---------------

No. 102

Notes on r e e f h a b i t a t s and gastropod molluscs of a

lagoon i s l a n d a t North Ma.le Atoll, Maldives

by
Alan J. Kohn

Issued by

THE PACIFIC SCIENCE BOARD

National Academy of S c i e n c e s - 4 a t i o n a l Resarch Council

Washington, D. C.

September 30, 1864

 Notes on r e e f h a b i t a t s and gastropod m o l l u s c s of a

lagoon i s l a n d a t North A t o l l , Maldives

E o r t n i w h l eh.to1-1 e v i n c e s most of t h o p e c u l i a r i t i e s c h a r a c t e r i s t i c
o f s e v e r 2 1 Waldivian a t o l l s b u t n o t found elsewhere. Its r i m , w i t h
maximum dimensions o l 5i x 37 km, i s formed. m i n l y by about 2 5 f a r o s ,
small a t o l l s w i t h i n t h e l a r g e r convlex. Also i n c l u d e d i n t h e r i m a r e
a few s i n g l e i s l a n d s p a r t i a l l y surrounded by f r i n g i n g r e e f s ; t h e l a r g e s t
o f t h e s e i s Nale I s l r i d , t h e s e a t of government. I n a d d i t i o n , more
t h a n 8 0 f a r o s m d s i ; ~ ~i sl lla n d s occur wit'nin t h e lagoon. The f a r o s and
i s l a n d s of t h e rim a r e s e p a r a t e d by about 30 l a r g e channels around i t s
e n t i r e c i r c u n f e r e n c e . The c h a n n e l s a r e 30 -
150 m deep, i . e . much deeper
t h a n t y p i c a l P a c i f i c a t o l l channels.

The l a r g e s i z e of t h e c h a n n e l s r e s u l t s from m e t e o r o l o g i c a l and

hydrographic c o n d i t i o n s a s s o c i a t e d with t h e r e g d l a r a l t e r n a t i ~ nof
n o r t h e a s t and s o u t h v ~ e s tmonsooos. The r e s u l t i n g i n c r e a s e d o c e a n i c
c i r c u l a t i o n i n and through t h e l a p o n i s t h e proximate cause of t h e
f a r o s and i n n e r o r lagoon i s l a n d s . The term c o m ~ o s i t ez t o l l a p p l i e s t o
North H a l e and some o t h e r Maldive a t o l l s because of t h e s e f e a t u r e s .

Origifi of Maldive A t o l l s

Although I have h e l d elsewhere (Kchn, 1961: t h a t Dwrwints subsid-

ence t h e o r y is t h e b e s t g e n e r a l e x p l a n a t i o n of t h e o r i g i n of a t o l l s ,
o t h e r p r o c e s s e s h m e probably been more i m p o r t a n t i n d e t e r m i n i n g t h e
p r e s e n t f e a t u r e s of t h e Maldive a t o l l s . C e r t a i n a s p e c t s of Naldive
a t o l l s s u p p o r t t h e a n t e c e d e n t p l a t f o r m t h e o r y (Hoffrneister and Ladd,
1944).

Formation of Lagoon I s l c m d s

The i s l a n d s w i t h i n some Naldive a t o l l l a g o o n s probably begin as

c o r a l growth on a p l a t f o r m a t s u i t a b l a depth (50-60 m). Grob~thi s
enhanced by t h e u n u s u a l l y l a r g e flow of o c e a n i c water through t h e
b r o a d , deep c h m n e l s , and c o n d i t i o n s f a v o r p e r i p h e r a l development of
t h e r e e f i n t o a f a r o ( s e e Koffmeister and Ladd, 1944). A s t h e r i m of
t h e f a r o grows t o w ~ dt h e s u r f a c e , i t s i n t e g r i t y i n c r e a s e s and i t s
l a g o o n , known as t h a v e l u , becomes i n c r e a s i n g l y p r o t e c t e d and a r e g i o n
of sediment d e p o s i t i o n . G r a d u a l l y , t h e f a r o i s transformed i n t o
i s l a n d with a f r i n g i n g r e e f . A s Agassiz (1903) n o t e d , a l l s t a g e s of
such t r a n s f o r m a t i o n s may be observed on Morth Male A t o l l .

Department of Zoology, U n i v e r s i t y of Washington.

Funidu I s l a n d

The i s l a n d studied i s not named on c h a r t s , and we have t r a n s c r i b e d

t h e name given us o r a l l y by t h e i n h a b i t a n t s a s Funidu. It is an e l l i p -
t i c a l i s l a n d (about 1%x 69 m) j u s t i n s i d e t h e main s o u t h e a s t channel
in.to North Piale lagoon and j u s t northwest of a l i n e between t h e north-
e a s t t i p of Male I s l a n d and t h e south t i p of Hulule I s l a n d . It is t h e
unnamed i s l a n d v i s i t e d i n 1901 o r 1902 and described by Alexander figassir,
(1903: p. 43, p l . 9, f i g . 2). Agassiz's d e s c r i p t i o n of t h e i s l a n d i s
as follows:

l 1 A s a type of an i n n e r i s l a n d , we examined a small well-

wooded i s l a n d which r i s e s i n t h e middle of t h e s o u t h e a s t
passage i n t o North Male with n e a r l y t h i r t y fathoms on e i t h e r
s i d e on i t . The i s l a n d i s e l l i p t i c a l , and i s placed on t h e
northwestern horn of an e l l i p t i c a l f l a t which s t r e t c h e s out
i n a s o u t h e a s t e r l y d i r e c t i o n . The f l a t s l o p e s very g e n t l y
t o t h e s e a , i s edged on t h e o u t e r r i a by a s i n k forming a
shallow d i t c h of varying width, flanked by c o r a l boulders o r
masses of beach rock extend-ing a s an i r r e g u l a r wall along
t h e g r e a t e r p a r t of the o u t e r edge of t h e r e s f f l a t . The
g r e a t e s t width of t h e r e e f i s from t t ~ ohundred and f i f t y t o
t h r e e hundred f e e t . It diminishes gradually i n width t o
t h e northwestern c o r n e r , where the i s l a n d i s s t e e p to.
Corals grow i n g r e a t abundance a t a depth of from f i v e t o
t h r e e fathoms upon t h e s t e e p slopes of t h e r e e f f l a t ; they
grow with l e s s profusion t o s i x o r seven fathoms, where they
a r e separated by wide l a n e s and patches of s a d which even-
t u a l l y cover the whole botton! a t a depth of from e i g h t t o
nine fathoms. From t h e three-fathom l i n e , they a l s o diminish
i n number towards the s u r f a c e and spread over t h e edge of
t h e f l a t , which i s p a r t l y bare a t low water; they extend but
a s h o r t way over i t , t h e g r e e t e r p a r t of the f l a t being
covered by dead c o r a l s overgrown with Nullipores. The sand
beaches surrounding t h e i s l a n d a r e s t e e p , from s i x t o seven
f e e t i n height. The c e n t r a l p a r t of t h e i s l a n d i s lower
than tl:e beaches which surround i t , forming a shallow s i n k
from twelve t o eighteen inches o r more i n d-epth. Consider-
a b l e moisture accumulates i n t h i s shallow sink, and i n t h e
r a i n y season a pool i s probably formed of more o r l e s s
brackish water. This s t r u c t u r e is most c h a r a c t e r i s t i c of
t h e i s l a n d s of t h e Maldives, whether they occur i n the in-
t e r i o r of t h e g r e a t s h e e t s of enclosed waters o r on t h e
o u t e r r i m s of t h e plateaus. The s i n k has been formed by
the washing up of t h e beaches round a c e n t r a l a r e a , a s we
have seen i t i n t h e Paumotus and elsewhere i n o t h e r P a c i f i c
a t o l l s . Before the vegetation became too dense, beach sand
w a s blown towards the i n t e r i o r and p a r t l y f i l l e d t h e c e n t r a l
a r e a , u n t i l t h i s was prevented by t h e growth of bushes and
shrubs, when t h e beaches merely increased i n height and t h e
sand of t h e upper r i d g e of t h e beach was driven s p a r i n g l y
 towards t h e c e n t r e of t h e i s l a n d , o r its-f u r t h e r passage
stopped by t h e b e l t o f d e n s e r v e g e t a t i o n which had come
up on t h e h i g h e r p a r t s of t h e c o r a l sand beach, The
i s l a n d o f PIale h a s g r a d u a l l y developed and been formed
much i n t h e same manner as t h i s i s l a n d ,

The account d e s c r i b e s t h e i s l a n d much as we observed i t i n 1957,

e x c e p t we do n o t r e c a l l t h a t t h e c e n t r a l ort ti on appeared lower t h a n
t h e uppermost beachec. Perhaps d u r i n g t h e i n t e r v e n i n g 55 y e a r s s u f f i -
c i e n t sand h a s blown onto t h e i s l a n d t o e l i m i n a t e t h e v e s t i g e s o f t h e
v e l u , as a p p a r e n t l y happened e a r l i e r on Male.

A g a s s i z l s o b s e r v a t i o n s of t h e f r i n g i n g r e e f around Funidu a r e con-

f i n e d t o t h e f o l l o w i 2 g n o t e on t h e c o r d s :

'?The c o r a l s growing on t h e s l o p e s of t h i s i s l a n d a r e
marked f o r t h e i r l ~ m r i a ~ c ethey ; grow n s abundantly as
t h e y do on t h e s e a f a c e of any a t o l l . T h i s i s i n s t r i k i n g
c o n t r a s t t o t h e i r s c c m t y developaent i n t h e i n t e r i o r of
t y p i c a l lagoons. It can r e a d i l y be e x p l a i n e d from t h e
g r e a t d e p t h s of t h e p a s s e s and t h e g r e a t mass and p u r i t y
o f t h e water p a s s i n g i n t o t h e i n t e r i o r of t h e enclosed
b a s i n o f North Male. .. The branching c o r a l s c o n s i s t
mainly o f s p e c i e s of Nadregores, of P o c i l l o p o r e s , and
Millepores. While t h e massive c o r a l s a r e u s u a l l y A s t r e a n s ,
P o r i t e s , and t h e l i k e , Naeandrinae a r e n o t common."

With D r . U i l l a r d D. Hartnan, I made f o u r t r i p s t o Funidu I s l m d ,

19-23 September 1957, d u r i n g t h e course of t h e Yale S e y c h e l l e s Expedi-
t i o n . Notes on h a b i t a t s and i n h a b i t a n t s were r e f e r r e d approxini!tely t o
t h e c o o r d i n s t e s i n d i c a t e d i n f i g . 1, as a c c u r a t e l y as could be done i n
t h e f i e l d ( s e e Kohn, 1956, f o r method of r e c o r d i n g d a t a ) . The d e n s i t y
of symbols i n f i g , 1 i n d i c a t e s t h e r e l a t i v e thoroughness of t h e observa-
t i o n s . The gastropod molluscs have been i d e n t i f i e d from c o l l e c t i o n s of
t h e Peabody Museum of N a t u r a l H i s t o r y , Yale U n i v e r s i t y . No a t t e m p t h a s
been made t o v e r i f y f u r t h e r t h e names used,

On t h e n o r t h s i d e of t h e i s l a n d ( q u a d r a t s B4-B8) t h e sand beach

is narrow and followed seaward by about 1 2 m of s l o p i n g beachrock
covered with a t h i n a l g a l mat, i n t h e i n t e r t i d a l zone. The s u r f a c e of
t h e beachrock i s smooth, b u t i t i s marked by d e p r e s s i o n s , p o t h o l e s , and
c r e v i c e s . Seaward of t h i s zone a t about t h e t i d a l datum (0: approxi-
mately mean low water s p r i r l g s ) , t h e beachrock beco~nesa very smooth l e v e l
bench covered with a t h i n l a y e r of sand. The width o f t h i s zone i s
mainly 7 m o r l e s s . Seaward i s a wide zone of l i v i n g c o r a l s i n v e r y
s h a l l o w w ~ t e r ,t h e t o p s of t h e heads a p p e a r i n g t o extend above datum.
Mitra l i t t e r a t a Lamarck and o t h e r s p e c i e s of Mitra were c o l l e c t e d i n
s h a l l o w c r e v i c e s i n t h e l i x e s t o n e and Trochus flanlmulatus Lamerck on
l i v e c o r a l i n t h e o u t e r zone as w e l l as on i n s h o r e beach rock a t about
0 - +1' t i d e l e v e l . I n B8, some r o c k s ( n a t u r e n o t recorded) extended
upward t o about +2' and perhaps s u g g e s t r a t h e r r e c e n t e l e v a t i o n .
 On t h e e a s t s i d e of t h e i s l a n d , a l a r g e t i d e pool is i n Cg. #The
f o l l o w i n g were c o l l e c t e d i n C 9 and D 9 on beach r o c k exposed - zt low t i d e :
-
T h a i s t u b e r o s a RtJding, N e r i t a h i s t r i o L., Zngina mendicaria L., Vasum
t u r b i n e l l u m I,., and Bursa sp.

Quadrats E'5-F8 a r e c h a r a c t e r i z e d by a f l a t , smooth l i n e s t o n e pave-

ment covered by a very t h i n l a y e r of s a d i m e d i a t c l y seaward of t h e
r a t h e r s t e e p , c o a r s e sand beach. L a t i r u s 6;?10.ra,~dula(L.) w a s u b i q u i t o u s
on t h i s t h i n l a y e r of sand. Drupa s p p . , liorulz spp.,
N a s s a r i u s sp., and Colurnbel& sp., were a l s o p r e s e n t . Mitra spp. were
c o l l e c t e d from c r e v i c e s i n t h e l i m e s t o n e rock. Seaward, t h e sand-covered
bench s u p p o r t s i s o l a t e d c o r a l heads. The sandy arsa i s p a d u a l l y reduced
and t h e c o r a l b e c ~ m e si n c r e a s i n g l y dense and s o l i d toward t h e o u t e r edge
( ~ 5G , 6 , H 4 , H 5 , EL).
0
I n F7, smooth beachrock s l o p e s o f f s h o r e a t about a 30 a n g l e j u s t
o f f s h o r e from t h e narrow sand beach. I n s:me a r e e s t h e r e a r e two t i e r s
of beachrock. L a t i r u s simragdula and Mitra l i t t e r a t a were c o l l e c t e d on
- t h i s beaclwocl;. Offshore p o r t i o n s of t h e r e e f on t h e e a s t s i d e could
n o t be i n v e s t i g a t e d because of t h e r a t h e r heavy s u r f . There a i ~ ~ z e a r e d
t o be much c o r a l r u b b l e and dead c o r a l i n t h i s r e g i o n . I n c r e a s e d abun-
dance of Drupa morum Rading was noted i n I;r and i s probnbly c o r r e l a t e d
w i t h t h e h e a v i e r s u r f i n t h i s r e g i o n . Morula t u b e r c u l a t a B l a i n v i l l e ,
n o t found i n calmer a r e z s t o t h e west, was a l s o p r e s e n t . It occurred
both i n t e r t i d a l l y and s u b t i d a l l y ; o t h e r s p e c i e s c o l l e c t e d i n t h i s q u a d r a t
a p ~ e a r e dt o be only s u b t i d a l . The substratum of F8 i s mainly beachrock
and c l o s e l y r e s e m b l r " ~F7.

A broader sand beach i s p r e s e n t on t h e west ~ i d . 2( ~ 3 3 , 3 ) , followed

seaward by a broad a r e a of t h i n sand on smooth, l e v e l l i m e s t o n e bench
which forms a channel s u i t a b l e f o r l a n d i n g a s m a l l b o a t i n C3. Specimens
-
of Turbo argyrostoma L., with e p i z o i c Eipponix s p . , Vasum turbinellurn,
--
N a s s a r i u s g a n i f e r u s ( K i e n e r ) , Cerithiuin s p . , %rombus m u t a b i l i s Swainson,
P o l i n i c e s s p . , M i t r a mitra L., Drupa g r o s s u l a r i a l o b a t a d l a i n v i l l e , and
P e r i s t e r n i a n a s s a t u l a Lmwxlc were c o l l e c t e d i n t h i s a r e a . S e w a r d (C2,
D 2 , E2) is a zone of dense growing c o r a l . The heads appear t o extend
s e v e r a l i n c h e s above datum, and many s p e c i e s c f c o r a l a r e :)resent. me
heads a r e i n t e r s p e r s e d with sandy a r e a s c o n t a i n i n g some r u b b l e , t h e sub-
s t r a t e being a t a b o a t -2.5'. I n some p l a c e s t h e r e i s a v e r y g e n t l e
s l o p e t o t h e r e e f edge, which d r o p s o f f p r e c i j j i t o u s l y i n t o b l u e wcter
(about 90 ni a c c . hdmircalty Chart 65b). Turbo argyrostorna were c o l l e c t e d
from l i v i n g c o r a l , t h e i r s h e l l s o f t e n b e a r i n g e p i z o i c Hipj:onix sp.
LamSis lambis (L. ) occurred on sand under overhanginc c o r a l heads, and
--
A t r i n a sp. occurred b u r i e d i n sand. Cerithium nodulosurr., Cerithium sp.,
--
P e r i s t e r n i a n a s s a t u l a occurred on sGmd amid l i v i n g and dead c o r a l ; Drupa
r i c i n u s ( L . ) , 2. g r o s s u l a r i a l o b a t a , and L a t i r u s s m a r w d u l a on dead c o r a l ;
-- -
and Morula d a t a B l a i n v i l l e , T h a i s h y s t r i x L., and Cy-xaee sp. c f . C.
--
$ o r a r i a L. on l i v i n g c o r a l heads. Mitra l i t t e r a t a , Mitra sp., Vasu;
t u r b i n e l l u m , and a h a l i o t i d a l s o occurred i n C2, 02, and Z2.

U n i d e n t i f i e d s p e c i e s of N a s s a r i u s and Mitra were c o l l e c t e d com-

p l e t e l y b u r i e d i n sand under c o r a l r o c k s i n u n i d e n t i f i e d quadrats.
 --
Conus a t .Funidu.
-- I am p r e s e n t l y p r e p a r i n g a more d e t a i l e d r e p o r t
on Conus i n t h e I " i d d i v e s , t o be published elsewhere, and w i l l t h e r e f o r e
o n l y b r i e f l y summarize t h e occurrence of t h i s genus a t F ~ m i d u . I n a l l ,
199 s p e c i n e n s of 16 s p e c i e s of Conus were col.lected. Conus was rnos,t
abundant on Seachrock, where average d e n s i t y w a s about one i n d i v i d u a l
p e r s q u a r e meter.

The Yale S e y c h e l l e s Z x p e d i t i o n , d u r i n g which t h e s e o b s e r v n t i o n s

were made, w a s s a d e p o s s i b l e by t h e g e n e r o s i t y of i?r. A l f r e d C. G l a s s e l l ,
Jr .
Literature cited

k g a s s i z , A. The c o r a l r e e f s of t h e :4alclives. i'.iern. ivius. Conip. Zool.

Hzrvard C o l l . , a: 1-168, 1?03.
Hoffraeister, J . X . , and H.S. Ladd. The a n t e c e d e n t p l a t f o r m theory.
J o u r . Geol. , 52: 3%-402, 1344.

Kohn, X.J. The ecology c o l l e c t i n g s a c k modified f o r marine orgaaisms.

Turtox Mews, 2:
33, 1956.
---------- The biology o f a t o l l s . B i o s , j2: 113-125, 1961.

Caption f o r map on f o l l o w i n g page:

Fuo;lidu I s l a n d and f r i n g i n g r e e f , North Male A t o l l , Maldive I s l a n d s

(4O10' N , 73'25' El. The substratum o f p o r t i o n s o f t h e r e e f f l a t examined,
and o t h e r f e a t u r e s , a r e i n d i c a t e d by symbols. Based on H.O. Chart 5664 and
o b s e r v a t i o n s by t h e a u t h o r , 19-23 September 1957.
 FUNIDU

ISLAND

smooth. sloping beachrak

with thin layer o f algae .'.'.
f l a t , smooth limestone
pavement with vergr
t h i n layer o f sand
living
CON
heads
a
t . dead coral
heads and
rubble

coral rubble apparently eroding tide tree

portion o f island pool @ stump