Inventory of the Puerto Rican coral reefs report

[From the U.S. Government Printing Office, www.gpo.gov]

INVENTORY OF THE PUERTO RICAN CORAL REEFS

cm collection

C. GOENAGA AND G. CINTRON

REPORT SUBMITTED TO THE COASTAL ZONE MANAGEMENT
OF THE DEPT. OF NATURAL RESOURCES
C014MONWEALTH OF PUERTO RICO
1979

US artMent of Commerce
Dep ces Center Libror.V
im& Coastal Servi son Avenue
22,34 South I-lob
Charleston, SC 29405-2413

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ACKNOWLEDGEMENTS

First we want to express thanks to Miguel Nieves

who provided invaluable help in the field and in the

preparation of graphic material.

We are indebted to Miguel Canals and Roberto Castro

for fruitful reef discussions.

We thank Frank Torres for providing personal reports

concerning the reefs of Puerto Rico.

Thanks are specially due to Eladio Rodriguez and
Gaspar P6rez, crew of the RV Jean "A", for their conti-

nuous help and cooperation during a cruise around the

island for the preparation of the inventory.

We want to acknowledge Jim Trumbull and the U.S.

Geological Survey for providing aerial photographic mate-

rial.

Other persons which assisted us at different stages

of the study are: Ram6n Martinez, Luis Negr6n, JOS6

Gonz&lez-Liboy,,Tommy Armstrong, Rafael Mosquera, Kurt

Grove and Barbara Cintr6n.

Gladys Rodriguez, Annie Fontanez, Ana Rita Goldilla

typed different parts of the manuscript and Actor Berrios

and Carmelo Montalvo did part of the art work.
H6ctor Merced developed and printed the photographs

in this report.

Funding for this project was provided by National

Oceanic and Atmospheric Administration, Coastal Zone

Management Program.

TABLE OF CONTENTS

Page

ACKNOWLEDGEMENTS .................................. iii

LIST OF FIGURES ................................... vi

LIST OF TABLES .................................... xii

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

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

WORLD WIDE CORAL REEF DISTRIBUTION ................ 7

CORAL REEF DISTRIBUTION WITHIN THE ATLANTIC OCEAN. 9

ECOLOGICAL ZONATION ............................... 12

Reef apron ................................... 12

Reef flat .................................... 1'30

Reef crest ................................... 14

Acropora zone ................................ 14

Buttress - Montastrea annularis zone ......... 15

Reef fore slope - gorgonian - head coral zone 15

DIFFERENT REEF TYPES .............................. 16

Rock reef .................................... 16

Fringing reef ................................ 16

Patch reef ................................... 16

Bank or ribbon reefs ......................... 16

Barrier reef ................................. 17

Atolls ....................................... 17

ECOLOGICAL CONTROLS OF REEF GROWTH ................ 18

Physical and Chemical Determi nants ........... 18
Light intensity ......................... 18

Temperature ............................. 19

Page

Salinity ................................ 19

Atmospheric exposure .................... 19

Water turbulence ........................ 20

Sedimentation .......................... 20

Storms .................................. 21

Pollution ............................... 21
Upland deforestation .... ........... 22

Sanitary effluents .................. 23

Thermal effluents .................. 23
Dredging ............................ 24
Bombing ............ o................ 24

Biological Determinants ................. 25

CORAL REEF DEVELOPMENT IN PUERTO RICO ............. 26

SUMMARY AND CONCLUSIONS ........................... 53

RECOMMENDATIONS ..................... o............... 59

BIBLI OGRA PHY. . o.......... o ......... o.o..o ......... 63
APPENDIX - DETAILED TECHNICAL DATA .............. o.... 69
Tables ........... o................. - ....... 69

Reef profiles .................................. 107
Figures ................ o........ 0 ............. 138

LIST OF FIGURES

Page

Figure 1 Diver making an underwater profile.... 138

Figure 2 Location of aerial photographs ........ 5

Figure 3 Average winter and summer surface iso-
therms in the Atlantic Ocean .......... 10

Figure 4 Average winter and summer surface iso-
therms in the Caribbean Sea ........... 10

Figure 5 Dense M. complanata stand on reef
crest ................................. 139
Figure 6 Reef crest ............................ 139
Figure 7 Abundant colonial anemones (zoanthids)
in the reef flat ...................... 139
Figure 8 A. palmata zone. Note 100% cover ..... 140

Figure 9 Large A. palmata colony sheltering
grunts and goat fishes ................ 140

Figure 10 A. palmata stand marginal to buttress
Zone .................................. 141

Figure 11 Distinct morphological shape of the
staghorn coral ........................ 141

Figure 12 M. annularis buttress ................. 142

Figure 13 Diver in buttress zone ................ 142

Figure 14 Brain coral in buttress zone. Observe
goby lying over the coral (middle) .... 143

Figure 15 Colony of the pillar coral Dendrogyra
cylindricus ........................... 143
Figure 16 School of surgeon fish in the buttress
zone. Observe buttress to the right
of the picture ......................... 144

Figure 17 Lobsters sheltering within crevice in
buttress zone ......................... 144

Figure 18 Large staghorn colony sheltering a
hamlet ................................ 145

Page

Figure 19 Gorgonian or soft coral in the fore
reef slope ............................ 145

Figure 20 Sponge and massive corals in the fore
reef slope ............................ 146
Figure 21 Turtle in the fore reef slope ......... 146
Figure 22 Vegetation denuded area ("haloll)
around reef or rock outcrop ........... 147

Figure 23 Line of rock reefs (Trending east to
west) offshore San Juan ............... 148
Figure 24 Heavily silted reefs off Punta Igle-
sias and Punta San Agustin (center).
Punta Vacia Talega can be seen at the
left of the picture ................... 149
Figure 25 Fringing reefs off Punta Miquill6and
Punta Picua ........................... 150

Figure 26 Punta Percha with fringing reef protec-
ting Luquillo Beach ................... 151

Figure 27 Reef fringing area from Cabeza Chiqui-
ta to Cabo San Juan ................... 152

Figure 28 Icacos, westernmost islet of La Cor-
dillera ............................... 153

Figure 29 Palominos and Palominitos (southern
sandy islet) off Fajardo .............. 154

Figure 30 Cayo Largo, off Fajardo ............... 155

Figure 31 Islets and reefs off Fajardo. From
the bottom Isla de Ramos, Cayo Ahogado
and Isleta Marina (Cayo Obispo and
Cayo Zancudo) are seen ................ 156

Figure 32 Reefs projecting east of Punta Barran-
cas and Punta Mata Redonda ............ 157

Figure 33 Isla Pifleros and Cabeza de Perro off
Medio Mundo, Ceiba .................... 158
Figure 34 Cayo Santiago (upper right) and Cayo.
Batata (center) off Humacao. Several
submerged patch reefs, including Bajo
Parse, can be seen south of Cayo San-
tiago ................................. 159

Vii

Page

Figure 35 a) Annular reef off Yabucoa Bay ...... 16o
b) Non vertical aerial photograph of
the same reef ..................... 160

Figure 36 Fringing reef off Punta Figueras ...... 161

Figure 37 Arrecife Guayama south of Punta Figue- -
ras ................... o ............... 162

Figure 38 Arrecife Las Mareas off Guayama (lower
right) ................................ 163

Figure 39 Cayos de Barca., Jobos ................. 164
Figure 40 Reefs off Salinas. ................. - 165

Figure 41 Reefs off Santa Isabel ................ 166

Figure 42 Cayo Berberia off Santa Isabel ........ 167
Figure 43 Isla Caja de Muertos off Ponce ..... - 168

Figure 44 Cayo off Ponce. Arrecife Ratones can
be seen at left .................. oo ... 169

Figure 45 Stressed reefs off Tallaboa ........... 170
45a Ship stationed between Cayo Rio and
Cayo Palomas in Tallaboa Bay.... ...... 171
Figure 46 Reefs off Punta Verraco, Guayanilla ... 172
Figure 47 Fringing and patch reefs off GuAnica.. 173

Figure 48 Reefs protecting Playa Ca?ia Gorda,
GuAnica .......... o ..................... 174

Figure 49 La Parguera. ........................... 175

Figure 50 Patch reefs off Isla Cuevas ........... 176

Figure 51 El Palo - Atravesado reef ............. 177

Figure 52 Margarita reef ................ o ....... 178

Figure 53 San Cristobal reef .................. o. 179

Figure 54 Laurel reef ...........;.......... o ..... 180

Figure 55 Media Luna reef ......... o ........ o .... 181

Figure 56 Enrique reef., .......... o ........... o. 182

viii

Pa ge

Figure 57 La Gata Caracoles reef ........................... 183

Figure 58 Enmedio reef ...................................... 184

Figure 59 Mario reef ........................................ 185

Figure 60 Spur and groove system. Observe high coral cover.. .. . 186

Figure 61 Idem ............................................. 186

Figure 62 Idem. Note whip antipatharians (black coral) ......... 187

Figure 63 Diver working at the shelf edge ...................... 187

Figure 64 Low relief sand channels north of the shelf edge ....... 188
Figure 65 Shelf edge. Note high coral cover . .................. 188

Figure 66 Fringing reefs of the west coast . .................... 190

Figure 67 Fringing - barrier reef off Boquer6n. 191

Figure 68 Offshore reefs of the west coast. Square marks off spur
and groove system. 192

Figure 69 Bajura caves ...................................... 193

Figure 70 Outer walls of the Bajura caves. Observe dense Agaricia
growth . .......................................... 193

Figure 71 Large dead A. palma colony inside cave. Its origin ...
remains to @e studied . ............................. 193
Figure 72 Reef off Dorado ........................... o ....... 194

Figure 73 Profile Northea st C oa st, C abo San jua n ................ 107

Figure 74 Profile Isleta Marina ............................... 108

Figure 75 Profile Cayo Zancudo (North) ........................ log

Figure 76 Profile Cayo Ahogado ............................... 110

Figure 77 Profile Palominitos (South) ........................

Pa ge

Figure 78 Profile Cayo Largo ................................. 112

Figure 79 Profile Ramos (West) ............................... 113

Figure 80 Profile Ramos (East) ................................ 114

Figure 81 Profile Pifieros .................................... 115

Figure 82 Profile Cayo Caribe ................................ 116

Figure 83 Profile Arrecife Las Mareas ......................... 117

Figure 84 Profile Ca yo Ra tones ............................... 118

Figure 85 Profile Cayo Alfeftique ........................... 119
Figure 86 Profile Cayo Cabuzasos ............................ 1.20

Figure 87 Profile Cayo Cardona .............................. 121

Figure 88 Profile Cayo Ratones ............................... 122

Figure 89 Profile Cayo Caribe ...... : ..........................

Figure 90 Profile San Crist6bal ........................... 0.. 124

Figure 91 Profile Enrique ................................... 125

Figure 92 Profile Cayo Laurel ............................... 126

Figure 93 Profile Cayo Turrumote I ........................... 127
Figure 94 Profile Cayo La Gata ............................... 128

Figure 95 Profile Cayo Margarita ............................ 129

Figure 96 Profile Cayo Ahogado .............................. 130

Figure 97 Profile Cayo Media Luna ........................... 131

Figure 98 Profile Cayo La Conserva .......................... 132

Figure 99 Profile Cayo Collado ............................. 133

Pa ge

Figure 100 Profile Enmedio ................................... 134

Figure 101 Profile Punta Guaniquilla ........................... 135

Figure 102 Profile Punta Ostiones ............................. 136

Figure 103 Profile Ratones .............................. 137

LIST OF TABLES

Page

Table 1. North east coast Cabo San Juan .......... 69
Table 2. Isleta Marina (East) .. ................. 70
Table 3. Zancudo (North) ......................... 71
Table 4. Ahogado ................................. 72
Table 5. Palominitos (South) ..................... 73
Table 6. Largo ................................... 74
Table 7. Ramos (West) ............................ 75
Table 8. Ramos (East) ............................ 76

Table 9. Piiieros ................................. 77
Table 10. Santiago ................................. 78

Table 11. Guayama ................................. . (9

Table 12. Cayos Caribe ............................ 80

Table 13. Las Mareas .............................. 81

Table 14. Pajaros ................................. 82

Table 15. Morrillos ............................... 83

Table 16. Ratones ................................. 84

Table 17. Alfeftique .... #......................... o 85
Table 18. Cabuzasos ................................ 86

Table 19. Cayo Frio ............. o............ o.... 87

Table 20o Cardona ................. o ............... 88

Table 21. Ratones ................................. 89

Table 22. Caribe ....... o...... o............... o... go

Table 23. San Cristobal .................. o....... o 92

Table 24. Enrique ....... o ......................... 93

A i

List of Tables (cont.) Page
Table 25. Laurel ................................. 94

Table 26. Turrumote I............................ 95

Table 27. La Gata ................................ 96

Table 28. Margarita .............................. 97
Table 29. Las Pelotas ............................ 98

Table 30. Cayo Ahogado ........................... 99

Table 31. Media Luna ............................. 100

Table 32. La Conserva ............................ 101

Table 33.. Collado ................................ 102

Table 34. Bajo Enmedio ........................... 103
Table 35. Guaniquilla ............................ 104

Table 36. Punta Ostiones ......................... 105

Table 37. Ratones ................................ 106

xiii

INTRODUCTION

The importance of the coral reef ecosystem has been

mentioned and documented on numerous occasions by several

writers. In summary:

a) Reefs are among the most biologically productive

ecosystems containing a great variety of benthic

organisms and providing a habitat for large num-
bers of juvenile fish of many species. Reefs

shelter and support the majority of fish and

crustaceans that are commercially extracted from

our coastal processes.
b) Reefs provide a buffer against seas pounding

shorelines and moderate currents thereby influen-

cing the deposit and maintainance of sand on

beaches. As an example, the calm waters and depo-

sited sands of Luquillo Beach would be lost if

the protecting reefs were to disappear.
c) Fragments from dead coral or from other calcium

carbonate producing organisms which inhabit reefs

are the principal components of many Puerto Rican

beach sands. Many islets are also formed by the

deposition of these.
d) Many organisms inhabiting.reefs,, such as algae
and certain soft corals, produce chemical subs-

tances with valuable medicinal properties.
e) Reefs are a recreational resource for skin and

SCUBA diving.

f) Due to the fact that reefs are the most exten-

sive coastal communities of the earth and due

to their complexity in terms of taxonomical

diversity and trophic relations, these are of

unlimited interest in the study of the dynamic

relations of biological processes.

The interest of this report is to give a general des-

cription, status and localization of our reef resource

and to stimulate in the beginner science students the

curiosity and desire to initiate ecological studies of

this most important and threatened ecosystem. We also

attempt to create a consciousness of the alarming rate

of degradation of this practically non-renewable resource

in the layman and to promote their right to publicly oppose

projects which may further endanger the latter. Finally,

we intend to delineate guidelines, based on the analysis
of our and others' observations, for the management of

the.coral reefs in the best social interest,

This document initially shows coral reef distribution

in the world and especially in the Caribbean Sea. Following

is a mention of the physical and biological conditions

necessary for the development of coral reefs. Then, the

most common zonation patterns and the different reef types

are discussed. Reef distribution along the Puerto Rican

coasts is described based on published literature, reports

and personal observations. Next are our' conclusions and

recommendations. Included as appendices are profiles of selected reefs

and tables containing detailed data.

METHODS AND MATERIALS

This reef inventory was carried out during a one

year period. Observations were limited to a maximum of

one day per reef and measurements were made only whenever

sea conditions, e.g. high water transparency and reduced

surge and wave action, permitted it.

Topographic profiles of the forereef were made with

a line marked every five meters. This line was laid,

perpendicular to the shore, from the closest area of the

reef flat that sea conditions permitted to the windward

base of the reef. Depth measurements and observations
were made at five meter-intervals (or less where pertinent)
and recorded on an underwater slate (Fig. 1).. Self con-
tained underwater breathing apparatus (SCUBA) was used

on areas where use of a snorkel was not appropriate.

The site of the profile was chosen after analyzing

aerial photography of the area and after making an under-

water reconnaissance of the reef. Occasionally a diver

was towed on an underwater sled to cover larger areas.

The area most representative of the forereef was selected.

Coral species diversity, equitability and living

cover were made with line transects according to a method
designed by Loya (1972) and later modified by Rogers (1977).

All transects were 10 m long and parallel to depth contours.

These were run with the aid of SCUBA.

I Black and white photographs were taken with a Nikonos

III camera with a 35 mm lens.

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Fig. 2. Location of aerial photographs.
AMCCI-0

Reefs where profiles were not made due to diverse

reasons, are described in the Coral Reef Development in

Puerto Rico section.

All aerial photography is vertical unless otherwise

specified. The locality and area of cover is shown in

Fig. 2. The approximate areas where underwater profiles

were made is shown with a line. Corresponding sections

of nautical charts are included with each vertical aerial

photograph.

Available north coast aerial photography was analyzed

.and areas where coral growth was suspected were spot-checked

in the field with the aid of a boat towed underwater sled.

The section Detailed Reef*Information includes:

1) The fore reef profiles showing the distribution of
the most common species within-each zone and 2) Tables

mentioning specific details of the zones or of the reef

as a whole.

The coral reef inventory does not include the islands

of Mona, Vieques, Culebra and Desecheo. For information

on the reef fauna of the first three the reader is reffered
to Cintro@n and Thurston (1975), Torres (1972) and Cintro'n
et al (1974) respectively.

WORLDWIDE CORAL REEF DISTRIBUTION

Hermatypic or reef building corals, which are the

main components of the coral reef, flourish in tropical
zones with seawater temperatures ranging from 25-290C.

These areas are situated within a belt roughly bounded

by the Tropics of Cancer and Capricorn, imaginary lines

which are drawn around the earth 23.5* north and south

of the equator. Within this 4800 kiiometer-wide belt,

coral reefs are abundant on the eastern shores of the

.Americas, Africa and Australia, but far less common off

.the western shores of these continents. Why is this? In

the Northern Hemisphere ocean currents are forced into

a clockwise circular movement by the earth's rotation

and associated wind distribution (Coriolis effect). The

opposite is true in the Southern Hemisphere. As a result,

warm water travels toward the poles along the eastern shores

of the continents. On these shores, therefore, there is

a much wider extension of warm water suitable for vigorous

coral growth. On the western shores the reverse is true.

Cold waters running toward the equator, combined with

upwellings of cold water from the depths, greatly restrict

the extent of shoreline favorable to coral growth.

Non hermatypic or non-reef building corals, contrary

to hermatypic onesare by no means'.restricted to the tropics

and can be found in the cold seas lying within the Artic

and Antartic circles as well as in the Norwegian fjords,

the rocky shores of the United States, Canada, and the

coasts of England and France. None of these cold-water

corals, however, grow so actively or to such a size as

the larger reef corals of the tropics. Their form is

usually small and delicate rather than large and massive
(Smith, 1948).

DISTRIBUTION WITHIN THE ATLANTIC OCEAN

Most coral reef development in the Atlantic Ocean

is restricted to the Caribbean and adjacent areas in the

southern Florida and the Bahamas. Although viable reef
production is nill north of southern Florida (due to low
winter temperatures), isolated hermatypic corals can
survive as far north as Cape Hatteras (35 N, see average

winter and summer temperatures in the Atlantic Ocean and

the Caribbean in Figs. 3, 4). One notable exception is

Bermuda which contains a surprisingly large number of

corals, but still lacks some of the most prolific West
Indian species such as Acropor palmata (Stoddart, 1969).

Also, these reefs appear to be only thin encrustations

over Pleistocene rock (Milliman, 1973). The Gulf of

Mexico is basically an area of terrigenous sedimentation,--

but some scattered marginal reef growth is present near

Veracruz, Mexico, together with many relict coral and
algal mounds throughout much of the Gulf shelf (Milliman,
1973). The northern porti on of this area, roughly that

area north of a line passing through Progreso, Mexico,

and Havanal Cuba, can be considered to be marginal tropics
in that winter temperatures commonly fall below 220C and

coral reef development is poor as compared with the central

and southern Caribbean.

Coral reef development in the Atlantic Ocean reaches

its southern limit off Rio de Janeiro, Brazil. These

communities are, how-ever, biologically quite distinct from
those of the Caribbean and., as in Bermuda, lack many of

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the principal constituents of the latter (Volcker, per-

sonal communication).

For a detailed summary of coral reef research in

the Caribbean, the reader is referred to Colin (1978).

ECOLOGICAL ZONATION

Identifiable ecological zones are created by the

degree to which the various physical and topographical

factors positively or negatively affect reef organisms

throughout the reef area. The characterization of each

zone may, therefore, be based on the name of one key or-
ganism and/or the name of the physical factor or feature

estimated to be of greatest importance to the ecology of

the zone. Overlap or combination of zones may occur

anywhere as a result of the varied effect of tidal currents,

wave action or bottom structure.

Following is a general scheme of reef zonation (lee-

ward to windward). It should be borne in mind that this

varies from location to location and even within the

same area.

(a) Reef apron

The reef apron is the area of sediment accumulation

leeward of the reef flat. It is generally a barren area

in the sense of epibenthic biota although in some cases

it is extensively burrowed by shrimps or holothurians

(sea cucumbers) which form mounds that sometimes cover

most of the barren sand areas (Mathews, 1974). It may
also contain beds of marine phanerogams (flowering plants)
such as turtle grass (Thalassia) and/or patch reefs.

Sand is primarily biogenic as in the case of Cayo Enrique,

La Parguera, where its principal constituents are mainly

coral and coralline algae fragments (Morelock et al,

1977).

(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 :@hough 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.

. During severe tropical storms, large quantities of

living coral are dumped into this area forming islets of

considerable heights above sea level.

As depth decreases in this 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 this area are generally sturdy

encrusting and small branching such as Porites asteroides,

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

Favia fragum, Diploria clivosa, Agaricia agaricites, and

Siderastrea radians. Corals present in this zoneare

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

to sedimentation and their distribution is influenced

by the presence of adequate substrates, such as coral

fragments, which favor their establishment.

Thalassia beds and their associated biota are also

common in this area.

This is the zone of highest wave energy and water

movement. Prominent organisms include Millepora compla-
nata (fire coral) and several species of zoanthids (co-
lonial anemones) (Fig. 7). Sometimes this zone extends

above the low tide level forming emergent communities.i

Under slightly lower energy regimes Millepora may be

replaced by the elk horn coral Acropora palmata.

Other organisms, very conspicuous in this are, include

the encrusting gorgonian Erythropodium caribaeorum, various

species of calcareous red algae and sea urchins.

Even though a discussion of this subject is out of

the scope of this report, it should be mentioned that

recent studies have confirmed the existence of emergent

algae ridges, similar to those present in several Indo-

Pacific reefs which replace the Millepora zone (Glynn,
1976). This algal ridge is best developed in areas of

very high wave energy.

(d) Acropora zone

This zone extends seaward from the bottom edge of

the Millepora zone and generally is monospecific contai-

ning, almost exclusively, the elk horn coral (Figs. 8,9,
10,11). Various forms of this coral are present depending

on the degree of exposure.

Other corals and different algae species may grow

on the protected substrate beneath the branches of the

elk horn coral.

(e) Buttress - Montastrea annularis zone

Seaward of the Acropora zone there is usually a

small drop off where massive heads or buttresses of the
star coral Montastrea annularis occur (Figs. 12, 13).

Very large isolated colonies of the elk horn coral are

also present thus creating an area of very high relief.
Soft corals or gorgonians, brain corals (Fig. 14 )and
pillar coral(Fig. 15) also abound here.

This area with all its steep and narrow channels,

canyons, tunnels and towering coral heads provide a great
variety of habitats for reef organisms (Figs. 16, 17) and

is generally the area of highest biotic diversity. Large
patches of A. cervicornis shelter numerous fish (18).
M Reef fore slope - gorgonian - head coral zone

Beyond the buttress zone, the reef flattens out
towards its base Gorgonians (Fig. 19) are usually the

most frequent life form in this area, sometimes forming

dense forests. Between the gorgonians encrusting coral

such as Montastrea cavernosa, M. annularis, Siderastrea

siderea, Diplori labyrinthiformis.and Porites asteroides

occur. Other corals present are Isophyllia multiflora,

Isophyllastrea rigida,, Mycetophyllia lamarkiana, and

Mussa angulosa. Sponges are also common here (Fig. 20).
A green turtle was observed in th is zone (Fig. 21). Usually,

corals growing near the base of the slope are heavily

silted and dead colonies are frequent.

Small patch reefs sometimes occur beybnd the base

of the slope. In areas where the reef base is shallower

than about seven meters a contiguous sea grass bed is
present. Normally a "halo" or non vegetated str-.ipe occurs

adjacent to the reef base as a consequence of reef fish
grazing (see Fig. 22).

Different reef types

(a) Rock reef

Rock reefs are shallow eolianite platforms thinly

veneered by stony corals. These will be discussed upon

the description of Puerto Rican north coast.
(b) Fringing reef

Fringing reefs grow marginal to the coast and

are separated from the latter by a shallow lagoon genera-

lly not exceeding a couple of meters and sometimes nearly

exposed to the atmosphere.

These are isolated coral colonies usually surroun-

ded by a sandy bottom and occuring.close to shore. They

are irregular in shape.

(d) Bank or ribbon reefs

Bank reefs are developed on calearenite cuestas

or on drowned synclines. These will be discussed in detail

in the description of the southwest coast.

(e) Barrier reef

These are usually emergent reefs separated from

and by a deep and wide lagoon.

W Atolls

Atolls are oval shaped reefs rising from deep

water and surrounding a lagoon in which there is little

or no land. They develop, usually but not always, on

the gradually subsiding cones of extinct volcanoes.

Barrier reefs and atolls are not found within the

Puerto Rico shelf or in its vicinity although these types

are found in the Caribbean. The best known and largest

barrier reef in the Caribbean flanks the coast of British

Honduras (Belize) which stretches for more than 200 km

and has lagoon depths of more than 20 m. The best known

Caribbean atolls are Hogsty Reef, Alacran Reef, Light house
Reef, Glover'.s Reef and Serrana Bank. These are found

off the coast of Honduras, Belize and the Yucatan Peninsula.

These definitions should be used carefully due to

the common occurrence of intermediate forms.

ECOLOGICAL CONTROLS OF REEF GROWTH

The environmental controls of local coral growth

and of general reef distribution have been summarized

by Stoddart (1969).

Physical and chemical determinants

(a) Light Intensity
Most hermatypic (reef building) corals grow

at depths of less than 25 m and maximum growth rates

appears in depths less than 10 m. This has been explaiqed

.in terms of the increasing concentration of suspended
sediments with depth (Wood-Jones, 1910) but the critical
control seems to be illumination (Gardiner, 1930). This

decrease in light intensity with depth limits photosyn-

thesis by the symbiotic algae present in coral endodermal

tissue which play animpprtantrole in the process of calcium

deposition.

It should be point ed out here that recent observa-
tions of deep reef -slopes by means of submersibles have
shown that certain species of hermatypic corals extend

up to 100 m depth though in general, they cease to be
framework constructors at about 70 m (Lang, 1974; Colin,
1974; Lang et al., 1975; Ginsburg and James, 1973, 1976;
Neumann and Ball, 1970; Porter, 1973).

Light penetration is limited by turbidity as well

as depth. Weak reef growth, especially in shallow waters

on leeward reefs, has been explained by turbidity 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 about 360C. 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 preclud ing coral growth or killing established

corals.

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 atmosphere 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

reef flat organisms coincident with midday, low water

exposures in La Parguera, Puerto Rico, has been reported
by Glynn (1968). As with other detrimental factors,

coral survival varies with the species.

(e) Water turbulence

Wave turbulence and energy are certainly impor-
tant controls of coral growth. Storr (1964), based on

studies of the Abaco reef tract, Bahamas, concludes that

wave thrust is the environmental factor which results in

ecological separation of the various reef organisms.

Apparently, extreme turbulence or extreme absence of it

is detrimental for certain species of corals. While water

movement has an important direct physical effect, it is

also responsible for bringing supplies of fresh water and
nutrients to corals. Removal of C02 is related also to
degree of water turbulence (Stoddart, 1969).

(f) Sedimentation

Sedimentation as a control of reef growth in
Puerto Rico has been stressed recently by workers such
as Cintron et.al (1973), Kolehmainen (1974), Loya (1976),
and Rogers (1977). Branching corals are relatively better

able to withstand se-dimentation than massive corals; others

especially with large polyps, have developed efficient
mucus and ciliary cleaning mechanisms (Yonge, 1935; Mar-
shall and Orr, 1931). The role of sedimentation in island-

wide coral distribution in Puerto Rico has been discussed

by Kaye (1959) and Almy and Carrion-Torres (1963).

(g) Storms

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 al (1965) reported on minor hurricane Edith in
1963 in Puerto Rico. Winds less than 90 km/hr. caused

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 coast and especially
on the southern coa"stal 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 f ormer.
Stoddart (1963) also reported very extensive reef

damage on the Britis'h 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

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

may 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

effect 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

of nutrient-laden 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

runoff has probably been the single greatest cause of reef
destruction historically (Johannes, 1975). Undoubtedly

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

that bad land management has greatly magnified the problem.
Maragos (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,

St. Thomas, most corals and other sessile animals have

been destroyed and conditions remain unsuitable for their

establishment due to sedimentation caused by bulldozing,

construction and the surfacing of land that drains into
the bay. A subsequent survey (van Eepoel et al., 1971)

indicated that conditions were rapidly worsening. Damage

to reef communities due to accelerated terrigenous sedi-
mentation has also been observed in Tanzania by Ray (1968)
and in the Seychelles by Vine (1972).

(2) Sanitary effluents

The relation between organic or nutrient enrich-

ment and lowered, stressful oxygen levels is well known.
Johannes (1975) has pointed out that tropical marine organisms

live closer, on the average, to their lower oxygen limit
than biota in colder waters, Kinsey (1973) observed that

oxygen levels artificially depressed only slightly below

normal levels on a healthy reef caused the death of many

reef inhabitants. Depressed oxygen levels in reef and

near reef environments subjected to sewage effluents have
been reported by Bathen (1968) and Wade et al. (1972).

(3) Thermal effluents

Tropical organisms live at temperatures only a
few degrees below their upper lethal limit (Mayer, 1914).

Consequently, the threat of destruction or alteration of

marine communities by overheating is greatest in the Tropics..

Heated effluent from a power plant in Guam led to extensive

destruction of reef corals (Jones and Randall, 1973).
Jokiel and Coles (1974) reported on the impact of thermal

effluent on corals in Hawaii. Nearly all corals in water
4-50C above ambient were killed.

(4) Dredging

Brock et al. (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 commu-

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

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 also 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
(Hubbard, 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

Vieques, P.R. is given by Rogers et al. (1978).

Biological determinants

Bioerosion or the presence of certain organisms

which bore into the reef framework have been shown to

control reef growth in the deep fore reef of Bahamas,
Jamaica and Belize (Lang, 1974; Colin, 1974; Neumann and

Ball, 1970; Porter, 1973; Lang et al., 1975; Ginsburg
and James, 1973, 1975). An example of this are the boring

sponges which weaken coral skeletons and hold fasts, thus

influencing the size and growth forms of corals which
live at their lower depth limit (Hartman and Goreau, 1970).

Other biological processes such as competitive inter-
actions, bioturbation (small scale distruptions) and

predation also assume an increasing influence.-on commu-

nity structure in deeper and more diverse reef assem-
blages (Glynn, 1976; Lang, 1973).

CORAL REEF DEVELOPMENT IN PUERTO RICO

Individual coral colonies grow almost everywhere in

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

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

the most important factor inhibiting coral growth. The

importance of salinity and light penetration 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 ne.xt, 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 Espifritu Santo. Kaye also mentions that because

of the large storm waves that rake the north coast from

time to time, reefs growing on sandy or muddy bottoms

in relatively shallow water have particularly unstable

foundations. Most known reefs in Puerto Rico are in sha-

llow water and a majority of them rest on a platform

with depths that are within the range of large storm

waves that would stir up the sand and mud and, in general,

both destroy the foundation of young reefs that are struggling

to establish themselves and smother them in the stirred-

up sedirqent.

Another factor may be the possible bevelling of the

shelf area by intense wave action which removed any pro-

jections above the bottom that would provide a su itable

place for reef development.

It should be mentioned here that intensive upland

deforestation during the last 30 years has probably in-

creased sediment runnoff aggravating this situation.
The possible presence of "luxurious deep reefs"

between 30 and 100 meters in the north coast of Puerto

Rico has been mentioned by Kclehmainen and Biaggi (1975)

based on the presence of these in the north coast of

Jamaica. We, however, question this assumption due to

the very different wave regimes and much lower precipi-

tation on the Jamaican north coast'..

East of San Juan lies a discontinuous chain of poorly
developed and heavily stressed rock reefs trending in a

general east-west direction and extending close to 1.5 km
offshore (Fig. 23). These, from their alignment with

eolianite ridges are interpreted to consist of a relatively

thin coral veneer growing on a shallow eolianite platform
which, in some cases (e.g. Isla Piedra, and Isla Cancora).
rise above tidal level (Kaye, 1959).
Off of Punta Las Marifas reef patches occur which

typically are moundlike and rise to within a couple of

meters of the surface. The center of th e top of these

small mounds consist of head corals rimmed on its borders

by the elkhorn coral (Acropora palmata) and on its lower

slopes by sea fans and other gorgonians.

Northwest of Boca de Cangrejos lies what was a well

developed reef system. Here, extensive coral formations

were common from the surface down to 10 meters in fairly-

clear waters. This reef was virtually destroyed by sedi-

ments derived from extensive dredging within Torrecilla

Lagoon and organic sediments discharged into this same

lagoon by sewage treatment plants. At present, barely

any living coral exists below approximately 1.5 m.

Stony corals are present on the rock reefs and beach
rock platform at Punta Vacifa Talega (Fig. 24). Most are

encrusting growths and not major contributors to reef

construction or maintainance. These are most abundant

along the northern side of the inner reef.,, Millepora

complanata is the most abundant coral near the surface

of these rock reefs. Diploria and Isophyllia are also

common on deeper areas. Soft corals are present on pro-

tected areas.

Scattered patch reefs breaking the surface are found

between Punta Iglesias and Punta San Agustin (Fig. 24).

Even though they do not form a continuous barrier, they

have formed an effective wave energy absorbing structure.

At present the patches adjacent to shore are dead probably

by siltation. Water conditions here are characterized

by high levels of suspended particles and low visibility.

Water quality and health of corals increases offshore,
living corals being present only in the shallow (1-3 m)

depths of the outermost reefs. Signs of sharp erosion

are evident at the base of the reefs.

Farther east, on the north and west side of Punta
Miquillo and on the north and east side of Punta Picuea,

are fringing reefs which average about a quarter of a
mile wide (Fig. 25). It is probable that both Punta

Miquillo and Punta Picua were formerly sand cays developed

from these reefs, which since have been tied to the main-

land by a broad marsh and narrow sand tombolos. These

two reefs, specially.that of Punta Miquillo, are in very

poor health situation with very low living coral cover

and diversity. The Punta Miquillo.reef has suffered

serious perturbations and partial destruction of its com-

ponents as a result of the dredging of a channel paralle I

to the shoreline which affected the structural integrity

of the reef framework and created a silty environment,

due to resuspension by wave action, which is detrimental

to corals. Punta Percha is part of the same system exhi-

biting similar conditons but slightly higher living

coral cover.
Ensenada Comezo'n (Fig. 25) is lined with numerous

patches covered principally by algae. Stony corals cover

these to a lesser extent. The patches, more than a couple

of meters in relief, present no distinct zonation. Coral L

species present are M. lamarckiana, h,. agaricites, M. squar-

rosa, P. asteroides, and P. strigosa. Surrounding waters

are generally very turbid.

Two large patch reefs occur offshore from the mouth
of RIfo Mameyes. Both are roughly circular (300-500 m. in-
diameter) and with an exposed shoal of coarse sand (mainly
Halimeda) and.broken pieces of coral. They consist mainly

of shallow grass beds incised by north-south trending

sand channels. The underwater edge of the island is

fringed with a narrow band of coral of which less than

20 per cent is alive. The seaward edge also contains en-

crusting coralline algae. The fore reef consists of a

steeply sloping coral pavement, sparsely covered with

living corals and terminating in a.barren sand flat. Corals

present are Acropora palmata, Montastrea annularis, @j. caver-

nosa., and Diplori strigosa. The general low diversity

here is apparently due to silting by the out flow of Rio

Mameyes. These islets and their surroundings were heavily

impregnated with Bunker oil during an oil spill in December

21) 1978.

East of this reef system there is a complex of barrier,

fringing and patch reefs which are responsible for the
formation of Luquillo Beach (Fig. 26). Some of these

probably rest on rock foundations with the exception of

those northwest of Luquillo which occur at considerable

distances.from rock out crops and therefore may be built

on sand or mud. These reefs have undergone various stages

of degradation. The fringing reefs surrounding the northern

and eastern end of the beach show degradation in the sea-

ward edge where growth has been limited to the upper

three meters.

East of Luquillo water transparency increases gra-

dually and reef exhibit slightly higher living coral cover.
East of Rio Juan Martien are a series of patch and
fringing reefs which have been described by Torres (1973).

Coral species diversity here is low,,@@. palmat , S. siderea,

D. strigosa, and M. cavernosa being the most common species.

The areas of highest abundance, namely the outer reef

flats, had 30 per cent coral coverage. Torres did not

observe significant coral growth below 3 m depth. Reduced

illumination, caused by silt particles in suspension,

appears to be the limiting factor. The seaward edge is

characterized by overhanging ledges. In some areas

these ledges have collapsedcreating crevices and cave-

like structures. The presence of uneroded dead coral

masses, mainly of A. palmata, suggest a recent death.

Pumping activities of water accumulated in sand extrac-

tion pits was observed by Torres. These activities are

an additional stress to these already stressed coral colo-

nies.

A reef system fringes the area comprised between
west of Cabeza Chiquita to Cabo San Juan (Fig. 27).

This reef is also undergoing rapid degradation from the

effects of siltation and also from systemic extraction

of corals for sale. The northernmost portion of this
bay has a very shallow reef platform (part 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,

in terms of diversity, high living coral cover, and extensive-

ness. The islets of La Cordillera, specially Icacos and
probably all (Kaye, 1959) are not reef constructions but

are composed of oolitic eolianite which was deposited and

partially submerged some time previous to the development

of the reefs. Most of these are high islands and generally
possess land and/or beach vegetation. Rock reef fringes

two thirds of Icacos north shore. Corals here cover less

than 50 per cent of the available surface area (Mckenzie.
and Benton., 1972). Off the southwestern shore of Icacos

Js a more protected area which exhibits fairly high coral

development. South of the main line of La Cordillera reefs

but still on the same platform there lie the Palominos
(Fig. 29) complex and Cayo Largo (Fig. 30), two other reefs

of high biotic quality and extensiveness. Between these

and the mainland lie some other islets (Fig. 31) posse-

ssing highest reef development on their eastern shores.

Palominos and Isla de Ramos are the tops of partially
submerged hills according to Kaye (1959). Ridges of the

former continue south and east as large shallow submerged

banks. The degree of development appears to be related

directly to the distance from the mouth of Rio Fajardo.

Palominitos,, Isleta Marina, and Cayo Ahogado have been

formed by wave deposited sand and coral fragments atop

a reef platform. These rise only less than 3 m above

sea level and are susceptible to occasional drastic wave

erosion, especially Palominitos and Ahogado.

Fringing the mainland, an extensive but dying reef

borders the coast from northeast Cabo San Juan to the

north end of Punta Sardinera (Fig. 27). The entrance
to Bahia Las Croabas is protected by this reef. From

Playa Sardinera to Punta Barrancas there are no coral

reefs fringing the coast probably because of the influence
of the Rlto Fajardo which carries significant amounts of

silt. Narrow coral reefs, however, project eastward about
450 m from Punta Barrancas and Mata Redonda (Fig. 32).
There is a shallow reef in the northern Bahia Demajagua

which rises abruptly from about 2 m to the reef crest

(about .3 m). Coral growth and reef development is not

extensive. Going westward there is a deep boulder coral

zone which merges into a Acropora zone and is followed

by the reef crest where fire coral predominates. West-

ward is a grass bed with thick mounds of P. porites and

extensive zoanthid carpets. Sea fans and other gorgonians
are present east of the boulder zone (McKenzie and Benton,

1972).

Off Medio Mundo,, Ceiba, is Isla Pifieros (Fig. 33)

with moderate coral growth on its northern and eastern

coasts and Cabeza de Perro. This latter islet was used

by the U.S. Navy for bombing practices and marine life

is wanting.

South of this point up to Punta Lima the coast is

fringed principally by Thalassia meadows although occasional

small fringing and patch reefs occur.

Some of these fringing reefs of the east coast pro-

bably rest on sand or mud foundations judging from their
location at the edge of tidal swamps (Kaye, 1959). These,

for the most part seem to grow from a 6-7 meters deep

platform. Also, many patch reefs that do not reach inter-

tidal level occur off this stretch of coast.

The high levels of light penetration, typical of

the south eastern end of the island, finds its limit at

Punta Lima. West of this point the coastal waters become

turbid due to a series of sediment laden rivers and creeks

that sharply reduce its transparency. The first signi-
ficant one is Rio Anton Rulfz which is occasionally dredged

and pumped. Another source of siltation is the recent
development of Palm-as del Mar at Humacao which has cleared@_-

the vegetation in extensive areas of steep slopes and loose

soil. Erosion here has been so extensive that deep scars,

which have resulted in damage to beach and coastal commu-

nities, are evident in the washed soil. Here occur several

islets such as Cayo Santiago and Cayo Batata (Fig. 34)

which present some coral growth specially in shallow
waters and in areas open to sea (facing south). Surpri-

singly dense 90 per cent living elkhorn coral stands inter-

mingled with gorgonian and head corals occur close to the

surface. Coral cover diminishes drastically with depth

giving way to areas of very sparse soft coral growth. It

appears that water movement caused by the incoming waves

is sufficient to keep coral colonies free of sediment

but at the same time water transparency is too low to

permit coral growth except in the shallow depths. Sub-

merged shoals with sparse coral growth also occur occa-

sionally off Humacao. An example of this is Bajo Parse
(Fig. 34), which consists of numerous gorgonians, small

head corals and extensive patches of the encrusting
sponge (Anthosigmella varians. Depth is never more than

about 5 m.

Adequate ecological conditions for the successful

growth of corals do not exist at present in most areas

of the Yabucoa Bay. The annular reef located at the
southern part of the bay (Fig. 35) is not an exception

and, in spite of th efact that a few living corals are

found there, several other biological indices from orga-

nisms thriving there indicate that environmental condi-

tions existing at present interfere with the normal

growth of corals. These conditions seem to be the influ-

ence of fresh water coming from the rivers, ravines, and

creeks which empty into the bay and resuspension of fine

sediments by propellers from the-heavy ship traffic.

Characteristics of this annular reef are: (1) the scattered

growth of A. palmata and many other dead corals, (2) the

macroscopic algae colonizing the dead or dying corals,
(3) the luxurious growths of large benthic marine algae
among the corals, (4) the establish of marine phanerogams,

mainly Cymodocea among the corals, (5) the scarcity of

reef Foraminifera and the presence of Foraminifera not
characteristic of coral reefs (Difaz-Piferrer, 1969, and
Seiglie, 1969). About 5.5 nautical miles east of Yabucoa

Bay there is a reef called La Conga by native fisherman.

This reef was not visitedbut presumably it forms part

of the submerged barrier reef which borders most of the

southern shelf of the Island which I am going to describe

later.

Sargeant Reef, '.3 km' southeastward of Punta Tuna

is 1.8 miles long and .@ .1 km wide at its widest point.

Because it breaks the force of the southeast swell, the

reef affords protection from the southeast for the shore-

line in the vicinity of Punta Tuna where the reef is from

.12 to .18kn- from shore. This reef is of a high quality

in terms of living coral cover and diversity. It presents

a reef flat with abundant A. cervicornis thickets. Following

south is an area of reduced A. palmata growth with high

encrusting algal cover. This area gives way to a very
dense Porites biotope (with patches larger than 100 square
meters in some areas) which alternates with overwash colo-

nies of A. palmata. Farther seaward is an area of dense,

100 per cent cover of A. palmat which is reduced gradually

south merging with an area where gorgonians predominate.

A fringing reef extends almost continually for four

miles along the coast between Cabo Mala Pascua to Puerto

Patillas. This is exposed at low tide and protects a low

sandy apron which lies at the foot of the Sierra de

Guardarraya. A similar highly stressed reef is respon-

sible for the seaward protection of Punta Figueras
(Fig. 36). Partially responsible also for the protection
of this area and Puerto Arroyo is Arrecife Guayama (Fig. 37)
about.6 to .9 km .off Punta Figueras. This reef is
very extensive (nearly 5 kilometers) but is partially

affected by siltation rendering an average low living

coral cover. The reef flat of Arrecife Guayama is frag-

mented in small buttressess with fire of elkhorn coral

on their tops depending apparently on the degree of expo-

sure. The finger coral is also an important component

of these structures. The Acropora palmat zone has a

very low cover and high'mortality of standing colonies.

Seaward is a zone of small sized gorgonians and dead stag-
horn coral (A. cervicornis) which increases its depth

gradually up to the shelf edge. Westward of this reef

are the'Corona and Algarrobo patch reefs which appear
relatively healthy and not much affected by siltation.

South of Las Mareas, Guayama, lies Arrecife Las Mareas
(Fig. 38) which is nearly totally devoid of living coral.

The death of this reef is probably due to siltation by
Guamanil River east of Punta Ola Grande. Tongues of silt

have been observed spreading predominantly westward from
the mouth of this river (Torres, 197@). High seas and

heavy surf action prevailing in this open beach maintain

the silt in suspension. Furthermore, the rock jetty

protecting the entrance to Puerto Las Mareas, blocks the

westward movement of the silted waters retaining them on

the eastern side. Extensive dredging operations took

place here for the construction of the artificial harbor

at Las Mareas and may have contributed to high sediment

levels.

Southwest of Punta Pozuelo extends a fringing-barrier

reef called Cayos Caribe for a distance of about 2.5 km.

As part of the same reef system, but divided from it and

.each other by shallow channels are Cayos de Barca and
Cayos de Pajaro (Fig. 39). These form a distinct arc
that effectively protects the entrance of Bahl'a de Jobos.

On the lee side of these reefs, specially Cayo Caribe,

are a score of narrow sand cays fringed by mangrove vege-

tation and oriented normal to the reef margin which are

separated from each other by drainage channels. These

channels drain the water that washes over the higher,
outer margin of the reef to the bay (Kaye, 1959). Living

coral cover is moderate and increases westward.

Numerous offshore keys occur south of Salinas (Fig. 40)
and Santa Isabel (Fig. 41). These are, west from Cayos

de Pajaros, Cayo Morrillos, Cayos de Ratones, Arrecife

Media Luna, Cayo Alfeftique, Cayos de Caracoles, and Cayo

Cabuzasos. These are in very healthy state with high

living coral cover and support rich benthic and nekton

41.

populations. Most of them except Media Luna have mangrove

vegetation in various stages of development. Crescent-

shaped Cayo Alfefiique, with its arms projecting to the

northwest, shows theimportance of the southeast winds and

the northwest and westerly currents in the molding of these
features (Kaye, 1959).
Cayo Berberifa (Fig. 42), west of Cayo Cabuzasos, has

an extensive fringing reef lying on its eastern and

southern shores. Coral development reaches its maximum

on the southern shore where the A. palmat zone reaches

a 95 per cent cover and there is an extensive fish fauna.

There is a relatively untouched mangrove forest on the

lee side of this cay. Southeast of Berberia is a small,

.submerged reef called locally Las Cervezas which has

extensive elkhorn coral coverage and dense gorgonian

stands. Fish life is especially abundant here.
About .6 km...-southeast off Ponce is Isla Caja de
Muertos (Fig. 43) which presents highest ree f development
on its northeastern shore. Specially notable of this

reef is its complex high relief lagoon which supports

a large variety of benthic and nektonic fauna.

Off Ponce lie numerous mangrove islets fringed by
coral reef (Isla del Frio, Isla de Cardona, Isla de Ratones,
and Cayo Cardona) (Fig. 44). These'present stands of

Acropora palmat surprisingly dense for these silt laden

waters. Scleractinian coral growth, though, is sparse

in deeper waters where gorgonians dominate.

About 2.5 km south of Ponce is also Bajo Tasmanian,

an area of prolific coral growth. Bajo Tasmanian con-
sists of a two leveled platform (Beach, 1975). The nor-

thern level ranges in depth from 6 to 12 meters and the

southern between 18 and 24 meters. The staghorn coral
(Acropor cervicornis) is particularly abundant over much

of thislower level. At the shelf edge of this lower

level are large shingle-like growths of various massive

corals.
The industrial development of Bahl'a Guayanilla and

Tallaboa have significantly altered the natural coastal

features and offshore reefs of the area. Reefs off

Tallaboa (Fig. 45) are, at present, under high stress

conditions due mainly to siltation by periodic dredging
of ship channels propeller stirring (Fig. 45a) and also

to the discharge of bilge water containing oily wastes

into the water as well as the industrial effluents which

are presently descharged into this bay. Living coral

cover in shallow reef areas off Tallaboa is close to zero

with some isolated heads of Acropora palmata and Millepora

complanata still surviving in the seaward side. Generally

these zones of dead coral are continued south by sparse

gorgonian growth and then by slightly higher living,

massive coral cover at the reef slope where the sediment
apparently does not accumulate. Off Punta Verraco (Fig. 46)

is a reef which has an extensive Thalassia and Syringodium

bed on its reef flat. Stony coral cover in the shallow

front reef is very reduced with the zoanthid Palythoa
covering most of the CaC02 framework. In the deeper fore
reef is an extensive and quite healthy community of the

possibly more tolerant soft coral or gorgonians.

An extensive submerged reef surrounds the coast from

Punta Ventana to Punta Vaquero, where it breaks the sur-

face here first as a fringing reef and later as a barrier

reef that protects Playa Tamarindo, Bahia de la Ballena
and Playa de CaTia Gorda (Figs. 47, 48). This reef is

almost totally devoid of living coral and huge living

carpets of the fast-growing colonial anemones Zoanthus.

and.Palythoa lie ov er the dead coral framework.

West of Punta Jorobado coral reefs become more pro-

lific and complex forming a series of provinces and brea-

king the surface as far as two nautical miles offshore.

These are the La Parguera reefs which not only protect

other important littoral communities such as mangroves

and Thalassia meadows but serve as their foundation

(Figs. 49-59).

Here, between the shelf edge and the coast, two

elongate reef systems, aligned approximately east-west

divide the shelf into an inner, middle and outer shelf.

Two theories have been presented to explain the pos-

-sible origin of La Parguera reefs, these are:

(1) The major geological features of La Parguera

are a result of deformation of upper Cretacous limestones

(with interbedded mudstones and volcanic rocks) into a

WNW-ESE trending syncline whose axis passes through Mague-
yes (Almy, 1969). The northern limb of the syncline is

represented by the La Parguera hills, and possibly, the

southern limb by the trend of coral reefs on the shelf.

A longer exposure of the south limb of the syncline to

attack by the surf zone at times of low sea levels would

result in a lower relief. With a rise in sea level follo-

wing the end of the last Pleistocene glaciation (Wisconsin),

the low limestone ridges on the shelf would have been gra-

dually submerged, providing preferred sites for coral
growth and subsequent reef form ation (Glynn, 1973).
(2) Kaye (1959) is of the opinion that the reefs ------

in these areas have developed on drowned, calcarenite

cuestas, which were formed as eolianite structures para-

llel to the shore during the Wisconsin glacial period.
Present seismic evidence (Morelock et al, 1977) fits
the origin proposed by Kaye (1959) for the inner shelf

provinces. However., there is no data for the outer lines

of reefs and they may have formed according to the first

theory presented.

These reef systems, which are considered the counter-

part of the reefs of La Cordillera, Fajardo, have been

subjected to comparably little pressure from industry or

development. Limited amount of rainfall, minimum runnoff

and great quantities of organic matter contributed by

fringing mangroves nearby have also contributed to the

formation of these highly developed systems. Increased

inland deforestation of La Parguera limestones, proposed

resort development, domestic waste discharges and close-

ness to heavy industrial areas, may impose in the near

future, a serious threat to these communities.

At this point, I will discuss one of the most spec-

tacular reef systems of Puerto Rico: the shelf edge reef.

A well developed submerged barrier reef borders much of

the shelf edge south of Puerto Rico. These syst.ems have

been most extensively studied in the southwestern shelf.

The top of this reef is shallower than further inshore

areas and at a depth of 17 to 25 m a sharp break in the

nearly level bottom occurs, dropping away at an angle of

up to 45 into the Caribbean. A buttressed spur and
groove formation (Figs. 60, 61, 62) has been observed for
more than 3 km on the shelf edge (Morelock et al, 1977).

Sand channels up to 6 m deep with vertical walls generally

less than 2 m wide and 20-30 m long are cut into the upper

insular slope. These have walls'covered with encrusting

coral growth, algae, and boring sponges and are separated

by a wide coral buttress dominated by massive coral and
Agaricia having lushest growth at the shelf edge (Fig. 63).

These appear to be some type of surge channel which allows
movement of sand from the outer shelf to the slope (Morelock
et -al. 1977). These sands form an obvious trail down the

slope below each channel and have been traced below 70 m.

Coral growth is so intense on the walls that they some-

times roof over the grooves providing an excellent habi-

tat for a great variety and number of fish. These grooves

continue northward forming shallow channels (Fig. 64)
containing calcareous sand and coral rubble (Quinn, 1972).

They tend to branch and meander and terminate gradually

in coral ridges which are aligned east to west parallel

to the shelf edge. These coral ridges rise slightly to

shallower depths and are covered with dense stands of

.gorgonians. Living coral cover is reduced here. Sand

flats, also parallel to the shelf edge, occur to the

north of the coral ridges. These are slightly deeper

than the coral ridges and exhibit ripple marks.

The slope, south of the groove and spur system,

consists of cemented or dead coral pavement with little

relief below approximately 30 meters. Zonation is dras-

tic,dense stands of gorgonians and antipatharians increa-

sing downwards in relation to a decrease in stony herma-

typic corals.
Along shelf edge surveys, Morelock et al,(1977)

observed areas where the upper.30 to 40 meters of the

slope are vertical. Where this occurs there are no grooves

and the general nature of the submerged reef is similar
to other Caribbean submerged reefs (MacIntire, 1972).

It is postulated that the reefs on the shelf edge

built up as barrier reefs during the Pleistocene low sea

stand (Goreau and Burke, 1966). The shelf edge was sub-

jected to intermittent surface drainage of water accumu-

lated by wave action or runoff during this periods when

subaerial erosion occurred thus forming drainage channels.

No growth occurred on the floor of these incisions as a

result of continued scourings. As the reefs were drowned

by the rapid eustatic sea level rise over the last 4,000

years, these erosional features were enhanced by coral.

growth forming the buttresses. Erosional processes con-

tinue at the present time, their rate being greater than
accretion (by coral growth) which is also affected by
boring sponges (Goreau, 1966). Currents of substantial
velocity (7.5 - 13.0 cm/sec.) move through the channels
preventing deposition of coral larvae (Arneson, undated).

The shoreward ridges and sand flats described earlier

are generally shallower than the shelf edge and probably--

were above the surf zone at the time of lower sea stand,

therefore not affected in the same way as the shelf edge

by contact with oncoming waves. They may be a more recent

phenomenon of coral growth and sediment entrapment in the
lower lying areas of the sand flats (Quinn, 1971). Goreau
(1966) has described a similar pattern in Jamaica. Quinn
(1971) also postulated that coral growth is lushiest near
the shelf edge (Fig. 65) on top of the spurs due to the

direct exposure to the oncoming wave trains which provide

a continual supply of nutrients.

Coral reef development along the west coast of Puerto

Rico can be considered from fair to moderate in relation

to either the northeast or the southwest coasts. Between

Cabo Rojo and Mayaguez there are various sporadic frin-
ging reefs (Figs. 66, 67) which appear to be suffering

from high turbidity of the water, unusually slight wave

action and heavy land drainage. The broad bank that lies

immediately offshore not only minimizes wave action against

the shore by reducing wave energy but also limits the

amount of ocean water available for diluting land drainage
(Kaye, 1959). Siltation, due to sugar growing activities

and especially,,to increasing industrial and housing develop-
ment and low salinities caused by the discharge of R110

Guanajibo during fall are other factors threatening these
reefs (Kolehmainen and Biaggi, 1975) and Morelock (per-

sonal communication).

These fringing reefs are generally wanting in stony
corals but possess spectacular dense "forests" of large
sized gorgonians (soft corals). Living stony corals are
presently being covered by mats of macroalgae (Kolehmainen,

1974).

Offshore reef areas (Fig. 68) include Escollo Negro,

Arrecife Tourmaline, Las Coronas, Escollo Rodriguez, Cayo

Fanduco, Manchas Interiores, Manchas Exteriores, Arrecife

Peregrina, and Gallardo.

Turbidity and sedimentation produced by resuspension

of local fine calcareous sediments during heavy, long-period

ground swells originating in the Atlantic Ocean mainly
during winter months were found by Cintro'n et al. (1973),

Kolehmainen (1974) and Loya (1974) to be the most impor-

tant factors affecting Arrecife Tourmaline during 1700

hour studies at an underwater habitat. Terrigenous sus-

pended sediments from the Guanajibo River and the pre-

sence of seston and plankton in the water column were

also found to increase water turbidity during this

study. Montastrea cavernosa, a coral species equipped

with many features that enable it to remove sediments

from its surface, was found to be the most important reef-

building coral in areas affected most by siltation. In

clear water reefs, this species is an important reef-buil-

ding species, but not the major one.

A low relief spur and groove system with abundant

and diverse encrusting coral growth characterizes the

area of Escollo Negro marked in Fig. 68. Water trans-

parency and living coral cover are high. Shoreward, the

spur and groove system diminish in relief gradually disa-

ppearing. Gorgonian cover increases, stony corals being.

then represented by small diameter head corals.

Las Coronas is a shallow (2-4 m) sand shoal colonized

principally by large sized gorgonians and occasional massive

corals. It extends east giving way to Cayo Fanduca which

is constituted essentially by the same fauna.

Manchas Interiores, Manchas Exteriores, and Arrecife

Peregrina also have low relief spur and groove systems

sloping more or less abruptly westward giving way to a

dense black coral-dominated fauna. Encrusting coral

growth with large pillar coral and gorgonians dominate

the shallower depths.
Escollo Rodr1guez, situated about 1.6 km west of

Cafio Corazones is composed of a series of elongated

patch reefs. Large dead coral heads, probably deposited

during storms, are permanently exposed. The rest of the

reef flat is exposed at low tide. This reef lacks any

zonation comparable to the other reefs mentioned. There

is no back reef slope or apron as characteristic of southern

reefs. Instead, the reef flat drops abruptly to deeper

back reef areas where large colonies of A. palmata occur

interspersed between clusters of various gorgonians.

Other stony corals are very rare or absent. Acropora

cervicornis and Dendrogyr cilyndricus are common to some

extent. The reef is better developed on the northern end

where a large Porites flat occurs. Fire coral and dead

Diploria and Montastrea heads overgrown by algae are

common. Very notable is the presence of crinoids in

rather shallow waters. To the west is also a rather abrupt

slope with very patchy coral growth. Even so, this is an

area of fairly high relief and fish life is abundant possi-

bly due to the abundance of sheltered areas. The reef,

in general, appears to be dying due to siltation by terri-

genous clays from the Guanajibo River. These observations
are based on a report by Schneidermann and Morelock (1973)

and on personal observations.

Bajo Gallardo is a well developed, relatively untouched

reef about 13 kilometers west of Punta Aguila, Cabo Rojo.

It exhibits luxurious elkhorn coral growth and an abun-

dant fish fauna.

North of Arrecife Peregrina to Punta HigUero the in-
sular shelf is very narrow (less than 1 kilometer) and

has well developed reefs.at its outer edge where the

bottom slopes steeply. Stony corals, unusual gorgonians

and black corals are abundant at depths of 15 to 40 m,

but water transparency is quite variable, being influenced

.by the local circulation and the discharge of the nearby
rivers (Colin, 1978, and personal observations)-.

Poorly developed fringing reefs, consisting primarily
of partially dead Acropora palmata (elkhorn coral) and
sparse gorgonians, occur on the north side of the Rinc;n@:@
Peninsula from Punta Higaero to Punta del Boqueron.

North of this point only scattered, undeveloped coral

growth occurs.

Off Bajura, Isabela, on the north coast there is

an underwater cave system. This system is possibly rela-

ted to dissolution of the Aymamon limestone which under-

lays a ridge of young cemented sand dunes of the area.

Although not considered a coral reef, coral growth, espe-
cially by Agaricia (lettuce coral),colonies, is quite
dense on the outer walls and ledges of the caves (Figs. 69, 70).

There appears to be a marked biotic zonation, possibly due

to changes in light and current conditions, from the va-

rious entrances to the interior of these caves.

North of the town of Dorado is an extensive but

highly stressed reef fringing the shore (Fig. 72). Its
reef flat is about 1-3 meters deep where sea fans (Gor-
gonia) are very abundant (up to 9 individuals per square
meter). Predominant stony corals are Diploria strigosa

and D. clivosa. The reef front is a high relief area

with many dead coral promontories overgrown either by

algae or by other coral species. Depth increases sea-

ward and about 100 meters north of the reef flat are

small patch reefs at a depth of 25 meters with abundant

fish fauna.

Except for patchy coral growth and several minor
coral assemblages in Arecibo (Torres, 1978) and submergeid@-
patch reefs off Camuy (Torres, 1978) and off Puerto de
Tortuguero (Roberto Castro, personal communication), reef

development on the north coast is reduced and cemented

.dunes are the most important feature protecting our shore-

lines from the severe buffeting of incoming high energy

waves.

SUMMARY AND CONCLUSIONS

1. Clockwise around the Island, starting at San

Juan, reefs of very high quality and extensiveness are
present in: (1) La Cordillera (including Palomino and
Cayo Largo), Fajardo; (2) Sargent Reef,,Maunabo; (3) all

offshore reefs between Bahia de Jobos and Santa Isabel

(including Berberia and Caja de Muertos); (4) Ratones,
Ponce; (5) offshore reefs of La Parguera, Lajas; (6) Tour-
maline and El Negro reef complex, Mayaguez; and (7) sub-

merged barrier reef at the edge of the southern and

western insular shelf.

2. Extensive coral reef degradation was observed
in (clockwise from San Juan): (1) all reefs from San
Juanto Las Cabezas de San Juan; (2) inshore Fajardo
reefs; (3) Humacao reefs; (4) annular reef off Puerto
Yabucoa; (5) inshore Ponce reefs; (6) all reefs off
Bahia Guayanilla and Bahia de Tallaboa; (7) all reefs
off and fringing GuAnica; (8) all west.coast inshore
reefs (from Boquer6n to Rinc6n); (9) reefs off Arecibo;
and (10) reefs off Dorado.

3. The most important stress appears to be silta-

tion possibly due to several causes such as upland vege-

tation clearing which leads to accelerated runnoff,

periodic inshore dredging, and alterations in sediment

dynamics. Other stresses may be caused by discharge of

untreated or partically treated sewage into the sea or

into streams and rivers and mechanical damage by boat

anchorage. Coral extraction, though localized, presents

a serious problem in the reefs off Fajardo.

4. Rapidly degrading and heavily stressed coral

communities of the rock reefs of the north coast, although

not nearly as complex and diverse as southern, eastern,

and western reefs, are important in the sense that they,

by active biological growth, protect the integrity of

the structures which they overlay.

5. The underwater caves off Bajura, Isabela, are

very interesting features from a biological and geolo-

gical point of view. Coral communities fringing the various

entrances to these caves are extensive and may play an

important ecological role.

6. Present observations show that even though

occurrence or distribution of certain species within

the reef are related to several principal determinants
(e.g. wave action, light intensity, etc.), zonation pat-

terns are very diverse around the island and even vary

within discrete locations. The principal zones of one

reef can be compressed, widened or simply non existent

in another. Species coverage within the same zone of

two different reefs can vary widely. Zones with very

low cover can be very diverse due to high equitability

or equal coral distribution.. Zones of very high coral

cover can and often do have very low diversities due to

the almost complete dominance by a single species. Ano-

ther factor that has to be taken into consideration is

vertical relief. Areas of very low coral species diver-

sity can still have high total diversity due to high

relief which creates a very diverse habitat. Examples

of this are the monospecific zones of A. palmata. Due

to the high relief caused by extensive branching of

this species, many other fish and invertebrate fauna

populate the underlaying areas. This situation has been
reported by Rogers (1977) for San Crist6bal reef at

La Parguera. Spur and groove areas also have a high

diversity due to their high relief.

-7. Observed stressed reefs are characterized by a.

general paucity of coral species and by areas overpopu-

lated by fast growing species such as the colonial ane-

mones or zoanthids. A clear example are the GuAnica

reefs. The shallow areas of these reefs are sometimes

100 per cent covered with either Zoanthus or Palythoa.

To a lesser extent Guayanilla reefs present similar con-

ditions.

8. Another characteristic we observed in stressed

reefs is the over abundance of the. sea urchin Diadema

antillarum. This condition is probably related to ample

supplies of algae which colonize the surfaces of dead
corals. Also, reduced predator populations (e.g. Balistes

vetula) related to a dying reef, may cause a Diadema

population explosion. An example of this are the inshore

reefs of Fajardo, such as Ahogado. In contrast to this,

was the outer non-stressed Cayo Largo which presented a

markedly reduced urchin population.

9. Colony death by bioturbation is common on heal-

thy reefs. Various stages of this process were observed

at Cayo Largo, Fajardo. Large sized parrot fish were

seen accidentally knocking off branches of individual

colonies of A. palmata. Consequently, "holes" were made

on the otherwise 100 per cent A. palmata covered zones.
These "holes" are later colonized by rapid growing spe-

cies such as M. complanata.

10. Gorgonian populations were observed to thrive

under conditions of heavy siltation and high turbidity.

Apparently healthy and dense gorgonian stands were ob-

served on the deeper zones of reefs off GuAnica, Guaya-

nilla, and Ponce.

11. Stony corals were also observed to thrive on

sloping surfaces in areas with high'rates of sedimentation.

Due to the nature of sloping surfaces, sediments appear

to flow by gravity to the reef base. This is apparent

on the fore reef slope of Cayo, Caribe, off Tallaboa Bay.

12. Inshore reefs of La Parguera, although apparently

underdeveloped, present areas of very high coral species

diversity as shown by the fore slope of Cayo Collado,

La Parguera. This, however, may be due to the develop-

ment of large numbers of small-sized colonies on availa-

ble, unoccupied substrates.

13. Areas of high water turbidity due to high amounts

of fine sediments in suspension permit coral growth on

shallow level surfaces. This is apparent in Humacao

where A. palmata growth is fairly abundant. Wave action

at the surface does not permit the sediments to settle

over the corals. Living coral cover, though, decreases

sharply with depth due to water turbidity.

14. During the end of the field data gathering period
(August 30, 1979) two tropical storms, one of them (David)

.of considerable magnitude buffeted the island causing

extensive reef damage especially in the shallower outer

reefs of the east and south coast. Some of these reefs

are under very heavy stress and their ability to recover

in the near future is seriously questioned by several

reef scientists.

15. We conclude that the acceptance of ecological

principles is mandatory if the environmental integrity

of the reef ecosystem is to be maintained in the face

of development. Taking into account the extensive damage

made by the two mentioned tropical storms, it seems rea-

sonable to state that an additional stress may seriously

impair the natural capacity of this ecosystem to recover.

RECOMMENDATIONS

1. Pollution research has always been weighted

towards public health aspects. It is not always suffi-

ciently rec,ognized by pollution scientists with a public

health background, that waters containing levels of

pollutants which do not threaten human health directly are
destructive to aquatic communities (Johannes, 1975).

Water pollution research in relation to benthic marine

communities should be promoted and sponsored by the De-

partment of Natural Resources in coordination with the

Environmental Quality Board and the University of Puerto

Rico. This research should aid in the modification, if

necessary, of our water pollution laws and regulations.

2. The enactment and effective enforcement of appro-

priate laws., and the public censure of polluters can

only be brought about when the public is made aware of

their value. The most important step in deterring both

the corporate and the individual polluter is thus educa-
tion (Johannes, 1975). We think it should be of high

priority to the Coastal Zone Management Division to pre-

pare televised educational programs for the public in

general in order to create consciousness of the impor-

tance of this and other important coastal ecosystems.

Several public information booklets have already been

prepared by this Division.

3. Reef areas which because of their quality and

extensiveness should be included among the Natural Reser-
ves are: (1) La Cordillera, Fajardo, (including Palomino
and Cayo Largo); (2) Sargent Reef, Maunabo; (3) all off-

shore reefs between Bahia de Jobos and Santa Isabel

(including Berberia and Caja de Muertos); (4) Cayo Rato-
nes, Ponce; and (5) Tourmaline and El Negro reef complex.

4. The underwater caves off Bajura, Isabela, should

be considered an important and potential touristic diving

attraction. Safety measures, such as marking its diverse

entrances and exits., should be taken by this Department.

5. Coastal Zone Management Division should prepare

booklets with the following recommendations to divers

and boaters:

(a) boat anchors These never should be dropped

on top of a reef. Damages by doing this

are the tearing up of coral when it hits,

the continued slashing of coral while

the boat is anchored and the tearing of

more coral when the anchor is hauled up.

A Danforth-type anchor should be used by

dropping it in a sandy patch and letting

the boat drift over the reef.

(b) sitting and standing on coral - This not

only causes breakage of branching corals

but abrades and injures tissue which can

then be infected by algae. Holding onto

coral can have the same consequences. Dead

coral and bottom rocks should be used for

this purpose.
(c) shell collection - Pry-bars should never

be used to dislodge and turn over coral

heads.

(d) spear fishizag - Coral injury from flying

spear shafts and working a fish in a cave

can be very severe.

6. Laws requiring the replanting of land cleared

of vegetation have been poorly enforced in the past. We

recommend that these actions be monitored by these Depart-

ment in-order to reduce reef siltation.

r(. In summary, we recommend the active involve-

ment of this Department in coming actions such as dred-

ging, upland deforestation, etc. which could result in

the serious impairment of the natural abilities of the

reef ecosystem to repair itself and recover after the

past tropicAl storms. The new erosion regulation being

prepared by the Coastal Zone Management Division should,

by all means, include the control of upland deforestation.

This active involvement is completely necessary if

the policy of this Department favors the preservation of

these most important and endangered Puerto Rican natural

resources.

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Table 1.

Peof Lo ca I i ty Zona t ion Diver- Eq it 1 ta- Corn] \fain
North east sity bility I cover :Component 0 B S F R V A T 1 0 N S
Cabo Fajardo
an SA i An
CCOT di n I tes
t . 18122. V
Lon p. 6 5'3 7. 2' Reef pa Ima t Colonies heavily silted. Gorgonlans common.
crest 5-10 and
M. complanata

palmata
zone L. Palmata Very sparse growth. Many dead and overturned colonies.

Slope :Head corals Colonies of the hydrocoral Stylaster common.
:and Actaricia

Heavily stressed reef probably due to discharge from Rio Fajardo.
PEIIARKS.

----------
Table 2.

T)ec- f i.o cn i i ty zonat ton I) i ver- rquita- Coral 'ma in 0 B S F R V A T 1 0 N S
sity bility I cover :Component
Isleta

t. 18-20.6' Reef M. compla- 90% of standing coral dead. P. asteroldes common.
Lon 1'. 65,37. 0' nata and M.
crest
:@-icicornis

Mix
and Massive Zones not well defined. A. palmata and Diploria labvrinthiformiS
palmata coral and competing for space (I ?)
palmata-
zone

Slope :Sparse
Reduced cover. A. agaricites and S. siderea common.
gorgonlans

Dead .coral fragments common. Many algal species present.
Reef
Thalassia P. porite abundant.
flat

I@eavily silted reef possibly due to the influence of Rio Fajardo and by construction activities on the island.
REIIARKS.

Reef Local i ty Zona t I on Diver- Equita- Coral Main
sity bility I cover :Component 0 B S F R V A T 1 0 N S
7anciido T-'njnrdo
(NO r r)
Coordinates
I-It. 18020.61 Reef
Lonp. 65*37.21 crest :A. almata Sparse growth
TA M. Small DiploTia strig0sa heads common.
cave7nosa

pa 3 ma t a
zone :A.pa3.mata :Sliphtly hipher cover.
:14. alcicornis common.

Mix zone
and
slope _7 Gorgonians :Aparicia sp., Colpophyllia natans common.

Consolida-
ted limes-
Reef :tone over-
flat grown by se:
verals-PCcies.
of algae.

REIIARKS: Heavily silted reef.

'72

Table

Pce. f Lo ca I i ty Zonat ion Diver- Fqiii ta- Coral: Main
sity bility I cover :Component 0 B S F R V A T 1 0 N S
Ahogi:do Fajardo

roorainatcs
I'a t 18, 19. 3'
Lon Reef
65037. 1' crest M, complErnta' A. palmata common.

palmata
zcn e A. palmata 40-55 % of standing @. palmata dead. Sea urchins very abundant
of top of these colonies.

Slope Gorgonlans
and heads Sparse growth

REIiARKS: Heavily silted reefs. Salinities close to M'have been measured on the surface of the reef flat during heavy rains. Colonies of Llcomplanata-
had expulsed zooxanthellas and died (as shown by the lack of stinging properties).

Table

P c P f Locil i ty 7 onn t i on Diver- T-qid ta- Coral ',,Ii 1 n
sity bility I cover Component 0 B S 1. R V A T I 0 N S
P=floini5itcPs -aJardo,
(solit I
These two zones are jifted. M . @Lornpl nata common but not
J'a t .jj_
Lon p nd predominant. D. clivosa, P . fragum D. qtokes
i_f, A. cervicornis
and P. asteroides. common. Very high cover although a significant
zone 4 A. _Eillmata number of colonies are partially broken.

Mix P. porites and A. cervicornis common. This zone descends up to
zone Gorgonians .7bout Gm wh.-re there are large colonies of A. paLinata at the fore
and M. edge. This is a zone of very high diversity. There Is a limited
annularis groove and spur system about 2 .5 m in relief.

Slope A. ija,2ric@tcs Area of high competition for space. Slope very steep. Flattened
and head :gro&,,h forms. Madracis sp. , P. @p5terc@ides_, E, P.Q@ites, El-milia
corals fastiqiata, Diplorla M. lamarcklana, 1. sinuosa, M. @@naulosa
common.

RElfARKS: The fore reef of Palorninitos Is heavily damaged by what appears to be boat anchorage. The abundance of Madracts decacti s is notable.
For a more detailed description of the benthic communities of this and other cast coast systems the writer Is reffered to Mckenzie and
Benton (1972).

Table 6.

Pce f I.oc n I i ty 7ona t i on Mve r- Eq u i t a - Coral Miin 0 B S F R V A T 1 0 N 5
sity bility I cover :Component
Largo Fajardo

Coordinates
l"i t .
I.on p. Reef Very abundant A. palmata with different shapes related to degree
65'34 .8' crest 0 0 100 palmata of exposure. Very occassional growth of M. complanata where
"holes" in the dense-A. palmata occur.

palmata Cover is slightly reduced. This area ends rather abruptly giving
zone 75 A. palmata way to the mix zone.

Mix Area of medium relief with very high number of coral species.
zone 1.58 0.72 25 Diversity lower than in the slope due to patchiness of coral
distribution. M. annularis, A. palmata, P_. porites, -M. complanata
and gorgonians common.
t

Slope 1.02 0. 63 35 A. cervicornis Zone of low relief dominated by L. cervicornis on upper slope and
and gorgo- by gorgonlans on lower slope. P. porites very common . Small
niaris head corals common.

Reef
flat P. porites Porites blotope very well developed covering extense areas.
Absence of Diadema and other urchins is notable.

REIIARKS: Reef terminates eastward with a bare sand "halo" about 8 meters wide after which a very healthy Thalassia bed starts. Large "halo., may be
due to an extensive ichtllyofauna. Numerous Queen Conch (strombu qiqas) were present in this bed. Paucity of sea urchins in the reef
flat may be due to very low coral mortality which are overgrown by algae and grazed by these echinoids . Within the reef crest sometimes is
apparent the effect of bloturbation (cor;il breakage by fish in this case). Apertures on the dense zone are then colonized by @4. complanata
and plate-like colonies of P. asteroides. Several stages of this process were observed. This is a relatively untouched reef of great
complexity which deserves further study.

7,
Table 7.

Peof 1,ocn I I ty Zona t I on N ver- rqui ta- Coral Main
Ramos sity bility I cover :Component 0 B S F R V A T 1 0 N S
(West) Fajardo
CoordInates
I'a t 18,18.91
T,o n 65,36.6

ReeF complanata absent. Small D. strigosa common. Large Mi.annularis
:colonies with dead areas on the 1pophyllia common.
crest palmata ir top parts. Co

Slope :Gorgonlans High gorgonian density. cervicornis and S. siderea common.

Reef
flat :algae Dead coral colonized mainly by several algal species. P. asteroides
common. Sparse A. palmata Increasing westward.

REIIARKS: West of the slope are dead A-cervicornis patches, a "halo" about 3m wide and a Thalassia Syringodlu bed. Siltation evident. Living
coral cover low.

Table

PC C, f Lo ca I i ty Znnn t i on M ve r- Fquita- Coral Ma in
sity bility I cover :Component 0 B S F R V A T 1 0 N S
R@,)Mos Fajardo
Ea S t)
Coordinates
1"I t
Lon p. Reef Dead colonies common. Halirreda, P. asteroldes and D. strigosa
crest :M.complanata common.

palmata Overturned and dead colonies common. Gorgonian common.
zone 10-30 A. palmata

Slope :Gorgonians :P. asteroides and head corals present.

REkIARKS: Very turbid water possibly by the outflux of Rio Fajardo. The crest zone merges with the reef flat which Is common to the west reef,also.

Table

P ce f Loc a l ity Zonation T)ivcr- Equita- Coral Main
Pircros Ceiba sity bility I cover :Component 0 B S F R V A T 1 0 N S

Coordinates
T-a t - 18*IS.31 Reef
T,onp. 6S.35.S, crest O.SS 0.51 60 M.complanat Sparse growth. No abrupt separation between this
zone and A. palmata zone.

52
pn1mata (live an@
zone 0.65 '0. 47 (lead butA.palmata @Iany colonies dead and standing. Low living cover (about
standinp- :20t) . D. striposa common. M. alcicornis common.
cover)

Slope Corgonians Tsolated M. annularis heads. Rarely any other coral
species.

Reef
flat Mialassia

REMARKS: Relatively u-tieveloped reef. Reef frinping the cast coast less developed. From the shore to sea there is a Thalassia
and yrinpo lum A.palmata
a* Ined followed by a 'hand, aboiit 20m wide of 95% dead coral. Farther east is a zone of 10% live
(20t-s-f`an_M_ngcover) increasing to '201 seaward and finally a band 13m wide of gorgonians and small head corals -. -ro-ral
reef development on neipliborinp island Cal-eza de Perro is virtually non-existent due to bombing practices by the
U.S. Navy.

Table 10.

Pcef Local i ty Zorn t ion Diver- P-qui t a - Coral Mlin
sity hility t cover :Component 0 B S F R V A T 1 0 N S
Santiago Humacao
---ro-ordinates
J'a t. 18109.5 Colonies of very diverse shapes and with no definite orientation.
Lon p palmata These alternate with gorgonlans. In shallower areas, M.
66,44.0' zone 60-70 A.palmata complanata. is common. Small head corals, common.

Slope gorgonians Large heads of M. annularis common.

PE4ARKS: Reef terminates with, n 'area of very sparse gorgonian growth.

Table 11.

Re e. f t.oci I ty zona t ion I) i ver- Equita- Coral Main
sity bility I cover :Component 0 B S F R V A T 1 0 N S
Guayama Arroyo
Coordinates Reef crest fragmented Into numerous patches where either of the
La t . 170 5 6. 5'
Lon P WO 1. 5 Reef pa I ma ta @wo principal components dominate according to the degree of
crest and M. com- exposure. P. porites an Important reef crest component A.
plana ta cervicornis and M. annularis common. 60% of A. palmata- dead.

palmata
zone 35 A. palmata Reduced cover and many colonies dead@.

Mix
zone Gorgonlans Gorgonlans very small in height. Dead A. cervicornis common.

Dead A.
cervicornis A.cervicornis: Very few gorgonlans present.
zone

PFfARKS: Southward of the dead A. cervIcornis zone the depth increases gradually to the edge of'the shelf. Sparse gorgonlans are present here.
This reef is considered as decaying and unbealthy.

Table 12.

Ree f Local I ty Zonation M ve r- Eqiii ta- I Coral @iain
cayos sity bility I cover :Component 0 B S F R V A T 1 0 N S
Punta Pozuelo
Caribo Jobos
--7-0 o r (F-nd-t-c s
1"I t 17* 5 S. 5' Reef Palythoa very common. Encrusting gorgonlan Erythro2odfum common.
66,12.5- crest M. complanata Calcareous algae common. Channels of 1-2 meters of relief.

palmata Sea fan (Gorgonia ) common. Large pieces of rubble encrusted
zone 39-49 with P. asteroldes, Palythoa, Erythropodium and Chondrilla.
ISzmant, A. palmata M. annularis-common.
:undated)

Buttress Zone of high relief due to large A. palmata colonies. Depth
zone 63 varies from 5-8 meters. Rich fish and. invertebrate area.
:(Szmant, A. palmata
undated)

Fore reef M. annularis, M. cavernosa, S. siderea, 1q. labyrinthiformis
slope Isophyllastrea rigida, Mycetophyllic and Mussa common.

Reef P.asteroldes Forms small channels and lagoons in some places between
flat :and overwash, mangrove stands. Depth up to Im, Usually a strong out flow
:A. palmata current occurs. P. porites, Favia Agari6ia and'sidera strea
common. Sparse Thalassia.

RE'fARKS:
Sea urchins (Dadema and Trypneustes common In the reef flat. High cover of Palythoa and Zoanthus inthis area. Size of A. palmata increases
with depth. Beyond the varied relief of the buttress zqne, the reef flattens out and slopes down to its base. Reef slope gradual down to about
13 m and then abrupt to 19m. Channels cut through reef slope In some areas.Aj@L@cla common in the wallsof these ch-3nnels.,

based on Szmant, undated.

Table 13.

Reef Locality 7onntion niver- Equita- Coral \,fain
sity bility I cover :Component 0 B S F R V'A T 1 0 N S
Las Mareas Guayama

Coordinntes
Lat. 17155.8'
Lorip. 66108.5'
Reef :Principal algal components are Dyctiopterls Hallmeda and Dvctlota.
flat Algae These cover almost 95% of the available substrate. Shoreward the
:cover is reduced.

Encru sting :Many over hanging ledges Indicating erosional processes. Where thls
Slope :sponges and :zone merges with the reef flat there are extensive pits made by the
algae *sea urchin Echinometra ILICunter.

"Fore Very abundant urchin pits. Promontories of high relief overgrown
reef Echinumetra to different extent. These probably represent collapsed ledges.
lucunter

REIIARKS - No living corals apart from Millepo faj@!_rro s aand Siderastrea radians were observed. Turbidity of the water was very high. No gorgonlans were
observed. Shoreward from the reef flat is a Thalassia bed followed by a Syringodiu bed . Between the latter bed and the shore is a
depression where large amount of broken off clig., fragments collect. (Cbservations based on Torres 1978 and personal visits to the area).

Table 14.

Peof Local i ty Zonatlon Di ve r. Equita- I Coral Ma in
sity bility I cover :Component 0 B S F R V A T 1 0 N S
Pajaros Salinas

Coordi na tes
T-It- 17055.5'
Lonp . 66015 .7' Reef Fragmented Into buttressess about 3m in relief and overwash.
crest M. compla-a ta D. clivosa and A. Palmata common. Ictyofauna abundant possibly
due to the highly sheltered area. Encrusting sponge Cliona common.

-palmata
zone A. palmata Well developed.

Mix zone gorgonians
and slope and head Similar to Morrillos, Salinas.
corals

@PIARKS: The north side of this cay is highly stressed with many dead and overturned colonies.

Table 15.

Peef 1.0 ca 3 i ty Zonn t ion DI ver- Eqiiita- Coral Main 0 B S F R V'A T 1 0 N S
sity bility I cover :Component
Morrillos Salinas

Coordinates
La t . 170 5 5. 8'
Lonp- 66115.3' Reef Very wide zone (from north to south). Buttressess about 2min
crest M.complanala: relief present. Coralline algae a very Important component
of this zone. Sand channels oriented N-S between th@e
buttressess. Dead coral present in the channels.

palmata A.palmat Slightly deeper sand channels In this zone. No dead coral, just
zone bare sand In the channels.

Mix zone Read corals r,,vernosa Meandrina meandrites, S. radians, 1. rigida, C..
and slope and gorgo- natans and D. strigosa common.
nians

REMUS:
Mangrove vegetation present.

Tab f6- i-6'.-

Peef Locality Zonation 111ver-: Equita- Coral: 'fain
sity bility I cover ;Component 0 B S E R V A T I 0 N S
Patones Salinas Ree

1,,,lt. 17"56.21
Pee f
LonP-66017.St crest 0.39 0.35 39 :M. compla- A. palmata common.
:nata

palmata
zone 0 . 3 6 0 . 2 2 40 :A. palmata jlalythoa oarse bare sand bctivcen crest
common. Zone of c
-an-E-IlLmata zone.

very abundant anddominant in upper mix zone.
Upper and 1.25 0.92 6 Gorgonians Massive corals dominant near reef base. Mycetophyllia,
lower mix 1.64 0.84 16 and and Oculina. common
zone massive
corals

REIIARKS: Seaward of the reef crest is an abrupt slope leading to a bare sand area with occassional isolated small head corals.
Farther south depth decreases asrapidly to a zone where medium relief mounds occur.
M. complanata. g-rows on top of these structures.

Tablb '17.

Reef Loca I i ty Zonn t I on nj vcr- Eqiii t a- Coril ','f 1 in
sity bility I cover -.Component 0 B S F R V A T 1 0 N S
AlfeAlque Sqnta Isabel

1-1 t - 17-55 .8'
Lonp, 66121.1' Reef M. complanata.- Blades oriented E-W.
crest 70

palmata A.palmata Many silted colonies.
zone 20-40 :and gorgorians@

Mix zone
and slope G)rgonians :A. palmata common. Head corals common.

Lagoon :Sparse Thalas-*
sia and dead
:Porites

RE@IARKS Zonation not well defined to this side (east) of the reef.. A . palmata zone not nearly as well developed as other nearby reefs. Spur and
channel system east of reef slope with orientation roughly north to south or parallel to the reef crest. Cover in this area Is mainly by
gorgonian In the spurs and bare sand on the bottom of the channels.

Table 18.

Peef Lo ca I i ty 7nnation D I ve r- Fquita- Co ra 3 Ma in
sity bility I cover :Component 0 B S r-. R V A T 1 0 N S
Cabuzosos Santa Isabel:

Coordinntes
1,1 t 1715 S. 4' Small Diplorla clivosa and D. strigosa forman Important component
Lonp. Reef of this ;irea. Calcareous ;@dgae also very common. Palytho common.
66'23. 1 crest 40 M. complanat About 30% of the available substrate is uncolonized

palmata
zone 45-75 :A.Palmata Cover increases seaward.

:A.cervicornis Both components very abundant. M. annularis. P. asteroldes,
Mix
and Dendrogyra cilyndricus and Col@ophylll natans common.
zone gorgonlans

Slope G org o n i a n s cavernosa, M . lamarcklana, D[chocoenta stokessli, 1. sinuosa
and asteroldes and several sponges common.
:A. agaricites

Reef Well developed Porites biotope and farther north a Thalassia
flat P. p2rites bed mixed with Porites.
and
:Thalassia

REIIARKS:

Table 19.

peef Locn1ity Zonation Diver-: Equita- Coral: 'Nain
sity bility I cover :Component 0 B S F R V A T 1 0 N S
Cayo Ponce

Coordinatcs
La t. 1715 7. 9'
Lon p W33 4'
Reef M. compknafa: Mounds about Im in height with coral on top. Coralline algae
crest :and M.alci- abundant.
:cornis

palmata
zone A.palmat D. clivosa an important component. Small M. annularis overwash
mounds.

Slope Gorgonians Large S. siderastrea heads.

RE@IARKS:_ Due to high water turbidity the southern limit of the gorgonian community was assessed up to a depth of Sm.

Table 20.

P e c f Local i ty Zonation DI ver- Equita- Coral M a in
sity bility I cover :Component 0 B S F R V A T 1 0 N S
Cardona Ponce

--76-57M n a t c s
1,;' t - ' 1715 7. S'
Lonp. Reef High cover. Low relief mounds with M. comrianata growing
66,38. V crest 75 -.M.complanatat on their tops at slightly deeper areas seaward of the crest.
Palytho common.

palmata High cover far high water turbidity present. Cover increases
zone 20-45 A,palmat seaward.

:Gorgonians
gorgonian 'and occassio-. Sparse growth.
zone :nal A. palmata

RE'iARKS: Gorgonian zone commences seaward of the fore reef slope and continues for an unknown distance. Small head corals occur here.

Table 21.

P e e f l.o ca i i ty zonation Diver- Equita- Co rn I main 0 B S F R V A T 1 0 N S
sity hility t COVQT !COMPOnent
Ratones Ponce
_-7-0o r d I na t e s
1,1 t - 1715 7 . V
Lonp- 66,40.8' Reef M.complanata: Ocassional A . palmata . Patythoa abundant. Small (1m) mounds with
crest LA. complanata- and D. strigosa on their tops.

Many dead colonies in northern zone. Living cover increases
palmata A. 2almata seaward. Sporadical gorgonians and annularis colonies
large buttressess with A. palmata at their tops about 3 meters
zone
In relief.

mix Lpalmata cover reduces gradually white gorgonian cover increases.
zone Gorgonlans Dead A. cer-vi--ornis common. M . annularls, M. cavernosa and
Siderastrea common.

Reef Thalassia Porites biotope not well developed.
flat and
podium

REIIARKS: Wide gorgonian zone (close to 100m.) Gorgonlan patches south of reef slope. Mangrove vegetation on reef flat.

Table 22.

Reef Local i ty Zonation Diver- Equi ta- Coral Main
sity bility I cover :Component 0 B S F R V A T 1 0 N 5
Caribc Ti I I aboa
T@ a Y
o r d i na t e s
Lat. 17?59.21 Dead :Area consistinp mainly of coral rubble with Palythoa
LonP-66044.01 coral Dead :and Nfillepora colonizing parts of the dead c@;Tal.
zone coral :Isolated 211mata colonies.

Former
mix Sporadical:90% of zone is bare sand.
zone gorgonians:

:rorgonian density increases in slope. M. cavernosa,
Slope Gorponian :M. annularis, 1). clivosa, and DendropyrT Ty-1-i-n-d-ricus-
:Eommon. ,ortaTity red ced.

Reef Vinlassia :Large bare sand areas. Mangrove islet present.
flat -577717-
:poriTes

PPfARKS: Shallow parts of this reef are dead possibly due to neaby dredging and ship traffic which stir up the sediments thus
SMOLhering corals. The slope, however, seems normal possibly because sediment runsdown the reef base without being
deposited on coral's surface. Isolated A. T),ilmata colonies in the dead zone may signify higher individual tolerance.
Fish life is sparse in the dead zone whiTe _qi_i-i_t_c-5"bundant on slope.

Table'"23'.

Reef Locality Zonation Diver-: Equita- Coral: 7,fain
sity bility I cover :Component 0 B S F R V A T 1 0 N S
Sa n La
rristobal P a rpue ra
-C-o -o-rTFn--,-i t e s Crest not solid but fragmented into buttresses of the
La t 17 0 5 6 . 7 Roof
two predominant species.
Lon 9. crest 'A. rinimata
67004 . 2 Tn d-rF-
COMPT'anata

:Northern most section with colonies having blades oriented
palmat :N-S. Farther south blades are thicker and with no
zone :A. palmata :definite orientation. Standing partially dead colonies,
: Ti n d - -u -nCe -r- 'common.
:laying dead'

:Other head corals, A. cervicornis and P. porites common.
Buttress
zone :%I. annularis

Slope :porgonian :M. annularis, Lf. cavernosa and Diploria labirynthiformis
:and A. common.
cervTcorn, is

Reef -.Thalassia mixed vith Zoanthus. Dyctiota common. Patches
flat :A.CCTViCM-rIi5:of -gorgonians and heaU -corals.
: -th IM7-e- t Tsa n d
:porponians : ,
RDIARKS: North of crest follows an area of dead coral fragments cemented by coralline alpae; Porites biotope; sparse
Thilassia with zoanthids; area o:F T)@,ctiota; patches of heads and porgonians. Reef Mi-textends to mangrove.
Apron are sand.

Table 24.

Pce f Lo ca I i ty Zonntion Nver- Fquita- Coral \Ia in
sity bility I cover :Component 0 B S F R V A T 1 0 N S
Enrique La parguera

Coordinates
]-a t - 17157.4-
Reef
Lon p 6P02.8'
crest .94 .68 19 M. com Plan ata: A. i3almata common. Dead, overturned A. palmata colonies, common.
Some coralline algae.

palmata
zone 18 .12 60 A. palmat Zone reduced due to abruptness of slope.

Mix zone
and A. cervicornis DIploria, M. annularis and A. agaricites abundant.
slope .51 .32 51 :and massive Occasional clumps of Oculina.
a I's Jln e
we
pe

Reef Thalassia Zoanthus common.
flat and mangrove:
isled

REIIARKS: Well developed platform In the front reef at about 8 meters depth. Dead coral common In this area. Width of platform Is reduced to the
west until It gradually disappears.

Table 25.

Peef I.o ca I I ty Zonation Di ver- Equita- Co ra I Main
sity bility I cover :Component 0 B S 1. R V A T 1 0 N S
Laurel La Parguera
--7.oordinites
1-1 t - 17-56.6-
I.onP' 67-03.8- Reef
crest M.complanat M. complan blades oriented E-W. A. palmata common.

palmata 100 :A. palmata Well developed zone with cover reduced south ward.
zone

Mix zone:
and :A. cervicornis Head corals common.
slope and gorgonons.

Micro
patch M.annularis
reefs and Agaricia High relief area.
south of sp.
slope

Reef
flat A. Dalmata with dead tips of branches.

REIiARYS' North of crest follows an area of A. palmata and P. pcxites; broader area of dead Porites.with sparse colonies of P. asteroides;
sparse Thalassia. Reef of consi-aerabTe -size and-devel-op-ment.

Table' 26.

Peof Local i ty Zona t i on I)ivcr- T-qiiita- Coral Ma in
sity bility I cover :Component 0 B S F R V A T 1 0 N S
Turrumote I La Parguera
Coordinates
7-1t- 17-56.3'
I.,np '67-01.2- A. palmata dominates over M. complanata. Branches of the
R&ef :A. palmata former, oriented N-S. Blades of M. complanata oriented E-W.
crest 1.15 0.63 58 :and M.corn-
:planata
palmata
zone 0 0 100 'A. palmata Solid cover by this species. Some growth can be delected on
the substrate beneath.

Mix
zone '1. 75 0.84 24.2 M. annularls Area of highest coral diversity even though cover is not
P. asteroides as high.
:and gorgcdans:

Buttress M. annularis
zone and Aga ricia, Zone of very high relief (over 2m).
sp.

Reef Sandy, some
flat Thalassia
m a ng rove
isledt

RE@IARKS: Buttress zone of very high relief. East of this reef Is a zone of even higher relief called locally "The Pinnacles" where there Is an
abundant Icthyofauna.

Table-

'.la in
peef Locality Zonation fliVCT-: Equita- Coral:
sity hi lity I cover tComponent 0 B S, F R V A T I 0 N S
La Gata La Parguera
Coordinates
La t . 1715 7'5 4
Lonp. 6702'6" Reef Many bear substrate mainly dead A. palmata M. complanata
crest 40% 'M.complanata oriented E-W. Stoic'hacthys common.

palmat 70-85 A. palmata
zone

Mix 55 M.-annularis Sparse palmata and gorgonlans common.
zone

Gorgonians M. cavernosa D. strigosa and M. annularls, Agaricia
Slope :and head common.
:corals

Reef :Thalassla and: Zoanthus and Palythoa, common.
flat :dead P. parl-
tes

REIIARKS: -South of the reef base Is a high turbidity zone with isolated partially dead gorgonian colonies. North of crest is a zone of dead coral
about Sm wide followed by sparseThalassia which gets denser close to the mangroves.

Table 28.

Peef I,Ocnl i ty Zonation Diver- EqMta- Co ra I 'Hain
n B S F R V A T 1 0 N S
sity hility I cover :Component
Margarita La Parguera

17055.2'
Lonp 67-6. 7' Reef
complanat Small sized A. palmata colonies, common. _Z@l @hoa @bundant.
crest 60

j?almata
zone 25-75 A. Palmata Wide (100m) zone

Buttress
zone M.annularis
:P. p2r@tes Gorgonians very abundant. Zone over 600m wide (N-S).
and gorgo-A3ns:

Reef Pcr-tially dead: Badly sorted sedimert .
flat P. Porite

Apron :Sandy; some
Thalassia

REIfARKS:
A. Palmata and mix zones very broad from north to south. Reef slopes gradually southward reaching a zone of dense gorgonian growth at about
130 m from the reef crest and dissipates at a depth of 16m and at a distance of over 600m from the reef flat. No mangrove present.

Table 29.

Pee f Loca I i-tx Zona t i on Diver- Eqiiita- Coral 1,1a i n
sity bility I COVOT Component 0 B S F R V A T 1 0 N S
Las La
Pelotas Parguera
Coordinn tes
T.'I t
Lonp. Reef Northern reef crest wanting In M. complanat P. porites common.
crest :A. palmat Southern crest with abundant Palythoa and some complanata .

palrr@ata
zone
60 -palmata Narrow zone

Mix Gorgonlans
zone and A. cervlcon@s:
slope :and M. com- A.cervicornis very abundant.
planata

Reef
flat :Thalassia P. porites and A. Palmata common.
:mixed with
-zoanthus

Apron Bare sand
with some
Thalassia

RE'iARKS: Beef disipates at about 13m depth. M. annularls patches occur southward of the 13m depth contour. Mangrove Islet present. South of
mangrove islet is an area of large sized (3. Sm diameter) L. palmat colonies. Eastward of this area, A. _palmata - cover Is reduced and
isolated heads of M. annularls of large size occur. Farther eastward is a z6ne of isolated A. cervicor-nis colonies. Subsediment dark in
this area (anoxic). Gorgonians abundant. North of this area the depth is reduced and there Is a dense A. palmata zone followed by a
complanata zone and later by a dense Thalassia meadow.

Table @0.

Pee f Lo ca I i ty Zona t ion Diver-: Fquita- Coral: 114a in
Cayo sity bility I cover :Component 0 B S F R V A T 1 0 N S
Ahogado La Parguera.
coordinates
La t.
Lonp. Reef A. -palmata
crest and M.
complanata

palmata
zone :A. palmata

Mix :M. annularis,:
zone M.alcicornis Isolated gorgonians just north of slope.
'@nd'gorgonbns:

Slope Small head
corals Many stressed colonies

Reef :Thalassia
flat and A. cervi- North of the crest Is a "halo" about Im wide after which the
cornis Thalassla meadow starts.
thickets

Zonation not well defined. No zoanthids in lagoon. Large aggregations of -sea urchin Diadema and Isolated gorgonlans where crest is absent.
REITARKS.
Crest dominated by L. palmata Instead of M. complanata possibly due to the protection of Enrique Reef. Very turbid mne south of the slope
with partially dead isolated head corals. Reef rises abruptly from sea floor.

100

Table 31.

Peef Locality Zonntion Diver-: Equita- Coral: @fain
sity t bility I COVeT COMPOntnt t 0 B S E P, V A T 1 0 N S
Media La
T.1inii - Parquera
70or(linatcs
17-56.4'
Lon p 67-02 . 8' Reef M. complanata
crest :and M.alci- Colonies oriented E-W
:corn

-palmat
zone 80 :L. palmat A. cervicornis common.

:A.cervicornts
Mix
zone :and gorgcrians

Slope :Massive
torals and Limited -spur and groove system.
gorgonians

Lagoon Sa-idy; some
Thalassia

RPIARKS:
North of the crest is a zone of P. _porite followed by a band where P. Isteroides dominates. Density Is reduced leeward where a zone
of dead A. Prolifera and gorgonlans occur. No mangrove vegetation. A. cervicornis, P. porites , and massive corals predominate in the
apron. Differences between Enrique and Media Luna aprons are due to different wave regimes (Morelock, 1977).

101

Table 32.

peer Lo ca I i ty Zona t i on Dive r- Equi ta- Coral @fa in
La La sity bility t cover :Component 0 B S F R V A T 1 0 N 5
Conserva Parguera,

Coordinates
La t
Reef
Lonp.
crest M. co mpla na ta: Exposed at low tide

palmata
A. palmat Very narrow zone
zone

Mix zone :A. prolifera Coverage very low. Many dead colonies of A-. palmata and A-.
and slope: :dead A. cer-vi-: cervicornis apparently due to siltation.
:cornis, M.
:annularis and
-00Fjonian

PPIARKS Reef not well developed possibly due to excessive protection to wave action. Mangrove islet present.

102

Table 33.

Ree F
Zonn t !on Diver- Equita- Coral Main
s@ty bility I cover :Component 0 B S F R V A T T 0 N S
Col lado Ln PargUera:
T__
Coor(l nates
compla- Palythol is a very important component on the inner reef
'Fat,1 and
Lonp. Ree F
1T,7,f-vti,o, flat.
Cres t :1 . 2 1 0.86
38 ca r IT a e o rum:

N I tra t n
Z -on 6-- *0. 1 0 S8 A. palmiti @one narrow and not well developcd.

@parse and sedimented Th -alassia bed between palmata and
Thnlassia @ix zone. Thalassia tTT!n iincFihort-bladed orl coarse
T ha In s s ja hiogenic sa-Eir-

tfix zone commences drasticallv south of Thalassia bed.
Coarse sediment. Sponges common. ColpojjLj1a natans
Mix Gorgonians common.
zolle :2.01 0.91 23 and massive:
corals

Thilassia,
Ree F I Ites ffalimeda zone divides crest and lagoon.
"or
flat in(! P.
asteroides

a hed hetween palmata and mix zone. Isolated pitches of A. cervicorn'
Pr-fARK,S: Anticlimactic Thalassi, is NI I inatn, Siderastrea
comp '
siderea and gorponinns oCCUr within this bed. ninderria common. about 1M_wi_(rj__F_e_t,,j_,n__@hT, bed and _PalTii_aCi_ione .
n_eF'_tv.'rminates pra(litally South Or MIX Zone. Miliprovc vegetation. Apron is bare sand.

103

Table 34.

Peef Locality Zonation I)ivcr-: Equita- Coral: Main
sity bility I cover :Component 0 B S F R V A T I 0 N S
Bajo Boquer6n
Eninedio
COORInatcs
t. Thalassla
180 0 1. 3'
Lon p. 670 1 26' iTed Thalassia Sparse to moderately dense.

mix.. Gorgonlans Lev&-d platforms with very densely gorgonlan growth alternating
zone and A. cer- with A. cervicornis patches. M. annularls, P. porites, M.
vicornis cavernosa and Colpophylia natans common.

Zone of high turbidity. Gorgontans of great size and abundance
Slope Gorgonlans as sole colonizers.

REMARKS: Gorgonian "forests" most espectacular feature of this and other west coast reefs. Turbidity fairly high.

lo4

Table 35.

Reef Local i ty Zonation Diver- Equita- Cora2 Ma in
sity bility I cover :Component 0 B S F R V'A T 1 0 N S
Guaniquilla Boquer6n

n .1 t c s
1,a t. 18102. 7'
LonP- 67'12.3' Reef M. complanata.
crest and
A. pa Imata

palmat
A. palmata Standing dead colonies common.
zone

Sparse
gorgontan Fine sediment and very sparse gorgonlan growth. Density of the
zone
Gorgonians latter Increases westward.

D3nsely populated by large sized gorgonians (close to 2m.).
Mix
zone and Large heads of M. annularis. C. natans alcicorni S.
slope 0.66 0.32 20 Gorgonians siderea, M. annularls, M. lamarchiana and P. porites common.
A. cervicornis common In mix zone.

Reef
flat Thalassia Seaward from shore Is a Thallasia bed followed by a dead coral
zone.

RPIARKS: Heavily silted fringing reef possibly due to discharge of nearby rivers. West of reef slope there is a zone of fine silt with occasional
gorgonlan patches.

105

Table 36.

Peof Locality Zonation Diver-: Eqiiita- Coral: Main
Punta sity bility I cover -.Component 0 B S E R V A T 1 0 N S
Ostiones Cabo Rojo

La t. 18005 .51
Loop.
6 7112. 0' Reef @Lcomplanata No A. palmata. P. asteroides common.
crest :and D. strkjosa:

Slope 'Gorgonians Abrupt slope. Head corals common. -Gorgonians large ard
abundant.

Reef Small leaved (5m) Thalassia. Considerable portion of reef
flat Thalassia flat bored by urchin Echinometra. Dyctlota- common. Palythoa
common.

PE4ARKS: Reef terminates abruptly. One meter"nalo" east of reef base after which starts a Thalas-sia bed. Slightly higher coral cover than Ratones, Joyuda.

1o6

Tabli@i*-37-.'-

Reef T.ocal I ty Zonation Diver- Eqiiita- Coral 'Ma in
Ratones Toyuda sity bility I cover :Component 0 B S F, R V A T 1 0 N S

oorainntcs
T-a t . 18,07. V
T,OnP. 67,11.3' Reef A
oalmata common. M-. alcicornis emergent or
crest M. abicornis overwash.

palmata 1.10 0.68 33 Crest and A. palmata zones intermixed. Lower palmata
zone (lower :(lower p@jl- (lower palA. palmata zone is the most diverse.
'palmata 'mata zone) :mata zone@

Slope Gorgonians Colpophylia natans M. cavernosa and M. annularis common..

Reef
flat Thalassia Sparse P. porite and overturned massive corals common. P.
asterotdes common close to crest.

RPIARKS: Small patch reefs continue west of the fore reef slope. Fine sediment mixed with Halimeda. Islet with extensive beach vegetation.

107

0- FA
\",Sea Level

NORTHEAST COAST CABO SAN JUAN
Fajordo P.R.

ACROPORA PALMATA

5-
MILLEPORA COMPLANATA

GORGONIAN

OVERTURNED A PALMATA

(L
ul
STYLASTER ROSACEA

HEAD CORAL

10-
r--\ -PORITES ASTEROIDES

AGARICIA AGARICITES

V. E.=3

Distance from Reef Flat In meters
I 1 10
0 15 30 45 ro

108

Figure 74 ISLETA MARINA tveto)
Fajardo P. Ft.

0
@Sea level

P
4ASTEROIDES
2 THALASSIA

CORAL RUBOL

3 MILLEPORA COMPLANATA

CL PORITES PORITES

4 an- MASSIVE CORAL

ACROPORA PALMATA

AGARICIA

MILLEPORA'ALCICORNIS
",-?-.G 0 R G 0 NIAN V.E. 3. 3
10 15 20 25 @O 35

Distance from Real Flat In motors

log

Figure 75

Cap Zoncudo (North)
Fajardo P.R.

3-
Consolidated dead coral
covered by algae-

4- -MASSIVE CORAL

ACROPORA PALMATA

MILLEPORA ALCICORNIS

GORGONIAN.

AGARICIA

V. E. 12.5

'0
21. 5 foo

110

Figure 76 CAYO AHOGA,DO
Faj a rd o, P.R.

N@',Saa Level

MILLEPORA COMPLANATA
E 3- V-ACROPO*RA PALMATA

GORGONIAN
CL

4-r'7\-/P%-MASsivr: CORAL

6 V.E 5

10 20 30 40 50

DIstance from Reef Flat In meters

Figure 77 'PALOMINITOS (South)
Fajardo) RR.

0- Sao Level

A. PALMATA
5

nn - M.COMPLANATA

- P. PORITES

@@"-NrASSIVE* CORAL

10-
5 - A. AGARICITES

- GORGONIAN

A, CERVICORNIS

15-

20-

251
10 2'0 3b 4'0 @O 6.0 7.0 60 0 160

DISTANCE FROM REEF FLAT IN METERS

CAYO LARGO
Faj a rdo R,R.
0 ",\Soo Lov*l

ACROPORA PALMATA

/)n(-MILLEPORA COMPLANATA
4 -

PORITES PORITES

MASSIVE CORAL
6

E GORGON IAN

CL ACROPORA CERVICORNIS
8

10 -

12 -

14- V. E. 5. 0

I 0 20 30 46 50 60 70 so 90 100 110

Distance from Reef Flat In meters

113

RAMOS (oeste)
Fojordo,P R,

0-
\,Sea Leval

2- ACROPORA PALMATA

-MASSIVE CORAL
E
3- -ACROPORA CERVICORNLS
CL
GORGONIAN

4- THALASSIA

6- V. E. 2.2

F
0 2 4 6 8 10 12 14 16 18 20 22 24 26

DIstonce from Reef Flat In meters

114

RAMOS,(este)
Fajardo, P. R.

00000 \IS*a Level
00

00000 - CORAL RUBBLE

4 - MILLEPORA COMPLANATA

E*,
M A S S I V E C 0 R A L

CL
ACROPORA PALMATA
6-

/--\-PORITES ASTEROIDES

8- GORGONIAN
A-OVERTURNED A PALMATA

V. E. 2. 5

10
0 10 20 30 40 .50

Distance from Reef Flat in meters
00

115

PINEROS
Ce I ba P.R.

'rs%Sea Level

-MILLEPORA COMPLANATA

3- -.ACROPORA PALMATA
E
GORGONIAN

-MASSIVE CORAL
4- (/7

V. E. 6

7
76
to 20 30 40 50 60

DIstance from Reef Flat- In meters

116

Coyo Corlbe'
Punta Pozuelo,P.R.
0- --Sao Level

MILLEPORA COMPLANATA

4- PALYTHOA

ACROPORA'PALMATA

MASSIVE CORAL:

GORGONIAN

V. E.5
7 10 15 20 25: 30 35 40 45
0 5

Distance from Reef Flat In meters

117

ARRECIFE LAS MAREAS
Guoyarno
.a- Sea Love I

-Dead reet flat colonized by algae---

@--Reet slope

-High relief botton

V. E. 4.4
101 110 20 2 5 3'0 3 5 410 4 5 5 0 515
0

118
CAYO RATONES
Sall nos .P.R.

Sea Level

10- nnn - MILLEPORA -COMPLANATA
15?-ACROPORA PALMATA
MASSIVE CORAL

GORGONIAN

20-

25-
V. E. 2.66

20 4'0 @O so

119

CAYO ALFENIQUE
Santa Isobel, P. R.

0-
Sao Love I

W-MILLEPORA COMPLANATA

ACROPORA PALMATA

7 GOR GO N IAN
fiF' ),K@MASSIVE CORAL

V. E.= 12

104,
0 io 160 150

DISTANCE FROM REEF FLAT IN METERS

120

CAYO CABUZASOS
Sania Isobeli P R.

'@"Ssa Level

Ann- MILLEPORA COMPLANATA
-MASSIVE CORAL
-ACROPORA PALMATA
_@,_GORGONIAN
AGARICIA AGARICITES
10-
-1"'-ACROPORA CERVICORNIS

V. E. 4

201
50 160 lio 2bO

DISTANCE FROM REEF FLAT IN METERS

Coyo Cardona
Ponce P. R,

0.
Sea Level

MILLEPORA COMPLANATA

PALYTHOA

CL
w ACROPORA PALMATA

4- 6 0 R G 0 N I A N

V. E. 6. 6
6

0 10 20 30 40 50 60 70

Distance from Reef Flat In meters

122

Cayo Ratones
Ponce, P R.
0- '*-Sao Love I

2 -

6- :MI-LLEPORA-COMPLANA-TA

PALYTHOA

ACROPORA PALMATA
8.
OVERTURNED ACROPORA PALMATA

10- GORGONIAN

MASSIVE CORAL

12 - ej-T4 ACROPORA CERVICORNIS

V.E. 10

14
0 40 80 120 160 200

MI.#.- 4- 0-4 el.,

123

CAYO CARIBE'
Tollaboaj RR. 'f",

0-
so Level

Dead Coral Zone
Isolated A.PALMATAand
M.COMPLANATA colonies.

iL
w

10- %

V. E. 4.6

10 25 @O 75 100 11,5

DISTANCE FROM REEF FLAT IN METERS

124

0-
Sea Level
SAN CRISTOBAL
Pargue'ra,P R.

M. COMPLANATA

A. PALMATA
5-

-M. ALCICORNIS

-A. CERVICORNIS

GORGONIAN

-HEAD CORAL

10-
M
(L

V. E.= 4

I
40 @O

125

0- SEA LEVEL-/'
ENRIQUE
Parquera, P. R.

5- Ann M. COMPLANATA

A. PAL MATA

MASSIVE CORAL

AGARICIA

w GORGONIAN

10-

V. E.= 2

15-

0 10 20 30

126

CAYO LAUREL
Porguero P. R.

Sea Level

5- n(in-MILLEPORA COMPLANATA
'4'1-?-ACROPORA PALMATA
-t-'*@-ACROPORA CERVICORNIS

MASSIVE CORAL

GORGONIAN

10-

CL
U.1

15-

V. E.z 4. 8

0 30 io do IL

DISTANCE FROM REEF FLAT IN METERS

127

CAYO TURRUMOTE I
Porguero P. R.

0-
Sea Level

5- non-MILLEPORA COMPLANATA

ACROPORA PALMATA

MILLEPORA ALCICORNIS

(ym- PORITES PORITES

a. MASSIVE CORAL
w AGARICIA AGARICITES
10-
GORGONIAN

V. E. 3.6

0 30 60 90

DISTANCE FROM REEF FLAT IN METERS

128

CAYO LA GATA
Parguera, P.

0-
Sea Level

01-MILLEPORA COMPLANATA

-ACROPORA PALMATA
E
-MASSIVE CORAL
cx AGARICIA SR

GORGONIAN

10-

15-

V. E 2.25

15 30 45 60

DISTANCE FROM REEF FLAT IN METERS-

129

CAYO MARGARITA
Cabo Rojo P R.

Sea Level

MILLEPORA COMPLANATA
'9? -.ACROPORA- PALMATA
/7N-HEAD CORAL
6' \):,@-GORGONIAN

8 V. E. 2.15
4
0 15 30 45 60 75 80 110 130

Distance from Reef Flat In meters

130

Ol
Sea Lfvol@@@@
CAYO AHOGADO
Porguero P.R.

A PALMATA

A@CICORNIS
6

7. -.HEAD CORAL

GORGONIAN

9. A. CERVICORNIS
CL
w

lo@

12.

13, V. E.=

14.

7L
18 1 4 6i 6 7 a 6 ib I'l it it 14 it 65 f7 16 6 io i I i2 i 3 24 5

131

CAYO MEDIA LUNA
Porguero i., P. R.

Sea Level

n ri

W)n-MILLEPORA COMPLANATA
5- 'YT -MILL EPORA ALCICORNIS
NT -ACROPORA PALMATA
'"@-ACROPORA CERVICORNIS
mm@ -MASSIVE CORAL
GORGONJAN
CL

10-

15-
V. E. 3.6 6

0 io 6.0 @O lio

DISTANCE FROM REEF FLAT IN METERS
7ACOMPLANATA

132

CAYO LA CONSERVA
Parguero,P.R.

0- \%-Sea Level

5.
E M. COMPLANATA

x
(L -A. PALM ATA
w
(,,@-MASSIVE CORAL
PROLIFERA.
C RVICORNIS
10- P.
AGARICIA S

GORGONIAN

V. E. 1.88

0 15 0 45

DISTANCE FROM REEF FLAT IN METERS

133 CAYO COLLADO
PARGUERA P. R.
0-
k"SEA LEVEL

PALYTHOA CARIBAERUM

m. COMPLANATA

-THALASSIA
(r--) HEAD OR MASSIVE CORAL
GORGONIAN

A. CERVICORNIS

SPONGE

ACROPORA PALMATA

10-

V. E. 4.7

0 30 45 60
fop'Nm qiad)f =I^# In @?,d)*mrv

134

ENIVIEDIO
Boqueron P.R.

0 *@Sea Level

JUL..

THALASSIA

6 GORGONIAN

ACROPORA CERVICORNIS

PORITES PORITES

MASSIVE CORAL

V. E. 10

20 40 60 80 100 120 140 ISO

Distance from East to West across the Roof

135

PUNTA GUANIOUILLA
Boqueron P. R.

0-
\Sea Level

THALASSIA

-HALIMEDA
7-ACROPORA PALMATA
x
CL
w ffiA -MILLEPORA COMPLANATA
\@-GORGONIAN
/P."\-MASSIVE CORAL
<-<--f-ACROPOR CERVICORNIS

V. E 7.5

0 15 30 45 60 75 @O 1@5 120 135 1@0

DISTANCE FROM SHORE IN METERS

136

Punta 001ones
Cabo Rojo, P.R.

0
77
IT k Sea Level

THALASSIA

CORAL'
,HEAD

2 PORITES ASTEROIDES
substrate bored
ATA
MILLEPORA COMPLAN by Echinometro

GORGONIAN

V. E.

6
0 30 45 60 75 90

Distance from shore In meters

137

Rcton6s
10 6 jayudo P.R.

0. -- - - - - - 04 ON a - A a woo

THALASSIA

2. PORITES,PORITES

HALIMEDA

OVERTUR'NED MASSIVE CORAL

P. ASTEROI DES
Cx
w
4- -MILLE PORA ALCICORNIS

ACROPORA PALMATA

GORGONIAN

-HEAD CORALS

V. E. r 10

6.
0 20. 40 60 80 100

Distance from shore in meters

138

'IN

VW7

41
7

Fig. 1. Diver making' an underwater profile. Observe
sea fans in gorgonian zone.

139

"-A -4-

Fig. 5. Dense M.
C- complanata on reef
R
e beginning
crest.
of reef flat in the
ound.
backgr

y7-
Fig. Reef crest. je

Fig. Abundant colonial
7
anemones (zoanthids) in
the reef flat. Several
hins (Diadema
black urc
antillarum. are present in
e background.

I A
ILI IL

14o

Fig. A. palmata zone. Note lOCr/5 cover and high ver-
fical relief.

Am@

Fig. 9. Large colony
of A. 1 ta helte-
rinig- g s h:emulon sp.)
and goat fishes (Mulloi-
Z- dichthys sp.)

14.1

.EEL-

h6
16

14W

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

...........

Fig. 11. Distinct morphological shape of the staghorn
coral palmata).

142

K,
IZ* @N

Fig. 12. M. annularis buttress. Note ldrge quantities
if planktiv6rous fish. Transect line can be
seen in the background.

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

14.3

Fig. 14. Brain coral in buttress
zone. Observe goby lying on the
coral.

Fig. 15. Colony of the pillar
coral Dendrogyra cylindricus.
Note large stand of the elkhorn
C
-al (A. cervicorn ) and a A-1
ol. is
gorgonian to the right of the
photograph.

40@7 I- I

144

Fig. 16. School of surgeon fish (Acanthurus sp.) in the
buttress-zone. Observe buttress the right
of the picture.

41%

soft*,-
IN

Fig. 17. Lobsters (Panul-'rus argus) wheltering within
crevice in the but:Eress zone.

145

..@Af

Mawr

Fig. 18. Large staghorn colony sheltering a hamlet
(Hypoplectrus sp.)

Fig. 19. Gorgonian or
soft coral in the fore
reef slope. Observe
the dominance of gorgo-
nians over scleracti-
nians (hard corals) in
this zone.

146

17-

77.

Fig. 20. Sponge (center) and massive coralg (fore and
background) in the fore reef slope.

Fig. 21. Turtle (probably Chelonia mydas) in the fore
reef slope.

FOTOGRAFIA AEREA 147.

.......... ...'Z: ...........
.... .....

. ... ......... .
............. .......... . .... :.

......................
..........

..........

. .... ....... .

(2) INTERPRETACION

AFLORAMIENTO

FRANJA SIN VEGETACION

PLANICIE ALGASEA

(3) COMO LO "VE" LA ECOSO@DA

c:) C3 c=m t=j c;:a C:@. C6 =3 lz> c=> co C=@ cz@ c- c:- cz) C=l
-_ __ -- - J. 'l -"

10 10 10

20 20 20

30 30 30

tf.'o c@b C=3 C=3 C3 C=3 CZ7 Cl c"a t=5 CZ> Z:3 ca cza

Fig. 22:
FRANJA DESNUDA DE VEGETACION ALREDEDOR DE UN AFLORAMIENTO
30 11 30 @3 0

ROCOSO SUBMARINO. FENOMENO CAUSADO POR EL- PASTOREO INTENSIVO
DE LOS PIECES RESIDENTES Y TURBULENCIA (after Lugo., 1978).

Ell
4Y
ig" 7 Pr
6nX Oj.
Aug'

Zi
"C ik
';4F"
;Ira
? -. Mnll
N

I Km

Fig. 23. Line of rock reefs (Trending east to west) offshore
San Juan.

149.

"Am
.. tn*

WU"Y",

2 Kms

Fig. 24. Heavily silted reefs Off FU%A1*a Iglesias and Punta San.
Agustin (center). Punta VAoia Talega can be seen at.
the left of the picture.

150

I K m

Fig. 25. Fringing reefs off Punta and Punta Picua.
Annular reef off Rio Mamey,@F is 6een to the center
right of the photograph.

151

K m

Fig. 26. Punta Percha with.fringing reel@,-"I')rotecting Luquillo Beach.

152

-re

t-A

2 Kms

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

153

I T
rv@

Fig. 28. Icacos,* westernmost islet of [-a Cordillera.

154

.71

N
A

Fig. 29. Palominos and Palominitos (sca-',I.-iern sandy islet) off Fajardo.

155

l'K m

Fig. 30. Cayo Largo, off Fajardo.
ASP'-
KORN

156

z@r

Z.
1J, JA

IL
jr

I K

Fig. 31. Islets and reefs off Fajai-do. From the bottom Isla de
Ramos, Cayo Ahogado and I-sleta Marina (Cayo Obispo and
Cayo Zancudo) are seen..
r,,:--WAIA&

157

I Km

Fig. 32. Reefs projecting east of Barrancas and Punta Mata
Redonda.

158

K m

Fig. 33. Isla Pigeros and Cabeza de Perro off Medio Mundo, Ceib'a.

159

I Km

Fig. 34. Cayo Santiago (upper right) and Cayo Batata (center) off
Humacao. Several submerged patc--L reefs, including Bajo
Parse, can be seen south of Cay,.-, Santiago.

16o

77s

-,r

I Km

Fig. 35a. Annular reef off Yabucoa Bay.

Fig- 35b. Non ve@rtical aerial photoGraph of the same reef.

161

1-1444

.'Z

Fig. 36. Fringine ree.f off Pai a. @,)_@_ueras.

162

I Km
4

Fig. 37. Arrecife Guayama south of Punta Figueras.

163

ire, 'y
L
I Km

Fig. 38. Arrecife Las Mareas off Guayama (lower right).

164

Z- Z.4

K ms

Fig. 39. Cayos de Barca, Jobos.

165

jr a .t

f:- xy

2 Kms

Fig. 40. Reefs off Salinas.

166

-wl .0

K m

Fig. 41. Reefs off.Santa Isabel.
- ar-

167

7-7

10- k

'-5A,

Kms

Fig. 42. Cayo Berberia off Santa Isabel- Highest reef
development in south coast.

168

T@F

47,

Z.-

2 Kms

Fig. 43. Isla Caja de Muertos off Ponce. Highest reef
development in eastern coast.

169

e n
YN'.
>

41- W

Z@A

777

man

Kms

Fig. 44. Cayo off Ponce. Arrecife Ratones can be seen at left.

170

-----------

t
ff v

e5o

.1 Km

Fig. 45. Stressed reefs off Tallaboa. Note oil refining
complex close to reef area.

171

- *W-.

Fig. 45a. Ship stationed between Cayo Rio and Cayo Palomas
in Tallaboa Bay.

172

@'A -k-', 11
KAI"

c NIV

JW,

RMS

Fig. 46. Reefs off Punta Verrac.o, Guayanilla.

173

Nr
j .I - . - @@- : t, ( #, - I@f. '. @ -V " -,-, ., .1k

.1 Km

Fig. 47. Fringing and patch reefs off GuAnica.

.174

A-
-7:
A.

IN.
N'

A,; t"
'UNIT
j - L

PN' I%w

2 Kms

Fig. 48. Reefs protecting Playa Cai-la Gorda, GuAnica.

175

IL
Jz-

K M'

Fig. 49. La Parguera.

176

b7. C
A;

-Nt.
li;MV@
A-M a

I K Mi

Fig. 50. Patch reefs off Isla Cueva.@,
ili'r J)A

177

-7@p`

N 1

I Km

Fig. 51. El Palo Atravesado reef.
ly

178

vv
jell,

K m

Fig. 52. Margarita reef.

179

v'-

.3k

IKm

Fig. 53. San Cristobal reef.

180

;,4

K m

Fig 54. Laurel reef .

181

I Km

Fig. 55. Media Luna reef.;-

182

_Km

Fig. 56. Enrique reef.

183

iv'f V,
T -A

Fig. 57. La Gata Caracoles reef.

184

7!1

ol.

IX4

I Km

Fig. 58. Erunedio reef

185

I Km

Fig. 59- Mario reef.

186

Fig. 60. Spur and groove system.
Observe high coral cover.

Fig. 61. Idem. Spur and groove
system. Note high coral cover,
specially Agaricia sp., lining
the sides of the grooves.

An@
I 4i

187

%'1" 49 +4

Fig. 62. Idem. Note whip antipa-
tharians (black coral). Spur and,
groove system. Note whip coral
(Stichopathes sp. in the fore-
ground. Accumilld sand can be
seen between spurs.

A
4

Fig. 63. Diver'working at
the shelf edge. Note insular
slope.
All

188

Fig. 64. Low relief sand channels north of the
shelf edge.

IL
U;

Fig. 65. Shelf edge. Note high coral cover. Insular shelf,
A

edge. Note high coral cover.

189

V
Ail
IA

2 Kms

Fig. 66. Fringing reefs of the west coast. Also note
Cayo Ratones west of Joyuda Lagoon (upper right).

190

2 Kms

Fig. 67. Fringing b&rrier,.-j@ecf,cff Boquer6n.

191

-,,.0

ell

2 Kms

Fig. 68. Offshore reefs of the west-coast. Square marks
off spur and groove system.

192

@Z. Ir

Fig. 69. Bajura caves. Fig-'70- Outer walls of the
Bajura caves. Observe_dense-...
Agaricia growth.

-Ak

Fig. 71. Large dead A. palmata colony-inside.
cave. Its origin remains to be studied'.

193.

4- "'t,

.tlv

71@

I Km

Fig. 72. Reef off Dorado.

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