c a l c o r d h ad n o t d r i e d . The u m b i l i c a l c o r d o f a t h i r d

one was d r y b u t h a d n o t d ro p p e d o f f . The u m b i l i c a l c o r d s o f t h e

l a s t two h a d d rm pped o f f b u t b o t h j u v e n i l e s w e ig h e d o n l y 10 g ,

i . e . 1 t o 1 . 5 g m ore t h a n t h e n e o n a t e s w i t h w e t u m b i l i c a l c o r d s

187

and 0.5 g less than the one with the dry umbilical cord in s i tu .

These observations suggest that the juveniles

may be born .outside the roost and that for the f i r s t few days

afte r the b irth they are carried by their mothers even when the

l a t te r are foraging. However, when the juveniles are a l i t t l e

older their mothers carry them out of the caves in the evening

but deposit them elsewhere while foraging.

15 » v . Mode of carrying young.

.Juvenile bats were suspended from one side of

the mother's abdomen. They were attached firmly by the mouth

to one of the mother's nipples (which, as in a l l bats, are

pectoral) and by the feet; one on each side of the mother's

abdomen'just anterior to her pelvis. The forelimbs- were not

essential for their attachment to the mother but there was a

strong tendency, especially in very young bats, to press the

wrists together thus gripping anything between thenu By placing

one wrist in the mother's ax illary region nearest the nipple

to which i t was attached and the other on the opposite side of

her chest, they obtained additional purchase and, probably,

s ta b i l i ty .

When adult females were caught carrying juveniles

the la t te r were removed for weighing-. When removed,the juveniles

showed a d is tinctive behavioural pattern : This consisted of

straddling the observer's finger (or any other object) v/ith their

feet while gripping i t between their wrists as they searched

with their mouths for any object they could grasp. They would

firmly clamp their jaws on any object of similar size to the

188

m o t h e r ' s n i p p l e . O f t e n t h e i r m ou ths h a d t o h e p r i s e d open by

i n s e r t i n g t h e p o i n t s o f a p a i r o f s c i s s o r s b e tw e e n t h e ja w s

and o p e n in g them i n o r d e r t o rem ove t h e young from t h e m o t h e r ' s

n i p p l e s ( o r a s u b s t i t u t e o b j e c t when t r y i n g t o r e t u r n them to

t h e m o th e r ! ) .

T h e se r e s p o n s e s w e re s t r o n g e s t ' i n t h e new b o r n

and v e r y young b a t s { w h ic h w e re e a s i l y r e t u r n e d t o t h e i r m o t h e r s

a f t e r w e i g h i n g ) . Among o l d e r j u v e n i l e s ( a s ju d g e d b y t h e i r

h e a v i e r w e i g h t ) t h e s e t e n d e n c i e s d e c l i n e d u n t i l y oung w e i g h in g

b e tw e e n 1 8 a n d 20 g ( a n d a l m o s t c a p a b l e o f f l i g h t ) showed v e r y

l i t t l e t e n d e n c y t o h o l d on t o t h e i r m o th e r a f t e r b e i n g d i s t u r b e d .

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

m o t h e r s a f t e r t h e y h a d b e e n rem oved f o r w e i g h i n g .

Hp .'WL. Growth o f y o u n g .\

No d a t a w e re o b t a i n e d t o show t h e a c t u a l g ro w th

r a t e s o f young E . s p e l a e a . H ow ever , m e a s u re m e n ts o f t h e v /e ig h t

an d t h e w ing d i m e n s i o n s r e v e a l some i n t e r e s t i n g f e a t u r e s . N e o n a te s

h a v e a f o r e a r m l e n g t h o f 22} t o 2 5 mm (n = 3 ) an d w e ig h 8 . 5 t o

9 . 0 g (n = 3 ) . A d u l t m a le s h a v e a f o r e a r m l e n g t h o f 60 t o 70 mm

(mean 6 6 . 6 + 5 . 8 , n = 9 2 ) an d w e ig h 2}6 t o 67 g (m ean 5 8 .2 5 +

2}.5, n = 9 3 ) . A d u l t f e m a le s a r e s m a l l e r w i t h a f o r e a r m l e n g t h

o f 58 t o 69 mm (mean 62}.73 + 2 . 2 , n = 1 7 2 ) . The v / e ig h t o f n o n ­

p r e g n a n t t o 1 / 3 p r e g n a n t f e m a le s v a r i e s from 2}0 t o 59 g (mean

4 9 . 9 7 + 4 . 0 , n = 130) b u t t h o s e n e a r i n g p a r t u r i t i o n a r e h e a v i e r ,

w e ig h in g 57 t o 67 g (mean 6 2 .1 5 + 2 . 9 , n = 2 0 ) .

Two young b a t s w i t h f o r e a r m l e n g t h s o f 4 8 . 5 and

49*5 mm w ere u n a b l e t o f l y b u t f o u r w i t h f o r e a r m l e n g t h s be tv /e en

1 89

5 0 . 5 t o 51 »0 mm c o u l d j u s t f l y . a n d t a t s w i t h a f o r e a r m l e n g t h

o f 53 mm f l e w s t r o n g l y (3 h a t s w i t h f o r e a r m l e n g t h s o f 53*0 mm

w e re c a u g h t 7 km from B a tu C aves u n d e r a f l o w e r i n g d u r i a n t r e e

( D u r io z l b e t h i n u s ) a t t h e F . S . C . , U lu Gombak). T h e r e f o r e , f i r s t

f l i g h t t a k e s p l a c e when t h e b a t s h a v e a f o r e a r m l e n g t h o f a b o u t

vnrn T'Vi rt*V> V* c* 4*vr ^ ^ ^ O l /•»-» r» ■!* 4* •£**) •? 4-

I n F i g . 73 r e g r e s s i o n s h a v e b e e n p l o t t e d f o r

t h e f o r e a r m l e n g t h a g a i n s t w e i g h t o f young E . s p e l a e a . The

r e g r e s s i o n s w e re draw n s e p a r a t e l y f o r j u v e n i l e s ( i n c a p a b l e o f

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

(p = 0 .0 0 1 ) b u t t h e i r s l o p e s a r e s i g n i f i c a n t l y d i f f e r e n t (p =

0 . 0 0 1 ) . The w e i g h t s o f s u b a d u l t s j u s t c a p a b l e o f f l i g h t ( a b o u t

20 g) i s 3 l+% o f t h e mean a d u l t m a le w e i g h t and 40% o f t h e mean

n o n - p r e g n a n t / e a r l y p r e g n a n t a d u l t - f e m a l e w e i g h t w h i l e t h e f o r e ­

arm; l e n g t h o f t h e same s u b a d u l t s ( a b o u t 50 mm) i s 70% o f t h e

mean a d u l t f o r e a r m l e n g t h . Thus from b i r t h t o f i r s t f l i g h t t h e

g ro w th r a t e o f t h e f o r e a r m com pared t o t h e r a t e o f w e i g h t i n c r e a s e

i s r e m a r k a b l y r a p i d b u t a f t e r f i r s t f l i g h t t h e g ro w th r a t e o f

t h e f o r e a r m d e c r e a s e s m a r k e d ly co m p ared t o t h e r a t e o f w e ig h t

i n c r e a s e so t h a t b o t h f o r e a r m and w e i g h t r e a c h a d u l t d im e n s i o n s

a t a b o u t t h e same t i m e .

I n F i g . 74 t h e w in g l o a d i n g i n d e x h a s b e e n p l o t t e d

a g a i n s t w e i g h t f o r young E . s p e l a e a . A l i n e a r r e g r e s s i o n l i n e

h a s b e e n f i t t e d t o t h e p o r t i o n o f t h e g r a p h from f i r s t f l i g h t

t o m a t u r i t y . ( I t i s a s t r a i g h t l i n e , p = 0 . 0 0 1 ) b u t i n s u f f i c i e n t

d a t a a r e a v a i l a b l e t o d raw a r e g r e s s i o n l i n e from b i r t h t o f i r s t

f l i g h t . A t b i r t h t h e w in g a r e a i s v e r y s m a l l an d c o n s e q u e n t l y

Fig. 73* Graph showing the forearm length of young

Eonycteris spelaea plotted against their weight.

Subadults (young capable of free fligh t) and

juveniles (bats too young to fly) have been

treated separately. Regression lines have been

f i t ted to the data of both age groups.

(

Fo

re

ar

m

m

m

7 0

6 0

50

4 0

'▲ A

O A d u l t s ,

A S u b a d u l t s , f l y i n g

A J u v e n i l e s , n o n - f l y i n g

m e a n3 0

- L6 0

1 I20 3 0 45

W e i g h t ( g r a m s )

F i g . 7*f. G raph s h o w in g t h e v/ing l o a d i n g i n d e x o f young

E o n y c t e r i s s p e l a e a p l o t t e d a g a i n s t t h e i r w e i g h t .

J u v e n i l e s ( b a t s t o o young t o f l y ) and s u b a d u l t s

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

a nd a r e g r e s s i o n l i n e h a s b e e n f i t t e d t o t h e

d a t a f o r s u b a d u l t s .

Win

g

load

ing

in

de

x0-2 7 s-

0-2 5

0-231-

A

0 - 2 1

0-19

0-17

0-15

0-13

0-51o

15

O Of

O O o O A d u l t s ,

O S u b a d u l t s , f I y i n g

A J u v e n i l e s , n o n - f l y i n g

m e a n

X

3 0 . . . 45W e i g h t i n g r a m s

6 0

190

t h e w i n g ' l o a d i n g i n d e x i s h ig h , . H o w ever , t h e w in g 'g r o w s v e r y

r a p i d l y com p ared t o th e ' w e i g h t u n t i l f i r s t f l i g h t . T h i s r e s u l t s

i n a d e c r e a s e i n t h e l o a d i n d e x u n t i l f i r s t f l i g h t a f t e r w h ich

i t a g a i n i n c r e a s e s b e c a u s e t h e g ro w th r a t e o f t h e w in g d e c r e a s e s

com pared t o t h e w e i g h t i n c r e a s e . -Thus f i r s t f l i g h t c o i n c i d e s

w i t h t h e l o w e s t w in g l o a d i n g i n d e x e x p e r i e n c e d b y t h e b a t s ,

S i n c e t h e w in g l o a d i n g i n d e x i s d i r e c t l y p r o p o r t i o n a l t o t h e

t r u e w ing l o a d i n g ( F i n d l e y , S t u d i e r and W i l s o n , 1 9 7 2 ) , f i r s t

f l i g h t c o i n c i d e s w i t h t h e l o w e s t - w i n g l o a d i n g e x p e r i e n c e d byI

t h e b a t s .

191

DISCUSSION.

'ECOLOGICAL DISTRIBUTION.

1 . E c o l o g i c a l d i s t r i b u t i o n o f M a c r o g lo s s u s s p p .

B a t s o f t h e g e n u s M a c r o g lo s s u s w e re f o r a l o n g

t im e c o n s i d e r e d uncommon i n M a l a y s i a ( e . g . C h a s e n , 1 9 35 ; Lim •

and Heyneman., 1 9 68 ; Medway, 1 965 )* H ow ever , w i t h t h e u s e o f m i s t

n e t s r e c e n t c o l l e c t o r s h av e o b t a i n e d more s p e c i m e n s ( e . g . Lim,

1 9 6 6 , 1973) an d t h e y b e g a n t o b e d e s c r i b e d a s common (L im , 1973;

Medway, 196 9 )* N e v e r t h e l e s s , k n o w le d g e o f t h e i r h a b i t a t r e q u i r e ­

m e n ts r e m a in e d l i t t l e known. C h asen (1 9 3 5 ) comm ented u n d e r t h e

h e a d i n g o f "M. l a g o c h i l u s l a g o c h i l u s " (= M. m in im us h e r e ) b u t

r e f e r i i i g t o t h e g e n u s a s a w h o le t h a t " t h e y a p p e a r t o b.e p a r t i a l

t o m angro ve sw am ps" . Lim (1 9 6 6 , 1973) r e c o r d e d b o t h s p e c i e s i n

m a n g r o v e s , l o w la n d f o r e s t and m o n tan e f o r e s t up t o 6 6 0 0 f t .

E x a m i n a t i o n o f t h e s p e c i m e n s on w h ich he b a s e d h i s r e c o r d s

r e v e a l e d t h a t many h a d b e e n m i s i d e n t i f i e d and h e ( p e r s , comm.)

a d m i t t e d d i f f i c u l t y i n s e p a r a t i n g t h e s p e c i e s . I n f a c t , a l l h i s

s p e c im e n s o f M. m in im us a r e from c o a s t a l l o c a l i t i e s and M. s o b r i n u s

from i n l a n d l o c a l i t i e s . Medway (1 9 6 9 ) r e c o r d e d M. s o b r i n u s f rom

f o r e s t i n t h e h i l l s and M. m in im u s a s common i n t h e m a n g ro v es

b u t a l s o p r e s e n t i n l a n d : The l a s t s t a t e m e n t i s p r o b a b l y b a s e d

on m i s i d e n t i f i e d s p e c i m e n s .

E x a m i n a t i o n o f h u n d r e d s o f museum s p e c i m e n s from

many l o c a l i t i e s an d e x t e n s i v e m i s t n e t t i n g d u r i n g t h e p r e s e n t

s t u d y c l e a r l y show ed t h a t i n P e n i n s u l a r M a l a y s i a b o t h s p e c i e s

a r e common. M. m in im us i s a c o a s t a l s p e c i e s a s s o c i a t e d w i t h

m a n g r o v e s . I t i s a l s o fo u n d on many o f f s h o r e i s l a n d s ( F i g . i+0) .

192

M. sobrinus- occurs in suitable habitats throughout the country,

from coastal regions, but not mangroves, to the tops of mountains

over 2000 m a . s . l . I t i s very strongly associated with the v/ild

banana species, Musa malaccensis and M. ,truncata. This association

explains many-apparent anomalies in i t s re la tive abundance in

g.g. no Macroglossus spp. have been caught

in the primary forest reserve at Pasoh, Pahang (D.R'.Wells, pers.

comm.) although many other bats have been mist-netted there as

part of an I.E.P. project. On Gunong Benom, Medway (1 972b) reported

catching only two specimens of M. sobrinus, both near base camp,

while on.Gunong Brinchang Lim (1973) reported catching 74 specimens

of Macroglossus and regarded i t as common there. (Re-examination

of his material showed a l l to be M. sobrinus.) On Gunong Benom

a l l the s i te s at which bats were mist-netted except base camp

were in primary forest (Medway, (1972a).Wild bananas do not grow

in primary forest except on land s lips and sometimes on river

banks. However, near base camp the forest had been disturbed

and in the photograph of the camp s i te (Medway,1 972a, Plate 2)

banana plants are clearly v is ib le . On Gunong Brinchang, especially

alongside the road, the forest has been disturbed and Musa truncata

i s extremely common (pers. obs.).

During the present study many f ru it le s s nights

were spent with mist nets set in primary or secondary forest

i before the association of M. sobrinus with some wild bananas was

realised. Thereafter, wherever mist nets were set near flowering

plants of these banana species, even in the otherwise biologically

s te r i le conditions prevailing in rubber esta tes , M.sobrinus was

usually caught.

193

E lse w h e re . , t h e d i s t r i b u t i o n o f l o c a l i t y r e c o r d s

i n S u m a t r a and J a v a (A p p e n d ix 1 ) s u g g e s t t h a t t h e r e a l s o

M* s o b r i n u s o c c u r s i n l a n d w h i l e M. m in im us i s a c o a s t a l s p e c i e s *

L o c a l i t y r e c o r d s f o r M_. s o b r i n u s i n T h a i l a n d ( H i l l and T h o n g lo n g y a ,

1 9 7 2 ) , S o u th V ie tn a m (v a n P e e n e n , 1968 ) and L aos ( P h i l l i p s ,

196?) a r e a i l from i n l a n d w h i l e t h e o n ly r e c o r d o f M. m in im us

from T h a i l a n d ( a s p e c im e n ex am in ed i n t h e S i n g a p o r e N a t i o n a l '

C o l l e c t i o n ) i s c o a s t a l f rom Nakom S r i Tamm arat i n P e n i n s u l a r

T h a i l a n d a s a r e two s p e c im e n s exam in ed from Thu D ue, n e a r S a i g o n ,

S o u th V ie tn a m an d one from P o u lo (= P u la u ) C ondor o f f t h e S o u th

V ie tn a m e s e c o a s t * The l a t t e r t h r e e s p e c im e n s a r e now d e p o s i t e d

i n t h e R o y a l O n t a r i o Museum. M’. m in im us h a s n o t p r e v i o u s l y b e e n

r e c o r d e d f rom V ie tn a m .

H o w ev er , i n E o rn e o w h e re M. s o b r i n u s i s a b s e n t ,

H* Minimus i s fo u n d b o t h i n t h e m a n g ro v es ( a s a t Bako N a t i o n a l

P a r k ) and i n l a n d ( H a r r i s s o n , 1966 ; Lira and H eyneman, 1 9 6 8 ; Medway,

1965; T an , 1 9 6 5 ) . H i l l ( 1 9 6 6 ) r e c o r d e d a s p e c im e n f rom 3 , 5 0 0 f t

a . s . l . i n t h e K e l a b i t H ig h l a n d s o f n o r t h e a s t e r n S a ra w a k . (S p e c im e n s

from i n l a n d d i s t r i c t s i n K a l im a n ta n h a v e a l s o b e e n ex am in e d ( F i g .

M ) . N o th in g i s known o f t h e i n l a n d h a b i t a t r e q u i r e m e n t s o f

t h i s s p e c i e s i n B o r n e o .

A s i t u a t i o n s i m i l a r t o t h a t i n B o rn e o o c c u r s i n

t h e P h i l i p p i n e s ( s p e c i m e n s o f M. m i n i mus from many i n l a n d l o c a l i t i e s

a r e d e p o s i t e d i n t h e R o y a l O n t a r i o Museum a nd t h e N atu rm useum

un d F o r s c h u n g s - I n s t i t u t S e n c k e n b e r g , F r a n k f u r t ) and i n S u la w e s e

(= C e l e b e s ) ( s p e c i m e n s o f M. m in im us from i n l a n d l o c a l i t i e s h a v e

b e e n c o l l e c t e d b y a U n i t e d S t a t e s N a v a l R e s e a r c h U n i t NAMRU-2).

. . 194

S p e c im e n s from a l l t h e s e a r e a s h a v e b e e n ex am in ed and t h e i r

i d e n t i t y c o n f i r m e d . ' • ’ ,

Further e a s t , ' in the Key, Aru and Amboina groups

of is landsN ew Guinea, northern Australia and the Bismarck

Archipelago, M. minimus, i s sympatric with the morphologically

very similar nectarivorous genus, G y co n y cLeris. Nothing i s known

of the habitat requirements of either genus there. In Papua™

New G u in e a b o t h g e n e r a a r e r e c o r d e d from a w id e v a r i e t y o f f o r e s t

t y p e s i n c l u d i n g m a n g ro v e s (McKean, 1972) b u t S y c o n y c t e r i s i s

a p p a r e n t l y more a b u n d a n t and w id e s p r e a d , ( a n o n , 1 9 73 ; A. M acW ill iam

p e r s . c o m m .) b u t van D e u s e n ( l 9 6 l ) r e c o r d e d M.m in im us o n l y i n r a i n

f o r e s t a nd S y c o n y c t e r i s from r a i n f o r e s t and m a n g r o v e s . McKean

( o p p . c i t . ) fo u n d t h a t i n Papua-New G u in e a M. m in im us was c a u g h t

£>nly b e lo w 260 m a . s . l . b u t S . c r a s s a e x t e n d e d t o much h i g h e r

a l t i t u d e s w h i l e on B o u g a i n v i l l e I s l a n d i i i t h e Solom on I s l a n d s

M* mi'n tm us was c a u g h t up t o 1155 m a . s . l . He t e n t a t i v e l y a t t r i b u t e d

t h e d i f f e r e n c e t o t h e a b s e n c e o f S y c o n y c t e r i s from B o u g a i n v i l l e .

D e e p e r i n v e s t i g a t i o n w ou ld p r o b a b l y r e v e a l the. o c c u p a t i o n o f

d i s t i n c t n i c h e s w h e re t h e s p e c i e s a r e s y m p a t r i c .

Ecological dis tr ibution of Eonycteris spelaea.

I n P e n i n s u l a r M a l a y s i a , E o n y c t e r i s s p e l a e a i s a

common b a t r e c o r d e d r e g u l a r l y from a w id e v a r i e t y o f h a b i t a t s

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

i n c l u d i n g m a n g r o v e s . l t - i s a l s o common i n many c u l t i v a t e d a r e a s .

I t was f r e q u e n t l y c a u g h t i n t h e same m i s t n e t s a s M a c r o g lo s s u s

m in im us i n c o a s t a l d i s t r i c t s o r M. s o b r i n u s i n l a n d .

I n Borneo and in the Philippines i t is sympatric

195

w i t h Eony c t e r i s ma.jor A n d e rse n ( i n c l u d i n g E . r o b u s t a M i l l e r and

l o n g i c a u d a T a y l o r ) * T a t e (1 9 ^ 2 ) a l s o r e c o r d e d E . ma.jor from

n o r t h P a g i j M en taw i I s l a n d s , o f f t h e w e s t c o a s t o f S u m a t r a . T h e re

a r e no r e c o r d s o f E . s p e l a e a from M entaw i I s l a n d s b u t E , s p e l a e a

i s t h e o n l y s p e c i e s r e c o r d e d i n S u m a t r a ( e . g . R o b in s o n and Boden

K l o s s , i 9 i 8 ; T a t e , 19^2) P o l l e n o f D uabanga m o lu c c a n a was fo u n d

i n t h e a l i m e n t a r y c a n a l o f one p r e s e r v e d s p e c im e n o f E . ma.jor

from K u c h in g , S a ra w a k ( p e r s , o b s . ) . E . ma.jor t h e r e f o r e d o e s a t

t i m e s e a t p o l l e n b u t i t i s a l i t t l e known s p e c i e s and i t s e c o l o g i c a l

r e l a t i o n s h i p t o E . s p e l a e a i s unknow n. ■

2 , Food s o u r c e s o f m a c r o g l o s s i n e b a t s i n P e n i n s u l a r M a l a y s i a ,

A l l t h r e e s p e c i e s o f M a l a y s i a n m a c r o g l o s s i n e b a t s

a r e p r i m a r i l y n e c t a r i v o r o u s . T h i s was r e a l i s e d l o n g ago ( e . g .

A n d e r s e n , 1 9 1 2 , van d e r P i j l , 1936) b u t a l m o s t n o t h i n g h a s b e e n

r e p o r t e d on t h e i r d i e t s i n M a l a y s i a . B e rw ic k (1 9 5 0 ) p u b l i s h e d

a p h o t o g r a p h o f E o n y c t e r i s s p e l a e a ( m i s i d e n t i f i e d a s Cynopt e r u s

b r a c h y o t i s ) f e e d i n g from t h e f l o w e r s o f C e ib a p e n t a n d r a . Lim (

1 9 6 5 j 1973) and Tan (1 9 6 5 ) r e p o r t e d p o l l e n i n t h e a l i m e n t a r y

c a n a l s o f M a c r o g l o s s u s s p p . U n o r i g i n a l r e f e r e n c e s t o f l o w e r s t h a t

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

v a r i o u s . - - F lo r a s a nd b o o k s on M a l a y s i a n b o t a n y , e ' . g . B a c k e r and

v an S t e e n i s ( 1 95^f) » C o rn e r (1 9 5 2 , i n c l u d i n g some o r i g i n a l

o b s e r v a t i o n s and s u g g e s t i o n s ) , H o l t t u m (19 6 5 ) an d W h itm ore ( 1 9 7 2 ) .

( H o l t t u m ( l 965) c l e a r l y a l l o w e d h i s i m a g i n a t i o n c o n s i d e r a b l e

f r ee d o m when h e d e s c r i b e d b a t s v i s i t i n g b a n a n a f l o w e r s f o r he

s t a t e s t h a t t h e y h a n g from t h e o p en ed b r a c t s b y t h e c la w s on

t h e i r w in g s .H e may h a v e o b s e r v e d t h e p u n c t u r e s made b y t h e c la w s

1 96

o f s u n b i r d s o r s p i d e r h u n t e r s ( N e c t a r i n i i d a e ) on t h e o pen ed b r a c t s

a f t e r t h e i r e a r l y m o rn in g v i s i t s ( F i g , 12 ) f o r t h e y do h an g

from t h a t b r a c t t o p r o b e t h e f l o w e r s . Knowing some Musa s p p .

t o b e v i s i t e d b y b a t s he c o u l d h a v e assu m ed t h a t t h e c la w m a rk s

Were t h o s e o f b a t s . )

M ost o f t h e s e u n o r i g i n a l s t a t e m e n t s h a v e b e e n

d e r i v e d from t h e t h o r o u g h an d m e t i c u l o u s o b s e r v a t i o n s o f van ••

d e r P i j l (1936 , 1 941 } 1956) i n J a v a , . He i d e n t i f i e d many o f t h e

c h i r o p t e r o p h i l o u s t r e e s t h e r e , i n c l u d i n g some e x o t i c s p e c i e s

g ro w in g i n t h e B o t a n i c G a rd e n s a t B ogor an d r e v i e w e d a l l t h e

e a r l i e r o b s e r v a t i o n s made b y n a t u r a l i s t s i n J a v a . T h e se e a r l y

o b s e r v a t i o n s h a v e a l s o b e e n r e v i e w e d b y B a k e r a n d H a r r i s (1957)*

A l th o u g h van d e r P i j l ' s w ork was i n v a l u a b l e i n i d e n t i f y i n g

c h i r o p t e r o p h i l o u s t r e e s i n M a l a y s i a h e (a n d m o s t o f t h e e a r l i e r

_ o b s e r v e r s ) e m p h a s i s e d t h e b o t a n i c a l a s p e c t s o f t h e r e l a t i o n s h i p

b e tw e e n t h e f l o w e r s and t h e b a t s . Van d e r P i j l (1936 d i d d e s c r i b e

some m o r p h o l o g i c a l a d a p t a t i o n s o f M a c r o g lo s s u s an d E o n y c t e r i s

f o r a d i e t o f n e c t a r and p o l l e n an d so m e t im e s r e c o r d e d w h ic h

s p e c i e s o f b a t v i s i t e d w h ich p l a n t . I n 1956 h e p u b l i s h e d a

p h o t o g r a p h o f M. s o b r i n u s v i s i t i n g a b a n a n a f l o w e r . He su m m a r ise d

h i s ' f i n d i n g s i n F a e g r i a nd v an d e r P i j l (1966) b y d e s c r i b i n g a

" b a t f l o w e r " sy nd ro m e and a " f l o w e r b a t " s y n d r o m e . S i m i l a r

sy n d ro m e s h a v e b e e n d e v i s e d o r c o p i e d b y v a r i o u s a u t h o r s , e . g .

B a k e r (1973)> P r o c t o r an d Yeo( l973)»

The c o n c e p t o f s u c h sy n d ro m es h a s b e e n a u s e f u l

one i n t h a t i t h a s d raw n t o g e t h e r a l l t h e a v a i l a b l e i n f o r m a t i o n

i n an a t t e m p t t o u n d e r s t a n d t h e c o r e l a t i o n s h i p s b e tw e e n t h e p l a n t s

and t h e a n i m a l s . H o w ever , t h i s s t u d y shows t h a t t h e r e i s a s e r i o u s

197

d a n g e r i n a t t e m p t i n g t o draw up g e n e r a l i s a t i o n s f rom l i m i t e d

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

k n o w le d g e h a s b e e n m o s t l y o n e - s id e d , .

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

( i n c l u d i n g d i e t s , r o o s t i n g , f o r a g i n g , f l i g h t an d m o rp h o lo g y ) o f

t h e t h r e e m a c r o g l o s s i n e b a t s fo u n d i n p e n i n s u l a r M a l a y s i a and

t h e s e a r e r e l a t e d t o i m p o r t a n t d i f f e r e n c e s i n t h e p l a n t s on w h ic h

t h e two s p e c i e s o f M a c r o g lo s s u s on one hand an d J5. s p e l a e a on

t h e o t h e r h an d r e l i e d .

The p l a n t s i d e n t i f i e d i n P e n i n s u l a r M a l a y s i a a s

p r o v i d i n g fo o d f o r m a c r o g l o s s i n e b a t s a r e d i v i s i b l e i n t o two

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

Group I . The s p e c i e s i n g r o u p I a r e a l l a s e a s o n a l , f l o w e r i n g

c o n t i n o u s l y ( a l t h o u g h i n d i v i d u a l p l a n t s may n o t do s o ) . T h i s

g ro u p i n c l u d e s a l l t h e m o n o c o t y l e d o n s , i . e . Musa t r u n c a t a .

M* m a l a c c e n s i s , t h e i r h y b r i d s , c u l t i v a t e d b a n a n a s . Cocos n u c i f e r a

and A ren g a s p p . a s w e l l a s two s p e c i e s o f d i c o t y l e d o n s , i . e .

S o n n e r a t i a c a s e o l a r i s an d D uabanga g r a n d i f l o r a ( b o t h S o n n e r a t i a c e a e ) .

T hese p l a n t s ( e x c e p t A ren g a ? ) s h a r e o t h e r a t t r i b u t e s b e s i d e s

t h e i r a s e a s o n a l i t y :E a c h f l o w e r i n g s p e c im e n p r o v i d e s a r e l a t i v e l y

s m a l l num ber o f f l o w e r s e a c h n i g h t b u t d o e s so f o r a l o n g t im e

o r c o n t i n o u s l y . They a r e f r e q u e n t l y g r e g a r i o u s .

Group I I . The s p e c i e s i n g ro u p I I make up t h e m a j o r i t y o f t h e

c h i r o p t e r o p h i l o u s s p e c i e s i d e n t i f i e d d u r i n g t h e s t u d y . They

f l o w e r s e a s o n a l l y o r i n f l u s h e s , w i t h a l l p l a n t s o f one s p e c i e s

i n one a r e a f l o w e r i n g t o g e t h e r . T h i s g ro u p i n c l u d e s M a n g i f e r a

s p p . , Bomb ax v a l e t o n l l , C e ib a p e n, t a n d r a , D u r io g r a v e o l e n s , D.

z l b e t h l n u s ( a n d a ny o t h e r c h i r o p t e r o p h i l o u s s p e c i e s o f D u r i o ) ,

p r o b a b l y B a r r i n g t o n i a ( t h e s p e c i f i c i d e n t i t y of. B a r r i n g t o n i a

p o l l e n i n t h e gu ano was n o t e s t a b l i s h e d b u t m o s t s p e c i e s i n

t h i s g e n u s f l o w e r i n t e r m i t t e n t l y ) s P a r k i a j a v a n i c a , P . s p e c l o s a

( a n d p r o b a b l y P . s i n g u l a r i s and P . s u m a t r a n a ) , E u g e n ia m a l a c c e n s i s

(and p r o b a b l y o t h e r c h i r o p t e r o p h i l o u s E u g e n ia s p p . ) , S o n n e r a t i a

B. l b s. 3.23. ci S . cvs.’ts.w

B e s i d e s t h e i r p h e n o lo g y m o s t p l a n t s i n G roup I I

d i f f e r from t h o s e i n Group I i n t h a t e a c h t r e e p r o v i d e s a l a r g e

t o enorm ous q u a n t i t y o f f l o w e r s p e r n i g h t ( e x c e p t S . a l b a and

S . o v a t a ) b u t o n l y d u r i n g a s h o r t s e a s o n , A l s o , t h e y a r e u s u a l l y

n o t g r e g a r i o u s ( u n l e s s t h e y a r e c u l t i v a t e d s p e c i e s p l a n t e d c l o s e

t o g e t h e r a n d , e x c e p t f o r S ; a l b a and S . o v a t a ) b u t t h i n l y d i s p e r s e d

o v e r v e r y l a r g e a r e a s . T h i s e v en a p p l i e s t o many c u l t i v a t e d

s p e c i e s .

O roxylum i n d i c u m h a s a t t r i b u t e s o f b o t h g r o u p s :

A l th o u g h a d e f i n i t e b u t e x t e n d e d f l o w e r i n g s e a s o n was r e c o r d e d

on t h e T a p a h - R i n g l e t r o a d i t was n o t e x c e p t i o n a l t o f i n d f l o w e r i n g

p l a n t s a t an y t i m e o f t h e y e a r . Each f l o w e r i n g p l a n t r e g u l a r l y

p r o d u c e s a few f l o w e r s e a c h n i g h t f o r s e v e r a l m o n th s a t a t i m e .

I n s e v e r a l p l a c e s i t w as fo u n d t o b e g r e g a r i o u s , e . g . a t a b o u t

m i l e 13 on t h e T a p a h - R i n g l e t r o a d , 16 t r e e s fo rm e d a n e a r l y mono-

s p e c i f i c s t a n d i n a ' c l e a r i n g b e s i d e t h e r o a d , b u t m ore o f t e n s i n g l e

t r e e s w e re f o u n d .

The u n i d e n t i f i e d p o l l e n t y p e s m u s t a w a i t i d e n t i f i c a t i o n

b e f o r e t h e y c a n b e a l l o c a t e d w i t h c e r t a i n t y t o o ne g ro u p o r t h e

o t h e r , b u t two r e q u i r e com m ent. The s p e c i f i c i d e n t i t y o f

A r t o c a r p u s s p p . . p o l l e n fo u n d i n t h e guano was n o t d e t e r m i n e d :

199

Many A r t o c a r p u s s p p , f l o w e r s e a s o n a l l y , e . g A. I n t e g e r , A.

h e t e r o p h y l l u s , A. e l a s t l c u s ( p e r s . o b s» ) b u t p o l l e n was p r e s e n t

i n a l l t h e E a t u C aves and Gua S a n d in g guano s a m p le s t h r o u g h o u t

t h e y e a r . T h e re a r e two p o s s i b l e e x p l a n a t i o n s ; e i t h e r i t came

from one (.or m ore) s p e c i e s t h a t f l o w e r e d c o n t i n o u s l y o r i t came

from a s e r i e s o f s p e c i e s w h ic h f l o w e r e d . s e a s o n a l l y i n a c o n t i n o u s

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

p o s s i b l e .

S e c o n d l y , t h e n a t u r e o f t h e o c c u r r e n c e o f T8 p o l l e n

( S a p o t a c e a e ) i n t h e guan o s a m p le s from B a tu C aves s u g g e s t s t h a t

i t s h o u l d b e i n c l u d e d w i t h G roup I I p l a n t s .

G e n e r a l l y , t h e two s p e c i e s o f M a c r o g l o s s u s f e d

from Group I p l a n t s b u t from d i f f e r e n t s p e c i e s w i t h i n t h a t g ro u p

w h i l e E . s p e l a e a u s e d b o t h g r o u p s ; m o s t o f t h e p o l l e n E . s p e l a e a

i n g e s t e d came from G roup I I p l a n t s b u t when t h e y w e re s c a r c e o r

u n a v a i l a b l e t h e b a t s f e d f rom G roup I p l a n t s . M a c r o g l o s s u s s o b r i n u s

f e d m o s t l y from t h e f l o w e r s o f w i l d b a n a n a s , Musa m a l a c c e n s j s ,

M* t r u n c a t a o r t h e i r h y b r i d s . T h i s c o n f i r m e d t h e o b s e r v a t i o n s

o f v an d e r P i j l ( 1 9 3 6 , 1956) i n J a v a . He fo u n d i t f e e d i n g on

c u l t i v a t e d b a n a n a s and v a r i o u s w i l d s p e c i e s i n c l u d i n g members

o f t h e M. a c u m i n a t a s p e c i e s c o m p le x an d M. t e x t l l i s ;N u r ( q u o t e d

from P u r s g l o v e , 1972) c o n f i r m e d v a n d e r P i j l ' s o b s e r v a t i o n s on

££• t e x t i l i s i n J a v a .

W ild b a n a n a s grow i n g r o v e s o r c lu m p s a n d so t h e r e

a r e n o r m a l l y s e v e r a l f l o w e r i n g p l a n t s w i t h i n an y l a r g e c lum p a t

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

q u a n t i t y o f f o o d e v e r y n i g h t f o r up t o f i v e m o n t h s , a g ro v e o f

200

b a n a n a s p r o v i d e s a d e p e n d a b l e an d c o n i i n o u s s u p p l y o f fo o d so

l o n g a s t h e c lum p s u r v i v e s . H ow ev er , t h e q u a n t i t y o f foo d

p ro d u c e d p e r p l a n t per- n i g h t i s l i m i t e d b e c a u s e t h e r e i s o n ly

one i n f l o r e s c e n c e p e r s t e m ,

. The o n l y o t h e r p o l l e n r e g u l a r l y fo u n d i n f a e c a l

s a m p le s f rom M a c r o g lo s s u s s o b r i n u s was t h a t o f D u aban ga g r a n d l f l u r a

(o n e o f t h e o n l y two d i c o t y l e d o n s i n Group I ) . B a t s c a u g h t u n d e r

f l o w e r i n g t r e e s o f P a r k i a s p e c i o s a and D u r io z i b e t h i n u s ( b o t h

Group I I t r e e s ) c o n t a i n e d a l m o s t no p o l l e n b e s i d e s t h a t o f Musa

s p p . o r D. g r a n d i f l o r a and no p o l l e n o f an y o t h e r G roup I I s p e c i e s

was fo u n d i n an y p o l l e n s a m p le s c o l l e c t e d from M. s o b r i n u s .

I t i s a l i t t l e s u r p r i s i n g t o h a v e fo u n d no e v i d e n c e

o f K. s o b r i n u s f e e d i n g from t h e f l o w e r s o f P ro x y lu m i n d l c u m w h ich

f r e q u e n t l y grew a l o n g s i d e Musa .m a l a c c e n s i s an d h a d many o f t h e

c h a r a c t e r i s t i c s o f G roup I p l a n t s . H ow ever , m o s t s t u d i e s o f

0 . in d i c u m w e re made on two t r e e s on t h e U n i v e r s i t y o f M alaya

campus w h ic h was o t h e r w i s e n o t s u i t a b l e , f o r M. s o b r i n u s b e c a u s e

t h e r e w e re v e r y few w i l d b a n a n a s an d no D uabanga g r a n d l f l o r a ••

t r e e s t h e r e . F u r t h e r s t u d i e s w h e re 0 , i n d i c u m g ro w s a l o n g s i d e

w i l d b a n a n a s , e . g . a l o n g t h e T a p a h - R i n g l e t r o a d may show t h a t

0 . I n d i c u m i s u s e d b y t h e s e b a t s .

The m o s t i m p o r t a n t p o l l e n s o u r c e s f o r M. m in im us

a t R a n ta u P&njang w e re m a n g ro v e s o f t h e g e n u s S o n n e r a t i a . The

r e s t r i c t i o n o f t h i s b a t i n P e n i n s u l a r M a l a y s i a t o t h e v i c i n i t y

o f m a n g ro v e s s t r o n g l y s u g g e s t s t h a t S o n n e r a t i a i s e s s e n t i a l t o

them t h e r e .

T h e re w e re t h r e e s p e c i e s o f S o n n e r a t i a a t R a n ta u

201

P a n j a n g . ' S . c a s e o l a r i s b e l o n g s t o Group I , f l o w e r i n g c o n t i n o u s l y

b u t p r o d u c i n g r e l a t i v e l y few f l o w e r s e a c h n i g h t . I t was v e r y

common on t h e r i v e r b a n k s . S . a l b a and S . o v a t a h a v e b e e n p l a c e d

i n Group I I on a c c o u n t o f t h e i r f l o w e r i n g i n f l u s h e s b u t ' a few

t r e e s c o u l d a lw a y s be fou nd i n f l o w e r . S . a l b a and S . o v a t a e a c h

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

t h e i r f l o w e r i n g s e a s o n s d i d n o t f u l l y o v e r l a p one s p e c i e s o r t h e

o t h e r was i n f l o w e r f o r a b o u t e i g h t m on ths i n 1972 and 1973*

'T hey b o t h r e s e m b l e d Group I p l a n t s i n o t h e r r e s p e c t s , e . g . t h e y

a r e g r e g a r i o u s and p r o d u c e r e l a t i v e l y few f l o w e r s p e r t r e e p e r ■

n i g h t com pared t o m o s t Group I I s p e c i e s .

I n t h e p a s t a t R a n ta u P a n j a n g , S . c a s e o l a r i s w ou ld

p r o v id e r , fo o d f o r t h e b a t s c o n t i n o u s l y and S , a l b a and S . o v a t a

b e tw e e n them f o r n e a r l y 3 / h o f t h e y e a r . H ow ev er , m o s t S . o v a t a

h a s b e e n e r a d i c a t e d b y man b u t i t h a s b e e n r e p l a c e d b y m ixed

c u l t i v a t i o n i n c l u d i n g s e v e r a l p l a n t s p e c i e s t h a t M a c r o g lo s s u s

m in im us c a n an d d o e s u s e f o r f o o d , p r i n c i p a l l y c o c o n u t p a lm s ,

Cocos n u c i f e r a , and c u l t i v a t e d b a n a n a s , Musa s p p . T h e se a r e b o t h

Group I p l a n t s , p r o v i d i n g fo o d c o n t i n o u s l y . T h e r e w e re a l s o

s e v e r a l Group I I p l a n t s grown t h e r e , a l l o f w h ic h M. m in im us f e d

from when a v a i l a b l e , i . e . C e lb a p e n t a n d r a , D u r io z i b e t h i n u s ,

E u g e n ia m a l a c c e n s i s and M a n g i f e r a s p p . T h ese w e re n o t t h o u g h t

t o b e i m p o r t a n t t o t h e b a t s .

The u s e o f so many a l t e r n a t i v e s t o S o n n e r a t i a ,

i n c l u d i n g b o t h G roup I and G roup I I p l a n t s c o n t r a s t s s t r o n g l y

w i t h t h e r e s t r i c t i o n o f M. s o b r i n u s t o Musa s p p . an d o c c a s i o n a l l y

D uabanga g r a n d i f l o r a and i t s u g g e s t s t h a t M. m in im us i s more

a d a p t a b l e t o v a r i a t i o n i n fo o d s u p p l i e s t h a n i s M. s o b r i n u s .

202

A d a p t a b i l i t y w ou ld b e a d v a n t a g e o u s t o M a c r o g lo s s u s

m in im us i n two s i t u a t i o n s . F i r s t l y , b e tw e e n f l u s h e s o f S o n n e r a t i a

a l b a and S . o v a t a t h e b a t s may h a v e t o l o c a t e a l t e r n a t i v e foo d

s o u r c e s u n l e s s S . c a s e o l a r i s -'can c a t e r f o r a l l t h e i r n e e d s , e . g .

a t T e lo k Asara, Bako N a t i o n a l P a r k , S a ra w a k , S . a l b a was t h e o n ly

t?"n0 c i o f * S o n n g ps . t is . s. v s i i2.5.13Xo* I t 1 1 ov,r0 r*sd. i n H nsiiG S Tlicx'G

w ere no c u l t i v a t e d s p e c i e s s u c h a s b a n a n a s ( e x c e p t f o r a v e r y

s m a l l num ber o f c o c o n u t p a l m s ) . I n May 1973 when v e r y few S . a l b a

t r e e s w e re s t i l l i n f l o w e r 5 6 . 2 5 % o f t h e 3 2 b a t s c a u g h t w ere

s u b a d u l t s , i . e . 1 . 3 t i m e s a s many . s u b a d u l t s a s a d u l t s and 2 . 6

t i m e s a s many s u b a d u l t s a s a d u l t f e m a l e s , T h i s c o m p a re s w i t h

9 . 1 % s u b a d u l t s and 8 t i m e s a s many a d u l t f e m a le s a s s u b a d u l t s

i n J a n u a r y t h e same y e a r . A l th o u g h a h i g h number o f b i r t h s a r e

t h o u g h t t o h a v e o c c u r r e d t h e r e i n t h e ' i n t e r v e n i n g t im e t h i s

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

May. The m o s t r e a s o n a b l e e x p l a n a t i o n f o r t h e d i f f e r e n c e i s t h a t

many a d u l t s h a d e m i g r a t e d i n t h e f a c e o f t h e im p e n d in g s h o r t a g e

o f f o o d . H ow ever , t h i s h a s n o t b e e n v e r i f i e d .

The v e r y much d e n s e r p o p u l a t i o n o f M a c r o g lo s s u s

m in im us a t R a n ta u P a n ja n g t h a n a t T e lo k Asam may b e e x p l a i n e d

by t h e en orm ous r e s e r v o i r o f foo d c o n t i n o u s l y a v a i l a b l e from

c o c o n u t p a lm s an d b a n a n a p l a n t s a t t h e f o r m e r s i t e b u t n o t a t

t h e l a t t e r .

S e c o n d l y , a d a p t a b i l i t y may b e o f v a l u e t o M. m inim us

where^M . s o b r i n u s d o e s n o t o c c u r , e . g . i n t h e P h i l i p p i n e s , B o rn e o

and S u l a w e s i ^ b e c a u s e i n t h e s e p l a c e s M. m in im us i s n o t r e s t r i c t e d

t o m a n g ro v es b u t c a n c o l o n i s e s u i t a b l e h a b i t a t s t h r o u g h o u t t h e

l a n d . I t s fo o d s o u r c e s f a r from m an g ro v e s i n t h e s e p l a c e s a r e

203

unknown t )u t Musa s p p . may b e e x p e c t e d t o b e o f c o n s i d e r a b l e

i m p o r t a n c e .

E o n y c t e r i s s p e l a e a . . (S in c e n o t h i n g i s known o f

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

by the ' b a t s , n e i t h e r t h e n u m e r i c a l a b u n d a n c e n o r t h e vo lum e ' o f

an y p o l l e n s p e c i e s i n t h e gu an o o f E . s p e l a e a n e c e s s a r i l y r e f l e c t s

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

r e f e r e n c e s t o t h e i m p o r t a n c e o f an y p o l l e n t y p e r e f e r o n ly to

t h e c o n t r i b u t i o n o f t h a t p o l l e n t y p e to t h e am oun t o f p o l l e n/

i n g e s t e d by. t h e b a t s . )

Most o f t h e i n f o r m a t i o n on t h e r e l a t i v e i m p o r t a n c e

o f v a r i o u s fo o d s o u r c e s t o E . s p e l a e a was d e r i v e d from an a n a l y s i s

o f p o l l e n i n t h e g uan o from B a tu C aves and Gua S a n d i n g . B e c a u s e

o f t h e w id e v a r i e t y o f fo o d p l a n t s ( t h e y w ere r e c o r d e d f e e d i n g

from a l l t h e c h i r o p t e r o p h i l o u s p l a n t s i n P e n i n s u l a r M a l a y s i a )

an d t h e enorm ous f e e d i n g r a n g e o f E . s p e l a e a . t h i s was t h e o n ly

m e th od b y w h ich t h e r e l a t i v e i m p o r t a n c e o f d i f f e r e n t fo o d s o u r c e s

c o u l d b e e s t a b l i s h e d . ;

The m a j o r i t y o f p o l l e n g r a i n s i n t h e guano from

B a tu C aves w e re o f D uab ang a g r a n d i f l o r a o r A r t o c a r p u s s p p . , b u t

t h e y b o t h h a v e v e r y s m a l l p o l l e n g r a i n s and t h e b u l k o f t h e p o l l e n

was a l l from Group I I s p e c i e s . The l a t t e r f l o w e r e d i n f l u s h e s o r

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

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

e x c e p t i o n was P ro x y lu m in d i c u m w h ic h , a t l e a s t on t h e T a p a h -

R i n g l e t r o a d , show ed a d i s t i n c t f l o w e r i n g s e a s o n ( a l t h o u g h

f l o w e r i n g p l a n t s w e re n o t uncommon t h r o u g h o u t t h e y e a r ) b u t w h ic h

o t h e r w i s e was c h a r a c t e r i s t i c o f Group I p l a n t s . Anyway 0 . In d ic u m

20'+

p o l l e n was n e v e r common i n t h e g u a n o . T h i s c a n p a r t l y b e

e x p l a i n e d b y " t h e f a c t t h a t a l t h o u g h i t i s a common s p e c i e s e a c h

p l a n t p r o d u c e s o n l y a few f l o w e r s e a c h n i g h t ' an d t h e y p r o v i d e

r e l a t i v e l y l i t t l e p o l l e n w h ic h i s d e p o s i t e d p r e c i s e l y on t h e

b a t ' s h e a d a nd n a p e .

Two s p e c i f i c e x c e p t i o n s o c c u r r e d . F i r s t l y , d u r i n g

a f l o w e r i n g f l u s h o f S o n n e r a t i a a l b a from A p r i l t o J u n e 1 972 t h e r e

was v e r y l i t t l e S . a l b a p o l l e n i n t h e B a tu C av es gu ano s a m p le s

sh o w in g t h a t b a t s from t h a t c o l o n y w e re n o t f e e d i n g on i t v e r y

m uch. The f l u s h v/as a p o o r one i n t h a t f e w e r f l o w e r s w e re p r o d u c e d

e a c h n i g h t t h a n i n a n o rm a l f l u s h and i t c o i n c i d e d w i t h a v e r y

h e a v y f l o w e r i n g s e a s o n o f P a r k i a s p e c i o s a . P a r k i a s p p . p o l l e n

made up t h e b u l k o f t h e p o l l e n i n t h e B a tu C aves g uano a t t h a t

t i m e . T h i s i s an ex am p le o f t h e b a t s sho w ing a p r e f e r e n c e f o r

a t r e e t h a t was f l o w e r i n g v e r y h e a v i l y i n t h e f o r e s t s a r o u n d B a tu

C aves o v e r a t r e e t h a t was n o t f l o w e r i n g h e a v i l y a n d grew 38 km

o r more from t h e i r r o o s t .

S e c o n d l y , E u g e n ia m a l a c c e n s j s f l o w e r e d f o u r t i m e s

d u r i n g t h e t w o ' y e a r s . Each t im e f l o w e r i n g c o i n c i d e d w i t h a p e a k

i n E u g e n ia s p p . p o l l e n i n guano from B a tu C aves e x c e p t i n March

1973* No d i f f e r e n c e i n t h e num ber o f f l o w e r i n g t r e e s n o r t h e

num ber o f f l o w e r s p e r t r e e was n o t i c e d a t t h a t t im e b u t t h e

m a j o r i t y o f t h e p o l l e n i n t h e s a m p le s a t t h a t t i m e was Du a b a n g a

g r a n d i f l o r a (w h ic h f l o w e r s c o n t i n o u s l y anyway) w h ic h g av e way

t o p e a k s i n - t h e p r o p o r t i o n o f D u r io s p p . p o l l e n an d a f t e r t h a t

an unknown E u g e n ia s p p . p o l l e n t h e f o l l o w i n g m o n th .

W henever p o l l e n from Group I I s p e c i e s v/as s c a r c e

I

205

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

p o l l e n becam e v e r y common i n t h e g uano so t h a t D^ gr a n d i f l o r a

anc* A r t o c a r p u s s p p . . a p p e a r t o b e '’i n s u r a n c e " s p e c i e s on v /h ich

t h e b a t s c a n f a l l b a c k when o t h e r fo o d s o u r c e s a r e i n s u f f i c i e n t

f o r t h e i r n e e d s . (T he n u m e r i c a l p r o p o r t i o n s o f t h e s e two p o l l e n

t y p e s w e re c l e a r l y r e l a t e d t o one a n o t h e r b u t , u n t i l t h e s p e c i f i c

i d e n t i t y o f t h e A r t o c a r p u s s p p . p o l l e n i s a s c e r t a i n e d and th e

d i s t r i b u t i o n , p h e n o lo g y and m o rp h o lo g y o f t h e t r e e s i n v e s t i g a t e d ,

no f r u i t f u l s p e c u l a t i o n on t h e n a t u r e o f t h e r e l a t i o n s h i p i s

p o s s i b l e . )

I t i s i n v i t i n g t o r e g a r d t h e r e d u c t i o n i n t h e

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

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

e v i d e n c e f o r a p r e f e r e n c e f o r t h e l a t t e r . U n d o u b te d ly , t h i s i s

so on some o c c a s i o n s a s t h e Group I I s e a s o n a l p o l l e n t y p e s w ere

so m e t im e s a l m o s t t h e o n l y p o l l e n s p e c i e s p r e s e n t i n t h e s a m p le s

i n n u m e r i c a l a n d , e v e n more s o , i n v o l u m e t r i c t e r m s , a l t h o u g h

g r a n d i f l o r a a n d , p r e s u m a b ly A r t o c a r p u s s p p . , w e re s t i l l

f l o w e r i n g .

I n t h e l e s s w e l l d e f i n e d c a s e s i t i s n o t n e c e s s a r i l y

t r u e t h a t a r e d u c t i o n i n t h e n u m e r i c a l p r o p o r t i o n o f one p o l l e n

t y p e c o r r e s p o n d i n g w i t h an i n c r e a s e i n t h e p r o p o r t i o n o f a n o t h e r

p o l l e n t y p e shov/s t h a t t h e b a t s a t e l e s s o f t h e f o r m e r t y p e .

T h i s i s b e c a u s e t h e n u m e r i c a l p r o p o r t i o n o f e a c h p o l l e n t y p e i n

a s am p le o f t h e g uan o i s n o t a m e a s u re o f t h e a b s o l u t e num ber o f

g r a i n s o f e a c h s p e c i e s i n g e s t e d b y t h e b a t s . F o r e x a m p le , i f a

g ro u p o f b a t s f e d p r i m a r i l y on D. g r a n d i f l o r a and P . s p e c i o s a

206

f o r two c o n s e c u t i v e w eeks b u t d u r i n g t h e s e c o n d w eek t h e number

£ • g r a n d i f l o r a g ' r a i n s i n g e s t e d v/as d o u b le d w h i l e t h e number

0 ^ £* s p e c i o s a g r a i n s i n g e s t e d was t r e b l e d , t h e n t h e n u m e r i c a l

p r o p o r t i o n s o f D. g r a n d i f l o r a i n t h e s e c o n d week w o u ld a c t u a l l y

b e lo w e r t h a n i n t h e f i r s t w eek b y a b o u t 20%.

T h i s i s an e x t r e m e h y p o t h e t i c a l e x am p le b u t i t

d o e s i l l u s t r a t e how an i n c r e a s e i n t h e a b s o l u t e num ber o f p o l l e n

g r a i n s o f a g i v e n s p e c i e s t h a t w ere e a t e n b y t h e b a t s c o u ld s t i l l

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

p o l l e n s p e c i e s i n g uano s a m p le s t h e s e c o n d w ee k .

H ow ev er , d e s p i t e t h i s l i m i t a t i o n , a r e d u c t i o n i n

n u m e r i c a l p r o p o r t i o n s o f one p o l l e n c o r r e s p o n d i n g w i t h an

i n c r e a s e i n a n o t h e r i s a m e a s u re o f t h e ch a n g e i n t h e r e l a t i v e

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

th e m . . '

T hus t o r e t u r n t o t h e h y p o t h e t i c a l e x a m p le ,

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

t h e ' s e c o n d week i n c r e a s e d , D . g r a n d i f l o r a was r e l a t i v e l y l e s s

i m p o r t a n t t o t h e b a t s t h a n P . s p e c i o s a d u r i n g t h e s e c o n d w eek .

/ S i m i l a r l y , t h e r e d u c t i o n s i n t h e n u m e r i c a l

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

w i t h e a c h p e a k i n t h e p r o p o r t i o n o f p o l l e n o f s e a s o n a l l y f l o w e r i n g

s p e c i e s do r e f l e c t c h a n g e s i n t h e r e l a t i v e i m p o r t a n c e o f t h o s e

p o l l e n s p e c i e s t o t h e c l u s t e r s o f b a t s from v /h ich t h e g uano was

c o l l e c t e d .

M ost o f t h e Group I p l a n t s w e re n o t i m p o r t a n t

s o u r c e s o f p o l l e n t o E o n y c t e r i s s p e l a e a ♦ H ow ever , t h e i m p o r t a n c e

o f Musa s p p . i s d i f f i c u l t - t o a s s e s s b e c a u s e c u l t i v a t e d b a n a n a s

207

yp r o d u c e no p o l l e n and p o l l e n o f w i l d s p e c i e s b r e a k s down r a p i d l y

i n t h e guano ( b u t n o t b e f o r e d e f a e c a t i o n ) and so was r a r e l y

e n c o u n t e r e d i n guano s a m p l e s . B a n an as f l o w e r i n g i n open s i t u a t i o n s

w e re o f t e n v i s i t e d b y E . s p e l a e a w h ic h was f r e q u e n t l y m i s t - n e t t e d

a d j a c e n t t o them b u t , e x c e p t f o r b a t s c a u g h t n e a r f l o w e r i n g

b a n a n a s , v e r y few E . s p e l a e a h ad Musa s p p . p o l l e n on t h e i r muz,z,ies

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

and f o r a g i n g b e h a v i o u r o f E . s p e l a e a and t h e n o r m a l g ro w th h a b i t s

o f Musa s p p . , b a n a n a s a r e n o t l i k e l y t o b e o f g r e a t i m p o r t a n c e

t o Eh s p e l a e a .

The p o l l e n o f Cocos n u c i f e r a was a r e g u l a r b u t

m in o r c o n s t i t u e n t o f t h e g uano and A re n g a s p p . p o l l e n was e x t r e m e l y

r a r e i n gu ano from B a tu C a v e s . H ow ever , S o n n e r a t i a c a s e o l a r i s

was s o m e t im e s an i m p o r t a n t c o n s t i t u e n t o f t h e g u a n o . I t s a b u n d a n c e

f l u c t u a t e d ; s m a l l p e a k s so m e t im e s o c c u r r e d b e f o r e o r a f t e r p e a k s

i n t h e p r o p o r t i o n o f S . a l b a p o l l e n i n t h e g u a n o . T h ese c o u ld

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

S . a lb .a and f e e d i n g on S . c a s e o l a r i s b e c a u s e f l o w e r s o f S . a l b a

w ere s c a r c e a t t h e b e g i n n i n g o r end o f a f l o w e r i n g f l u s h . The

r e a s o n s f o r p e a k s i n t h e p r o p o r t i o n o f S . c a s e o l a r i s p o l l e n a t

o t h e r t i m e s a r e n o t c l e a r b u t p o s s i b l y E. s p e l a e a u s e d S . c a s e o l a r i s

more when o t h e r fo o d s o u r c e s w e re s c a r c e .

T h e re was a m arked d i f f e r e n c e i n t h e p r o p o r t i o n s

S o n n e r a t i a a l b a and S . c a s e o l a r i s p o l l e n p r e s e n t i n guano

c o l l e c t e d from B a tu C aves and from Gua S a n d i n g . The p o l l e n o f

S . a l b a was v e r y common i n s a m p le s from B a tu C aves e s p e c i a l l y

d u r i n g f l o w e r i n g f l u s h e s o f S . a l b a w h e re i t c o n t r i b u t e d from

6.65% o f a l l t h e p o l l e n on T ra y I to 1 k ,0 k% o f a l l t h e p o l l e n

208

on T ra y IX^ I n s a m p le s from Gua S a n d in g i t was a lw a y s r a r e and

c o n t r i b u t e d o n l y 0 o f a l l t h e p o l l e n g r a i n s c o u n t e d . On t h e

o t h e r h a n d , S . c a s e o l a r i s was much l e s s common i n B a tu C aves t h a n

i n Gua S a n d in g w h e re i t was t h e commonest s i n g l e p o l l e n s p e c i e s .

The q u a n t i t y a nd p r o x i m i t y o f t r e e s o f e a c h s p e c i e s

t o t h e tv/o b a t c o l o n i e s p r o b a b l y a c c o u n t s f o r t h e s e i n i t i a l l y

p u z z l i n g r e s u l t s . B e tw een 38 and 50 km from B a tu C aves t h e r e a r e

v e r y e x t e n s i v e a r e a s w h e re S . a l b a grow s b u t t h e c l o s e s t p o i n t

t o Gua S a n d in g a t w h ic h i t i s l i k e l y t o grow i s 60 km from t h e

c a v e . H o w ever , t h e r e i s o n l y a l i m i t e d am oun t o f s u i t a b l e h a b i t a t

a l o n g t h e K la n g r i v e r ( a s m a l l r i v e r ) f o r S . c a s e o l a r i s t o grow

c l o s e t o B a tu C aves b u t many m i l e s o f t h e t i d a l r e a c h e s - o f t h e

h u g e P e r a k r i v e r an d some o f i t s t r i b u t a r i e s w h ic h a r e p r o b a b l y

s u i t a b l e f o r S . c a s e o l a r i s , a r e w i t h i n 50 km o f Gua S a n d i n g .

( S . c a s e o l a r i s g ro w s a b u n d a n t l y a t T e lo k Anson on t h e P e r a k r i v e r

( p e r s . o b s . ) . )

O t h e r m arked d i f f e r e n c e s i n t h e p o l l e n c o n t e n t

o f guano from Gua S a n d in g and B a tu C aves c o n c e r n e d A r t o c a r p u s

s p p . w h ic h was n o t common i n g uano from Gua S a n d in g b u t was

common i n gu ano from B a tu C a v e s , and A ren g a s p p . w h ic h was a

common and r e g u l a r s p e c i e s i n Gua S a n d in g b u t v e r y r a r e i n B a tu

C a v e s . T h e re a r e two p o s s i b l e r e a s o n s f o r t h e d i f f e r e n c e s o f

e a c h o f t h e s e s p e c i e s . E i t h e r t h e t r e e s w ere more common i n t h e

f e e d i n g r a n g e o f one c o lo n y t h a n t h e o t h e r o r . t h e y w e re e q u a l l y

common i n t h e f e e d i n g r a n g e s o f b o t h c o l o n i e s b u t t h e b a t s from

B a tu Caves r a r e l y f e d on. A re n g a s p p . p o l l e n and t h e b a t s from

Gua S a n d in g f e d on A r t o c a r p u s s p p . p o l l e n l e s s o f t e n t h a n t h e

209

b a t s from B a tu Caves,, -B e fo re t h e s e q u e s t i o n s c a n b e r e s o l v e d t h e

s p e c i f i c i d e n t i t y o f t h e p o l l e n m us t b e e s t a b l i s h e d and t h e

d i s t r i b u t i o n and p h e n o lo g y o f t h e t r e e s from w h ic h t h e p o l l e n

comes m u s t b e s t u d i e d . .

D e s p i t e t h e s e d i f f e r e n c e s i n t h e p r o p o r t i o n o f

some p o l l e n t y p e s fo u n d i n t h e g uano from B a tu C aves an d Gua

S a n d in g , ( k e e p i n g i n m ind t h e l i m i t a t i o n s t o v /h ich c o m p a r i s o n s

a r e p o s s i b l e b e c a u s e o f d i f f e r e n c e s i n t h e m e th o d s b y w h ich t h e y

w e re c o l l e c t e d ) t h e r e i s a b r o a d s i m i l a r i t y i n t h e p o l l e n fo u n d

i n t h e two c a v e s , w i t h p o l l e n from Group I I p l a n t s p r e d o m i n a t i n g

d u r i n g t h e i r f l o w e r i n g s e a s o n s and c o n t r i b u t i n g t h e b u l k o f t h e

p o l l e n i n b o t h c a v e s . B e c a u s e c o l l e c t i o n s from Gua S a n d in g w e re

n o t c o n t i n o u s i t c a n n o t b e shown w h e th e r a G roup I p o l l e n ,

s u c h a s D uabanga g r a n d i f l o r a becam e nu m erou s i n t h e g uano t h e r e

when p o l l e n was u n a v a i l a b l e from Group I I p l a n t s ( a s v/as t h e c a s e

i n B a tu C av es ) b u t t h i s p r o b a b l y h a p p e n e d .

I n sum mary, t h e d i e t s o f M a c r o g l o s s u s min im us

and M. s o b r i n u s do n o t o v e r l a p i n P e n i n s u l a r M a l a y s i a e x c e p t

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

a d d i t i o n t o t h e i r u s u a l d i e t s . M. m in im us i s more a d a p t a b l e t o

a w id e v a r i e t y o f f o o d ' t h a n i s M. s o b r i n u s an d t h i s h o l d s c e r t a i n

a d v a n t a g e s . H o w ev er , b o t h s p e c i e s r e l y p r i m a r i l y on p l a n t s i n

Group I , i . e . s p e c i e s t h a t f l o w e r t h r o u g h o u t t h e y e a r , e a c h p l a n t

p r o v i d i n g r e l a t i v e l y l i t t l e fo o d e a c h n i g h t b u t f o r a lo n g t i m e .

The p l a n t s a r e u s u a l l y g r e g a r i o u s . S o n n e r a t i a a l b a and S . o v a t a

a r e e x c e p t i o n a l i n t h a t t h e y do n o t f l o w e r c o n t i n o u s l y . H ow ever ,

b e tw e e n them t h e y p r o v i d e fo o d f o r much o f t h e t i m e an d t h e y a r e

g r e g a r i o u s .

£10

Eonycteris spelaea feeds on the same plant; species

but they are not. of very great' importance except when other food

sources are unavailable. (Sonneratia caseolaris was exceptional

to the Gua Sanding colony where the pollen regularly occurred

in abundance in the guano,)

Most of the pollen in the guano of E. spelaea

was from interm ittently flowering species. These trees were

usually thinly distributed over very large areas but they did

provide enormous quantities of food during their short flowering

seasons. /

3 . ROOSTING BEHAVIOUR IN RELATION TO FOOD SOURCES.

M a c r o g lo s s u s sp p „

V ery l i t t l e i s known o f t h e r o o s t i n g h a b i t s o f

e i t h e r s p e c i e s o f M a c r o g lo s s u s i n t h e w i l d . Lim (1 9 6 6 ) r e c o r d e d

M* m in im us r o o s t i n g i n g r o u p s o f f i v e t o t e n i n p a lm t r e e s .

H ow ev er , h e was c l e a r l y u n a b l e t o d i s t i n g u i s h M. m in im us from

M. s o b r i n u s a nd t h i s r e c o r d m i g h t a p p l y to e i t h e r s p e c i e s . I n

P e n i n s u l a r M a l a y s i a one M_. s o b r i n u s was c a u g h t h a n g i n g i n a

c lum p o f b am b o o s . ( D a ta on l a b e l o f a s p e c im e n i n t h e IMR

c o l l e c t i o n ) I n t h e P h i l i p p i n e s , B a r b o u r , L aw rence and P e t e r s '

(1 9 3 9 ) r e p o r t e d t h a t one M. m in im us v/as " s h a k e n o u t o f a r o l l e d

up hemp l e a f " b u t t h a t " w i t h f i v e men a t w ork o n l y t h i s s i n g l e

i n d i v i d u a l was c a u g h t d u r i n g an e n t i r e d a y ’s s e a r c h ; a n o t h e r

p r e s u m a b ly t h e same s p e c i e s e s c a p e d " . VJalker ( “[SSh) r e p o r t e d

them t o b e s o l i t a r y and r e c o r d e d them s h e l t e r i n g u n d e r t h e

b r a n c h e s o f t r e e s and u n d e r r o o f s b u t " t h e p r e f e r r e d d a y t im e

//

211

r e t r e a t seem s t o b e i n t h e r o l l e d l e a v e s o f hemp an d b a n a n a p l a n t s " .

He d o e s n o t g i v e r e f e r e n c e s o u r c e s b u t h i s s t a t e m e n t may h a v e

come i n p a r t f rom H o r s f i e l d (1 8 2 2 ) who w r o te t h a t " d u r i n g t h e

day i t r e m a i n s s u s p e n d e d u n d e r t h e b r a n c h e s o f t r e e s o r i t r e t i r e s

u n d e r r o o f s o f o l d h o u s e s an d s h e d s " . I f t h i s i s t h e o r i g i n o f

W a l k e r ' s ( l 9 o u ) s t a t e m e n t r e g a r d i n g t r e e s and r o o f s i t s h o u l d

b e t r e a t e d w i t h c a u t i o n a s H o r s f i e l d o b v i o u s l y c o n f u s e d M a c r o g lo s s u s

(= h i s P t e r o p u s r o s t r a t u s ) w i t h o t h e r p t e r o p i d s , e . g . i n t h e

p r e c e d i n g s e n t e n c e s h e s t a t e s t h a t i t com m its " s e r i o u s i n j u r y ;

among, t h e p l a n t a t i o n s and f r u i t t r e e s " and t h a t " i t f e e d s on

f r u i t s o f e v e r y d e s c r i p t i o n b u t p a r t i c u l a r l y i n f e s t s v a r i o u s \

s p e c i e s o f E u g e n ia ................. . . . " . T h i s i s n o t t r u e o f M a c r o g lo s s u s

i n J a v a (v a n d e r P i j l , 1936) o r P e n i n s u l a r M a l a y s i a ( p e r s . o b s . ) .

N e v e r t h e l e s s , t h e s e f a l l a c i e s a r e a l s o r e p o r t e d i n W a lk e r (1 9 6 * 0 .

A l th o u g h d i l i g e n t s e a r c h e s w ere made i n r o l l e d

up b a n a n a l e a v e s t h e o n l y b a t s l o c a t e d t h e r e w e re M y o t l s m y s t a c i n u s

f o r w h ich t h a t i s a u s u a l r o o s t s i t e (Medway, 1 969 )#

The p o s t u r e and b e h a v i o u r i n c a p t i v e M a c r o g lo s s u s

o f b o t h s p e c i e s a r e t y p i c a l o f man^r o t h e r s m a l l p t e r o p i d s w h ic h

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

( 1 9 7 2 ) , R o s e v e a r (1 9 6 5 ) an d p e r s . o b s . J o n e s ( 1 972 ) fo u n d t h a t

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

and p o s t u r e o f c a p t i v e b a t s w e re s i m i l a r t o t h o s e o b s e r v e d f o r

t h e same s p e c i e s i n t h e w i l d . T h e re i s no r e a s o n t o b e l i e v e t h a t

t h i s i s n o t t r u e f o r M a c r o g lo s s u s s p p .

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

o f M a c r o g lo s s u s s p p . c an b e a c c e p t e d w i t h c o n f i d e n c e . •

Ii

21 2

1 . They r o o s t s i n g l y o r i n s m a l l g r o u p s ,

2., They r o o s t s u s p e n d e d from a "b ranch , pa lm f r o n d o r s i m i l a r

o b j e c t , w i t h one f o o t e i t h e r s i d e o f t h e s u p p o r t , and w i t h

t h e i r b o d i e s e n v e lo p e d i n t h e i r w in g s ( o r p o s s i b l y t h e y som e­

t i m e s r o o s t i n t h e u n f u r l e d l e a v e s o f b a n a n a s ) ,

3» They r e s t o r s l e e p m o s t o f t h e d a y .

4 . Some r o o s t r e p e a t e d l y i n t h e same v i c i n i t y ( p o s s i b l y i n t h e

same s i t e ? ) . ( T h i s was shown b y r e c a p t u r e o f t h e same

i n d i v i d u a l s o f M. m in im us a t t h e same n e t s i t e s v e r y e a r l y -

i n t h e e v e n i n g on s e v e r a l o c c a s i o n s a t R a n ta u P a n j a n g and

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

t h e i r c a g e . I t was e m p h a s i s e d b y t h e one c a p t i v e b a t w h ic h ,

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

pa lm t r e e t o w h ic h i t h a d b e e n a c c u s to m e d i n i t s c a g e a l t h o u g h

t h e l a t t e r h a d b e e n rem o ved 2 0 0 m i n t o t a l l y u n f a m i l i a r

s u r r o u n d i n g s .

B o th s p e c i e s o f M a c r o g lo s s u s o b t a i n fo o d from

s o u r c e s t h a t a r e a v a i l a b l e t h r o u g h o u t t h e y e a r . T h e i r r e s p e c t i v e

fo o d p l a n t s grow g r e g a r i o u s l y and a b u n d a n t l y i n s u i t a b l e h a b i t a t s .

The b a t s do n o t h a v e t o t r a v e l 3.ong d i s t a n c e s i n s e a r c h o f e v e r

c h a n g in g s u p p l i e s ( e x c e p t t h a t M. m in im us f e e d i n g on S o n n e r a t i a

a l b a i s o l a t e d from t h e l a n d an d o t h e r s p e c i e s o f S o n n e r a t i a m u s t

h a v e an a l t e r n a t i v e fo o d s o u r c e , o r f a c e a fo o d s h o r t a g e , b e tw e e n

f l u s h e s ) .

R o o s t s o f i n d i v i d u a l s o r s m a l l g r o u p s d i s p e r s e d

t h r o u g h o u t t h e a r e a i n w h ich t h e i r foo d p l a n t s grow a r e t h e r e f o r e

s u i t a b l e t o t h e s e b a t s .

B e in g s m a l l w i t h a c o r r e s p o n d i n g h i g h s u r f a c e

213

a r e a t o vo lum e r a t i o and l i v i n g i n a h o t a t m o s p h e r e i t i s

p r o b a b l y i m p o r t a n t t o them t o m a i n t a i n a low r a t e o f m e t a b o l i c

a c t i v i t y by d ay i n o r d e r t o c o n s e r v e e n e r g y and w a t e r . I f

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

t h e f o l l o w i n g d ay ( p e r s . o b s . ) ; K o k ( l 973) o b s e r v e d t h e r e c t a l

t e m p e r a t u r e of_M. m in im us s u b j e c t e d t o i2 ° C i n a r e f r i g e r a t o r .

(H e r s u b j e c t s w e re c a u g h t by me a t a b o u t 2 3 .0 0 h r s , i . e . a f t e r

f i v e h o u r s i n w h ic h t o f e e d . E x p e r im e n t s b e g a n a t a b o u t 2 4 .0 0

h r s t h e same n i g h t . ) She fo u n d t h a t i n i t i a l l y t h e y t r i e d to

m a i n t a i n t h e i r b o d y t e m p e r a t u r e b u t a f t e r a few h o u r s i t f e l l

r a p i d l y an d t h e b a t s d i e d so o n a f t e r w a r d s . She s u p p o s e d ( p e r s .

comm.) t h a t t h e su d d e n f a l l i n t e m p e r a t u r e came a b o u t when a l l

a v a i l a b l e e n e r g y r e s e r v e s ha d b e e n u s e d up t r y i n g t o m a i n t a i n

a n o rm a l bod y t e m p e r a t u r e . I f h e r s u p p o s i t i o n i s c o r r e c t i t

shov/s t h a t M. m in im us h a s a v e r y l i m i t e d e n e r g y r e s e r v e .

I f b a t s c a r r y i n g t h e e x t r a b u r d e n o f p r e g n a n c y

o r l a c t a t i o n a r e t o s u r v i v e even when h e a v y r a i n may p r e v e n t

f e e d i n g i h t h e f i r s t few h o u r s o f t h e e v e n i n g , t h e n i t may b e

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

t h e a c t i v i t y and s o c i a l s t r e s s commonly o b s e r v e d i n t h e .v a s t

c o l o n i e s o f some o f t h e l a r g e r p t e r o p i d s . T h i s t h e n may b e an

■ a d d i t i o n a l r e a s o n f o r h a v i n g s m a l l r o o s t i n g g r o u p s i n w h ic h t h e

b a t s r e m a in i n a c t i v e a l l d a y .

E o n y c t e r i s s p e l a e a .„

The r o o s t i n g b e h a v i o u r o f E . s p e l a e a i s d i f f e r e n t

i n a l m o s t a l l r e s p e c t s from t h a t o f M a c r o g lo s s u s s p p . They

n o r m a l l y r o o s t i n c a v e s , so m e t im e s i n h uge num b ers ( e . g . Medway,

214

195.8, 1 969 ; W a lk e r , 1964) • W i t h in t h e r o o s t s t h e b a t s w e re v e r y

r e s t l e s s , c o n s t a n t l y c h a n g in g p o s i t i o n s , g ro o m in g o r j o s t l i n g

f o r f o o t h o l d s on t h e r o o f * They hung i n t i g h t g r o u p s , t h e .

i n d i v i d u a l s i n c o n t a c t w i t h t h e i r n e i g h b o u r s .

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

t o l o c a t e , t h i s i s a common b a t w here no c a v e s e x i s t , e . g . i n

S i n g a p o r e ( H a r r i s o n , 1 9 6 6 ) ; t h e r e a r e no known c a v e r n i c o l o u s

c o l o n i e s o r even s u i t a b l e c a v e s s o u t h o f B a tu C a v e s , 300 km t o

t h e n o r th , w e s t . Lira (1 9 6 6 ) r e c o r d e d a r o o s t s h a r e d b y C y n o p te r u s

b r a c h y o t i s an d C. h o r s f i e l d i i n a c o c o n u t p a lm an d t h e a b o r i g i n e

c o l l e c t o r s w o rk in g f o r t h e I . M .R , h a v e l o c a t e d s m a l l g r o u p s o f

50 t o a few h u n d r e d i n d i v i d u a l s r o o s t i n g among b o u l d e r s and i n

r o c k c r e v i c e s (L im , 1966 and p e r s . comm.)

A common f e a t u r e o f a l l t h e c a v e r o o s t s t h a t h a v e

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

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

t o t h e t h r e e c a v e s ex am in ed i n P e n i n s u l a r M a l a y s i a a r e a l l i n

t h e s i d e s o f c l i f f s w e l l a b o v e t h e g ro u n d (a n d t h u s a b o v e t h e

ca n o p y o f t h e s u r r o u n d i n g f o r e s t b e f o r e i t v/as rem o v ed ) . The

i n a c c e s s i b i l i t y o f t h e e n t r a n c e t o Gua Anak T a k u n , a l i t t l e

n o r t h o f B a tu C a v e s , e x p l a i n s t h e a b s e n c e o f E . s p e l a e a from

t h a t c a v e ( s e e Dunn, 196 5 )* The e n t r a n c e i s b a r e l y one m e t r e

h i g h an d o n l y a l i t t l e w i d e r . I t i s h i d d e n b e h i n d a h u g e b o u l d e r

an d f u r t h e r o b s t r u c t e d b y t h e r o o t s o f a s t r a n g l i n g f i g ( F i c u s

s p p . ) . F u r t h e r m o r e , i t i s a t g ro u n d l e v e l and s u r r o u n d e d by

t h i c k v e g e t a t i o n .

Ward an d Z a h a v i (1 9 7 3 ) h a v e p u t f o r v /a rd s t r o n g

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

215

among b i r d s i s t h e d i s s e m i n a t i o n among members o f t h e r o o s t o f

i n f o r m a t i o n on t h e l o c a t i o n o f f o o d . A l th o u g h t h e s p e c i e s

i n v o l v e d h a v e a v a r i e t y o f d i f f e r e n t d i e t s and come from many

f a m i l i e s , t h e a u t h o r s p o i n t o u t t h a t a l l s p e c i e s s h a r e one

c h a r a c t e r i s t i c : They a l l h a v e t o t r a v e l l o n g d i s t a n c e s i n

s e a r c h Of fo o d s u p p l i e s w h ic h a r e w id e ly d i s p e r s e d , l o c a l i s e d

and i n t e r m i t t e n t b u t may b e t e m p o r a r i l y e x t r e m e l y r i c h .

The r o o s t s a r e u s u a l l y t r a d i t i o n a l and p e r s i s t

ev en when fo o d i s a b u n d a n t l y a v a i l a b l e . T h e re i s . u s u a l l y a h i g h

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

d e p a r t f rom t h e r o o s t s , and f o r a g e , i n f l o c k s .

The n a t u r e o f t h e d i e t and t h e r o o s t i n g b e h a v i o u r

s p e l a e a p a r a l l e l v e r y c l o s e l y t h o s e o f t h e com m unally

r o o s t i n g b i r d s . F u r t h e r m o r e Ik s p e l a e a f o r a g e s i n f l o c k s . Many

o f t h e t r e e s p e c i e s from w h ic h t h e y f e e d a r e t h i n l y d i s p e r s e d

o v e r w id e a r e a s . ' They f l o w e r f o r s h o r t i n t e r v a l s b u t when i n

f l o w e r e a c h t r e e p r o v i d e s a v e r y r i c h s o u r c e o f f o o d . I t i s

l i k e l y t h a t l a r g e communal r o o s t s i n t r a d i t i o n a l s i t e s s e r v e

t h e same p u r p o s e f o r E_. s p e l a e a a s t h e y do f o r some b i r d s , i . e .

t h a t o f d i s s e m i n a t i o n o f i n f o r m a t i o n on t h e w h e r e a b o u t s o f f o o d .

I t i s n o t e w o r t h y t h a t l a r g e communal r o o s t s a r e

t y p i c a l o f some o t h e r l a r g e p t e r o p i d s , e s p e c i a l l y R o u s e t t u s

(McCann, 1 94 1 ; M u t e r e , 1 9 6 8 , W a lk e r , 1 964 ; and p e r s . o b s . ) ,

R o u s e t t u s i s v e r y c l o s e l y r e l a t e d t o E o n y c t e r i s ( A n d e r s e n , 1912)

I t i s a l s o n o t e w o r t h y t h a t many s p e c i e s o f R o u s e t t u s t r a v e l

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

s o u r c e s , e . g . W a lk e r ( 1 9 6 4 5 P« 187) and Kock ( p e r s . comm.) f o r

R o u s e t t u s s p e c i e s i n A f r i c a . ; McCann (1 9 4 1 ) s u g g e s t e d t h a t t h e

216

c o lo n y o f R. a e g .y p t i a e u s G e o f f r o y l i v i n g on t h e b a r r e n i s l a n d

o f Kishra i n t h e P e r s i a n G u l f f l e w 20 m i l e s t o t h e m a i n l a n d to

f e e d e a c h n i g h t . T h i s i s a s y e t u n s u b s t a n t i a t e d b u t i t i s t h e

o n ly r e a s o n a b l e e x p l a n a t i o n o f how t h e y o b t a i n f o o d .

The m ethod by w h ich i n f o r m a t i o n on fo o d s o u r c e s

i s com m u n ic a ted t o o t h e r b a t s i n t h e r o o s t ( a s s u m i n g t h i s t o

b e a f u n c t i o n o f t h e r o o s t ) m u s t r e m a in s p e c u l a t i v e a t t h i s

s t a g e . Ward and Z a h a v i (1 9 7 3 ) s u p p o r t t h e s u g g e s t i o n t h a t i n

many b i r d s p e c i e s t h o s e i n d i v i d u a l s w h ich f a i l e d t o l o c a t e

s a t i s f a c t o r y fo o d s u p p l i e s one d ay f o l l o w t h o s e t h a t d i d when

t h e l a t t e r l e a v e t h e r o o s t t h e f o l l o w i n g d a y . I f t h e same i s

t r u e f o r E . s p e l a e a t h o s e b a t s t h a t fo u n d fo o d t h e p r e v i o u s d a y

c o u l d co m m u n ica te t h e f a c t i n an y o f s e v e r a l w a y s , e . g . b y t h e i r

b e h a v i o u r i n t h e r o o s t ( l e s s o r more a c t i v e t h a n t h o s e t h a t

f a i l e d t o f i n d f o o d ? ) ; by l e a v i n g t h e r o o s t e a r l y t h e f o l l o w i n g

n i g h t ; b y d e p a r t i n g from t h e r o o s t p u r p o s e f u l l y i n a s e t d i r e c t i o n

o r b y some o t h e r w a y s .

One p o s s i b l e m e tho d b y w h ich i n f o r m a t i o n c o u ld

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

on t h e f u r o f b a t s ( t h a t h ad f e d s u c c e s s f u l l y t h e n i g h t b e f o r e )

by o t h e r members o f t h e r o o s t . T h i s w ou ld o b v i a t e t h e n e e d f o r -

v i s u a l s i g n a l s when t h e b a t s l e a v e t h e r o o s t ( a s e n v i s a g e d i n

one form o r a n o t h e r by Ward and Z a h a v i (1 9 7 3 ) f o r b i r d s ) . V i s u a l

s i g n a l s g i v e n when t h e b a t s l e a v e t h e r o o s t may b e o f l i m i t e d

v a l u e a f t e r d a r k .

I f t h e b a t s i n t h e r o o s t a r e a b l e t o i d e n t i f y

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

217

e x p e r i e n c e t h e y know l o c a l i t i e s a t w h ic h e a c h s p e c i e s o f foo d

p l a n t g ro w s , t h e n , e i t h e r by t a s t i n g p o l l e n on t h e f u r o f

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

f u r o f o t h e r s , b a t s c o u l d d i s c o v e r w h ic h t r e e s p e c i e s t h e o t h e r s

h a d o b t a i n e d fo o d from and t h e y c o u l d t h e n s e a r c h f o r t h e same

s p e c i e s . I n t h i s way k n o w le d g e o f t h e o n s e t o f new f l o w e r i n g

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

t h e new fo o d s o u r c e f u l l y u t i l i s e d . I n f o r m a t i o n on how much fo o d

one b a t h a d o b t a i n e d f rom a p a r t i c u l a r s o u r c e c o u l d e a s i l y b e

co m m u n ica ted b y v a r i o u s r e f i n e m e n t s , e . g . b y i t s b e h a v i o u r o r

by t h e q u a n t i t y o f p o l l e n on i t s f u r . ( P r e s u m a b l y a b a t t h a t

h a d s a t i s f i e d i t s a p e t i t e w o u ld b e l e s s i n c l i n e d t o i n g e s t a l l

t h e p o l l e n from i t s f u r t h a n one t h a t h a d f a i l e d t o f i n d enough

f o o d . )

T h i s m e tho d f o r d i s t r i b u t i o n o f k n o w le d g e o f ^ f o o d

s o u r c e s i s h y p o t h e t i c a l . H ow ev er , t h e r e i s c i r c u m s t a n t i a l

e v i d e n c e t o show t h a t i t c o u l d o p e r a t e . Many E . s p e l a e a r e t u r n i n g

t o t h e r o o s t a t Gua S a n d i n g h a d p o l l e n on t h e i r f u r ; t h e f i r s t

two b a t s c a p t u r e d on t h e i r r e t u r n t o t h e c a v e a t 0 2 . 0 0 h r s h a d /

t h e i r b o d i e s so h e a v i l y c o a t e d w i t h p o l l e n o f Bombax v a l e t o n i i

t h a t - t h e i r f u r a p p e a r e d y e l l o w i s h i n s t e a d o f t h e n o r m a l b row n

c o l o u r . S t a r t (1 9 7 2 ) found, t h a t members o f a c a v e r n i c o l o u s

c o lo n y o f R o u s e t t u s a e g y p t i a c u s i n K enya h ad a v e r y h e a v y d e p o s i t

0:f A d a n s o n ia d i g i t a t a p o l l e n on t h e i r f u r a t a t i m e when many

o f them w e re f e e d i n g on t h e f l o w e r s o f A. d i g i t a t a .

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

i n g e s t e d p o l l e n from th e f u r o f a d u l t s ( p r o b a b l y t h e i r m o t h e r s )

l a t h e r o o s t came from a n a l y s i s o f p o l l e n i n t h e a l i m e n t a r y

c a n a l s o f s u b a d u l t s c a u g h t i n B a tu C a v e s : : O ver 90 % o f t h e

p o l l e n r e c o v e r e d from m o s t o f them was D uabanga g r a n d ! f l o r a

p o l l e n s u g g e s t i n g t h a t t h e y h a d f e d i n t h e h i l l s t o t h e e a s t

o f B a tu C a v e s , y e t m o s t o f them a l s o c o n t a i n e d a l i t t l e (u p t o

5%, se ld o m m ore) S o n n e r a t i a a l b a p o l l e n , S . a l b a grew 38 km o r

more t o t h e w e s t o f B atu . C a v e s . S u b a d u l t s w e re s e ld o m c a u g h t

i n t h e m a n g ro v e s b u t t h e r e was a f l u s h o f S . a l b a f l o w e r s a t

t h e t im e t h e s e b a t s w e re c a u g h t and m o s t o f t h e a d u l t f e m a le s

h a d b e e n f e e d i n g on S . a l b a p o l l e n t h e n i g h t b e f o r e t h e y w ere

c a u g h t .

F o r a g i n g b y E. s n e l a e a i s n o t a lw a y s ran d o m .

The r e c a p t u r e o f b a t s a t p r e c i s e l y t h e same s i t e s a t w h ich t h e y

w ere c a u g h t w eeks o r m on th s e a r l i e r a t R a n ta u P a n j a n g and t h e

F . S . C . , U lu Gombak (w h e re one was c a u g h t f o u r t i m e s i n t h r e e

m o n th s ) show s t h a t some b a t s r e t u r n t o t h e same f e e d i n g a r e a s

on more t h a n one n i g h t . T h i s i n d i c a t e s t h a t t h e y may h a v e a

p r e c i s e k n o w le d g e o f t h e i r f e e d i n g r a n g e s o r a t l e a s t p o r t i o n s

o f them an d so ca n r e l o c a t e p o t e n t i a l fo o d s o u r c e s w i t h o u t

random s e a r c h . I f t h e b a t s c a n i d e n t i f y p o l l e n on t h e f u r o f

o t h e r s i n t h e r o o s t a p r e c i s e k n o w le d g e o f t h e p o s i t i o n o f t r e e s

o f t h a t s p e c i e s i n a f a m i l i a r f e e d i n g r a n g e w o u ld e n a b l e t h e

b a t s t o u t i l i s e t h a t s p e c i e s im m e d i a t e l y t h e y l e a r n e d t h a t i t

was f l o w e r i n g and w i t h o u t h a v i n g t o s e a r c h r a n d o m ly f o r i t .

Where the food source may be a long way from the roost this

ab i li ty would be of enormous value.

219

k . FORAGING METHODS.

The f o r a g i n g m e th o d s o f t h e two M a c r o g lo s s u s

s p p . and E . s p e l a e a a r e a l s o i n m arked c o n t r a s t t o one a n o t h e r .

I n e a c h g e n u s t h e m e th o d s em p lo yed a r e r e l a t e d t o t h e n a t u r e

o f t h e i r fo o d s o u r c e s . Thus t h e two M a c r o g lo s s u s s p p . w h ic h h ad

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

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

e a c h n i g h t . They p r o b a b l y become v e r y f a m i l i a r w i t h t h e i r

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

m oving from one t o a n o t h e r o f t h o s e a l r e a d y know n. Thus t h e y

a r e a b l e t o f e e d s i n g l y and f e e d i n g f l o c k s w ou ld b e u n n e c e s s a r y

..or e v e n d i s a d v a n t a g e o u s w h e re t h e fo o d s u p p l y was n o t v e r y r i c h .

S i n c e t h e i r r o o s t s i t e s a r e c l o s e t o o r i n t h e i r f e e d i n g r a n g e s

t h e y a r e a b l e t o ' b e g i n f e e d i n g a s so o n a s d a r k n e s s f a l l s . No

d i f f e r e n c e s i n t h e f e e d i n g r a n g e s o f m a le s an d f e m a l e s o r a d u l t s

an d s u b a d u l t s w ere n o t i c e d .

. I n t h e s e r e s p e c t s , t h e y a r e t y p i c a l o f some o t h e r -

s m a l l p t e r o p i d s ( J o n e s , 19 7 2 ; van D eusen and P e t e r s o n , 1 958 ;

p e r s . o b s . ) . H o w ev e r , no d e t a i l e d s t u d i e s o f an y o t h e r s m a l l

p t e r o p i d s w e re fo u n d i n t h e l i t e r a t u r e and t h i s l e a v e s a w ide

f i e l d o f r e s e a r c h t h a t u r g e n t l y r e q u i r e s a t t e n t i o n .

E . s p e l a e a , on t h e o t h e r h a n d , s o m e t im e s h a s t o

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

e . g . t h o s e b a t s f rom B a tu C aves f e e d i n g on S o n n e r a t i a a l b a m u s t

h a v e t r a v e l l e d a t l e a s t 38 km b e f o r e t h e y c o u l d b e g i n f e e d i n g

a l o n g t h e c o a s t ( a l t h o u g h t h e y so m e t im e s f e d from o t h e r p l a n t s

en r o u t e ) . The s c a r c i t y o f S . a l b a p o l l e n i n a l l t h e Gua S a n d in g

g uano s a m p le s s u g g e s t s t h a t S . a l b a grew to o f a r ( a b o u t 60 km)

220

from the cave for the bats to regularly feed from i t .

Alternatively, they may not have had to travel so far i f

S. caseolaris could provide a l l their food requirements when

other plants were insuffic ien t for their needs. (The pollen

of S. caseolaris was more abundant in Gua Sanding guano than

T> ~ 4 - , . n 4 - 1 * ----------- 4 ^ T ~ 4- « « « « h r , V s * •C 'rsv .i J C.l/U V /a v o o ^ucwAU CU1U. irU O i c JLO ca. x .u u m u x o o u. .l. o au n -U x Gjl

_S. c a s e o l a r i s t h a n f o r S . a l b a c l o s e t o Gua S a n d i n g . )

M os t o f t h e fo o d s p e c i e s o f E . s p e l a e a p r o d u c e

l a r g e n u m b ers o f f l o w e r s and when i n b loom many b a t s can- f e e d

s i m u l t a n e o u s l y from, one t r e e . Thus E . s p e l a e a i s a b l e t o f e e d

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

t h a t f r e q u e n t l y h a s t o s e a r c h Jiuge a r e a s o f l a n d f o r s c a t t e r e d

f l o w e r i n g t r e e s b e c a u s e by s e a r c h i n g a s members o f a f l o c k ( a l l

o f w h ich c a n s h a r e a fo o d s u p p l y l o c a t e d b y one o f them ) t h e y

c a n im p ro v e t h e e f f i c i e n c y o f t h e s e a r c h i n g .

T h e r e a r e v e r y few r e f e r e n c e s i n t h e l i t e r a t u r e

t o p t e r o p i d s f o r a g i n g i n f l o c k s b u t t h a t i s no i n d i c a t i o n t h a t

many s p e c i e s do n o t do s o , p a r t i c u l a r l y t h o s e t h a t t r a v e l l o n g

d i s t a n c e s . McCann ( 1 9 M ) i m p l i e d t h a t l a r g e n u m b ers o f R o u s e t t u s

l e s c h e n a u l t i f e d f rom v a r i o u s t r e e s i n I n d i a . I r e c o r d e d upw ard s

o f 100 R . a e g y p t i a c u s f e e d i n g from one b a o b a b ( A d a n s o n ia d i g i t a t a )

i n K enya ( S t a r t , 1 9 7 2 ) . A l th o u g h I n e i t h e r saw t h e b a t s a r r i v i n g

o r d e p a r t i n g d u r i n g my o b s e r v a t i o n t h e l a r g e num ber o f b a t s

g a t h e r e d a r o u n d one t r e e s u g g e s t s t h a t ^ w e r e members o f one o r

more f l o c k s .

N o t h in g i s known o f t h e c o m p o s i t i o n o f t h e f l o c k s

£ • ' s p e l a e a e x c e p t t h a t t h e y may c o n t a i n a d u l t s o f e i t h e r s e x

a n d s u b a d u l t s . W h e th e r t h e c o m p o s i t i o n o f t h e f e e d i n g f l o c k s

221

r e m a in s c o n s t a n t o v e r n i g h t o r from n i g h t t o n i g h t and w h e th e r

t h e y i n v o l v e b a t s from t h e same r o o s t i n g g r o u p s w ou ld a l l b e

o f p a r t i c u l a r i n t e r e s t t o know.

One p o i n t i s c l e a r . U n l ik e many o f t h e g r a m i n i v o r o u s

an d some o f t h e o t h e r b i r d s w h ic h Ward and Z a h a v i (1 9 7 3 ) d e s c r i b e d ,

E . s p e l a e a f l o c k s do n o t e x h a u s t a f o o d s u p p l y b e f o r e t h e y move

on i n s e a r c h o f o t h e r s . Thus one f l o c k a f t e r a n o t h e r was a b l e

t o f e e d f o r a w h i l e from t h e d u r i a n t r e e s a t t h e F . S . C . , U lu

Gombak. A t R a n ta u P a n j a n g t h e i m p r e s s i o n g a i n e d w as t h a t f l o c k s

° f E . s p e l a e a p a s s e d a l o n g t h e dyke o r t h e r i v e r o r t h r o u g h t h e

c u l t i v a t e d b a n a n a s (w h ic h d i d n o t grow i n g r o v e s a s t h i c k a s

t h o s e o f many w i l d s p e c i e s ) p a u s i n g b r i e f l y t o v i s i t f l o w e r s

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

b e f o r e m oving o f f . I t was a l s o o f i n t e r e s t t h a t a t t h e F . S . C .

when two f l o c k s m et an d f e d s i m u l t a n e o u s l y f rom one d u r i a n t r e e

t h e y d i d n o t a l l l e a v e t h e t r e e t o g e t h e r b u t one g ro u p l e f t

b e f o r e t h e o t h e r . I t w o u ld b e v e r y i n t e r e s t i n g t o know i f an y

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

s u c h t i m e s .

B e c a u s e t h e y h a v e t o t r a v e l some d i s t a n c e t o ■

t h e i r f e e d i n g g r o u n d s , E . s p e l a e a do n o t u s u a l l y b e g i n f e e d i n g

f o r an h o u r ( e . g . a t t h e F . S . C . ) o r an h o u r an d a h a l f ( e . g . a t v

R a n ta u P a n j a n g ) a f t e r d a r k . Thus e v en w h e re t h e y f e e d from t h e

p l a n t s u s e d b y e i t h e r M a c r o g l o s s u s s p p . t h e l a t t e r s p e c i e s c an

f e e d f o r some t i m e i m m e d i a t e l y a f t e r d a r k w i t h o u t an y c o m p e t i t i o n

from E . s p e l a e a . S i n c e t h i s i s t h e m o s t a c t i v e f e e d i n g t im e f o r

t h e two M a c r o g l o s s u s s p p . i t may b e o f c o n s i d e r a b l e v a l u e t o

th e m . The g r e a t e r m o b i l i t y o f E . s p e l a e a and i t s more v a r i e d

222

diet are of obvious advantage when i t i s in competition with

'the two Macroglossus spp.

A f u r t h e r a d v a n t a g e o f f o r a g i n g i n f l o c k s w ou ld

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

t h e l o c a t i o n o f v a r i o u s fo o d s o u r c e s an d t h e g e o g r a p h y o f t h e

f e e d i n g r a n g e o f th e c o l o n y . T h i s w o u ld b e u n u e e c e s s a r y for-

M a c r o g lo s s u s w h ich h a v e o n ly t o l e a r n r e l a t i v e l y s m a l l f e e d i n g

r a n g e s w h ic h a r e c l o s e t o o r s u r r o u n d t h e i r r o o s t s i t e .

An i n t e r e s t i n g f e a t u r e o f t h e r o o s t i n g and f e e d i n g

b i o l o g y o f E. s p e l a e a i s t h e s e g r e g a t i o n o f a g e an d s e x c l a s s e s .

E v id e n c e from t h e r a t i o s o f m a l e s , f e m a le s and s u b a d u l t s c a p t u r e d

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

a l i m e n t a r y c a n a l s o f b a t s c a u g h t a t E a t u C aves c l e a r l y shows

t h a t more a d u l t f e m a l e s t h a n m a le s o r s u b a d u l t s go t o t h e c o a s t

t o f e e d and t h a t s u b a d u l t s r e m a i n f a i r l y c l o s e t o B a t u C a v e s .

S i n c e young b a t s a r e p r o b a b l y n o t a s s t r o n g f l i e r s a s a d u l t s

i t i s t o b e e x p e c t e d t h a t t h e y do n o t - g o so f a r . H ow ever , t h e

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

E v id e n c e o f s e g r e g a t i o n i n t h e r o o s t i s n o t so

s t r o n g e x c e p t v/here a g ro u p o f s u b a d u l t s was s e e n t o r e g u l a r l y

r o o s t i n one p a r t o f Gua S a n d i n g . H ow ever , t h e s e x r a t i o s o f

t h e b a t s s h o t i n B a tu C aves and r e c o r d e d i n t h e I . M . R . r e g i s t e r

a r e s t r o n g l y i n d i c a t i v e . McCann (1 9 ^ 1 ) fo u n d t h a t t h e s e x e s o f

R o u s e t t u s l e s c h e n a u l t i w ere s e g r e g a t e d i n a c a v e r o o s t on S a l s e t t e

I s l a n d , Bombay.

S e g r e g a t i o n o f t h e m a l e s , f e m a le s an d s u b a d u l t s

i n t o d i f f e r e n t r o o s t i n g g r o u p s i n C av e rn C, B a tu C aves c o u ld

e x p l a i n why t h e p o l l e n c o n t e n t o f t h e guano a t the . f o u r c o l l e c t i o n

223

s i t e s was d i f f e r e n t . Thus T ra y I , w h ic h was V ery d i f f e r e n t from

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

i t h a d v e r y l i t t l e S o n n e r a t i a s p p . p o l l e n e s p e c i a l l y S . a l b a

b u t more p o l l e n f rom f o r e s t s p e c i e s t h a n t h e o t h e r t r a y s . T ra y I I

c o u l d h a v e b e e n u n d e r a g ro u p o f m a le s w h ic h f e d from S . a l b a

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

o t h e r w i s e . Hence t h e r e was a l o t o f S . a l b a p o l l e n b u t v e r y

l i t t l e S . c a s e o l a r i s i n t h e g u a n o . The b a t s a b o v e T ra y s I I I and

IV c e r t a i n l y c o n t a i n e d some a d u l t f e m a le s f o r two a b o r t e d em bryos

w e re fo u n d on T r a y I I I an d t h r e e on T ra y IV . T h e re was a l o t o f

S . a l b a an d S . c a s e o l a r i s on b o t h t h e s e t r a y s

T h i s e x p l a n a t i o n o f t h e d i f f e r e n c e s i n t h e p o l l e n

on t h e t r a y s i s p u r e l y c o n j e c t u r a l and t h e a c t u a l c o m p o s i to n o f

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

c o n c l u s i o n s c a n b e d ra w n .

5'. FLIGHT.

The m e th o d s o f F i n d l e y , S t u d i e r a n d W i l s o n (19 7 2 )

w e re u s e d t o com pare t h e w ing c h a r a c t e r i s t i c s o f M. m in im u s ,

M. s o b r i n u s an d E . s p e l a e a b e c a u s e t h e r e s u l t s a r e b a s e d onr(J til

s t a n d a r d m e a s u re m e n ts ( w e i g h t , f o r e a r m , 3 d i g i t and 5 d i g i t )

w h ic h c a n b e t a k e n from l i v e b a t s i n t h e f i e l d o r f rom museum

s p e c im e n s ( e x c e p t w e i g h t , u n l e s s t h e c o l l e c t o r n o t e d t h e w e i g h t

a t t h e t i m e o f c a p t u r e ) . They can t h u s be co m pared w i t h any

o t h e r s p e c i e s o f b a t f o r w h ic h d a t a o b t a i n e d from t h e same

m e a s u r e m e n t s , i s a v a i l a b l e . F i n d l e y , S t u d i e r and W i l s o n (1 9 7 2 )

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

22Zf

i n c l u d i n g M e g a l o g l o s s u s w oerm ann l P a g e n s t e c h e r ( a n A f r i c a n

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

t h e G l o s s o p h a g i n a e ( n e o t r o p i c a l n e c t a r i v o r o u s b a t s ) .

D a ta o b t a i n e d by o t h e r r e s e a r c h e r s u s i n g v a r i o u s

c o m p l i c a t e d u n s t a n d a r d i s e d t e c h n i q u e s v a r y from one a u t h o r t o

a n o t h e r , e . g . S t r u h s a k e r ( l 9 b l ) an d F a r n e y and F l e h a r t y ( i y 6 y )

b o t h m e a s u re d t h e w in g a r e a o f b a t s ( i n c l u d i n g t h e i n t e r f e m o r a l

membrane) by t r a c i n g t h e o u t s t r e t c h e d w in g s o n to g r a p h p a p e r

and m e a s u r in g t h e a r e a w i t h a p l a n i m e t e r . S t r u h s a k e r i n c l u d e d

t h e h e a d a n d b o d y i n h i s m e a s u re m e n ts o f t h e t o t a l w in g a r e a

and m e a s u re d t h e w ing sp a n from t i p t o t i p . F a r n e y and F l e h a r t y

d i d n o t i n c l u d e t h e h e a d and bo dy and m e a s u re d w in g l e n g t h o f

one w ing from t h e s i d e o f t h e b o d y t o t h e t i p . O t h e r w i s e , t h e i r

m e th o d s w e re e s s e n t i a l l y s i m i l a r . H ow ever , t h e i r r e s u l t s f o r

t h e same b a t s p e c i e s d i f f e r e n o r m o u s ly , s o m e t im e s b y a f a c t o r -

g r e a t e r t h a n two!

M* m in im us an d M. s o b r i n u s h a v e v e r y s i m i l a r w ing

c h a r a c t e r i s t i c s . T h e i r w in g s a r e l o n g and b r o a d . T h i s g i v e s them

a v e r y low a s p e c t r a t i o an d v e r y low w ing l o a d i n g . T h e i r w in g

t i p : f o r e a r m r a t i o i s h i g h .

The low w ing l o a d i n g e n a b l e s them t o f l y s l o w l y

w i t h o u t , h a v i n g t o w ork e x c e s s i v e l y h a r d t o p r e v e n t s t a l l i n g .

H ow ever , d r a g i n c r e a s e s a s t h e w in g l e n g t h , t h e s u r f a c e a r e a

an d t h e v e l o c i t y i n c r e a s e . T h e r e f o r e , t h e l o n g w in g a nd l a r g e

s u r f a c e a r e a p r e v e n t fas^t f l i g h t b e c a u s e t h e w ork r e q u i r e d t o

overcom e d r a g b ec o m e s p r o h i b i t i v e a t h i g h v e l o c i t y .

F i n d l e y , S t u d i e r and W il so n (1 9 7 2 ) p o i n t o u t t h a t

225

a h i g h w ing t i p : f o r e a r m r a t i o com b ined w i t h a low a s p e c t r a t i o \

i s c h a r a c t e r i s t i c , o f b a t s t h a t h o v e r . (They l i s t t h e G l o s s o p h a g i n a e ,

M u r in in a e an d K e r i v o u l i n a e a s e x a m p l e s . ) M e g a l o g l o s s u s w o e rm a n n l ,

l i k e o t h e r p t e r o p i d s t h e y s t u d i e d , h a d n e i t h e r a m a r k e d ly h i g h

w ing t i p : f o r e a r m r a t i o n o r low a s p e c t r a t i o and t h e y a s k "why

a r e t h e r e n o . s m a l l , h o v e r i n g f l o w e r b a t s i n t h e p a l e o t r o p i c s ? "

I n f a c t , M ac ro g l-o s su s s p p . a r e s m a l l f l o w e r b a t s t h a t c an h o v e r .

L ik e t h e G lo s s o p h 'a g in a e t h e y h a v e l o n g w ing t i p s and t h e a s p e c t -

r a t i o i n d i c e s o f M a c r o g lo s s u s s p p . a r e c o n s i d e r a b l y lo w e r t h a n

t h e mean a t t r i b u t e d t o t h e G l o s s o p h a g i n a e by F i n d l e y , S t u d i e r

and W i l s o n .

T h e i r w in g s h a p e t h u s a l l o w s b o t h M a c r o g lo s s u s

s p p . t o h o v e r an d t o f l y i n e n c l o s e d s u r r o u n d i n g s , e . g . w i t h i n

t h i c k b a n a n a g r o v e s and th r o u g h g a p s i n t h e f o l i a g e o f m angrove

t r e e s . They a r e h o w ev e r p r e v e n t e d from f l y i n g f a s t . ' ' S i n c e t h e y\

do n o t h a v e t o m i g r a t e ( a s many G l o s s o p h a g i n a e do ) t h e y c an

a f f o r d t o h a v e a l o n g e r w ing t i p and lo w e r w in g l o a d i n g t h a n

even t h e G l o s s o p h a g i n a e . I t i s s t r a n g e t h e r e f o r e t h a t m os t

G l o s s o p h a g i n a e c a n and on o c c a s i o n do f e e d fro m f l o w e r s w h i l e

h o v e r i n g i n f r o n t o f them b u t t h a t M a c r o g lo s s u s s p p . l a n d on

t h e f l o w e r s . (Com pare F i g . 4 5 o f M. m in im us s e t t l e d on a c u l t i v a t e d

b a n a n a i n f l o r e s c e n c e w i t h t h e p h o t o g r a p h i n W a lk e r ( 1 9 6 ^ , p . 2 6? )

o f G lo s s o p h a g a s o r i c i a n a P a l l a s h o v e r i n g i n f r o n t o f a c u l t i v a t e d

b a n a n a i n f l o r e s c e n c e w h i l e f e e d i n g . ) A p o s s i b l e e x p l a n a t i o n i s

o u t l i n e d on p . 2 3 9 *

S* spelaea i s again very different from the two

Macroglossus spp. in i t s f ligh t requirements, fligh t and wing

characteristics . I t has a re la tive ly short wing, a high aspect

226

ra tio (and so a high wing loading) but an average v/ing tip :

forearm ratio* The short v/ing and high aspect ra t io are

characteristic of long distance fligh t since they induce less

drag than long or broad wings would do. The resu ltan t high

wing loading prohibits slow f ligh t without s ta l l ing unless the

bats exert a great deal of work. However, the average wing tip:

forearm ra t io does indicate a potentially powerful propulsive

wing beat.

The shape and size of their wings allows

.§• spelaea to travel for long distances and at high velocities

but they prevent the bats from approaching flowers slowly or

hovering in front of them. They also prevent them taking off

d irectly from a flower a fte r feeding. Hence the bats resort to

the complex method of braking when approaching a flower (as

observed when they landed on Proxylum indlcum flov/ers) and

ro ll ing backwards and fa ll ing from a flower on departing so as

to gain suffic ien t speed to fly . Baker and Harris (1 9 5 7 )

observed Nanonycteris veldkampi (Jentink) performing the same

manoevures when departing from Parkia clappertoniana Keag flov/ers

in Ghana. (K. veldkampi i s usually a frugivorous species (Baker

and Harris, 1 9 5 7 ) . )

■v

6. REPRODUCTION.

At Rantau Panjang where food was continously

available, there was no noticeable seasonal variation in the

reproductive rate of M. minimus. Data was limited in the la t te r

part of the year and in January but pregnancy and lactation

occurred throughout the year. The l i t t l e evidence available

227

on t h e r e p r o d u c t i o n o f t h i s s p e c i e s i n Bako N a t i o n a l P a r k

s u g g e s t s t h a t i t may b r e e d s e a s o n a l l y t h e r e , p o s s i b l y i n

r e s p o n s e t o t h e f l o w e r i n g o f S o n n e r a t i a a l b a o r some o t h e r

p l a n t s p e c i e s . T h i s w o u ld r e p a y f u r t h e r w o rk .

Too few E . s p e l a e a w e re c a p t u r e d e a c h m onth t o

show w h e th e r o r n o t t h e r e was s e a s o n a l v a r i a t i o n i n t h e

r e p r o d u c t i v e r a t e : H ow ev er , t h e y to o b r e e d a l l y e a r . Lim and

B eck (1 9 7 3 ) r e c o r d e d two p e a k s i n t h e p r o p o r t i o n o f b a t s t h a t

w e re p r e g n a n t i n B a tu C aves on two s u c c e s s i v e y e a r s . Each p e a k

l a s t e d one t o two m o n th s ( a l t h o u g h t h e y r e c o r d a g e s t a t i o n

l e n g t h o f " s l i g h t l y more t h a n s i x m o n th s , p o s s i b l y a s l o n g a s

200 d a y s " ! ) . H o w ever , some o f t h e i r m o n th ly s a m p le s w e re v e r y

s m a l l an d t h e y m i s c a l c u l a t e d t h e d a t a on w h ic h t h e y drew t h e i r

g r a p h ( F i g . 2 , p . 2 5 5 ) and b a s e d t h e i r c o n c l u s i o n s . (T hey o n ly

r e c o r d e d t h e r e p r o d u c t i v e s t a t u s o f 8 0 2 b a t s ( p . 2 5 5 ) h u t

c a l c u l a t e d t h e p e r c e n t a g e o f p r e g n a n t and l a c t a t i n g f e m a le s

u s i n g t h e t o t a l num ber o f f e m a le s c a p t u r e d , i . e . 1060 ( T a b l e 1 ,

p . 2 5 4 ) . ) I f t h i s e r r o r i s t a k e n i s t a k e n i n t o a c c o u n t t h e p e a k s

i n t h e p r e g n a n c y r a t e a r e e m p h a s i s e d .

L i t t l e d a t a i s a v a i l a b l e on o t h e r M a l a y s i a n

p t e r o p i d s : Lim (1 9 7 0 ) d e s c r i b e d t h r e e s m a l l p e a k s i n t h e

r e p r o d u c t i v e r a t e o f C y n o p te r u s b r a c h y o t i s : T h e se a r e n o t

s i g n i f i c a n t . D a ta g a t h e r e d on C. b r a c h y o t i s d u r i n g t h e p r e s e n t

s t u d y f a i l e d t o c o n f i r m L i m 's f i n d i n g s . C. h o r s f i e l d i and

R o u s e t t u s a m p l e x i c a u d a t u s w e re fo u n d t o b r e e d t h r o u g h o u t t h e

y e a r t o o .

The e s t i m a t e d l e n g t h o f g e s t a t i o n o f M. m in im u s ,

228

a b o u t 120 d a y s , i s p r o b a b l y a c c u r a t e t o w i t h i n + 10 d a y s . I t

i s t h u s a b o u t e q u a l t o t h e l e n g t h o f g e s t a t i o n o f C y n o p te r u s

s p h i n x (80, t o 100 d a y s ; Moghe, 1 9 5 6 ) , R o u s e t t u s l e s c h e n a u l t l

( c a . 1 2 5 d a y s ; C h a u d h a r i i n P a t i l , 1 9 6 8 , q u o te d from R a c e y ,

1973) an d R. a e g y p t i a c u s ( c a . 120 d a y s ; K u l z e r , 1 9 5 8 , q u o te d

* r\/C c \ *u ___ _____J- X V14I n o o c v c a i ) | y v s J • U U W C I f O A y l / U O O O W G l / O C U. C? I I I U X C O J U c U i l U l i i '

t i m e s t h e w e i g h t o f M. m i n im u s . .

The e s t i m a t e d s p e c i f i c f o e t a l g ro w th v e l o c i t y o f

M. m in im us ( a = 0 . 0 1 7 ) i s lo w e r t h a n t h a t r e c o r d e d f o r any

o t h e r mammal a l t h o u g h i t i s o f t h e same o r d e r o f m a g n i tu d e a s

o t h e r b a t s (R a c e y , 1 9 7 3 ) . H o w ever , S c h m id t ( l 9 7 h ) r e c o r d e d a

g e s t a t i o n l e n g t h o f 20 5 d a y s f o r Desmodus r o t u n d u s (D esm odon-

t i d a e ) a n d W im s a t t a n d T r a p i d o (1 9 5 2 ) r e c o r d e d a b i r t h w e i g h t

o f 11 .5 g . U s in g t h i s d a t a t o com p u te a , a = 0 . 0 1 3 7 f o r D.

r o t u n d u s : H ow ev er , W a lk e r (1 9 6 ^ ) q u o te d a g e s t a t i o n l e n g t h o f

90 t o 120 d a y s f o r t h a t s p e c i e s and so t h i s v a l u e o f a s h o u l d

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

c o n f i r m e d . R acey (1 9 7 3 ) c a l c u l a t e d a f o r o n l y o ne o t h e r p t e r o p i d ,

P t e r o p u s g l g a n t e u s B runn ( a = 0 . 0 2 7 ) . D a ta d e r i v e d from K u l z e r ' s

( 1 9 5 8 , q u o te d from R o s e v e a r , 1965) r e s u l t s a r e s u f f i c i e n t t o

c a l c u l a t e a f o r R o u s e t t u s a e g y p t i a c u s a s a = 0 . 0 2 9 .

B eck a nd Lim (1 9 7 3 ) b a s e d t h e i r e s t i m a t e d l e n g t h

o f g e s t a t i o n o f E . s p e l a e a on two r e c a p t u r e r e c o r d s , n e i t h e r

o f w h ic h g i v e v e r y p r e c i s e e s t i m a t e s . No d a t a w as o b t a i n e d

d u r i n g t h e p r e s e n t s t u d y t o c o n f i r m o r r e f u t e t h e i r e s t i m a t e .

The s p e c i f i c f o e t a l g ro w th v e l o c i t y , a , c a l c u l a t e d u s i n g t h e i r

d a t a l i e s b e tw e e n 0 .0 1 3 and 0 . 0 1 k* T h i s i s ev e n lo w e r t h a n

t h a t o f _M. m in im us b u t i n v iew o f t h e i m p r e c i s e n a t u r e o f t h e

I!

229

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

No d a t a on t h e a c t u a l g ro w th r a t e s o f y oung o f

an y s p e c i e s w e re o b t a i n e d b u t d a t a p r e s e n t e d b y B eck and Lim

(1 9 7 3 ) f o r E . s p e l a e a a r e i n c l o s e a g r e e m e n t w i t h t h o s e o b t a i n e d

b y K u l jc e n ' ( 1 9 5 8 , q u o te d from R o s e v e a r , 1 965) f o r t h e s l i g h t l y

l a r g e r R. a o g y p t r a c u s . B o th a u t h o r s fo u n d g ro w th t o b e v e r y

r a p i d i n i t i a l l y b u t t o s lo w down a f t e r t h e f i r s t f l i g h t . I n d i r e c t

e v i d e n c e from t h e d u r a t i o n o f l a c t a t i o n an d t h e g ro w th r a t e o f . x

f r e e f l y i n g y ou ng M. m in im us s u g g e s t s a s i m i l a r g ro w th p a t t e r n

i n t h a t s p e c i e s .

E a r l y i n d e p e n d e n c e i s o b v i o u s l y o f c o n s i d e r a b l e

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

t h a t E_. s p e l a e a ^ w h i c h may t a k e a y e a r cbr two t o r e a c h s e x u a l

m a t u r i t y , c a n f l y when s i x w eeks o l d (B eck and Lim, 1973) w e ig h in g

o n l y 3 k t o k0% o f t h e a d u l t w e i g h t . To a c h i e v e t h i s , t h e i r w in g s

grow much f a s t e r t h a n t h e i r b o d i e s b e tw e e n b i r t h an d f i r s t

f l i g h t so. t h a t t h e f o r e a r m s h a v e r e a c h e d 70% o f a d u l t l e n g t h

a t f i r s t f l i g h t . F i r s t f l i g h t w o u ld a p p e a r t o b e g o v e r n e d by

w ing s i z e , n o t w e i g h t , and i t ' c o i n c i d e s w i t h t h e l o w e s t w ing

l o a d i n g e x p e r i e n c e d b y t h e b a t s . (D a v i s (1 9 6 8 ) fo u n d t h e same

f o r a m i c r o c h i r o p t e r a n b a t , A n t r o z o u s p a l l i d u s Le C o n te ,

V e s p e r t i l i o n i d a e , . • ,

V ery few l a c t a t i n g f e m a le M. m in im u s o r E , s p e l a e a

w e re c a u g h t i n t h e f e e d i n g ' a r e a s c a r r y i n g y oung ( a n d no M .s o b r i n u s

w e re c a u g h t c a r r y i n g y o u n g ) . Young t h a t w ere c a r r i e d w e re

e s t i m a t e d t o b e o n l y one o r two d a y s o l d . H o w ever , l a c t a t i n g

f e m a le s c a u g h t l e a v i n g B a tu C aves and Gua S a n d in g i n t h e e v e n i n g s

c a r r i e d t h e i r young W ith th e m . What becam e o f t h e y ou ng i s n o t

230

known b u t t h e y m u s t h a v e b e e n d e p o s i t e d , som ew here o u t s i d e t h e

c a v e . They may h a v e b e e n t a k e n t o t h e f e e d i n g g r o u n d s and

d e p o s i t e d t h e r e w h i l e t h e a d u l t s f o r a g e d . T h i s i s c o n t r a r y to

t h e f i n d i n g s o f B eck an d Lim (1 9 7 3 ) who c a u g h t l a c t a t i n g f e m a le s

c a r r y i n g young l e a v i n g B a tu C aves an d i n " f e e d i n g a r e a s " ,

r o a s i o x y t h e b a t s the,y c a u g h t i n " f e e d i n g a r e a s " w e re i n f a c t

en r o u t e t o o r f ro m f e e d i n g a r e a s and s t i l l c a r r y i n g t h e i r

y o u n g . K u l z e r ( 1 9 5 8 , q u o te d from R o s e v e a r , 1965) o b s e r v e d t h a t

R o u s e t t u s a e g y p t i a c u s f e m a le s c a r r y t h e i r young a l m o s t c o n s t a n t l y

f o r s i x w eeks o n ly s u s p e n d i n g them t e m p o r a r i l y on some f a m i l i a r

p e r c h w h i l e f e e d i n g .

A t w h a t a g e t h e s u b a d u l t s becam e i n d e p e n d e n t i s

n o t known b u t when t h e p o l l e n i n t h e a l i m e n t a r y c a n a l s o f

E . s p e l a e a c a u g h t i n B a tu C aves was a n a l y s e d , yo un g o n ly j u s t

c a p a b l e o f f l i g h t w e re fo u n d t o c o n t a i n b o t h m i lk and D uabanga

g r a n d i . f l o r a p o l l e n a s w e l l a s a s m a l l q u a n t i t y o f S o n n e r a t i a

a l b a p o l l e n . M ost a d u l t f e m a le s h ad f e d on S . a l b a and i t was

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

s u b a d u l t s was from t h e f u r o f a d u l t s , p o s s i b l y t h e i r m o t h e r s ,

w h ich h ad f e d fro m S . a l b a . I f t h i s i s t h e c a s e , i t w ou ld s u g g e s t

t h a t s u b a d u l t s t h a t w e re n o t y e t w eaned b u t w e re c a p a b l e o f

f o r a g i n g f o r t h e m s e l v e s d i d n o t n e c e s s a r i l y f o r a g e i n t h e same

a r e a s a s t h e i r m o t h e r s , . . . .

7 . BAT FLOWERS AND FLOWER BATS.

I n ' a r e v i e w o f c h i r o p t e r o p h i l y F a e g r i . and van

d e r P i j l ( l 966) t a b u l a t e d t h e f e a t u r e s t h e y r e c o g n i s e d a s t y p i c a l

o f " b a t f l o w e r s " and r e l a t e d them t o t h e " f l o w e r b a t s " b y w h ich

231

t h e y w ere p o l l i n a t e d . I t i s now c l e a r t h a t t h e t a b l e i s o v e r -

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

h a v e fo rm ed t h e b a s i s o f l a t e r d i s c u s s i o n s on c h i r o p t e r o p h i l y

( e . g . B a k e r , 1 9 7 0 , 19 7 3 ; P r o c t e r and Yeo, 1 97 3 ; V o g e l , 1 96 8 ,

1969) and t h e y fo rm a c o n v e n i e n t b a s i s f o r d i s c u s s i n g how t h e

b a t f l o w e r s i n P e n i n s u l a r M a l a y s i a a r e r e l a t e d t o t h e b a t s

w h ich f e e d f rom th e m . - •

N o c t u r n a l a n t h e s i s .

I n m o s t o f t h e p l a n t s - s t u d i e d a n t h e s i s was

n o c t u r n a l b u t t h e f l o w e r s o f many s p e c i e s op en ed j u s t b e f o r e

d a r k and s u r v i v e d i n s i t u a f t e r dawn so t h a t a n i m a l s b e s i d e s >

b a t s , e s p e c i a l l y b i r d s ( N e c t a ^ r i n i i d a e ) and b e e s ( M e la p o n id a e )

w e re a b l e t o v i s i t th e m . E x c e p t f o r Musa s p p . an d p o s s i b l y f o r

P a r k l a s p p . an d D u r io s p p . t h e b i r d s and b e e s w e re n o t t h o u g h t

l i k e l y t o a f f e c t p o l l i n a t i o n . B a k e r , C ruden an d B a k e r (1 9 7 1 )

and B a k e r (1 9 7 3 ) h a v e p a i d p a r t i c u l a r a t t e n t i o n t o s u c h v i s i t o r s

w h ic h t h e y r e g a r d a s p a r a s i t e s o f l i t t l e c o n s e q u e n c e t o t h e

p l a n t s a l t h o u g h t h e p r o v i s i o n o f fo o d by t h e p l a n t s may b e

i m p o r t a n t t o t h e a n i m a l s w h ic h v i s i t them and t h e y i n t u r n may

b e i m p o r t a n t p o l l i n a t o r s o f o t h e r p l a n t s p e c i e s .

T h e re w e re two e x c e p t i o n s t o n o c t u r n a l a n t h e s i s ,

i . e . Musa m a l a c c e n s i s ( a n d c u l t i v a t e d b a n a n a s ) an d Cocos n u c i f e r a .

i-i* ma-3-a c c e n s i s may e x p o s e f l o w e r s b y d ay o r b y n i g h t ; b y d ay

s u n b i r d s , e . g . A n t h r e p t e s and s p i d e r h u n t e r s , A r a c h n o t h e r a

( N e c t a r i n i d a e ) f r e q u e n t l y v i s i t e d th em . One v a r i e t y , M. m a l a c c e n s i s

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

(Y e l lo w i s t y p i c a l l y an e n to m o p h i l o u s c o l o u r a l t h o u g h t h e

232

o r n l t h o p h l l o u s Musa ( C a l l i m u s a ) b o r n e e n s j s I s a l s o y e l l o w . )\

T h is may d e n o t e an i n c i p i e n t d e p a r t u r e o f M. m a l a c c e n s i s from

c h i r o p t e r o p h i l y t o o r n o t h o p h i l y 6 r e n t o r a o p h i l y . Be t h a t a s i t

may, M. m a l a c c e n s i s i s v i s i t e d b y b a t s , , b i r d s an.d b e e s , a l l o f

w h ich c o l l e c t p o l l e n a nd a r e p r o b a b l y c a p a b l e o f p o l l i n a t i n g

i t s i n c e t h e y a l s o come i n c o n t a c t w i t h t h e s t i g m a . I t , i s n o t

s u r p r i s i n g t h e r e f o r e t h a t w h e re M. m a l a c c e n s i s i s s y m p a t r i c

w i t h o t h e r Musa s p p . h y b r i d i s a t i o n i s com m on-(C hen, 1 972 ;

Sim monds, 1 9 3 5 ) . T h e r e f b r e , i n a n s w e r t o van d e r F i j i ' s q u e s t i o n

"how ............. i s g e n e t i c a l p u r i t y m a i n t a i n e d b e tw e e n r e l a t e d

c h i r o p t e r o p h i l u s s p e c i e s . . . . ( o f ) . . . . w i l d Musa?" (v a n d e r P i j l ,

1 9 5 6 ) } g e n e t i c a l p u r i t y i s n o t a l w a y s ' m a i n t a i n e d w h e re s u c hI

s p e c i e s a r e s y m p a t r i c . (

L i t t l e i s known o f t h e a n t h e s i s o f C ocos n u c i f e r a

(W h i tm o re , 1973) b u t f l o w e r s w e re o b s e r v e d i n a n t h e s i s by day

and by n i g h t . S u n b i r d s ( N e c t a r i n i i d a e ) h a v e commonly b e e n r e p o r t e d

f e e d i n g among t h e i n f l o r e s c e n c e s o f c o c o n u t s ( S y m t h i e s , 1960)

an d b e e s ( M e la p o n id a e ) w e re f r e q u e n t l y s e e n am o n g s t th em . I n

v iew o f t h e u b i q u i t o u s d i s t r i b u t i o n o f c o c o n u t s i n t h e t r o p i c s ,

p a r t i c u l a r l y on t r o p i c a l i s l a n d s w h e re t h e y s e t f r u i t a l t h o u g h

b a t s a r e a b s e n t , c l i i r o p t e r o p h i l y c a n n o t b e e s s e n t i a l t o them .

(Van d e r F i j i (1 9 5 6 ) r e j e c t e d o l d r e p o r t s o f b a t s f e e d i n g on

c o c o n u t f l o w e r s b u t v an D eu sen (1 9 6 9 ) r e c o r d e d two c a s e s o f

P t e r o p u s s p p . f e e d i n g from them and showed t h a t c o c o n u t f l o w e r s

h a v e am ple r e s e r v e s o f fo o d w h ich he t h o u g h t w o u ld b e a t t r a c t i v e

t o b a t s .)

I

233

C o l o u r .

W h i t e , m a ro o n , l i v e r - c o l o u r e d o r p u r p l e a r e

t y p i c a l c o l o u r s o f h a t f l o w e r s i n t h e p a l e o - an d t h e n e o t r o p i c s

( e . g . B a k e r an d H a r r i s , 1 95 7 ; van d e r P i j l , 1 9 5 6 ; V o g e l , 1 9 6 8 -

6 9 )« T h i s i s t r u e o f s p e c i e s exam ined d u r i n g t h i s s t u d y e x c e p t

Cocos n u c i f e r a and M a n g i f e r a s p p , , n e i t h e r o f w h ic h a r e depehckaht

on c h i r o p t e r o p h i l y and b o t h o f w h ich h av e c ream ' to o r a n g e ( o r ,

M a n g i f e r a , s o m e t im e s r e d ) c o l o u r e d f l o w e r s . The f l o w e r c o l o u r

A re n g a s p p . i s d in g y y e l l o w .

W h i te i s o b v i o u s l y a s u i t a b l e c o l o u r f o r f l o w e r s

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

e y e - s i g h t t o l o c a t e f l o w e r s s u c h a s m a c r o g l o s s i n e b a t s . H ow ever ,

no e x p l a n a t i o n h a s b e e n p u t f o r w a r d f o r t h e m aroon and r e l a t e d

c o l o u r s commonly fo u n d on b a t f l o w e r s . To a human o b s e r v e r

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

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

g r o u n d . The f u n c t i o n o f p u r p l e o r m aroon c o l o u r s on b a t f l o w e r s

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

f o r t h e b a t s t o s e e th e m .

I n P e n i n s u l a r M a l a y s i a c h i r o p t e r o p h i l o u s p l a n t s

w i t h d a r k c o l o u r e d f l o r a l members i n c l u d e O royy lum i n d l c u m,

P a . j a n e l i a I on g i f o l i a , Musa s p p , and E u g e n ia m a l a c c e n s i s . E x c e p t

f o r E . m a l a c c e n s i s t h e I n f l o r e s c e n c e s o f a l l t h e s e s p e c i e s

a r e l o c a t e d on l o n g p e d u n c l e s t h a t h o l d them w e l l away from

t h e l e a v e s o f t h e p l a n t s . The i n f l o r e s c e n c e s a r e t h u s l i k e l y

t o b e s e e n a s s i l h o u e t t e s . The w h i t e r e s i n o u s pow d er on t h e

u n d e r s i d e s o f t h e l e a v e s o f some Musa s p p . may e n h a n c e t h e

Z3h

s i l h o u e t t e e f f e c t o f t h e i n f l o r e s c e n c e w h ic h h a n g s b e lo w them

b y p r o v i d i n g a p a l e b a c k g r o u n d a g a i n s t w h ic h t h e b a t s c an s e e

t h e i n f l o r e s c e n c e s . (B o th M a c r o g lo s s u s s p p . s e a r c h f o r b a n a n a

f l o w e r s from b e lo w an d t h e i r e y e s , e s p e c i a l l y i n M. s o b r i n u s

a r e d i r e c t e d p a r t i a l l y u p w a rd s : They a l s o h a v e b i n o c u l a r v i s i o n . )

On a l l t h e s e p l a n t s , e x c e p t E u g e n ia ma1a c c e n s i s

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

e n a b l i n g them t o d e t e r m i n e t h e p o s i t i o n and o r i e n t a t i o n o f t h e .

f l o w e r s o n c e t h e y h a v e l o c a t e d t h e i n f l o r e s c e n c e . I n P ro x y lu m

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

F a - . j a n e l i a l o n g i f o l i a t h e i n n e r s u r f a c e o f t h e p e t a l s i s

w h i t e an d i n c h i r o p t e r o p h i l o u s Musa s p p . t h e f l o w e r s t h e m s e l v e s

a r e w h i t e a g a i n s t a d a r k e r b r a c t . I n t h e m o s t s t r i c t l y c h i r o p ­

t e r o p h i l o u s s p e c i e s o f M u sa . M. t r u n c a t a , t h e a d a x i a l s u r f a c e

o f e a c h b r a c t i s w h i t e b u t t h e a b a x i a l s u r f a c e i s d a r k e r t h a n

i n an y o f t h e o t h e r s p e c i e s ( F i g . 1 3 ) .

I n P e n i n s u l a r M a l a y s i a c h i r o p t e r o p h i l o u s p l a n t s

t h a t h a v e p a l e c o l o u r e d o r w h i t e f l o w e r s i n c l u d e m o s t s p e c i e s

D u r i o , Bombax v a l e t o n i i , C e ib a p e n t a n d r a , a l l P a r k i a s p p . ,

D u ab ang a g r a n d i f l o r a , S o n n e r a t i a a l b a and jS. o v a t a . I n m o s t Of

t h e s e s p e c i e s t h e f l o w e r s h a v e s h o r t p e d u n c l e s an d t h e y a r e

e i t h e r p r o d u c e d on t h e b r a n c h e s ( D u r io s p p . , B. v a l e t o n i i ,

C. p e n t a n d r a ) o r t h e y a r e t e r m i n a l , s u r r o u n d e d b u t n o t e n c l o s e d

b y l e a v e s (D . g r a n d i f l o r a , S . a l b a , S . o v a t a ) . T h e se . f l o v / e r s

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

w h e re s i l h o u e t t e w o u ld u s u a l l y b e o f l i t t l e v a l u e i n a s s i s t i n g

t h e b a t s t o l o c a t e th e m . H ow ev er , t h e i r w h i t e c o l o u r d o e s show

up a g a i n s t a d a r k b a c k g r o u n d a t n i g h t and i t i s p r o b a b l y t h e

235

m o s t s u i t a b l e c o l o u r f o r t h e s e f l o w e r s . ( Bombax v a l e t o n i i and

so m e t im e s Co i b a p e n t a n d r a f l o w e r when t h e t r e e s a r e l e a f l e s s

and s i l h o u e t t e i s p r o b a b l y an i m p o r t a n t a d d i t i o n a l a i d t o t h e i r

l o c a t i o n by b a t s . )

P a r k i a s p p . a r e e x c e p t i o n a l b e c a u s e t h e i r p a l e

c o l o u r e d i n f l o r e s c e n c e s a r e i n a p o s i t i o n w h e re s i l h o u e t t e

w ou ld b e a u s e f u l a i d t o t h e i r l o c a t i o n b y b a t s and t h e y m ig h t

b e e x p e c t e d t o h a v e d a r k c o l o u r e d f l o w e r s . H ow ever , i t may be

s i g n i f i c a n t t h a t t h e A f r i c a n P a r k i a c l a p p e r t o n i a n a , w h ic h i s

c h i r o p t e r o p h i l o u s , d o e s h a v e d e e p r e d c o l o u r e d f l o w e r s (B a k e r

and H a r r i s , T957)* I n i t s mode o f n e c t a r p r o d u c t i o n an d s t o r a g e

i t i s a l s o b e t t e r a d a p t e d t o c h i r o p t e r p p h i l y t h a n a r e t h e

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

t h e i n f l o r e s c e n c e i n w h ic h a p o o l o f n e c t a r g a t h e r s (B a k e r and

H a r r i s , 1957)= M a l a y s i a n s p e c i e s l a c k t h e d e p r e s s i o n and h a v e

no s p e c i a l m echan ism f o r r e t a i n i n g a l a r g e vo lum e o f n e c t a r ,

S o n n e r a t i a c a s e o l a r i s f l o w e r s a r e b i - c o l o u r e d ,

t h e b a s a l h a l v e s o f t h e s t a m e n s a r e r e d and t h e d i s t a l p a r t s

a r e w h i t e ; S . c a s e o l a r i s f l o w e r s a r e l o c a t e d a t t h e e n d s o f1lo n g d r o o p i n g t w i g s , u n l i k e S . a l b a and S . o v a t a w h ic h f l o w e r

on s h o r t t w i g s ; S . c a s e o l a r i s f r e q u e n t l y h a s a t h i n crow n w i t h

f e w e r and s m a l l e r l e a v e s t h a n S>. a l b a and S . o v a t a . S i l h o u e t t e

may t h e r e f o r e p l a y a p a r t i n t h e l o c a t i o n o f S . c a s e o l a r i s

by b a t s and t h e r e d p o r t i o n s o f t h e s t a m e n s may e n h a n c e t h i s

w h i l e t h e w h i t e p o r t i o n o f t h e s t a m e n s can f u l f i l l t h e d o u b le

r o l e o f m ak in g t h e f l o w e r v i s i b l e i n s i t u a t i o n s w h e re s i l h o u e t t e

w ou ld n o t show up o r i n a i d i n g t h e b a t s t o d e d u c e t h e o r i e n t a t i o n

236

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

O d o u r .

None o f t h e c h a r a c t e r s o f b a t f l o w e r s a r e more

c o n f u s e d t h a n o d o u r . Many a u t h o r s e m p h a s i s e t h e r o l e o f odour^

F a e g r i an d van d e r F i j i (1 9 6 6 ) w r i t e " t o an i n v e s t i g a t o r w i t h

some e x p e r i e n c e t h e s m e l l o f b a t b lo s s o m s i s m o s t d e f i n i t e . I t

h a s much i n common w i t h t h e s m e l l o f t h e a n i m a l s t h e m s e l v e s .

............ The same k i n d o f o d o u r . . . . i s fou nd i n f r u i t s t h a t a r e

d i s p e r s e d b y b a t s ( e . g . t h e g u a v a ) The f a l l a c y t h a t b a t

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

l e d t o many o v e r s t a t e m e n t s . B a t f l o w e r s I ex am in e d w ere n o t

f r a g r a n t b u t I fo u n d no s i m i l a r i t y b e tw e e n t h e o d o u r s o f e . g .

S o n n e r a t i a s p p . (w h ic h a l l h a v e a s i m i l a r b u t n o t a p o w e r f u l

s c e n t ) , Musa s p p . , Dur.io s p p . o r g u a v a f r u i t s . H o l t t u m ( l 965)

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

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

C o rn e r (1 9 5 2 ) w r o t e o f O roxylum in d l c u m t h a t " t h e c o r o l l a b e g i n s

t o open a b o u t 10 pm and t h e h a r s h o d o u r e s c a p e s from th em .

By m i d n i g h t t h e l u r i d mouth g a p e s w i d e l y and i s f i l l e d w i t h

s t i n k " . A l th o u g h t h e y do s m e l l u n p l e a s a n t t o t h e human s e n s e s ,

t h e o d o u r i s n o t a s s t r o n g a s t h e s t a t e m e n t s u g g e s t s , e . g . w h i l e

p h o t o g r a p h i n g E o n y c t e r i s s p e l a e a v i s i t i n g t h e f l o w e r s I was

v / i t ’n i n 5 m o f e l e v e n f l o w e r s on tv/o t r e e s b u t I was u n a b l e t o

d e t e c t t h e i r o d o u r .

Macnae (1 9 6 8 ) s t a t e d t h a t S . a l b a f l o w e r s on

t h e E a s t A f r i c a n c o a s t h a v e no o d o u r and E a k e r (1 9 7 3 ) showed

t h a t m o th s a r e a t t r a c t e d t o f l o w e r s w i t h f r a g r a n t o d o u r s and

237

f l o v / e r s w i t h a " b a t s m e l l ’1. The dogma s u r r o u n d i n g t h i s t o p i c

i s b e g i n n i n g t o b re 'a k down t o more r e a l i s t i c p r o p o r t i o n s b u t

t h e r e i s l i t t l e t h a t c a n b e proved, u n t i l t h e n a t u r e an d t h e

c h e m i c a l c o m p o s i t i o n o f t h e s u b s t a n c e s p r o d u c i n g t h e " b a t o d o u r s ”

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

o f t h e D a ts t o them h a v e b e e n i n v e s t i g a t e d . I n t h e m ean t im e

c a r e m u s t b e e x e r c i s e d when d e s c r i b i n g t h e o d o u r o f b a t f l o w e r s .

F lo w e r s t r u c t u r e .

Two d i s t i n c t b u t v a r i a b l e t y p e s o f b a t f l o w e r

h av e b e e n r e c o g n i s e d b y m o s t a u t h o r s ( e . g . B a k e r , 1 97 3 ; F a e g r i and

d e r F i j i , 1 966 ; V o g e l , 1 9 6 8 -6 9 )« T h ese a r e t h e " s h a v i n g b r u s h "

o r " s ta m e n b a l l " f l o v / e r s and t h e " c a m p a n u la t e " - f l o w e r s .

They d i f f e r p r i n c i p a l l y ‘i n t h a t t h e f l o w e r s o f

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

i s l i b e r a l l y and g e n e r a l l y d e p o s i t e d on a b a t when i t l a n d s on

them w h e r e a s t h o s e o f t h e c a m p a n u la t e t y p e p r o d u c e r e l a t i v e l y

l i t t l e p o l l e n b u t t h e y d e p o s i t i t a c c u r a t e l y on a s p e c i f i c p a r t

o f t h e v i s i t i n g b a t ’ s b o d y . F lo w e r s o f t h e s h a v i n g b r u s h ty p e

can h a v e s i n g l e f l o v / e r s w i t h l a r g e num bers o f s t a m e n s ( e . g .

Bomb ax s p p . , S o n n e r a t i a s p p . ) o r many s m a l l f l o v / e r s i n a s i n g l e

i n f l o r e s c e n c e ( e . g . P a r k i a s p p . ) . A s p e c i a l v a r i e t y i s r e p r e s e n t e d

by D u r io s p p . and Ce i b a p e n t a n d r a i n w h ich i n d i v i d u a l f l o w e r s

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

t h u s m u l t i p l y i n g t h e e f f e c t o f e a c h f l o w e r ( F i g 2 5 ) •

The c a m p a n u la t e sy nd ro m e u s u a l l y i n v o l v e s s i n g l e

c a m p a n u la t e f l o w e r s . Musa s p p . a r e a b e r a n t members o f t h i s t y p e :

T h e i r f l o w e r s a r e n o t c a m p a n u la t e b u t t h e y d e - p o s i t a l i t t l e

238

p o l l e n a c c u r a t e l y on t h e m u z z l e s o f t h e b a t s t h a t i n s e r t t h e i r

m u z z le s i n t o t h e f l o w e r i n s e a r c h o f n e c t a r . Musa s p p , . f l o w e r s

a r e p ro d u c e d i n hands:: Thus e a c h f l o w e r c o n t r i b u t e s t o t h e

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

i n f l o r e s c e n c e a s a w h o le .

A l l t h e c h i r o p t e r o p h i l o u s f l o w e r s ex am in ed i n

P e n i n s u l a r M a l a y s i a a r e o f t h e s h a v i n g b r u s h t y p e e x c e p t Musa

s p p , , O roxylum in d i c u m and P a . j a n e l i a l o r i g i f o l i a . T h i s c o n fo rm s

w i t h t h e v ie w s e x p r e s s e d b y B a k e r (1 9 7 3 ) t h a t t h e s h a v i n g b r u s h

t y p e o f b a t f l o w e r i s more common t h a n t h e c a m p a n u la t e t y p e i n

t h e p a l e o t r o p i c s ( b u t n o t i n t h e n e o t r o p i c s ) . F lo v /e r s o f t h e

s h a v i n g b r u s h ty p e a r e more s u i t e d t o b a t s t h a t l a n d on t h e

f l o v /e r s t o f e e d v /h i l e t h e c a m p a n u la t e t y p e o f f l o w e r s a r e s u i t e d

t o b a t s t h a t h o v e r i n f r o n t o f t h e f l o v / e r s t o f e e d ( B a k e r , 19 73 )*

V o g e l ( 1 9 6 8 - 6 9 ) o b s e r v e d t h a t p a l e o t r o p i c a l

c a m p a n u la t e f l o w e r s a r e more r o b u s t and h av e d e e p e r c o r o l l a s

t h a n t h e i r n e o t r o p i c a l c o u n t e r p a r t s . A l s o , t h e p a l e o t r o p i c a l

c a m p a n u la t e f l o w e r s p r e s e n t t h e b a t s w i t h a l a n d i n g p l a c e

u n l i k e n e o t r o p i c a l f l o w e r s . T h ese o b s e r v a t i o n s a r e t r u e o f t h e

M a l a y s i a n . c a m p a n u la te b a t f l o w e r s t h a t w ere e x a m in e d , b e a r i n g

i n mind t h a t t h e f l o v / e r s o f Musa s p p . a r e many b u t s m a l l and

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

t h e b a t s i n p l a c e o f a r o b u s t c o r o l l a .

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

t h e f l o w e r s t h e y f e e d from i n P e n i n s u l a r M a l a y s i a an d a l l t h e

b a t s t h a t h a v e b e e n o b s e r v e d f e e d i n g from f l o w e r s i n A f r i c a ■

a l s o l a n d e d on t h e f l o w e r s t o f e e d ( e . g . B a k e r , 1 9 7 3 ; B a k e r

239

and. H a r r i s , 1 9 5 7 ,1 9 5 9 ; H a r r i s and B a k e r , 1 9 5 8 , 1 95 9 ; J a e g e r ,

19^+5; S t a r t , 1 9 7 2 ) . H o w ever , many n e o t r o p i c a l n e c t a r i v o r o u s

B a t s h o v e r i n f r o n t o f t h e f l o w e r s t o f e e d ( e . g . B a k e r , 1 9 7 0 ,

1973 ; Cockrum and H ay w ard , 1 962 ; Hayward and C ockrum , 1 971 ;

V o g e l , 1 9 6 8 - 6 9 ) .

B a k e r (1 9 7 3 ) was p r o b a b l y c o r r e c t when h e

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

and t h e r o b u s t b u i l d o f t h e few c a m p a n u la t e f l o v / e r s among t h e

p a l e o t r o p i c a l c h i r o p t e r o p h i l o u s p l a n t s w ere r e l a t e d t o t h e

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

f l o w e r s t o f e e d . H o w ev er , h i s s t a t e m e n t t h a t p a l e o t r o p i c a l

n e c t a r i v o r o u s b a t s a r e l a r g e r and l e s s a c t i v e ( B a k e r , 1973)

i s n o t s t r i c t l y t r u e o f M a c r o g lo s s u s s p p . They a r e no l a r g e r

t h a n some n e o t r o p i c a l s p e c i e s , e . g . C h o e r o n y c t e r i s m e x ic a n a

(10 t o 20 g) and L e p t o n y c t e r i s s p p . (18 t o 30 g) ( w e i g h t s

q u o te d from W a l k e r , 1 96^) and t h e y a r e c a p a b l e o f h o v e r i n g

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

f o l l o w i n g r e a s o n

H e i t h a u s , Op.ler and B a k e r (197^-) fo u n d t h a t

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

B a u h a n ia p a u l i e t a P e r s . ( C a e s a l p i n i a c e a e ) i n C o s t a R i c a . The

l a r g e r s p e c i e s i n v a r i a b l y l a n d e d on t h e f l o v / e r s an d t h e i r

w e i g h t was s u f f i c i e n t t o b e n d t h e b r a n c h e s , s o i n v e r t i n g t h e

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

s p e c i e s u s u a l l y h o v e r e d i n f r o n t o f t h e f l o w e r s t o f e e d . When

t h e y d i d l a n d t h e i r w e i g h t was i n s u f f i c i e n t t o i n v e r t t h e

f l o w e r s an d t h e y h a d t o l a p t h e n e c t a r .

2M5

S i n c e M a c r o g lo s s u s m in im us i s h e a v y enough to

i n v e r t , t h e f l o v / e r s o f S o n n e r a t i a s p p . when i t l a n d s on them

i t ‘m a y 'h e a d v a n t a g e o u s f o r them t o l a n d i n s t e a d o f h o v e r i n g

i n f r o n t o f t h e f l o v / e r s t o f e e d .

N e c t a r and P o l l e n .

A l l t h e c h i r o p t e r o p h i l o u s f l o v / e r s ex am in ed i n

P e n i n s u l a r M a l a y s i a p r o d u c e v e r y l a r g e q u a n t i t i e s o f p o l l e n

and n e c t a r b u t n o t a s much a s h a s b e e n r e p o r t e d f o r some

c h i r o p t e r o p h i l o u s f l o v / e r s e l s e w h e r e ( Ochroma l a g o p u s '] m l;

'■2.° g r a n d i f l o r a 15 ia l( F a e g r i and van d e r P i j l , 1 966) . F a e g r i

and v an d e r P i j l (1 9 6 6 ) a ssu m ed t h a t n e c t a r p r o v i d e d t h e

e n e r g y r e q u i r e m e n t s o f n e c t a r i v o r o u s b a t s and p o l l e n was t h e i r

’’s o l e s o u r c e o f p r o t e i n ’* b u t t h i s i s n o t t r u e o f E jp jnyc i^r is

s p e l a e a o r M a c r o g lo s s u s m i n im u s .

Guano o f E . s p e l a e a c o l l e c t e d f rom E a t u C aves

o f t e n c o n t a i n e d p l a n t f r a g m e n t s b e s i d e s p o l l e n . They w ere

p a r t i c u l a r l y common when few Group I I p l a n t s w e re i n f l o w e r

a n d t h u s a t t i m e s when t h e b a t s w ere p r o b a b l y s h o r t o f p o l l e n

and n e c t a r s u p p l i e s . I t i s n o t known w h a t p l a n t t i s s u e s t h e y

a t e b e s i d e s p o l l e n b u t p o s s i b l y t h e y a t e f r u i t . The r e m a i n s o f

a b a s i d i o m y c e t e f r u i t i n g bod y i n t h e s to m ach o f one E. s r e l a e a

s e r v e s t o e m p h a s i s e t h e i r a d a p t a b i l i t y t o o t h e r fo o d s o u r c e s .

The p r e s e n c e o f i n s e c t s c a l e s , f r a g m e n t s o f

c h i t i n an d am o rp h o u s m a t e r i a l o f t e n c o n t a i n i n g o i l d r o p l e t s i n

t h e s to m a c h c o n t e n t s and f a e c e s o f E . s p e l a e a and e s p e c i a l l y

M. m in im us s u g g e s t t h a t t h e y f e d from l a r g e i n s e c t s , p r o b a b l y

m o ths ( L e p i d o p t e r a ) . T h i s u n d o u b t e d l y s u p p l e m e n t s t h e i r p r o t e i n

241

i n t a k e from p o l l e n . M e g a c h i r o p t e r a n s f e e d i n g on f r u i t s o r

l e a v e s u s u a l l y chew t h e foo d i t e m s i n g e s t i n g t h e l i q u i d and

r e j e c t i n g t h e f i b r o u s t i s s u e t h a t r e m a in s \.van Som m eren , 1 97 2 ;

W a lk e r , 1964* p . 1 8 7 ; p e r s , o b s . ) I t i s r e a s o n a b l e t o a ssum e

t h a t t h e y w ou ld t r e a t m oths i n t h e same w ay , t h u s e x p l a i n i n g

why o n l y s c a l e s , odd f r a g m e n t s o f c h i t i n and u n i d . e n t i f i a . b l e

am o rp ho us m a t t e r r e m a i n e d . W h e th e r o r n o t t h e s m a l l a r t h r o p o d s

s u c h a s m i t e s and T h r i p s t h a t t h e b a t s i n g e s t e d c o n t r i b u t e d

a n y t h i n g t o t h e i r n u t r i t i o n i s unknown b u t t h e i r e x o s k e l e t o n s

w e re u s u a l l y undam aged and t h e y may n o t h a v e b e e n o f a n y v a l u e

t o t h e b a t s .

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

t o h a v e o r i g i n a t e d from i n s e c t i v o r o u s a n c e s t r a l s t o c k and so

i t i s n o t s u r p r i s i n g t h a t many o f them s t i l l a c t i v e l y f o r a g e

f o r i n s e c t s t o s u p p l e m e n t t h e i r d i e t ( e . g . B a i t e r , 1973) b u t i t

was n o t e x p e c t e d t h a t i n s e c t s m ig h t b e c a p t u r e d an d e a t e n by

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

from f r u g i v o r o u s a n c e s t r a l s t o c k .

P o s i t i o n o f t h e i n f l o r e s c e n c e .on t h e p l a n t .

T h i s t o p i c h a s b e e n commented on b y m o s t r e v i e w e r s

b u t e s p e c i a l l y v an d e r P i j l (1 9 4 1 ) who e m p h a s i s e d t h e f r e q u e n c y

o f c a u l i f l o r y an d f l a g e l l i f l o r y i n t r o p i c a l p l a n t s . He s u g g e s t e d

t h a t m o s t i n s t a n c e s w e re an a d a p t a t i o n t o b a t s , e i t h e r f o r

c h i r o p t e r o p h i l y o r f o r c h i r o p t e r o c h o r y . (He f u r t h e r d i s c u s s e s

t h e a d a p t a t i o n s o f t h e p l a n t s f o r c h i r o p t e r o c h o r y i n van d e r

P i j l , 1 9 6 9 . )

E x am p le s o f b o t h t h e s e c o n d i t i o n s w e re fo u n d

22*2

d u r i n g t h e p r e s e n t s t u d y ( e . g . o f c a u l i f l o r y D u r lo g r a v e o l e n s

and o f f l a g e l l i f l o r y a l l P a r k l a s p p . ) b u t t e r m i n a l i n f l o r e s c e n c e s

and r a m i f l o r y a r e more common. T e r m i n a l i n f l o r e s c e n c e s w e re

e i t h e r on s h o r t l e a f y tw ig s ( e . g . S o n n e r a t i a a l b a ) o r on a lo n g

r a c h i s ( e . g . P ro x y lu m i n a i c u m ) .

A l th o u g h m o s t a u t h o r s c o r r e c t l y d e s c r i b e t h e

p o s i t i o n o f t h e i n f l o r e s c e n c e a s b e i n g i m p o r t a n t i n a l l o w i n g

a c c e s s f o r t h e b a t s t o f l y up t o i t and t o d e p a r t a f t e r f e e d i n g

F a e g r i an d van d e r P i j 1,(1 966) a l s o commented on i t s im p o r t a n c e

i n e n a b l i n g t h e b a t s t o l o c a t e t h e i n f l o r e s c e n c e ( a l t h o u g h t h e y

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

l o c a t e t h e i r f o o d ) . T h i s h a s b e e n e m p h a s i s e d i n t h i s s t u d y i n

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

p o s i t i o n .

B e f o r e co m m enting on t h e a c c e s s i b i l i t y o f

v a r i o u s i n f l o r e s c e n c e s t o n e c t a r i v o r o u s b a t s , p a r t i c u l a r c a r e

m u s t b e t a k e n t o e x a m in e , n o t j u s t t h e p o s i t i o n o f t h e

i n f l o r e s c e n c e on an i s o l a t e d p l a n t , b u t t o e x a m in e t h e n o rm a l

mode o f o c c u r r e n c e o f t h e p l a n t and t o e s t a b l i s h t h e f l i g h t

c a p a b i l i t i e s o f b a t s t h a t a r e l i k e l y t o v i s i t i t . The n e e d f o r

s u c h c a r e i s i l l u s t r a t e d b y Musa s p p . t h a t a r e v i s i t e d b y b a t s

i n P e n i n s u l a r M a l a y s i a : The i n f l o r e s c e n c e s o f t h e s e Musa s p p .

a r e p e n d u l o u s an d b o t h M a c r o g lo s s u s s p p . and E o n y c t e r i s s p e l a e a

a r e a b l e t o l a n d on an d f e e d from th em . H o w ever , w i l d Musa s p p .

u s u a l l y grow i n d e n s e g r o v e s and E . s p e l a e a i s i n c a p a b l e o f

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

o f m a n o e v e u r in g s u f f i c i e n t l y s l o w l y and d e x t r o u s l y t o f l y

b e tw e e n t h e b a n a n a p s e u d o s t e m s .

C o n c l u s i o n s ,

The u n i v e r s a l i t y o f t h e f e a t u r e s t h a t

c h a r a c t e r i s e h a t f l o w e r s i s w e l l i l l u s t r a t e d h y t h e f a c t t h a t

c h i r o p t e r o p h i l o u s p l a n t s from t h e p a l e o t r o p i c s , w h e re t h e y a r e

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

o f t h e s u b o r d e r M i c r o c n i r o p t e r a when t h e y a r e t r a n s p l a n t e d t o

t h e n e o t r o p i c s ( e . g . D u r io z i h e t h i n u s (B o m b aca ceae ) from S o u th

E a s t A s i a w h e re i t i s v i s i t e d h y M e g a c h i r o p t e r a n s , was v i s i t e d

hy h a t s i n H o n d u ra s w h e re t h e r e a r e no M e g a c h i r o p t e r a n s ( B a k e r ,

1970) an d one n e c t a r i v o r o u s h a t f rom M ex ico , M u s o n y c t e r i s

h a r r i s s o n i S c h a ld a c h and M c L a u g h l in ( P h y l l o s t o m a t i d a e ) h a s

o n l y b e e n c a u g h t i n a s s o c i a t i o n w i t h b a n a n a s ( h e n c e i t s g e n e r i c

name j W a lk e r , 1 9 6 4 ) J y e t t h e r e a r e no i n d i g e n o u s b a n a n a s i n

t h e new w o r l d ; t h e y h a v e b e e n i n t r o d u c e d from S o u th E a s t A s ia

(S im m onds, 1 96 2 )!

C o n v e r s e l y , n e o t r o p i c a l c h i r o p t e r o p h i l o u s p l a n t s

t r a n s p l a n t e d t o t h e p a l e o t r o p i c s a r e v i s i t e d t h e r e h y Mega­

c h i r o p t e r a n s , e . g . v an d e r P i j l (1 9 3 6 ) r e c o r d e d h a t s v i s i t i n g

Agave a n g u s t i f o l i a ( A m a r y l l i d a c e a e ) and C r e s c e n t i a c u . j e t e

( B i g n o n i a c e a e ) i n J a v a .

N e v e r t h e l e s s , t h i s s t u d y shows t h a t i n M a l a y s i a

d i f f e r e n t s p e c i e s o f n e c t a r i v o r o u s b a t s f e e d from d i f f e r e n t

s p e c i e s o f p l a n t s and b o t h t h e b a t s and t h e p l a n t s a r e ad ap . te d

t o one a n o t h e r . M ost p a p e r s i n w h ich t h e c h a r a c t e r s o f h a t

f l o w e r s and f l o w e r h a t s a r e d i s c u s s e d o v e r s i m p l i f y t h e f e a t u r e s

t h a t c h a r a c t e r i s e t h e t a t s and t h e f lo w e r s - b e c a u s e t h e y f a i l t o

t a k e i n t o a c c o u n t t h e d i f f e r e n t r e q u i r e m e n t s and c a p a b i l i t i e s

o f t h e v a r i o u s s y m p a t r i c s p e c i e s o f n e c t a r i v o r o u s b a t s . T h i s

i s p r o b a b l y b e c a u s e v e r y l i t t l e w ork h a s b e e n c a r r i e d o u t on

t h e s e s p e c i e s o f b a t s . I n f u t u r e r e v i e w s o f t h i s t o p i c , t h e

s p e c i f i c r e q u i r e m e n t s and a t t r i b u t e s o f b o t h t h e b a t s and t h e

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

SUMMARY . 2i| ,5

The d i e t s and some r e l a t e d a s p e c t s o f t h e b i o l o g y

o f t h r e e s p e c i e s o f n e c t a r i v o r o u s b a t s w ere s t u d i e d i n M a l a y s i a

from J a n u a r y 1972 t o J a n u a r y 197^-. M a c r o g l o s s u s m in im us G e o f f r o y

v/as r e s t r i c t e d t o c o a s t a l d i s t r i c t s and r e l i e d h e a v i l y on t h e

m angrove g e n u s S o n r i e r a t i a f o r fo o d b u t a l s o u s e d many o t h e r p l a n t s .

M a c r o g l o s s u s s o b r l n u s (A n d e r s e n ) f e d m o s t l y from w i l d b a n a n a s

(Musa s p p . s e c t i o n Eumusa) an d v/as c a u g h t w h e r e v e r t h e s e b a n a n a s

g re w . M ost p l a n t s p e c i e s u s e d b y b o t h M a c ro g lo s s x is s p p . f l o w e r

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

E o n y c t e r i s s p e l a e a Dobson f e d on a w id e v a r i e t y o f

p l a n t s i n c l u d i n g t h o s e u s e d b y b o t h M a c r o g lo s s u s s p p . , b u t m a i n ly

from s e a s o n a l l y f l o w e r i n g t r e e s t h a t p r o v i d e d l a r g e q u a n t i t i e s o f

fo o d f o r l i m i t e d p e r i o d s .

B o th M a c r o g l o s s u s s p p . w ere p re s u m e d t o r o o s t i n

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

a l o n e , d i d n o t t r a v e l f a r and w ere c a p a b l e o f g r e a t d e x t e r i t y i n

f l i g h t . They h a v e l o n g , b r o a d w in g s . E . s p e l a e a r o o s t e d i n v a s t

c o l o n i e s and f o r a g e d i n f l o c k s u p t o 38 kra o r m ore from t h e r o o s t .

They h a v e s h o r t n a r r o w w in g s p e r m i t t i n g f a s t f l i g h t and l o n g

d i s t a n c e t r a v e l b u t l i m i t i n g t h e i r d e x t e r i t y . T h e se f a c t o r s a r e

a l l r e l a t e d t o t h e fo o d r e s o u r c e s u s e d b y e a c h s p e c i e s . The

communal r o o s t s o f E ' . ' s p e la e a may s e r v e a s c e n t r e s f o r t h e d i s s e m i n ­

a t i o n o f i n f o r m a t i o n on t h e l o c a t i o n o f fo o d s o u r c e s .

B r e e d i n g o c c u r s t h r o u g h o u t t h e y e a r i n a l l t h r e e

s p e c i e s , an d i n M.m in im us no s e a s o n a l v a r i a t i o n was d e t e c t e d i n

P e n i n s u l a r M a l a y s i a , b u t may h a v e o c c u r r e d i n B o r n e o . I n s u f f i c i e n t

d a t a w e re o b t a i n e d t o show an y p e a k s i n t h e r e p r o d u c t i v e r a t e o f

M .s o b r i n u s o r E . s p e l a e a . E a r l y p r e g n a n c y o v e r l a p p e d l a c t a t i o n i n

a l l t h r e e s p e c i e s and a p o s t p a r t u m e s t r o u s was t h e r e f o r ^ s u s p e c t e d .

ACKNOW LED G EM EN TS.

I am s i n c e r e l y g r a t e f u l t o D r . A . .G .M a r s h a l l ,

D r . P . S . A s h t o n an d D r . D .R .W e l l s f o r . s u p e r v i s i n g me d u r i n g

t h e s t u d y , f o r t h e i n t e r e s t an d e n c o u ra g e m e n t t h e y showed

and f o r t h e t im e t h e y a l l g a v e so g e n e r o u s l y . To a l l o f them

T r s t , X ^ »t m u 1 - t lJL Vtxoa KsKJ C K J XJUCUXrV JUUi ' •

Numerous o t h e r p e o p l e h a v e g i v e n a d v i c e o r

h e l p e d me i n many w ays d u r i n g t h e p a s t t h r e e y e a r s . D r . J , : . .

M u l l e r , R e j k s h e r b a r i u m , L e id e n ; D r . G .T h a n i k a im o n i , P o n d i ­

c h e r r y I n s t i t u t e , I n d i a an d D r . E .S o epad m o , U n i v e r s i t y o f

M a la y a , a l l ex am in ed p o l l e n on my b e h a l f . D r . Soepadmo a l s o

l e n t me p h o t o g r a p h i c e q u ip m e n t and l a b o r a t o r y f a c i l i t i e s f o r

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T h i s s t u d y v/as s u p p o r t e d i n p a r t b y a K i l g o u r

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1

A p p e n d ix 1 ,

Taxonomy of the genus Macroglossus.

Introduction.

T h e re a r e two d i s t i n c t i v e s p e c i e s o f M a c r o g lo s s u s

i n P e n i n s u l a r M a l a y s i a . S i n c e t h e r e v i e w s o f A n d e r s e n (1911*

1912) t h e y h a v e b e e n known a s M a c r o g lo s s u s m in im us s o b r l n u s

A n d e rs e n an d M. l a g o c h i l u s l a g o c h i l u s M a t s c h i e , H o w ever , i n

t h i s s t u d y t h e y h a v e b e e n r e f e r r e d t o a s M. s o b r i n u s (A n d e r s e n )

and M. m in im us ( G e o f f r o y ) r e s p e c t i v e l y . The r e a s o n s f o r t h i s

c h a n g e w i l l b e p u b l i s h e d i n a r e v i e w o f t h e w h o le g e n u s ( S t a r t ,

i n p r e p . ) b u t f o r c l a r i t y t h e y a r e a l s o d e t a i l e d b e l o w .

E c o l o g i c a l d i s t r i b u t i o n s .

I n P e n i n s u l a r M a l a y s i a , b a t s r e f e r a b l e t o

M* l a g o c h i l u s l a g o c h i l u s ( s e n s u A n d e r s e n , 1911) w e re fo u n d t o

b e a s s o c i a t e d w i t h m a n g ro v e s o f t h e g e n u s S o n n e r a t i a and t h e i r

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

s h o r e i s l a n d s . I n c o n t r a s t , b a t s r e f e r a b l e t o M. m in im us s o b r i n u s

( A n d e r s e n , 1911) o c c u r r e d i n l a n d from t h e l o w l a n d s t o o v e r

1800 m a . s . l . b u t n e v e r i n m an g ro v es ( i d e n t i f i c a t i o n s b a s e d

on c h a r a c t e r s d e f i n e d by A n d e r s e n , 1 9 1 2 , s e e b e lo w ) (m ain t e x t ) .

R e c o rd s o f M. l a g o c h i l u s f rom Cameron H ig h l a n d s (L im , 1 9 6 6 ,

1973) were based on misidentified specimens (main text ) . How­

ever, in Borneo where M. m. sobrinus does not occur, M. lagochilus

lagochilus occurs in the mangroves but also a t ' many inland

lo ca li t ie s too (main te x t ) .

As M. l a g o c h i l u s ha d n o t b e e n r e c o r d e d from

J a v a I w i s h e d t o e s t a b l i s h t h e p r e s e n c e o r a b s e n c e o f M. m in im us

i n t h e m a n g ro v e s t h e r e . H ow ev er , I v/as p u z z l e d by t h e c u r i o u s l y

m ixed d i s t r i b u t i o n o f t h e two r a c e s o f M. m in im us ( i . e . M. m.

m in im us G e o f f r o y and H. m. s o b r i n u s A n d e rs e n ) r e c o r d e d on t h e

i s l a n d b y A n d e r s e n (1911? 1 9 1 2 ) . and by t h e a b s e n c e o f M. l a g o c h i l u s

w h ich h a d c o l o n i s e d a l l th e l a r g e r g r o u p s o f i s l a n d s an d many

s m a l l e r o n e s from S u m a t r a i n t h e w e s t t o t h e Solom on I s l a n d s

i n t h e e a s t and P h i l i p p i n e s i n t h e n o r t h t o N o r t h e r n A u s t r a l i a

i n t h e s o u t h b u t n o t J a v a . I t h e r e f o r e ex am in e d t h e r e l a t i o n ­

s h i p s o f t h e two J a v a n t a x a and M. l a g o c h i l u s .

Taxonomic h i s t o r y o f t h e g e n u s .

1 8 1 0 . G e o f f r o y - S t ~ H i l a i r e d e s c r i b e d P t e r o p u s m in im us from two

s k i n s ( a p p a r e n t l y v / i t h o u t s k u l l s ) c o l l e c t e d i n J a v a by

L e s c h e n a u l t de l a T o u r . The t y p e s p e c i m e n s a r e no l o n g e r

i n t h e P a r i s Museum ( A n d e r s e n , 1912 ; R o d e , 1 9^1) an d a r e

a p p a r e n t l y l o s t *

182/f . C u v ie r c r e a t e d t h e g e n u s M a c r o g lo s s u s f o r P . m in im us

G e o f f r o y .

1 8 37 - Temminck. A c c o r d in g t o A n d e rs e n (1 9 1 2 ) Temminck "v/as w e l l

a w a re o f t h e e x i s t e n c e o f l o n g e r and s h o r t e r m u zz led

i n d i v i d u a l s o f M. m in im u s” . He c o n c lu d e d t h a t t h e y w e re

" l o c a l ( r a c i a l ) r a t h e r t h a n i n d i v i d u a l d i f f e r e n c e s " . He

e x am in e d m a t e r i a l from S u m a t r a w i t h l o n g m u z z l e s and

from T im or w i t h s h o r t m u z z l e s . J a v a n s p e c i m e n s w ere

s i m i l a r t o t h o s e from S u m a t r a b u t w i t h s h o r t e r m u z z le s

t h a n t h e l a t t e r , Temminck was t h e r e f o r e t h e f i r s t p e r s o n

t o r e c o g n i s e more t h a n one form o f M a c r o g lo s s u s b u t he

d i d n o t name a n y o f th em .

1 8 6 7 . P e t e r s d e s c r i b e d M. m in im us v a r a u s t r a l i s from Q u e e n s l a n d ,

A u s t r a l i a .

1 8 8 8 . Thomas r e f e r r e d s p e c im e n s from A o la , Solom on I s l a n d s t o

M* a u s t r a l i s w h ic h he r e g a r d e d a s s p e c i f i c a l l y d i s t i n c t

from M. m in im us ( G e o f f r o y ) from J a v a . H is d e c i s i o n was

b a s e d on t h e s h o r t e r f o r e a r m o f t h e A o la s p e c im e n s

co m pared w i t h J a v a n s p e c im e n s and on d i f f e r e n c e s i n t h e

g r o o v e s o f t h e r h i n a r i a : I n t h e A o la s p e c i m e n s t h e r e was

a d i s t i n c t i n t e r n a r i a l g ro o v e r e a c h i n g t o t h e m a r g in o f

t h e u p p e r l i p . T h i s was n o t so i n t h e J a v a n spec im en s , he

h a d e x a m in e d .

1899• M a t s c h i e c r e a t e d t h e g e n u s S y c o n y c t e r i s f o r M, m in im us

v a r a u s t r a l i s P e t e r s an d d e s c r i b e d two new s p e c i e s o flb,

M a c r o g l o s s u s , M. l a g o c h i l u s from B u ru I s l a n d ( u n d e r w h ich

name h e i n c l u d e d Thom as ' ( 1 8 8 8 ) s p e c im e n s f rom A o la ) and

and M. n a n u s from t h e Aru I s l a n d s , B i s m a r c k A r c h i p e l a g o .

1911 • A n d e r s e n r e c o r d e d two fo rm s o f M. m in im us ( G e o f f r o y ) i n

J a v a . He r e f e r r e d t h e s m a l l e r , s h o r t m u z z le d fo rm , known

t o h im from J a v a , M adura and t h e K angean I s l a n d s , t o

M* B* m i n im u s , com m enting t h a t i t " i s p r o b a b l y t h e

t r u e l y i n d i g e n o u s J a v a n r a c e " . The s e c o n d , l a r g e r , l o n g e r

m u z z le d fo rm h e named M. m. s o b r i n u s , t y p e l o c a l i t y i n

P e r a k , M alay P e n i n s u l a r . I t .was knov/n t o h im from " m a in ­

l a n d S o u t h E a s t A s i a w hence i t h a s s p r e a d t o S u m a t r a

an d J a v a " . He a l s o d e s c r i b e d two more s u b s p e c i e s o f

H* l a g o c h i l u s M a t s c h i e , v i z . M. _1. pygmaeus and M. l . m i c r o t u s ,

and i n c l u d e d M. n a n u s M a t s c h i e w i t h i n t h i s s p e c i e s a s

M. 1 . n a n u s ,

1 9 1 2 . A n d e r s e n e x p a n d e d c o n s i d e r a b l y on t h e s t a t u s o f M. m in im u s ..

He e x am in e d t h e m e a s u re m e n ts o f a l l t h e M. m in im us

s p e c im e n s a v a i l a b l e t o him c l e a r l y s h o w in g t h a t a d i v i s i o n

i n t o two J a v a n e s e fo rm s ( i . e . M. m. m in im us and M. m.

s o b r i n u s i n h i s w ork) was j u s t i f i e d . H ow ev er , h e c o n c e d e s

t h a t t h e t y p e d e s c r i p t i o n o f M. m in im us ( G e o f f r o y ) i s to o

v a g u e t o i n d i c a t e w h ic h fo rm G e o f f r o y r e f e r r e d . The ty p e

m a t e r i a l b e i n g l o s t , h e e x p l a i n e d t h a t " i n t h e c i r c u m ­

s t a n c e s i t i s e x p e d i e n t t o f i x t h e name m in im us on t h e

s m a l l e r fo rm o f t h e s p e c i e s " w h ic h he r e g a r d e d a s t h e

" p r e d o m i n a n t " and " t r u e l y i n d i g e n o u s fo rm i n J a v a " .

He m a i n t a i n e d M. l a g o c h i l u s M a t s c h i e w i t h i t s

v a r i o u s s u b s p e c i e s a s a s p e c i e s d i s t i n c t f rom M, m. min im us

( G e o f f r o y ) an d M. m. s o b r i n u s A n d e r s e n .

1 9 2 7 . C h asen an d B oden K l o s s d e s c r i b e d a w e l l d e f i n e d fo rm o f

M. m i n im u s , M. m. f r a t e r n u s from t h e M entaw i I s l a n d s ,

d i s t i n g u i s h e d from M. m. s o b r i n u s A n d e r s e n b y i t s g r e a t e r

s i z e .

1 9 3 k • T a y l o r d e s c r i b e d a new s p e c i e s M. f r u c t i v o r o u s from t h e

P h i l i p p i n e s .

1 9 4 0 . C h asen r e t a i n e d A n d e r s e n ' s a r r a n g e m e n t f o r t h e M a l a y s i a n

fo rm s o f M a c r o g l o s s u s b u t n o t e d t h a t t h e r a n g e s o f

M• m. m in im us and M. m. s o b r i n u s "seem c u r i o u s l y m ixed

i n J a v a . M a t e r i a l i s n o t a v a i l a b l e t o j u s t i f y an y

a l t e r a t i o n " ,

5

The f o l l o w i n g i s a s y n o p s i s o f t h e named form s

a c c e p t e d t o d a t e .

M a c r o g lo s s u s m in im u s m in im us E . G e o f f r o y . R ange J a v a ,

i n c l u d i n g M adura an d K angean

I s l a n d s .

M. m. s o b r i n u s A n d e r s e n . R ange B a l i , J a v a , S u m a t r a , M alay

P e n i n s u l a r , T h a i l a n d , B urm a, Laos

and S o u th V ie tn a m .

M. m. f r a t e r n u s C hasen and K l o s s . Range S i p o r a an d S i b e r u t

I s l a n d s , Men.tawi I s l a n d .

M. l a g o c h i l u s l a g o c h i l u s M a t s c h i e . R ange M alay P e n i n s u l a r ,

S u m a t r a , B o rn e o , P h i l i p p i n e s ,

S u l a w e s i , B u r u , C eram , A m boina ,

B an d a and many o t h e r I n d o n e s i a n

i s l a n d s b u t e x c l u d i n g J a v a .

M. 1 , n a n u s ( M a t s c h i e ) . R a n g e M y s o l , New I r e l a n d , A d m i r a l t y

I s l a n d s , Aru I s l a n d s , B i s m a rc k

A r c h i p e l a g o and New G u in e a .

M. 1 . pygm aeus A n d e r s e n . R ange M u rray I s l a n d , T o r r e s S t r a i t s .

M. 1 . m i c r o t u s A n d e r s e n . Range Solom on I s l a n d s .

M. f r u c t l v o r o u s T a y l o r . Range P h i l i p p i n e s

A c o m p l e t e synonoiny i s g i v e n by A n d e r s e n ( 1 9 1 2 ) .

The s t a t u s o f M. m, f r a t e r n u s , M. 1 . n a n u s , M. 1 . p y g m a e u s ,

M. 1 . m i c r o t u s an d M. f r u c t i v o r o u s a r e n o t r e l e v a n t t o t h i s

s t u d y and a r e n o t c o n s i d e r e d i n d e t a i l b e l o w . The f o l l o w i n g

d i s c u s s i o n c o n c e r n s t h e s t a t u s o f M. m. m in im u s , M. m. s o b r i n u s

an d M. l a g o c h i l u s l a g o c h i l u s .

Points used by Andersen (1912) to distinguish M. minimus sobrinus

Andersen from M. lagochilus Matschie.

Andersen noted the following points

1 . Size, M. m. sobrinus i s altogether larger than M. lagochilus

although considerable overlap occurs (p. 768 - 770 in

w o u j t y \ j

2. Nares in M. m, sobrinus are "directed much more outward

than forward; in te rna r ia l groove (perhaps for that

reason) shallower than in M. lagochilus ............ some­

times d is tinc tly traceable as a fein t groove right

to the margin ’of the l ip , but more often quite obselete,*

la te ra l grooves.... i l l-defined or even practically

absent." In M. lagochilus, . . . ."na res directed halfgroove

outward half forward; internarial^deep and always

continued downward as a sharply defined groove right

to the margin of the upper lip; la te ra l groove ............

always well pronounced, . . . "

3 . Colour. Similar in M. minimus and M. lagochilus but with a

tendency for the underparts and the base of the fur

of the upper side of M. lagochi lus to average ligh ter.

Applying these three factors to West Malaysian

specimens, I have found that

1. Size. As stated by Andersen. Range of measurements more or

less as he gives, in the- fie ld and in specimens in

alcohol the re la tive length of the rostrum is very

noticeable, more so than measurements suggest.

2. In ternaria l grooves and la te ra l grooves. In live animals,

as stated by Andersen. However, particula rly in skins

and i n s p e c i m e n s i n a b s o l u t e a l c o h o l , t h e g r o o v e s may

b e p a r t i a l l y o r t o t a l l y o b s c u r e d ( e . g . i n ”M. l a g o c h i l u s 11

from C a re y I s l a n d , S e l a n g o r i n t h e I . M . R . c o l l e c t i o n ,

K u a la Lumpur N o s . 8 7 9 7 2 , 87975* 87976 ( a l l a d . )

an d No. 95625 from E a t u L a u t , K u a la L a n g a t , S e l a n g o r

t v i w T T v s A O O f N C n n a v i \.J.Ai AU j1 U Vy U U J. Uli | /

N .B . C hasen (1 9 ^ 0 ) a l s o fo u n d t h a t f o r some s k i n s ” a s

t h e n o s e s ................ a r e much s h r i v e l l e d , t h e i d e n t i f i c a t i o n s

.............. a r e n o t a b s o l u t e l y c e r t a i n . ”

The p o s i t i o n o f t h e n a r e s i s s u b j e c t i v e , n e v e r c l e a r

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

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

3* C o l o u r . ” M. m. s o b r i n u s ” i s c o n s i s t e n t l y d a r k e r t h a n ”M.

l a g o c h i l u s ” . ”M. l a g o c h i l u s ” i s much more v a r i a b l e

t h a n ”M. m. s o b r i n u s ” . A r a r e b u t d i s t i n c t c ream

c o l o u r e d form o f ”M. l a g o c h i l u s ” a l s o e x i s t s .

C o lo u r i s s e ld o m a s a f e d e f i n i t i v e c h a r a c t e r , p a r t i c u l a r l y i n

p r e s e r v e d m a t e r i a l w h e re t h e d i f f e r e n c e s b e tw e e n t h e s p e c i e s

a r e s m a l l . I n p r e s e r v e d m a t e r i a l t h e d e g r e e o f d e v e lo p m e n t o f

t h e g r o o v e s on t h e r h i n a r i u m c a n n o t a lw a y s b e u s e d w i t h

c e r t a i n t y t o s e p a r a t e t h e s e s p e c i e s . S i z e i s u s e f u l and i s n o t

a f f e c t e d s i g n i f i c a n t l y b y p r e s e r v a t i o n . H ow ev er , t h e r e r e m a i n s

a c o n s i d e r a b l e r e g i o n o f o v e r l a p .

The f o l l o w i n g p o i n t s do s a f e l y s e p a r a t e a l l

t h e a d u l t s p e c i m e n s from P e n i n s u l a r M a l a y s i a t h a t I h a v e s e e n .

1 . C o l l a r . I n a d u l t m a le '|M. l a g o c h i l u s ” t h e r e i s a w e l l d e f i n e d

t h i c k e n e d g l a n d u l a r c o l l a r a c r o s s t h e t h r o a t , w h ich

v a r i e s from b r i g h t r e d . t o d e e p c h o c o l a t e b r o w n ,

c o n t r a s t i n g s t r o n g l y w i t h t h e p a l e c o l o u r o f t h e s k i n .

In- b o t h a l c o h o l s p e c im e n s and d r i e d s k i n s t h i s c o l l a r

r e m a i n s d i s t i n c t , th o u g h l e s s so t h a n i n l i f e . Even

i n new b o r n young t h e i n c i p i e n t 15 c o l l a r i s v i s i b l e b u t

i t ri6V6r t h i c k e n s o r become bright?L.y c o l o u r s d U n t i l

a d u l t h o o d . Some a d u l t m a le *'M. m. s o b r i n u s " a l s o

p o s s e s s a f a i n t t r a c e o f a c o l l a r . I t i s n e v e r t h i c k e n e d

o r b r i g h t l y c o l o u r e d . T h i s d o e s n o t h o l d f o r J a v a n e s e

m a t e r i a l w h ere c o l l a r s a r e ev en p r e s e n t on fe m a le

"M. m. £ o b r i n u s n (Thomas and W ro u g h to n , 1 9 09 )*

M a n d i b u la r a p e x . I n *'M. ra. s o b r i n u s 11 t h e m a n d i b u l a r s y m p h y s i s

i s p r o j e c t e d f o r w a r d i n t o a d i s t i n c t c h i n ( F i g . 1 ) . The

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

t o t h e l o w e r m a r g in o f t h e jaw and p a s s i n g t h r o u g h t h e

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

i n c i s o r a nd a t t h e c a n i n e . I n "M. l a g o c h i l u s " t h e r e

may b e a m e n t a l r i d g e a t t h e m a n d i b u l a r s y m p h y s i s b u t

i t d o e s n o t p r o j e c t a s c h i n and t h e d e p t h o f t h e m a n d i b l e

i s c o n s i d e r a b l y g r e a t e r a t t h e c a n i n e t h a n a t t h e f i r s t

i n c i s o r . ( T h i s i s a l i t t l e l e s s p ro n o u n c e d i n s u b a d u l t

"M. m. s o b r i n u s 1*. )

I n l i v e a n i m a l s and s p e c im e n s p r e s e r v e d w i t h

t h e s k u l l i n s i t u t h i s i s r e f l e c t e d i n t h e d i s t i n c t

c h i n o f "M. m. s o b r i n u s 11 w h i l e "M, l a g o c h i l u s " a p p e a r s

c h i n l e s s a l t h o u g h t h e lo w e r l i p p r o j e c t s b e y o n d t h e

u p p e r i n b o t h s p e c i e s . The c h i n f u r t h e r e m p h a s i s e s t h e

a p p e a r a n c e o f a l o n g m u z z le i n "M. m. s o b r i n u s *1. T h i s

f e a t u r e h o l d s f o r s p e c im e n s ex am in ed from a l l p a r t s

o f t h e r a n g e s o f b o t h t a x a .

3* P o s i t i o n o f t h e l a c h r y m a l p i t . The a n t e r i o r e d g e o f t h e

l a c h r y m a l p i t i s a lw a y s a n t e r i o r t o t h e p o s t e r i o r

?edge o f m" ( u s u a l l y t h e l a s t m o la r b u t an e x t r a one

i s n o t i n f r e q u e n t ) i n "M. l a g o c h i l u s " b u t f r e q u e n t l y

p o s t e r i o r t o i t i n "M. m. s o b r i n u s 18.

From a c o m b i n a t i o n o f t h e s e c h a r a c t e r s ,

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

a b s e n c e o f an i n t e r n a r i a l g ro o v e e x t e n d i n g t o t h e l i p , t h e

f o r e a r m l e n g t h and t h e p r o f i l e o f t h e m a n d i b u l a r a p e x a l l

s p e c i m e n s ca n b e i d e n t i f i e d .

P o i n t s b y w h ic h A n d e r s e n (1 9 1 2 ) d i s t i n g u i s h e d "M. m. m in im us

from M. l a g o c h i l i t s .

A n d e r s e n (1 912) w r o t e t h a t ’’ t h e l e s s o u t w a r d l y

d i r e c t e d n a r e s a n d s t r o n g l y d e f i n e d v e r t i c a l g r o o v e on th e

u p p e r l i p " ( o f M. l a g o c h i l u s ) " a p p e a r t o b e t h e o n l y c h a r a c t e r s

b y w h ich t h i s fo rm ca n b e s a f e l y d i s t i n g u i s h e d f rom t h e s m a l l e r

r a c e o f M. m in im us ( i . e . M. m. m in im us ( G e o f f r o y ) i n h i s w o r k ) .

"The r e l a t i v e l e n g t h o f t h e r o s t r u m , t h e b r e a d t h o f t h e t e e t h

and t h e g e n e r a l s i z e o f t h e a n im a l seem s t o b e EXACTLY ALIKE"

(my c a p i t a l s ) " i n t h e two f o r m s . Any d i f f e r e n c e i n t h e p a l a t e

r i d g e s . . . . . d o e s n o t e x i s t . ( M a t s c h i e ' s s t a t e m e n t t o t h e

c o n t r a r y m u s t h a v e b e e n b a s e d on i n s u f f i c i e n t m a t e r i a l . ) "

I t h a s a l r e a d y b e e n shown t h a t i n p r e s e r v e d

s p e c im e n s t h e p r e s e n c e o r a b s e n c e o f t h e i n t e r n a r i a l g ro o v e

may b e d i f f i c u l t t o d e t e r m i n e and t h a t t h e d i r e c t i o n i n w h ich

1°

t h e n a r e s f a c e c a n b e a l t e r e d b y p r e s e r v a t i o n . I t t h e r e f o r e

seem ed p o s s i b l e t h a t M. m. m in im us G e o f f r o y an d M. l a g o c h i l u s

l a g o c h i l u s M a t s c h i e w e re sy n o ny m ou s . T h i s p o s s i b i l i t y was

s t r e n g t h e n e d when t h e d i s t r i b u t i o n s o f t h e two J a v a n e s e t a x a

was mapped ( F i g . 1 ) . M. m. s o b r i n u s v/as r e c o r d e d o n ly from t h e

h i n t e r l a n d and M. m. m in im us o n ly i n c o a s t a l l o c a l i t i e s o r o f f ­

s h o r e i s l a n d s .

Examination of museum specimens.

T h ro u g h t h e k i n d n e s s o f Mr, B o e a d i o f MuseumA

B o g o r e n s e , D r . D .R .W e l l s v/as a b l e t o exam in e a l l t h e Ma c r o g l o s s u s

m a t e r i a l i n t h a t c o l l e c t i o n on my b e h a l f . T h e re w e re n i n e

s p e c i m e n s from Bawean I s l a n d an d t h i r t y n i n e from J a v a . Those

from Bawean I s l a n d a r e c l e a r l y r e f e r a b l e t o M. l a g o c h i l u s

M a t s c h i e . ( F o r e a r m s 4 0 . 7 t n 4 2 . 0 mm; a l l w i t h s h o r t m u z z l e s ;

d e e p m e d ia n i n t e r n a r i a l g r o o v e s and t y p i c a l jaw p r o f i l e s . )

T h i k t y one o f t h e J a v a n e s e s p e c im e n s r e p r e s e n t '*M. m. s o b r i n u s 11

( f o r e a r m s 43*0 t o 4 6 . 5 mm; g r e a t e s t s k u l l l e n g t h 2 7 . 0 t o 2 9 . 2

mm, n — 11 ; c f . 2 8 . 5 t o 2 9 . 5 mm r e c o r d e d b y A n d e r s e n , 1 9 12 ;

l o n g m u z z l e s ; no i n t e r n a r i a l g r o o v e s and t y p i c a l jaw p r o f i l e s . )

The r e m a i n i n g e i g h t s p e c im e n s from J a v a a r e i n s e p a r a b l e from

l,M. l a g o c h i l u s 1* ( f o r e a r m s 41 .2. t o 4 4 «0 mm, n = 7 ; g r e a t e s t

s k u l l l e n g t h 2 5 . 4 t o 2 7 . 2 mm, n = 8 ; a l l h a v e s h o r t m u z z l e s ,

d e e p i n t e r n a r i a l g r o o v e s and t h e jaw p r o f i l e i s t y p i c a l o f

t h a t s p e c i e s i n a l l e x c e p t o n e , No. 9327 i n w h ich i t i s d e f o r m e d ) .

T h e re a r e no d e r m a l c o l l a r s on a n y o f t h e t h i r t y one s p e c im e n s

• .2* s o b r i n u s b u t on t h r e e o f t h e f o u r m a le M. l a g o c h i l u s

t h e r e i s a d i s t i n c t c o l l a r , t h e f o u r t h i s s u b a d u l t . F u r t h e r m o r e ,

F i g . 1 «, a . Map o f J a v a t o show t h e d i s t r i b u t i o n o f M acro '

g l o s s u s s o b r i n u s ( A n d e r s e n ) and M a c r o g lo s s u s

m in im us m in im us ( G e o f f r o y )

b . Jaw p r o f i l e s o f M a c r o g lo s s u s s o b r i n u s (A n d e r s e n )

and M a c r o g lo s s u s m in im u s m in im us ( G e o f f r o y ) (x 2

a p p r o x . ) t o show t h e " c h i n 1* o f M. s o b r i n u s t

a b s e n t f rom M. m in im u s m in im u s .

♦

M. s o b r i n u s200M. minimus minimusscale km

M. s o b r i n u s

t \ N s n n g

M. m. minimus

I

i

11

t h o s e r e f e r a b l e t o M. m. s o b r i n u s w e re from i n l a n d and t h o s e

r e f e r a b l e t o M. l a g o c h i l u s from c o a s t a l d i s t r i c t s .

T h e re w e re no s p e c i m e n s r e f e r a b l e t o M» m» m in im us

s e n s u A n d e r s e n ( 1 9 1 2 ) .

M r. J . E . K i l l and L o rd Medway b o t h ex am in e d a l l

t l i 0 -Tsvsnoso Eisit03?xs.1 i n "tli6 co!L 1 0 c tx o i i o f t l i s B r i t i s h Mus013.n1

( N a t . H i s t . ) u s i n g t h e c h a r a c t e r s d e f i n e d a b o v e t o d i s t i n g u i s h

t h e d i f f e r e n t t a x a . They c o n f i r m e d ( i n l i t . ) t h a t t h e r e w e re

o n ly two fo rm s r e p r e s e n t e d f rom J a v a and t h a t t h e s m a l l e r ,

f o r m e r l y r e f e r r e d t o M. m in im us m in im us ( G e o f f r o y ) b y A n d e rse n

(1 9 1 1 , 1912) was i n d i s t i n g u i s h a b l e from M. l a g o c h i l u s l a g o c h i l u s

M a t s c h i e .

I h a v e s u b s e q u e n t l y b e e n a b l e t o c o n f i r m t h i s

and t o exam ine two s p e c im e n s c o l l e c t e d b y t h e U .S . N a v a l R e s e a r c h

U n i t , NAMRU - 2 i n J a v a . They a l s o a r e i n d i s t i n g u i s h a b l e from

M* l a g o c h i l u s l a g o c h i l u s M a t s c h i e from P e n i n s u l a r M a l a y s i a .

I t i s c l e a r t h a t M. m in im us m in im us ( G e o f f r o y )

( s e n s u A n d e r s e n , 1 9 1 1 , 1912) i s synonym ous w i t h M. l a g o c h i l u s

l a g o c h i l u s M a t s c h i e and t h a t , a s i n o t h e r p l a c e s , t h i s t a x o n

i s s y m p a t r i c w i t h M. m in im us s o b r i n u s i n J a v a b u t t h e fo r m e r

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

The t a x o n o m ic i m p l i c a t i o n s .

T h e re i s no d o u b t t h a t t h e r e a r e two s p e c i e s o f

M a c r o g lo s s u s i n J a v a , t h e t y p e l o c a l i t y o f t h e g e n u s . H ow ever ,

t h e t y p e d e s c r i p t i o n o f t h e t y p e s p e c i e s i s t o o v ag u e t o d e f i n e

w h ich s p e c i e s was d e s c r i b e d , p o s s i b l y one o f e a c h s p e c i e s was

i n v o l v e d . The t y p e m a t e r i a l i s now u n t r a c e a b l e . a n d so i t i s

12

i m p o s s i b l e t o d e t e r m i n e w h i c h t a x o n i s t h e c o r r e c t t y p e s p e c i e s

o f t h e g e n u s . A n d e r s e n ( 1 9 1 1 » 1 9 1 2 ) b e l i e v e d o n l y o n e s p e c i e s

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

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

i . e . M a c r o g l o s s u s , m i n i m u s m i n i m u s . T h e r e a s o n s h e g a v e f o r h i s

d e c i s i o n n n c n o i v n l i c l n n d o x * b l i e I n i e n n o . i s i o n n l K n l e s 0" * 2 o ^ 3 o Q i c ^

N o m e n c l a t u r e .

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

t h e s e c i r c u m s t a n c e s .

1 . B y a p p l i c a t i o n t o t h e I n t e r n a t i o n a l C o m m i s s i o n o f Z o o l o g i c a l

N o m e n c l a t u r e ^ t o s u p p r e s s t h e n a m e m i n i m u s w i t h r e f e r e n c e t o

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

r e p l a c e i t b y t h e n e x t a v a i l a b l e s y n o n y m . S i n c e m i n i m u s i s t h e

s p e c i f i c n a m e o f t h e t y p e s p e c i e s o f t h e g e n u s t h i s w o u l d

n e c c e s s i t a t e t h e s u p p r e s s i o n o f t h e g e n e r i c n a m e a s w e l l

( A r t i c l e s 2+2b a n d 6 7 a o f t h e I n t e r n a t i o n a l C o d e o f Z o o l o g i c a l

N o m e n c l a t u r e ) . T h i s w o u l d b e u n d e s i r a b l e b e c a u s e t h e g e n e r i c

n a m e i s w e l l e s t a b l i s h e d a n d b e c a u s e t h e n e x t a v a i l a b l e

g e n e r i c n a m e , O d o n t o n y c t e r i s J e n t i n k , i s b a s e d o n a n a b e r e n t

s p e c i m e n .

2 . T o r e g a r d t h e l a r g e r t a x o n ( M . m . s o b r i n u s A n d e r s e n , 1 9 1 1 )

a s t h e t y p e s p e c i e s a n d t o d e s c r i b e , a n e o t y p e u n d e r t h e

p r o v i s i o n s o f A r t i c l e 7 9 o f t h e I n t e r n a t i o n a l C o d e o f Z o o l o g i c a l

N o m e n c l a t u r e . I f t h i s o p t i o n w a s a d o p t e d t h e e a r l i e s t a v a i l a b l e

n a m e f o r t h e s m a l l e r t a x o n ( M . m . m i n i m u s s e n s u A n d e r s e n , 1 9 1 1 ,

1 9 1 2 ) w o u l d b e M a c r o g l o s s u s r o s t r a t u s ( H o r s f i e l d ) f o r w h i c h a

c o t y p e e x i s t s i n t h e B r i t i s h M u s e u m ( N a t . H i s t . )

13

3 . To r e g a r d t h e s m a l l e r J a v a n e s e t a x o n (M. m. m in im us s e n s u

A n d e r s e n , 1911) a s t h e t y p e s p e c i e s and t o d e s c r i b e a n e o t y p e .

I f t h i s o p t i o n w e r e a d o p t e d , t h e n a m e M . l a g o c h i l u s M a t s c h i e

w o u l d b e c o m e a s y n o n y m o f M . m i n i m u s ( j e o f f r o y . M . m , s o b r i n u s

A n d e r s e n w o u l d b e r a i s e d t o s p e c i f i c r a n k a s M . s o b r i n u s

( A n d e r s e n ) *

T a x o n o m i c s t a b i l i t y , i n c o n f o r m i t y w i t h t h e

p r i n c i p l e s o f t h e I n t e r n a t i o n a l R u l e s o f Z o o l o g i c a l N o m e n c l a t u r e

(• Mayr» 1969) i s b e s t s e r v e d b y a d o p t i n g t h e t h i r d o p t i o n s i n c e

t h e g e n e r i c n a m e i s r e t a i n e d a n d n o s p e c i f i c n a m e s n e e d b e

r e s u r r e c t e d f r o m t h e j u n i o r s y n o n o m y o f a n y t a x a . T h i s i s t h e

c o u r s e t h a t h a s b e e n a d o p t e d .

14

B I B L I O G R A P H Y .

Andersen, K. 1 9 1 1 . Six new f ru i t bats of the genus Macro­

glossus and Syconycteris. Ann. Mag, nat. I-Iist..; 8:

641 - 6 4 3 .

A n d e r s e n , K . 1 9 1 2 . C a t a l o g u e o f t h e C h i r o p t e r a i n t h e c o l l e c ­

t i o n o f t h e B r i t i s h Museum. V o l . 1 ♦ M e g a c h i t o p t e r a .

p . 1 - 8 5 4 . B r i t . Mus. p u b l . L ond on .

C h a s e n , F .N . 1 9 4 0 . A h a n d l i s t o f M a l a y s i a n mammals. A s y s t e m -

■ a t i c l i s t o f t h e mammals o f t h e M alay P e n i n s u l a r ,

S u m a t r a , B o rn e o an d J a v a , i n c l u d i n g t h e a d j a c e n t s m a l l

i s l a n d s . B u l l . R a f f l e s M us. . 1J>: 22 - 5 9 .

C h a s e n , F .N . an d K l o s s , B .C . 1 9 2 7 . S p o l i a M e n t a w i e n s i a . -

Mammals. P r o c . Z o o l . S o c . L o n d o n , 5 3 : 797 - 8 4 0 .

C u v i e r , M .F . 1 8 2 5 . Des d e n t s d e s m a m m ife re s , c o n c i d e r e e s

comme c a r a c t e r e s z o o l o g i q u e s . L e . Norm.ant, P a r i s .

G e o f f r o y - S t . - H i l a i r e , E . 1 8 1 O. D e s c r i p t i o n d e s r o u s s e t t e s

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

d e c h a u v e s - s o u r i s . A n n . M u s . n a t n . H i s t . ’n a t . , P a r i s ,

1 § : 87 - 1 0 8 .

I n t e r n a t i o n a l c o d e o f Z o o l o g i c a l n o m e n c l a t u r e a d o p t e d b y t h e

XV I n t e r n a t i o n a l C o n g r e s s o f Z o o l o g y , L o n d o n . P u b l i s h e d

1961. R e v i s e d e d i t i o n 1 9 6 4 •

Lim B . L . 1 9 6 6 . A bundance an d d i s t r i b u t i o n o f M a l a y s i a n b a t s

i n d i f f e r e n t e c o l o g i c a l h a b i t a t s . F e d n . Mus. J ., 11 :

61 - 7 6 .

L i m B . L . 1973* B r e e d i n g p a t t e r n , f o o d h a b i t s a n d p a r a s i t i c

infestation of bats in Gunong Brinchang. Malay. Nat. J ..

2 6 : 6 - 1 3 »

M a t s c h i e , P „ 1 8 9 9 . D ie F led e rm au .se d e s B e r l i n e r Museums f u r

N a t u r k u n d e . I L i e f e r u n g d i e M e g a c h i r o p t e r a . p p . 1 - 102 . .

B e r l i n .

M ayr , E . 1969* P r i n c i p l e s o f S y s t e m a t i c Z o o lo g y , p p . 1 - 4 2 8 .

M c G ra w -H il l Book Co. New Y o rk , T o r o n t o , L on do n , S y d n e y .

P e t e r s , W. 1 8 6 7 . U eb e r d i e F l e d e r h u n d e , P t e r o p i , u nd I n s -

b e s o n d e r e u e b e r d i e A r t e n d e r g a t t u n g P t e r o p u s s . s .

M o n a t s b e r . A kad . W is e . B e r l i n » : 483 - 4 8 7 .

R od e , P . 1 9 4 1 . C a t a l o g u e d e s t y p e s de m am m ife res du Museum

N a t i o n a l d ' H i s t o i r e n a t u r e l l e , I I . O rd re d e s C h i r o p t e r e s .

B u l l . Mus. n a t n . H i s t , n a t . , s e r i e s 2 e , ij5: 73 - 1 0 5 .

T a y l o r , E .H . 1 9 3 4 . P h i l i p p i n e Land Mammals. M onograph 3 0 .

B u re a u o f S c i e n c e , M a n i l a .

Temminck, C . J , 183 7• M o n o g ra p h ie s d e s m am m alo g ie . V o l . 2 : 49

- 1 1 2 . D u fo u r and D' O c a g n e , P a r i s .

Thomas, 0 . 1 8 8 8 . The mammals o f t h e Solomon I s l a n d s b a s e d

on a c o l l e c t i o n made by Mr. C .M .W oodford d u r i n g h i s

s e c o n d e x p e d i t i o n t o t h e a r c h i p e l a g o . P r o c . Z o o l . S o c .

L o n d o n . 1 8 8 8 : 470 -• 4 8 4 .

Thomas, 0 . an d W ro u g h to n , R .C . 1909* On a c o l l e c t i o n o f

mammals from v / e s t e r n J a v a p r e s e n t e d t o t h e N a t i o n a l

Museum by Mr. W . E . B a l s t o n . P r o c . Z o o l . S o c . L o n d o n .

1 909 : 371 - 3 9 2 .

Appendix 2. Data relating to accuracy of pollen analysis methods.

Table 1.Two separate counts each of 500 pollen grains were made in Gua Sanding, Perak on 14th May 1972 and from each of

from one slide prepared from guano collected

seven slides prepared from guano collected inB a tu C a v e s , S e l a n g o r

c o u n t s .

( s a m p le Nos • 7,, 8 , 9:> 1 0 , 4 9 , 53 and 5 7 ) • The t a b l e shows t h e r e s u l t s o f t h e

Sam ple No. Gua S a n d in g 7 8 9 10 49 53 57

Coun t No. 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2

S o n n e r a t i a a l b a - - 0 . 2 0 .2 1 ..4 2 .5 5 . 9 7 . 5 3 . 2 4 . 8 3 . 6 6.1 0 . 6 0 . 6 3 . 2 5 . 6

S . c a s e o l a r i s 3 6 . 0 33 .1 - - - 0 .2 4 .1 2.1 0 . 2 0 . 2 2 . 6 3 . 5 0 . 2 1 .4 7 . 4 7 . 4

D u r io s p p . 16 . 4 21 .9 1 .4 1 .4 2,.4 4 .3 1 .8 1 .7 0 . 6 0 . 4 1 6 . 2 18.1 0 . 2 0 . 6 0 . 2 -

E u g e n ia s p p . 1 5 -4 1 9 .0 - - - - - - - 0 . 2 0 . 7 - 0 . 4 0 . 4 1 .4 0 . 2

A ren g a s p p . 7 . 4 9 . 2 - - - - - - - - - - - - - -D uabangag r a n d i f l o r a 1 0 .0 2 . 0 8 5 . 0 85 ,2 78,.6 77 .8 5 4 .0 5 2 .8 5 5 .8 5 6 .8 3 . 0 1 .6 6 . 2 7 . 0 3 . 0 1 . 4

P a r k i a s p p . 6 . 4 3 . 7 0 . 5 0 .6 1 ,.0 - 1 .4 1 .3 1 .2 1 .0 7 . 7 9 . 4 2 0 . 8 26 .1 4 5 . 0 46 .1

T8 3 . 0 5 . 5 0 . 2 0 .5 14,.9 13 .8 6 . 3 4 .1 8 . 0 7 . 2 5 . 3 3 . 2 2 2 . 0 21 .0 3 . 2 2 . 6

A r t o c a r p u s s p p . 0 . 8 3 . 5 8 . 7 7 .7 1 ,.7 1 .0 2 0 . 4 2 0 . 3 2 3 .4 21 .8 5 0 .3 5 2 .8 4 8 . 4 4 2 . 2 3 6 . 0 3 5 . 6

C e ib a p e n t a n d r a 0 . 8 - 1 .6 - 0 .5 9 . 0 9 . 0 ^ • 2 4 . 8 2 .1 3 . 7 - - - -

Orox.ylum in d ic u m _ _ _ . . _ _ - 0 . 4 - - _ _ _

T10 - - 3 . 4 2 . 7 - - 0 . 2 - 2 . 4 2 . 2 - - - - - -

T3 - - 0 . 4 - - - 1 .2 1 .1 0 . 2 0 . 4 - - 0 . 2 0 . 2 -

M a n g i f e r a s p p . - - - - - - 0 . 2 - 0 . 2 0 . 2 - - - - 0 . 2 -

Musa s p p . - - - - - - - - - - 2 .1 1 . 6 0 . 4 1 .0 0 . 6 0 . 6

? - - - - - - - - 0 . 2 0 . 2 0 . 2 0 . 2 0 . 4 - - -

The counts from each slide were compared by the any of the pairs of counts. This shows that the

t test. There was method used for

no significant difference (p > 0.05) between counting was adequate.

Table 2 .

Three subsam ples were c o l l e c t e d from guano cample

No. 6 from E a tu Caves. S l i d e s were p r e p a re d

i n d e p e n d e n t l y from each o f th e t h r e e subsam ples and

over 1000 g r a i n s were counted on each s l i d e . The

4* r > V , 1 r> V » r i « r n 4 - V i v * o i > "1 4 - r * r \ P +■ V i r \ c* rs r* P tr*V Ca. W Vli W X u u U>k WO W A VAX vy kJ W U VMU V>U |

S p e c ie s Subsample 1 Subsample 2 Subsample 3

S o n n e r a t i a a l b a7 2 2 ( 6 2 . 0 ) 7 86(64 .7 ) 7 59(65 .4 )

S. c a s e o l a r i s J . /Ceiba o en tan d ra 3 0 3 (2 5 .9 ) 332(27 .3 ) 2 9 2(25 .2 )

A r t o c a r r u s so p . 11 6 ( 1 0 . 0 ) 83( 6 .9 ) 99( 8 .5 )

P a r k i a sp p . 5( 0 .4 ) 4( 0 .3 ) 4( 0 .3 )

T2 p r e s e n t - -

T3 2 ( 0 . 2 ) 0 0

TZ* 1 6 ( 1 .4) 1 0 ( 0 . 8 ) 6 ( 0 . 5 )

Durio sp p . - - 1 ( 0 .1 )

T o t a l 11 64 121 5 11 61

Note F i g u r e s i n b r a c k e t s a r e p r e c e n t a g e s .

The t h r e e c o u n ts were compared by th e t t e s t .

There was no s i g n i f i c a n t d i f f e r e n c e (p > 0 . 0 5 ) . Th is shows

t h a t th e method used to c o l l e c t subsamples from th e samples

was a d e q u a te .

+30 -|

+ 2 0 -

co£ + 1 0 H

LUU

? 0

z oU - 1 0 - 4

- 20 -

-3 0- n r2 5 50 100

I200

“ 1— 4 00

“ I500

I3 0 0

n

F i g . 1 . G raph s h o w in g t h e 95% c o n f i d e n c e l i m i t s f o r n w h ere

n i s t h e num ber o f p o l l e n g r a i n s o f a s p e c i f i c t y p e

i n a c o u n t o f 5 0 0 p o l l e n g r a i n s made from a s am p le

c o n t a i n i n g more t h a n one p o l l e n t y p e . D a ta c a l c u l a t e d

from nomograms d e v i s e d fro m Maher ( 1 9 7 2 ) .

R e f e r e n c e .

M ah e r , L . J . j n r . 1 9 7 2 . Nomograms f o r c o m p u t in g 0 . 9 5

c o n f i d e n c e l i m i t s o f p o l l e n d a t a . R e v . P a l e o b o t .

P a l y n o l . , i j : 85 - 9 3 .

A p p en d ix 3 .

Data, r e l a t i n g t o w e e k ly guano c o l l e c t i o n s from

B a tu C a v e s , .S e l a n g o r .

A b b r e v i a t i o n s u s e d : -

Wo. = Sam p le n u m b er .

FW = F r e s h w e i g h t i n g r a m s .

%W1 = E s t i m a t e d d r y w e ig h t f o r w h o le s a m p le .

A = A bundance ( s u b j e c t i v e e s t i m a t e ) ,

+ 4 4 = v e r y a b u n d a n t ,

++ = a b u n d a n t ,

4. = l e s s a b u n d a n t

.CW = W a te r p e r c o l a t i n g i n t o t h e c a v e ,

+++ ? c a v e v e r y w e t ,

++ = c a v e w e t ,

+ = c a v e d r y

R a in = R a i n f a l l f o r e a c h w eek r e c o r d e d by t h e M e t e o r o l o g i c a l

D e p a r tm e n t a t B a tu C a v e s ,

W eekly Max. = Maximum r a i n f a l l r e c o r d e d o v e r 2 k h o u r s d u r i n g

t h e w e e k .

s e e p a g e 2 7 f o r d e f i n i t i o n s .

s e e p a g e 2 7 f o r

d e f i n i t i o n s ,

N o t e ; - A and CW a r e o n ly r e c o r d e d w i t h t h e d a t a f o r T ray I ,

No. F.V DW Date Rain V/eekly Max. A CW

3 - — — 19/2 - 2 5 /2 /7 2 - —

7 — — — 25/2 - 6 /3 ___

11 — — — 6/3 -1 3 /3 — --

15 — — — 13/3 -2 0 /3

19 53 20 /3 -2 7 /3 0 . 1 0 0 .0 5 +

23 27 /3 -4 /4 T .39 0 .92

27 66 4 / 4 -10 /4 - -

31 375 10/4 -1 7 /4 2 . 1 0 1 .12 +++

35 210 17/4 -2 4 /4 1 .15 0 .32 +++

39 278 24 /4 - 1 / 5 0 .67 0 .2 5 444

43 238 1 /5 - 8 / 5 - _ 44

47 232 8 /5 -2 2 /5 3 .13 0.71 44 4 4 4

51 286 2 5 .7 22 /5 - 2 9 /5 0 .0 9 0 . 0 6 +++ ++

55 268 29 .8 2 9 /5 - 5 /6 0 .84 0 . 4 8 44 ++

59 268 3 0 . 0 5 /6 -1 3 /6 1 .12 1 .07 4 +

— — — 13/6 - 2 0 / 6 1 .32 0 .63 4 +++»

66 50 4 .0 2 0 / 6 - 2 6 / 6 0 .1 5 0 . 1 2 4 +

70 340 2 4 .3 26 /6 - 3 /7 - - 44 +

74 450 4 4 .5 3 / 7 - 1 0 /7 0 .05 0 .0 5 ++ +

- _ 10 / 7 -1 6 /7 - - 44 +

8 l 318 1 1.1 3 5 .3 16/7 -2 6 /7 0 . 4 8 0 .2 5 ++ +

85 390 11 .6 4 5 .2 26 /7 - 3 1 /7 - - 4 +

89 520 12.25 6 2 .9 31 /7 - 6 / 8 o i l ? 0 .3 7 44 +

93 575 8 .7 5 0 . 0 6 / 8 - 1 4 /8 0.41 0.31 + + 4

97 390 11 .9 4 6 .4 14/8 - 2 i / 8 0 . 0 2 0 . 0 2 4 4 4 44

101 380 11 .9 45 .2 2 1 / 8 - 2 8 / 8 0 .61 0.41 +++ 44

105 350 13.1 4 5 .8 2 8 / 8 - 4 / 9 0 .82 0 . 8 0 ? ?

109 381 1 6 . 8 64 .0 4 / 9 - 1 1 /9 0 .53 0 .34 44 44

Tray I c o n td .No. F'.V DW Date Rain Y/eekly

Max. A CW113 317 1 8 . 0 3 7 . 0 11/9 - 1 9 /9 1 .1 9 0 .4 9 7 7

117 3k 5 14 .9 51 .4 19/9 -2 6 /9 0.1:3 0 . 2 8 + + +

121 1146 4 .4 50.4 2 6 /9 - 2 / 1 0 1 .29 1 .05 + + ++

125 1 672 3 .0 50.2 2 / 1 0 - 9 / 1 0 1 .33 1 .05 ++ ++

125 576 7 .6 4 3 .8 9/ 10-1 6 / 1 0 0 .6 7 0 .5 9 ++ + +

133 1 120 4 .4 4 9 .3 1 6 /1 0 -2 3 /1 0 0 .9 5 0 . 2 7 ++ + +

137 810 5.1 41 .5 2 3 /1 0 -3 0 /1 0 1 .47 0 .62 + + ++ +

141 810 4 .2 3 3 .7 3 0 / 1 0 - 6/11 1 .07 0 .6 9 ++ ++

145 325 3 .6 11 .7 6/11 -13/11 1 .15 0 . 4 0 + +++

149 - No dal a 13/11 - 20/1 1 1 . 6 8 0 .8 3 + ++ +

153 603 4 . 8 2 9 . 2 20/1 1 -27/1 1 0 .36 O. 2 3 + + +

157 885 3 . 5 31 .1 27/11 -4 /1 2 1 . 2 8 0 .5 4 + ++

1 61 648 5 .5 35 .4 4 /1 2 -1 1 /1 2 1 .07 0 . 6 2 ? ?

165 495 9.4 4 6 .5 1 1 / I 2-1 8 / 1 2 0 .42 O. 1 6 + ?

1 69 - No d a t a 1 8 / 12 - 2 5 / y ? ' 1 .43 1 .09 + + + + +

173 272 10.1 2 7 .4 2 5 / 1 2 - 1/1 /73 0 . 2 5 0 . 1 4 + ++

177 248 7 .6 1 8 .9 1/1 - 8/1 0 .53 0 .21 + +

1 81 331 9 .6 31 .9 8/1 -15/1 1 .22 0 .62 + +

1 85 790 5 .4 4 2 .9 15/1 - 22/1 1 .49 1 .09 + +

1 89 1250 3 .2 3 9 .6 22/1 - 29/1 0 . 1 4 0 .14 + +

193 1024 3 .4 35.1 29/1 - 5 /2 - - + +

1 97 1000 3 .4 34.4 5 / 2 - 1 2 / 2 0 . 0 8 0 . 0 8 + + +

201 502 6 . 2 31 .0 1 2 / 2 -1 9 /2 0 . 0 6 0 . 0 4 + +

205 258 11 .9 3 0 .8 1 9 / 2 -26 /2 1 .56 0.51 + +

209 - No d a t a 26 /2 - 5 / 3 0 . 0 8 o .o 3 + +

213 953 4 . 6 4 3 .9 5 /3 -1 2 /3 0 . 6 2 0 . 4 0 + +

217 1300 4 .4 57.3 12/3 -1 9 /3 0 .3 6 0 . 1 4 + +

Tray I co n td ,No. 17/ % m DW Date Rain weekly . A CW221 542 4 . 2 2 ^ . 9 19/3 -2 6 /3 1 .28 0.&0 4 4

225 210 8 .6 1 8 .0 2 6 /3 - 2 /4 1 .99 1 .42 ++ 44

229 691 5 .7 3 9 .5 2 / 4 - 9 /4 1 .97 1 .32 44 444

233 613 8 .8 53 .9 9 / 4 - 1 6 / 4 0.61 0 .3 8 + + 444

237 950 5 .7 54.2 16/4 -2 3 /4 0.41 0.1 7 + 44

241 910 3 .8 3 4 .9 23 /4 -3 0 /4 1 . 24 0.71 ? ?

245 - No dat a 3 0 /4 - 7 / 5 1 .38 0 .93 + 444

249 545 9 .9 53.8 7 /5 - 1 4 /5 1 .22 0 .79 + 44

253 540 6 .0 3 2 .6 1 4 /5 - 2 1 /5 1 .28 0 . 8 0 4 444

257 1 80 7.3 13.2 21 /5 - 2 8 /5 3 .7 5 1 .06 4 444

261 730 4 .4 2 9 .5 28 /5 - 4 /6 1 .1 4 0 .7 6 + 444

265 1340 3 . 6 4 7 .9 4 /6 - 1 1 / 6 0 .37 0.1 5 + 444

269 1055 3 . 6 3 7 .7 11/6 - 1 8 / 6 0 .0 9 0 .0 9 4 44

273 1000 2 .0 19*6 1 3/6 - 2 5 / 6 0 .34 O.24 4 44

277 407 5 .8 2 3 .5 25/6 - 2 /7 0 .5 5 0 .33 + + ?

281 330 6 .4 21 .0 2 /7 - 9 /7 - - + + ?

285 530 3 . 9 2 5 .O 9/7 - 1 6 / 7 0.04 0 .04 44 4

289 625 - - 1 6 / 7 - 2 3 /7 0 .06 0 . 0 3 44 4

293 600 5 .0 2 9 .9 23 /7 -3 0 /7 0 .34 0.1 5 44 4

297 490 9.1 4 4 .7 3 0 /7 - 6 /8 1 .61 0 .7 9 44 4

301 480 10.3 4 9 .4 6 /8 -1 3 /8 0 .43 0 .40 44 4

305 330 - - 13/8 -2 0 /8 O. 2 5 0 .20 4 4

309 380 - - 2 0 / 8 - 2 7 / 8 0 .43 0.21 4 4

313 770 - - 2 7 / 8 - 1 0 /9 2.31 1 .1 8 44 44

317 1337 5 .7 76.2 10/9 -1 7 /9 0 .60 ■\ 4 44

321 838 9 .2 77.1 17/9 -2 4 /9 1 .79 44 44

325 725 - - 2 4 /9 -1 /1 0 0.71 0 .6 5 44 44

329 360 - - 1 /1C-8/10 2.11 1 .06 44 4 44

3 3 3 - -“

8 / 10-1 5/10 1 .06 0 .68 44 444

i r ay No.

A. com P\V

✓ u *

%D’.V DW Date Rain WeeklyMax. A CW

337 - - - 15/10 -22 /10 2 .70 1 .22 +++ +++

341 - - - 22/10 - 2 9 / 1 0 0 .9 7 0 .3 5 ++ 44

345 - - - 29/10 -5/11 1 .27 0 .5 7 44 44

349 - No l a t a 5/11 -12/11 1 .46 0 .4 8 + 44

353 /* /> /\ DUU - - I £./ 1 1 —1 y / I | 0 .9 5 0 .5 7 4 44

357 1000 - - 19/11 -26/11 1 .37 0 .8 8 + 44

36l 500 - - 26/11 -3 /12 1 .77 0 .7 9 + ++

365 - - - 3/12 -10 /12 2 .3 0 1 .12 + + 444

369 - - - 10/12 -17 /12 0.31 0 . 3 0 ++ 44

373 - - - 17/12 -2 if / l2 0 .82 0 .5 5 ++ 44

377 - - - 24/12 -31 /12 0 .0 7 0 .0 7 ++ 4

381 - - - 31/12 - 7 /1 /7 4 - - + 4

385 - - - 7/1 -14/1 - - ■++ 4-4

389 - - - 14/1 -21/1 - - + + 4

393 - - - 21 / I -28/1 - - 44 44

T r a y I I .

No. FW %DW DW Date Rain Week lyMax.

4 - - - 19/2 - 2 5 /2 0 .13 0 . 0 8

8 - - - 25/2 -6 /3 - -

I 2 - - - 6 /3 “ 13/3 0 . 0 8 0.0416 - - - 13/3 -20 /3 - -

20 ?1 - - 2 0 / 3 - 2 7 / 3 0 .1 0 0

24 - - - 2 7 / 3 - 4 /4 1 .89 0 .92

28 22 - - 4 /4 -1 0 /4 - -

32 12 - - 10/4 -17 /4 2 .10 1 .12

36 89 - - 17/4 -24 /4 1 . 15 0 .32

40 49 - - 24 /4 - 1 /5 0 .67 0 .2 5

44 72 - - 1 /5 - 8 /5 - -

48 114 - - 8 /5 - 2 2 / 5 3 .13 0.7152 46 - 5 .8 2 2 /5 -2 9 /5 0 .09 0 .0 6

56 70 - 7 .9 29 /5 -5 /6 0 .84 0 .4 8

60 115 — 14 .7 5 /6 -1 3 /6 1 .12 1 .07

63 110 — 11 .6 13/6 -2 0 /6 1 .32 0 .63

67 74 — 7 .8 20 /6 -2 6 /6 0 .1 5 0 .12

71 80 — 9 .0 26 /6 - 3 /7 - -

75 150 - 11 .1 3 / 7 -1 0 /7 0 .05 0 .0 578 89 10,0 8 .9 10/7 - 1 6 / 7 - -

82 90 8 .3 7 .4 1 6 / 7 - 2 6 /7 0 .48 0 .2 5

86 39 6.1 2 .4 26 /7 -3 1 /7 - -

90 45 11 .2 5 .0 3 1 /7 - 6 /8 0 .3 7 0 .3 7

94 31 11 .7 3 .6 6 /8 -1 4 /8 0.41 0.31

98 16 11 .1 1 .9 r— \ 00 1 PO \ 00 0 .02 0 .02

1 02 12 1 6 .9 2 .0 2 1 / 8 - 2 8 / 8 0 .61 I 0.41

106 14 U . 1 1 .9 28 /8 - 4 /9 0 .82 0 . 8 0

T r a y l l c o n t d .

No. F.V (/oD\" DW Date Rain WeeklyMn*.

110 "3i 14.4 4 . 5 4 /9 - 1 1 /9 0 .53 0 .34

114 54 17.4 4 .6 11 /9 - 1 9 /9 1 .1 9 0 .4 9

11.8 31 1 6 .3 4 .2 19/9 -2 6 /9 0 .43 0 . 2 8

122 51 14.0 7.1 2 6 /9 - 2 / 1 0 1 .29 1 .05

1 26 90 1. 0 , 5 9 .5 2 / 1 0 - 9 / 1 0 1 .33 1 . 05

130 118 6 .7 7 .9 9 / 10-1 6/1 0 0 .6 7 0 .5 9

134 94 7.1 6 .7 1 6 / 1 0 - 2 3 / 1 0 0 .9 5 0 .27

138 200 3.1 6 . 1 2 3 /1 0 - 3 0 / 1 0 1 .47 0 . 6 2

142 254 3 .2 8 . 2 3 0 / 1 0 - 6/11 1 .07 0 .6 9

146

oo<\i 3 . 0 8 .4 6/11 -13/11 1 .1 5 0 .4 0

1 50 249 3 .6 9.1 13 / 1 1 - 20/11 1 .68 0 .83

154 60 10.7 6 .4 20/1 1 - 27/1 1 0 .36 0 .23

158 167 5.2 8 . 6 27/1 1 -4 /12 1 . 2 8 0 .54

162 305 2.1 6 .4 4 / 1 2 - 1 1 / 1 2 1 .07 0 . 6 2

1 66 147 5 .7 8 .3 11 / 1 2 - 1 8 /1 2 0.42. 0 . 1 6

1 70 65 1 1 .0 7.2 1 8 / 1 2 - 2 5 / 1 2 1 .43 1 .09

174 97 6 .7 6 .5 2 5 / 1 2 - 1/1 / 7 3 O.25 0 .14

1 78 53 8 . 6 4 .6 1/1 - 8/1 0 .53 0 .21

182 63 10.3 6 .; 5; 8/1 -15/1 1 .22 0.62

186 135 4 .8 6 .4 15/1 - 22/1 1 .49 1 . 0 9

1 90 157 8 .5 13.3 22/1 - 29/1 0 .14 0.14

194 450 , 2 . 8 1 2 . 8 29 /1 - 5 /2 - -

1 93 260 4 .3 1 1 .2 5 / 2 - 1 2 / 2 0 . 0 8 0 . 0 S

202 154 5 .3 8 . 2 1 2 / 2 -19 /2 0 . 0 6 0 .04

206 302 2 . 8 8 .4 19/2. - 2 6 / 2 1 .56 0 .51

210 1 87 4 . 6 8 . 6 2 6 / 2 - 5 / 3 0 . 0 8 0 . 0 8

214 - No d a t a 5/3 -12 /3 0 . 6 2 0 .4 0

Tray II contd.No. FW %DW 1)7/ Date Rain Week]y

Mav.21 8 2 b k 5 .2 8 .5 12/3 -1 9 /3 0 .36 0 . 1^

222 203 3 . 0 6.1 19/3 -2 6 /3 1 . 2 8 0 . 6 0

226 1 85 4 .2 7 .8 26 /3 -2 /4 1 .99 1 .42

230 273 4 .3 11 .8 2 /4 - 9 /4 1 .97 1 .32

2.34 245 3 . 8 9 .2 9/4 -1 6 /4 0 . 6 i 0 .3 8

238 233 6 .7 1 5 .6 16/4 -2 3 /4 0.41 0.1 7

242 163 5 .2 8 .5 23 /4 -3 0 /4 1 .24 0.71

246 1 50 4 . 9 7 .3 3 0 /4 - 7 / 5 1 . 38 0 .93

250 135 4 . 7 6 .3 7 /5 - 1 4 /5 1 .22 0 .7 9

254 1 02 0 . 8 0 . 8 14/5 -2 1 /5 1 . 28 0 . 8 0

258 390 2.1 8 . 2 2 1 / 5 - 2 8 / 5 3 .7 5 1 .06

262 270 3 .0 8 . 0 2 8 /5 - 4 /6 1 .14 0 .76

266 250 3 .4 8 .5 4 / 6 - 1 1 / 6 0 .37 0 .1 5

270 225 2 .9 6 . 6 1 1 / 6 - 1 8 / 6 0 .09 0 .0 9

274 210 4.1 8 .7 1 8 / 6 - 2 5 / 6 0 .34 0 .24

278 300 1 .9 5 .6 25 /6 - 2 / 7 0 .5 5 0 .33

282 ~Z.> > 6 .3 5 .0 2 /7 - 9 / 7 - -

286 200 1 .8 3 .0 9 /7 - 1 6 /7 0 .04 0 .04

290 80 5 .7 4 . 6 16/7 - 2 3 /7 0 . 0 6 0 .03

294 120 4 . 8 5 .8 2 3 /7 - 3 0 /7 0 .34 0.1 5

298 - No ci a t a 3 0 /7 - 6 / 8 1 .61 0 .7 9302 - No d a t a 6 / 8 - 1 3 /8 0 .43 0 .4 0

306 58 - - 13 /8 -2 0 /8 0 .25 0 . 2 0

310 40 - - 2 0 /8 -2 7 /8 0 .43 0 .21

314 137 - - 2 7 /8 -1 0 /9 2.31 1 .1 8 .

31 8 65 0.71 0 . 5 10/9 -1 7 /9 0 . 6 0 0 .3 7322 31 - - 17/9 -2.4/9 1 .79 1 .07

T r a y I I c o n t d .

No. FW %DW DW Date Rain WeeklyMs y .

326 - - - 24 /9 - 1 / 1 0 C .71 0 .6 5

330 66 - - 1/ 1 0 - 8 / 1 0 2.11 1 .0 6

334 - - - 8 / 10-1 5 / 1 0 1 .0 6 0 . 6 8

338 - - - 1 5/1 0 - 22/1 0 2 .70 1 . 22.

342 - - - 09 /1 A 90 /1 A4_ i— / i v_/ “*<— y / | v r\ r\n v • 7 ( 0 .3 5

346 - - - 29 /10-5 /11 1 .46 0 .5 7

350 100 - - 5 /11-12/1 1 0 .95 0 .4 8

3 3 k - - - 1 2/1 1 -> 9/1 1 1 .37 0 .5 7

3 3 8 - - - 1 9/11 - 2 6 / 1 1 1 .77 0 . 8 8

362 - - - 26/1 1 - 3 / 1 2 1 .77 0 .7 9

366 - - - 3 /1 2 -1 0 /1 2 2 .3 0 1 .12

370 ■ - - - 1 0 / 12-1 7 /1 2 0.31 0 .3 0

3 7 k - - - 1 7 /1 2 -2 4 /1 2 0 . 8 2 0 .5 5

3 7 8 - - - 24 /1 2 -3 1 /1 2 0 .0 7 0 .0 7

382 - - - 3 1 /1 2 -7 /1 /74 - -

386 - - - 7/1 -14 A - -

390 - - - 14/1 - 21/1 - -

3 9 k - - 21/1 - 28/1 - -

No. FW DW Date Rain Y/eekiy Max.

5 - - - 19/2 -2 5 /2 0 .13 0 . 0 8

9 - - - 25/2 - 6 /3 - -

13 - - *- 6/3 -1 3 /3 0 . 0 8 0 .04

1? - - - c* 1 ro 0 - _

«■■» <• c.1 1 1 7. i. I -/T on /7 on /? 0 . i 0 0 . 0 5

25 - - - 27/3 - 4 /4 1 .89 0 .92

29 143 - - 4 /4 -1 0 /4 - -

■SV 99 - - 10/4 -1 7 /4 2 .10 1 .12

37 131 - - 17/4 -2 4 /4 1 .15 0 . 3 2

41 98 - - 2 4 /4 - 1 / 5 0 .67 0 . 2 5

45 154 ' - - 1 /5 - 8 / 5 - -

49 98 - - 8 / 5 - 2 2 / 5 3 .1 3 0.71

53 201 - 2 9 . f- 2 2 /5 - 2 9 /5 0 .0 9 0 .0 6 v

57 240' - 2 9 .5 2 9 /5 - 5 /6 0 .84 0 .4 8 '61 220 - 3 0 .8 5 /6 -1 3 /6 1 . 12 1 .07

64 310 - 3 3 .5 13/6 - 2 0 / 6 1 .32 0 .6363 174 - 23 .8 2 0 / 6 - 2 6 / 6 0 .1 5 0 . 1 2

72 176 - 2 5 . S 2 6 /6 - 3 /7 - -76 184 - 14 .7 3 / 7 - 1 0 /7 0 .0 5 0 .0 5

79 1 80 13.6 2 4 .5 1 0 / 7 - 1 6 / 7 - -

83 135 1 5 . 0 2 0 . 2 1 6 / 7 - 2 6 /7 0 .4 8 0 . 2 5

87 94 21 .1 1 9 . 8 2 6 /7 - 3 1 /7 - -91 124 17.1 21 .2 3 1 /7 - 6 / 8 0 .37 0 .3 7

95 144 1 5 .9 2 2 . 8 6 / 8 - 1 4 /8 0.41 0.31

99 88 16.1 14.2 14/8 - 2 i / 8 . 0 . 0 2 0 . 0 2

103 140 14 .4 2 0 . 2 2 1 / 8 - 2 3 /8 ' 0.61 0.41

107 201 1:5.1 30.4 2 3 /8 - 4 / 9 0 .82 0 .3 0

Tray I I I contd.

No. FW %un DV/ Dat e RainWeekly

Max.111 2 2 5 1 3 . 0 4 0 . 5 4 / 9 - 1 1 / 9 0 . 5 3 0 . 3 4

115 236 1 7 . 4 41 .1 1 1 / 9 - 1 9 / 9 1 .1 9 0 . 4 9

119 24 9 1 6 . 3 4 0 . 6 1 9 / 9 - 2 6 / 9 0 . 4 3 0 . 2 8

123 264 1 6 . 9 4 4 . 6 2 6 / 9 - 2 / 1 0 1 . 2 9 1 . 0 5

1 on I < - { o r et- y y •t n ni ; • i 1. /. 7 O /1 O n /i A w i ^ ' y/ i v 1 77• • -x 1 nc;

1 • v-/ y

131 227 1 5 . 8 3 5 . 9 9/ 1 0 - 1 6 / 1 0 0 . 6 7 0 . 5 9

135 25 0 1 5 . 7 4 5 . 5 1 6 / 1 0 - 2 3 / 1 0 0 . 9 5 0 . 2 7

139 210 1 5 .0 31 . 5 2 3 / 1 0 - 3 0 / 1 0 1 . 4 7 0 . 6 2

143 322 11 . 8 3 7 . 9 3 0 / 1 0 - 6 / 1 1 1 . 0 7 0 . 6 9

147 263 1 2 . 0 31 . 5 6 / 11 - 1 3 / 1 1 1 . 1 5 0 .4 0

151 2 6 9 1 0 . 4 2 8 . 0 1 3 / 1 1 - 2 0 / 1 1 1 . 6 8 0 . 8 3

1 55 314 1 0 . 4 3 2 . 5 20/1 1 - 27 /11 0 . 3 6 0 . 2 3

159 4 5 5 8 . 8 4 0 . 1 2 7 / 1 1 - 4 / 1 2 1 . 2 8 0 . 5 4

163 5 6 5 7 . 7 2 8 . 2 4 / 12-1-1 /1 2 1 . 0 7 0 . 6 2

167 3 00 5 . 2 1 5 . 5 1 1 / 1 2 - 1 8 / 1 2 0 . 4 2 0 . 1 6

•171 301 8 . 5 2 5 . 7 1 8 / 1 2 - 2 5 / 1 2 1 . 43 1 . 0 9

175 163 1 7 . 2 2 7 . 9 2 5 / 1 2 * 1 A / 7 3 0 . 2 5 0.1-4

179 277 8 . 6 3 4 . 1 1/ 1 - 8 / 1 0 . 5 3 0.2.1

1 S3 2 8 7 1 0 . 3 2 8 . 1 8/1 - 1 5 / 1 1 . 2 2 0 . 6 2

1 87 47 0 5 . 2 21 . ? 15 / 1 - 2 2 / 1 1 . 4 9 1 . 0 9

191 3 6 0 8 . 0 2 8 . 9 2 2 / 1 - 2 9 / 1 0 . 1 4 0 . 1 4

1 95 3 6 7 7 .8 2 3 . 7 2 9 / 1 - 5 / 2 - -

199 3 2 8 1 0 . 6 3 4 . 6 5/2- - 1 2 / 1 0 . 0 3 o.oS

203 3 2 7 1 0 . 4 3 4 . 0 1 2 / 2 - 1 9 / 2 0 . 0 6 0 . 0 4

207 48 3 8 . 9 4 2 . 8 1 9 / 2 - 2 6 / 2 1 .56 0 . 5 1

21 1 5 5 5 1 0 . 0 3 5 . 5 2 6 / 2 - 5 / 3 0 . 0 8 0 . 0 8

2 1 5 35 6 8 . 4 2 9 . 8 5 / 3 - 1 2 / 3 0 . 6 2 0 . 4 0

Tray I I I con i d .N o. FW %D'U DW Date Rain Weekly

Max.21 9 458 7.4 7 7 , R _• • o 12/3 -19/3 O.56 0.14

223 265 9.1 24.2 19/3 -26/3 1 .28 0 .60

227 305 12.2 37.5 26/3 -2/4 1 .99 1 .42

231 392 9.7 37.8 2/4 -9/4 1 .97 1 .32

235 >8o 8 .8 33.5 9/4 -1 6/4 0.61 0.38

239 400 8.7 34.8 16/4 -23/4 0.41 0.17

243 335 10.6 35.5 23/4 -30/4 1 .24 0.71

247 240 14.9 35.7 30/4 -7/5 1 .32 0.93

251 280 1 5.3 42.9 7/5 -14/5 1 .22 0.79

255 353 7.4 26.1 14/5 -21/5 1 .28 0 .80

259 470 8.5 39.9 21/5 -28/5 3.75 1 .06

263 400 7.7 30.0 28/5 -4/6 1 .14 0.76

267 266 4.7 12.4 4 /6 -11 /6 0.37 0.15

271 510 4.4 22.5 11/ 6-18/6 0.09 0.09

275 470 6.9 23.8 18/6 -25/6 0.34 O.24

279 340 1 .9 30.9 25/6 -2/7 0.55 0.33

283 430 6.3 27.0 2/7 -9/7 - -

287 350 5.7 20 .0 9/7 -16/7 0.04 0.04

291 390 6.7 26.0 16/7 -23/7 0 .06 0.03

295 260 10.9 2 8 .0 23/7 -30/7 0.34 0.1 5

299 330 10.9 3 6 .0 30/7 -6/8 1 .61 0.79

303 210 12 .8 2 6 .8 6 /8 -13/8 0.43 0.40

307 21 7 - - 13/8 -20/8 0.25 0 .20

311 225 - - 20/3 -27/8 0.43 0.21

31 5 1 00 - - 27/8 -10/9 2.31 1 .13

319 - - - 10/9 -17/9 0 .60 0.37

323 320 - - 17/9 -24/9 1 .79 1 .'■'7

Tray TIT contd.N o . F/V %WI DW Date Rain Weekly

^ r

215 - - 24 /9 -1 /1 0 0.71 0 .6 5

531

K\COAJ - - 1 /1 0 -8 /1 0 2.1 1 1 .06

335 - - - 8/10-1 5/10 1 .06 0 .68

339 - - - 15 /1 0 -2 2 /1 0 2.70 1 .22

343 - - - 22/1 0 - 2 9 / 1 0 0 .97 0 .35

347 - - - 29/10-5 /11 1 .27 1 .57

351 150 - - 5/11 -1 2/11 1 .46 0 . 4 8

355 300-, - - 1 2/1 1 -1 9/11 0 .9 5 0 .5 7

359 200 - - 1 9/1 1 -26/11 1 .37 0 .8 8

363 1 70 - - 2 6 /1 1 -3 /1 2 1 .77 0 .7 9

367 1 60 - - 3 /1 2 -1 0 /1 2 2 .30 1 .12

371 - - - 10/12-1 7/12 0.31

O

r< \ •0

375 - - - 1 7 /12 -24 /12 0.82 0 . 5 5

379 - - - 2 4 /1 2 -3 1 /1 2 0 .07 0 .0 7

383 - - - 3 1 / 1 2 -7 /1 /7 4 - -

387 - - - 7/1 -14/1 - -

391 - - - 14/1 -21/1 - -

395 - - - 21/1 -28/1 - *1

4-

Tray IVNo. FV %m DW Date Rain Weekly

Mo v1 id/. •

6 - - - 19/2- 25/2 0.13 0.08

10 - - - 25/2 -6/3 - -

14 - - - 6/3 -13/3 0.08 0 .04

18 - - - 13/3 -20/3 - -

22 76 - - 20/3 -27/3 0.10 0 .0 5

26 - - - 27/3 -4/4 1 .89 0 .9 2

30 43 - - 4/4 -10/4 - -

34 62 - - 10/4 -17/4 2 .10 1 .12

38 105 - - 17/4 -24/4 1 .15 0.32

42 81 - - 24/4 -1/5 0.67 0.25

46 87 - - 1/5 -8/5 - -

50 HO - - 8/5 -22/5 3.13 0.71

3k 95 - 8.0 22/5 -29/5 0.09 0 .0 6

58 72 - 6.5 29/5 -5/6 0.84 0.48

62 9k - 7.9 5/6 -13/6 1 .12 1 .07

65 106 - 7.3 13/6 -20/6 1 .32 0.63

69 290 - 19.4 20/6 -26/6 0.15 0.12

73 105 - 8.8 26/6 -3/7 - -

77 162 - 8.4 3/7 -10/7 0.05 0.05

80 1 50 8 . 5 12.7 10/7 -16/7 - -

84 75 9.3 7.0 16/7 -26/7 0.48 0.25

88 98 9.3 9.1 26/7 -31/7 - -

92 128 8.9 11 .4 31/7 -6/8 0.37 0.37

96 120 7.6 9-2 6/8 -14/8 0.41 0.31

100 118 9.6 11 .3 14/8 -21/8 0.02 0.02

104 285 10.1 10.6 21/8 -28/8 0.61 0.41

108 1 7k 6 .3 10.9 28/8 - 4 / 9 0.82 0 .8 0

Tray IV contd.No. F.7 %D'N DW Date R ain Weekly

Fax.112 2 0 5 7.1 14.6 4/9 -11/9 0.53 0.34

1 1 6 171 8.3 14.2 11/9 -19/9 1 .1 9 0.49

120 272 4.9 13.3 19/9 -26/9 0.43 0.28

124 1 06 10.5 13.0 26/9 -2/10 1 .29 1 .05

128 480 3.5 16.8 2./10-9/10 1 .33 1 .05

1 ■*,?i *— 1 p p0 n-' 1 x cI • w n / 1 r\ i C. /* r\J / 1 ~ 1 Kj / I n C nW • O f 0.59

T36 483 2.9 1 4 .0 16/ 10-23/10 0.95 O.27

140 700 1 .9 13.1 23/10-30/10 1 .47 0 .62

144 960 1 .1 10.4 30/10-6/11 1 .07 0.69

148 148 1 .4 2.1 6/11 -13/11 1 .1 5 0.40

152 832 1 .2 10.1 1 3/1 1 -29/1 1 1 .68 0.83

156 807 1 .0 8.1 20/1 1 -27/1 1 0.36 0 .25

1 60 156 7.7 12.1 27/11-4/12 1 .28 0.54

1 64 476 1 .9 9.1 4/12-1 1 /1 2 1 .07 0.62

168 864 1 .9 1 6.7 11/12-1 8/12 0.42 0 .16

172 58 9.3 5.4 1 8/ 12-25/12 1 .43 1 .09

176 48 13.2 6.3 25/ 12-1/1 /73 0.25 0 .14

1 80 1 08 - - 1/1 -8/1 0.53 O.21

1 84 140 - - 8/1 -1 5/1 1 .22 0 .62

188 195 4 .4 8.5 15/1 -22/1 1 .49 1 .09

1 92 700 0.9 6.23 22/1 -29/1 0 .14 0.14

1 96 105 9.6 10.0 29/1 -5/2 - -

200 65 6.4 4.2 5/2 -12/2 0.08 0.08

204

ooK\ 1 .5 4.59 12/2 -19/2 0.06 0.04

208 212 1 .7 3.5 19/2- 26/2 1 .56 0.51

212 - - - 26/2 -5/3 0.08 0.08

216 275 1 .9 5.1 5/3 -12/3 0.62 0 .4 0

Tray IV contd.No. F.7 SfflDW DW Date Rain V'eekiy

' f a x .220 210 2 . 2 4 .6 12/3 -1 9 /3 0 . 5 6 0 .14

224 206 2.1 4 .4 19/3 -2 6 /3 1 . 2 8 0 . 6 0

228 175 3 . 7 6 .4 26/3 -2 /4 1 .99 1 .42

23 2 246 1 .9 4 .7 2 /4 - 9 /4 ' 1 .97 1 .32

236 100 3 .8 3 . 8 9 / 4 - 1 6 / 4 0 .61 0 .38

240 850 0 .4 3 . 6 16/4 -2 3 /4 0.41 0.1 7

244 250 - - 23 /4 -30 /4 1 .24 0 .71

248 215 4 .6 9 .9 30 /4 - 7 /5 1 .38 0 .93

252 250 1 .8 4 . 5 7 /5 - 1 4 /5 1 .22 0 .7 9

236 350 1 .5 5 .4 14 /5 - 2 1 /5 1 .28 0 .80-

260 110 4 . 7 5.1 2 1 / 5 - 2 8 /5 3 .7 5 1 .06

264 654 2.1 13 .5 2 8 /5 - 4 /6 1 .14 0 .7 6

268 - - - 4 /6 -1 1 /6 0 .3 7 0.1 5

272 - - - 1 1 / 6 -1 8 /6 0 .0 9 0 .0 9

276 70 5 .7 4 .0 1 8 / 6 - 2 5 / 6 0 .3 4 0 .2 4

280 - - - 25 /6 - 2 / 7 0 .5 5 0 .33

284 330 7 .6 2 5 . 0 2 /7 - 9 /7 - -

288 270 7 .7 2 0 .8 9 /7 -1 6 /7 0 .0 4 0 .04

292 280 9 .7 2 7 . 2 16/7 -2 3 /7 0 .06 0 .03

296 240 10.1 24 .2 23 /7 -3 0 /7 0 .3 4 • 0.1 5

300 365 9.4 3 4 .5 30 /7 - 6 /8 1 .61 0 .7 9

304 490 4 .9 2 4 . 2 6 /8 -1 3 /8 0 .43 0 .40

000f-'* 307 - - 13 /8 -2 0 /8 0 .2 5 0 .20

312 655 - - 20/S -2 7 /3 0 .43 0.21

m 475 - - 27 /8 -1 0 /9 2.31 1 .18

320 - - - 10/9 -1 7 /9 0 .6 0 0 .3 7

324 490 8 .2 7 0 O 17/9 -2 4 /9 1 .79 1 .07

I

Tray IV contd.No. F.V %Wl DV/ Date Rain V/eekly

> 'a v-.28 310 - - 24 /9 - 1 / 1 0 0 • / I 0 .65

332 457 - - 1 / 10 - 8 / 1 0 2 .11 1 .0 6

336 260 - - 8 / 10-1 5 / 1 0 1 .06 0 . 6 8

340 - - - 1 5/1 0 - 2 2 / 1 0 2 .7 0 1 .22

344 - - - 22/1 0-29/1 0 0 .9 7 0 .3 5

348 - - - 29 /10-5 /11 1 . 2 7 1 .57

352 350 - — 5/1 1 -1 2/1 1 1 .46 0 .48

356 350 - - 12/11-19/11 0 .9 5 0 .5 7

360 400 - - 1 9 / 1 1 - 26/11 1 .37 0 . 8 8

364 300 - - 26/11 - 3 / 1 2 1 .77 0 .7 9

368 - - - 3 / 1 2 - 1 0 / 1 2 2 .3 0 1 .12

37 2 - - - 10 / 12-1 7 /1 2 0.31 0 . 3 0

376 - - - 17/1 2 - 21^/12 C ,82 0 .5 5

380 - - - 2 4 /1 2 -3 1 /1 2 0 .0 7 0 .0 7

3 CM - - - 3 1 /1 2 -7 /1 /74 - -

388 - - - 7/1 -14/1 - -

392 - - - 14/1 - 21/1 - -

396 - - - 21/1 - 28/1 - -

A p p en d ix b -

D e t a i l s o f a l l p o l l e n c o u n t s made from s a m p le s

p r e p a r e d from g uano c o l l e c t e d e a c h w e e k i n B a tu C a v e s , S e l a n g o

A b b r e v i a t i o n s u s e d : -

N o . — S am ple n u m b e r » F o r o t h e r d a t a on e a c h s a m p le s e e

A p p e n d ix 3«

D .g . D uabanga g r a n d i f l o r a .

S . a . = S o n n e r a t i a a l b a .

S . c . = S o n n e r a t l a c a s e o l a r l s

P . = P a r k i a s p p .

O . i . = Orox.ylum i n d l c u m .

D.1 D u r io t y p e 1 ( i . e . w i t h

C . p . = C e ib a p e n t a n d r a .

B . v . = Bombax v a l e t o n i i .

E . = E u g e n ia s p p .

C . n . = Cocos n u c i f e r a . '

A. ~ A re n g a s p p .

A r t . = A r t o c a r p u s s p p .

T3 a ?

Tb = ?

T8 = ?

= M a n g l f e r a s p p .

T10 = ?

R. = R e m a in d e r ( a l l r a r e p o l l e n t y p e s , w h e t h e r i d e n t i f i e d

, o r n o t a r e i n c l u d e d i n t h i s colum n i n t h i s A p p e n d i x ) .

TRAY 1.

No. D.g. S.a. S.c. p. O.i. D.1 C.p. E. v. E, C.n. A. Art. T3 T4 T8 M. T10 R • o

3 - 16.5 0.1 0.4 - - 0.1 - - - - 82.7 0.2 - _ _ _ -

7 85.1 0.2 - 0.6 - 1 .4 0.8 - - - - 8.2 0.2 - 0.2 - 3.0 ■- -

11 82.4 - 5.6 1 .2 - 2.6 0.2 - - •• - 4.0 0.2 - 1 .2 - 2.6 - -

15 67.2 0.4 - 2.2 0.4 6.0 0.6 - 0.6 0.6 - 1 7 0 - - 0.2 - 4.6 0.2 -

19 60.6 0.4 - 2.4 0.2 9.2 0.4 - 0.2 - - 26.4 0.2 - - - - - -

23 85.8 0.6 0.2 0.6 - 2.2 0.2 - 0.2 - 9.6 0.4 - - 0*2 - - -

27 77.0 0.8 C .6 1 .0 0.2 0.8 0.2 - 15.6 0.2 3.2 - - - 0.2 0.2 - -

31 90.4 - - 0.6 0.2 0.2 - - - - - 8.2 0.2 - - 0.2 - - -

35 72.8 1 .0 - 1.8 0.2 0.6 0.2 - - 0.4 - 20.8 0.2 - - - 1 .8 0.2 -

39 59.2 1 .2 0.2 2.4 - 0 .8 0.6 - 0.4 0.2 - 18.6 0.2 - 15.8 0.4 - - -

43 17.2 0 ;2 - .3.8 0.4 0.8 - 0.2 0.4 - - 5.6 - - 71 .4 - - - -

47 43 *6 0.2 0.4 16.8 0.4 5.8 - 0.2 2.2 0.4 - 13.8 0.4 - 15.0 0.4 - 0.4 -

51 41 .0 0.8 0.2 30.0 0.8 1 .6 - 0.2 0.6 0.6 - 8.0 0.2 - 15.8 0.2 0.2 - -

55 25.0 4.6 0.4 47.2 1 .0 2.0 - - - 0.4 - 5.2 0.2 - 11.8 2.2 - - -

59 10.4 1 .6 - 83.2 2.2 - - - - - - 2.2 - - 0.4 - - - -

66 16.3 3.4 4.2 56.4 6.2 0.2 - - 1 .0 0.8 - 9.4 1 .2 - 0.8 0.2 - - -

70 2.0 2.0 0.2 89.2 2.4 - - - 0.2 - - 3.6 - - - 0.4 - - -

7b 7.4 1 .2 27.2 54.6 4.6 - - 0.2 - 0.2 - 3.2 0.2 - 0.8 0.4 - - -

81 46.8 - 3.6 1 6.4 2.0 - - 0.6 3.2 1- 27.0 0.4 - 0.2 - - - -

85 41 .2 1 .0 2.6 9.4 1 .6 - 0.2 0 0.2 1 .4 - 41 .2 0.8 0.4 - - - -

89 47.6 0.6 0.8 7.4 0.6 0.2 - - 13.4 2.0 - 26.4 - - 1 .0 - - - -

93 16.6 0.8 0.6 2.8 0.4 0.2 - - 58.4 2.2 - 17.0 0.6 - 0.6 - - - -

97 6.4 6.4 0.2 0.2 - - - - 83.0 0.2 - 3.4 - - - 0.2 - - -

101 24.2 38.2 0.2 0.2 - - - 31 .6 - - 5.2 0.4 - - - - - -

105 31 .4 55.8 0.6 0.8 - 1 .4 - - 2.2 - : - 6.4 0.4 - 0.6 0.4 - - -

109 13.0 63.2 0.2 1.8 - 6.6 0.2 - - - 12.4 2.4 - 0.2 - - - -

113 63.2 6.0 0.6 2.6 0.6 1 S.6 - - 1.6 0.4 i - 4.8 1 .4 - - 0.2 - - -

117 70.2 2.6 0.4 4*6 0.4 10.8 - - 1.0 0.2 0.4 8.0 0.4 - 1 .0 - - - -121 71.6 2.0 1 .4 10.0 0.4 8.4 - - - 0.6 0.2 3.6 1 .0 - 0.8 - - - - '

125 66.0 0.6 1 .0 11 .4 0.2 15.2 - - 0.6 - - 3.0 0.4 - 1 .6 - - - -

129

COir\ 0.2 0.2 19.2 0.2 7.6 1 .0 - 9.2 0.6 - 5.4 0.8 - 1 .0 - - - -

133 32.2 0.6 0.4 10.4 - 2.6 - - 26.0 - - 27.4 0.4 - - - - - -

137 50.4 0.8 1 .4 25.0 0.2 1 .2 - - 14.4 - - 5.0 0.6 - - - - - -

141 61 .2 5.2 1 .0 24.6 0.2 2.8 0.4 - 0.4 0.4 - 3*6 - - 0.2 - - - -

145 49.0 26.8 2.;.' 12.6 - 0.2 0.2 0.4 0.4 - - 7.2 0.6 - - - - 0.2 -

X JA/l X I

No. D .g . S #cl • S .c . P . O.i. D .1 C.p. r . v. S. C.n. A. A rt . T3 T4 T8 M. T10 P . 7

153 3 7 . 4 3 4 . 9 3 . 7 13.1 0 . 2 0 . 2 - 1 . 2 - 0 . 2 _ 8 . 2 1 . 0 - - - - - -■

157 4 5 . 4 4 . 2 1 . 8 11 . 6 0 . 8 - 1 .0 3 . 0 0 . 4 - 2 5 . 2 2 . 4 - 4 . 2 0 . 2 - 0 . 2 0 . 4

1 61 7 4 . 6 1 . 6 0 . 8 6 . 2 0 . 4 - 1 . 2 2 . 4 0 . 4 - - 8 . 8 3 . 4 - - 0 . 2 - - -

165 6 5 . 8 1 .6 - 6 . 4 0 . 8 - 0 . 8 1 .4 1 .2 0 . 4 - 1 6 . 4 4 . 4 - 0 . 2 - 0 . 2 0 . 4 -

173 5 0 . 6 1 .2 0 . 8 5 . 8 0 . 2 0 . 2 4 . 2 2 . 6 0 . 6 0 . 2 - 3 0 . 2 3 . 4 - - - - - -

177 4 5 . 8 0 . 8 - 3 . 0 0 . 2 - 7 . 7 1 .7 - 0 . 7 - 3 8 . 8 1 .2 - - - - 0 . 2 -

181 2 3 . 0 0 . 4 0 . 4 0 . 4 - 0 . 2 4 . 8 1 .8 1 0 .2 0 . 2 - 5 0 . 6 1 .6 -■ 0 . 2 - - - 6 . 4

189 8 2 . 6 - 1 .0 0 . 6 - - 3 - 4 2 . 0 - 0 . 2 - 8 . 6 1 .2 - 0 . 2 0 . 2 - - -

193 7 0 . 0 2 . 7 0 . 2 0 . 2 - - 2 . 2 1 .1 7 .3 - - 1 5 .2 0 . 6 - - - - - -

197 7 3 .2 7 . 2 1 .0 0 . 2 - - 1 . 8 - - 0 . 2 - 1 3 .2 0 . 4 - 2 . 8 - - - -201 6 8 . 0 11 .0 0 . 4 0 . 4 - - 0 . 8 0 . 2 3 . 2 - - 1 2 . 4 - - 3 . 2 - - - -

205 7 8 .4 9 . 6 0 . 4 0 . 8 - - 1 .6 - 0 . 2 0 . 2 - 7 . 6 - - 0 . 4 - - - -

213 7 8 . 2 - 1 .4 1 .0 - 0 . 4 1 .8 - - 0 . 4 - 6 . 2 0 . 4 - 2.0 - 8 . 2 - -

217 8 8 . 0 0 . 8 - 0 . 4 0 . 2 2 . 0 1 .4 - - 0 . 4 - 5 . 8 - - - 1 . 2 - -

221 8 3 . 0 0 . 2 - 0 . 4 - 0 . 2 1 .0 - - - - 6 . 4 0 . 2 - - - 3 . 6 - -

225 6 5 . 0 0 . 2 - 2 . 2 0 . 2 9 . 2 1 .2 - 2 . 6 0 . 2 - 1 6 .4 0 . 6 - 0 . 4 0 . 2 2 . 0 - -

229- 5 7 . 8 0 . 2 - 2 . 2 - 2 9 . 0 0 . 6 - 5 . 0 - - 4 . 8 - - 0 . 2 - 0 . 2 - -

233 61 . 8 0 . 2 . - 0 . 4 - 1 6 . 6 - - 6 . 0 0 . 2 - 1 2 . 8 1 .0 - 0 . 6 - 0 . 2 - -

237 6 3 . 4 0 . 4 0 . 2 1 .2 0 . 4 1 0 . 2 - - 7 .0 - - 1 6 . 2 0 . 6 - 0 . 2 0 . 2 - - -

241 4 9 . 2 4 . 4 2 . 4 7 . 8 0 . 4 11 .4 - - 6 . 4 3 . 6 - 1 1 . 8 0 . 8 - 0 . 4 0 . 2 1 . 2 - -

2 49 2 3 . 6 3 4 . 2 3 . 0 0 . 4 2 . 4 8 . 4 0 . 4 2 . 4 - 3 . 4 1 3 . 4 0 . 2 - 0 . 6 1 . 0 - - -

253 2 6 . 6 2 6 . 4 8 . 6 1 4 . 4 0 . 8 4 . 0 - 0 . 4 1 .2 1 .0 - 1 4 . 8 0 . 8 - 0 . 4 - - 0 . 2 -

257 8 . 8 1 8 . 2 4 . 4 5 5 .0 1 .2 0 . 6 0 . 6 - - 0 . 8 - 00 CO - - 1 .6 - - - -

265 2 9 . 4 1 0 . 8 4 . 8 31 .8 - 0 . 6 0 . 2 - 1 . 4 0 . 2 - 1 9 . 0 - - 0 . 6 1 . 2 - - -

269 3 8 . 2 6 . 0 1 .8 1 0 . 0 1 . 8 1 .4 - 0 . 2 0 . 6 0 . 6 - 3 8 . 4 0 . 4 - 0 . 2 0 . 2 - - -

273 1 3 . 0 0 . 8 6 . 6 2 8 . 4 2 . 8 0 . 2 - - - 0 . 4 - 4 4 . 2 - - 1 .6 1 .4 0 . 2 - -

277 1 5 . 0 1 .2 4 . 6 1 9 . 2 2 . 0 0 . 2 - - 0 . 6 1 . 0 _ 55.4 0 . 2 - 0 . 2 0 . 4 - 0 . 2 -

281 2 0 . 4 1 . 0 6 . 3 11 .4 2 . 2 0 . 4 0 . 2 - 1 . 2 1 . 8 5 3 . 8 0 . 2 - 0 . 2 0 . 4 - - -

285 4 7 . 0 1 .4 4 . 0 4 . 0 2 . 6 - 0 . 2 - 0 . 6 0 . 2 _ 3 9 . 2 0 . 2 - 0 . 6 - - 0 . 2 -

289 5 1 . 8 2 . 2 4 . 0 5 . 4 2 . 0 - - - 1 .0 - _ 3 3 . 0 - - 0 . 2 - - - 0 . 4

293 2 8 . 2 1 3 . 2 2 . 0 3 . 4 1 .0 0 . 2 - - 0 . 6 0 . 2 - 51 .0 - - - 0 . 2 - - -

297 1 4 . 8 2 5 . 2 0 . 8 1 .0 0 . 8 - - - 0 . 4 - - 5 7 . 0 - - - - - - -

301 2 6 . 0 1 8 . 4 0 . 2 1 . 2 0 . 6 - - - 1 .8 0 . 6 - 5 1 . 0 - - - 0 . 2 - - -

305 3 8 . 8 3 . 2 2 . 2 1 .2 1 .0 0 . 2 - - 2 1 . 4 - - 3 2 . 0 0 i 2 - - - - - -

TRAY 1 con td .

No. D.g. S .a. S.c. P. O.i. D.1 C.p. E .v . E. C.n. A. A rt . T3 T4 T8 M. T10 R. ?

309 25.6 - - - - 0.2 - - 59.4 - - 14.8 - - - - - - -

313 18.6 1 .2 - 0.4 0.4 - - - 42.2 1 .8 - 34.3 0.6 - - - - - -

317 59.4 1 .4 0.6 2.4 0.2 11 .4 - - 2.2 - - 22.0 0.4 - - - - - -

321 39.8 0.2 4.4 6.0 0.4 14.2 - 0.2 0.4 - - 32.0 0.4 - - - - - -

325 57.0 6.0 3.0 6,0 0,8 1 .8 - - 1 .0 - - 0 3 # Q - - A 1. w •M’ - - A A -

332 42.2 42.6 7.2 3.0 - 0.6 - - 0.4 - - 4 . 0 - - - - - - -

337 62.4 17.6 3.2 10.8 - - - - 0.2 - - 5.2 0.6 - - - - • - -

341 80.2 3.0 1 .6 8.2 0.4 - - - 0.8 0.2 - 5.0 0.6 ' - - - - - -

345 88.6 1 .0 0.2 4.0 0.2 0.2 - - 0.6 0.2 - 4.6 0.4 - - - - - -

333 77.8 6.2 2.6 7.8 - 9.2 - 0.6 - - - 3.4 1 .0 - - - 0.2 0.2 -

357 75.0 6.0 0.6 4.2 0.2 0.4 0.6 2.8 0.6 0.2 - 8.2 1 .2 - - - - - -

361 50.2 8.8 4.6 8.6 - - 1 .6 1 .4 0.2 3.4 - 17.3 3.4 - - 0.2 - 0.2 -

365 73.8 0.4 0.4 4.6 - - 2.2 1 .4 0.2 0.2 - 15.8 0.8 - - 0.2 - - -

369 61 .0 0.2 0.8 2.8 0.8 - 0.4 1.8 21 .6 - - 8.2 2.4 - - - - - -

373 71 .4 - 1 .2 2.6 - - 1 .2 3.0 4.8 0.4 - 13.0 2.4 - - - - - -

377 75.4 0.2 2.0 2.4 - - - 1 .2 0.4 0.6 - 16.6 1 .0 - - - - 0.2 -

381 69.8 ■0.8 0.6 1 .2 0.2 - 2.4 4.2 0.2 0.2 - 19.8 0.6 - - - - -

385 62.4 4.6 1 .2 ■ 0.6 0.2 4.2 2.4 - 3.0 - 20.4 1 .0 - - - - - -

393 38.8 0.2 - 1 .4 - - 3.0 1 .6 - - - 4.4 0.6 - - - - - -

TRAY I I

No. D.k. S .a. S. c. p t O.i. D.1 C . p . B . v . 17 C.n. A. Art. T3 T4 T8 M. T10 R. ?

4 - 10.2 - 0.6 - - - - - - _ 74.6 - - - - - - -

8 78.6 1.3 0.2 0.1 - 3.1 0.2 - - - - 1 .4 - - 14.5 - - - -

12 78.4 - 6.6 1 .2 - 4.6 0.6 0.2 - - - 4.0 0.4 - 0.8 - 3.2 -

16 86.2 - - 1 .2 - 9.8 0.6 - 0.2 - - 1 .2 0.2 - - - 0.6 - -

20 88.4 - - 0 £ - 4 r\ r\ 1 m\J - - - 0.2 - 0.2 r* 1. v-/ • *+ - n O \ j • i- - - - -

24 63.4 0.6 0.4 6.0 - 10.0 0.4 - - 0.2 - 18.6 0.2 - - 0.2 - - -

28 95.6 0.2 0.6 0.8 0.2 - - - - 0.6 - 1 .4 0.4 - - - - 0.2 -

32 51 .6 2.4 0.2 1 .2 - 1.0 0.4 - 0.2 6.4 - 41.8 0.4 - - 6.4 - - -

36 39.0 7.2 9.6 6.8 0.2 0.4 - - 0.4 4.2 - 29.4 - - 0.6 1 .4 - 0.4 0.4

44 1 .6 0.8 1 .0 1.8 - 3.0 - - 0.2 0.2 - 9.6 0.4 - 81 .4 - - - -

48 5.2 0.8 6.0 12.6 0.2 13.8 - - 0.8 1 .0 - 33.6 0.4 - 25.6 - - - -

32 0.4 1 .0 26.8 11 .4 - 0.4 - - 2.2 0.6 - 50.0 - - 6.6 0.6 - - -

56 2.2 4.2 1 .2 31 .4 - 0.2 - - 0.2 - - 57.2 - - 3.0 0.4 - - -

60 1 .6 0.2 0.2 92.0 - - - - - - - 5.8 - - 0.2 - - - -

67 0.8 11 .5 0.2 84.2 - - - - - 0.3 - 1 .7 0.7 - 0.2 0.5 - - -

71 1 .0 0.8 5.2 85.6 - - - - - - - 6.4 - - - 1.0 - - -

75 2.9 0.9 0.3 60.0 - - - - 0.3 - - 34.8 - - - - - - -

78 5.6 - - 75.6 - - - - - 0.6 - 18.2 - - - - - -

82 16.4 "1 .4 33.2 12.8 0.2 - - - 0.2 7.2 - 27.8 0.6 - - 0.2 - - -

86 10.0 0.8 39.8 22.2 1.8 - - - 0.4 11 .8 - 13.2 - - - - - - -

90 20.6 2.0 15.6 9.8 0.6 1 .0 - - 0.4 3.8 - 46.0 - - - - - 0.2 -

94 20.0 15.0 2.0 5.8 - 0.4 - - 17.2 4.8 - 34.4 - - 0.4 - - - -

98 3.4 25.4 0.6 0.4 - - - - 66.6 1 .2 - 2.2 - - - 0.2 - - -

114 56.4 8.4 1 .2 1 .0 0.2 11 .0 0.2 - 0.4 2.4 - 12.6 - - - 0.2 - - 6.2

122 16.6 4.8 4.8 6.0 - 6.2 - - 0.2 0.4 - 61 .0 - - - - - - -

126 28.2 4.8 0.2 16.4 - 5.4 - - - - - 45.0 - - - - - - -

130 18.3 1 .3 - 19.5 - 1 .0 - - - - - 58.3 1 .3 - - - - - -

134 4.0 0.2 - 7.6 - - - - - - - 87.6 - - - - - - -

138 2.6 2.A - 22.6 - 0.4 - - - - - 72.0 - - - - - - -

142 22.0 1 1 .0 5.8 14.2 - - - - 0.4 0.2 - 44.2 1 .6 - - - - -

146 11 .4 77.9 - 5.1 - - 0.4 - - - - 5.1 - - - - - - -

150 2.2 86.2 3.2 3.0 - - - 3.0 - - - 1.8 0.4 - 0.2 - - - -

154 34.2 44.0 5.4 7.0 - - - 0,2 0.2 - - 9.0 - - - - - - -

158 30.6 1 7.6 12.6 1 6 . 2 - - - 1 .4 1 .0 - - 20.0 0.2 - 0.4 - - - -

TRAY I I con td .

Mo. D.g. 3 .a • S.c. F. C . i . D.1 C.p. r.v. E • C.n. A. Art. T3 T4 T8 M. T10 R. ?

162 31.6 13.0 20.0 1 4.6 - - 1 .2 8.2 0.2 0.6 - 7.2 1 .6 - - - - 0.2 1 .2

166 60.2 1 .2 8.6 13.8 - 1 .2 1 .4 0.2 2.0 - 7.0 4.4 - - - - 0.2 -

170 5.6 1 .2 2.4 1 .0 - - 5.2 0.4 0.4 - - 78.0 5.8 - - - - -

174 35.6 2.2 1.0 3.0 - - 8.0 1 6.4 0.8 2.0 - 27.0 4.0 - - - - - -

1 78 33.4 0.6 1 .4 6.6 - - 5.8 7.0 - i .6 - 39.4 2.8 - - - - 0.4 1 .0182 40.6 - 0.6 2.2 - - 9.4 4.2 0.4 7.0 - 33.6 1 .0 - - - - 1 .0 -

186 63.6 3.6 0.6 0.8 - - 10.0 3.2 - 1 .0 - 17.8 0.2 - - - - - -

190 69.0 6.2 2.4 0.2 - - 5.0 0.8 0.8 3.8 - 10.2 0.6 - 0.6 - - - -

194 60.4 1 5.0 0.2 0.2 - - 3.0 1.0 0.4 3.8 - 15.2 - - 0.8 - - - -

198 63.8 7.8 - 0.4 0.6 - 2.2 - 0.2 1 .4 - 22.8 0.2 - 0.6 - - - -

202 61 .8 9.2 - 0.2 - - 1 .0 - 0.2 0.8 - . 23.8 - - 2.8 - - - 0.4

206 33.8 22.6 - 1 .6 - - 3.4 - - 0.2 - 37.0 - - 0.8 - - - -

210 54.0 0.2 1 .6 0.2 - - 2.6 - - - - 23.8 - - 5.4 - 7.6 - -

218 37.6 - 0.2 0.4 - 1 .4 - - - 1.8 - 7.8 0.2 - - - 0.4 0.2 -

222 77.4 0.4 1 .4 1 .2 - 1 .4 0.8 - 0.4 0.8 - 10.0 0.4 - - - 5.8 - -

226 32.0 - - 3.0 - 4.6 - - 0.4 - - 6.8 0.4 - - - 2.6 - -

230 22.4 - 1 .0 5.4 - 61.0 - - 0.4 0.2 - 8.2 0.8 - - - 0.6 - -

234 7.2 - - 0.6 - 17.0 0.2 - 66.4 - - 8.4 0.2 - - - - - -

238 0.2 0.2 - 0.2 - 2.0 - - 96.8 0.4 - 0.2 - - - - - - -

242 34.2 11 .2 1 .8 12.0 - 8.2 0.2 - - 2.8 - 28.8 0.2 - - 0.6 - - -

246 4.0 5.6 - 5.0 - 1 .6 - - 0.4 0.4 - 82.8 - - - 0.2 - - -

250 7.6 40.2 1.8 10.2 - 6.4 - - 0.4 4.6 - 28.4 - - - 0.4 - - -

254 8.4 42.2* 9.8 20.6 - 0.6 - - 1 .0 8.2 - 5.3 - - 0.3 - - -

258 0.2

COITNr\i - 44.6 - 0.2 - - 0.8 - - 27.0 - - 1 .2 0.2 - - -

262 0.8 6.6 0.2 31 .0 0.2 - - - - - _ 61 .2 - - - - - - -

266 1.0 4.3 - 42.4 - - - - 2.0 0.8 - 4S.0 - - 1 .0 - - - -

274 0.8 9.8 2.4 51 .8 - - - - - 1.0 - 34.2 - - - - - - -

278 2.4 0.2 2.6 40.0 - - - - 0.2 5.8 - 48.6 - - - - - 0.2 -

282 3.2 12.6 13.8 17.6 - 0.4 - - - 1 .2 - 50.8 0.2 - - - - 0.2 -

286 2.1 26.8 1 .1 11.8 - 0.3 - - 1 .1 1 .1 - 55.7 - - - - - - -

290 0.8 17.2 0.2 7.8 - - - - - 0.2 - 66.0 - - - 7.8 - - -

294 3.6 21 .8 0.2 3.0 - - - - C.2 0.2 - 71 .0 - - - - - - -

306 34.^ 1 9.6 - 0.4 - 0.4 2.8 - 2.0 0.4 - 39.4 0.2 - - 0.4 - - -

310 11 .4 13.8 1 .0 2.0 - 0.2 0.6 - 43.2 1 .6 - 26.2 - - - - - - -

TRAY I I con td .

No. D.g. S.a. S.c. P. O.i. D.1 C . p . E .v . E. C .n . A. A rt. T3 T4 t3 M. T1 0 R. ?

314 14.0 29.4 0.2 - - 0.4 - - 2.4 5.8 - 47.8 - - - - - - -

318 7.6 2.0 0.2 0.2 - 54.6 - - 0.8 - - 34.2 - - - 0.2 - - 0.2

322 59.6 10.4 - 0.4 - - - - 0.2 - 11 .6 - - - 0.8 - “ —

330 1 .2 91 .6 5.4 0.4 - - - - - - - 1.0 - - - 0.4 — — "

334 0.6 96.8 0.6 - - - - - - 0 2 1.8 - - - - - “

338 1 .0 85.0 4.4 - 3.8 - - - - - - 5.0 0.8 - - - -

342 25.6 20.8 18.4 9.0 - 1 .0 0.8 - - 1 .2 - 21 .6 1 .2 - - - - 0.4

346 7.4 22.8 1 .6 7.6 - - - - - 4.4 - 54.4 1 .6 - - - - 0.2

350 12.6 20.2 1 .6 4.6 - 0.2 0.4 - 0.2 0.2 - 57.2 2.4 - - 0.2 - 0.2 “

354 6.2 64.4 2.6 17.2 - - 1 .6 1 .2 - - - 5.2 1 .6 - - - - -

358 3 2 . 2 32.4 1 .6 12.0 - - 1.8 1 .0 0.4 4.4 - 9.6 3.8 - - 0.6 - 0.2 “

370 36.8 3.4 10.4 14.0 - - 2.0 6.2 1 .2 1 .0 - 21 .8 3.2 - - 0.2 - - “

374 35.8 13.6 0.8 10.4 - - 5.2 6.2 0.8 0.8 - 20.0 6.0 - - 0.2 - 0.2

378 30.6 2.0 5.4 6.8 - - 4.0 14.2 0.2 1 .6 - 31 .8 2.2 - - 0.6 - 0.6

382 16.6 0.4 5.6 5.4 - 0.2 18.4 0.8 - 1 .4 - 43.8 6.8 - - 0.6 - -

386 14.4 1.8 - 4.2 - - 4.4 1 .2 - 5.0 - 57.8 11 .0 - - - - 0.2 -

394 51.0 6.8 1 .6 7.8 - - 9.8 1.8 - 0.4 - 20.4 0.4 “ ~ ~ - "*"

TRAY I I I

No. D.g. S. a. 0 .c P. O.i. D.1 C . p . P V E. C . n . A. Art. T3 T4 mQ M. T1 0 R. ?

5 - 71 .0 - 1 .1 - - 10.9 - - - _ 14.6 2.4 _ _ - - - -

9 51.2 6.7 3.0 1 .3 - 1 .7 9.0 - - - - 20.2 1 .1 - 5.2 0.1 0.1 - -

13 68.8 - 10.2 1 .4 - 10.4 3.8 - - 0.6 - 4.4 0.2 - 0.4 - 0.8 - -

17 62.2 - 4.4 0.2 - 12.2 1 .0 - 0.6 0.6 - 14.0 - - 0.4 - 4.2 - 0.221 56.6 - 0.8 0.4 0.8 4.6 0.2 - 4.0 2.8 - 27.0 0.2 - i .3 - - - -

23 6l .8 - 0.8 - 0.2 0.8 0.4 - 5.6 1 .6 - 24.4 - - - 1 .2 - -

29 79.8 0.2 1.2 0.4 0.2 0.2 0.2 - - 1 .4 - 1 6.4 - - - - 0.2 - 0.2

33 39.0 - 1 .4 1 .0 1.0 0.4 0.4 - 0.2 10.2 - 44.8 0.6 - - 0.6 - - 0.4

37 37.2 4.5 0 .9. 2.8 - 1 .2 - - 0.2 - - 49.4 0.2 - 3.4 0.6 - - -

41 5.0 .0.8 0.4 1 .2 0.6 0.6 - - 0.2 15.0 - 71 .8 - - 3.6 0.4 - - 0.4

49 2.3 4.8 3.0 8.2 0.2 17.0 2.9 - 0.4 - - 54.5 - - 4.2 - - 1 .9 0.2

53 ^ 6.6 0.6 0.8 23.5 - 0.4 - - 0.4 - - 45.3 0.1 - 21 .5 - - 0.7 -

57 ' 2.2 4.4 7.4 45.6 - 0.1 - - 0.8 - -

OOIPs - - 2.9 0.1 - 0.7 -

61 1.4 - 2.4 91 .8 - 0.2 - - - - - . 3.4 - - 0.2 - - - 0.6

64 - 0.1 1 .8 84.1 - - - - - - - 14.0 - - - - 0.1 -

68 5.5 0.1 4.1 46.3 0.4 0.9 0.7 - 0.5 8.0 - 33.3 - - - - - - -

72 13.7 0.6 8.4 56.7 4.3 0.4 0.1 - 0.1 - - 15.7 - - - - - - -

76 2.6 0.5 7.6 53.2 0.2 - - - - - - 35.7 - - - - - - -

79 7.3 5.3 21 .1 42.2 - 0.1 - - 0.3 - - 23.5 - - - 0.2 - - -

83 2.2 0.3 9.4 6.1 0.2 - 0.1 - 0.2 13.1 - 68.4 - - - - - - -

87 0.5 0.1 , 1 .6 94.3 - - - - - - - 3.5 - - - - - 0.1 0.1

91 5.8 0.2 40.0 1 .2 - - - - 1 .4 - - 48.1 - - - - - 3.2 -

95 8.6 1 .2 7.4 1 .6 - - - - 48.0 - - 33.8 0.2 - - - - - 2.0

99 8.2 46.2 11 .6 0.2 - - - - 28.0 - - 8.0 - - - - - - 0.2

103 6.2 82.2 8.4 0.4 0.2 - - - 0.2 - - 1.8 0.2 - 0.2 - - - -

107 1 .4 82.6 12.6 0.2 - 1 .4 - - - - - 2.0 - - - - - 0.2 -

111 23.6 21.0 16.0 1 .4 - 14.2 - - 0.2 - - 23.0 - - - - - 0.2 0.4

115 24.S 0.8 7.8 0.4 0.4 39.8 0.2 - - - - 25.8 - - - - - - -

119 48.0 - 6.3 2.6 9.8 - 0.2 - - - 32.6 - - - - - - -

123 48.3 - 3.7 11 .7 - 7.5 - - - - - 28.5 0.5 - - - - - -127 23.2 0.4 2.2 11 .8 - 10.4 0.2 - - - - 51 .6 0.4 - - - - - 0.6

131 37.8 0.4 3.8 16.0 - 3.4 0.4 - 0.2 - - 38.2 0.2 - - - - - -

135 44.3 0,1 0.3 10.6 - 1 .4 0.4 - - - - 42.1 0.3 - - 0.1 - - 0.1

139 30.6 0.6 7.4 14.2 0.2 - 0.8 - 0.2 - - 44.0 0.8 - - - - - 1 .2

TRAY I I I con td .

Ho. D.g. 3 • ci • S.c. P. O.i. D.1 c.?. B.v. E. C.n, A. Art. T3 T4 T8 V T1 0 R. 0

143 23.4 7.6 12.4 14.6 - 0.2 0.4 - - - - 27.2 1 .2 - - - - - 3.0

147 6.8 55.4 13.2 3.4 - - 0.2 - - 0.2 - 20.6 - - - - - 0.2

151 2.0 66.6 18.4 3.2 - - 0.2 - - - - 9.6 - - - - - - -

155 14.4 40.6 19.0 3*2 - 0.2 1 .8 1 .0 3.6 0.4 - 15.4 0.4 -O

- - - -

159 34.2 4.6 17.2 8.0 - - 2.0 3.4 0.4 - " 23.6 3.2 — 1 - “ 0.6 0.4

163 53.4 1 .0 16.8 4.2 - - 1 .6 5.6 - 3*2 - 11 .4 1 .8 - 0.6 - - 0.4

167 46.4 3.0 4.2 10.0 - - 7.6 3.4 0.2 4.0 - 13.8 6.8 - - 0.2 - 0.2 0.2

171 57.4 0.4 6.6 3.8 0.2 0.2 6.4 5.0 0.2 0.4 - 11 .4 6.8 - - - 0.2 0.4 0.6

175 42.0 1 .0 6.8 2.0 0.2 7.6 2.0 0.2 2.0 - 33.2 3.0 - - - - - -

179 17.2 0.2 2.2 1 .0 - 0.4 3.0 1 .6 - 0.4 - 12.4 1 .6 - - - - - -

183 49.2 0.4 2.8 0.4 - - 4.6 2.2 - 2.2 - 36.6 1 .2 - - 0.2 - - 0.4

187 40.0 3.2 1 .6 0.8 - - 6.4 2.2 - 5.0 - 40.8 0.4 - - - - - 0.2

191 53.4 2.2 6.0 0.6 - - 5.0 1 .8 0.8 2.3 - 27.4 0.2 - - - - - 0.2

195 59.2 9.2 1 .6 0.4 - - 1 .8 1 .2 1 .6 3.8 - 20.8 0.4 - - - - - -

199 39.6 35.2 2.4 - - - 2.0 - 0.4 1.6 18.6 0.2 - 0.6 - - - -

203 38.8 27.4 2.0 0.6 - - 1 .4 - 12.0 0.2 - 13.4 - - 4.2 - - - -

207 51 .8 2Z}.2 1 .8 1 .0 - - 3.4 - 0.2 0.2 - 16.6 - - 1 .2 0.2 - - -

211 41 .4 4.6 1 .4 1 .6 - - 5.0 - 0.2 1 .6 - 40.6 0.2 - 3.4 - - - -

215 66.0 - 1 .6 2.6 - 0.2 1.6 - - 0.6 - 19.6 0.2 - 0.6 - 7.4 - -

219 87.2 - 1 .0 0.8 - 1 .6 1 .4 - 0.2 0.4 - 6.2 0.2 - - - 1 .2 - -

223 74.4 - 0.8 1.0 - 1 .2 0.6 - - - - 14.4 - - - - 7.2 - -

227 55.4 1 .6 0.2 1.0 - 18.2 - - 0.4 0.2 - 8.8 - - 1 .0 - 13.6 - -

231 42.8 - - 1 .2 - 34.2 0.2 - 3.4 0.2 - 16.8 - - - - 1 .2 - -

235 15.6 0.2 0.2 1 .0 - 26.0 0.4 - 16.6 - - 40.0 - - - - - - -

239 8.2 1 .2 2.8 1 .6 - 5.6 0.4 - 23.6 0.8 - 54.6 0.6 - - 0.6 - - -

243 4.4 5.8 7.4 4.0 - 2.4 0.4 - 0.4 6.0 - 68.2 0.6 - - 0.2 - 0.2 -

247 4.4 1 5.6 10.2 2.0 - 0.4 - - 0.4 3.6 - 62.8 - - - 0.4 0.2 - -

251 1 .6 35.8 23.0 1 .0 0.2 0.6 0.4 - 0.2 6.8 - 30.2 - - - - - 0.2 -

255 3.2 46.6 23.8 9.6 - 0.2 - - 0.2 7.2 - 12.6 0.2 - - 0.6 - - -

259 0.8 37.0 17.4 28.2 0.2 0.2 - - 0.4 3.2 - 7.2 - - 4 .2 0.4 - 0.8 0.2

263 0.8 6.2 15.2 56.6 _ _ 0.2 _ 0.6 - 1 6.0 - - 0.8 0.2 - 0.2 -

267 0.4 5-6 17.8 34.6 _ 0.6 0.2 _ 0.2 5.0 - 34.6 - - 0.4 0.2 - 0.2 0.2

271 4.4 1 .2 27.1 17.8 0.6 0.1 - 0.3 0.2 1 .7 - 46.4 - — 0.2 0.1 _ 0.1 —

TRAY I I I con td .

No. D.g. S . a . 3. c. P. O.i. D.1 C .p. E .v. E. C n A. Art, T3 T4 T8 M • T10 R. ?

275 0.6 1 .6 30.2 54.2 - - - - - 5.4 - 8.0 - - - - - - -

279 5.0 0.4 19.8 14.2 0.4 - 0.2 - - 8.6 51 .2 - - - - - 0.2 -

283 3.2 1 .0 32.2 9.4 0.2 - 0.2 - 0.4 7.0 - 46.4 - - - - - - -

287 1.8 3.2 19.0 0.6 - - 0.2 - 0.6 7.4 - 67.4 - - - - - - -

291 0.8 9.2 9.2 3.4 0.4 - 0.2 - 0.8 2.2 - . 73.6 - - - 0.2 - - -

295 5.0 15.4 15.0 0.8 - - 0.2 - - 1 *2 - 62.2 - - 0.2 - - - -

299 1 .0 45.6 11 .2 0.2 - - - - - 1 .2 - 40.6 - - - 0.2 - - -

303 6.8 30.0 4.6 0.2 - - - - 1 .4 0.4 - 56.6 - - - - - -

307 4.4 1 .0 10.2 C.4 - 1 .4 0.2 - 31.0 0.2 - 51 .0 - - - - - - -

311 4.2 - 5.2 0.2 - 5.2 - - 67.8 0.8 - 20.6 - - - - - - . -

315 2.2 - 6.8 - 0.2 1 .0 - - 0.2 6.0 - 82.6 - - - - - ~ 0.8

319 11 .0 1 .6 11 .6 1 .2 - 62.4 - - 0.2 - - 11 .6 0.4 - - - - -

323 3k A 1 .2 10.8 1.8 - 10.8 - - 0.6 - - 24.0 0.4 - - - - - -

327 5.6 27.0 13.8 3.2 - 1 .6 - - 1 .8 0.2 - 46.2 0.4 - - 0.2 - - -

331 3.6 86.2 4.6 1 .0 - - 0.8 - 0.2 0.2 - 3.0 - - 0.2 - 0.2 -

335 4 .3 80.4 6.8 0.8 - 0.2 - 0.4 - - - 6.2 0.4 - - - - •- -

339 3.6 63.2 18.8 1 .0 - 0.8 0.4 - 0.2 - - 12.0 - - - - - - -

347 . 47.2 6.8 13.6 9.4 - 1 .4 3.2 - 0.2 0.6 - 15.0 1.8 - - 0.6 - 0.2 -

351 40.4 14.0 9.0 6.6 - 0.2 4.2 - 0.4 4.2 - 13.6 2.0 - - 0.4 - - -

355 36.0 40.0 7.6 8.0 - - 0.6 0.6 - - - 5.0 2.0 - - 0.2 - - -

359 48.0 13.2 3.0 9.4 - - 5.0 0.6 - 2.6 - 14.8 3.2 - 0.2 - - - -

3b5 74.4 1 .2 0.2 7.0 - 0.2 2.6 1 .4 - - - 12.0 1 .0 - - - . - - -

367 51.6 1 .2 0.8 10.6 - - 1 .0 2.4 2.4 4.6 - 21 .0 4.4 - - - - - -

371 24.8 - 3.0 7.0 - 0.2 2.6 2.4 41 .0 0.3 - 14*6 3.2 - - - - 0.4 -

375 15.2 1 .6 6.4 12.0 - - 3.2 2.8 33.4 2.2 - 19.2 4.0 - - - - - —

379 18.4 0.8 2.S 6.0 - - 4.2 4.4 12.8 4.4 - 41 .0 5.2 - - - - ~ -

333 46.0 0.6 4.2 2.4 - - 12.2 0.2 0.2 1.6 - 31.0 1 .4 - - 0.2 - - -

387 15.8 16.8 1 .4 1 .2 - - 10.0 4.6 - 11 .8 - 35.4 2.8 - - 0.2 - - -

395 23.6 0.2 3.8 ■2. Q,D • 0 - - 11 .2 0.4* - 1 .0 - 55.6 0.4 — “ - —

TRAY IV

No. D.g. S.a. S.c. P. C . i . D.1 C.p. B.v. E. C.n. A. Art. T3 T4 T8 M. T10 R. ?

6 _ 62.0 - 0.4 - - 27.4 - - - - 10.0 0.2 - - - - - -

10 56.3 4.0 0.2 1 .1 - 0.5 4.5 - 0.1 - - 22.6 0.3 - 7.6 0.1 2.3 0.2 0.2

14 CO 00 0.4 2.8 0.8 - 5.2 3.4 0.2 0.2 2.2 - 2.0 0.4 - - - 0.6 - -

18 78.8 0.2 0.2 1 .0 0.2 14.0 0.4 - 0.2 0.2 - 3.4 - - - 0.2 1 .2 - -

22 56.0 0.2 0.2 0.2 - 23.4 1 .0 - 5.4 0.4 - *0 0.8 - - 0.2 1 .2 - -

26 70.2 2.0 0.6 1 .2 - 7.0 0.2 0.2 0.6 - - 16.0 0.4 - - 1 .4 - - 0.2

30 1.6 0.6 - 0.6 - 0.6 - - - - - 96.6 - - - - - - -

34 2.4 - 7.4 0.4 - - 0.2 - - 0.2 - 89.4 - - - - - - -

33 12.6 2.0 0.8 1 .4 - 1.0 0.2 - 0.2 - - 81 .0 - - - 0.8 - - -

42 15.2 3.6 3.4 1 .0 1.8 2.0 0.4 - 0.6 - - 68.6 - - 0.2 - - 1 .2 1.8

46 8.8 3.8 0.2 6.0 0.4 0.6 - 0.4 0.8 - - 8.4 - - 71 .0 - - - ~

5° 11 .2 3.8 0.4 11 .4 0.2 1 8 . 8 0.2 - 0.2 0.8 11 .0 0.4 - 40.6 1 .0 - - -

54 6.8 1 .4 0.8 22.6 - 1 .2 0.2 - 0.8 1.2 - 27.0 - - 37.4 0.6 - - -

58 3.4 5.2 3.6 60.8 - 0.4 - - - - - •8.8 - - 17.8 - - - -

65 3.6 0.2 0.4 93.0 - 0.2 0.4 - - - - 1.2 0.2 - 0.8 - - - -

69 7.6 2.0 5.6 41 .8 - - - - 13.6 0.4 - 28.6 0.2 - - 0.2 - - -

73 2.2 1 .0 0.4 47.6 0.4 - - - - - - 48.0 - - - 0.4 - - -

77 2.6 0.5 7.6 53.2 0.2 - - - - - - 35.7 - - 0.2 - - - -

80 2.6 0.2 5.0 78.2 0.4 - - - 0.8 - - 12.2 0.2 - - 0.2 - - 0.2

84 1 .2 1 .8 4.4 7.0 - - - - 0.4 1.8 - 83.4 - - - - - -

88 41 .9 0.2 2.4 6.1 1 .5 - - - 2.8 1 .5 - 40.4 4.3 - - - - - -

92 13.2 0.8 0.6 1 .6 0.2 0.2 - - 16.4 2.6 - 55.4 9.0 - - - - - -

96 0.6 0.6 0.2 0.4 - 0.2 - - 79.8 1 .0 - 16.8 0.4 - - - - - -

100 1 .4 21 .0 2.4 0.8 - 0.2 - - 61.0 0.8 - 11.8 0 .8 - - - - - -

104 1 .0 35.6 - 0.4 0.2 - - - 9.6 - - 3.0 - - - - - - 0.2

108 5.4 91 .3 - 0.6 - 0.2 - - - - - 2.0 - - - - - - -

112 70.6 10.6 1 .4 1 .4 0.4 8.2 - - 2.2 0.2 - 4.4 - - 0.4 0.2 - - -

116 15.0 7.2 3.0 13.0 0.4 47.2 0.6 - 1 .0 1 .2 - 9.4 1 .2 - 0.6 0.2 - - -

120 25.4 2.8 0.8 26.6 - 5.6 - - 0.4 0.2 - 31.2 3.4 - 0.4 0.4 - - 3.0

128 15.8 0.2 - 13.4 - 5.0 1 .4 - - - - 63.8 0.2 - - - 0.2 - -

132 17.8 0.6 - 19.6 - 3.0 - - - - - 57.6 0.2 - - - - - 1 .2

136 41 .2 - - 17.2 - 0.2 - - - - - 41 .4 - - - - - - -

140 47.6 1 .0 0 . 4 9.6 - 0.4 - 0.8 0.2 - - 39.6 0.4 - - - - - -

144 43.0 1 .4 14.6 13.4 - 0.2 - - 0.2 ■ 0.2 - 26.6 0.2 - 0.2 - - - -

TRAY IV c o n t d .

No. D.g. S . a . S.c. P . O.i. D.1 C.p, E.v. ■u 9 C.n. A. Art. T3 T4 T8 M. T10 R. ?

148 19.0 35.6 22.4 8.8 - 0.4 - - - - 13.6 0.2 - - - - - -

156 1 .6 53.0 35.6 4.4 - 0.2 0.4 0.8 - - 3.8 0.2 - - - - - -

160 7.0 3.4 30.6 17.0 - - 2.0 4.4 0.2 0.2 - 29.8 3.2 - 2.2 - - - -

164 9.2 2.8 29.6 22.0 - 0.2 5.0 16.0 - 0.4 - 8.8 5.3 - - - - 0.2 -

168 10.8 0.8 2.2 6.6 - - 1 13.6 1 .2 1 .6 - 56.2 5.0 - 0.2 0.4 - - -

172 43.4 1 .6 1 .8 7.4 - - 2.6 20.2 - 0.2 - 17.6 5.4 - 0.2 - - - -

180 4.0 0.4 1 .6 0.8 - - 6.8 0.8 0.2 - - 83.0 0.6 - - - 1 .8 -

184 6.2 - 0.8 - - - 8.4 0.6 - - - 83.2 - - - - - 0.8 -

188 8.2 1 .0 6.8 0.6 - - 35.0 2.4 0.4 1.8 - 40.2 3.4 - - - 0.2 - -

192 13.8 0.4 4.4 - - - 5.6 1 .8 0.2 2.2 - 70.6 0.6 ~ 0.4 - - - -

196 22.3 3.5 2.2 0.2 - - 4.3 0.2 0.2 0.4 - 66.0 0.2 - 0.4 - - - -

200 63.8 5.4 - - - - 0.8 - 2.0 1 .0 - 26..0 - - 1 .0 - - - -

204 59.0 5.4 0.2 1 .0 - 0.2 1 .0 - 1 .2 0.2 - 24'. 2 - - 7.6 - - - 0.2

208 18.8 7.8 2.6 0.2 - 0.2 2.4 - 2.6 - -

0COirv - - 7.4 - - - -

21 6 18.2 0.4 - 0.4 - - 2.6 - - 1 .4 - 71 .8 - - 3.8 - 1.8 - -

220 83.4 0.2 0.2 0.4 - 0.2 0.2 - - - - 14.0 - - - - 1.4 - -

224 44.4 1 .2 7.4 3.2 - 6.4 3.8 - 5.4 0.2 - 19.4 - - - 0.6 8.2 - -

228 30.8 0.4 - 5.4 0.4 20.8 3.6 - 0.8 0.6 - 33.6 0.6 - 0.2 - 2.6 0.2 -

232 14.8 1 .4 0.2 2.2 0.2 71 .6 1 .k - 1.2 - - 5.4 0.2 - - - 1 .4 - -

236 4.0 1 .0 0.4 4.0 - 76.0 0.4 - 0.8 1 .0 - 10.6 0.6 - - - - - -

240 20.8 2.0 0.2 1 .4 - 9.0 0.4 - 0.8 4.0 - 60.2 0.6 - 0.2 0.2 - - -

244 19.8 3.4 1.8 3.2 - 7.2 - - 0.4 0.8 - 61.2 1 .6 - - 0.2 0.2 - -

248 10.6 9.6 19.4 4.2 0.4 16.8 - - - 1 .6 - 36.4 0.2 - 0.6 0.2 - - -

252 2.8 23.4 30.4 2.8 0.2 5.3 0.2_ - 1 .0 2.0 - 30.6 0.2 - 0.6 - - - -

256 7.6 26.2 16.0 5.8 0.4 2.6 0.4 0.2 - 3.0 - 37.6 - - 0.2 - - - -

26i* 6.6 9.4 17.6 46.0 0.4 0.6 0.2 - 0.2 3.6 - 13.0 0.4 - 1 .2 0.4 - 0.4 -

276 3.2 1 .6 24.0 56.2 - 1 .2 - - - - - 12.0 0.6 - 0.8 0.4 - - -

284 0.6 4.8 48.0 1.8 - - - - 7.4 - 37.2 - - - 0.2 - • - -

288 1 .4 7.8 39.0 0.6 - - 0.2 - - 3.6 - 47.0 - - - 0.2 - - -

292 1 .8 5.0 25.6 0.8 - - - - 0.6 1 .0 - 65.4 - - - - - - 0.2

296 0.4 30.2 12.0 0.2 - - - - 0.6 1 .6 - 55.0 - - - - - - -

300 - 56.6 2.4 - - - - - 0.8 0.2 - 40.0 - - - - - - -

304 2.0 34.0 . 6.6 - - - 0.2 - 1 .0 3.8 - 52.4 - - - 0.2 - - “

VjJ 0 CO 2.8 1 .2 7.8 1 .2 - 1 .4 1 .0 - 56.8 0.2 - 27.8 - —"

II

TRAY IV c o n t d .

No. D .e . S . a . S .c . p . 0 . 1 . D.1 C .p . E .v . E. C , n . A. Art . T3 T4 T8 M. T10 R. ?

312 1 .4 1 .4 3 .6 _ 0 .2 _ 68.4 0 . 8 - 24 .8 - - - - - - 0 .2

316 13 .4 0 . 8 7 .4 0 .2 - - 0 .2 - 10.4 8 .6 - 53.4 - - - - - - 0 .2

324 11 .6 6 .0 2 8 .6 2 . 0 - 1 7 .4 - - 3 .6 2 .0 - 2 8 .2 - - - 0 .4 —

328 13 .6 3 4 .6 2 8 .8 0 .2 - 0 . 4 - - 0 .4 1 .2 - 2 0 .8 - - - - - “ •

332 2 .4 9 0 .2 4 .2 O. p - 0 .4 - - - - - 2 2 - - - - “ - -

336 4 .2 8 7 .0 5. 2 1 .0 - - 0 .2 - - - - 2 .4 2 .2 - - ~ —

340 4 .6 6 9 .8 6 . 4 1 .6 - - - - - 0 .2 - 17.2 - - - 0 .2 - —

344 6 .8 3 4 .4 18 . 2 3 . 6 0 .2 2 . 8 2 .0 0 .2 0 .6 1 .4 - 2 9 .4 0 . 6 - - - - - —

348 1 8. 6 7 .8 2 6 .0 4 .4 - 0 .6 5 .4 1 .2 0 .4 4 .4 - 3 0 .0 0 .2 - - 0 .6 - 0 .2 -

352 2 2 .6 1 9 .2 16 .8 3 . 6 - 0 .2 3 .4 0 .2 0 .2 1 .8 - 3 0 .0 1 .2 - 0 .2 0 .6 - 0 .2 -

356 9 .4 5 2 .2 16 . 6 2 .2 - - 1 .4 1 .2 1 .2 - - 15.2 0 .2 - 0 .2 0 .2 - - -

360 1 3 .8 2 3 . 8 8 .2 3 .8 - - 7 .8 1 .0 - 6 .6 - 3 4 .2 0 .4 - - 0 .2 - -

364 12 .6 2 . 4 4 .4 5 .0 - 0 .2 7 .0 3 .4 0 .4 5 .3 - 56.4 1 .0 - 1 .2 0 .2 - - -

368 8 .2 1 .0 7 .2 4 . 6 - - 3 .6 0 .2 2 .8 11 .4 - 60.2 0 .6 - - - “ 0 .2

372 11 .4 1 .0 6 .6 7 .2 - - 2 .2 1 .0 12.2 4 .6 - 52.4 1 .6 - - - —

376 16. 2 0 . 4 10.6 4 . 2 - 5 .6 0 .2 2.2 7.6 - 52 . 6 0 .6 - - - ~ —

380 2 5 .3 0 . 2 4 .8 3 .0 - 7 .4 1 .6 1 .0 8.C - 4 7 .6 - - - 0 .2 ~ — 0 .4

384 3 8 .0 1 .0 8 .0 1 .2 - - 2 0 .0 2 .4 - 1 .0 - 28 .0 0 .4 - - - “ “

388 8 .4 0 . 2 11 .0 1 .4 - - 1 2 .4 0 .2 0 .2 9 .2 - 56.2 0 . 4 - - 0 .2 - -

396 3 3 .0 0 . 2 2 .8 1 .4 - - 2 0 .6 - - 2 .4 - 3 9 .0 0 .2 ~ - 0 .4"