inventory of the puerto rican coral reefs

INVENTORY OF THE PUERTO RICAN CORAL REEFS

C . GOENAGA AND G . CINTRON

REPORT SUBMITTED TO THE COASTAL. ZONE MANAGEMENT OF THE DEPARTMENT OF, NATURAL RESOURCES

COMMONWEALTH OF PUERTO RICO

DOCUMENTATION PAGE DEPARTMENT OF NATURAL RESOURCES

- ?.ABSTRACT ( 2 0 0 WORDS OR LESS) An Inventory of 3 ~ e r t o Ricen reefs shows t h a t well developed reef sys t em a r e found o f f Fzjzrao (La CordLllera), ?;Iaunabo (Sargent Ree f ) , Bznia de Jobos and Santa I sabe l , Ponce (Ratones), L a ?argueray Lajas, Hayaguez ( T o u ~ a l i n e and E l Negro) and along the edge of ti.e southern and western in su la r she l f .

REPORT NUMBER

-T ITLE:

I n v e n t o ~ j of the h e r t o Rican Coral 3eef:;

AUTHOR f S)

Carlos Goenaga, Gilbert0 Cintrbn AGENCY FOR WHICH T HIS REPORT WAS PREPARED

Cff i ce of Coastzl Zone -3knagezent NCAA

STUDY OR PROYECT TITLE

z o s t Inshore and North Coast offshore reefs e r e degra- ded. -tenslve degradation was observed i n a l l r ee f s from San Juan t o Sas Cz.bezzs de San Juzn, the inshore 'aj'ardo -! heefs, I iu~zcao Xeefs, ree fs o f f i ue r to Yabucoz, inshore Tcnce r ee f s , a l l ree fs off 3ahia GuayanLllA and 3 a h h de TsllaSoa, a l l ree fs off and f r ing ing Guanica, a l l west coast inshore reefs and those ree's o f f Xrecibo and 2orado.

2- DATE:

28 Dec 1979

4 - T Y P E O F REPORT PRELIMINARY n FI NAL El O T H E R

?-PER100 COVERED BY THIS REPORT.

O C ~ 1978/oct 197.9 9-NUMBER OF PAGES

S i l t z t i o n due t o upland&fores ta t ion 2nd s o i l erosion seem t o be t h e most important s t r e s so r a f fec t - Puerto -7ican r ee f s . Other s t r e s s e s z3ect i -ng lnsnore r ee f s z r o cozs t a l eut rophicat ion and dredging. Coral ex t r zc t ion and boat znchor damge z r e addi t iona l but l o c ~ l i z e d problem.

Inventory of t he Puerto Zican Corzl Seeif 1'93 - >*KEYWORDS - - -

?uerto 3 i c o g / ~ o r z l XeeFs*/?rTnging 3eef/3ank Reef s/?.ibSon Beefs/3ock 3eefs/?atch 3eefs/Shelf edge/Spur and Groove systezs/Stressed r ee f s .

l e r i o d i c occurrence of storm ;lraves i s an tmportmt f a c t o r con t ro l l i ng reef s t ruc ture . Storm waves can cause

AIC.NO. 3 8 N O V . 7 9 - - .

extens ive and t h i s natural as s i l t z t i o n

widespread reef event may be impa and p o l l u t a n t s .

damage. . i r ed by

Xecovery of r ee f s from cron ic s t r e s s o r s such

KNOWLED GEMENT S

s t we want t o

i n v a l u a b l e h

e x p r e s

. e l p i n

s t h a n k s

t h e f i e 1

t o Miguel Nieves who

d and i n t h e p r e p a r a t i o n

of g r a p h i c m a t e r i a l . ?I &

4 I I T . .. - We a r e i n d e b t e d t o Miguel C a n a l s and Rober to C a s t r o

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

We t h a n k Frank T o r r e s and J a c k Morelock f o r p r o v i d i n g

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

Thanks a r e s p e c i a l l y due t o E l a d i o Rodr iguez and Gaspar

PErez, c rew o f t h e RV J e a n A , f o r t h e i r c o n t i n u o u s ' h e l p and . c o o p e r a t i o n d u r i n g a c r u i s e a round t h e i s l a n d f o r t h e

p r e p a r a t i o n of t h e i n v e n t o r y .

We want t o acknowledge J i m Trumbull and t h e U.S. Geo- - - , . . . - . l o g i c a l Survey f o r p r o v i d i n g . a e r i a l p h o t o g r a p h i c , m a t e r i a l . . .

Other p e r s o n s which a s s i s t e d u s a t d i f f e r e n t s t a g e s o f - -

t h e s t u d y a r e : Ram6n M a r t i n e z . L u i s ~ e g r ~ n , J o s e Gonzalez-

L iboy , Tommy Armstrong, R a f a e l Mosquera, Kur t Grove and

Gladys Rodrzguez, Annie Fon tgnez , Ana R i t a G o l d i l l a and

Amir ica Guzm5n typed d i f f e r e n t p a r t s of t h e m a n u s c r i p t and

H s c t o r B e r r i o s and Carmelo Montalvo d i d p a r t of t h e a r t work.

HEctor Merced d e v e l o p e d and p r i n t e d t h e p h o t o g r a p h s i n

t h i s r e p o r t .

Funding f o r t h i s p r o j e c t was p r o v i d e d by N a t i o n a l Ocean ic

and Atmospher ic Adminis t ra t ion , C o a s t a l Zone Management Program.

J:- ' .:. ,;.y&J: 'y: ;: ,I ;<;,; , :

TABLE OF CONTENTS , . - - - " -... ,,. .__. .- ~, . .. ,- . a I. *

WLEDGEMENTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . iii b

LIST OF FIGURES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . - e t v

LIST OF TABLES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ix

INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

METHODS AND MATERIALS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

WORLD WIDE CORAL REEF DISTRIBUTION . . . . . . . . . . . . . . . . . . . CORAL REEF DISTRIBUTION WITHIN THE ATLANTIC OCEAN . . . .

.. . - . ECOLOGICAL ZONATION . . . . ... . . : . . . . . .-..-. . .. :. . . ... . . . . . . . . . . . . . - - . . 7 .

1 3 ' .

Reef Apron . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 3

Reef f l a t . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

Reef c r e s t . . . . . . , . . . . . . . . . . . . . . . . .. . . . . . . ,.. . f . . . - 1 5 - . . ... - . -: -:

. , Acrppora z o n e . . . ; .... . ... . . . . . . . . . . . . . . . . . . . . . . . . . 1 5

B u t t r e s s - M o n t a s t r e a a n n u l a r i s zone . . . . . . . . . . . Reef f o r e s l o p e - g o r g o n i a n - head c o r a l zone . . 19

DIFFERENT REEF TYPES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rock r e e f . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F r i n g i n g r e e f . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P a t c h r e e f . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bank of r i b b o n r e e f s . . . . . . . . . . . . . . . . . . . . . . . . . . B a r r i e r r e e f . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A t o l l s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

ECOLOGICAL CONTROLS OF REEF GROWTH . . . . . . . . . . . . . . . . . . . P h y s i c a l and Chemical D e t e r m i n a n t s . . . . . . . . . . . . .

L i g h t i n t e n s i t y . . . . . . . . . . . . . . . . . . . . . . . . . . . T e m p e r a t u r e . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

TABLE OF CONTENTS .... (CONT.)

Salinity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Atmospheric exposure . . . . . . . . . . . . . . . . . . . Water turbulence . . . . . . . . . . . . . . . . . . . . . . . Sedimentation . . . . . . . . . . . . . . . . . . . . . . . . . . Storms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pollution

Upland deforestation . . . . . . . . . . . . . . . Sanitary effluents . . . . . . . . . . . . . . . . . Thermal effluents . . . . . . . . . . . . . . . . . .

' . Dredging ...........................

. . . . . . . . . . . . . . . . . . . . . . . . . . . . Bombing

. . . . . . . . . . . . . . . . .. Biological . Determinants . . .... .............. CORAL REEF DEVELOPMENT IN PUERTO RICO

. . . . . . . . . . . . . . . . . . . . . . . . . . . . SUMMARY AND CONCLUSIONS . . .. . . . . . . . . . . . . . RECOMMENDATIONS c . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . BIBLIOGRAPHY

APPENDIX . DETAILED TECHNICAL DATA . . . . . . . . . . . . . . . . . Tables . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Reef profiles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

LIST OF FIGURES

F i g u r e 1 D i v e r maki:: a n u n d e r w a t e r p r o f i l e . . . . . . . F i g u r e 2 L o c a t i o n of a e r i a l p h o t o g r a p h s . . . . . . . . . .

F i g u r e 3 Average w i n t e r and summer s u r f a c e i s o t h e r m s i n t h e A t l a n t i c Ocean . . . . . . . . . . . . . . . . . . . 11

F i g u r e 4 Average w i n t e r and summer s u r f a c e i s o t h e r m s i n t h e C a r i b b e a n Sea . . . . . . . . . . . . . . . . . . . . 11

F i g u r e 5 Dense M . - complana ta s t a n d on r e e f c r e s t . . 1 6

F i g u r e 6 Reef c r e s t . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

F i g u r e 7 Abundant c o l o n i a l anemones ( z o a n t h i d s ) i n t h e r e e f f l a t . . . . . . . . . . . . . . . . . . . . . . . . 16

F i g u r e 8 A . - p a l m a t a zone . Wote'100Qo .cpver . . . . . . . 1 7 . . - . .

F i g u r e 9 ' L a r g e A . p a l m a t a c o l o n y s h e l t e r i n g grunts -and g o a t f i s h e s . . . . . . . . . . . . . . . . . .

F i g u r e 10 A . - p a l m a t a s t a n d m a r g i n a l t o b u t t r e s s zone . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

I . -.. - -

F i g u r e 11, D i s t i n c t m o r p h p l o g i c a l s h a p e . ' d f t h e . . . . . . . . . . . . . . . . . . . . . . . . . . s t a g h o r n c o r a l

F i g u r e 1 2 M . a n n u l a r i s b u t t r e s . . . . . . . . . . . . . . . . . . . -

F i g u r e Dive r b u t t r e s s zone

F i g u r e 14 B r a i n c o r a l i n b u t t r e s s zone . Observe goby l y i n g o v e r t h e c o r a l ( m i d d l e ) . . . . . . .

F i g u r e 1 5 Colony o f t h e p i l l a r c o r a l Dendrogyra c y l i n d r i c u s . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

F i g u r e 16 Schoo l o f s u r g e o n f i s h i n t h e b u t t r e s s zone. Observe b u t t r e s s t a t h e r i g h t o f t h e p i c t u r e . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2

F i g u r e 17 L o b s t e r s s h e l t e r i n g w i t h i n c r e v i c e i n b u t t r e s s zone . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2

F i g u r e 18 L a r g e s t a g h o r n c o l o n y s h e l t e r i n g a h a m l e t . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 3

LIST OF FIGURES (CONT) Page

r e 19 Gorgonian o r s o f t c o r a l i n t h e f o r e . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r e e f s l o p e 23

,F igu re 20 Sponge and mass ive c o r a l s i n t h e f o r e . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r e e f s l o p e 26

F igu re 2 1 T u r t l e i n t h e f o r e r e e f s l o p e . . . . . . . . . . . ': 26

F igu re 2 2 V e g e t a t i o n denuded a r e a ("halo") around r e e f o r rock ou t c rop . . . . . . . . . . . . .

Figu re 23 L ine o r rock r e e f s (Trending e a s t t o wes t ) o f f s h o r e San Juan . . . . . . . . . . . . . . . . . 4 0

F igu re 24 Heav i ly s i l t e d r e e f s o f f Punta I g l e s i a s and Punta San Agust in ( c z n t e r ) . Punta Vacia Talega can be s een a t t h e l e f t

. . . . . . . . . . . . . . . . . . . . . . . . . . . o f . t h e p i c t u r e 4 1 . . .

~ i ~ u i e 2.5 ringing r e e f s o f f punt%. M i q u i l l o and Punta Picua . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43

F igu re 26 Punta Percha w i t h f r i n g i n g r e e f p r o t e c - t i n g L u q u i l l o Beach . . . . . . . . . . . . . . . . . . . . . 46

Fi-gGre. 27 Reef f r i n g i n g aria from ~ a b e z a Chi-qui ta . '

. . t o Cabo San Juan . . . . . . . . . . . . . . . . . . . . . . . . * 48

F igu re 28 I c a c o s westernmost i s l e t of La C o r d i l l e r a - 4 9

F i g u r e 29 Palominos and Pa lomin i tos ( s o u t h e r n . . . . . . . . . . . . . . . . s a n d k i s l e t ) o f f Fa j a rdo

F igu re 30 Cayo Largo, o f f Fa j a rdo . . . . . . . . . . . . . . . . . ' Figu re 31 I s l e t s . a n d - r e e f s o f f Fa j a rdo . From t h e

bottom I s l a d e Ramos, .Cay0 Ahogado and - P s i e t a Marina (Cayo Obispo and Cayo Zancudo) a r e s een . . . . . . . . . . . . . . . . . . . . . . . .

Figu re 32 Reefs p r o j e c t i n g e a s t o f Punta Bar rancas and Punta Mata Redgnda . . . . . . . . . . . . . . . . . . .

F i g u r e 33 I s l a Pifieros and Cabeza d e P e r r o o f f Medio Mundo, Ceiba . . . . . . . . . . . . . . . . . . . . . . .

F i g u r e 34 Cayo San t i ago (upper r i g h t ) and Cayo B a t a t a ( c e n t e r ) o f f Humacao. S e v e r a l submerged p a t c h r e e f s , i n c l u d i n g Bajo P a r s e , c an be s een s o u t h of Cayo S a n t i a g o . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . -

LIST OF FIGURES (CONT . ) Page

p 6.4 I . . . . . . . F i g u r e 3 5 a ) Annular r e e f o f f Yabucoa Bay

b) Non v e r t i c a l a e r i a l p h o t o g r a p h o f t h e same r e e f . . . . . . . . . . . . . . . . . . . . . . .

F i g u r e 3 6 F r i n g i n g r e e f o f f P u n t a F i g u e r a s . . . . . . . F i g u r e 37 A r r e c i f e Guayama s o u t h of Pun ta F i g u e r a s

F i g u r e 38 A r r e c i f e Las Mareas o f f Guayama ( lower r i g h t ) . . . . . . . . . . . . . . . . . . . . . . . . . .

F i g u r e 3 9 Cayos d e Barca . J o b o s . . . . . . . . . . . . . . . . . . F i g u r e 4 0 Reefs o f f S a l i n a s . . . . . . . . . . . . . . . . . . . . . . . F i g u r e 41 Reefs o f f S a n t a I s a b e l . . . . . . . . . . . . . . . . . F i g u r e 4 2 Caya Berce rFa o f f S a n t a . I s a b e l .......:. . . 7 1

F i g u r e 4 3 I s l a C a j a d e Muer tos o f f Ponce . . . . . . . . . F i f u r e 4 4 Cayo o f f Ponce . A r r e c i f e Ra tones c a n

be s e e n a t l e f t . . . . . . . . . . . . . . . . . . . . . . . . ..

F i g u r e 4 5 ' S t r e s s e d ' i e e f s o f f ' T a l l a b o a . . . . . . . . . . . . . F i g u r e 4 5 a S h i p s t a t i o n e d be tween Cayo RZo and

Cayo Palomas i n T a l l a b o a Bay . . . . . . . . . . . - F i g u r e 46 Reef s o f f P u n t a V e r r a c o . G u a y a n i l l a . . . . F i g u r e 4 7 F r i n g i n and p a t c h r e e f s o f f Gugnica . . . . F i g u r e 48 Reefs p r o t e c t i n g P l a y a Cafia Gorda. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gu5nica

F i g u r e 4 9 La Parguera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F i g u r e 50 P a t c h r e e f s o f f I s l a Cuevas . . . . . . . . . . . . . F i g u r e 51 E l P a l o . A r e v e s a b r e e f . . . . . . . . . . . . . . . . F i g u r e 5 2 M a r g a r i t a r e e f . . . . . . . . . . . . . . . . . . . . . . . . . . F i g u r e 5 3 San C r i s t 6 b a l r e e f . . . . . . . . . . . . . . . . . . . . . F i g u r e 54 L a u r e l r e e f . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F i g u r e 5 5 Media Luna r e e f . . . . . . . . . . . . . . . . . . . . . . . . . F i g u r e 56 'Enr ique r e e f . . . . . . . . . . . . . . . . . . . . . . . . . . . .

LIST OF FIGURES (CONT . ) " > * ' , < *

* . . . A . &sk*w-&-; *-

~ F i g u r e 57

' F i g u r e 58

F i g u r e 59

F i g u r e 60

F i g u r e 61

F i g u r e 6 2

F i g u r e 6 3

F i g u r e 6 4

. . F i g u r e - 6 5

F i g u r e 6 6

F i g u r e 67

F i g u r e 68

F i g u r e 69

La Gata - C a r a c o l e s r e e f . . . . . . . . . . . . . . . 8 9

Enmedio r e e f . . . . . . . . . . . . . . . . . . . . . . . . . . . 90

Mario r e e f . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 1

.I Spur and g r o o v e sy s t em. Observe h i g h q

c o r a l c o v e r . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 4

Idem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 4

Idem. Note whip a n t i p a t h a r i a n s (black coral) 9 5

Diver work ing a t t h e s h e l f edge . . . . . . . . 9 5

Low r e l i e f sand c h a n n e l s n o r t h o f t h e s h e l f edge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97 . S h e l f edge . Note h i g h ' c o r a l cove r . . . . . : -97 . . .

F r i n g i n g r e e f s o f t h e wes t c o a s t . . . . . . . 100

Fr i ng ing -ba r r i e r r e e f o f f Boquer6n . . . . . . 101

. O f f s h o r e r e e f s o f t h e wes t c o a s t . S q u a r e -- . ,marks o f f s p u r .and g roove sys tem . . . . . . . . . 1 0 2

B a j u r a c a v e s . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 0 6

C u t e r w a l l s of t h e , B a j u r a c a b e s . Observe d e n s e A g a r i c i a g rowth . . . . . . . . . . . . . . . . . . 1 0 6

F i g u r e 7 1 Large dead A . pa lma ta c o l o n y i n s i d e c a v e . I t s o r i g i n r ema ins t o be s t u d i e d .. .. . . . .

F i g u r e 7 2 Reef o f f Dorado . . . . . . . . . . . . . . . . . . . . . . . . . F i g u r e 7 3 P r o f i l e N o r t h e a s t C o a s t Cabo San J u a n . . . F i g u r e 7 4 P r o f i l e I s l e t a Marina . . . . . . . . . . . . . . . . . . . . F i g u r e 7 5 P r o f i l e Cayo Zancudo ( N o r t h ) . . . . . . . . . . . . . F i g u r e 7 6 P r o f f l e Cayo Ahogado . . . . . . . . . . . . . . . . . . . . F i g u r e 7 7 P r o f i l e P a l o m i n i t o s (Sou th) . . . . . . . . . . . . . F i g u r e 7 8 P r o f i l e Cayo Largo . . . . . . . . . . . . . . . . . . . . . . F i g u r e 7 9 P r o f i l e Ramos (West) . . . . . . . . . . . . . . . . . . . .

LIST OF TABLES (CONT .)

T a b l e 24 E n r i q u e . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146

T a b l e 25 L a u r e l . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147

T a b l e 26 T u r r u m o t e I .. . . . . . . . . . . . . . . .. . . . . . . . . 148

T a b l e 27 La Ga ta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149

T a b l e 28 M a r g a r i t a . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150

T a b l e 29 Las P e l o t a s . . . . . . . . . . . . . . . : . . . . . . . . . . 1 5 1

T a b l e 30 Cayo Ahogado . . . . . . . . . . . . . . . . . . . . . . . . . 152

T 'ab le 31 Media . Luna ...-., .... -...; . . . . . . . . . . . . . . . . - - 7 . , .

T a b l e 32 La Conse rva . . . . . . . . . . . . . . . . . . . . . . . . . . T a b l e 3 3 C o l l a d o . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 5 5

T a b l e 3 4 B a j o Enmedio . . . . . . . . . . . . . . - , . ... . . .,. . . . - 1 5 6 - , . ._. -

- % - -

. , , T a b l e 35 G u a n i q u i l l a . . . . . . . . . .". . . . . . . . . . ; . . . . . 157

T a b l e 36 P u n t a O s t i o n e s . . . . . . . . . . . . . . . . . . . . . . . 158

T a b l e 37 R a t o n e s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159

The importance o f t h e c o r a l reef e c o s y s t e m has been

mentioned and documented on numerous occasions by several

writers. In summary:

a) Reefs are among t h e n o s t b i o l o g i c a l l y produccix7e

e c o s y s t e m s c o n t a i n i n g a g r e a t variety of benthic

organisms and p r o v i d i n g a habitat for large num-

b e r s of juvenile f i s h of Eany species. Reefs

shelter and support t he majority of f i s h and - - , crustaceans that are commercially extracred f rom

our coastal processes.

b) Reefs prov ide a buffer against s;as pounding

shorelines and moderate currents- &ereby i n f l u e n -

c i n g the d e p o s i t and maintainance of sand on

beaches. A s an example, the calm w a t e r s and d q o -

s i t ed sands of Luquillo Beach would be l o s t if

the protecting reefs were to dksappear.

c ) Fragments from dead c o r a l a r f r o m o t h e r calcium

carbonate producing organisms which inhabit r e e f s

a r e t h e p r i n c i p a l components o f many Pue r to RFcan

beach sands. Yany i s l e t s are also formed by the

d e p o s i t i o n of these .

d ) Hany organisms inhabiting r e e f s , suck as a l g a e

and c e r t a i n s o f t corals, produce chez ica l subs-

rances with valuable medicinal properties.

e ) Reefs are a recreational -resource f o r s k i n and

SCLXA d i v i n g .

f! Due t o t h e fact , c?,at r e e f s a r e t 5 e most exten-

s i ve coas ta l comrnmicies of the earth and due

t o t h e i r complexity in terms of taxonomical

diversity and t ropP. ic relations, these a r e of

unlimited Interest in the study of t h e dynamic

relations o f biological processes .

The i n t e r e s t o f t h i s r e p o r t i s to 'g ive a g e n e r a l des-

cri?rion, s t a t u s and localization of our reef resource

- and to stimulate in rhe beginner s c i e n c e students the : . -

curiosity and d e s i r e t o initiate e c o l o g i c a l studies of

t h i s most impor tan t and threatened ecosys tem. Ke a l s o

attempt t o c r e a t e a consc iousness o f t h e a l a r m i n g ra te .- .. I '

o f degradation o f t h i s practically non-renewable r e s o u r c e

i n - t h e layman and to promote r h e i r r i g h t t o ~ublicly oppose

p r o j e c t s which may further endanger t h e l a t t e r . F i n a l l y ,

ke i n t e n d t o d e l i n e a t e g ~ i d e l i n e s , based on t h e ana lys i s

of o u r and o t h e r s ' observations, f3r the managemen t o f

t h e coral r e e f s in the b e s t soc ia l i n t e r e s t .

T h i s document initially shows c o r a l r e e f disiribution

in t h e w o r l d and e s p e c i a l l y in t h e Caribbean Sea . Folloui::

is a mention of ?he physical and b i o l o g i c a l conditions

n e c e s s a r y for t h e development of c o r a l reefs . Then, the

most common zonation p a t t e r n s and the different r e e f types

are discussed. Reef distribution a l o n g t h e Puerto Rican

c o a s t s is descr ibed based on published literature, r e p o r t s - and per sona l observations. Y e x t ale ou: ~oncLusions and

r e c o m r n e n d a t i ~ r . ~ . Includ-::.l 2s a p p e n d i c e s a r e profiles o f

s e l e c t e d r e e f s and t a b l e s c o n z a i n i n g detailed d a t a .

F i g . 1. 3iver =king an w.derwa2er p r o f i l e . Observe sea fans In gorgocian zone.

YETI-rJDS Ah4 MATERIALS

Tkis r e e f i n v e n t 0 7 was c a r r i e d out d . l r i ng a one

y e a r p e r i o d . O S s e n - a t i o n s were l i m i t e d to a maximum of

one day p e r r e e f and measurements were made o n l y whenex-er

sea c o r - d i t i o n s , e . g. PLlgh water transparency and reduced

surge and wave a c t i o n , p e m i t t e d it.

T o p o g r a p h i c profiles o f t h e f o r e r e e f were made w i t h

a l i n e marked every f i v e a e t e r s . This ' l i n e vas l a i d ,

per7endicular to t h e s h o r e , from the c l o s e s t area of the

.- r e e f f l a r t h a r sea conditions permitted t o the win=ward

base o f tb,e r e e f . D e ~ t h measurements and observations

were made at five meter-intervals ( o r l e s s where p e ? t i n e n t )

, . .. . . and recorded on an u n d e m a t e r s l a t e (Fig. 1). S e l f con-

t a l n e d undewaier b r e a t h i z g appa ra tus ( S C L W - ) was used

on areas where use of a snorkel was n o t a p p r o p r i a t e .

The site o f t h e profile uas chosen af r e r -ana lyz ing

a e r l a l photography o f t h e area ar.d after aaking under-

water reconnaissance o f t h e ree f . Occasionally a diver

'%as toned on XI underwater s l e d t o cover l a r g e r areas.

The area most representative of t he f o r e r e e f was s e l e c r e d .

Coral spec i e s diversity, equitability and livizg

cover were made with Line trahsects according to a mechod ,

designed by Loya ( 1 9 7 2 ) and l a t e r modified by Rogers ( 1 9 7 7 )

.41: transeccs were IG m long and paraliel t o depth contours

These xere run with t h e a i d of SCUBA.

S l a c k and white photographs were taken with a Nikonos 'C

I11 c m z r a with a 35 mrn l e n s .

Reefs where p r o f i ! were n o t m a z e due to diverse

reas2ns , a r e descr ibed :- t h e Cora l ? e e f Development in

Pue r to R i c o section.

A 11 aerial pho tography is v e r t i c a l unless o t h e r d i s e

s p e c i f i e d . T h e l o c a i i t g and a r e a o f cover i s shown i n

F i g . 2 . The approxiaate a r e a s where underwater profiLes

w e r e made is shown with a line. Corresponding s e c t i o n s

o f nautical char t s a re included w i t h each v e r t i c a l a e r i a l

photograph.

Available n o r t h c o a s t a e r i a l p h o t o g r a p h y was analyzed

and areas where c o r a l growth was suspected w e r e spot-checked

in the f i e l d w i t h the a i d o f a boat t o w e d underwarer sled.

The s e c t i o n Detailed Reef Information includes: -

The f o r e r ee f p r o f i l e s the distribution

t h e m o s t common s p e c i e s within each zone a d 2 ) Tables

mentioning specific details of the zones o r of the reef

as a whole.

The c o r a l reef inventory does n o t include the i s l a n d s

o f Xona, Vieques, Culebra and Desecheo. For i n fo rma t ipn

on t h e reef fauna of the f i r s t t h r e e t h e reader is r e f f e r e d

t o ~ i n t r b n and Thurston ( 1 9 7 5 1 , T o r r e s ( 1 9 7 2 ) and Cin t rbn

e t a1 (19747 respectively. -

REEF CLSTRIEUTION

Herma.c;.-pic o r reef building c o r a l s , which are t h e

m a i n components of the coral r e e f , f l o u r i s h in t r o p i c a l

zones with seawater temperatures r a n g i n g from 2 5 - 2 9 ' ~ .

These areas I r e situated within a belt rough ly bounded

by t h e T r o p i c s of Cancer and C a p r i c o r n , imag ina ry lines

which are drawn aromd t h e e a r t h 23.5' north and s o u t h

of the equaror. Vithin this 4800 k i l o m e t e r - w i d e b e l t ,

cora l r e e f s are abundant on the eastern shores of the

Americas, A f r i c a and A u s t r a l i a , but f a r less common, o f f

t h e western shores of these c o n t i n e n t s . k'hy is t-his? I n

the Northern Hemisphere ocean currents a r e fo rced into

a c l o c k w i s e circular movement by t h e earth's r o t a t i o n . ..

- and a s s o c j a t e d wind distribution ( C c r i o l i s effect). The

oppos i te is true in the Sou the rn Hemisphere. A s a result,

w a r m water--travels toward t h e p o l e s along t h e e a s t e r n shores

of t h e c o n t i n e n t s . On these s h o r e s , t h e r e f o r e , there i s

a much wider extension of warm water su i t ab le f o r v i g o r o u s

coral growth. On t he wes te rn shores t h e reverse is t rue.

Cold waters running toward the equator, combined w i t h

upwellings of c o l d water from t h e d e p t h s , g r e a t l y r e s t r i c t

the extent of shoreline favorgble to c o r a l growth.

Yon hermatypic or non-reef building c o r a l s , contrary

to hermatypic ones, are by no means restricted to t h e t r o p i c s

a d can be found in the c o l d s e a s lying within t h e A r t i c

and Antartic c i r c l e s as w e l l as in t h e Nomegiar, i j l ? r d s ,

, ,..#? -r rocky shares of the United States, Canada,, zr:' t h e

c e a s t s of England and France. ?lone cf these co ld -wa t e r

corals, however, grow so actively o r to such a s i z e as

:tLe I z r g e r reef c o r a l s of t h e t r o p i c s . Their form is

us~ally sm.z!lI and d e l i c a t e rather than l a r g e and massive

!Smith, 19L8) .

DISTRIBUTION W I T H I U T I E ATLA." IC OCEAN

Mast c o r a l r e e f development in t ? ~ Atlantic Ocean

is r e s t r i c r e d to t he Caribbem and adjacent areas in the

southern Florida and t h e Bahamas. Altk-ough v i a b l e reef

producrion is nil1 n n r t h o f sou thern fiorida (due to low

winter temperatures) , i s o l a t e d h e r m a t g i c corals can

survive as far n o r t h as Cape Hatteras ( 3 5 N , see average

w i n t e r and s m e r temperatures in t h e 4 t l a n t i c Ocean and

the Caribbean in F i g s . 3 , 4 ) . One n o t a b l e exception is

Bermuda which conta ins a s u r p r i s i n g l y large number o f

corils, but s t i l -1 lacks some of the most p r o l i f i c West

Indian s p e c i e s such as A c r o ~ o r a palmata (stoddart, 1 9 6 9 1 . A L S O , these r e e f s appea r to be on ly tPLin encrustations

over ~ l e i s t o c k n - e rock (Mil1iman;- -1973) .' The Gulf oE

Mexico is b a s i c a l l y an area of terrigenous sedimentation,

but some scattered marg ina l r e e f growth is p r e s e n t n e a r

V e r a c r ~ z , Mexico, together vith many r e l i c t co ra l and

a l g a l mounds thrqughout much of t he Gulf shelf (Milliman,

1 9 7 3 ) . The n o r t h e r n p o r t i o n of t h i s area, roughly that '3

area north of a l i n e passing through Progreso , H e x i c o , b

and Havana, Cuba, can be considered to be marginal tropics

in t h a t winter temperatures c~mmonly fall be lov 1:'~ and

coral reef development is poor as compared ~ 1 r h t h e central

and southern Caribbean.

Coral r e e f development i n t h e Atlantic Ocean r eaches

i t s southern l i m i t o f f Rio d e J a n e i r o , B r a z i l . These

Cornunities a r e , however, b i o l o g i c a l l y quite distinct from

those of the Caribbean and, as in Sermuda, l a c k many 2'

Fi(. 3 A r r n g e r+?!rc (-) and ~urnmrc ( - - ) ~ u t f s c r Isnrhrrm~ ( ' C ) In lh AfLntlc C h i n (s l trr S-crdmp n d., I S I ? ) . A m u r i !h prclr.nbrd r r ;uurn lo

Ionpr.turn Irrs I'M Z2.C a r i t h ,!me rrpmutn I r u ~ h n 18'. at brrt r l U hsvc a &;.tupnle ml p p w l ~ t i ~ m . I'rnodac InIlr?sr~ ol m l d r . s 0 . r s from 1l.e Cuinu

Cuncut &A-r hclird to wrr,rlr 1;rnil rm-1 Jcrrlcrymcnt eE tropi-l hltica. ,

rincipa

commun

For a d

.s of the lat

.ry of coral

ter (Volcker,

reef research "* ,i.the Caribbean, the reader is referred to Colin (1978).

"9:. *

E C O L O G I C A ~ ZC, QTPOW - ~ < ~ ~ ~ G -4 5

I d e n t i f i a b l e e c o l o g i c a l zozes a r e c r e a t e d by t h e

degree t o which t h e v a r i o u s p h y s i c a l and topograph ica l

f a c t o r s p o s i t i v e l y o r n e g a t i v e l v a f f e c t r e e f organisms

throughout t h e r e e f a r e a . The c h a r a c t e r i z a t i o n of each

zone may, t h e r e f o r e , be based on t h e name of one key o r -

ganism and /o r t h e name of t h e p h y s i c a l f a c t o r o r f e a t u r e

e s t i m a t e d t o be of g r e a t e s t importance t o t h e ecology of

t h e zone. Overlap o r combination of zones may occur

anywhere as a r e su l t ' ' o f t h e v a r - i e d e f f e c t o f t i d a l c u r r e n t s , . - 7 .

wave a c t i o n o r bottom s t m c t u r e .

Following i s a g e n e r a l scheme of r e e f zona t ion ( l e e -

ward to- windward) . I t should be borne i n wind t h a t t h i s I ..> - . - -

v a r i - e s from l o c a t i o n t o l o c a t i o n - a n d even w i t h i n t h e .

same a r e a :

( a ) Reef apron - The r e e f apron i s t h e a r e a of sediment accumulation

leeward of t h e r e e f f l a t . I t i s g e n e r a l l y a ba r ren a r e a

i n t h e sense of e p i b e n t h i c b i o t a a l though i n some c a s e s

i t i s e x t e n s i v e l y burrowed by shrimps o r h o l o t h u r i a n s

( s e a cucumbers) which form mounds t h a t sometimes cover

most of t h e b a r r e n sand a reas ' (Mathews, 1974) . I t may

a l s o c o n t a i n beds of marine phanerogams ( f lower ing p l a n t s )

such as t u r t l e g r a s s ( T h a l a s s i a ) and/or pa t ch r e e f s .

Sand i s p r i m a r i l y b iogen ic as i n t h e case of Cayo Enr ique ,

L a Parguera , where i t s p r i n c i p a l c o n s t i t u e n t s a r e mainly

and cora1lb:~e algae fragments (Morelock et al,

(b) Reef flat - low wave action zone This is a shallow area of sand and rubble that lies.,

behind the reef front. The depth may vary from 0 to 1 m.

Usually even though wave pounding is minimal, a very

strong current is created by the outrushing of the water

accumulated by incoming waves which come over the reef

front.

. Durjng evere tropical storms, large quantities of - _ . - - 1 .

llvkg coral are dumped into this area forming islets of

considerable heights above sea level.

As depth decreases ,in his area, an important litto- - . . . . ral community-,--the mangrove forest, may become established

creating a new habitat for many species of birds, fishes -

and invertebrates. Mangrove trees may, at the same time,

influence the surrounding reef habitat by providing a

nutrient subsidy by decomposition and exportation of leaves.

Corals present in thss area are generally sturdy

encrusting and A l l branching such as Porites asteroides,

Porites porites and, to a lesser degree, Manicina areolata,

Favia fragum, Diploria clivosa, Agaricia aparicites, and a - Siderastrea radians. Corals present in this zone are

sometimes, as in the case of 2. radians, very tolerant

to sedimentation and their distribution is influenced

by the presence of adequate substrates, such as coral -

fragm&&, which favor t f i e i r establ ishment . .'J

Thalass ia beds and t h e i r a s soc i a t ed b i o t a a r e a l s o

common i n t h i s a rea .

( c ) Reef c r e s t ; Mil lepora - zoanthid zone; Breaker

zone (F ig s . 5 , 6 ) .

This i s t he zone of h ighes t wave energy and water 1 movement. Prominent organisms inc lude Mil lepora compla- 1 na t a ( f i r e c o r a l ) and s eve ra l spec ies o f -zoan th ids (co- I - l o n i a l anemones) (F ig . 7 ) . Sometimes t h i s zone extends

above t h e low t i d e l e v e l forming emergent c o m m ~ i t i e s . . Under s l i g h t l y lower energy regimes ~ i l l e p o r a may be

replaced by t he e lk horn c o r a l Acropora palmata.

Other organisms, very conspicuous i n t h i s a r e , inc lude . , - . -.. -

t he encrus t ing gorgonian E,wthropodium c a r i b a e o m , va r ious . .

species of ca lcareous red a lgae and sea urchins . - Even though a d i scuss ion of t h i s sub jec t i s out of

t he scope of t h i s r e p o r t , should be mentioned t h a t

recent s t u d i e s have confirmed the ex i s tence of emergent

a lgae r i dges , similar t o those p resen t i n severa l Indo-

Pac i f i c r e e f s which rep lace t h e Mil lepora zone (Glynn,

1976). This a l g a l r i dge i s b e s t developed i n a r e a s of

very h igh wave energy.

( d ) Acropora zone

This zone extends seaward from the bottom edge of

the Mil lepora zone and gene ra l l y i s monospecific con ta i -

n ing, almost exc lus ive ly , t he e lk horn c o r a l (F igs . 8 , 9 , t

1 0 , ) Various forms of t h i s c o r a l a r e present depending

f i g . 6. Reef c r e s t . ez

Fig. 5. 2ense M.

of reef f l a t i n t h e background.

F ig . 7. Abundant c o l o n i a l anemones (zoanth ids) i n t h e reef f l a t . Severz l black urchins (Diadem a n t i l l a r u m a r e p re sen t i n the background.

~ i g . 8. almata zone. Note 100$ cover and high ver- t i c h e f . 7 .

Fig. 9 . Large colony of A . pa1,yt; shel te- ring g m s haemulon and goat f i shes (Mullo dichthys sp . )

SP - 4 ' A -

Fig. 10. - A. palmata stand marginal to buttress zone.

Fig. 11. Distinct morphological shape of the staghorn coral (A. - ~almata) .

on t h e degree of exposure.

Other c o r a l s and d i f f e r e n t a lgae spec ies may grow

on t h e p ro tec ted s u b s t r a t e beneath t h e branches of t h e

e l k horn c o r a l .

* ( e ) B u t t r e s s - Montastrea a n n u l a r i s zone

Seaward of t h e Acropora zone t h e r e i s usua l ly a

small drop of f where massive heads o r b u t t r e s s e s of the

s t a r c o r a l Montastrea a n n u l a r i s occur (F igs . 1 2 , 13) .

Very l a r g e i s o l a t e d colonies of t h e e l k horn c o r a l a r e

also:pr-e;en.t t h i s c r e a t i n g an area, -of very high r e l i e f .

S o f t c o r a l s o r gorgonians, b r a i n c o r a l s (Fig. 1 4 ) and

p i l l a r c o r a l (Fig. 15) a l s o abound here .

- , . . T.his a r e a wi th a l l i t s - s t d e p and narrow channels , . --

canyons, tunne l s and towering c o r a l heads provide a g r e a t

v a r i e t y of h a b i t a t s f o r reef organisms (F igs . 1 6 , 17) and -

i s genera l ly t h e a r e a of h ighes t b i o t i c d i v e r s i t y . i a r g e

patches of A_. c e n r i c o r n i s s h e l t e r numerous f i s h (18) .

( f ) Reef f o r e s lope - gorgonian - head c o r a l zone

Beyond t h e b u t t r e s s zone, t h e r ee f f l a t t e n s out

towards i t s ' ~ a s e . Gorgonians (Fig. 19) a r e u s u a l l y t h e

most f requent l i f e form i n t h i s a r e a , sometimes forming

dense f o r e s t s . Between t h e gbrgonians encrus t ing cora l

such as Montastrea cavernosa, M_. a n n u l a r i s , S i d e r a s t r e a

s i d e r e a , D i p l o r i a l a b y r i n t h i f o m i s and P o r i t e s a s t e r o i d e s

occur. Other c o r a l s present a r e I s o p h y l l i a m u l t i f l o r a ,

I ~ o p h ~ l l a s t r e a r i ~ i d a , Mycetophyllia lamarkiana, and

Fig. 12. - M. arinularis bGttress. Note laFge quantities of planktivorous fish. Transect line can be seen in the background.

Fig. 13. Diver making observations in buttress zone. Observe large variety of invertebrates.

Fig. 14. Brain zone. Observe cora l .

c o r a l ( A , cerv icorn i s ) and a gorgonizn t o t h e r i g h t of t h e - photograph.

c o r a l i n buttress*" goby ly ing on t h e

'ig. 15. Colony of t he p i l l a r c o r a l Dendropyra cy l lndr lcus . Note l a rge stand of t h e elkhorn

Fig. 16. School of surgeon fish (Acanthurus sp.) in the buttress zone. Observe buttress to the ri:ht - of the picture.

Fig. 17. Lobsters (Panulirus argus) wheltering within crevice in the butfress zone.

- F t g . 18. .-Large s ta o rn colony shel ter- a hamlet - i (Hypoplec r u s - s p . )

Flg. 19. Gorgonlan o r s o f t c o r a l I n t h e f o r e reef s lope . Observe t h e dominance of gorgo- n lans over s c l e r a c t i - nlans (hard c o r a l s ) i n t h i s zone.

mssa a n g u l o s a . Sponges a r e a l s o common h e r e ( F i g . 20) . - &.I* 5

A g r e e n t u r t l e was o b s e r v e d i n t h i s zone ( F i g . 21) . U s u a l l y ,

corals growing n e a r t h e b a s e of t h e s l o p e a r e h e a v i l y s i l t e d

and dead c o l q n i e s a r e f r e q u e n t .

Small p a t c h r e e f s sometimes occur beyond t h e base of . t h e s l o p e . I n a r e a s where t h e r e e f b a s e i s s h a l l o w e r t h a n

about seven m e t e r s a c o n t i g u o u s s e a g r a s s bed i s p r e s e n t .

Normally a "halo" o r non v e g e t a t e d s t r i p e o c c u r s a d j a c e n t

t o t h e r e e f b a s e a s a consequence of r e e f f i s h g r a z i n g

(See F i g . 22).. .

DIFFERENT REEF TYPES

a) Rock r e e f

Rock r e e f s a r e shal low e o l i a n i t e p la t forms t h i n l y

veneered by s tony c o r a l s . These w i l l be d i scus sed upon

the d e s c r i p t i o n of Pue r to Rican n o r t h c o a s t .

b) Fr ing ing r e e f

Fr ing ing r e e f s grow marginal t o t h e c o a s t and a r e

separa ted from t h e l a t t e r by a shal low lagoon g e n e r a l l y no t

exceeding a couple of meters and sometimes n e a r l y exposed

t o t h e atmosphere. . . .

t ) Patch r e e f :

These a r e i s o l a t e d c o r a l c o l o n i e s u s u a l l y surrounded

by a sandy bottom and occur ing c l o s e t o shore . They a r e - ,

' i rYegflar i n shape. . - > - - - . . . . .

d ) Bank or r i bbon r e e f s

Bank r e e f s a r e developed on c a l c a r e n i t e c u e s t a s o r

on drowned s y n c l i n e s . These w i l l be d i scus sed i n d e t a i l i n

the d e s c r i p t i o n of t he southwest c o a s t .

e ) B a r r i e r r e e f

These a r e s u a l l y emergent r e e f s s epa ra t ed from and

by a deep and :ride lagoon.

f ) A t o l l s

A t o l l a r e ' o v a l hsaped r e e f s r i s i n g from deep water

*nd surrounding a lagoon i n which ahere i s l i t t l e o r no l a n d .

deve lop , - u s u a l l y bu t not a lways, on t h e g r a d u a l l y sub-

s id ing cones of e x t i n c t vo lcanoes .

Fig. 20. Sponge ( c e n t e r ) and massive co ra l s ( fo r e and background) I n the fo re reef s lope .

Pig- 21. Tur t le (probably Chelonia mydas) I n the fo re reef s lope. -

41) FOTOGRAFIA AEREA

( FRANJA S I N VEGETACION I

I I I I

I NTERPRETACION I I

I * .--- I

i - . 1 ,

* . - - . . I . . I , .

1 I I 7

PLANlClE ALGASEA

Fig. 22: -

p R 4 * ~ 4 DESNUDA D E VEGETACION ALREDEDOR DE UJ' A F L O R A w I E N T O R O ~ 0 3 0 SUBMARINO. FENOMENO CAUSADO P O R EL- P A S T O R E O INTENSIVO DC L O S PECES R E S ~ D E N T E S Y TURBULENCIA (a f te r ~ u g o , 1978).

AFLORAMIENTO - 4

. > - -

B a r r i e r r e e f s and a t o l l s a r e no t found w i t h i n t h e - - " 5 '

P u e r t o Rico s h e l f o r i n i t s v i c i n i t y a l t hough t h e s e t ypes

a r e found i n t h e Car ibbean . The b e s t known and l a r g e s t

b a r r i e r r e e f i n t h e Car ibbean f l a n k s t h e c o a s t o f B r i t t i s h

Honduras ( B e l i z e ) which s t r e t c h e s f o r more t h a n 200 km and . has lagoon d e p t h s o r more t h a n 20 m. The b e s t known Caribbean

a t o l l s a r e Hogsty Ree f , Alacran Reef , L igh t House Ree f ,

G l o v e r ' s Reef and S e r r a n a Bank. These a r e found o f f t h e coast

of Honduras, B e l i z e and t h e Yucatgn Pen insu l a .

These d e f i n i t i o n s should be used c a r e f u l l y due t o t he - common occu r r ence of i n t e r m e d i a t e f8;ms i y

ECOLOGICAL CONTRES OF REEr GROWTH

The environmental con t ro l s of l oca l cora l growth

and of general reef d i s t r i b u t i o n have been summarized

by Stoddart (1969).

Physical and chemical determinants

( a ) Light I n t e n s i t y

Most hematyp ic ( reef building) cora l s grow

a t depths of l e s s than 25 m and maximum growth r a t e s

appears i n depths l e s s th& 1 0 - - m . T h i s has been explained . - . . . i n terms of t h e ' incredsing concentrat ion of suspended

sediments with depth (Wood-Jones, 1910) but the c r i t i c a l

control seems t o be i l lumina t ion (Gard-iner, 1930). _ This - , .._. - . ,decrease i n l i g h t i n t e n s i t y with- depth l i m i t s photosyn-'

t he s i s by the symbiotic a lgae present i n cora l endodemal - t i s s u e which play an important r o l e i n the process of calcium

deposi t ion.

I t should be pointed out here t h a t recent observa-

t i ons of deep reef s lopes by means of submersibles have

shown t h a t c e r t a i n species of hermatypic co ra l s extend

up t o 100 m depth though i n general , they cease t o be

framework const ructors a t about 70 m (Lang, 1974; Colin,

1974; Lang e t a l . , 1975; Ginsburg and James, 1973, 1976;

NeUmann and B a l l , 1970; Po r t e r , 1973).

Light penetra t ion i s l imi ted by tu rb id i ty as well

as depth. Weak reef growth, especia l ly i n shallow waters R

bn leeward r e e f s , has been explained by tu rb id i ty of

the water.

(b) Temperature

The effect of cold water currents on coral reef

distribution has already been discussed in the previous

section.

Corals also have an upper temperature tolerance

limit. Vaughan (1919) found that the highest temperature

endurable by West Indian reef corals is abobt 3 6 ' ~ . Edmon-

son (1929) presented similar data for Hawaiian species.

Temperatures close to these sometimes occur in stagnant

waters of shallow reef lagoons during extreme low tides,

thereby precluding coral growth or killing established

c6ral s. . .

(c) salinity

Torrential rain .- on wide shallow reef flats or

coincident with low spring tides may lower local salini-

ties and lead to physiological damage or death. Goenaga

and Canals (1979) observed mass mortalities of Millepora

complanata (fire coral) during heavy rains on the Puerto

Rican east coast.

(d) Emersion

Exposure to the atdosphere during tidal cycles,

especially if coincident with midday sunshine or with

rainfall, may 'lead to coral death (Edmonson, 1929). Dura-

tion of emersion is obviously an important factor (Sto-

ddart, 1969). Mass mortalities of echinoids and other .t

reef f l a t organisms coincident with midday, low water

exposures i n L a Parguera, Puerto Rico, has been reported

by Glynn (1968). A s with o t h e r det r imenta l f a c t o r s ,

cora l surv iva l v a r i e s with t he species .

- (e ) Water turbulence

Wave turbulence and energy a r e c e r t a i n l y impor-

tant con t ro l s of co ra l growth. S t o r r (1964), based on

s tudies of t he Abaco reef t r a c t , Bahamas, concludes t h a t

wave t h r u s t i s the environmental f a c t o r which r e s u l t s i n - . . ecological separa t ion of the var ious reef organisms. . - - .

Apparently, extreme turbulence o r extreme absence of i t

i s det r imenta l f o r c e r t a i n spec ies of co ra l s . While water

movement has an important d i r e c t physical e f f e c t , - i t t 1s ... _.

also responsible f b r br inging suppl ies of f r e s h water and . .

n u t r i e n t s t o co ra l s . Removal of C02 ' i s r e l a t e d a l s o t o . ..-

degree of water turbulence (S toddar t , 1969).

( f ) Sedimentation

Sedimentation as a cont ro l reef growth

Puerto Rico has been s t r e s s e d ' r e c e n t l y by workers such

a s ~ i n t r 6 n e t a1 (1973), Kolehmainen (1974), Loya (19761, - .- and Rogers (1977). Branching c o r a l s a r e r e l a t i v e l y b e t t e r

able t o withstand sedimentat ion than massive co ra l s ; o the r s

espec ia l ly with l a r g e polyps, have developed e f f i c i e n t

mucus and c i l i a r y cleaning mechanisms (Yonge, 1935; Mar-

sha l l and O r r , 1931). The r o l e of sedimentation i n i s land-

wide c o r a l d i s t r i b u t i o n i n Puerto Rico has been discussed

by &ye ( 1959) and Almy and ~ a r r i h n - ~ o r r e s (1963).

(g) Storms I. The major cause of catastrophic coral mortality

on reefs is destruction during tropical storms. This

destruction is mostly mechanical. Colonies are uprooted,

carried above sea level or into deep water or fragmented

in situ by wave action. In some cases, though, corals

may survive the storm but later succumb to changed envi-

ronmental conditions resulting from it (Stoddart, 1969).

Glynn et a1 (1965) reported on minor hurricane Edith in

1963 in Puerto Rico.. Winds 'less than 90 h/hr. caused . - . - - 7 .

extensive coral destruction, especially of branching

corals. During the course of this study, we observed the

disastrous effects of two tropical storms (David, -Frede- . - - , . ... - - ric,, Sept. 1979') on, the outer east.'&a.st and especially

on the southern coastal reefs. Damage was most obvious

at the shallow A _ . palmata zone where these corals were

ripped off and overturned causing damage at the same time

to massive corals which were--'extensively bruised by the . - -- .

. landing of the farqr. -

Stoddart (1963) also reported very extensive reef

damage on the British Honduras reefs resulting from ano-

ther tropical storm in 1961. . (h) To this list, other not so natural environmental

controls may be added. These are different kinds of man-

made pollution. Johannes (1975) has given an extensive

account' of forms of environmental degradation caused by .I

man which are detrimental to coral growth. Among others,

erosion caused by upland deforestation, sanitary effluents,

thermal effluents, dredging, chemical spills and bombing

have been pointed out as unquestionable degraders of coral .

reef communities.

The environmental controls mentioned in this list

say act in isolation (as single factors) but most frequently

synergistically (in conjunction) where the sublethal effects

of one are aggravated by the presence of another. The

~ffe=t of these -may be summarized as follows: . - - . . (1) Upland deforestation

Natural erosion is an essential process without

. which we would have no soil, and the- delivery by .rivers . - . . , .L --

of nutrient-laderi soil to the ocean enhances marine .pro-

ductivity. In excess, however, and due to excessive upland - deforestation or bad land management, siltation of erosion

products leads to decreased productivity of coral reef

communities and, ultimately, to their destruction. Exposure

of reefs to brackish, siltladen water associated with flood

-off has probably been the single greatest cause of reef

destruction historically (Johannes, 1975) . Undoubtedly

some damage is natural, but aiso there can be no doubt

that bad land management has greatly magnified the problem.

Xaragos (1972) has reported extensive reef damage in southern

Kaneohe Bay, Hawaii by terrigenous sediments. Van Eepoel

and Grigg (1970) report that in large areas of Lindberg Bay, *

been destroyed and condi t ions remain unsu i tab le f o r t h e i r

establishment due t o sedimentation caused by bulldozing,

construct ion and the sur fac ing of land t h a t d ra ins i n t o

the bay. A subsequent survey (van Eepoel e t a l . , 1971)

indicated tha t condi t ions were rap id ly worsening. Damage

t o reef communities due t o acce le ra ted ter r igenous sedi-

mentation has a l s o been observed i n Tanzania'by Ray (1968)

and i n the Seychelles by Vine (1972).

(2) San i ta ry e f f l u e n t s . . - The r e l a t i o n between organic o r n u t r i e n t enrich-

i ment and lowered, s t r e s s f u l oxygen l e v e l s i s well known. ,

t Johannes (1975) haps pointed out t h a t t r o p i c a l marine organisms*

. . - 3 . - . l i v e cl 'oser, on the average, , t o t h e i r lower oxygen - l i m i t

than b i o t a i n colder waters. Kinsey (1973) observed t h a t i oxygen l e v e l s a r t i f i c i a l l y depressed bnly s l i g h t l y below i

normal l e v e l s on a heal thy reef caused the death of many

reef inhab i tan t s . Depressed oxygen l e v e l s i n reef and 4 near reef environments subjected t o sewage e f f luen t s have i been reported by Bathen (1968) and Wade e t a l . (1972).

(3) Thermal e f f l u e n t s I

Tropical o r g a i sms -live a t temperatures only a ?

i few degrees below t h e i r upper l e t h a l l i m i t (Mayer, 1914).

Consequently, the t h r e a t of des t ruc t ion o r a l t e r a t i o n of

marine communities by overheating i s g r e a t e s t i n t he Tropics.

Heated e f f luen t from a power plant i n Guam l e d t o extensive -

destruction of reef cora's (Jones and Randall, 1973).

Jokiel and Coles (1974) reported on the impact of thermal

effluent on corals in Hawaii. Nearly all corals in water

4-5O~ above ambient wbre killed. T

(4) Dredninq

Brock &. (1966) give a detailed account

of the destruction of corals and reduction of fish and

echinoderm population at Johnston Island due to siltation

brought about by dredging. Deterioration of reef comu-

nities continued for at least a year after dredging ceased - . - because of continual resuspension of sediments. - Grigg

and van Eepoel (1970) observed the destruction of hard ji corals due to sedimentation associated with the release

- - , ... - . of clays brought about by the 'dredge removal . . of overlying. '-.

sand in Water Bay, St. Thomas. Apart from siltation,

dredging can alsd cause an increase in chemical oxygen

demand which may constitute an additional stress (Johannes,

1975). Also, alterations of reef topography through

dredging, filling or underwater construction will alter

current velocity and directions which influence reef

zonation and may alter settling and survival patterns

(~ubbard, 1974) . Examples of, reef damage through dredging

activities are numerous in the literature.

(5) Bombing

An account of the disastrous effect of Naval

maneuvers on coral reefs off northern and southern eastern

Bio log ica l determinants

Bioerosion o r the presence of c e r t a i n organisms

which bore i n t o the reef framework have been shown t o

con t ro l r ee f growth i n t he deep f o r e ree f of Bahamas,

Jamaica and Bel ize (Lang, 1974; Col in , 1974; Neumann and

B a l l , 1970; Po r t e r , 1973; Lang e t a l . , 1975; Ginsburg

and James, 1973, 1975). An example of t h i s a r e the bor ing

sponges which weaken c o r a l ske le tons and hold f a s t s , t hus

infl-uenc'ing th9' s i i e and growth forms of c o r a l s which . - .

l i v e a t t h e i r lower depth l i m i t (Hartman and Goreau, 1970).

Other b io log i ca l processes such a s competit ive i n t e r -

a c t i o n s , b io tu rba t ion (smal l Scale d i s t r up t i ons ) and - , . -.. -

.. - -

predat ion a l s o assunie a n inc reas ing inf luence on commd-

n i t y s t r u c t u r e i n deeper and more d ive r s e reef assem- -

blages (Glynn, 1976; Lang, 1973).

" ., I a, CORAL REEF DEVELOPMENT IN PUERT.1 R I C O

Individual coral colonies grow almost everywhere in

all four coasts of Puerto Rico. Coral reefs, as such, P

however, are present only where favorable conditions

are present.

Reef development along the western two thirds of the

north coast of the Island is poor. Kaye (1959) summarized

the possible factors affecting the distribution of coral

reefs on the north coast. He pointed out that the water-

shed of the'.north coast- is the largest of the Island, in

both area and volume of discharge, and the large volume

'of silt-laden waters from the north coast rivers may be F

- the most importarit >factor inhibiting cbral growth. The ., --

,bportance of salinity and light penetxation for coral

growth has been mentioned earlier in this report. Flood ,

discharges from the several large rivers reduce the sali-

nity of the coastal waters near their mouths. Alter

, heavy rains the long plume of muddy river water off the

mouths of the rivers (generally diverted to the west)

may interconnect one major river mouth with the next, and

form a widespread apron of turbid, low salinity water

along much of the coast. The possibility that these

turbid waters are responsible for inhibiting reef growth

is given some support by the fact that well formed reefs

occur only east of the mouth of the eastern most major

river ~ s ~ i r i t u Sanm. Kaye also mentions that because

of the l a rge storm wave. fha t rake the nor th coast from

time t o time, r e e f s growing on sandy o r muddy bottoms

i n r e l a t i v e l y shallow water have p a r t i c u l a r l y unstable

foundations. Most known . e e f s i n Puerto Rico a r e i n sha-

lldw water and a major i ty of them r e s t on a platform

with depths t h a t a r e wi thin t he range of l a rge s t o m

waves t h a t would s t i r up the sand and mud and, i n general ,

both des t roy the foundation of young r e e f s t h a t a r e struggling

t o e s t a b l i s h themselves and smother them i n the s t i r r e d - - up sediment. - . - -

Another f a c t o r may be the poss ib le bevel l ing of the

shel f a r ea by in tense wave ac t ion which removed any pro-

j ec t ions above the bottom t h a t would pr-ovide.,a su i t ab l e - - - -

- place f o r reef development.

I t should be mentioned here t h a t in tens ive upland -

defores ta t ion during the l a s t 30 years has probably in-

creased sediment runnoff aggravating t h i s s i t ua t ion . I

The poss ib le presence of " luxur ious deep reefs" i between 30 and 100 meters i n t he nor th coas t of Puerto

Rico has been mentioned by Kolehmainen and Biaggi (1975) i !

based on the presence of these i n the nor th coast of

Jamaica.. We, however, quest ion t h i s assumption due t o

the very d i f f e r e n t wave regimes and much lower precipi -

t a t i o n on the Jamaican nor th coas t .

East of San Juan l i e s a discontinuous chain of poorly

developed and heavi ly s t r e s sed rock r e e f s trending i n a

p'", I - - -. -

. , genera i east-west d i r e c t i o n and extending c l o s e t o 1.5 1-1

o f f s h o r ~ (Fig. 2 3 ) . These, from t h e i r alignment wi th ,-

e o l i a n i t e r i d g e s a r e i n t e r p r e t e d t o c o n s i s t of a r e l a t i v e l y

t h i n c o r a l veneer growing on a shallow e o l i a n i t e platform. .a,

which, i n some c a s e s ( e . g . I s l a P iedra , and Isla ~ i n c o r a )

r i s e above t i d a l l e v e l (Kaye, 1959).

Off of Punta Las arias reef patches occur which

t y p i c a l l y a r e moundlike and r i s e t o wi th in a couple of

meters of t h e su r face . The cen te r of t h e top of these

g a l l mounds c o n s i s t of &ad 'corals- ; i y n ~ d on i t s borde.rs . - . ' . by t h e eikhorn c o r a l (Acropora palmata) and on i t s lower

s lopes by sea f a n s and o t h e r gorgonians.

Northwest of Boca de Cangrejos l i e s what w a s a we l l - I . . . - . -

' - developed, r e e f system. Here, ex tens ive c o r a l formations

were common from t h e s u r f a c e down t o 10 meters i n f a i r l y

c l e a r wa te r s . Th i s r e e f w a s v i r t u a l l y destroyed by sedi -

ments de r ived from ex tens ive dredging wi th in T o r r e c i l l a

Lagoon and organic sediments discharged i n t o th is same

lagoon by sewage t r ea tmen t p l a n t s . A t p resen t , b a r e l y

any l i v i n g c o r a l e x i s t s below approximately 1.5 m .

Stony c o r a l s a r e p r e s e n t on t h e rock r e e f s and beach

rock p la t form a t Punta ~ a c i a Talega (Pig . 24). Most a r e

enc rus t ing growths and n o t major c o n t r i b u t o r s t o r ee f

c o n s t r u c t i o n o r maintainance. These a r e most abundant

along t h e nor the rn s i d e of t h e i n n e r r e e f . Mil lepora

complanata' i s t h e most abundant c o r a l nea r t h e su r face

Fig. 83. Line of rock reefs (Trending east t o west) off shore San Juan.

2 Kms - Fig. 24. Heavily s i l t e d reefs off Punta Ig les ias and Punta San

Agustin (center ) . Punta Vacia Talega can be seen a t the l e f t of the picture.

of these rock r e e f s . D ip lo r i a and I sophy l l i a a r e a l s o

common on deeper a r eas . Sof t c o r a l s a r e present on pro-

t ec ted a reas . .-. ,. - - -

- - Sca t t e r ed ;&if r breaking the surf6ce- are fo-Ad

'%--.- - -. ?

between Punta, --. W e s i a s and Punta San ~ ~ u s t b - i ~ i ~ . 24) . z -

- - . - . ~> .

Everi though thel.db-no .- . . t f o m a cont i n ~ o u ' $ ~ & r r i e r , ...- they - - -. - .,-. . - have formed S ' b f f e t t i G e - w a v e k e r g y a b s r b i n g s t r u c t u r e .

A t present t he pa tches - . - - adjacent- t o shore a r e d a d probably "

--=--7 A - - - -- - by s i l t a t i o n . Water kondi t i o n s here are 'characterized-_ - -- - . . . . ) _ _

b y h i g h l e r e l s o f i ~ u - ~ ~ e n d e d p a r t i c l e s and-low.ui~ii!&liG:~ . - - . . :.:-- -. - , . . .-

_.A. ? .

Water q u a l i t y and hea l th of c o r a l s increases . . . offshore-$ ~. .~-

. . . +* 'a_ I TI. - ' - - 5 . . .

l i v ing corab being present only i n the-:$t;al%w::E-3 -~ - -- . . h) -: . -- .. . _ .i . *_- - .- - - - - -. .- - - . . .~

depths of the outermost r ee f s . S i g n s - ~ + ; - p ~ ~ . . e , ~ ~ i o n ~ - - -.. - . -.-..>..:--:- . , .: .. . - - . - .. - . .-a . . - . > - -

. , . . ' a r e evident , a t t he base of the reef-s. - . - . . ' .

.. - .

- Far ther e a s t , on the nor th anh w e i t s ide of Punta

Miquillo and on the nor th and eas t s ide of Punta PicGa,

a r e f r i ng ing r e e f s which average about a quar te r of a

mile wide (Fig. 25). I t i s probable t h a t both Punta

Miquillo and Punta'PicGa were formerly sand cays developed

from these r e e f s , which s ince have been t i e d t o the main-

land by a broad marsh and narrow sand tombolos. These

two r e e f s , spec i a l l y t h a t of 'Punta Miquil lo, a r e i n very

poor hea l th s i t u a t i o n with very low l i v ing cora l cover

and d i v e r s i t y . The Punta Miquil lo reef has suffered

ser ious per tu rba t ions and p a r t i a l des t ruc t ion of i t s .com-

ponents a s a r e s u l t of t he dredging of a channel p a r a l l e l

Fig. 25. Fringing reefs off Punta Miquillo and Punta Picua. Annular reef off Rfo Mameyes i s seen t o the cen te r r igh t of the photograph.

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

of the r e e f framework and c r e a t e d a s i l t y environment,

due t o resuspens ion by wave a c t i o n , which i s de t r imen ta l

t o c o r a l s . Punta Percha i s p a r t of t h e same system exhi-

b i t i n g similar condi tons bu t s l i g h t l y h igher l i v i n g

c o r a l cover .

Ensenada ~omezbn ( F i g . 25) i s l i n e d wi th numerous

patches covered p r i n c i p a l l y by a lgae . Stony c o r a l s cover

these t o a l e s s e r e x t e n t . The pa tches , more than a couple

of meters i n r e l i e f , p r e s e n t no d i s t i n L c t zonat ion. ~ o r a i - . - . - .

species a r e M_. lamarckiana, -4. a g a r i c i t e s , M_. squar-

rosa P. a s t e r o i d e s , and P . s t r i g o s a . Surrounding waters - 9 - a r e g e n e r a l l y ve ry t u r b i d .

- I ... - . Two l a r g e pa tch r e e f s occur o f f s h o r e from t h e mouth

of R ~ O Mameyes. Both a r e roughly c i r c u l a r (300-500 m i n ~ -

- diamet-er) and wi th an exposed shoal of coa r se sand (mainly

Halimeda) and broken p i e c e s of c o r a l . They c o n s i s t mainly

of shallow g r a s s beds i n c i s e d by north-south t rending

sand channels . The underwater edge of t h e i s l a n d i s

f r inged wi th a narrow band of c o r a l of which l e s s than

20 per c e n t i s a l i v e . The seaward edge a l s o c o n t a i n s en-

c r u s t i n g c o q a l l i n e a lgae . The f o r e r ee f c o n s i s t s of a

s t eep ly s loping c o r a l pavement, spa r se ly covered wi th

l i v i n g c o r a l s and t e rmina t ing i n a ba r ren sand f l a t . Corals

Present a r e Acropora palmata, Montastrea a n n u l a r i s , M_. caver-

nosa, and D i p l o r i a s t r i g o s a . The genera l low d i v e r s i t y

Mameyes. These i s l e t s and t h e i r surroundings were h e a v i l y E P impregnated with Bunker o i l during an o i l s p i l l i n December I

2 1 , 1978. I $

East of t h i s r e e f system t h e r e i s a complex of b a r r i e r , ,

f r i n g i n g and patch r e e f s which a r e r e spons ib le f o r t h e

formation of Luquil lo Beach (Fig . 2 6 ) . Some of t h e s e i probably r e s t on rock foundat ions with t h e except ion of b

I those northwest of Luquil lo which occur a t cons ide rab le

. . . . ' d i s t a n c e s frgm rock out crops and t h e r e f o r e may be b u i l t

on sand o r mud. These r e e f s have undergone v a r i o u s s t a g e s

of degradat ion. The f r i n g i n g r e e f s surrounding t h e nor thern

and eastekn end of -the beach show degradat ion i n t h e sez - - - ward edge where growth h a s been l i m i t e d t o t h e upper

t h r e e meters. -

East of Luquil lo water t ransparency i n c r e a s e s gra-

d u a l l y and reef e x h i b i t s l i g h t l y h igher l i v i n g c o r a l cover .

East of R ~ O Juan arti in a r e a s e r i e s of patch and

f r i n g i n g r e e f s which have been descr ibed by Torres (19735.

Coral spec ies d i v e r s i t y he re i s low, A_. palmata, 2. s i d e r e a ,

D . s t r i g o s a , and M_. cavernosa being the most common species . - The a r e a s of h ighes t abundanbe, namely t h e o u t e r r e e f

f l a t s , had 30 per cen t c o r a l coverage. Torres d i d not

observe s i g n i f i c a n t c o r a l grdwth below 3 m depth. Reduced

i l lumina t ion , caused by s i l t p a r t i c l e s i n suspension,

appears t o be the l i m i t i n g f a c t o r . The seaward edge i s -

Punts Percha with fringing r.eef protecting Luquillo Beach.

characterized by overhanging ledges. In scme areas

these ledges have collapsed' creating crevices and cave-

like structures. The presence of uneroded dead coral A

masses, mainly of A_. palmata, =suggest &recent death. - Pumping activities of W e r accumulated in sand extrac- . -- - tion pits was observed by Torresc These activities are

- an additional stress to these already.stressed coral co%-

- nies.

A reef system fringes the area comprised between

--west of Cabeza Chiquita to C a k - S a a Juan - ( ~ i g . 27). ' , - - -:

- This reef .is also undergoing rapid-degradation from' the .- --Cr . - -

effects -. of siltation and. also: F~ .* from .sy;temic -~ extract ion ~.

--' a. - - - &*. : ...-

-> . . of corals 'for salx, The - horChernwst portlon, of this. -

.-ba;-<as a very shallow reef platfdrm -(*art-'of the mentioned

system) which terminates at the beach after merging into -

a shallow Thalassia and algal bed. -Corals are actively

growing along an'eastern channel edge and back reef,

Northward of the reef crest is a steep slope terminating

abruptly in a sand flat. Fire coral is very abundant

in this reef' s crest.

The best reef development on the northeast coast is

found in the fringing reefs formed around the string of

islets which overlie La Cordillera, a shallow, narrow

submarine ridge approximately 18 miles long which trends

east-southeast from the northeastern most tip of Puerto

Rico to Culebra (Fig. 28). These reefs are of high quality, I-

Fig. 27. Reef fringing area from Cabeza Chiquita to Cabo San Juan. Observe also reef frlnging eastern coast of Las Cabezas.

Fig. 28. Icacos, nesternmost i s l e t of La Cordi l lera .

i n terms of d i v e r s i t y , h igh Living c o r a l cover , and extensive-

ness . The i s l e t s of L a C o r d i l l e r a , s p e c i a l l y I cacos and

probably a l l (Kaye, 1959) a r e not reef c o n s t r u c t i o n s bu t -

a r e composed o f o o l i t i b e o l i a t ~ i t e which was depos i t ed and

p a r t i a l l y shbmerged some the previous t o t h e development *

of t h e r e e f s . Most of t h e s e - a r e high i s l a n d s and g e n e r a l l y - - % -.

possess landLand/or beach vegeta t ion . Rock r e e f f r i n g e s

two t h i r d s ' o f I c q o s n o r t h shore. Corals he re cover l e s s - " -- -A s

than 50 per cent of t h e - a v d l a b l k sur face a r e a (Mckenzie +

- . -and B'entm, 1972J. , .Off - t h e -&uthwestern shore of I cacos -

i s a more p ro tec ted a r e a which - e x h i b i t s f a i r l y h igh c o r a l -

development. South o f ' t h e main l i n e of L a C o r d i l l e r a r e e f s

but s t i l l on-,th& same p l a t f o m t h e r e l i e t h e Palominos - % - - . .

(Fig . ' 29) complex and Cayo ~ a r g o (Fig . 30) ', two o t h e r r e e f s

of high b i o t i c q u a l i t y and extensiveness . Between t h e s e - and t h e mainland l i e some o t h e r i s l e t s (F ig . 31) posse-

ss ing h ighes t r ee f development on t h e i r e a s t e r n shores .

Palominos and Isla de Ramos a r e the tops of p a r t i a l l y

submerged h i l l s according t o Kaye (1959). Ridges of the

former cont inue south and e a s t as l a rge shallow submerged

banks. The degree of development appears t o be r e l a t e d

d i r e c t l y t o the d i s t a n c e froai t h e mouth of R ~ O F a j a ~ d o .

Palomini tos , I s l e t a Marina, and Cayo Ahogado have been

formed by wave depos i ted sand and c o r a l fragments a top

a reef platform. These r i s e only l e s s than 3 m above

sea l e v e l and a r e suscep t ib le t o occas ional d r a s t i c wave -

Fig. 29. Palominos and Palominitos (southern sandy islet ) of r' Fa jardo.

Flg. 30. Cayo Largo, o f f Fajardo.

Fig. 31. I s l e t s and reefs off Fajardo. From the bottom I s l a de Ramos, Cayo Ahogado and I s l e t a Marina (Cayo Obispo arrd Cayo Zancudo) a re seen.

eros ion , esp ,?c ia l ly Palominitos and Ahogado.

Fr inging the mainland, an extens ive but dying reef

borders t h e coas t from n o r t h e a s t Cabo San Juan t o t h e

nor th end of Punta Sardinera (Fig. 27). The ent rance r PI

t o - ~ a h i a Las Croabas i s p ro tec ted by t h i s r e e f . From

Playa Sardinera t o Punta Barrancas t h e r e a r e no c o r a l

r e e f s f r i n g i n g the coas t probably because of the in f luence

of the R ~ O Fajardo which c a r r i e s s i g n i f i c a n t amounts of

s i l t . Narrow c o r a l r e e f s , however, p r o j e c t eastward about

- - . - 450 m from Punta Barrancas and Mata Redonda (Fig . 32) . . . - There i s a shallow ree f i n t h e nor the rn ~ a h i a Demajagua

which r i s e s ab rup t ly from about 2 m t o t h e r ee f c r e s t

(about . 3 q). Coral growth and r e e f development i s n o t - -

*'extensive. Going westward t h e r e i s a deep boulder c o r a l

zone which merges i n t o a Acropora zone and i s followed

by t h e r e e f c r e s t where f i r e c o k predominates. West-

ward i s a g r a s s bed wi th t h i c k mounds of P. p o r i t e s and

extensive zoanthid c a r p e t s . Sea f a n s and o t h e r gorgonians

' a r e p resen t e a s t of the boulder zone (McKenzie and Benton,

1972).

Off Medio Mundo, Ceiba, i s I s l a Pifieros (Fig . 33)

with moderate c o r a l growth on i t s nor the rn and e a s t e r n

coas t s and Cabeza de Perro. This l a t t e r i s l e t was used

by t h e U.S. Navy f o r bombing p r a c t i c e s and marine l i f e

i s wanting.

South of t h i s po in t up t o Punta L i m a t h e coas t i s

f r inged p r i n c i p a l l y b y g ~ h a l a s s i a meadows although occas ional

Ftg. 32. Reefs projecttng east of Punta Barrancas and Punta Mata Radonda .

Fig. 33. Is la Plfieros and Cabeza de Perro off Medio Elundo, ' ~ e i b a .

small f r i n g i n g and patch r e e f s occur .

Some of these f r i n g i n g r e e f s of t h e e a s t coas t pro-

bably r e s t on sand o r mud foundat ions judging from t h e i r

loca t ion a t the edge of t i d a l swamps (Kaye, 1959). These, ~ c p . 9 - n .

f o r the most p a r t seem t o grow from a 6-7 meters deep

platform. Also, many patch r e e f s t h a t do not reach i n t e r -

t i d a l l e v e l occur o f f t h i s s t r e t c h of c o a s t .

The high l e v e l s of l i g h t p e n e t r a t i o n , t y p i c a l of

t h e south e a s t e r n end of t h e i s l a n d , f i d s i t s l i m i t a t

Punta L i m a . West of t h i s poin t t h e c o a s t a l waters become . .

t u r b i d due t o a s e r i e s of sediment laden r i v e r s and creeke

t h a t sharp ly reduce i t s t ransparency. The f i r s t s i g n i -

f i c a n t one i s R ~ O ~ n t o n R U ~ Z which i s o c c a s i o n a l l y dredged - .

and pumped. not her source of s i l t a t i o n i s t h e r ecen t

development of Palmas d e l Mar at Humacao which has c l e a r e d

the v e g e t a t i o n i n extens ive a r e a s of s t e e p s lopes and loose

s o i l . Erosion he re has been so extens ive t h a t deep s c a r s ,

which have r e s u l t e d i n damage t o beach and c o a s t a l commu-

n i t i e s , a r e evident i n t h e washed s o i l . Here o c c u r - s e v e r a l

i s l e t s such as Cayo Sant iago and Cayo B a t a t a (F ig . 3 4 )

which p resen t some c o r a l growth s p e c i a l l y i n shallow

waters and i n a r e a s open t o sea ( f a c i n g sou th ) . S u r p r i -

s i n g l y dense 90 pe r c e n t l i v i n g elkhorn c o r a l s t ands i n t e r -

mingled wi th gorgonian and head c o r a l s occur c l o s e t o t h e

sur face . Coral cover diminishes d r a s t i c a l l y with depth

giving way t o a r e a s of very spa r se s o f t c o r a l growth. I t

appears t h a t w a G r movement caused by t h e incoming waves

Fig. 34. Cayo Santiago (upper r i g h t ) and Cayo Batata ( cen te r ) off Humacao. Several submerged patch r e e f s , LnclutiLng Bzjo Parse, can be seen south of Cayo Santiago.

i s a f f i c i e n t t o keep c o r a l c o l o n i e s f r e e of sedimer.,

but a t t h e same time water t ransparency i s too low t o

permit c o r a l growth except i n t h e shallow depths . Sub-

mergpd. shorls with spa r se cdPBl alsa oc_cuF occa-

?i@ially o f f Humacao. An-exa&le of t h i s i s ~ a j o Parse

(Fig . 34) , which consiSt s of numerous- gorgqnians, small

head c o r a l s a h d s x t e n s i v e pa tches of - the enc rus t ing -.

sponge ( A n t h o s i m e l l a varians. Depth i s "ever more than

- - - - about 5 m. - + - - - - - . . ~.

- : . ~ ; & e ~ i t ~ " e c o l o g i c a l conditi-&S . . . . ..- f o r . t h e . - -. succe.ssfu1 . ,

growth of c o r a l s do no t - e x i s t a t p resen t i n most a r e a s - -- - - =

of t h e Yabucoa Bay. The annu la r r e e f l o c a t e d a t t h e #

- s o d t h e m p a r t - o f t h e bay . (F ig . 35) i s n o t an except ion . --

and, i n s p i t e o f - t h e fa& t h a t a f e & l i v i n g -co ra l s a r e * =r

found t h e r e , s e v e r a l o t h e r b i o l o g i c a l i n d i c e s from orga- -

nisms t h r i v i n g t h e r e i n d i c a t e t h a t environmental condi-

t i o n s e x i s t i n g a t p resen t i n t e r f e r e wi th t h e normal

growth of c o r a l s . These cond i t ions seem t o be t h e i n f l u -

ence of f r e s h water coming from t h e r i v e r s , r a v i n e s , and

creeks which empty i n t o t h e bay and resuspension of f i n e

sediments by p r o p e l l e r s from t h e heavy s h i p t r a f f i c .

C h a r a c t e r i s t i c s of t h i s annular r e e f a r e : (1 ) the s c a t t e r e d

growth of A_. palmata and many o t h e r dead c o r a l s , ( 2 ) t h e

macroscopic a lgae coloniz ing t h e dead o r dying c o r a l s ,

(3) t h e luxur ious growths of l a r g e ben th ic marine a l g a e

among t h e c o r a l s , ( 4 ) t h e e s t a b l i s h of marine phanerogams, *

. - . -T - 1 K m . J - *

Fig. 35a. Amiular r e e f off Yabucoa Bay.

Fig* 35b. Non v e r t i c a l z e r 4 a l photograph :ke same r ee f .

mainly Cvmodocea among t h e c o r a l s , (5) t h e s c a r c i t y of

reef Foraminifera and t h e presence of Foraminifera n o t

c h a r a c t e r i s t i c of c o r a l r e e f s ( ~ { a z - ~ i f e r r e r , 1969; and

S e i g l i e , 1969). ..About 5.5 n a u t i c a l mi les e a s t of Yabucoa

Bay t h e r e i s a reef c a l l e d L a Conga by n a t i v e fisherman.

This r ee f was not v i s i t e d but presumably i t forms p a r t

of t h e submerged b a r r i e r r e e f which borde r s most of t h e

southern she l f of the I s l a n d which I am going t o d e s c r i b e - l a t e r .

. Sargeant Reef, .3 h. southeastward of Ebnta Tuna ' .

i s 1.08 Ean long and .1 km wide a t i t s widest p o i n t .

Because i t breaks t h e f o r c e of t h e sou theas t swe l l , t h e

- . * reef a f f p r d s p r o t e c t i o n from t h e southeas t f o r the shore- . .

l i n e i n t h e v i c i n i t y of- Punta Tuna where the r e e f i s from ,

. 1 2 t o 18 Ean from shore. This r e e f i s of a h igh q u a l i t y - i n terms of l i v i n g c o r a l cover and d i v e r s i t y . 1t p r e s e n t s

a reef f l a t with abundant A_. c e r v i c o n i s t h i c k e t s . Following

south i s an a r e a of reduced A_. palmata growth wi th high

encrus t ing a l g a l cover. Th i s a r e a g ives way t o a ve ry

dense P o r i t e s b io tope (wi th pa tches l a r g e r than 100 square

meters i n some a reas ) which a l t e n a t e s wi th overwash colo-

n i e s of A . - palmata. F a r t h e r +seaward i s an a r e a of dense,

100 p e r cent cover of A_. palmata which i s reduced g r a d u a l l y

south merging with an a r e a vhere gorgonians predominate.

A f r i n g i n g reef extends almost c o n t i n u a l l y f o r f o u r

mi les a long the coast between Cabo Mala Pascua t o Puerto Ir

Pati l las . This i s ex-posed a t low t i d e and p r o t e c t s a low

sandy apron which l i e s a t the foo t of t h e S i e r r a de

Guardarraya. A s i m i l a r h igh ly s t r e s s e d r e e f i s respon-

s i b l e f o r the seaward p r o t e c t i o n of Punta F igueras

(Fig. 36) . P a r t i a l l y r e spons ib le a l s o f o r t h e p r o t e c t i o n

of t h i s a r e a and Puerto Arroyo i s A r r e c i f e Guayama ( F i g . 37)

about .6 to - 9 km o f f Pun ta Figueras . This reef i s

very extens ive ( n e a r l y 5 k i lometers ) but i s p a r t i a l l y

a f f e c t e d by s i l t a t i o n rendering an average low l i v i n g

c o r a l cover. T'ne r ee f f l a t of A r r e c i f e Guayama i s f r a g -

-mented in,-small b u t t r e s s e s s wi th f i r e of e lkhorn c o r a l

" on t h e i r tops depending apparent ly on t h e degree of expo-

sure. The f i n g e r c o r a l i s a l s o an important component

I of - these s t r u c t u r e s . The Ac-ropora palmata zone has a -: --

. . very low cover and h igh m o r t a r i t y of s tanding ko lon ies .

Seaward i s a zone of small s ized gorgonians and dead s tag- - 1

horn c o r a l ( c e r v i c o r n i s ) - which i n c r e a s e s i t s depth

gradual ly up t o t h e she l f edge. Westward of t h i s r e e f

a r e the Corona and Algarrobo pa tch r e e f s which appear

r e l a t i v e l y hea l thy and no t much a f f e c t e d by s i l t a t i o n .

South of Las Mareas, Guayama, l i e s A r r e c i f e L a s Mareas

(Fig. 38) which i s n e a r l y t o t a l l y devoid of l i v i n g c o r a l .

The dea th of t h i s r ee f i s probably due t o - s i l t a t i o n by

Guamani River e a s t of Punta Ola Grande. Tongues of s i l t

have been observed spreading predominantly westward from

the mouth of t h i s r i v e r (Tor res , 1978) . High s e a s and

heavy su r f a c t i o n p r e v a i l i n g i n t h i s open beach maintain -

Fringing reef off Punta Figueras.

Fig . 37. A r r e c i f e Guayama south of Punta F i g u e r a s .

Fig. 38. Arrecife Las Mareas off Guayam (lower r i g h t ) .

t h e s i l t i n suspens:m. Furthermore, t h e rock j e t t y

p r o t e c t i n g t h e entrarace t o Puer to Las Mareas, b locks t h e

westward movement of the s i l t e d wa te r s r e t a i n i n g them on

t h e e a s t e r n s ide . Extgqsive d r e d g f n g , o p u a t i o n s took - - L .- -

~ l a c e h e r e f o r t h e castruction-or-t~.-artificial harbor , : . - -

&- - . . . - .. . - . " . .

a t Las E ; a s and.may h a ~ r - i o n m i ~ & t i d . . to h i g h r e d i n h t . . . - ~ ~

- -- - l e v e l s . LL - -

Southwest of Punta Pozuelo extends-a f r i n g i w b a r r i e r *

- . '~ , . - - .

reef c a l l e d Cayos Caribe f o r - a d i - . ~ ~ . ~ f &bout 5 . 5 lan -- . - . . - - -~-+ -. . - . ~ . .

. .. 2

A s p a r t bf th:.& same -,. reef ry'sta,~~but-; . - W i d e d - ,from i t - - - C Z . . - - -r L -., - 7 . -: .. - , ..- . .: .. -. .

-,s: - :- -7 each o t h e r :>by shall* , . channklk e e ? ~ a y u r - - d e - .- Bar& q d -.--. -&

.. ---.- - . . ,- -- - -*--gz -~ - . . . - -. . .- - . . - - .s -- .. .. . - - -

Cayos .:de P&jaroz(~fg~, 3 9 ) - . . . - z~!&&-fa-~ . : -a . di'&noE&rc - -L .. - - . . . . ..--.* - :- - ~-- - =

. - z,., =,:. . :---L-. - -. = . -' a:>. ; - . . -- -. _ . _ . __i_A& - . - -;- " C

C - . . ghat *f f . .. ec t i ~ e l ~ ~ ~ ~ o t e c ~ ~ ~ ~ ~ g ~ ~ ~ ~ ~ f ; ~ - ~ - - .- ., - --.-. ., . - de . . . ~oboG:- - - ;z- ' -*.--. ." .- - , . ... - . , 4 - < . - . - - - ..&. -- 5:,a-. &-.-- + ,;- .*?-- ~- :--+-- - - ' . .-*A. . ., . - 2%- -

z,:* t h e l e k ~ : - o & ~ . ~ e . & e e ~ & a ~ & ~ f ~ ~ a , - ~ ~ b b e , . ..-- - -. . -. - , ..-. :----.= , - - -- - ,--,~ - . . - _.--,

, - . - - . . i-- - - - - . - -. - ? - -

-n ' .- - *-5 -.I{~ :-. ~ .. . d i . = - a r e a score of narrow sand c a y - s i - f r i n g ~ ~ by nangrove vege-

- t a t i o n and o r i e n t e d normal t o t h e r e e f margin 'which a r e

separa ted from each o t h e r by d ra inage channels . These

channels d r a i n the water t h a t washes over the h i g h e r ,

ou te r margin of t h e r e e f t o t h e bay (Kaye, -1959). Living

c o r a l cover i s moderate and i n c r e a s e s westwai-d.

Numerous o f f shore keys occur south of S a l i n a s (Fig . 40)

and Santa I s a b e l (F ig . 41). T b s e a r e , west from Cayos .

de ~ i j a r o s , Cayo M o r r i l l o s , Cayos de Ratones, A r r e c i f e

Media Luna, Cayo Alfefiique, Cayos de Caracoles , and Cayo

Cabuzasos. These a r e i n ve ry h e a l t h y s t a t e wi th h igh

l i v i n g c o r a l cover and support r i c h ben th ic and nekton t

Fig. 39. Cayos de Barca, Jobos.

2 K m s

Fig. 40. Reefs off Salinas.

Fig. 41. Reefs off Santa Isabel.

populations. Most of them except Medic. Luna .have mangrove

vegeta t ion i n va r ious s t ages of development. Crescent-

shaped Cayo Alfefiique, wi th i t s arms p r o j e c t i n g t o t h e - _.--_ -

wr thwes t , shows t h e importance of the soli bind ;=- and , , a.

the northwest and wes t e r l y cu r r en t s i n the molding of these . .

f ea tu r e s (Kaye, 1959.). - .. . C. -.-' -.

cay* ~ e r b ~ = r i . { f i ~ . 42)-,- titiit : b f E i y o ~abuzat i6 ' s , has

an 'extensive f r i n g i n g r ee f ly ing on i t s e a s t e r n and +i>=. . --. . - -- , - .

south<rn shores .=-.- ~ o ; f r ~ . . d6velopment' . - reaches . . . :its m q i m u m - .. -- .- -3- . -. . .

:.:. , .-- . . . : .;i- - on t h e s0u the . a &ore where theA;: bdima$&.;-zone- r&hG.~t- _ - - -~ . .

. =. . . --= . - ,*:.-. : . ... . . -- . . - . . . - - . . - a 95 .per..:cen<;cbve~ & d - t h e r e & - s e x ~ n l i ~ e fi&hJauna.

.. - - - .. . - 11*. - . t:=- - -. .- -. - . -. . . .

- - ~ K- . ~ *- =.+ .- --.I-- A. L . - , -A*. .-.*i- --. ? - ' Y _- . r '

.?&re -. 4.g . --&elat .*- : :.=-. i v e l y 'mfouched'm~~rp.t!~ ...-. lgor6st=+-th&.-r'--- .s .-, -- 5 ---:- :--. - &- L- -.- -;-;; - . -,: ----- 3-% .-.=T- - .. - - . .

7 & ' & . .-. - ..<A. -,- .<E--,;.. ":- ;----.--- - ' . - - . - ' 2- i ;-@ e*. o ~ ~ ~ ~ ' ~ ~ ~ - - . . ~ ~ u t a - ~ r ~ ~ r -B&-* ;f s-. - - . : 7 . - - ' -.-.. - rr'

< .- , . . ~

- - - - -- P a '--ma , .. z -- . -. . . . - - -+.' -- ' ~ 1 , .--a.: . - . - - , - . . 7 - . - - . .*. .~ &_ .. , _ .- --._ , - - . _*,+ .'* 1. ~-'--..z-- .>:: , . , f iket ea ,= eer::a~1 =d ~ ~ ~ ~ l ~ 2 w . e & r y e ~ s,+x G-6% + ; -

_ ._ . . I . .-...-,.+ 0 . . . . . .- ~

- - . - .~ - .... . -- .e- - . - - ~ A -- . -A L$-g-h&e,m .- .;#-: a . . - - .... . - - * - . - * . .?' ...-. . .. - -,

extik-si~e tlRliorn ~ti.ral~'ei5vereige'~and deiisbgotgonf%n - stands. F i sh l i f e i s e s p e c i a l l y abundant here .

About .6 km southeas t o f f Ponce i s I s la Caja de

Muertos (F ig . 43) which p r e sen t s h ighes t r e e f development

'on i t s no r theas t e rn shore. Spec ia l ly no t ab l e of t h i s

reef i s i t s complex high r e l i e f lagoon which suppor ts

a l a rge v a r i e t y of benthic and nektonic fauna.

Off Ponce l i e numerous mangrove i s l e t s f r inged by

co ra l r ee f ( I s l a d e l F r i o , I s l a de Cardona, I s l a de Ratones,

and Cayo Cardona) (Fig . 44 ) . These p resen t s t ands of

Acropora palmata s u r p r i s i n g l y dense f o r these s i l t laden

waters. S c l e r a c t i n i a n c o r a l growth, though, i s sparse IL

Fig . 42. Cayo Berber ia off Santa I s a b e l . Highest reef development i n south coes t .

' Fig . 43. I s l a Caja de bfuertos. off Ponce. Highest reef development i n eastern coast .

2 K m s

! Fig. 44. Cayo off Ponce. Arrecife Ratones can be seen a t left.

i n deeper w a t e r s where gorgonians dominate.

About 2.5 km south of Ponce i s a l s o Bajo Tasmanian,

an a r e a of p r o l i f i c c o r a l growth. Bajo Tasmanian con-

s i s t s of a two .. -*., l eve led p la t form (Beach, 1975). The nor- = - t h e m l e v e l ranges i n depth from G.m 3 2 meters and t h e -

southern between 18 and 24 meters . - The s taghorn c o r a l

(Acropora c e r v i e f i i s ) i s p a r t i m l a r l y &un$ant dver much . - of t h i s l o w e r l e v e l . A t t h e s h e l f edge of t h i s lower

l e v e l a r e l a r g e s h i n g l e - l i k e growths o a r i u s massive - % -If . - .- .- . ..& ' % . * - . %. .- . - . . c o r a l s . . . ' *; . . .. .. . :...

- . e-. . - .. ~ - . . ? . . -. - . - .

. . The i n d u s t r i a l development oli33ah:a G w y a n i l l a and:'

. -. - ... - - : 4- .

Tallaboa h$ye s i g n i f i c a n t l y a1 t e red t h & + t - o r a ~ ~ ~ k s t a f ~ =-. - - - -.. - .=* : :- . .r> - --- - -- -'-. -- . _'.. - -

f e a t u r e s . --- and of f k h r e . - .-, f e@f k - 2 ~ h . f . - .. ,. Qff ~ .:- . . --: .. , :--s ?.& - L. . - .. . - . . -.<- - - ..'i - ' - . .: ..a- -

P~~~~~ - . ( r i g ~ ~ < - ~ s $ ~ ~ ~ ~ ; ~ ..%. -. oc m ~ & & ~ ~ " , - ~ ~ ~ : ~ gkr' 'st- s s . . , .. - .- .-; :: 2- L . T .

- . . .- * . . . - - - -. - .-

- .*-. . - - - .- .-.;I -* -* ~ -ri

c o n d i t i o n s due mainly t o s i l t a a ~ - - b ? p e r i b d i c dredging ..A

of sh ip channels p r o p e l l e r s t i ' r r i n g (F ig . 45a) and a l s o

t o the d i scharge of b i l g e water con ta in ing o i l y wastes

i n t o t h e water as w e l l as t h e i n d u s t r i a l e f f l u e n t s which

a r e p r e s e n t l y descharged i n t o t h i s bay. Living c o r a l

cover i n shallow r e e f a r e a s o f f Tal laboa i s c l o s e t o zero

wi th some i s o l a t e d heads of Acropora palmata and Y i l l e p o r a

comolanata s t i l l su rv iv ing i n t h e seaward s i d e . General ly

these zones of dead c o r a l a r e cont inued south by spa r se

gorgonian growth and then by s l i g h t l y h i g h e r l i v i n g ,

massive c o r a l cover a t t h e r e e f s lope where t h e sediment

apparen t ly does n o t accumulate. Off Punta Verraco (Fig . 46)

. Flg. 45. St ressed r ee f s off Tallaboa. Note oil re f in ing complex c lose ' t o reef a r ea .

Fig. 45a. Ship stationed between Cayo Rio and Cayo Palomas in Tallaboa Bay.

a Fig. 46. Reefs o f f Punta Verraco, GuayanFlla.

i s a r e e f which h a s an ex tens ive Tha lass i a and Syringodium

bed on i t s r e e f f l a t . Stony c o r a l cover i n t h e shallow

f r o n t r e e f i s v e r y reduced w i t h t h e zoanthid Palythoa

covering most of the CaC02 f m u r e m r k ; ~ I n t h e deeper f o r e - re* i s an ex tens ive and q u i t e heal thy- community of t h e

poss ib ly more t o l e r a n t s o f t c o r a l o r gorgonians. ? -

-. An ex tens ive submerged r e e f surr&ds t h e c o a s t from

Punta Ventana t o Punta Vaquero, where i t breaks t h e sur-

f a c e h e r e f i r s t as a f r i n g i n g ree f and l a t e r as a b a r r i e r

reef t h a t pro tec- t s playa. ~ a m a r i i d o , ~ a h i a de $-a -a - ~ a l - l e n ; : - . . R

and Playa de C d a Gorda (Figs . 47, -68). This r ee f i s .. . .- .

almost t o t a l l y ; devoid of l i v i n g - . . + r a t a n d ..-% huge l i v i n g . : . - . , . -- . - - --.--

c a r p e t s of t h e fast ,-growing ..-.. cp1ciriia1 , -... ? & x e g ~ ~ s ~ - . Zoanthus - -. .

and Paly thoa l i e over t h e deaa c6ras-framework.

West of Punta Jorobado c o r a l r e e f s become more pro-

l i f i c and complex forming a s e r i e s of provinces and brea-

I king the s u r f a c e as f a r as two n a u t i c a l m i l e s o f f shore . i

i These a r e t h e L a Parguera r e e f s which no t only p r o t e c t !

f o t h e r important l ' i t t o r a l c o r n u n i t i e s such as mangroves

and T h a l a s s i a meadows bu t serve a s t h e i r foundat ion

(F igs . 49-59).

Here, between t h e s h e l f edge and t h e c o a s t , two

e longate r e e f systems, a l igned approximately east-west

d i v i d e t h e s h e l f i n t o an i n n e r , middle and o u t e r s h e l f .

Two t h e o r i e s have been presented t o exp la in t h e pos-

s i b l e o r i g i n of L a Parguera r e e f s , t h e s e a r e :

. Fig . 47. Fringing and patch reefs off Guhice .

2 K m s r

i . Fig. &8. Reefs pro tec t ing Playa C a b Gorda, G u h i c a . i

. Fig. 49. La Pargaera.

Fig . 50. .Patch reefs off I s l a Cuevas.

. Fig. 51. El Palo - Atravesado reef.

Ftg. 52. Margarita reef .

Fig. 53 . San Cristobal reef.

~ i g . 54. Laurel reef.

Fig . 55. . Media Luna reef.

Fig. 56. Enrique reef .

Fig-. 57. La Gata - Caracoles reef.

Fig. 58. Enmedio reef.

Fig. 59. Mario reef.

(1) The major geo log ica l f e a t u r e s of La Parguera

a r e a r e s u l t of deformation of upper Cretacous l imestones 6 -

(wi th in terbedded mudstones and volcanic rocks) i n t o a - . - *-

W - E S E t r end ing s y n c l i n e whose a x i s pa%ses through Mague- P A - .-- !S w*a

yes (Almy , 1969) .- : The northenT'1imb of t h e --cline i s

r ep resen ted by t h e ~ a - . ~ a r ~ u e ; a h i l l s , and p o s s i b l y , t h e

southern l imb-by t h e t r e n d of c o r a l r e e f s on t h e s h e l f . *-

A longer exposure of t h e south limb of t h e syncl ine t o -

a t t a c k by t h e sur f zone a t t imes of low sea l e v e l s would

r e s u l t i n a n lower r e l i e f . With a ' r i s e i n sea l e v e l folio- - . - T .

wing t h e end of t h e l a s t P le i s tocene g l a c i a t i o n isco cons in),

t h e low l imestone r i d g e s on t h e she l f would have been gra-

d u a l l y submerged,q'rou)dinHp~eferred s i t e s f o r c o r a l - - - .* - - .. -_.. I .

gro;th and subsequ* - _ - r ee f - fo rmat ion (Clam, -- T 1973). --3 -

( 2 ) Kaye (1359). i s - ?~f the" opinion t h a t t h e r e e f s

' i n t h e s e a r e a s have developed on drowned, c a l c a r e n i t e

c u e s t a s , which were formed as e o l i a n i t e s t r u c t u r e s para-

l l e l t o t h e shore during3the Wisconsin g l a c i a l per iod. . -

Present seismic evidence (Morelock e t a l , 1977) f i t s

the o r i g i n proposed by Kaye (1959) f o r t h e inner she l f

provinces . However, t h e r e i s no d a t a f o r the ou te r l i n e s

of r e e f s and they may have fsrmed according t o the f i r s t

theory presented .

These r e e f systems, which a r e considered the counter-

p a r t of t h e r e e f s of La C o r d i l l e r a , Fa jardo , have been

subjec ted t o comparably l i t t l e p ressu re from indus t ry o r

development. Limited amount of r a i n f a l l , minimum runnoff

and g r e a t q u a n t i t i e s of organic mat ter cont r ibuted by

f r i n g i n g mangroves nearby have a l s o con t r ibu ted t o the

formation of t he se h igh ly developed systems. Increased

i n l and d e f o r e s t a t i o n of La Parguera l imestones, proposed

r e3o r t development, domestic waste d ischarges and close-

ness t o heavy i n d u s t r i a l a r e a s , may impose i n the near

f u t u r e , a s e r i o u s t h r e a t t o these communities.

A t t h i s p o i n t , I w i l l d i scuss one of t h e most spec- *

t a c u l a r r ee f systems of Puerto Rico: t he she l f edge r e e f .

A we l l developed submerged' b a r r i e r r ee f borders much of . - l .

t h e she l f edge south of ~ u e r t o Rico. These systems have

been most ex t ens ive ly s tud ied i n t t s southwestern s h e l f .

The top of th - i s r ee f i s shallower than f u r t h e r inshore . .. - . - -

a r e a s and a t a depth of 1 7 t o 25 m 3 &alp break i n the

nea r l y l e v e l bottom occurs , dropping away a t an angle of

up t o 45 i n t o t he Caribbean. A bu t t r e s s ed spur and

groove formation (Figs . 60, 61, 62) has been observed f o r

more than 3 km on t he she l f edge (Morelock e t a l , 1977).

Sand channels up t o 6 m deep with v e r t i c a l walls genera l ly

l e s s than 2 m wide and 20-30 n long a r e cu t i n t o the upper

i n s u l a r s lope . These have walls covered wi th encrust ing

c o r a l growth, a l g a e , and boring sponges and a r e separated

by a wide c o r a l b u t t r e s s dominated by massive co ra l and

~ n a r i c i g having l u shes t growth a t t he she l f edge (Fig. 63) .

These appear t o be some type of surge channel which allows

movement of sand from the ou t e r she l f t o t he slope (Morelock IL

e t a l , 1977). These sands form an obvious t ra i l down the

~ i g . 60. Observe

Spur and groove high doral cover

system.

Fig . 61. Idem. Spur and groove system. Note high c o r a l cover, s p e c i a l l y A a r i c i a sp . , l i n i r g + t h e s i d e s of e grooves.

Fig . 63. Diver working at t h e she l f edge. Note i n s s lope .

. 62. Idem. Note whip antib;? F#arians (black c o r a l ) . Spur end

groove system. Note whip c o r a l - ( ~ t i c h o ~ a t h e s sp . ) i n the fo re - ground. Acdwnulzted ssnd can be - seen batween- spurs .

Fig . 63. Diver working a t ' t h e she l f edge. Note i n su

s lope . i

*w <*M . 62. Idem. Note whip a n t i l a - F%arians (black c o r a l ) . Spur 2nd

groove system. Note whtp c o r a l (Stichopathes s p . ) i n t h e fo re - ground. Acdwnulzted sand can be - seen batween- spurs .

s lope below each channel and have been t r aced below 70 m .

Coral growth i s so in tense on the wal ls t h a t they some-

t imes roof over t he grooves providing an exce l l en t habi- - - tat f o r a g r e a t v a r i e t y and o u m b e ~ of f i s h . These grooves

-- . .-. .- - con t i nue n o r t h w a d fo.mihgcsh.allb~-&annels ( ~ i ~ . 64) % . . .. . . - ,- conta in ing ca lcareous sand and coyal rubble . . @ ~ i n n , 1972).

They tend t o branch and meander and terminate gradual ly 8

in-:*oral r i dges which a r e a l igned ea s t tqrwest p a r a l l e l

t o t h e she l f edge. These c o r a l . - r idges r i s e s l i g h t l y - t o - - - - - -

g&gdniws. Living c o r a l couer- i s . ..*- red*ed- . h e r e . --$-and -

flats-, -al so p a r a l l e l t o the shel f edge, occur t o the - : - - - -

nor-Fh d- t h e . ... G t ~ l - . b d g e s . These &e s l i g h t l y deeper - -

e 68n -Xhe c o r a l rtdgas and exh ib=tm-r ipp le marks.

The s l ope , sou th o f - t h e groove and spur system,

c o n s i s t s of cemented o r dead c o r a l pavement with l i t t l e

r e l i e f below approximately 30 meters. Zonation i s dras-

t i c , d e n s e s tands of gorgonians and an t i pa tha r i an s increa-

s ing downwards i n r e l a t i o n t o a decrease i n stony herma-

t yp i c c o r a l s .

Along she l f edge surveys, Morelock e t a1 (1977) -- observed a r e a s where t he upper 30 t o 40 meters of the

s lope a r e v e r t i c a l . Where t h i s occurs t h e r e a r e no grooves

and t he genera l na tu re of the submerged reef i s s im i l a r

t o o t h e r Caribbean submerged r e e f s (MacInt ire , 1972).

I t i s pos tu l a t ed t h a t the r e e f s on t h e shelf edge .r

b u i l t up a s b a r r i e r r e e f s during the Ple is tocene low sea

Fig. 64. Low- r e l i e f - sand channels north of .- shelf edg'e. . - . - - . - - .- ' . *-

Shelf edge. Note high cora l cover. I n su l a r edge. Note high c o r a l cover.

s tand (Goreau and Burke, 1966). The shel f edge was sub-

j ec ted t o i n t e r n i t t e n t su r face drainage of water accumu-

l a t e d by wave a c t i o n o r runoff during t h i s periods when

subae r i a l e ros ion occurred thus forming drainage channels.

No growth occurred on t h e f l o o r of these i nc i s i ons a s a

r e s u l t of continued scourings. As the r e e f s were drowned

by t he r ap id e u s t a t i c sea l e v e l r i s e over the l a s t 4,000

yea r s , these e ros iona l f e a t u r e s were enhanced by c o r a l

growth forming the b u t t r e s s e s . Erosional processes con-

t i nue a t the present t ime, t h e i r r a t e being g r ea t e r than

' a c c r e t i o n (by. c o r a l growth) which i s a l s o a f f ec t ed by . - T .

boring sponges ( b r e a u , 1966). Currents of s u b s t a n t i a l

v e l o c i t y (7 .5 - 13.0 cm/sec.) move through the channels

prevent ing deposi t ion of c o r a l l a rvae (Arne-son, undated) . - I . . . - . - -

, The shoreward. r i dges and sand- ' f la t s described e a r l i e r

a r e gene ra l l y shallower than t he shel f edge and probably e

were above t he su r f zone a t t he time of lower sea s tand,

t he r e fo r e no t a f f e c t e d i n t he same way a s the shel f edge

by con tac t wi th oncoming waves. They may be a more recent

phenomenon of c o r a l growth and sediment entrapment i n the

lower l y ing a r e a s of the sand f l a t s (Quinn, 1971). Goreau

(1966) ha s descr ibed a s im i l a r p a t t e r n i n Jamaica. Quinn

(1971) a l s o pos tu la ted t h a t c o r a l growth i s l u s h i e s t near

the she l f edge (Fig. 65) on top of the spurs due t o the

d i r e c t exposure t o the oncoming wave t r a i n s which provide

a con t inua l supply of n u t r i e n t s .

Coral r ee f development along the west coas t of Puerto

Rico can be considered from f a i r t o moderate i n reTation

\T A:.,-" - - to' e i t h e r t h e n o r t h e a s t o r t h e southwest ~ o a s t E ~ ~ 8 e t k e e n

Cabo Rojo and Mayaguez t h e r e a r e v a r i o u s sporad ic f r i n -

g ing r e e f s ( F i g s . 66, 67) which appear t o be s u f f e r i n g

I from h i g h t u r b i d i t y of t h e wa te r , unusua l ly s l i g b t wave 1 !

! a c t i o n and heavy l and dra inage . The broad bank t h a t l i e s

immediately o f f s h o r e n o t on ly minimizes wave a c t i o n a g a i n s t >

t h e shore by reducing wave energy but a l s o l i m i t s t h e

amount o f ocean wa te r a v a i l a b l e f o r d i l u t i n g land d ra inage . (Kaye, 1959). S i l t a t i o n , due t o sugar growing a c t i v i t i e s

and e s p e c i ~ l l y , t o i n2 reas ing i n d u s t r i a l and housing -develop- . . ment and low s a l i n i t i e s caused by t h e d i scha rge of R ~ O

Guanajibo dur ing f a l l a r e o t h e r f a c t o r s t h r e a t e n i n g t h e s e

r e e f s .(.&olelynainen and B iagg i , 1975) *and .Morelock (pe r - -: --

. , sona l c o m m ~ i c a t i o n ) . These f r i n g i n g r e e f s a r e g e n e r a l l y wanting i n s tony

- c o r a l s b u t posses s s p e c t a c u l a r dense " f o r e s t s " of l a r g e -

s i z e d gorgonians ( s o f t c o r a l s ) . Living s tony c a r a l s a r e

p r e s e n t l y be ing covered by mats of macroalgae (Kolehmainen,

1974).

Offshore r ee f a r e a s (F ig . 68) i nc lude Esco l lo Negro,

A r r e c i f e Tourmaline, Las Coronas, Esco l lo ~ o d r i ~ u e z , Cayo

Fanduco, Manchas I n t e r i o r e s , Manchas E x t e r i o r e s , A r r e c i f e

1 P e r e g r i n a , and Gal la rdo .

I T u r b i d i t y and sedimentat ion produced by resuspension

1 of l o c a l f i n e ca l ca reous sediments du r ing heavy, long-per iod

ground s w e l l s o r i g i n a t i n g i n t h e A t l a n t i c Ocean mainly Ir

dur ing w i n t e r months were found by ~ i n t r 6 n &.. ( 1973) ,

5 - .-

2 Kms - Fig. 66. Fringing reefs of the west coast . Also note

Cay0 Ratones west of Joyuda Lagoon (upper r i g h t ) .

Fig . 67. Fringing - barrier reef off Boquer6n.

2 Kms - Fig. 68. Offshore reefs of the west coast. Square marks

off spur and groove system. *

Kolehmainen (1974) andk toya (1974) t o be t h e most impor- '

t a n t f a c t o r s a f f e c t i n g A r r e c i f e Tourmaline during 1700

hour s t u d i e s a t an underwater h a b i t a t . Terrigenous sus- 3

pended sediments from t h e Cuanajibo ~ i v e r - d t h e pre-

sehce of ses ton and p lanktan i n t h e water column were

a l s o found t o inc rease water t u r b i d i t y during t h i s

s tudy. Montastrea v- a c o r a l spec ies equipped

wi th many f e a t u r e s t h a t . e n a b l e i t t o remove sediments

from i t s s u r f a c e , was found t o be the most importqpt r e e f -

.-bu&.lding c6;af - i n a r e a s a f f e c t e d most by s i l t a t i o n . :In . . , . .. . .-

c l e a r water r e k f s , t h i s . ~ spec ies i s ' a n important:-reef-buil- .- - - . -

ding spec ies ,.-but not^-the majog . . one.,- .. . - ;= .. * - - - . - - -. .. *- %-, a--

. -9 . . . A + . - . .. ?. . - . -. _..._-_ - ;i 7

, . ... - A low relief :-ur- arid' gr& systehr-uith . .. .&yndant . . - .

- . .. .;-- . . .= . ,

a n d d i v e r s e encrus t ing - c o r a l growth csa iac t e r i z e s t h e

a rea of Escol lo Negro marked i n Fig. 68. Water t r a n s - -

parency and l i v i n g c o r a l cover a r e high. shoreward, the

spur and groove system diminish i n r e l i e f gradual ly d i s a -

ppearing. Gorgonian cover i n c r e a s e s , s tony c o r a l s being

then represented by small diameter head c o r a l s .

Las Coronas i s a shallow (2-4 m) sand shoal colonized

p r i n c i p a l l y by l a rge s i zed gorgonians and occas ional massive

c o r a l s . I t extends e a s t g iv ing way t o Cayo Fanduca which

i s c o n s t i t u t e d e s s e n t i a l l y by t h e same fauna.

Manchas I n t e r i o r e s , Manchas E x t e r i o r e s , and A r r e c i f e

. Peregrina a l s o have low r e l i e f spur and groove systems

s loping more o r l e s s ab rup t ly westward g iv ing way t o a -

dense b lack coral-dominated fauna. Encrust ing c o r a l

growth wi th l a rge p i l l a r c o r a l and gargonians dominate

the shallower depths.

Escol lo ~ o d r i ~ u e z , s i t u a t e d about 1 .6 lon west of

C&o Corazones i s composed of a s e r i e s of elongated r 2 C I I.**

patch r e e f s . Large dead c o r a l heads, probably deposi ted

during storins, a r e permanently exposed. The r e s t of the

reef f l a t i s exposed at low t i d e . This reef l acks any

zonation comparable t o the o the r r e e f s mentioned. There - i s no back ree f slope or apron a s c h a r a c t e r i s t i c of southern

- r e e f s . f n s t e a d , the reef f l a t drops abrup t ly t o deeper . - A

back- ree f a r e a s where l a rge colonies of A_. palmata occur

i n t e r spe r s ed between c l u s t e r s of va r ious gorgonians.

, - . Other s tony c o r a l s a r e very - rare o r absent . Acropora ,

. . . ce rv i co rn i s and Dendroavra c i l ynd r i cus a r e cornmop t o some

ex ten t . The reef i s b e t t e r developed on the nor thern end -

where-a l a rge Po r i t e s f l a t occurs. F i r e co r a l and dead

D ip lo r i a and Montastrea heads overgrown by a lgae a r e

common. Very notable i s t he presence of c r i no id s i n

r a t h e r shallow waters. To the west i s a l s o a r a t h e r abrupt

s lope w i th very patchy c o r a l growth. Even so, t h i s i s an

a r e a of f a i r l y high r e l i e f and f i s h l i f e i s abundant poss i -

b l y due t o t he abundance of she l t e red a r ea s . The r e e f ,

i n genera l , appears t o be dying duel t o s i l t a t i o n by t e r r i -

genous c l a y s from the Guanajibo River. These observat ions

a r e based on a repor t by Schneidermann and Morelock ( 1 9 7 3 )

and on personal obse rva t i ons . ' 5

Bajo ~allar&I"l'k' a w e l l developed, r e l a t i v e l y untouched

ree f a b o u ~ 1 3 k i lometers west of Punta Agui l a , Cabo Rojo.

I t e x h i b i t s luxur ious elkhorn c o r a l growth and .an abun- . - - . . . . . . - , . .~

. . . ..- .% .. -. &

C . , . .Z.+>

.. dant f i s h fauna; - ; . - .- -. . . -

. . d -- .. -. ~ ~ r i h o f . A ~ n e a i f ~ . - . ~ e r & i n a to'-.Rmta ~ i * e f o - t h e in- -- ..- ,. . .

- -. . . . -.-- :% '-&lar &elf- $&'%en, na&w- ( l e s s thak~-17ki lometer ) and

. . - - . . .;---- - -.

.bas w e l l d ~ i i o * ' tdefs 4 its oute'r 'edge w h e r i the . .

bottom slopth,fleeply; A - Stony c o r a l s , unusual gorgonians 't

. - . - 4

--aiid. black - - coidlidlz . . - &--abundant - -- . a t depths of 1 . 5 t o 40 m , . . - . -- - . .. -- . . ~. . .-

- - . - . - z- .:-.+. . -

- : . - - :b* .,. vatge; f r - a s p a r h c y ..is-q&te-%ariable,, being inf luenced . . - . - . - . . . - - . t -. , - ' . C .-- .... -.. .~

. - - -- -:by - - the local .-- ..-. - &~rcuiagi--:+hd:the ..- . - . . . ~. . d i qchaarge of t&-nearby

-, ' ... . - - - . , . : . - - - -5% 5 ~ , * . ' - . ---.-. - --.. .L--. x ;-- - -

r i m s ; : . .-=4~. ( ~ o l c i __._ .197&,:%d. p e r ~ o n 8 ~ . o b s ~ ~ t ' C o n s ) - $ =- .. -

-. - . - - - . - . .,' ---- --.- ,. .. .- -,- -_L. ~

, + _..__: ->a - - . - - . . -- - . --. .. .- - " ;t. . --. - -. -. : - - -%,r%.

- . . : . ~ ,&f 3-.- l y e v e J o j d - : f - . . . . --:.-- ringing--=eefk;-~ntisting - . .- --.. . - . - . . . .. . - . p r i m a r i l y . . - " - . , - - .- ?. - .. _-. :- -2 .:pa: =iorLFL *.'i - -

1.- -. .- .. . . - ~,=a- iraba-bra - ~ f ' , ~ o & ' c b r a l \ .-and *= .*- . ,i.- .. . -4 >... . .* - - .r+- , -

.-- -,.: . . - . '*

- 78 - - - . -.. ..-: ,

spa r se 'gorgcmians , - k c c u r - o w h t h side -bf t he - ~ i n c b n - Peninsula from Punta ~ i g G e r o t o Punta d e l Boquer6n.

North of t h i s po in t only s c a t t e r e d , undeveloped c o r a l

growth occurs .

Off Bajura , I s a b e l a , on the nor th c o a s t t h e r e i s

an underwater cave system. This system i s poss ib ly r e l a -

t ed t o d i s s o l u t i o n of t h e ~ymamhn limestone which under-

l a y s a r i d g e of young cementgd sand dunes of t h e a r e a .

Although not considered a c o r a l r e e f , c o r a l growth, espe-

c i a l l y by Agar ic i a ( l e t t u c e c o r a l ) c o l o n i e s , i s q u i t e

dense on the o u t e r w a l l s and ledges of t h e caves (Figs . 6 9 , 70) .

There appears t o be a marked b i o t i c zonat ion , poss ib ly due -

. , - - ~

- - .- -

- .%+=.-&,=., 0 , . . .me$,:- -113 of t h e &ma: G i v e s . . .

Observe dense '

Agarlcta growth'.

'>. , L *

Fig. 71. Large dead A . palmata colony i n s i d e czve. I t s o r i g i n remains t o be s tud ied .

L --- --. -- -. --"

t o changes i n l i g h t and c u r r e n t c o n d i t i o n s , from t h e v a r i o u s ? A - d

e n t r a n c e s t o t h e i n t e r i o r of t h e s e caves .

North of t h e town of Dorado i s an e x t e n s i v e bu t h igh ly

s t r e s s e d r e e f f r i n g i n g t h e sho re (F igu re 7 2 ) . I t s r e e f -. 7 ' . - .-

.- . - .&K ' : . >.- - .. - - - . .--. -. . - .

f l a t is about deep w l e t t i e - - f a n i : . ( ~ o r g o & a ) . - . . a r e -..

ve ry abundant---(up to 9 indivi- l r p e r squ*rs m & e t ) . . - F

Predominant s tony c o r a l s a r e D i p l o r i a s t r i g o s a and D . c l i v o s a . -. . - . r .7

- *- - .

The r e e f f r o n t i s a h igh r e l i f e d a r e a w i t h many dead c o r a l

promontor ies ovirgrown,either by a l g a o o r by o t h e r c o r a l , *.. *- .

species . ~ e b t b "increases seaward and, aFout 100 meters nor th .. - _ . .. . > . . . -. %az. S f - . ~. 2 ..L1 - - - . .. .- - - : ----- :. - - . - -

of the r;ef f l a t ;J a r e s m ? ~ ; p a t c h r e e f e, a dep th of t ~ ~ e t e r s - . . . . -. - + .- -- . . . z-za-.: . . ..,?

(Roberto C a s t r o , pe r sona l communication)- ,--reef development

on t h e n o r t h c o a s t i s reduced and cemented dunes a r e t h e

most impor tan t f e a t u r e p r o t e c t i n g our s h o r e l i n e s from t h e

s eve re b u f f e t i n g of incoming high energy waves.

. SUMMARY AND CONCLUSI ONS

C b [ 1. Clockwise a round t h e I s l a n d , s t a r t i n g a t S a n J u a n , h L

r e e f s of v e r y h i g h q u a l i t y and e x t e n s i v e n e s s a r e p r e s e n t

La Cord ( i n c l u d i n g Plamino and ayo L a r g o

&: F a j a r d o ; (2) S a r g e n t Reef , Maunabo; (3) a l l o f f s h o r e r e e f s

between BahEa d e J o b o s and S a n t a I s a b e l ( i n c l u d i n g B e r b e r l a i

; and C a j a d e Muer tos) ; (4) Ra tones , Ponce ; ( 5 ) o f f s h o r e r e e f s

: of La P a r g u e r a , L a j a s ; (6) Tourmal ine and E l Negro r e e f 1

L complex, Mayaguez; and ( 7 ) submerged b a r r i e r r e e f a t t h e

t h e sou t h e r n a nd w e s t e r n i n s u l a r s h e

2.: E x t e n s i v e c o r a l ' . r e e f d e g r a d a t i o n was obse rved in. ( c l o c k -

w i s e from San J u a n ) : (1) a l l r e e f s from San J u a n t o Las

P Cabezas d e San J u a n ; (2) i n s h o r e F a j a r d o r e e f s ; ( 3 ) Humacao

. l a r r e e f -of f Puer t o . ~ a b u c ) i n s hore

kt- once r e e f s ; (6) a l l r e e f s o f f Bahia G u a y a n i l l a and Bahia

I T a l a b o a ; ( 7 ' ) a l l r e e f s o f f a nd --f r i n g ) a l l

w e s t c o a s t i n s h o r e r e e f s (from Boquerdn t o Rincdn) ; (9) r e e f s

r e e f s o f f Dorado.

t 3 . The most i m p o r t a n t s t r e s s a p p e a r s t o b e s i l t a t i o n p o s s i b l y

&due t o s e v e r a l c a u s e s such a s u p l a n d v e g e t a t i o n c l e a r i n g

a c c e l e r mated runno f f , p e r i

' and a l t e r a t i o n s i n sed iment dynamics. Other s t r e s s e s may

;be caused by d i s c h a r g e o f u n t r e a t e d o r p a r t i c a l l y t r e a t e d A

bewage i n t o t h e s e a of i n t o s t r e a m s and r i v e r s and m e c h a n i c a l b idamage by b o a t a n c h o r a g e . C o r a l e x t r a c t i o n , though l o c a l i z e d

r e s e n t s a s e r i o u s vroblem i n t h e r e e f s o f f F a j a r d o .

.A X 4J l-l

k c cd

Q, Z C Q,P, r w

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Q, G t'

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

c r e a t e s a v e r y d . i v e r s e h a b i t a . Examples o f t h i s a r e t h e

m o n o s p e c i f i c zones o f A . - p a l m a t a . Due t o t h e h i g h r e l i e f

caused by e x t e n s i v e b r a n c h i n g o f t h i s s p e c i e s , many o t h e r

f i s h and i n v e r t e b r a t e f a u n a p o p u l a t e t h e u n d e r l a y i n g areas':

T h i s s i t u a t i o n h a s been r e p o r t e d by Rogers (1977) f o r San

C r i s t 6 b a l r e e f a t La P a r g u e r a . Spur and g roove a r e a s a l s o

have a h i g h d i v e r s i t y d u e t o t h e i r h i g h r e l i e f .

7 . Observed s t r e s s e d r e e f s a r e c h a r a c t e r i z e d by a g e n e r a l

p a u c i t y of c o r a l s p e c i e s and by a r e a s o v e r p o p u l a t e d by f a s t . growing s p e c i e < s u c h a s t h e c o l o n s a l anemones o r z o a n t h i d s .

A c l e a r example a r e t h e Gugnica r e e f s . The s h a l l o w a r e a of

t h e s g r e e f s a r e somet imes 100 p e r c e n t c o v e r e d w i t h e i t h e r

Zoanthus or- h l y t h o a . To a l e s s e r - % - - e x t e n i GuHy::nifla i e e f s - . ,

. . ' p r e s e n t sirnil& c o n d i t i o n s .

8 . Another c h a r a c t e r i s t i c we obse rved i n s t r e s s e d r e e f s i s -

t h e over abundance of t h e s e a u r c h i n Diadema a n t i l l a r u m .

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

. which c o l o n i z e t h e s u r f a c e s of dead c o r a l s . A l s o , r educed

p r e d a t o r p o p u l a t i o n s ( e . g . B a l i s t e s v e t u l a ) r e l a t e d t o a

d y i n g r e e f , may c a u s e a Diadema p o p u l a t i o n e x p l o s i o n . An

example o f t h i s a r e t h e i n s h o r e r e e f s o f F a j a r d o , such a s

. Ahogado. I n c o n t r a c t t o t h i s , was t h e o u t e r n o n - s t r e s s e d . Cayo Largo which p r e s e n t e d a markedly r educed u r c h i n

p o p u l a t i o n .

9 . Colony d e a t h by b i o t u r b a t i o n i s common on h e a l t h y r e e f s .

Var ious s t a g e s o f t h i s p r o c e s s were observed a t Cayo Largo,

F a j a r d o . Large s i z e d p a r r o t f i s h were s een a c c i d e n t a l l y

knocking o f f b ranches o f i n d i v i d u a l c o l o n i e s of A . - pa lma ta .

Consequen t l y , "ho les" were made on t h e o the rw i se 100 p e r

c e n t A . - pa lmata covered zones . These "holes" a r e l a t e r

c o l o n i z e d by r a p i d growing s p e c i e s such a s M . - complanata .

10 . Gorgonian p o p u l a t i o n s were obse rved t o t h r i v e under

c o n d i t i o n s of heavy s i l t a t i o n and h igh t u r b i d i t y . Apparent ly

h e a l t h y and dense go rgon i an s t a n d s were observed on t h e

d e e p e r zones of r e e f s o f f Gugnica, Guayan i l l a , and Ponce. - 11. S tony c o r a l s ~ & e - ~ l s o o b s e r i i d t o . t h r i v e o n ~ 3 l o p i n g

s u r f a c e s i n a r e a s w i t h h i g h r a t e s of s ed imen ta t i on . Due t o

t h e n a t u r e of s l o p i n g s u r f a c e s , sed iments appear t o f low by

g r a v i t y to- the: r e e f ba se . T h i s is. .: a p p a r e n t -- dn t h e f o r e r e e f . ,

s l o p e o-f Cayo c a r i b e , o f f T a l l a b o a Bay.

1 2 - I n s h o r e r e e f s of La P a r g u e r a , a l t hough a p p a r e n t l y under -

deve loped , p r e s e n t a r e a s o f v e r y h igh c o r a l s p e c i e s d i v e r s i t y

a s shown by t h e f o r e s l o p e o f Cayo Co l l ado , La Pa rgue ra .

T h i s , however, may b e due t o t h e development of l a r g e numbers

of s m a l l - s i z e d c o l o n i e s on a v a i l a b l e , unoccupied s u b s t r a t e s -

1 3 . Areas of h igh wa t e r t u r b i d i t y due t o h igh amounts of

f i n e s ed imen t s i n s u s p e n s i o n p e r m i t c o r a l growth on sha l low

l e v e l s u r f a c e s . T h i s i s a p p a r e n t i n Humacao where A. palmat2

growth i s f a i r l y abundan t . Wave a c t i o n a t t h e s u r f a c e does

n o t p e r m i t t h e sed iments t o s e t t l e o v e r t h e c o r a l s . Living

c o r a l c o v e r , though, d e c r e a s e s s h a r p l y w i t h dep th due t o

w a t e r t u r b i d i t y . -

1 4 . Dur ing t h e end of t h e f i e l d d a t a g a t h e r i n 3 p e r i o d

(August 3 0 , 1979) two t r o p i c a l s t o r m s , one o f them (David)

of c o n s i d e r a b l e magni tude b u f f e t e d t h e i s l a n d c a u s i n g

e x t e n s i v e r e e f damage e s p e c i a l l y i n t h e s h a l l o w e r o u t e r

r e e f s of t h e e a s t and s o u t h c o a s t . Some of t h e s e r e e f s ; & e

under v e r y heavy s t r e s s and t h e i r a b i l i t y t o r e c o v e r i n t h e

n e a r f u t u r e i s s e r i o u s l y q u e s t i o n e d by s e v e r a l r e e f s c i e n t i s t s .

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

i s mandatory i f t h e e n v i r o n m e n t a l i n t e g r i t y of t h e r e e f

ecosys tem i s t o be m a i n t a i n e d i n t h e f a c e of deve lopment .

~ a k i n g i n t o a c c o u n t t h e e x t e n s i v e damdgk made b y t h e two-

ment ioned t r o p i c a l s t o r m s , i t seems r e a s o n a b l e t o s t a t e

< h a t an a d d i t i o n a l s t r e s s may s e r i o u s l y i m p a i r t h e n a t u r a l

c a p a c i t y q f - t h i s ec~ ;~s t tSm t o r e c o v e r . . , - % --

RECOMMENDAT IONS

1. P o l l u t i o n r e s e a r c h h a s a lways been w e i g h t e d towards

p u b l i c h e a l t h a s p e c t s . I t i s n o t a lways s u f f i c i e n t l y

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

background, t h a t w a t e r c o n t a i n i n g l e v e l s o f p o l l u t a n t s

which do n o t t h r e a t e n human h e a l t h d i r e c t l y a r e d e s t r u c t i v e

t o a q u a t i c communi t ies ( J o h a n n e s , 1975) . Water p o l l u t i o n

r e s e a r c h i n r e l a t i o n t o b e n t h i c m a r i n e communi t i e s s h o u l d

be promoted and sponsored by t h e Depar tment o f N a t u r a l . .

7 . ~ e s o u i c e s i n c o o r d i n a t i o n w i t h t h e . E n v i r b n m e n t a l Q u a l i t y - Board and t h e U n i v e r s i t y o f P u e r t o R i c o . T h i s r e s e a r c h

shou ld a i d i n t h e m o d i f i c a t i o n , i f n e c e s s a r y , of o u r w a t e r

- , . -.. - - - p o l l u t i o n l a i r s and k e g u l a t i o n s . . .

2 . The enac tment and e f f e c t i v e e n f o r c e m e n t of a p p r o p r i a t e

l a w s , and- t h e p u b l i c c e n s u r e of p o l l u t e r s c a n o n l y be brought

a b o u t when t h e p u b l i c i s made aware of t h e i r v a l u e . The most

i m p o r t a n t s t e p i n d e t e r r i n g b o t h t h e c o r p o r a t e and t h e i n d i -

v i d u a l p o l l u t e r i s t h u s e d u c a t i o n ( J o h a n n e s , 1 9 7 5 ) . We t h i n k

i t shou ld b e of h i g h p r i o r i t y t o t h e C o a s t a l Zone Management

D i v i s i o n t o p r e p a r e t e l e v i s e d e d u c a t i o n a l programs f o r t h e

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

impor tance of t h i s and o t h e r i m p o r t a n t c o a s t a l ecosys tems .

Severa 1 p u b l i c i n f o r n a t i o n b o o k l e t s h a v e a l r e a d y been p repared

b y t h i s D i v i s i o n .

3 . Reef a r e a s which b e c a u s e of t h e i r q u a l i t y and e x t e n s i v e n e s s

s h o u l d be i n c l u d e d among t h e N a t u r a l Reserves a r e : (1) La

C o r d i l l e r a , Fa j a r d o , ( i n c l u d i n g Palomino and Cayo Largo) ;

( 2 ) S a r g e n t R e e f , Maunabo; ( 3 ) and o f f s h o r e r e e f s between

BahEa d e Jobos and S a n t a I s a b e l ( i n c l u d i n g B e r b e r i a andc:

Ca j a d e Muertos) ; ( 4 ) Cayo R a t o n e s , Ponce; and (5) Tourmal ine

and E l Negro r e e f complex.

4 . The unde rwa te r c a v e s o f f B a j u r a , I s a b e l a , shou ld be

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

a t t r a c t i o n . S a f e t y measu re s , such a s marking i t s d i v e r s e - - e n t r a n c e s ' and e x i t s , s h o u l d be t a k e n gy this Department .

5. C o a s t a l Zone Management D i v i s i o n shou ld p r e p a r e b o o k l e t s

w i t h t h e f o l l o w i n recommendations t o d i v e r s and b o a t e r s : b

a ) Boat ancho r s - These n e v e ~ ' ' s ~ o u l d be dropped qp t o p -

. . of a r e e f . Damages by do ing t h i s a r e t h e t e a r i n g

up 05 c o r a l when i t h i t s , t h e con t i nued s l a s h i n g

of c o r a l w h i l e t h e b o a t i s anchored and t h e t e a r i n g

of more c o r a l when t h e anchor i s hau led up. A

D a n f o r t h - t y p e anchor shou ld be used by dropping i t

i n a sandy p a t c h and l e t t i n g t h e boa t d r i f t over

t h e r e e f .

b) S i t t i n g and s t a n d i n g on c o r a l - T h i s n o t o n l y c a u s e s

b r eakage of b r a n c h i n g c o r a l s b u t ab r ades and i n j u r e s

t i s s u e which c a n t h e n be i n f e c t e d by a l g a e . Holding

o n t o c o r a l c a n have t h e same consequences . Dead

c o r a l and bottom r o c k s shou ld be used f o r t h i s purpose .

c) Shell collection - P r y - b a r s s h o u l d never be used t o Z

d i s l o d g e and t u r n over c o r a l heads .

d ) Spear f i s h i n g - C o r a l i n j u r y from f l y i n g s p e a r

s h a l t s and working a f i s h i n a c a v e can be v e r y

s e v e r e .

6 . Laws r e q u i r i n g t h e r e p l a n t i n g of l and c l e a r e d of v e g e t a t i o n

have been p o o r l y e n f o r c e d i n t h e p a s t . We recommend t h a t

t h e s e a c t i o n s be moni to red by t h e s e Department i n o r d e r t o

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

7 . I n summary, we recommend t h e a c t i v e involvement o f t h i s - Depar tment i n foming . a c t i o n s : such -as d r e d g i n g , up land d e -

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

of t h e n a t u r a l a b i l i t i e s of t h e r e e f ecosystem t o r e p a i r

i t s e l f and . .recover a f t e r t h e p a s t t r o p i c ; l siorms.- The new - . --

e r o s i o n r e g u l a t i o n h e i n g p r e p a r e d by t h e C o a s t a l Zone Management

D i v i s i o n s h o u l d , by a l l means, i n c l u d e t h e c o n t r o l of upland - d e f o r e s t a t i o n .

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

p o l i c y of t h i s Department f a v o r s t h e p r e s e r v a t i o n of these most

i m p o r t a n t and endangered P u e r t o Rican n a t u r a l r e s o u r c e s .

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Table 1. - ---

P c c f : I .ocal i t y : Z o n a t i o n : Diver - : Er111i t n - : \ C o r a l : Hnin

North cas t : s i t y : b i l i t y I c o v c r : C o ~ n p o n c n t : O D S ~ R \ I A T I O N S I .

C a ho ralardo : . . : ] . a t . 10 '22 .1 ' : Reef , :&. palmata : Colonles l ~ e a v l l y s l l t ed . Gorgonlans comlnon.

I-nnl'. 65'37.2' : cres t .-- , -- I S - 1 0 ' : and

. .. :A. palmala :Very sparse growth. Many dead and overturned c o l o n l e s .

: Slope

I llend corals ~ A l o n l e s of the hydrocoral Stvlaeter common. :and Aqarlcla :

8: , .

,kleavlly s t res sed rccf probably due t o disclmrye from R l a F3)ardo.

.puwls, all, uo sa l~ l r \ l ior uoll3rulsuo= ~q pue o p ~ e l e j olu jo e~uun l ju l aql 01 enp ~ ( q l s s o d jaaJ pellls X [ 1 ~ w a j ~ 'SXYV'JY - ..

. . : I . . . .uowwoo ea~apls -3 pue Xm.iim 'q' '~ar \oa pa3npsu : sueluob~ob : . .

, . - - - - - - a s ~ e d ~ i . . ' : -- . . e e ~ mi ., - - 1 - - - -

(i ?) aoeds JOJ hul~adu~oa PUP [PJO:, . . > : X F i i e a 1

slur~ojlq~upAqe[ e w i a pue Q ~ E U I I Q ~ . j -pauljap [IaM lou s o u o ~ : D A I S S P W : . . plte : I XIW :

- ( s a l l ~ a c l o ~ d hu[bu l l& jo y3el aq) Aq UMOI{S w) palp ptre sellaqluexooz paslndxa peq @ ] e u e ( d l u o 3 ~ JO ~ 3 1 ~ 0 1 0 3 'SUIQJ h e a q 6<11~1\p )e l l jaaJ aqi J O o3o:Jns a111 uo poJnSeJlU uaoq o ~ o l ; ~ ~ ; ~ 01 esol3 salllu(lv:; ' s j a a l pal l ls A[lnrral[

~ ~ M O J ~ J as~ec's I . i speatl pue I. e . . sue!uohro3 : : - - I -

- s 3 1 L ' U I p ~ o o ~

: Op~el@J OpBbOllV

S N O I ' J . V A l 1 3 S f f U : auauodwoj : J J A U ~ I X a ! l ! q : X I ~ S : u : v : - 1 A : U O ~ ~ L ' U O Z : X 1 r [ e ~ o ' l : j d a c l

. . : 1 . .

. -- - ---- ---. .- ---

. I -- A .. -.--.---- .---. ----

-UOuILUDJ : . -. esn[ iZZ iu *H 7iFs~iiCuiC '~TeiT5ietTiui .-hi ' F i i o T l 8 l ~ i ~ i ~ ~ b l j : i & ~ ~ ' , : . EleJ03 :, -- ..

elll~iiSTI-3 '3a)l~od ';I ~ ' 5 s ~ j o ~ l i s G -2 ' 'ds ki3C~pi;$y .sulloj 1 1 ~ ~ ~ 1 : ~ s peaq pus! : . .-- pdu")tu[,[ 'daals AJ~IA o*.lu[s 'asvds JOJ ~.~c~l l l lu i l~~~u:, q51q J'J v . 1 1 ~ . ssll5l1665 'v-: - - : - - - - . i a d o ~ ~ f .. - -- - -- - .- - - ~- .---- -_-;

'Ja([JJ u l LU s' z anoclr, Itr.lcsAs J ~ J S (>ire a~oc-816 : ~i . ru lnutru : . . pallcul[ rB s l o ~ o q ~ ' A j l s ~ d ~ l p qblq &>A J" ouo" P !;I sltl.l, 'aG11.u ' '-N- put. : .

a101 DL~I IY OW~i iLcr . v lo saluolos o b ~ s l ;IJV :.IJ.P~I~ ~JL~I~,,\ ,119 IILCMIV suvluo5.1o~ : -- - .. -- auoz 01 dn spuassnp ouo:c slll j, 'uoutluos B~Gi i i . ' ! r \ i~m . i j p 1 1 ~ T 5 l j F i d 'h : I :

- . -- - - - . . . . ,

.uaqoJq A[lul l~&,cl aJc saltrolo:, jo .loclllllr(l : Fiii*i-iCi .v: , , - - _ _ . 1 .

euos : I I I Y ~ ~ J I U ~ ~ S Y q6notllle tlhlq Alah 'UOIII~IIO:, 3ijj) l i j j i i i . ; f .;i p",y : . . ., . : e l r ~ i ~ ( i ~ t f : S~.U.I~F;W~:> .Q WiiFitijk .ij rw3GSj + 3 e;;o~~ifi (( ~ l u ~ ~ t ~ l l u ~ ~ p ~ ~ t ' ~ ~ , ~ , . . : pub' 1SJJ.J : , O ' P C - S 9 ' Juo-,

lob 311q uoucu,~:, aGb~;icliiiKT . fi . p a s l * ~ ~ n . 1 ~ si~rloz om1 os.~~l, l , : J;I;JU ' , S ' O Z e O I * l t . l ~ -- , . - -------- - - - -. . . . - - S 4 l V 11 u! !'-? J..-- --

a : I lll,-l'i

, a 0pJCfL2~ 50l('ll~~lfBll),j - .~ . S N O I J . V A t I i [ S t I U : l u ~ u o d c t o ~ : 1 3 ~ 0 3 1 X l ! [ t ( l : Xl !s : - - -- - -. - . ..

* . 1 u [ \ : I - 1 : UO!)I:IIUL : X~!(VJIJ.I : j ~ 3 , l - I . . . . - .- . - .--. - - -

1 ' 5 aTquJ,

P c c f : l . o c n l i t y : 2011.i t ion : Diver-: E q l ~ i t n - : \ Coral : l a i n : s i t y : h i l i t y I covcr :Component :

. Reef . Very sl>untlantJ. pda!~. w l t h different sh,~pcs re la ted l o degree : crest : 0 : 100 : A. j a l r n a t a : OF cx.posure. Very o c c . ~ ~ s s l o n d growth o f M. cornplanata where

: "holes" i n the dense _ A . pcrlnlata occur.

: ' . i ~ l n l a k i .. Cover I s s l l y h t l y redr~ced. Thls area ends rather abrupt ly g l v l n g . : zone - - -_ : 75 , ;_A. p~lma~t 1 way to the IIIIX zone.

. . : M I X : Area o f lrledluln r e l l e f wl l l r very h l g h number of c o r a l spec le r . : zone : 1.58 : 0 . 7 2 : 25 : - : D l v e r s l ~ y lower t l ~ a n i n l l ~ c slope due l o patch iness o f cora l

: d is l r l bu l ion . M_. fl?~!l{!rls. A. palmata, P, &o_rl!m, MJ compIann!a- : and gorqonlans common.

: Slope : 1.02 0 . 6 3 : 35 : A_. eel v l rnrnk: ' Zone o f l ow r e l i e f dorn lnn~ed b y A_. s!v_icon!& o n uppkr s lope and : and qnrgo- I)y goryonl,~ns on lower slope. P. p o r i t c s very commcn . Small

. : nlans : head C O ~ ~ I ~ S common. ! . r

- -- . : Reef . : f la t . - - : P. p q r l t c s i Porl tcs blotope very w c l i developed covcrlng extense a leas.

; ~ ' b s e n c e o f Diagema and other urchlns I s no lable.

R E ~ ~ A R R S : .Reef termlnetes cas lward wlt l r 3 bare sand "halo" almut 0 meters wit le after wl i ich a very heal thy .rl, ~ l a s s ( q bed starts. 1.arye "halo" may be due t o an cxtens lve i c h l l ~ y o f a r ~ n a . N u r l ~ c r o ~ ~ s ~ u & n Conch (s!rg!!btll; '~lt~a) were p~.csr?.nl i n 1111s bed. Pauclty o f sea urchins I n l l l e reef f l a t may be due to vcry low cora l mor ta l i ty which arc ovsrgrown b y nlcjae anti qi.i~zetl I)y t i ~ c s c e c l ~ i n o i d s . W i t h i n [he r c c l c res t so&etlmes I s apporent the ef fect o f b lo lurbal lon (cot- l brcakayc b y f l s h I n th i s c,ase). 4perltlres or1 (tic clense zorlc ere then co loniznd b y M. d m p l a n a t a and p late- l lke co lon ies o f P, as_!eroldes. Several slages o f th l s process were observed. This I s a relatively untouclled reef o f great complexity whlch deserves lur lher study.

Table 7. . .

P c c f : 1 , o c n l l t y : Z n n n t i o n : P i v c r - : E q i i i t a - : \ C o r a l : blain -- i .

Ramos : s i t y : h i l i t y I covcr : C o a p o n c n t . : w e s t ) Fajardo :

--- C o o r d l n n t c s

I . n t . 18'18.9'

: Red & complanata allsdnt. Small Q. s!riqosa common. Inrge N;.fiitn~rl. . -- : cres t . -- . -- :A. palrnota ~ c o l o n l e s with dead a r e a s on thelr top parts. Colpo~t~ylle common.

: Slope . -- : -- : -- :Gorgonlans : lilgh gorgonlan dens l ty . _A. cervlcornls and S . siderea common.

: Reef . . flat . -- . -- - - . Dcad coral colonlzcd rnalnly b y several a lgal s p e c l e s . P. asteroicle: : common. Sparse _A. pa11na ta lncreaslnq westward.

RE'IAAKS: West of the slope a re dead&.cervlcornls patclies, a " t ~ a l o " about 3m wlde and a T l ~ a l a s s l a - Syrlnc~oclluni bed. Siltation evident. Llvlng . ( coral cover low.

o c o +?.- L G . " L h ( 5 2 al v '-,t

C C v c C _ o C N ,. ' 4 - . r ; e

L Z 6 N C F ' W .-- C - C , C C C: c K t .- . r -- .- c, L C t c

C Z I K W

C E C . .- - k - c m L u c m o v L 2 Z C L L C

- - u L - - c.- C; =.- 0 LC c c -

Table 10.

~ c c f : I.acnlity : Z o n n t l o n : P i v c r - : E q l ~ l t n - : \ ~ o i n l : ) ( ; t in - : s i t y : h i l i t y I cover -:Component : ~ U S L N V A T I O N S

Santlayo Humacao : - - -

r o o r d i n n t e s I . n t . 18-09.5 : Colonlcs of very dlvcr s o shapes and wl 111 no deflnlte orlentatlon.

: palrr!ata : I,onr 66.44.0' . : Thesc allernale wlth cjorgonlanr. In sk~allower areas, 4 : zone . - - . -- : 60-70 : t .pa11nata : complanata Is common. Small head corals, common.

. . - - - - - - : goryonlans : Large heads of M. 2n1111larls common.

.' . . -

. . . ' - I

. . * . . -

, . . . -?. *..

(' >3 . . Reef termlnatee w1lh n'area of very sparse gorqonlan growfh. , ,,

. ', - 0

1 . c. C7 .3

. ... -..- ( I .

Table 11. - - - -

P c c f : 1.ocnl i ty : Z n n i l t i o n : D l v e r - : E q u i t n - : I C o r a l : :I.iin : s i t y : b i l i t y I c o v c r :Cornponcnt :

G u a y n ~ n a Arroyo : . . d -- - -

T o r d i n n t e s : Rccf c r e s t fr,lgmcntod l n ~ o nunaorous palc l les where e l t t w r o f tttc 1.3 t . ]7*56.5' : Reef

Iaonp. 66'01.5 . A.pelmata : !wo prlnclpal c o ~ n p o n e n t s domlnato according to t h e d e g r e e of . . - - - - - -

: c r e s t . _ _ . - - : a n d &I. SOL- : exposure . P. p g l ! e s .,n Important reef c r c s t component 9. . . : p a n a : cc rv lco rn l s a n d M. r ~ n u l a r l g c o m m o n . 6016 o f A . pa lma ta d e a d .

: @!!!h~ta : - - , z o n e . - - : 35 _A.pnlmata Rnctluced cove r and ntany c o l o n l e e d e a d .

. : MIX .. . : -- : -- : -- : z o n e 5 : Coryon lans : Gorgonlans very smal l In he igh t . Dead A_. _cervlcornls common.

. ' . . . a:* . I

: cc rv lco rn l s : -- . -- : z o n e

. '. . . . -- I A.cervicornls : Vdry few gorgonlans p resen t . 1 ,:

R E ' I A R K S : . Southward of the d e a d A. ce rv lco rn l s z o n e tlic dcp th l r i c reases g radua l ly to (he e d g e of the s h e l f . Spa r so go rgon lans a r e p r e s e n t he re . Th l s reef Is consl<Icred a s decay lng a n d r~ r~ t rca l thy .

p c e f : L o c r l i t ~ : Z o n a t i o n : Diver - : E q t t i t a - : I C o r n l : f fair^ : s i t y : h i l i t y I c o v c r :Comltoncnt :

Las Mareas Guayama : O B S E R V A T I O N S --

L o n ~ . 6Q00.5' : Reef :Prlnclpal a lgal cotllponentv aro Dyctlopteds, Hallmeda and Dyctlota,

: f lat . -- . -- .-- :Algae ';l'hese cover a ln iod 95% of the d v ~ l l ~ l ~ l e substrdle. Shoreward the :coGer Is reduced.. ',

: Encrusting :Many over henglng ledges lndlcatlng eroslonal processes. Where $13 . -- : Slope . -- .-- :sponges and :zqne merges w l l h the reef f lat there arc extznslve p i ts made by the :algal: :sea urchln _EcIilnol~~etra lucuntw . I . '

I f ' . I

. "Fore : reef" . -- . -- .--

. . Very abundant urchln p l ts . Promontories of hlgh re l lef overgrown

: Echlnumetra : to dlfforent extent .. These probal~ly represent collapsed ledges. : luc~n_tsr-

RE' 'ARKS: 'NO l lv lng corals apart from Mll lepora _ s ~ r ~ o _ s ~ a n d ~ d ~ r . ~ s t r - e f i radlans were observed. ~u;hlt l l ty of tho water was very hlgh. No gorqonlans were observed. Shoreward from the rccf f lat Is a Gy!!issla hctl followed by n ,$./~III~~I!O hetl . Uelwecn 1118 lat lor bed ,ntl the shore Is a depression where large an~ount of broken of f ~ ~ C J J ! frdonlents c0llcc:t. (Obscrv,trlot~s basctl on 'Porrcs 1970 dntl ~ ) c r l ; o ~ ~ . ~ l v ls l ts to Iha rtron).

Table 14.

P c r f : L o c a l l t y : Z n n n t l o n : I l i v c r - : I :c \u i ta- : \ C o r ~ 1 : > l a i n : s i t y : h i l i t y I c o v e r :Componen t :

PaJnros S a l l n a s - -- - C o o r d l n a t c s

I." t . 17'55.5' : Reef I . on r . 66*15.7' : rrayn,cntatrl Into buttrc~:;sess abou t 3m In re l lef a n d o v e r w a s h .

: c r e s t . -- . -- . -- : M . c o r ~ ) p l a ~ ~ h : Q . c l b ? : ~ a n d _A.J~!I~II& common. lc tyolauna abundan t possll,ly : d u e t o t h e hlqhly she l t e red a r e a . Encrustlng sponge Gllona common

, z o n e - - - - - - : & . p I m a ta : Wel l deve loped .

: MIX z o n e : . . : gorgon lans : ; a n d s l o p e : - - . -- . -- : a n d head : ~ l m l l a r l o Morr l l los , Sa l lnas . ::

c o r a l s . ' . .. - . .

R E ' f A R K S : The north s l d e of th la c a y i s h lqhly s t r e s s e d wlth many d e a d d b d over turned c o l o n l e s . #

P c c f : I . o c n l i t r : Znnn t lon : l l i v c r ~ : E q r l l t a - : \ C o r n l : t h i n . . : s i t y : b i l i t y 1 c o v c r :Component : ' O B S E R V A T I O N S

Monl l los Sallnae :

1.3 t . 17~55.8' . . : : Reef : . Very wide zone (from nor111 ro south). Ptrttresress *bout 2m I n l , o n ~ . 66'15.3' . - - a

: crest .- - , - - : M-complana:! rel ief present. Coralline alyae avery itliportent component

: of ~ h l s znne, S , ~ n d r h . ~ n n r l s orlented rJ-S hc~ween the : bullresress. I lcad cc~ral present In the channels.

< , : pal!^!* : -- a - - - _ : A . ~ l r n a t a : Sllyhtly deeper sand rlrannels I n thls zone. No dead coral. just '

1 zone . . bare sand I n tile cl iar~nels.

.. . < :

. 1 , . . ' : M I X zone : : Hoad corals '?, M . .. clv~rnos. ~ 4 ~ ! 1 ~ l ~ ' : n q . n ; e a n d r l t e s . &. radla ns. I. rigIda. C . : and slope : -- . - - . - - : and goryo- : natans and Q. ~ & o _ s ~ c o r n r n o n .

nlans r :' ---- . T i

RE81hRKS: --- Mangrove vegetallon present

Table 16.

P e r f : 1 . o c a l l t y : Zona t i on : P l v c t - : Eq l l l t e - : I C o r a l : : la in : s l t y : h i l l t y I c o v c r :Component :

P a t a n c s S n l i n n s , : Rce V A T 1 O N S

a . h a t . 17'56.2' ,. . . : Peef 1 a n * : " o n r ' ~ 6 ! 1 7 . ~ @ . c r e s t 1 0 . 3 8 0 .35 I 39 :?. c o n p l a - &., pa lmn ta common.

: n r t : n

, . 1 :

m a t I . , . zone : 0 .36 1 0.22 40 It. pn lma ta thna cnrnrnon. Zone o f c o a r s e b a r e s and I ,ctvcen c r c s t ? : a n 8 1 ~ i - ~ a t a zone.

. - 5 . I ~ o r ~ v o n i a n s \,cry n l ~ t ~ n t l a n t a n l d o n ~ i r ~ a n t i n upper m i x zone. : I!ppcr and : 1 .25 : 0.82 6 :Coryonians : ' l :~ss ive c o r a l s dominant n e a r r e e f b a s e . F l y c c t o p h y l l i a

: I o w c r m i x : 1.64 : 0.84 1 16 : and :and Oc111ina common ' zone . . :mass ive

: c o r a l s I .

. . . . .

R E ' I A R K S : Seaward o f t h o r e r f c r c s t i s an a b r u p t s l o p e lcac l inr t o a ] ) a r c sand a r e a w i , t l ~ o c c ; ~ s s i o n a l i s o l a t e d s m a l l head c o r a l s . F a r t h e r s o u t h d c p t h d e c r c a s c s Y r n p i c l l y t o a zone where medium r e l i e f mounds o c c u r . 9 :. complannta grows on t o p o f t h c s c s t r u c t ~ ~ r e s . '

Table 18. . 4

P c c C : I . o c n l i t y : Z n n a t l o n : n l v c r - : E q u i t a - : \ C o r a l : : ! ~ i n - : s i t y : h i l i t y I c o v c r :Component :

Cabuzosos a n t e isabel -- -. -

C o a n ~ n t cs 1.a t . 7-55 . 4 1

. . : $laal l D I j ~ l o r l a cllvosi!_and D. QrJgS~for in an Inaportsnt component

I.on p . : Reef : o f I h l s ?rr?8. C a ! c i l r c o ~ ~ s dgae a l s o very cornmon. P m corlllnon. 66'23.1 : crest . -- : 40 :M.cor~~p!anata: About 30% o f the sva l lab le substrate I s uncolonlzed

. ' : la!n!ala_ : - - - - : zone : 4 5 - 7 5 1A.palrnata 1 Cover increases seaward.

:&. c_ervicor n l s : 00th cornporlents very abundant. M. ~ n n ~ j l g f i s _ . 1. g s k r o l d e s , : M I X , : and : Dendroqyra _cllyndrlcus and Co lpophy l l l s natans common. --

.a zone . --

- - ' ~ o r g o n i a n s IM. C=_~W"-OI, @. lgp.11-ckLa.g. DM?_oc111a stokess'l i. L. Hnupsg, . . end : .P. a s t e r o ! d ~ a n d several sponges comlnon.

':A. rna_r!c!= :

: Reef .. : W e l l develol,ed Po~j!cs-l,lotope and farther north a Thalass la . - - . -- : f la t :~.po~o!gs bed mlxed w l t h Porl tes. . 1 .- I I and *

: ~ h a l a s s i a .; * - -- - --

Table 20. I . , I

Perf : I . o c a l i t y : Zona t i on : nivcr- : E q ~ ~ i t a - : 1 Corn] : b la in - . . : s i t y : b i l i t y I covcr :Component : ' O B S E R V A T I O N S

Cnrdona Ponce : -6iToTtJl n o t c s .4

l.". ' 17'57.5' I.on~. 66-38. 1'

. I i igh covcr. Low rcllc,f mounds ~ l l h M. gzmrlanotd growing : crest : -- : -- I 75 I &l.compl,~na~~ on t l ~ o i r tops at s l ly l r l ly deeper areas seaward o f tho crest.

. . ! Palyt l~oa common. - . - . . -

P?!?% i : 111g1) covcr far h lyh water turbldlty present. Cover Increases . . . zone . -- - - : 20-45 I A. pal~nata scawatd.

-- . i :Goqjonlans : : gorgonlsn : :and occasslo-: Sparse yrowth. . -- :nsl A. pal~nata: I . . .

. . R ~ b I A R I ; S : Gorgonlsn zone commences seaward of the fore reef slope and continues for an unknown distance. Small head corals occur here.

' 1 ~ 1 ) JaaJ uo uoll*~abo,\ a~o1burLy 'adols j o a ~ JO qlnos. soi[,wcd U Q ~ U O ~ J ~ > ~ ( - W O O [ 01 asol3) auoz UPJUO~JOB e p l ~ : S I l V l 1 3 d

. u o w t u o s 5 S j S 5 7 ~ f ~ : -. . - - -

PUP Q S O C I J ~ A Q ~ 'fi ; I i i G i K u l v fi -IJOUI~IIOJ ~ ~ U J O : ! A . I ~ ~ - V peal1 : sueluob~oyj : 2- - - - - . auoz : 'SaSeaJ3Ul JaAozJ U P ~ U O ~ J O ~ a J J q M A[[enpu~6 Sa3llp;>J JeAOZI - ~ i ? ; ~ , l r d ' ~ :

Table 22.

P c c f : local it^ : Z n n a t i o n : D i v e r - : Ec111itn- : 1 C o r n l : Main ; s l t y : t y I c o v c r :Conlponcnt : O B S E R V A T I O N S

c a r i h c T f i l l n h n a : nay -

7 0 x 1 nn t c s - . 1 . 3 ~ . 1 7 * S t l . Z t :'Arra c o n s i s t i n p r n i n l y o f c o r n 1 rulbhle w i t h P n l y t h o a l - o n 1 - . ~ ~ * ~ ~ . 0 * ' 1 Dcnd :;10d ~ ! - l q ~ o r a c o l o n i z i n l ! p a r t s o f t h e d e a d c o r a l . - - - - : c o r a l :Isolated k T a 1 1 1 1 n t a c o l o n i e s .

: Former : , . . - : mix - - - - a zone : S p o r n d i c a l o f zone i s l ~ a r c s a n d .

: p o r r o n i a n s :

: C o r r o n i n n t l cns i t y i n c l - c i ~ s e s i n s l o p e . PI. c n v u r n q s a , : S l o p e : - - : - - . - - : Gorponinn : ! I . a n n ~ ~ l a r i s , D. c l i v o s a , and D c n d r o p y r a c y 7 m u s - . :?omSon. ' l o r t a T i t y r c x e d .

. : ..

; Rccf : Th. , lnss i a 1P,,arpc bare s a n d a r c n s . blanp:-ove i s l e t p r e s e n t . . * - - - . - - : f l a t

PE&IARKS: S l la l low p a r t s o f t h i s r c c f a r e d e a d p o s s i h l y ~ I I C t o ncnll). (lrrtlpi.r!r And s l ~ i p t r a r f i c which s t i r ~ r p tlrc d&J.'tm"ts t l ~ u s s m o t h e r i n g c o r a l s . The s l o p e , h o w r v c r , scel*s normal ~ ~ o s s i l ~ l y I , c c n ~ ~ s o s c c l i l ~ ~ c n t r ~ ~ r ~ s ( l o v n thLc; jeef I)nsce w i t h o u t h e i n g d c p o s i t c c l on c o r a l ' s s ~ ~ r f ; l c c . I s o l a t c d A . p n l i i ~ t a c n l c t n i c s i n t h e dcnd zone may 5 i l : n i f y h P ~ h e r i n d i v i d u a l t o l e r a n c e . F i s h l i Ce i s s p a r s e i n t h e d c a d zone w11iTc q u i t d , g l , ~ ~ n d a n t on s l o p c .

.E s 4 z P a c"' - z -3 0 0 0 .

3 aJc G 2 f i m z 2 - . .t " a . - E u X =?.! w ,TI 2 - 0 c z;z

-, " W " 2 E : 7: =+ - 1: c . ; ", f - * CIa =! 5: % GI U 7. :I; 7 ; <: aJ - . P2 - 2 ' c,> -, - .li . - .: <I = 0

I2 - 0-0: 2 ; -

4 - E g o

Q w c > ;

, " w o * c ; $ :- 3 ,- c '0 -

8: c c E 3 L 0 c z ' - - -0 0 .: - 271 5 0' V u =' :, c -; c c P: ' " a

V1 - 01 V)

% L, c: " C

C 0 m '.

= a ' = &.o 9 2 % 2

-w I . C) L .i

8 . a:

.'. . . ' ~ u a s a ~ d la ls l a ~ o J 6 u e w 'uol13e aAew 08 u o l ~ 3 a l o ~ d eAlssa3xs 08 anp Xlqlssod p a d o l a ~ e p [ law IOU Jaau ' : s ~ ~ v \ t ~ ~ ,

. . sur? OD. . , pu c 3 i r . . , 3 i ~ i ~ i 'UOIIPIIIS 01 anp X l ~ u a ~ e d d e ~ C F V ~ S - p -- I -- : -- :edols pue :

'3 pug JO saluo(o? peap X u t ? ~ 'mol A 1 a ~ ~ G ~ J J A O ~ : VGJjjo~a.3: : auoz x l ~ : . . . - :

- - - - ' euoz : :B IP lu (a :

o:', : u k .n .- 21: 2

E - L - 2 C

Y % cr: = .- I t -

.cL - u c 2 Y -

: L b , . . , In- : 0 -

& L , - a

< > - - L;

V) - - C

u C U C

c 0 .- r, e E .'I 0

c2 - C L. L C 0 L >.

.... - I 4 X

- U L ! - - - - C - " - ... - C

L U X > - d * Cur

.................... c d

....................

s Y c P,

? P a U1 - E 8 - 0 - '=I 51 C 0 E E 0

21 . ,z! 5 =i 9 0 . > <! 0

.................... ",I - E 0 U - 3

.................... 21

...............

.................... I I

$ i C U

z -- 7 9

Figure 7 4 il

ISLETA MARINA (*at@)

F a j a r d o , P.R.

--&uImQlEs 2 . IIth\ -THALASSlA

..*..*-

3 - nnn- MILLEPORA COMPLANATA

- PORITES PORITES

4 . f l A- MASSIVE CORAL

- ACAOPORA PALMATA

5 7 - pGARlClA

f - MILLEPORA ALClCORNlS

y-GORGONIAN V . E . ~ 3 . 3 6 1 I I I I vv 1

5 10 15 2 0 25 3 0 35

Olstancr f r o m ~ ; ; f F Ig t I n meters

Figure 75

Cayo Z ancudo (North) Fa Jardo , P. R.

Conro l ido led dead c o r a l covore d by algae

f l OM ASSIVE CORAL

, ACROPORA PALYATA

3 A G A R I C I A

V . E . 1 1 2 . 5

nnn - MILLEPORA ~ O M P L A N A T A

, PORITES PORITES

V . E . = 5 . 0

10 2 0 30 40 5 0 60 70' 80 90 100 . 110 w

2 \ Olrtonce from Reef Flot in h e t e r r CT . U1

Q I . .

. , ' . .

0.0.. - CORAL RUBBLE

v- A C R O P O M PALMATA

Dlstance from Reef f l o t in meters

gure 8 1

- MILLEPORA COMPLANATA

V -ACRO;ORA PALMATA 3-

q- e o ~ o o r i ~ ~ ~

~ ~ - M A S S l V E CORAL -

V. E . = 6

7 -4 I 1 I I I I 0

I 10 20 30 . . 4 0 5 0 60 70

r I 0'

C A Y 0 C A B U Z A S O S Santa Isabal, R I?.

4,: .., .

..:s - ,..

1 ". p $l

50 100 I50 200 . - DISTANCE FROM REEF FLAT IN METERS

Figure 87 \

I Coy0 Cordono Ponoe , P. R.

fin(\ - MI LLEPORA COUP LANATA

I;.., - PALYTH,OA L

'v - ACROPORA PALMATA . .

' - 0 0 R 6 0 N I A N , 4 -

I ' . V.E. = 6 6

0 10 20 30 40 SO 6 0 70

Figure 102 c;:- ! 'p~dfo 4 0 ~ ~ i ~ f i ~ ~ ; :., ,.I* ;\.

Cobo RoJo, P.R.

- HEAD CORAL

PORITES ASTEROIDES

V. E. 10

0 IS 3 0 ' 45 " 60 7 5

D l r l a n c e from rhore in k t e r r

Figure 103 2 .

I -- s k-otonor

Joyuda , P.R. '

2. - P O R l T E S 8 P O R l T E S

- w- ,OVERTURNED MASSIVE CORAL I - f - P. ASTEROIDES Q Y

4 - - M I L L E PORA ALCICORNIS

-HEAD CORALS e m V . E . t l O .

6 7 I I I I

0 2 0 4 0 6 0 8 0 180 I

1 C . r Wl"s -I*-*( .-. lm a- ".U~alcz .jlh Dlrionrw~frorn rhore ,in metera*. * ' '

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