6 . VEGETATION OF LITTLE CAYMAN
D . R . S t o d d a r t
Althouc~h t h e v e g e t z t i o n 3f L i t t l e Caxpan h a s n o t bccn p r e v i o u s l y d e s c r i b e d , t h e l s l a n d b e l o n g s t o r_he c l a s s o f lo..i, d r y , l i m e s t o n e i s l a n d s ir, t h t C a r i b b e a n which h a s zl t t r a c t e d inuch a t t e n t i o ~ l . a e a r d
(1944, 1949, 1955) p r o v l d e s a c l a s s i f i c a t i o n o f t h e v e g e t a t i o n and d e s c r i p t i o n s o f t h e Windward and Leexard I s l a n d s which many subsequerlk workers have u s e d a s a model. S i m i l a r t r e a t m e n t s have been p r o v i 2 e d f o r l i m e s t o n e a r e a s i n Jamaica (Asprey and Robbins 1953, L o v e l e s s and Asprey 1957, Asprey and L o v e l e s s 1 9 5 8 ) , Cuba ( S e i f r i z 1943, Howard and B r i g g s 1 9 5 3 ) , A n t i g u a ( L o v e l e s s 1 9 6 0 , H a r r i s 19651, Barbuda ( H a r r i s 1965, S t o d d a r t , i n l i t t . ) , A n g u i l l a ( H a r r i s 1 9 6 5 ) , and Anegada (D'Arcy 1 9 7 1 , 1 9 7 5 ) . C o a s t a l s t r a n d v e g e t a t i o n s i m i l a r t o t h a t o f L i t t l e Cayman h a s b e e n w i d e l y d e s c r i b e d i n t h e C a r i b b e a n a n d t h e Gulf o f M e x ~ c o ( S a u e r 1959, 1967, P o g g i e 1962)
.
D r J . D . S a u e r h a s i n c l u d e dt h r e e L i t t l e Cayman t r a n s e c t s i n a s t u d y o f c o a s t a l v e g e t a t i o n o f t h e Cayman I s l a n d s which h e began i n 1962 and which w i l l i n due c o u r s e l e a d t o a m a j o r p u b l i c a t i o n .
T h i s c h a p t e r i s d i v i d e d i n t o f o u r p a r t s on a l a r g e l y e m p i r i c a l b a s i s . The f i r s t t r e a t s i n l a n d v e g e t a t i o n communities on l i m e s t o n e , t h e second c o a s t a l communities, and t h e t h i r d mangrove communities.
B e a r d ' s scheme i s f o l l o w e d f o r t h e l i m e s t o n e a r e a s , though it seems l e s s a p p r o p r i a t e f o r c o a s t a l v e g e t a t i o n and c o v e r s mangrove communities o n l y i n b r o a d o u t l i n e . The f o u r t h s e c t i o n o u t l i n e s a c l a s s i f i c a t i o n o f t e r r e s t r i a l h a b i t a t s o f b r o a d e c o l o g i c a l s i g n i f i c a n c e on L i t t l e Cayman ( F i g u r e 21)
.
A . I n l a n d communities on l i m e s t o n e
The v e g e t a t i o n o f L i t t l e Cayman, o t h e r t h a n t h e c o a s t a l and swamp communities, f a l l s w i t h i n t h e g e n e r a l c l a s s o f Dry E v e r g r e e n F o r m a t i o n s r e c o g n i s e d b y Beard (1944)
,
e x t e n d e d and d e f i n e d b y Beard (1949) , and r e v i s e d b y Beard ( 1 9 5 5 ) . Dry Rain F o r e s t , o r i g i n a l l y termed X e r o p h y t i c R a i n F o r e s t ( B e a r d 1 9 4 4 ) , i s n o t found i n t h e Cayman I s l a n d s . Four o t h e r t y p e s c a n , however b e r e c o g n i s e d : Dry E v e r g r e e n F o r e s t , DryA t o l l R e s e a r c h B u l l e t i n No. 2 4 1 : 53-70, 1 9 8 0 .
E v e r g r e e n Woodland, Dry E v e r g r e e n T h i c k e t , a n d E v e r g r e e n B u s h l a n d . I n a c o n t i n u a l l y v a r y i n g and a l s o l i t t l e known a n d g e n e r a l l y i n a c c e s s i b l e v e g e t a t i o n s u c h a s t h a t o f L i t t l e Cayman, where c o m m u n i t i e s a r e
f l o r i s t i c a l l y d i v e r s e b u t where many s p e c i e s a r e r e p r e s e n t e d i n d i f f e r e n t v e g e t a t i o n t y p e s , i t i s d i f f i c u l t t o a s s i g n l i m i t s t o t h e s e c a t e g o r i e s . H a r r i s ( 1 9 6 5 ) f o u n d a s i m i l a r d i f f i c u l t y i n A n t i g u a , A n g u i l l a a n d
B a r b u d a ( t h o u g h a l l t h e s e i s l a n d s have v e g e t a t i o n much m o d i f i e d by m a n ) , a n d combined t h e Dry E v e r g r e e n Woodland a n d E v e r g r e e n B u s h l a n d i n a s i n g l e c a t e g o r y o f E v e r g r e e n Woodland. Some c o n f u s i o n a l s o a r i s e s from t h e f a c t t h a t A s p r e y a n d Robbins ( 1 9 5 3 , 380-383) d e s c r i b e a Dry L i m e s t o n e S c r u b F o r e s t w i t h B u r s e r a a s an e m e r g e n t , which c l e a r l y h a s p a r a l l e l s i n L i t t l e Cayman, b u t e q u a t e i t w i t h B e a r d ' s Semi-Evergreen S e a s o n a l
F o r e s t , one o f h i s S e a s o n a l r a t h e r t h a n Dry E v e r g r e e n F o r m a t i o n s . A v a r i a n t o f t h e i r c a t e g o r y o f Dry L i m e s t o n e S c r u b F o r e s t i s an A r i d L i m e s t o n e S c r u b F o r e s t on d i s s e c t e d c o a s t a l l i m e s t o n e ( 1 9 5 3 , 3 8 3 - 3 8 4 ) , w h i c h a 1 s o h a s C a p a n c o u i ? t e r p a r t s .
1. Dry E v e r g r e e n F o r e s t
B e a r d ( 1 9 4 9 , 8 0 ) d i d n o t o r i g i n a l l y d e s c r i b e t h i s t y p e i n d e t a i l b e c a u s e " t h e b u l k o f t h e woodland which a p p e a r s t o b e l o n g t o t h i s t y p e i s v e r y h e a v i l y damaged and i t i s n o t p o s s i b l e t o form a n y r e l i a b l e c o n c e p t i o n o f t h e o r i g i n a l s t r u c t u r e " ; h e a s s o c i a t e d i t w i t h mean a n n u a l r a i n f a l l s o f 1270-2000 mm, h i g h e r t h a n t h o s e o f L i t t l e Cayman.
T a l l woodland i n t h e i n t e r i o r o f L i t t l e Cayman, s o u t h o f Sparrowhawk H i l l ( P l a t e 3 7 ) , m e e t s B e a r d ' s ( 1 9 5 5 , 92) d i a g n o s i s o f "crowded, s l e n d e r t r e e s r e a c h i n g 1 2 t o 20 m e t r e s i n h e i g h t above which a n o c c a s i o n a l
e m e r g e n t r e a c h e s 30 m e t r e s . L e a v e s a r e s i m p l e and e v e r g r e e n , t h e m a j o r i t y b e i n g s t i f f a n d f l e s h y , a h i g h p r o p o r t i o n w i t h l a t e x o r
e s s e n t i a l o i l . B a r k - s h e d d i n g may b e a c o n s p i c u o u s f e a t u r e . The s h r u b and h e r b l a y e r s a r e p o o r l y r e p r e s e n t e d . " The f o r e s t o c c u r s O i l a f l a t - l y i n g l i m e s t o n e a r e a a b o u t 4-5 m above s e a - l e v e l . The g r o u n d s u r f a c e i s u n d i s s e c t e d , and h a s a c o n s i d e r a b l e s o i l and l i t t e r c o v e r . The d o m i n a n t b i g t r e e s a r e B u m e l i a s a l i c i f o l i a , C a l y p t r a n t h e s p a l l e n s a n d C h i o n a n t h u s s p . , f o r m i n g a canopy a t 8-10 m . I n s p i t e o f t h e i r h e i g h t , m o s t o f t h e t r e e s a r e s l e n d e r a n d c l o s e l y - s p a c e d . A number o f s h r u b b y
s p e c i e s a r e p r e s e n t , b u t t h e f o r e s t i s r e l a t i v e l y open and e a s y t o walk t h r o u g h . The f o l l o w i n g s p e c i e s were
T r e e s : B u m e l i a s a l i c i f o l i a B u r s e r a s i m a r u b a C a l y p t r a n t h e s p a l l e n s Canel 1 a w i n t e r a n a C h i o n a n t h u s s p . S h r u b s : A l l o p h y l u s c o m i n i a
A m y r i s e l e m i f e r a A r q y t h a m n i a p r o c t o r i i C h i o c o c c a a l b a
C o l u b r i n a e l l i p t i c a H e r b s : L a s i a c i s d i v a r i c a t a
n o t e d :
C r o t o n n i t e n s
E r y t h r o x y l um r o t u n d i f o l i u m H y p e l a t e t r i f o l i a t a
M y r c i a n thes f r a q r a n s T a b e b u i a h e t e r o p h y l l a C r o t o n l u c i d u s
J a c q u i n i a b e r t e r i i Ma1 v a v i s c u s a r b o r e u s Ma y t e n u s b u x i f o l i a R a n d i a a c u l e a t a Paspalurn b l o d q e t t i i
S u c c u l e n t : A g a v e s o b o l i f e r a
E p i p h y t e s : S c h o m b u r g k i a b r y s i a n a T i l l a n d s i a s p p .
The A g a v e i s r a r e , and c a c t i a r e a p p a r e n t l y a b s e n t , a s a r e f e r n s . The l i m i t s o f t h e f o r e s t a r e o n j e c t u r a l , b u t p r o b a b l y c o i n c i d e w i t h t h e 20 f t c o n t o u r s o u t h o f Sparrowhawk H i l l on t h e 1 : 2 5 , 0 0 0 map. Northwards t h e f o r e s t e n d s i n a l i m e s t o n e r i d g e c o v e r e d w i t h Dry E v e r g r e e n T h i c k e t ; t o t h e s o u t h and e a s t it p a s s e s i n t o d i s s e c t e d l o w - l y i n g l i m e s t o n e
t e r r a i n w i t h C o n o c a r p u s and L a g u n c u l a r i a s c r u b . 2. Dry E v e r g r e e n Woodland
T h i s i s d e s c r i b e d by Beard (1955) a s a t w o - s t o r e y woodland o f d e n s e l y p a c k e d a t t e n u a t e d t r e e s f o n n i n g a canopy a t 6-12 m , w i t h e m e r g e n t s r e a c h i n g 20 m; i t i s n o t v e r y c l e a r l y d i s t i n g u i s h e d from Dry E v e r g r e e n F o r e s t . I n 1955 Beard e q u u t e d i t w i t h L i t t o r a l Woodland, b u t i t i s more u s e f u l h e r e t o m a i n t a i n t h e l a t t e r a s a s e p a r a t e
component o f c o a s t a l v e g e t a t i o n ,
Dry E v e r g r e e n Woodland i s used h e r e t o d e n o t e a more open and lower woodland t h a n Dry E v e r g r e e n F o r e s t , c h a r a c t e r i s t i c a l l y found on g e n t l y s l o p i n g t e r r a i n a s s o c i a t e d w i t h t h e Marl F a c i e s o f t h e I r o n s h o r e
F o r m a t i o n ( P l a t e 3 9 ) . Much o f t h e s u r f a c e i s b a r e r o c k , b u t i t i s r e l a t i v e l y u n d i s s e c t e d , and where t h e Marl Formation g i v e s way t o d i s s e c t e d B l u f f Limestone t h e r e i s a n a b r u p t t r a n s i t i o n from Dry E v e r g r e e n Woodland t o Dry E v e r g r e e n T h i c k e t . The woodland i s a l s o found i n more p r o t e c t e d s i t u a t i o n s , s u c h a s n o r t h w e s t , w e s t and s o u t h - f a c i n g s l o p e s . The woodland i s found on t e r r a i n r i s i n g t o 6 . 5 m above s e a - l e v e l a t P a r a d i s e E n d ( 0 4 5 7 8 1 ) , t o 1 2 . 5 m i n l a n d from J a c k s o n ' s Bay
( 9 7 7 7 6 7 ) , and t o 1 3 m i n l a n d from S a l t Rocks ( 9 4 4 7 4 9 ) . The most c o n s p i c u o u s t r e e s p e c i e s i s B u r s e r a s i m a r u b a . C h a r a c t e r i s t i c s p e c i e s o f Dry E v e r g r e e n T h i c k e t , s u c h a s P l u m e r i a o b t u s a , C e p h a l o c e r e u s and A g a v e , a r e i n c o n s p i c u o u s i n t h e woodland. I n l a n d from S a l t Rocks, t h e woodland e x t e n d s down t o t h e c o a s t ( i n most a r e a s i t i s s e p a r a t e d from t h e c o a s t b y a zone o f l a g o o n , swamp and b e a c h r i d g e ) , and t h e f o l l o w i n g s p e c i e s were n o t e d :
T r e e s : B u r s e r a s i m a r u b a C a n e l l a w i n t e r a n a
C i t h a r e x y l u m f r u t i c o s u m C o c c o l o b a u v i f e r a
C o r d i a g e r a s c a n t h u s C o r d i a sebes t e n a C r o t o n n i t e n s S h r u b s : A d e l i a r i c i n e l l a
A r g y t h a m n i a p r o c t o r i A t e r a m n u s l u c i d u s C a l y p t r a n t h e s p a l l e n s C a p p a r i s f l e x u o s a
C o c c o t h r i n a x jamai c e n s i s C o l u b r i n a e l l i p t i c a
F i c u s a u r e a G u a p i r a d i s c o l o r M e t o p i um t o x i fe r u m P i c r o d e n d r o n b a c c a t u m T a b e b u i a h e t e r o p h y l l a T r i chi 1 i a g l a b r a S w i e t e n i a m a h a g o n i J a c q u i n i a b e r t e r i i
M a l v a v i s c u s a r b o r e u s v a r
.
c u b e n s i s
Ma y t e n u s b u x i f o l i a
P h y l l a n t h u s a n g u s t i f o l i u s S a v i a e r y t h r o x y l o i d e s
V i n e s : C a p p a r i s f l e x u o s a Ipomoea acumina t a
Ipomoea ma c r a n t h a
G r a s s : L a s i a c i s d i v a r i c a t a
Many o f t h e t r e e s p e c i e s a r e i d e n t i c a l w i t h t h o s e o f t h e Dry E v e r g r e e n F o r e s t ; some, n o t a b l y C o c c o l o b a u v i f e r a and C o r d i a s e b e s t e n a , a r e p r e s e n t o n l y b e c a u s e o f t h e c o a s t a l s i t u a t i o n o f t h i s l o c a l i t y and a r e n o t p r e s e n t e l s e w h e r e i n Dry E v e r g r e e n Woodland.
3. Dry E v e r g r e e n T h i c k e t
T h i s v e g e t a t i o n t y p e , one o f t h e most common on B l u f f Limestone o f L i t t l e Cayman, d i d n o t a p p e a r i n B e a r d ' s e a r l i e r c l a s s i f i c a t i o n s and was f i r s t r e c o g n i s e d by him i n 1955; i n t h a t p a p e r , however, h e d i d e q u a t e i t w i t h h i s e a r l i e r c a t e g o r i e s ( 1 9 4 4 ) o f L i t t o r a l T h i c k e t and L i t ~ o r a l Hedge, though i t i s more c o n v e n i e n t i n t h e c a s e o f L i t t l e Cayman t o t r e a t t h e s e l a t t e r s e p a r a t e l y a s component o f c o a s t a l v e g e t a t i o n . Beard d e s c r i b e d Dry E v e r g r e e n T h i c k e t a s a t w o - s t o r e y t h i c k e c " w l t n a low, open, o r dense canopy and a d e n s e o r s p a r s e
undergrowth". The canopy i s a t 6-12 m , w i t h o c c a s i o n a l e m e r g e n t s , and t h e t r e e s a r e s l e n d e r . Dry E v e r g r e e n T h i c k e t h a s been f u r t h e r
d e s c r i b e d from s o u t h J a m a i c a , w i t h a canopy a t 5-11 m and emergents t o 1 6 m ( L o v e l e s s and Asprey 1957, 813-815), and from A n t i g u a , where i t c o m p r i s e s " a d e n s e s h r u b l a y e r a b o u t 2-4 m (7-13 f t ) h i g h , o v e r t o p p e d by s c a t t e r e d b u t more o r l e s s u n i f o r m l y d i s t r i b u t e d t r e e s r e a c h i n g a h e i g h t o f 7-8 m (23-26 f t )
" ,
w i t h A g a v e , O p u n t i a and C e p h a l o c e r e u s ( L o v e l e s s 1 9 6 0 , 5 1 1 - 5 1 3 ) . I n A n t i g u a i t o c c u r s i n a r e a s w i t h a mean a n n u a l r a i n f a l l o f l e s s t h a n 900 mm.Dry E v e r g r e e n T h i c k e t a s p r e v i o u s l y r e c o g n i s e d i s t h u s a r a t h e r v a r i a b l e e n t i t y , and on L i t t l e Cayman i t a l s o v a r i e s i n h e i g h t , d e n s i t y and c o m p o s i t i o n , a t l e a s t p a r t l y i n r e s p o n s e t o a s p e c t and e x p o s u r e
( P l a t e 3 8 ) .
( a ) Seaward s l o p e s o f B l u f f Limestone r i d g e s
Along much o f t h e n o r t h - f a c i n g c o a s t o f L i t t l e Cayman t h e r e i s a f l a t t e r r a c e o r g e n t l y s l o p e o f d i s s e c t e d B l u f f Limestone 150-200 m wide and r i s i n g up t o 5 m above s e a - l e v e l , s i t u a t e d between t h e c o a s t a l swamp and t h e B l u f f Limestone r i d g e , c o v e r e d w i t h open Dry E v e r g r e e n T h i c k e t . T r e e s a r e o n l y 3-4 m t a l l , and a s i n t h e c a s e o f B u r s e r a s i m a r u b a may b e s e v e r e l y deformed above t h i s l e v e l . T h r e e o t h e r v e r y c h a r a c t e r i s t i c t a l l e r s p e c i e s a r e P l u m e r i a o b t u s a , C e p h a l o c e r e u s
s w a r t z i i and Agave s o b o l i f e r a ( P l a t e 4 0 ) . G u a p i r a d i s c o l o r i s a v e r y common t r e e , and S a v i a e r y t h r o x y l o i d e s t h e most a b u n d a n t s h r u b . T h i s open t h i c k e t i s v e r y s i m i l a r t o t h a t d e s c r i b e d from t h e n o r t h e r n L e s s e r A n t i l l e s , e . g . Barbuda, e x c e p t t h a t t h e s p h e r i c a l c a c t i M e l o c a c t u s i n t o r t u s a n d Manunillaria n i v o s a a r e a b s e n t from t h e Cayman I s l a n d s .
The f o l l o w i n g s p e c i e s were n o t e d i n t h i s h a b i t a t on t h e n o r t h e r n s i d e o f Sparrowhawk H i l l (003781) :
T r e e s :
S h r u b s :
S u c c u l e n t s :
G u a p i r a d i s c o l o r G u e t t a r d a e l l i p t i c a M y r c i a n t h e s f r a g r a n s A m y r i s e l e m i f e r a B a u h i n i a d i v a r i c a t a C a l y p t r a n t h e s p a l l e n s C r o ton l i n e a r i s
Croton l u c i d u s E r i t h a l i s f r u t i c o s a A g a v e s o b o l i f e r a
P l u m e r i a ob t u s a
T a b e b u i a h e t e r o p h y l l a
E v o l v u l us. a r b u s c u l a J a c q u i n i a b e r t e r i i P h y l l a n t h u s n u t a n s R a n d i a a c u l e a t a
S a v i a e r y t h r o x y l o i d e s S t r u m p f i a m a r i t i m a C e p h a l o c e r e u s s w a r t z i i ( b ) D i s s e c t e d B l u f f Limestone r i d g e
The B l u f f L i a e s t o n e r i d g e i s a l i n e a r L'eatvre, p a r a l l e l t o t h e n o r t h c o a s t , r e z c h i n g i t s g r e a t e s t e l e v a t i o n s o f 6-13 x z b o u t 2 0 0 - 4 0 0 n i n l a n d . I t i s d e e p l y d i s s e c t e d , w i t h many d e p r e s s i o n s more t h a n 2 m d e e p , and w i t h c l i f f f a c e s o f t h e same a m p l i t u d e . The t h i c k e t i s more d e n s e t h a n on he seaward s l o p e s , and t h e r e a r e more t r e e s p e c i e s
i d e n t i c a l w i t h t h o s e o f Dry Evergreen F o r e s t and Dry E v e r g r e e n Woodland.
The l i t t o r a l s h r u b S t r u m p f i a m a r i t i m a d i s a p p e a r s , b u t B u r s e r a , P l u m e r i a
,
C e p h a l o c e r e u s and A g a v e remain c h a r a c t e r i s t i c . The f o l l o w i n g s p e c i e s were n o t e d a t Sparrowhawk H i l l ( 0 0 4 7 8 0 ) :
T r e e s :
S h r u b s :
S u c c u l e n t s : E p i p h y t e s :
B u r s e r a s i m a r u b a C a n e l l a w i n t e r a n a C o r d i a g e r a s c a n t h u s C o r d i a sebes t e n a G u a p i r a d i s c o l o r G u e t t a r d a e l l i p t i c a B a u h i n i a d i v a r i c a t a C a l y p t r a n thes p a l 1 ens E r i t h a l i s f r u t i c o s a J a c q u i n i a b e r t e r i i A g a v e s o b o l i f e r a S c h o m b u r g k i a b r y s i a n a
H y p e l a t e t r i f o l i a t a M y r c i a n t h e s f r a g r a n s P l u m e r i a ob t u s a
S c h o e p f i a c h r y s o p h y l l o i d e s T a b e b u i a h e t e r o p h y l l a
P h y l l a n t h u s a n g u s t i f o l i u s S a v i a e r y t h r o x y l o i d e s T o u r n e f o r t i a vol u b i l i s
C e p h a l o c e r e u s s w a r t z i i T i l l a n d s i a s p p .
( c ) I n l a n d s l o p e s o f B l u f f Limestone r i d g e
The i n l a n d s l o p e s o f t h e B l u f f Limestone r i d g e a r e o f v a r i a b l e s t e e p n e s s and h e i g h t , and i n p l a c e s c l i f f y . T h e i r main c h a r a c t e r i s t i c , however, i s t h a t t h e y a r e much more p r o t e c t e d t h a n s e a w a r d - f a c i n g
s l o p e s , and t h i s r e s u l t s i n t a l l e r , d e n s e r t h i c k e t w i t h many more e p i p h y t e s . The v e g e t a t i o n o f t h i s h a b i t a t a t Sparrowhawk H i l l i s a d j a c e n t t o t h e Dry E v e r g r e e n F o r e s t o f t h e c e n t r e o f t h e i s l a n d and i t i n c l u d e s t a l l t r e e s o f s p e c i e s common t o t h a t woodland ( B u m e l i a , A m y r i s , C h i o n a n t h u s )
.
S p e c i e s p r e s e n t i n c l u d e :5 8
T r e e s : A1 1 o p h y l u s c o m i n i a Bumel i a g l o m e r a t a A m y r i s e l e m i f e r a B u m e l i a s a l i c i f o l i a A n t i r h e a 1 u c i d a C h i o n a n t h u s s p
.
F i c u s a u r e a P l u m e r i a o b t u s a
G u a p i r a d i s c o 1 or T a b e b u i a h e t e r o p h y l l a Gue t t a r d a e l l i p t i c a Z u e l a n i a q u i d o n i a M y r c i a n t h e s f r a g r a n s
S h r u b s : B a u h i n i a d i v a r i c a t a G y m i n d a l a t i f o l i a B o u r r e r i a v e n o s a J a c q u i n i a b e r t e r i i C o l u b r i n a a s i a t i c a P h y l l a n t h u s n u t a n s C r o t o n l i n e a r i s R a n d i a a c u l e a t a
Herbs: P a s p a l um b l o d g e t t i i S a l v i a o c c i d e n t a l i s S u c c u l e n t s : A y a v e s o b o l i f e r a ( r a r e )
E p i p h y t e s : ,Schonrburgk.la b r y s i a n a T i l l a n d s i a spp ,
4 . Evergreen Bushland
T h i s c a t e g o r y was i n t r o d u c e d by Beard i n 1949 ( 8 2 - 8 0 ) , t a k i n g t h e v e g e t a t i o n o f Barbuda a s a t y p e example. Beard q u o t e d Warming's (1909) d e s c r i p t i o n o f " a g r e y , d e s o l a t e , u s e l e s s and s c o r c h i n g b u s h l a n d between whose t h o r n y t a n g l e d s h r u b s and low t r e e s one c a n n o t p e n e t r a t e w i t h o u t
t h e u s e o f an a x e " ( a d e s c r i p t i o n which a l s o a p p l i e s t o much o f t h e Dry E v e r g r e e n T h i c k e t ) . E v e r g r e e n Bushland forms a low woodland, 3-9 m
t a l l , c o n s i s t i n g o f a " d e n s e i m p e n e t r a b l e growth o f g n a r l e d , l i t t l e t r e e s and b u s h e s w i t h h a r d e v e r g r e e n l e a v e s " . On Barbuda t h e t r e e s i n c l u d e T a b e b u i a p a l l i d a , B u c e r a s b u c i d a and G u e t t a r d a s c a b r a
,
w i t h b r o m e l i a d s ( t h o u g h no mosses o r f e r n s ) , A g a v e and c a c t i . D'Arcy (1975) a s s i g n s t h e v e g e t a t i o n of Anegada to t h i s c a t e g o r y , though i t h a s been h e a v i l y d i s t u r b e d by g r a z i n g ; a c c o r d i n g t o Beard (1949) t h e sub-climaxes r e s u l t i n g from d i s t u r b a n c e o f Evergreen Bushland a r e v a r i o u s t h o r n y t h i c k e t s . E v e r g r e e n Bushland i s a l s o d e s c r i b e d by L o v e l e s s and Asprey( 1 9 5 7 , 809-813) from s o u t h Jamaica, where i t h a s a uniform h e i g h t o f 4 m , w i t h emergents ( m a i n l y B u r s e r a ) r e a c h i n g 7-8 m , and by Loveless (1960) from A n t i g u a .
Evergreen Bushland c o v e r s much o f t h e w e s t e r n end o f L i t t l e Cayman, on u n d u l a t i n g b u t g e n e r a l l y low and m o d e r a t e l y d i s s e c t e d t e r r a i n ; v e r y low and h e a v i l y d i s s e c t e d l i m e s t o n e i n c o n t r a s t i s covered w i t h
C o n o c a r p u s s c r u b . I t i s w e l l s e e n a l o n g t h e t r a n s - i s l a n d t r a c k n o r t h o f Blossom V i l l a g e , where t h e l a n d l i e s a t 2-3 m above s e a - l e v e l . T h i s a r e a h a s c e r t a i n l y been much m o d i f i e d by g r a z i n g and c u l t i v a t i o n , b u t i t i s n o t p o s s i b l e t o e s t i m a t e t h e e f f e c t o f t h i s on t h e v e g e t a t i o n .
P l u m e r i a , A g a v e and B u r s e r a a r e p r e s e n t b u t much l e s s conspicuous t h a n e l sewhere. S h r u b s i n c l u d e J a c q u e m o n t i a b e r t e r i i
,
E v o l v u l u s a r b u s c u l a and S t r u m p f i a m a r i t i m a , t o g e t h e r w i t h many weedy s p e c i e s . Clumps o f t h e l e a t h e r f e r n A c r o s t i c h u m a u r e u m n e s t l e i n s o l u t i o n h o l e s ( P l a t e 41) i n t h e l i m e s t o n e i n e x a c t l y t h e same way a s Fosberg (1971) h a s d e s c r i b e d f o r Aldabra A t o l l ; u s u a l l y t h i s f e r n (which i s t h e o n l y one r e c o r d e d f o r L i t t l e Cayman) i s a s s o c i a t e d w i t h s e a - l e v e l mangrove swamps.B . C o a s t a l v e g e t a t i o n
I n a d d i t i o n t o swamp v e g e t a t i o n , mainly mangrove, i n c o a s t a l
s i t u a t i o n s , Beard (1949, 84-87) d i s t i n g u i s h e d a s one o f h i s Dry Evergreen F o r m a t i o n s t h e c a t e g o r y o f L i t t o r a l Woodland a s " a s e r i e s o f c l o s e l y a s s o c i a t e d f o r m a t i o n s " , i n c l u d i n g L i t t o r a l Hedge, w i t h s e v e r a l sub- c l i m a x e s ( s a n d dune, s a l t f l a t s , rock pavements, rocky s l o p e s )
.
S u b s e q u e n t l y he e q u a t e d (1955) L i t t o r a l Woodland w i t h Dry Evergreen Woodland, and L i t t o r a l T h i c k e t and L i t t o r a l Hedge w i t h Dry Evergreen T h i c k e t . Asprey and L o v e l e s s (1958) i n n o r t h Jamaica s e p a r a t e d
c o a s t a l rock from c o a s t a l sand communities, w i t h i n t h e g e n e r a l c a t e g o r y o f L i t t o r a l Woodland. T h i s d i v i s i o n was b r o a d l y f o l l o w e d by L o v e l e s s
(1960)
,
i n A n t i g u a , who, however, c o n s i d e r e d s t r a n d communities t o be e d a p h i c c l i m a x e s r a t h e r t h a n s e r a 1 s t a g e s o f t h e k i n d e n v i s a g e d by Davis (1942),
Asprey and Robbins (1953) and Beard (1949) , The Asprey and L o v e l e s s scheme i n c l u d e d :On sandy s u b s t r a t e s : Herbaceous p i o n e e r S t r a n d s c r u b
S t r a n d woodland On r o c k y s u b s t r a t e s :
L i t t o r a l rock pavement L i t t o r a l hedge
L i t t o r a l e v e r g r e e n bushland L i t t o r a l palm t h i c k e t
The d i s t i n c t i o n between L i t t o r a l Hedge and S t r a n d Scrub i s n o t a c l e a r one on L i t t l e Cayman; and a s i n g l e c a t e g o r y f o r L i t t o r a l Rock Pavement v e g e t a t i o n , though a d o p t e d by Howard (1950) i n v a r i a b l e t e r r a i n , c a n n o t p r o p e r l y encompass t h e t o p o g r a p h i c and v e g e t a t i o n a l v a r i a t i o n found i n
rocky c o a s t a l s i t u a t i o n s . Here we u s e a p u r e l y e m p i r i c a l scheme s p e c i f i c t o L i t t l e Cayman:
S t r a n d community on sandy beach S t r a n d community on cobble beach C o a s t a l s a n d f l a t s
Rock pavement community S t e e p r o c k y c o a s t community 1. S t r a n d communitv on sandv beach
Beaches on L i t t l e Cayman a r e v a r i a b l e i n w i d t h and h e i g h t , b u t most a r e p r o t e c t e d by o f f s h o r e r e e f and a r e low and narrow. O f f s h o r e
beachrock i n d i c a t e s l i m i t e d beach r e t r e a t i n r e c e n t y e a r s round much o f t h e c o a s t . Ten sand beach a r e a s were s t u d i e d ( ~ i g u r e 3 ) , and t h e f o l l o w i n g s p e c i e s were r e p r e s e n t e d :
6 0
T r e e s :
S h r u b s :
Herbs :
A v i c e n n i a u e r m i n a n s L a g u n c u l a r i a r a c e m o s a C a s u a r i n a e q u i s e t i f o l i a M o r i n d a c i t r i f o l i a Coccol o b a u v i f e r a R h i z o p h o r a m a n g l e C o c o s n u c i f e r a T e r m i n a l i a c a t a p p a C o r d i a s e b e s t e n a T h r i n a x r a d i a t a B o r r i c h i a a r b o r e s c e n s S c a e v o l a p l u m i e r i C a e s a l p i n a s p . S u r i a n a m a r i t i m a
C o l u b r i n a a s i a t i c a M a l l o t o n i a g n a p h a l o d e s C o n o c a r p u s e r e c t a
A m b r o s i a h i s p i d a I p o m o e a m a c r a n t h a C a k i l e l a n c e o l a t a I p o m o e a p e s - c a p r a e
C a n a v a l i a r o s e a P h i l o x e r u s v e r m i c u l a r i s C a s s y t h a f i l i f o r m i s S a l i c o r n i a b i g e l o v i i E u p h o r h i a m e s e m b r i a n t h e m i fol i a
H y m e n o c a l l i s l a t i f o l i a S e s u v i um p o r t u l a c a s t r u m G r a s s e s and C e n c h r u s e c h i n a t u s
s e d g e s : C y p e r u s s p p .
S p a r t i n a pa tens
S p o r o b o l u s v i r g i n i c u s Where t h e n a t u r a l v e g e t a t i o n h a s n o t Seen c l e a r e d , t h e beach c r e s t i s t y p i c a l l y o c c u p i e d by a ' L i t t o r a l H e d g e ' , which a s d e f i n e d by Beard
(1949, 84) " c o n s i s t s o f a d e n s e m a t t e d , and i n t e r l a c e d , woody growth o f g n a r l e d b u s h e s u s u a l l y o f d i s t o r t e d form. T h e i r h e i g h t i s v a r i a b l e , from a few i n c h e s t o s e v e r a l f e e t , depending upon t h e f o r c e o f t h e wind a n d a v a i l a b l e s o i l d a p t h " . A s used b y Asprey and L o v e l e s s (1958) i n n o r t h J a m a i c a , t h e term i s c o n f i n e d t o rocky s h o r e s , where t h e hedge c o m p r i s e s many s p e c i e s , i s 60 cm t o 2 m t a l l , i s wind-sheared i n p r o f i l e , and c u t i n t o windrows i n p l a n . ' L i t t o r a l h e d g e ' , however, d e s c r i b e s p e r f e c t l y t h e a p p e a r a n c e o f t h e zone o f C o c c o l o b a u v i f e r a a t t h e beach c r e s t , r e a c h i n g 5 m i n h e i g h t S u t u s u a l l y l e s s t h a n 50 m wide.
I t i s wind-sheared i n p r o f i l e b u t n o t c u t i n t o windrows ( P l a t e 4 2 ) . I t forms a d e n s e and i m p e n e t r a b l e s c r u b , w i t h t h e ground s u r f a c e d e v o i d o f o t h e r p l a n t s b u t c o v e r e d w i t h d r y i n g C o c c o l o b a l e a v e s . I t
c o r r e s p o n d s t o and i n a p p e a r a n c e c l o s e l y r e s e m b l e s t h e S c a e v o l a t a c c a d a hedge d e s c r i b e d from many I n d o - P a c i f i c a t o l l s ( e . g . S t o d d a r t 1971, 1 3 5 ;
1975, 98)
.
T r e e s o f T e r m i n a l i a c a t a p p a , C o r d i a s e b e s t e n a and Cocos a r e o c c a s i o n a l l y found i n t h e C o c c n l o b a , b u t t h e hedge i s e s s e n t i a l l y m o n o s p e c i f i c , i n c o n t r a s t t o t h e a r r a y o f s p e c i e s o r i g i n a l l y d e s c r i b e dfrom t h i s community b y Beard ( 1 9 4 9 , 85) ; some o f B e a r d ' s s p e c i e s ( C h r y s o b a l a n u s i c a c o , E r i t h a l i s f r u t i c o s a ) a r e p r e s e n t i n L i t t l e Cayman, b u t n o t a s a component o f t h e l i t t o r a l hedge.
Between t h e C o c c o l o b a hedge and t h e s e a t h e r e may b e a s h r u b zone o f v a r i a b l e w i d t h , dominated b y S u r i a n a m a r i t i m a ; t h i s may b e o n l y t h e w i d t h o f a s i n g l e s h r u b , o r i n p r o t e c t e d s i t u a t i o n s ( a s a t West P o i n t )
it forms a s c r u b 2 m t a l l and 50-100 m wide. T o u r n e f o r t i a ( M a l l o t o n i a ) and S c a e v o l a p l u m i e r i a r e o c c a s i o n a l l y p r e s e n t a s s c a t t e r e d s h r u b s
1-2 m t a l l , b u t S o p h o r a t o m e n t o s a h a s y e t t o b e r e c o r d e d on t h e i s l a n d , t h o u g h p r e s e n t on Grand Cayman.
Where u n d i s t u r b e d , t h e C o c c o l o b a woodland and S u r i a n a s c r u b r e a c h down t o t h e swash l i n e , and p i o n e e r h e r b a c e o u s s p e c i e s a r e a b s e n t . A t Sandy P o i n t i n t h e s o u t h e a s t ( 0 8 6 7 8 5 ) , however, t h e beach i s a g g r a d i n g and t h e r e i s a more open mosaic o f s h r u b s ( S c a e v o l a , T o u r n e f o r t i a , S u r i a n a , C a e s a l p i n i a ) , o c c a s i o n a l C o r d i a , and a p a t c h y ground cover of p i o n e e r C a k i l e l a n c e o l a t a , S e s u v i um p o r t u l a c a s t r u m and S p o r o b o l u s v i r q i n i c u s , w i t h , i n l a n d , Ipomoea m a c r a n t h a , E u p h o r b i a
m e s e m b r i a n t h e m i f o l i a , H y m e n o c a l l i s l i to r e a , and C e n c h r u s ( P l a t e 4 3 )
.
Where t h e C o c c o l o b a hedge has been c l e a r e d , a s i m i l a r d i v e r s e mosaic o f s h r u b s , h e r b s and g r a s s e s i s developed. Cornrnon s p e c i e s a r e S e s u v i u m , Ipomoea p e s - c a p r a e , S p a r t i n a p a t e n s and S p o r o b o l u s a t t h e beach c r e s t , a n d , on t h e berm, S e s u v i u m , P h i l o x e r u s , A m b r o s i a , Ipomoea m a c r a n t h a , C e n c h r u s , H y m e n o c a l l i s , and t h e f o l l o w i n g s h r u b s : S u r i a n a , S c a e v o l a , T o u r n e f o r t i a , B o r r i c h i a a r b o r e s c e n s , C o n o c a r p u s e r e c t a , and low
C o c c o l o b a . T h ~ r e a r e y o u n g t r e e s o f C a s u a r i n a e y u i s e t i f o l i a a l o n g t h e b u a c h
a t
Wearis Bay, b u t no mature g r o v e s o f t h e s e t r e e s on L i t t l e Caymn, i?c o n t r a s t t o Grand Cayman where i t forms a d i s t i n c t i v e though r e s t r i c t e d c o a s t a l wood1 and.
Notable v a r i a n t s o f t h e above p a t t e r n s o c c u r a t two main l o c a l i t i e s , round Bloody Bay on t h e n o r t h c o a s t , and b o t h e a s t and w e s t o f D i g g a r y ' s P o i n t on t h e s o u t h , a s w e l l a s o c c a s i o n a l l y e l s e w h e r e . Here, i n
p r o t e c t e d s i t u a t i o n s , t h e c o a s t i s formed of a l t e r n a t i n g s m a l l low p r o j e c t i o n s , c o n s i s t i n g o f r u b b l e o r r o c k y o u t c r o p s , and i n t e r v e n i n g b a y s w i t h narrow low sandy b e a c h e s . The p r o m o n t o r i e s s u p p o r t s m a l l p a t c h e s o f s c r u b o f C o n o c a r p u s and L a q u n c u l a r i a , w i t h o c c a s i o n a l A v i c e n n i a and R h i z o p h o r a , and a h e r b mat o f S e s u v i u m , P h i l o x e r u s and S a l i c o r n i a . The sandy i n t e r v e n i n g bays have an i r r e g u l a r s c r u b o f B o r r i c h i a , C o n o c a r p u s and S u r i a n a , w i t h S e s u v i u m and S p a r t i n a p a t e n s . S i m i l a r v e g e t a t i o n i s found i n t h e e a s t e r n p a r t o f S o u t h Hole Sound where t h e c o a s t i s n o t f r i n g e d by mangroves.
2. S t r a n d community on c o b b l e beach
T h i s i s d i f f i c u l t i n many p l a c e s t o d i s t i n g u i s h from t h e sand beach community, because i n many p l a c e s a modern sand beach f r o n t s an o l d e r c o b b l e and r u b b l e r i d g e , s o t h a t f r e q u e n t l y t h e r e i s a z o n a t i o n o f
sandy communities c l o s e s t t o t h e s e a and o f c o b b l e s a few m e t r e s i n l a n d . S u r i a n a m a r i t i m a i s o f t e n a s s o c i a t e d w i t h t h e sandy s t r i p s o r p o c k e t b e a c h e s , and C o c c o l o b a w i t h t h e cobble r i d g e s . G r e a t e s t d i v e r s i t y i s found on t h e sand p a t c h e s , which c a r r y A m b r o s i a , I p o m o e a , S e s u v i u m ,
H y m e n o c a l l i s and B o r r i c h i a . A s s o c i a t e d w i t h C o c c o l o b a on c o b b l e s a r e M o r i n d a , T h e s p e s i a and H y m e n o c a l l i s . Where t h e s c r u b h a s been c l e a r e d from t h i s rough t e r r a i n , t h e v e g e t a t i o n u s u a l l y c o n s i s t s o f t h i c k
d e c e p t i v e mats o f S e s u v i u m , A m b r o s i a and C a s s y t h a
.
3. C o a s t a l sand f l a t s
These form a l a t e r a l l y c o n t i n u o u s zone ( e x c e p t f o r t h e B l u f f Limestone o u t c r o p a t E a s t End and t h e S a l t Rocks I r o n s h o r e a t t h e w e s t end) round t h e e n t i r e i s l a n d . The s u r f a c e s l o p e s g r a d u a l l y i n l a n d from t h e b e a c h - r i d g e c r e s t , and may be i r r e g u l a r b e c a u s e o f t h e p r e s e n c e o f e i t h e r blown sand o r r u b b l e l o b e s . The f l a t s a r e u s u a l l y 50-100 m wide,
and n e v e r more t h a n 200 m , and t h e y a r e u s u a l l y l e s s t h a n 2.5 m above s e a - l e v e l . E x c e p t f o r a s e c t i o n o f t a l l woodland a t S p o t Bay ( 9 5 0 7 5 7 ) , a l l t h e c o a s t a l f l a t s h a v e b e e n c l e a r e d a t some s t a g e and a r e now
o c c u p i e d by s e c o n d a r y s c r u b , c o c o n u t p l a n t a t i o n , o r newly c l e a r e d g r o u n d . The S p o t Bay woodland i s p r o b a b l y t h e l a s t s u r v i v i n g r e m n a n t o f L i t t o r a l Woodland on any o f t h e t h r e e Cayman I s l a n d s , a n d i s t h u s o f c o n s i d e r a b l e i n t e r e s t . I t i n c l u d e s t a l l t r e e s o f S w i e t e n i a mahagoni ( r e a c h i n g 1 5 m )
,
and B u r s e r a s i m a r u b a , w i t h D a l b e r g i a e c a s t a p h y l l u m , C o r d i a s e b e s t e n a , T e r m i n a l i a , C o c c o l o b a u v i f e r a , and J a c q u i n i a b e r t e r i i . C o c c o l o b a f o r m s a h e d g e a l o n g t h e seaward s i d e .
The s e c o n d a r y s c r u b c o n s i s t s o f s t r a n d s h r u b s a n d t r e e s ( S u r i a n a , T o u r n e f o r t i a , C o c c o l o b a , C o r d i a , C h r y s o b a l a n u s , C o n o c a r p u s ) , t o g e t h e r w i t h a b u n d a n t L a n t a n a , C o l u b r i n a a s i a t i c a , G o s s y p i u m , C a e s a l p i n i a , and
some C o m o c l a d i a d e n t a t a
.
A t J a c k s o n'
s Bay (976771) C o n o c a r p u s r e a c h e sd h e l g h t o f 5 m , S u r i a n a and C o c c o l o b a 3-4 m , and C a e s a l p i n j a 2 m. In p l a c e s t h i s s e c o n d a r y s c r u b i s e x t r e m e l y d e n s e . E l s e w h e r e i t i s
i n t e r r u p t e d b y more open g r o u n d , where s t r a n d h e r b s and g r a s s e s ( E u p h o r b i a m e s e m b r i a n t h e m i f o l i a , Tpormea p e s - c a p r a e , C y p e r u s s p p . , A m b r o s i a , Ernodea l i t t o r a l i s , C a s s y t h a , C a n a v a l i a , S p o r o b o l u s ) a r e
j o i n e d by weedy s o e c i e s s u c h a s B r y o p h y l l u m , A b r u s , P o r t u l a c a and S t a c h y t a r p h e t a . R h y n c h e l y t r u m r e p e n s i s c o n s p i c u o u s a l o n g r o a d s i d e s , e s p e c i a l l y i n t h e n o r t h e a s t , and i s e v i d e n t l y a r e c e n t c o l o n i s e r s i n c e i t was n o t r e c o r d e d by t h e Oxford E x p e d i t i o n i n 1 9 3 8 .
Coconut p l a n t a t i o n s a r e l i m i t e d t o t h e w e s t o f Blossom V i l l a g e , t o Crawl Bay, t o Gwen I s l a n d , and t o a few g r o v e s i n o t h e r l o c a l i t i e s . B u d - r o t d i s e a s e h a s been p r e s e n t on L i t t l e Cayman s i n c e 1 9 1 3 , a n d many t r e e s a r e d e a d a n d d e c a p i t a t e d , e s p e c i a l l y n e a r Blossom V i l l a g e and on Owen I s l a n d ( P l a t e s 4 4 and 4 5 ) . Gwen I s l a n d i t s e l f ( F i g u r e 22) i s c o v e r e d p a r t l y b y c o c o n u t p l a n t a t i o n , p a r t l y b y a d e n s e s c r u b o f
d i s c r e t e p a t c h e s o f C o c c o l o b a (up t o 8 m t a l l )
,
L a g u n c u l a r i a and C o r d i a , w i t h C o n o c a r p u s a n d T h e s p e s i a and s c a t t e r e d T h r i n a x . A r e a s n e a r t h e s h o r e s h a v e a low s c r u b o f B o r r i c h i a , T o u r n e f o r t i a , C o n o c a r p u s and C o l u b r i n a , a n d t h e r e a r e s m a l l g r o v e s o f R h i z o p h o r a and A v i c e n n i a on s h e l t e r e d b e a c h e s . The h e r b a c e o u s g r o u n d c o v e r i n t h e c o c o n u t p l a n t a t i o n i s d i v e r s e . M e l a n t h e r a a s p e r a and W e d e l i a t r i l o b a t a a r e v e r y common, a s a r e t h e f o l l o w i n g s p e c i e s :A m b r o s i a h i s p i d a Ipomoea m a c r a n t h a
B o e r h a v i a e r e c t a Ipcmoea p e s - c a p r a e
C a n a v a l i a r o s e a M e l a n t h e r a a s p e r a
C e n c h r u s s p . P h i l o x e r u s v e r m i c u l a r i s
C y p e r u s s p . P o r t u l a c a o l e r a c e a
E u p h o r b i a m e s e m b r i a n t h e m i f o l i a S e s u v i u m p o r t u l a c a s t r u m H y m e n o c a l l i s li t o r e a W e d e l i a t r i l o b a t a
R e c e n t l y c l e a r e d g r o u n d i s e x t e n s i v e a l o n g t h e s h o r e s o f Bloody Bay, from Bloody Bay P o i n t t o J a c k s o n ' s Bay, and o f W e a r i s Bay. The g r o u n d i s b e i n g c o l o n i s e d by weedy s p e c i e s ; c l e a r i n g o f t r e e s and s h r u b s h a s b e e n c o m p l e t e , e x c e p t f o r m a t u r e t r e e s o f S w i e t e n i a mahagoni which have been l e f t s t a n d i n g .
A l l human s e t t l e m e n t s on L i t t l e Cayman a r e l o c a t e d on t h e c o a s t a l sand f l a t s , and w i t h a v e r y few e x c e p t i o n s t h e h o u s e s a r e grouped i n Blossom V i l l a g e . There i s a common assemblage of i n t r o d u c e d d e c o r a t i v e p l a n t s around t h e h o u s e s , i n c l u d i n g C r i n u m , Z e p h y r a n t h e s , B o u g a i n v i l l e a and C a t h a r a n t h u s , w i t h t a l l t r e e s of T e r m i n a l i a , Cocos and C a s u a r i n a . 4 . Rock pavement ( I r o n s h o r e o u t c r o p )
" L i t t o r a l r o c k pavement" v e g e t a t i o n was d e s c r i b e d by Beard (1955) and s u b s e q u e n t l y by Asprey and L o v e l e s s (1958) i n n o r t h Jamaica and by L o v e l e s s (1960) i n A n t i g u a . Somewhat i n a p p r o p r i a t e l y , however, t h e y a p p l i e d t h e term " r o c k pavement" t o s t e e p rocky s l o p e s , l e d g e s and t e r r a c e s , o f t e n forming c l i f f s o r ramps, r a t h e r t h a n h o r i z o n t a l rock p l a t f o r m s . The term i s much more s u i t a b l e f o r t h e two main a r e a s of f l a t I r o n s h o r e pavement which o u t c r o p on t h e L i t t l e Cayman c o a s t : one souch of t h e a i r s t r i p , r h e o t h c r a t S a l t Rocks. There i s a l s o a l e s s c o n s p i c u o u s narrow pavement e a s t o f Mary's Bay on t h e n o r t h e a s t c o a s t . The maximum w i d t h o f t h e two main pavements i s a b o u t 50 m , and t h e h e i g h t a b o u t 2 . 5 m . S i m i l a r pavements a r e much more widespread on b o t h Grand Cayman and Cayman B r a c .
The o u t e r 15-20 m o f t h e pavement h a s no macroscopic v e g e t a t i o n o t h e r t h a n s c a t t e r e d p a t c h e s of S e s u v i u m , P h i l o x e r u s , C e n c h r u s and S p o r o b o l u s i n d e p r e s s i o n s . The i n n e r p a r t of t h e pavement i s covered by t h e f o l l o w i n g z o n e s , from s e a t o l a n d :
a f l e s h y mat o f S e s u v i u m ;
a zone o f dwarfed and p r o s t r a t e s h r u b s , i n c l u d i n g C o n o c a r p u s , B o r r i c h i a , R h a c h i c a l l i s a m e r i c a n a and S t r u m p f i a m a r i t i m a ; a t r a n s i t i o n a l zone w i t h t h e s e dwarf s h r u b s and s c a t t e r e d S u r i ana nzari t i m a ;
an i n n e r zone o f t h i c k bushy S u r i a n a , 3-4 m t a l l ;
a hedge o f dense C o c c o l o b a a t t h e i n n e r edge o f t h e pavement, where i t p a s s e s b e n e a t h a sand o r c o b b l e beach.
T h i s d e s c r i p t i o n a p p l i e s t o t h e pavement a t West End P o i n t ; Sauer (1976, 5 ) h a s p u b l i s h e d a t r a n s e c t diagram f o r t h e same pavement somewhat f u r t h e r n o r t h .
The s o u t h e r n p l a t f o r m i s s i m i l a r e x c e p t t h a t L a g u n c u l a r i a and t a l l C o n o c a r p u s a r e a l s o p r e s e n t and t h e C o c c o l o b a woodland i s l e s s developed.
The z o n a t i o n may be compared w i t h t h a t d e s c r i b e d on h i g h e r and l e s s r e g u l a r c o a s t a l r o c k s i n A n t i g u a , where S t r u m p f i a i s dominant ( L o v e l e s s 1 9 6 0 ) , and n o r t h Jamaica (where R h a c h i c a l l i s i s dominant (Asprey and L o v e l e s s 1958) )
.
5 . S t e e p rocky c o a s t
B l u f f Limestone o u t c r o p s f o r a d i s t a n c e o f 1 . 2 5 km a l o n g t h e c o a s t a t t h e e a s t end o f t h e i s l a n d , between S n i p e P o i n t and E a s t P o i n t ; t h e s u r f a c e r i s e s t o a h e i g h t o f 6-7 m w i t h i n 50 m of t h e s h o r e , and t h e r o c k i s d e e p l y d i s s e c t e d , w i t h many p o t h o l e s 3-4 m deep. The o f f s h o r e r e e f i s narrow and g i v e s l i t t l e p r o t e c t i o n , and t h e lower s l o p e s a r e i n t h e s p l a s h zone. T h i s i s t h e k i n d of d i s s e c t e d c o a s t a l l i m e s t o n e d e s c r i b e d a s S o b o r u c o o r d i e n t e d e p e r r o by Howard and B r i g g s (1953) i n Cuba and by a v a r i e t y of o t h e r v e r n a c u l a r names ( m a k a t e a , f e o ,
c h a m p i g n o n ) i n o t h e r p a r t s of t h e world; i t i s a l s o t h e same a s t h e L i t t o r a l Rock Pavement of Beard (1955) and Asprey and L o v e l e s s ( 1 9 5 8 ) .
On L i t t l e Cayman t h e lower s p l a s h zone i s d e v o i d o f p l a n t s e x c e p t f o r p a t c h e s of S e s u v i u m i n p o t h o l e s . A t 2-3 m above s e a - l e v e l t h e r e i s a p r o s t r a t e s c r d of dwarf skxubs, i n c l u d i s g C o ~ o c a r p u s , E o r r i c h i a and S t r u m p f i a m a r i t i m a . A t r a t h e r h i g h e r l e v e l s S u r i a n a becomes e x t e n s i v e , t o g e t h e r w i t h C a p p a r i s f l e x u o s a , E r i t h a l i s f r u t i c o s a , and Ph y l l a n t h u s a n g u s t i f o l i u s , Low C o c c o l o b a becomes dominant a b o u t SO rn from the s e a , forming a c o n t i n u o u s zone 40-50 m wide, e x c e p t f o r d e e p e r p o t h o l e s , many o f which have m e e s o f F i c u s a u r e u s growing i n them.
F u r t h e r i n l a n d , away from t h e s e a , B u r s e r a , T h e s p e s i a
,
C o r d i a , P l u m e r i a,
T h r i n a x and C e p h a l o c e r e u s become c o n s p i c u o u s . T a l l e r t r e e s of B u r s e r a a r e commonly deformed i n t o a L-shape where t h e y a r e emergent. T h i s v e g e t a t i o n i s v e r y s i m i l a r t o t h a t o r i g i n a l l y d e s c r i b e d by Beard (1955) f o r Barbuda, where t h e s c r u b on s i m i l a r t e r r a i n c o n s i s t s of C o n o c a r p u s , C o c c o l o b a , S u r i a n a , E r n o d e a , E r i t h a l i s , S t r u m p f i a , C r o t o n , O p u n t i a and s p h e r i c a l c a c t i .
Sandy p a t c h e s a l o n g t h e rocky s h o r e a t E a s t End c a r r y mats o f S e s u v i u m a n d P h i l o x e r u s , w i t h Ipomoea p e s - c a p r a e , I. m a c r a n t h a and H y m e n o c a l l i s , and o c c a s i o n a l i s o l a t e d t r e e s of C o n o c a r p u s up t o 8 m
t a l l ,
C . Mangrove communities
T h i s c a t e g o r y mainly comprises swamp communities dominated by mangroves. Some mangrove a r e a s a r e n o t swampy, however, and n o t a l l L i t t l e Cayman swamps i n c l u d e mangroves; f o r convenience h e r b a c e o u s swamps dominated by S e s u v i u m a r e i n c l u d e d h e r e . L i t t l e Cayman h a s no c o u n t e r p a r t t o t h e T y p h a swamps o f Grand Cayman. We d i s t i n g u i s h c o a s t a l mangroves; mangroves of c o a s t a l ponds and s a l t f l a t s ; i n l a n d
mangroves; and herbaceous swamps.
1. C o a s t a l mangroves
A p a r t from s c a t t e r e d b e a c h - f o o t t r e e s of A v i c e n n i a , L a g u n c u l a r i a and R h i z o p h o r a on rock and c o b b l e h e a d l a n d s i n Bloody Bay and Wearis Bay, c o a s t a l mangroves a r e r e s t r i c t e d t o a s t r i p a few m e t r e s wide a l o n g t h e n o r t h s h o r e o f The B i g h t i n S o u t h Hole Sound and on t h e nearby
leeward s h o r e of Owen I s l a n d . T h i s c o n s i s t s o f t r e e s o f R h i z o p h o r a and A v i c e n n i a 5-8 m t a l l ; t h e mangrove i s n o t s u f f i c i e n t l y well-developed t o e x h i b i t z o n a t i o n . Bare ground on t h e landward margin o f t h e s e
mangroves i s c o l o n i s e d by S e s u v i u m . B a t i s m a r i t i m a i s p r e s e n t b u t i s v e r y r a r e , i n c o n t r a s t t o i t s l u x u r i a n t development i n A v i c e n n i a swamp on Grand Cayman.
2 . Mangroves o f c o a s t a l ponds and s a n d f l a t s
E x t e n s i v e l i n e a r ponds a r e f r e q u e n t between t h e c o a s t a l s a n d r i d g e and t h e l i m e s t o n e u p l a n d s on L i t t l e Cayman; t h e y a r e s h a l l o w and r o c k - f l o o r e d , w i t h a v a r i a b l e s e d i m e n t c o v e r . The ponds v a r y g r e a t l y i n t h e e x t e n t o f mangroves on t h e i r s h o r e s . Some, s u c h a s t h o s e a t Blossom V i l l a g e and P i r a t s ' s P o i n t , have m a r g i n a l A v i c e n n i a , L a y u n c u l a r i a and C o n o c a r p u s ; o t h e r s , notcibly Tarpon Lake, have t a l l m a t u r e R h i z o p h o r a woodland ( P l a t e 4 6 ) . I n t h e Blossom V i l l a g e pond, dwarf A v i c e n n i a i s e x t e n s i v e , e s p e c i a l l y a t t h e wesE e n d , where b l e a c h e d stumps i n d i c a t e an even g r e a t e r former e x t e n t , presumably i n t e r r u p t e d b y h u r r i c a n e damage. I n t h e P i r a t e ' s P o i n t pond t h e r s i s a l s o s p u r e s t a n d o f
k c r o s t i c . , l i u n ~ a u ~ e u r i l 2 m mil. T h i s pond s l s o c o c t a i n e d a mat o f R u p ~ i a
-
- m a r i t i m a i n 1 9 7 5 , i n a d d i t i o n t o m a r g i n a l mzts o f S e s u v i u ~ n a1.dS a l i c o r n i a .
The mangrove a t Tarpon Lake i s t h e most e x t e n s i v e on t h e i s l a n d , o c c u p y i n g an a r e a 2 . 5 km !-ong ( e a s t - w e s t ) and a b o u t 1. krn w i d e ; i t !?as a maximum h e i g h t o f a b o u t 25 m. The t a l l e s t woodland c o n s i s t s o f R h i z o p h o r a m a n g l e and L a g u n c u l a r i a r a c e m o s a , w i t h A v i c e n n i a and C o n c a r p u s m a r g i n a l on t h e b e a c h - r i d g e s i d e and A v i c e n n i a a l s o forming p a r t o f t h e l a k e s h o r e ( P l a t e s 4 7 - 4 9 ) . There a r e no f e r n s , b u t on t h e landward s i d e many o f t h e t r e e s a r e f e s t o o n e d w i t h R h a b d a d e n i a b i f l o r a . The mangrove i s c o n t i n u o u s b e h i n d t h e b e a c h r i d g e f o r 2-3 km t o t h e e a s t o f t h e l a k e , b u t t h e w i d t h o f t h e swamp n a r r o w s t o 50-200 m and t h e h e i g h t o f t h e t a l l e s t t r e e s d e c l i n e s t o o n l y 6-8 m.
3 . I n l a n d mangroves
Mangroves a r e a l s o found w i t h i n some o f t h e l a r g e r i n l s n d ponds w i t h i n t h e B l u f f Limestone. One such un-named pond (063786) a t t h e e a s t end i s f r i n g e d w i t h a narrow zone o f L a g u n c u l a r i a , and t h e r e i s a wide zone o f t w i s t e d C o n o c a r p u s on r o u g h l y d i s s e c t e d l i m e s t o n e m a r g i n a l t o t h e p o o l .
The most e x t e n s i v e i n l a n d mangroves, however, a r e n o t d i r e c t l y a s s o c i a t e d w i t h open s t a n d i n g w a t e r . T h e r e a r e t h r e e s u c h e x t e n s i v e a r e a s i n t h e w e s t e r n p a r t o f t h e i s l a n d ( a t 957752, 975761 and 9 8 8 7 6 6 ) . The s e c o n d o f t h e s e i s e n c l o s e d t o n o r t h and s o u t h by B l u f f Limestone r i d g e s r i s i n g t o 6-13 m above s e a - l e v e l , b u t i t s s u r f a c e l i e s c l o s e t o s e a - l e v e l . The c e n t r e o f t h e mangrove c o m p r i s e s dwarf R h i z o p h o r a , v a r y i n g i n h e i g h t from 1.5-1.9 m , w i t h l o w e s t f o l i a g e 65-90 cm above t h e ground ( P l a t e 5 0 ) . These mangroves a r e d i s t i n g u i s h e d by d e n s e r o o t s y s t e m s and s m a l l l e a v e s . Root d e n s i t i e s a v e r a g e a b o u t 80 p e r s q m , w i t h a mean r o o t d i a m e t e r 50 cm above t h e ground of 2 . 3 cm. The mean l e n g t h and b r e a d t h o f 50 l e a v e s measured 1 0 . 1 and 3 . 8 cm,
r e s p e c t i v e l y , and t h e maxima 12.2 and 4.6 cm.
On t h e s o u t h e r n s i d e o f t h e dwarf R h i z o p h o r a t h e r e i s a zone o f s l i g h t l y t a l l e r R h i z o p h o r a i n t e r m i x e d w i t h L a g u n c u l a r i a a n d A c r o s t i c h u m . Mean t r e e h e i g h t i n t h i s zone i s a b o u t 2 m , and t h e h e i g h t o f t h e l o w e s t f o l i a g e above t h e g r o u n d i s 0 . 8 - 1 . 1 5 m. R o o t d e n s i t y i s a b o u t 55 p e r s q m , and mean r o o t d i a m e t e r 5 0 cm a b o v e t h e g r o u n d i s 2 . 6 cm.
T h e s e v e r y u n i f o r m i n l a n d R h i z o p h o r a swamps r e s e m b l e b u t a r e much more e x t e n s i v e t h a n t h e i n l a n d dwarf mangroves d e s c r i b e d from s o u t h e r n B a r b u d a ( S t o d d a r t , Bryan a n d G i b b s 1973)
.
L a r g e a r e a s o f l o w - l y i n g d i s s e c t e d B l u f f L i m e s t o n e i n t h e i n t e r i o r o f L i t t l e Cayman a r e a l s o c o v e r e d w i t h d e n s e , t w i s t e d C o n o c a r p u s and L a g u n c u l a r i a . The t e r r a i n i s r u g g e d , w i t h s a n d - f i l l e d d e p r e s s i o n s
i n h a b i t e d by C a r d i s o m a . B o t h E r i t h a l i s a n d S t r u m p f i a a r e f o u n d i n some l o c a l i t i e s m a r i n g a l t o t h e s e C o n o c a r p u s t h i c k e t s .
4 . Werbaceous swamp
A f u r t h e r t y p e o f i n l a n d m a r s h , a l s o a t s e a - l e v e l , i s f o u n d a b o u t 500 IT: from t h e s e a a t khe e a s t e1:d o f t h e i s l a n d ( 0 7 5 7 9 5 ! , s l ~ ~ r r o u n d e c ? b y d e e p l y d i s s e c t e d B l u f f L i m e s t o n e r i d g e s 2-4 m h i g h . I t c o n s i s t s o f a t h i c k maL o f s u c c u l e n ~ S e s u v i u m , w i t h o c c a s i o n a l r i d g e s a n d p i n n a c l e s o f t h e u n d e r l y i n g B l u f f L i m e s t o n e e m e r g i n g t h r o u g h i t . I n August 1 9 7 5 t h e whole o f t h i s marsh was w a t e r l o g g e d ( P l a t e 5 1 1 , a n d t h e r e w e r e some s m a l l p o o l s o f s t a n d i n g w a t e r .
D . C l a s s i f i c a t i o n o f h a b i t a t s
J o h n s t o n ( 1 9 7 5 ) d e f i n e d a number o f " e c o l o g i c a l f o r m a t i o n s " i n t h e Cayman I s l a n d s , b a s e d o n " m a j o r a n d m i n o r p l a n t a s s o c i a t i o n s ,
d i s t i n c t i v e t o p o g r a p h i c f e a t u r e s , and p e r t i n e n t g e o l o g i c , e d a p h i c , a n d h y d r o l o g i c c o n d i t i o n s " ( 1 9 7 5 , 2 4 3 ) . T h e s e f o r m a t i o n s w e r e l a r g e l y d e f i n e d i n t e r m s o f t h e h a b i t a t r e q u i r e m e n t s o f b i r d s , b o t h t e r r e s t r i a l a n d a q u a t i c . The main u n i t s r e c o g n i s e d w e r e :
F r i n g i n g r e e f s and l a g g o n s . S a n d - c o r a l b e a c h .
S e a Grape-Almond Woodland.
Mangrove Swamp.
P a s t u r e s a n d c u l t i v a t e d a r e a s .
Logwood F o r e s t o r s c r u b woodland ( s e c o n d a r y ) . L i m e s t o n e F o r e s t .
I n l a n d l a g o o n s a n d p o n d s . Town a n d h o u s e s i t e s .
The c a t e g o r i e s adopted have i n some c a s e s ( e . 9 . t y p e s 6 and 9) g r e a t e r r e l e v a n c e f o r Grand Cayman t h a n f o r t h e o t h e r two i s l a n d s , and i n o t h e r c a s e s , n o t a b l y t y p e 7 , a s i n g l e u n i t c o v e r s a wide v a r i e t y o f t o p o g r a p h i c and v e g e t a t i o n a l forms, e s p e c i a l l y on L i t t l e Cayman.
I t i s t h e r e f o r e , a p p r o p r i a t e t o conclude t h i s c h a p t e r w i t h a summary of t h e main h a b i t a t s t h a t can be r e c o g n i s e d on L i t t l e Cayman. I t i s based on topography, geology and v e g e t a t i o n , and i s i n t e n d e d a s a framework f o r d i s c u s s i o n of o t h e r a s p e c t s o f t h e t e r r e s t r i a l ecology and a l s o f o r c o n s i d e r a t i o n o f c o n s e r v a t i o n and management p r o c e d u r e s . A s it s t a n d s t h e c l a s s i f i c a t i o n i s e m p i r i c a l and i s n o t n e c e s s a r i l y a p p r o p r i a t e f o r t h e o t h e r Caynan I s l a n d s .
1, C o a s t a l h a b i t a t s --
i b ) Cobble beaches wl. th C o c c o l o b ~ l i t t o r a l hcdge
.
( c ) L i t t o r a l woodlan6 o f c o a s t a l sand f l a t s ( a l m o s t e n t i r e l y c l e a r e d )
.
( d ) S t r a n d s c r u b , secondary s c r u b , and c l e a r e d and c u l t i v a t e d l a n d o f c o a s t a l sand f l a t s .
B. Rocky s h o r e s .
( a ) Dwarf s c r u b of h o r i z o r - t a l I r o n s h o r e pavements.
( b ) Dwarf scrub o f c o a s t a l h l u f f s and c l i f f s .
C . Marine mangroves.
( a ) Rhizophora and Avicennia c o a s t a l f r i n g e s .
2. Non-marine Manaroves
A . Mangrove woodland;
( a ) T a l l Rhizophora woodland (Tarpon L a k e ) . ( b ) Low Rhizophora woodland on s a l t - p o n d margins.
( c ) Conocarpus and h e r b a c e o u s v e g e t a t i o n of r o c k f l a t s round s a l t ponds (Sesuvium sward, S a l i c o r n i a sward, A c r o s t i chum clumps)
.
B. Mangrove s c r u b .
( a ) Dwarf i n l a n d R h i z o p h o r a s c r u b i n e n c l o s e d b a s i n s .
( b ) C o n o c a r p u s - L a g u n c u l a r i a s c r u b o f low d i s s e c t e d i n t e r i o r B l u f f L i m e s t o n e .
C . H e r b a c e o u s marsh.
( a ) S e s u v i u m marsh o f i n t e r i o r b a s i n s i n B l u f f L i m e s t o n e . 3 . I n t e r i o r l i m e s t o n e a r e a s
A. B l u f f L i m e s t o n e .
;a) Dry E v e r q r e e n T h i c k e ~ on d i s s e r t e d l i m e s t o n e r i d g c s . ( b ) E v e r g r e e n Bilshland on low d i s s e c t e d I.iniestone
( c ) S e e c a t e g o r y 2 . B , b , B. I r o n s h o r e F o r m a t i o n .
( a ) Dry E v e r g r e e n F o r e s t on low u n d i s s e c t e d Marl F a c i e s .
( b ) Dry E v e r g r e e n Woodland on g e n t l e s l o p e s o f low u n d i s s e c t e d Marl F a c i e s r i d g e s .
( c ) E v e r g r e e n B u s h l a n d on low u n d i s s e c t e d Marl F a c i e s ( d i s t u r b e d ? j
.
R e f e r e n c e s
A s p r e y , G.F. a n d L o v e l e s s , A . R . 1958. The d r y e v e r g r e e n f o r m a t i o n s o f J a m a i c a . 11. The r a i s e d c o r a l b e a c h e s o f t h e n o r t h c o a s t .
J . E c o l . 4 6 , 547-570.
A s p r e y , G.F. and R o b b i n s , R . G . 1953. The v e g e t a t i o n o f J a m a i c a . E c o l . M o n o g r . 23, 359-412.
B e a r d , J . S . 1944. Climax v e g e t a t i o n i n t r o p i c a l America. E c o l o g y , 25, 127-158.
B e a r d , J . S . 1949. The n a t u r a l v e g e t a t i o n o f t h e Windward and Leeward I s l a n d s . O x f o r d F o r e s t r y Mem. 2 1 , 1-192.
B e a r d , J . S . 1955. The c l a s s i f i c a t i o n o f t r o p i c a l American v e g e t a t i o n - t y p e s . E c o l o g y , 36, 89-100.
D'Arcy, W.G. 1971. The i s l a n d o f Anegada and i t s f l o r a . A t o l l R e s . B u l l . 1 3 9 , 1-21.
D'Arcy, W.G. 1975. Anegada I s l a n d : v e g e t a t i o n and f l o r a . A t o l l R e s . B u l l . 1 8 8 , 1-40.
D a v i s , J . H . 1 9 4 2 . The e c o l o g y o f t h e v e g e t a t i o n and t o p o g r a p h y o f t h e s a n d k e y s o f F l o r i d a . P a p . T o r t u q a s L a b . 3 3 , 113-195.
F o s b e r g , F.R. 1 9 7 1 . The f e r n v e g e t a t i o n o f A l d a b r a . Am. F e r n J . 6 1 , 97-101.
H a r r i s , D . R . 1 9 6 5 . P l a n t s , a n i m a l s , and man i n t h e O u t e r Leeward I s l a n d s , West I n d i e s : an e c o l o g i c a l s t u d y o f A n t i g u a , B a r b u d a , and A n g u i l l a . U n i v . C a l i f . P u b l s G e o g . 1 8 , i - i x , 1-164.
Howard, R . A . 1950. V e g e t a t i o n o f t h e B i m i n i I s l a n d g r o u p , Bahamas, B . W . 1 . E c o l . Monogr. 2 0 , 317-349.
Howard, R . A . and B r i g g s , W.R. 1953. The v e g e t a t i o n o f c o a s t a l d o g t o o t h l i m e s t o n e i n s o u t h e r n Cuba. J . A r n o l d A r b . 34, 88-96.
J o h n s t o n , D . W . 1 9 7 5 . E c o l o g i c a l . a n a l y s i s o f t h e Cayman I s l a n d a v i f a u n a , B u l l , F l a S t . Mus. h i o l . S c i , 1 9 , 235-300.
L o v e l e s s , A.R. 1 9 6 0 . The v e g e t a t i o n o f A n t i g u a , Wes.t I n d i e s . J . E c o l . 4 8 , 435-527,
L o v e l e s s , A . R . and A s p r e y , G.F. 1 9 5 7 , The d r y e v e r g r e e n f o r m a ~ i o n s o f J a m a l c a . I . Tne l i m e s t o n e h i l l s o f t h e s o u t h c o a s t . J . E c o l . 4 5 , 799-822.
P o g g i e , J . J . 1 9 6 2 . C o a s t a l p i o n e e r p l a n t s and h a b i t a t i n t h e Tampico r e g i o n , Mexico. C s t l S t u d . I n s t . , La S t . U n i v . , Tech. R e p t . 17A, 1-62.
S a u e r , J . D . 1 9 5 9 . C o a s t a l p i o n e e r p l a n t s o f t h e C a r i b b e a n and G u l f o f M e x i c o . Madison: U n i v e r s i t y o f W i s c o n s i n , D e p a r t m e n t s o f Botany and Geography.
S a u e r , J . D . 1967. G e o g r a p h i c r e c o n n a i s s a n c e o f s e a s h o r e v e g e t a t i o n a l o n g t h e Mexican Gulf C o a s t . C s t l S t u d . I n s t . La S t . U n i v . C s t l S t u d . S e r . 21, i - x , 1-59.
S a u e r , J , D . 1976. Problems and p r o s p e c t s o f v e g e t a t i o n a l r e s e a r c h i n c o a s t a l e n v i r o n m e n t s . Geoscience a n d Man, 1 4 , 1 - 1 6 ,
S e i f r i z , W . 1 9 4 3 . The p l a n t l i f e o f Cuba. E c o l . Monogr. 1 3 , 355-426.
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A t o l l R e s . B u l l . 1 9 0 , 87-116.
S t o d d a r t , D . R . , B r y a n , G . W . , and G i b b s , P.E. 1973. I n l a n d mangroves and w a t e r c h e m i s t r y , B a r b u d a , West I n d i e s , J . n a t . H i s t . 7 , 33-46.
Warming, E . 1909. O e c o l o g y o f p l a n t s : a n i n t r o d u c t i o n t o the s t u d y o f p l a n t c o m m u n i t i e s . O x f o r d : U n i v e r s i t y P r e s s .
