The Coral Reefs of the Dominican Republic

Home , Acropora prolifera, Boca Chica, Isophyllia, Juan Dolio, Las Terrenas, Punta Cana

The coral reefs of the Dominican Republic

Francisco X. Geraldes

Centro de Investigaciones de Biologia Marina, Universidad Aut6noma de Santo Domingo and Fundaci6n Dominicana Pro - Investigaci6n y Conservaci6n de los Recursos Marinos, P.O. Box 748, Santo Domingo, Dominican Republic

ABSTRACT: The Dominican Republic has a land area of 48,484 km2, with a coastline of 1,389 km, of these 166 km or 11% are coral reefs. The continental shelf, averages 7.5 km wide and covers 8,130 km 2. There are two submerged offshore banks, two barrier reefs, as well as many fringing reefs. Dominicans in general recognize the importance of coral reefs as they provide safe ports, shelter and habitat for biodiversity, food and beaches. The first report of reefs in the Americas was from Hispaniola by C. Columbus in 1492, calling them "baxas" or "roqueiros". Other early naturalist worked with specimens from this island, and research continues until our days. The reef setting varies depending on the location and distance form the numerous river discharges. Dry areas and shallow platforms are favorable for reef growth at: Montecristi, Macao-Punta Cana, Parque Nacional del Este, Parque Nacional Jaragua, as well as the Silver Banks located some 170 km to the north of the island. The coral coverage varies from 40% to 9%, reflecting not only natural causes, but also anthropogenic impacts. There are reports of 64 coral species. The coastal marine habitats including the reefs of all the marine protected areas in the country have been studied, cataloged and mapped, the information produced have been important for their management and conservation. Some reefs are under threat by development of ports such is the case of Boca Chica, the most studied reef site in the island. Sedimentation along the coast has increased, and has become a threat to reef growth, occurring on 1/3 of the coastline, and now reaching reef sites such as Juan Dolio and Barahona. Coral bleaching has been found mainly on areas near urban development. The 1980's mass mortalities occurred as in the rest of the Caribbean. The reefs of the southern and eastern coasts of the Dominican Republic are usually exposed to hurricanes. The reef tracks near urban development are more impacted by habitat degradation due to physical damage and from nearby sources of pollution transported by currents. Non-adequate beach use in some tourists centers have caused reef degradation in the past, lessons learned have induced the tourism sector to become involved in reef conservation. The main problem reefs are facing is overfishing of several essential species such as Strombus sp., Panulirus sp., and fishes of the Serranidae, Lutjanidae, and Scaridae families. Several non-official institutions as well as the recent created Secretaria de Medio Ambiente y Recursos Naturales (Ministry of the Environment and Natural Resources) have programs for conservation of marine and coastal habitats, communities and species.

1. INTRODUCTION

The coral reefs of the Dominican Republic are an important resource, and it is acknowledged by the citizens that they provide safe ports, shelter, and food. Coral reefs have recently been associated with the tourism industry. While in the past, Dominicans took coastal and marine resources for granted, and considered them everlasting, there is

Latin American Coral Reefs, Edited by Jorge Cortrs 2003 Elsevier Science B.V. All rights reserved. 78 F.X. Geraldes today an awareness of their fragility, and of the fact that they need care and good management to ensure their health and function. At present, coastal and marine environments, as well as forest habitats, have national economic, political and aesthetic value. Nevertheless, overfishing, pollution and urban growth still pose very relevant and real threats to these ecosystems. In order to minimize these, the country has set aside four large coastal regions as marine protected areas: Parque Los Haitises, Parque Montecristi, Parque del Este and Parque Jaragua. The last three are located downstream of oceanic currents and receive minimal river influence. In all coral reefs, seagrasses, and mangroves constitute important habitat reserves. Other coastal features of the Dominican Republic include emerged reef terraces, shorelines of terrigenous origin, estuaries and sandy beaches. This chapter includes a description of the coral reefs of the Dominican Republic, as well as maps of the most important reef sites: two barrier reefs (Montecristi and Macao- Bavaro-Punta Cana), and several fringing, hard base, high energy marine-coastal communities. Finally, a short review of the environmental hazards that coral reefs face in the Dominican Republic is presented.

1.1. Natural history The island of Hispaniola, situated at 17040 ' and 19~ and 68020 ' and 70~ is the second largest in the Caribbean (78,000 km2), and located in the north-central boundary of the Caribbean Sea. The deep Windward Passage (4,000 m) separates it from Cuba, to the north-northwest, and the Jamaica passage separates it from Jamaica, to the west-southwest (3,000 m). It is separated from Puerto Rico, to the east, by the shallow Mona Passage (350-400 m). Oceanic currents and winds are primarily governed by the easterly trade winds. Hispaniola is politically divided into two countries: Haiti to the west, and the Dominican Republic to the east (Fig. 1). The Dominican Republic has a land area of 48,484 km 2, with a coastline of 1,389 km. Of these, 376.7 km (or 27%) are mangroves, and 166 km (or 11%) are coral reefs. Emerged reef terraces and cliffs, are the main coastal features found along the coast, especially on the southeastern region of the island. The continental shelf has a mean width of 7.5 km, and covers an area of 8,130 km 2. There are two submerged offshore banks: La Navidad and La Plata, 70 and 150 km 2 respectively, located north of Cabo Saman~ (Fig. 1).

1.2. Geography The island topography is diverse, with three large valleys and four mountain chains. These have a directional trend northwest to southeast, with broad valleys in between. Two outstanding features of the Dominican Republic are: that it contains the deepest zone in the Caribbean (the Valle de NeybaJLago Enriquillo, at 48 m below sea level), and the highest peak (Pico Duarte/La Pelona, 3,087 m above sea level) (De la Fuente 1976). The island also has a very complex tectonic and geological history, being seismically active.

1.3. Climate and oceanography The climate of Hispaniola is considered to be tropical marine dry, with an annual average temperature ranging from 18 to 32~ at the lower elevations. There are regional variations in climate and rainfall, influenced by the predominant northeasterly trade winds, The coral reefs of the Dominican Republic 79

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Fig. 1. Dominican Republic coral reef localities. as well as cold fronts from the northwest, and sporadic hurricanes and summer storms from the southwest. Rainfall tends to decrease from east to west. The annual average rainfall is 980 mm, with May and November having the highest precipitation, and December to April having the lowest. The oceanic circulation patterns in coastal waters are dominated by the Northem Equatorial Current, which flows westward and divides itself into northem and southem branches at Mona Passage (Metcalf et aL 1977). Counter currents, usually associated with tides, are common near to the coast. Tides are semi-diumal, with mean spring tidal ranges of 90 and 30 cm on the northern and southem coasts respectively.

1.4. Culture, population, and development The people of the Dominican Republic form an agricultural and farming society, which is presently moving towards light industry and tourism. Historians indicate that deforestation and human intervention in the natural setting has occurred since the 15 th century. Severe environmental changes occurred during the early to middle 20 th century, when the sugarcane industry boomed and large areas of forest were cut down, increasing soil erosion and sedimentation, and hence ultimately affecting the nearby marine ecosystems. 80 F.X. Geraldes

Another major impact to the environment has been the recent population rise, from 2.5 million in the 1960's to the present 8.5 million today. This rise has increased the degradation of watersheds, river systems, and marine and coastal habitats.

1.5. Coral reef research The study of reefs in Hispaniola began in 1492, when Christopher Columbus described these submerged structures as "roqueiros", "baxas" and "restringas" (confinements) in his navigation log (Col6n 1492). Later, in 1525, Gonzalo Femfindez de Oviedo (in Chardon 1949) mentions these strange formations on his Sumario de la Natural Historia de las Indias. Several early naturalists, such as Carlos Plumier (1695 in Chardon 1949) and George L. Leclerc (1770 in Chardon 1949) made collections for the Muse6 de Historie Naturelle de Paris. In 1776, Luis Nicolson published his Essai sur l'Histoire Naturelle de St. Domingue (Chardon 1949), where he mentions madreporaria, corals, crustaceans, and other invertebrates. Also worth mentioning is the work of Moreau de Saint-M6ry during 1796 (Chardon 1949) and his Description Topographique et Politique de la Partie Espagnole de L'Ile de Saint Domingue. Later in the 19th century, more works followed such as those of William Gabb (1868, in Chardon 1949), who published "On the Topography and Geology of San Domingo". In the early 1900's, other scientists such as Vaughan (Vaughan 1900; Vaughan et al. 1921), Wells (1956), and William Hassler, who published "From Sea Base to Mountain Top at Santo Domingo" (1933, in Chardon 1949), collected and worked either on living and fossil reefs of the Dominican Republic. More recent studies, including those of Sir W. Halcrow and Partners's reports on the environmental impacts at Boca Chica (1976); Geister's (1980), and Schubert and Cowart's (1980) work on the paleontology of reef terraces on the south coast of the Do- minican Republic; the studies of Galzin and Renaud-Mortand (1983) on pollution effects on coral reefs; and the work on the reef conditions after hurricanes (Barnwell 1983). Dominican contributors to reef studies include Bonnelly de Calventi (1974), who published a taxonomic coral list. Gonzfilez-Nufiez (1974) located reefs sites and lists species collected in "Operaci6n Madre Perla". In 1973, F. X. Geraldes began his research on coral reefs, publishing in the local newsletters of the Museo Nacional de Historia Natu- ral, Centro de Investigaciones de Biologia Marina, and the Herbario de la Universidad de Santo Domingo. These early works included the first Dominican scientific reef study with descriptions of reef types, species, and locations, the results of which were later summarized and published (Geraldes 1976, 1978). Rathe (1981) produced the first sys- tematic study of sponges in Dominican coral reefs. Geraldes (1982) also studied the effects of hurricanes on Dominican reefs. More recent studies include reef characteriza- tions, ecological assessments, and species lists (Geraldes 1994a, 1996a, b, c; Vega 1994; Vega et al. 1994, 1997; Geraldes and Vega 1995a, b; Geraldes et al. 1997), as well as reef conservation efforts made by creating volunteer networks for reef monitoring (Cintr6n et al. 1994; Geraldes 1994b).

2. DESCRIPTION OF CORAL REEF AREAS

Most coral reefs of the Dominican Republic are fringing reefs. There are also two barrier reefs, numerous patch reefs, and four large offshore banks; their distribution is associated with the coastal profile and depth of the ocean platform. In places like Macao The coral reefs of the Dominican Republic 81

-Puma Cana and Momecristi, broad platform barrier reefs are found. Not so in Palenque, Saman~ Bay, and the Bahia Escocesa, where reefs are unable to establish due to the high turbidity caused by numerous river discharges. There are also unnatural conditions that are affecting the reefs such as increasing coastal development, pollution, untreated waste, water discharges, and beach erosion. This chapter describe the reefs found in the coasts of the Dominican Republic, starting with the offshore bank reefs (Silver Banks). The coastal reefs are described beginning at the northwestern shores bordering Haiti at Rio Masacre (Montecristi), and moving clockwise around the island to the southwestern border with Haiti at Rio Pedernales.

2.1. Offshore bank reefs: The Silver Banks, National Sanctuary of Whales

The Silver Banks (Fig. 1), at 20~ 69~ l'W, and 140 km north of Puerto Plata in the Dominican Republic, have an area of 3,740 km 2. In the northern portion of this bank the barrier is composed of a series of patch reefs bound together near the surface. This area is shallower and shaped like a triangle. The reef extends for 30 km to the southeast; it is exposed at low tide: on its ocean side to a great depth, over a distance of less than 100 m. The coral patches are pillars of cemented coral skeletons ascending from the rubble and sandy base 15 to 25 m up the surface. The living coral species found here follow the zonation pattems described by Goreau (1959). A. palmata is found occupying the top portion of the reef down to the 6 m depth contour. Below that zone, most corals are typical of the lower palmata and buttress zones. The substrate of the Silver Banks is mainly sand and coral gravel. At the southern portion of the breaker zone, the mean depth is 40 m, and corals grow only where suitable substrate is present. The turbidity of the water tends to increase to the south and away from the reef crest. A reduction of coralline columns or pillars is obvious to the south of the reef crest. The mean coral cover in the Silver Bank is 40%. There is a low density of sponges (2%). Turf algae covers 51% of the sampled substrate. The rugosity of this reef is relatively high (1.3%) due to the magnitude of the coral columns or pillars, some of which reach the surface of the water.

2.2. Coastal reefs

2.2.1. Montecristi barrier reef (Fig. 2.). The Montecristi region in the northern coast of the Dominican Republic has the largest reef formation of the country with a length of 64.2 km, which grows on the nearshore areas of the Montecristi Shoals (1,181 km2). The coastline consists of a low mountain range of sedimentary (Miocene) origin. At the northwestern end is the landmark of E1 Morro (273 m) and the city of Montecristi, and 2 km to the west is the Yaque del Norte River estuary with its large mangrove forest. The climate and land ecology of the area is dry to very dry tropical forest, conditioned mainly by the lack of river runoff and the steady easterly winds (15 to 30 knots). The reef begins at E1 Morro and extends westward reaching Punta Rucia. Along the coast and protected by the barrier and lagoon, extensive nearshore seagrass beds and frondose growths of red mangroves (Rhizophora mangle) thrive. The reef system of Montecristi can be considered as a barrier reef in active expansion. High relief features and large living coral colonies with sizes exceeding 10 m in diameter are common. The deep reef 82 F.X. Geraldes

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Reef lagoon: The reef lagoon can be between 20 m and 2 km wide, and up to 20 m deep. In association with it there are sandy beaches, mangrove forests and terrigenous cliffs. In hard base areas (1 to 5 m deep), small coral patches (5 to 800 m 2) are found with Gorgonia spp. and Plexaura spp. as the dominant soft corals, associated with the M. annularis complex, Diploria spp., Manicina spp., A. cervicornis, Porites spp. and Millepora spp. The main feature in this lagoon is the dense to sparse seagrass beds of Thalassia and Syringodium on a sandy or sand-mud base. Other coral species found in the reef lagoons are: P. divaricata, M. areolata, S. siderea, S. radians and A. cervicornis.

Reef flat and back reef: There is an extensive and diverse back reef followed by a reef fiat. In areas closer to tidal channels a more vigorous growth of corals occurs: Porites porites, Porites divaricata, the M. annularis complex, A. cervicornis, A. palmata and Millepora complanata are present with octocorals and sponges as well. Near to the reef crest a harder, more consolidated base is found, where calcareous algae Amphiroa spp. and Jania spp. are dominant. Thalassia and Syringodium grow in patches.

Reef crest: Skeletal remains of poorly lithified Acroporids form the reef crest. Millepora is dominant although a few young A. palmata are also found. This change in dominance patterns could be attributed to the Acroporid disease, with Millepora substituting the previously dominant A. palmata and Porites spp. structures. In most of the reef area, the crest is narrow and crossed by tidal channels. Here, the waves from oceanic swells barely reach the reef. This may be due to the effect of the precedent shoal which reduces their force. The energy here is mainly represented by wind-generated waves with a short lifespan producing choppy seas, which conditions a low energy environment. On its ocean side the crest abruptly drops to 6 - 10 m in less than 30 m, allowing a clear view of its basal structure, which is composed of large skeletons of A. palmata, A. cervicornis, Porites spp. and Montastraea spp. Some of these skeletons have broken loose and lie at the base of the crest, serving as suitable substrates for future colonization, as well as refuge for other reef inhabitants.

Outer reefs: In exposed areas, there is evidence of a lower Palmata zone consisting mostly of large, dead colonies of A. palmata. Seaward is a low relief spurs and grooves formation, with large colonies of the dominant Montastraea complex are also found. There are variations to this zonation pattern: when tidal channels divide the reef crest, there is usually a portion of the breaker zone, facing away from the predominant forces (wind and waves); this creates a very calm and protected portion on the reef. In this setting, the reef crest can sometimes abruptly drop into a sandy channel with seagrass, often down to 30 m deep. As this portion receives some of the ocean's energy, the coral growth can be found forming patches 10 to 5,000 m 2 in size. This set-up then continues either towards the shore where it connects to the backreef, or seaward, where it is followed by spurs and grooves or hard grounds. In the majority of the cases for Montecristi, the reef crest slopes towards a hard base (10 m deep), where octocorals are 84 F.X. Geraldes

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0 The coral reefs of the Dominican Republic 85 dominant. The hard base ends in a wall, dropping to the 25 m contour where large coral colonies are found. The M. annularis complex, M. cavernosa, Diploria spp. and Agaricia spp. amongst others cover most of the bed, with a total coral cover exceeding 60%. In these places, the colonies form gigantic structures, some occupying up to 100 m E, formed by coral heads of multiple colonies forming a single structure. Among these, broad caves and deep crevices (14 m) are common. In the deepest portion (30 m), coral pinnacles are found. Live corals and other hermatypic organisms use the large boulders and debris that have precipitated down from shallower regions as substrate to grow vertically in a pillar-like fashion. When several of these pinnacles grow close together, they sometimes join to create a larger base. In most cases the dominant species is Montastraea, and it covers all previous growth. In other cases some of these are dominated by octocorals and coralline algae such as Halimeda goreaui, Halimeda discoidea and Dyctiota spp. The encrusting octocoral, Erythropodium caribaeorum, and the red calcareous algae Porolithium caribeaeum are also common features of the pillars at the Montecristi reefs. Another type of reef formation found here are deep reefs, which form buttress systems. These are characterized by large to very large dominating colonies of the M. annularis complex, as well as M. cavernosa, Siderastrea spp., D. strigosa, D. labyrinthiformis, D. stokesiL Madracis spp., M. meandrites, A. agaricites, A. cervicornis and the sponge Cliona langae. These buttress systems are usually found in association with the tidal channels located throughout the extension of the reef.

Offshore keys Punta Rucia keys. In the eastern end of the Montecristi Barrier Reef at Punta Rucia, there are 16 offshore coral keys. These are small (800 to 5,000 m2), submerged, mount- like features that rise from a sandy platform 45 to 100 m deep. They have an arch-like shape, with the convex side facing the incoming winds. There is a significant difference in coral growth pattern between the exposed and protected sides of these keys. On the protected side of the keys, at 15 m depth, there is sparse marine vegetation, with Syringodium and fleshy algae dominating; as this reaches the flat, a back reef may form. Here P. caribaeorum, and the corals P. astreoides, D. strigosa and M. annularis are found with small colonies. In these places the long-spined black sea urchin Diadema antillarum is common, hiding amongst the crevices of the healthy reef found here. Moving towards the reef crest, on its windward side and at 2 m deep, a breaker zone is evident; species diversity increases, as do organisms adapted to stronger waves and currents. On some of the keys the breaker zone occupies the entire key. The dominant species found here are: A. palmata, Millepora spp. the, M. annularis complex and D. strigosa. In some of the other keys in these high-energy areas, there are large areas of hard substrate without any coral growth; instead, large algae mats of Lobophora sp. and other algae cover most of the area. The frontal reefs usually end at approximately 12 m depth, sometimes followed by a sand plateau or by a forereef slope to deeper waters. Hermatypic corals dominate this reef slope. Numerous healthy colonies of the M. annularis complex, A. palmata, M. cavernosa, P. porites, A. agaricites, together with the sponges Cliona langae and Aplysina fistularis are found. Due to currents induced by heavy swells at the base of the exposed side (forereef slope), a detrital area is found. 86 F.X. Geraldes

Here octocorals such as Plexaura homomalla, P. americana and P. flexuosa and the coral M. alcicornis become dominant species. Siete Hermanos keys. These keys are not part of the Montecristi barrier reef system. They are located 25 km west of El Morro, at the western and distal end of the Montecristi Shoals, on a ocean floor rise (5-30 m deep) and at the edge of the Manzanillo submarine canyon (>800 m deep). The keys occupy an area of 10 km 2, and vary in size between 3,000 to 40,000 m z. All are composed of sand and unconsolidated coral debris with arid vegetation, where nesting populations of Tula leucogaster (buN) are commonly found. Water turbidity and salinity show influences of the Yaque del Norte river estuarine system about 20 km upstream. The sea bed is sand-mud. Despite these conditions, the shallowness of these seven shoals offer enough light penetration for fringing reefs to form on the windward sides, as well as patches on the leeward sides of the keys. In shallow areas between the keys, there are numerous patch reefs. In some cases these are vaguely connected and form large structures, especially between Cayo Rata and Cayo Muerto. Coral reef formation is least at Cayo Tourur6, this being the most easterly key and thus closer to the estuary. Nevertheless, the river plumes can reach as far away as Cayo Arenas, located furthest to the west. It is considered that water quality, mainly its turbidity and salinity, have influenced this region to conform a unique reef formation. The coral structure on these keys is of a l~inging reef type down to 20 m, consisting of a sandy beach followed by a narrow reef flat 10 to 50 m wide. The species composition on the back reefs is similar to Punta Rucia back reefs. The breaker zone, however, presents differences between the keys. Most of them rise from a sand-mud base at 10 - 12 m. In keys facing surf and currents, the hard base substrate is bare and covered by cementing and boring sponges, and octocorals. In those keys where the currents and surf are not as strong, vertically growing species are found together with large to gigantic coral colonies. These grow sparingly, and are surrounded by finger-like corals such as Porites spp. Sometimes, gigantic round coral forms can be found which comprise most of an area, with very few if any other accompanying benthic species.

Biodiversity of the Montecristi barrier reef Recent studies done in Montecristi (Geraldes 1996a, b, c; Geraldes et al. 1997) have produced updated information about this area, a community and substrate map (1:40,000) of the coastal region, and the first biodiversity list for the region's marine and coastal habitats. The biodiversity sampled here includes 22 Classes, 285 Families, 525 Genera, and 742 species. The highest species richness is found in the hard base communities, which is related to substrate rugosity and complexity. Of these hard base community types, coral patches, high relief spur and grooves, and reef keys represent the refuges for biodiversity.

2.2.2. Luper6n reef, Luper6n Bay, Puerto Plata. Eastward of Montecristi, the coast is characterized by a diversity of habitats: a reef terrace extends from Punta Rucia to Luper6n, and the coast is then mainly of terrigenous origin until it reaches Puerto Plata. Luper6n Reef is a fringing reef located 250 m offshore. It grows over a narrow drowned reef terrace, which borders a shallow submarine canyon at the entrance of a bay (Bahia de Luper6n), where mangroves and a small town are found. The reef grows on a hard base, with low relief and high gorgonian cover up to 10 m deep. The reef slope ends at The coral reefs of the Dominican Republic 87

40 m depth near a submarine canyon. Coral cover is 12%, with algae dominating (67%). The growth is mainly of hard and smooth substrates, with very little reefrugosity (1.04%).

2.2.3. Playa Dorada reef, Puerto Plata (Fig. 4.) Patchy coral growth with low cover by live corals, dominated by encrusting turf and fleshy algae, are characteristics of a sedimentary coastal region of calcareous origin, which is washed by numerous torrential streams that form sedimentary plumes; rip currents are common here. Offshore (18 m deep) a hard basal community grows on top of a sandstone substrate. In areas where coastal features provide shelters, like Playa Dorada, the coast was originally fringed by healthy mangroves and marshes (now replaced by golf courses and hotels), which acted as sediment and nutrient barriers, and allowed the formation of patch reefs. These often formed small mounts and pillars, with rose vertically from a 15 m base to the surface. A small fringing reef of A. palmata and Porites sp. developed. These structures protected the coast and created a large stretch of beach, which is now intensely utilized by the tourism industry. At present, this reef is severely degraded due to environmental misuse by the resorts (over 5,000 rooms) and the golf course built here: all these infrastructures have exceeded their sanitary infrastructures capacity, thus streams and ground water pollution, together with nutrient runoff, have seriously affected the nearby reefs and coastal regions. There is little hope of recovery unless the activities that maintain this present level of pollutants are controlled. Studies shows that there is an 80% coral mortality, and that 92% of the basal and reef substrate is covered by algae, especially Gracilaria spp., Dyctiota spp., Turbinaria spp. and Codium spp. Eastward of Playa Dorada, the coastal features of the coast change to reef terraces followed by a narrow flinging reef that extends eastward for 3 km. The distance from shore to the reef crest reef varies from 10 to 200 m away, where the breakers begin; A. palmata skeleta are found here covered with algae. Millepora spp. now dominates this breaker system. In deeper water (15 - 25 m), irregular sandstone structures with many crevices and pillars, covered with scattered coral growth, rise up to 5 m high from the sandy beds.

2.2.4. Sosfia reef, Puerto Plata. SosOa reef is located in a small bay open at the northeast. The base is composed of beach rock. The calm waters and white sandy beaches off the coast make this a preferred tourist destination. High reef terraces encompassing a unique view surround the beach. Underwater, a patch reef surrounded by sand occupies approximately 30% of the center of the small bay. A prairie of octocorals with sparse growth of A. palmata, M. alcicornis, D. clivosa, and D. cylindrus begins at the 4-m contour. Deeper, at the 10 m contour, the diversity and cover by coral species increases, covering 28% of the substrate. Intense visitation by tourists has caused severe impacts on this site. Continuous visitation creates large sediment plumes. Algae cover is 43%, possibly due to some undetermined nutrient input. In terms of species diversity, the absence of large predatory fish species is noticeable. The sponges occupy 6.4%, and are mainly of the encrusting and burrowing types.

2.2.5. Reefs of the northern shore of the Samanfi Peninsula Reef Las Ballenas keys. This reef surrounds a carbonate outcrop in the coast of Las Terrenas, on the northern shore of the Samanfi peninsula. The base is a hard pavement 88 F.X. Geraldes

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M The coral reefs of the Dominican Republic 89 eroded relict reef, where sparse medium-sized coral colonies, mainly of Diploria spp. and Meandrina, establish themselves in crevices and grow. There are also octocorals, and a thick carpet of fleshy and turf alga covers this basic substrate. The species richness is not high, and comprises 11 species of algae, 9 sponges, 9 octocorals and 2 hard corals. E! Portillo reef. This reef is located on the northern shore of the Saman~i peninsula. The coastline is composed of high and steep carbonate mountain slopes and terraces. The climate is very humid. Since the 1940s, most of the coastal lands have been turned into coconut farms, deforesting the area in the process. The reefs here are fifnging, growing very close to shore in sparse patches with a tendency to close at the breaker and to form lagoonal environs. On these lagoons, and very close to the shore in shallow waters, the patches have coral remains of A. palmata, the M. annularis complex, Millepora spp., and Diploria spp. Most are fully covered by fleshy and turf algae; very few, if any, live corals are found on them (Geraldes and Vega 1995b). Further out to sea, approximately 9 km offshore, there are numerous shoals (15 m deep), that rise from the surrounding oceanic waters. These are carbonate terraces, eroded either by bioerosional processes or by exposure to weathering during past geological times. The species of corals found in these shoals are few and low in coverage (11.7%). More dominant is turf algae (44%), fleshy algae (20%), and encrusting sponges (18%) (Geraldes and Vega 1995b). The lack of large predators and the scarcity of large herbivores and sea urchins are affecting coral recruitment on these reefs, giving the opportunity for more active erosional processes to Occur. Puerto Escondido. This site is the northermost shore in the east of the Saman~i peninsula. It is located very close to a terrigenous rocky shore with falling boulders, and to the edge of the narrow continental shelf. Corals here grow without any apparent anthropogenic influences. Runoff and springs in the past, which are now reduced, have created small, narrow, protected coves where corals flourish, and where A. palmata grows vigorously. In deeper waters, on a terrigenous base, a young fringing reef can be found with 25 coral species, 15 species of sponges, 11 species of octocorals and 18 species of algae (Geraldes and Vega 1995b). Cabo Cabr6n reef, Las Galeras. The reef near Cabo Cabr6n offers a spectacular wall dive. It is located at the tip of the Saman~ peninsula. The coastal region is formed by Tertiary rocks (marble), and is steep and high (400 m). The water depth right off the coast surpasses 150 m. In some areas where landslides have occurred, narrow terraces can be found formed by large boulders. In those places coral grows from the surface to 50 m deep in a 30 ~ incline. The coral forms are of the encrusting and massive type, and coral cover is approximately 40%. Tube sponges follow with a 28% coverage (Geraldes and Vega 1995b). Large fish are frequently found here. During winter, humpback whales are often encountered, here and in the Portillo and Terrenas reef sites.

2.2.6. Miehes reef. This region on the eastern coast of the Dominican Republic is very humid. Numerous rivers and streams loaded with sediments join the ocean near Miches. The Yuna river system discharges to the west in Saman~ bay, and together with the Los Haitises and the Sabana de la Mar watersheds, they form the largest estuarine system of the Caribbean islands. The waters in the entire region are generally murky due to the high loads of sediments, limiting coral growth. Near the town of Miches, at Punta Hicacos, a 90 F.X. Geraldes

Punta 68115,

Swamps Atlantic Ocean

..... o .... ,. '",,,, -pry F

SwarrMp~ -~ , Reef Terraces Bavaro

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Fig. 5. Macao- Bavaro- Punta Cana. small fringing reef has developed. To the west there are small patch reefs. Extending towards the center of the bay to the north, there are offshore shoals, which are dangerous for navigation and where important 16 th century wrecks are found (Tolosa and Concepci6n). This coral region has not been documented.

2.2.7. Bfivaro- E! Macao- Punta Cana barrier reef system (Fig. 5). On the Eastern shores of the Dominican Republic, facing the Mona Passage, is the B~ivaro-E1 Macao- Punta Cana barrier reef system. This portion of the island has a configuration resembling a bold arrowhead. B~ivaro faces northeast into the Atlantic, E1 Macao is to the east (Mona Passage), and Punta Cana to the southeast (the Caribbean). The coastline is sandy, with mangroves, coastal lagoons, and swamps behind the coastal dune. These humids drain into the sea through numerous outlets or underground springs. The watershed is a coastal plain in the B~varo region. At E1 Macao and Punta Cana, reef terraces are usually found close to shore mainly at Cabo Engafio and near the airport. The reefs of E1 Macao, B~ivaro and Punta Cana extend for 70 km. There are marked structural differences between them. While E1 Macao and B~varo face northeasterly winds and high swells, Punta Cana faces southeasterly winds and waves. Thus E1 Macao and B~ivaro are high energy reef complexes with hard bases and eroded profiles, while Punta Cana has the characteristics of a low relief reef. The coral reefs of the Dominican Republic 91

El Macao (Arena Gorda) reef. This reef is located in the north central portion of the Mona Passage, close to Cabo Engafio, and has Atlantic reef characteristics. Its ap- pearance is similar to the one described for Puerto Plata and E1 Portillo in Saman~t. The smooth and solid limestone rises from the sandy surroundings up to 10 m, forming a high relief reef. There are no spur and groove formations here: the rocky formations are more like reef relicts from the late Quaternary period. The substrate where the reef has established itself is covered with algae (23%, of which 1.3% are encrusting and boring algae). The basal reef rocks of this reef have become brittle through bio-erosional processes, forming sediments and sand and creating temporary sediment plumes that affect coral recruitment and growth. This explains the low coral coverage (5.5%), most of which consist of small colonies no larger than 20 cm in diameter. B~ivaro reef. The breaker zone at B~ivaro can be as far as 3.5 km from shore, creating a wide lagoon (2-5 m deep) with coral patches and an extensive seagrass bed, followed by a broad porous and shallow back reef that reaches gradually onto the reef fiat. Coral species commonly found in this area are P. porites, P. astreoides, S. radians, M. complanata, A. cervicornis, Diploria spp., C. natans and the M. annularis complex. A. palmata skeletons covered with algae in association with Millepora sp. dominate the windward side of the breaker zone, which is narrow and steep. At 4 m, there are large dead stands of A. palmata as well as large boulders of the M. annularis complex and Diploria sp. Reaching the 13 m depth there is an irregular and wide sand channel, ending on its seaside in a rise of 8 m, which continues to form a smooth sandstone shoal, mostly covered by turf algae; during stormy events, this area becomes the first breaker region on the reef complex. These shoals are uneven, with crevices 2-5 m deep. Between them, shallow sand pockets appear and interconnect with others. Towards the deeper regions and on the frontal face of this shoal, the 10 m contour is reached with a gentle slope. The reef base then flattens and low-relief spurs (1.5 rn high) with sand and rubble-filled grooves begin to extend for 800 m or more, to about the 18 m depth. The coral cover for this portion of the reef is 16%. Punta Cana Reef. This reef is located in the southern portion facing the Mona Passage, and begins on its northern side following the cornering of Cabo Engafio by deep seas. Here, isolated patch reefs may be found close to shore. The Punta Cana reef grows closer to shore, characteristic of the fringing reef type, and is oriented towards the southeast. This zone is frequently hit by hurricanes. The Punta Cana sandy beaches are interspersed with low coral cliffs, which tum into high escarpments towards the south where the reefs end and there are deep nearshore waters. The reef lagoon region is shallow with rubble and sparse seagrasses. Several freshwater springs discharge underwater, thus influencing the type of biological diversity found here. The breaker zone at 5 m is narrow and composed of large, compacted skeletons of A. palmata; algae cover is high and few live corals are present. Seaward of the breaker zone, there is a sand and rubble area, with large boulders comprising the base of this frontal structure. The spur and groove area is of low profile and highly eroded. The 50 rn contour line is very close, and there are some areas where it is possible to find features used as dive destinations. The Macao-B~ivaro-Punta Cana Barrier Reef is now facing large impacts and threats from a steadily growing tourism industry. In a 30 km stretch of coast, about 11,000 hotel rooms have been constructed. The intense use of some hotel front sites and dive destinations has inflicted obvious damage on the reefs. In those areas close to shore, coral cover- 92 F.X. Geraldes

I 68o36 '

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Fig. 6. Parque Nacional del Este. age is less than 5%, and the seagrass beds as well as the lagoon patch reefs and backreefs are clearly degraded. There has been a rapid and unplanned development, without environmental impact assessments on the potential impacts of several activities, such as dredging and intense beach use creating sediment plumes, which mainly affect the backreefs, the reef flats, and the reef crests at Macao, B~ivaro and Cabeza de Toro. There are other impacts that have affected this reef, such as anchors, environmentally unconscious diving and snorkeling, and increased fishing pressure. The result of these actions has induced the algae growth, which has reached to 58% of substrate cover. Be- side the sediment effect described above, there exists no control over the release of detergents and other water treatment chemicals that are usually associated with hotel operations. The evidence of a stressed system is clear, and there is an unusually noticeable presence of the black band disease, affecting several coral species. The coral reefs of the Dominican Republic 93

2.2.8. Reefs of Parque Naeional del Este (PNE) (Fig. 6). The reefs of this protected area are basically low relief systems, found either as small, fringing, deep (20-30 m deep) patches, or as banks. Most of them are on the leeward side, protected by a land mass of Pleistocene and Recent reef terraces. Southeasterly trade winds are dominant. The reef on the leeward side can be divided into two distinct areas: that along the southern coast of Saona Island (influenced by oceanic currents,) and that along the western side of the Catuano Passage (more protected). The bases of the Saona reefs are consolidated hard bases and octocorals and sponges dominate the benthic communities. Hard corals are abundant only at the specific places where they concentrate, forming small, dispersed coral patches. Here the waves and currents are strong, and are in part responsible for sculpturing the reefs. The reefs west of Catuano mainly have sandy bases with patch reefs. Large amounts of sediment and biogenic sands are transported from the Catuano Passage and deposited along this coastline towards the west, with large seagrass meadows covering most of the nearshore areas. Corals mostly grow in patches from 12 to 30 rn deep. Further to the west, away from the influences of the Catuano Passage, coral patches increase in frequency and grow as deep-water fringing reefs, these being the most common reef structures of the southern coast of the Dominican Republic. Parque Nacional del Este (PNE) is the most studied marine site in the Dominican Republic (Vega 1994; Vega et al. 1994, 1997). Six categories of hard base substrate have been identified for this area: low relief spur and groove formations, reef flats, transitional reef communities, patch reefs, low relief rocky shoals, and rocky coasts. The basal substrate for these formations is consolidated carbonate reef, in addition to sediments and rubble.

Fringing Reefs Catalinita reef (Fig. 7.). This reef is located at the eastern end of a channel that separates the mainland f~om Saona Island. The base rises abruptly at the edge of the channel. In the deep portions, high relief spurs and grooves with large, rounded coral forms are common, while in shallower areas a hard base virtually without sand deposits, serves as a substrate for a large octocoral prairie. Following the 10 m contour depth, the base is covered by a wide frontal section of large skeletons ofA. palmata which project to the surface. In some areas where live colonies still exist, patches of Montastraea spp. and M. complanata form, mainly on the leeward side of the breaker zone and near Catalinita Island, which is a deposit of coral debris. Arreeife del Troneo (Catalinita reef). This is a leeward reef located to the north of Pasa Grande and south of Catalinita Island. Porites sp. is the dominant species at 0.5 m depth. At 3 m coral diversity increases. Algae (27 species) cover 50% of the area, sponges (16 species) occupy 5% of the area, octocoral (7 species) growth is sparse, and corals (14 species) cover 25% of the benthos. P. fureata is the dominant species. This setting forms a low frontal wall that ends in a narrow sand-gravel base, colonized by a dense seagrass bed of T. testudinum. Reef Crest, Catalinita Reef. Due to the act that this reef geographically faces the westbound currents, it is common to fred large amounts of solid waste from the heavy traffic of the Mona Passage and other offshore territories to the east. Dominating the top of the crest in 0.2 rn of water, is the short leaf type of Thalassia. There are also some live colonies of A. palmata, A. cervicornis, M. complanata, D. strigosa, M. areolata and S. 94 F.X. Geraldes

! 18o12'

Hard bottom 0 1.5kin

Punta Alglbe

68o27 . -- , _- .7 - _- c.;;" ...... ,'" Reef l~ Terraces ,iiiiiiiiiiii!iii!iiiii ; , Isla Catalinita [:~ Breakers [~] Reef fiats Back Reefs 20 m .....::::::::::::::::::::::::::::: ~] l',,lecushore Corrcr~nities [~ Mangroves ~ Deep reefs ;q~-'iiiiiiiiiiiii!ii!i!i?ii?ii?ii!: Seagrass ~ Son.mud/algae 7~ Hard bottom fJ//f///d//JJ////J/P corrcnunities J//

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ISLA SAONA

Fig. 7. Catalinita reef, Parque Nacional del Este.

radians. Below this, large, unconsolidated pieces of A. palmata lie like non-cemented tiles. At the 2 m deep, a patchy seagrass bed is established on top of coarse gravel (>25 cm diameter) substrate. In this zone, 8 species of algae cover 50% of the base. Sponges and octocorals are rare: There are 5 species of sponge and 3 species of octocoral. Hard corals are represented by 10 species, growing sparsely as small colonies. Adamanay Reef. This narrow fringing reef starts at E1 Faro to the West, and runs very close to shore along the village of Mano Juan, ending at Canto de la Playa in the central southern portion of Saona Island. The reef crest is separated by a shallow and narrow (30-50 m) sandy channel with seagrasses. Sporadic coral aggregations form the narrow reef fiat, which then continues into a Acropora-Montastraea zone; this is the basal structure of the breaker. In the seaward direction lies a hard carbonate platform, where gorgonians and coral grow in depths and configurations similar to other Saona Island sites. The Adamanay Reef protects the only human settlement in Saona Island, which dates back to the 1700s as a pirates' village, and is now a fishing settlement and a tourist destination. The coral reefs of the Dominican Republic 95

Low relief spur and groove communities The low relief spur and groove communities are mainly found to the west of Mano Juan, in a somewhat protected region of PNE, and in the south of Saona Island. They are located between 0.5 - 3 km away from the coast at 15 to 30 m deep. Sponges are dominant, as are octocorals and algae. Coral colonies are medium-sized and scarce. MAMMA's reef. This reef begins 18 m in front of Punta E1 Faro. The reef system is a continuation of a patch reef near the coast, in 3 to 8 m of water, dominated by live A. palmata and the M. annularis complex. Going seaward from this patch reef, a broad sand channel occurs, ending where the spur and grooves begin, with an orientation facing East towards the incoming waves and winds. Coral cover is low (7.5%), octocorals are the dominant benthic forms (17% coverage), and algae occupy 25% of the base. Parque Naeional. This reef is located westward of the mainland, at 15 m depth. It has small, narrow spurs (6-9 m wide and a length up to 25 m), and the grooves are narrow (3-5 m wide) and filled with fine sediments. The basic orientation of this reef is east to west. The algae are dominant with 15 species, forming patches with Halimeda opuntia and H. tuna as the dominant species. Twenty-nine species of sponge have been documented, with ,4gelas, Xestospongia and Verongula constituting the most common genera. Octocorals are abundant with 10 species, Pseudopterogorgia acerosa and P. americana being the most common species. Twenty-five species of hard corals have been observed, with relatively low coverage. The most common species are medium sized colonies of D. labyrinthiformis, S. siderea and M. cavernosa. El Pefi6n (CARICOMP/site). E1 Pefi6n has a hard base, with sparse sand-clay sediments at 15 m. This eroded spur and groove formation gently slopes towards a 2 m cut reminiscent of ancient shores, and into a sand channel where seagrasses grow. The dominant benthic fauna are the octocorals, with 18% coverage and 16 species, Briareum, Eunicea and Pseudopterogorgia being the most common genera present. Sponge coverage is 11%, with a diverse sponge composition (36 species), Agelas, Verongula, and Xestospongia being the most common. The corals account for 8% of basal coverage with 26 species (the highest species count for the whole PNE). Algae cover is significant; 24%, with 15 species represented. This location has been chosen as the CARICOMP permanent reef monitoring station for the country. Arrecife de Rub6n. This reef lies on the 20 m contour line that extends from Mano Juan westward. The reef's loose substrate is formed by sand, clay, and rubble. The benthic biodiversity is dominated by algae, with 14 species covering 25% of the substrate. Sponge diversity is also high, with 34 species covering in some cases up to 50% of surveyed areas. The most common genera present are again ,4gelas, Verongula and Xestospongia. Coral coverage is low (5%), and only 18 species have been documented. El Toro. This reef is found in the southwestern shore of Saona Island, at the 20 m contour depth. The orientation and position of this site faces the incoming currents and seas from the southeast. A spur and groove system forms its basic design, ending abruptly in the cut of an old shore terrace at 26 m depth, where large gravel and reef debris accumulate from the bioerosional processes occurring in the living portion of the reef. Algae cover (50%) is dominant, followed by octocorals with 22 species. Corals (25 species) grow, but cover no more than 5% of the base. 96 F.X. Geraldes

Bayahibe Reef. This reef is located at the northwestern limit of Parque Nacional del Este, 800 m away from shore, on top of a submerged terrace 18 m deep. It has a well- def'med spur and groove system, with relief up to 2 m high. The reef orientation is east- west, perpendicular to the shore. The coverage by corals is 34%, with 25 species; sponges 11% with 37 species; and algae 35%, with 16 species.

Hard base carbonate reef flat communities This reef type is characterized by a hard base flat carbonate substrate. These reefs are low relief, and are associated with high energy seas and currents. They are most common in the eastern portion of the park facing the incoming surge and waves from the open seas. In terms of diversity, turf and brown algae, and/or a co-dominance of algae and corals dominate them. Pasa Grande (Catalinita Reef). Algae, with 36 species, dominate this hard base. The most conspicuous genera are Dictyota, Turbinaria, Stypopodium and Halimeda. Coral colonies grow sparsely forming large individuals. Octocorals are few in number, represented by Gorgonia ventalina, Pseudoplexaura poros and Plexaura flexuosa: all are species adapted to high-energy conditions. Seven sponge species are present, encrusting and boring forms (Chondrilla nucula and Cliona langae) dominating. The corals are more diverse with 12 species, the most common being A. palmata, D. clivosa, P. astreoides and P. porites.

Transitional reef communities E! Faro. To the northwest of the Adamanay reef crest, somewhat protected by an extension of the southern coast of Saona Island, a low relief-high energy occurs at 7 m depth, mostly colonized by: algae, corals, and octocorals. It may be considered a transitional reef with accumulations of sediments and rubble. There are 24 species of algae, with Halimeda, Dictyota and Amphiroa dominating, and a cover from 25 to 50%. There are 22 octocoral species, with Eunicea, Plexaura, Plexaurella and Pseudopterogorgia being the most abundant, and encompassing 25% of the basal coverage. Hard corals are present, with 23 species growing scattered along the fiat bottom, covering 5% of it.

Patch reef communities These are located in protected waters in the western portion of the leeward side of the park, and inside the Catuano Passage, protected by the fringing reef and reef crest of Catalinita Island. Arrecife del Angel 1. This reef is located in shallow water (1.8-5.4 m), in the western entrance of the Catuano Passage, and is surrounded by a large seagrass bed. The structure is dome-shaped, with a diameter of 30 m, and is collapsed in its middle portion; hence the name that reminds one of a halo. In this middle portion Porites rubble accumulates. In its most exposed portion, large coral heads are found, with a tendency to link to the back reef by a rubble and deposition bed. Algae is the dominant form here with 50% coverage representing 21 species, the common ones being Halimeda opuntia, Caulerpa racemosa, Titanoderma sp., Stypopodium zonale, Amphiroa tribulus and Dictyota sp. Sponges here are scattered with a 5% coverage, but with 20 species present, of which Cliona langae and Iotrochota birotulata are the most obvious ones. Octocorals are very sparse, with 7 species found only in the periphery of the patch structure. Corals The coral reefs of the Dominican Republic 97 represent >5% coverage with 11 species present. P. porites, the M. annularis complex and M. cavernosa are the most common. Arrecife del Angel 2. Again in the Catuano Passage, at 5 rn depth, there is a patch reef with low relief, surrounded by a seagrass bed, sand, and gravel, with spotted and patchy coral colonies. Algae are dominant with 50% coverage and 15 species. Sponges have only 5% coverage with 18 species, with C. langae, Amphimedon compressa, I. birotulata and Aplysinafistularis being the most commonly seen species. Besides algae, the most dominant group is octocoral, with 13 species and large colonies (>50cm) of Eunicea, Plexaurella, Pseudoplexaurella and Pseudopterogorgia. Here coral cover is >5% of the substrate, with 15 species. There are moderate to large colonies (> 25 cm) of the M. annularis complex, M. cavernosa, D. labyrinthiformis and S. siderea. There are also acroporids present. Los Flamencos. This reef is located near E1 Faro reef, in the protected waters of the small cape which the coastline forms here. The reef is at 6 m depth and consists of a series of coral patches separated by sediments and rubble. The physical relief is medium to low in a hard substrate, where corals congregate to form outcrops of growth with large heads intercepted by the sand channels surrounding them. In some cases A. cervicornis is found, initiating the settling process and patch formation. Algae are represented by 21 species, dominated by Dyctiota sp. Octocoral fauna is common, with Eunicea, Plexaurella, Pseudoplexaurella, Pseudopterogorgia and Pterogorgia being the most commonly seen. Also common are the hard corals with 23 species. There are large (>2 m diam.) colonies of A. palmata serving as a basal structure for other species to settle on, such as A. tenuifolia and the rare M. squarrosa. Large numbers of Montastraea, Colpophyllia and Dendrogyra are also found in these coral patches.

Hard base carbonate platform communities Arrecife de Los Cocos. At 4 m depth, in the western portion of the Catuano Channel, there is a tidal channel frequently washed by strong currents generated from tides and winds, which structures the benthic community. Due to the strong currents, the dominant biota found are algae and octocorals, and in a lesser quantity, sponges and corals. Algae cover is above 50%, with 29 species present, the most common being Hypnea cervicornis, Acanthophora spicifera, Jania rubens and Laurencia intrincata. Of these, H. cervir represents more than 25%. Halimeda opuntia, H. tuna, Coelothryx irregularis, Amphiroa brasiliana and Galaxaura oblongata are also common. It is notable that the red algae are dominant in this location. Sponges are sparse around this reef site, occupying 5% of the base with 28 species. It is interesting to note that some of them can reach sizes up to 1.2 m, and the most commonly found species here are Amphimedon compressa, Pandaros acanthifolium and Callyospongia vaginalis. Octocorals are the most conspicuous group on this reef, with some individuals reaching up to 2 m high. They are well represented by 23 species, the largest number of species found in the park nevertheless their basal attach- ment or foot only covers 5% of the base. The common species are: P. acerosa, E. clavigera and E. calyculata, representing 60% of the total. Corals cover 1% of the base, with small colonies (<100 cm 2) of 14 species growing mainly in aggregations surrounded by rubble. The dominant species found are: M. alr P. astreoides, D. stokesii and D. labyrinthiformis, which represent 90% of the population. 98 F.X. Geraldes

2.2.9 Playa de la Isla Catalina reef, La Romana. On the southern coast of the Dominican Republic, to the south of the town of La Romana is a small island named Catalina, with two reef sites: one is a leeward reef, the other is a wall. The leeward reef lies seaward after a seagrass bed, sloping to 5 m depth; then a spur and groove system begins, ending at 14 m. Below this is a wide sandy area with coral patches. Recently this area has been turned into a cruise ship and tourist port, impacting the reefs there. Open water anchoring structures were deployed to tie these large vessels. Assessment studies have found a reduction of more than 80% in the overall benthic cover, due to the physical effects of these structures. Here, coral cover has been reduced from 12% to 4%; similar values have been found for sponge and octocoral cover, and algal cover increased from 15 to 27%. The solid reef platform has been transformed into rubble substrate. In general terms, this reef site is one of the most affected and damaged in the country. In nearby undisturbed areas, the average coral cover is 8%, with 31 species; octocorals cover 3%, with 14 species; sponges cover 8%, with 33 species; and algae cover 21%, with 19 species (Geraldes 1994b). The wall site is located on the northern tip of Catalina island, and extends some 500 m along the shore. It is separated from shore by a narrow (20 m) and shallow (2-3 m) zone. At 2-5 m depth a dense and healthy coral conglomerate is found, where A. palmata, the Montastraea complex, Diploria spp., M. decactis, Porites spp., M. miriabilis, many sponges such as Xestospongia spp. and Cliona spp. and octocorals cover most of the base. The wall starts abruptly at the eastward margin, and vertically extends over 40 m, ending at a sandy base. On the wall Halimeda spp. and plate forms of Agaricia spp., together with P. astreoides and Montastraea spp., are common. In the deeper areas, anthipatharian and octocorals are also common, in crevices and along the vertical hangings.

2.2.10. Reefs of Juan Dolio - Guayacanes, San Pedro de Macoris On the south coast and 25 km to the west of the Higuamo and Soco rivers is the fringing reef of Juan Dolio-Guayacanes, extending for 10 km, close to the shore. This reef protects a narrow sandy beach on a low-laying terrace. Due to the proximity of major cities, tourist installations have been developed along this coast in excess of its carrying capacity. In turn, beach intervention has deteriorated the reef setting and created conflict between users of this region. Tunnel Reef. The coastline near this reef is composed of porous and calcareous reef terraces, and sandy shores. The Higuamo River influences the area. This reef is made up of a crest, with sparse patches. Seaward from the reef crest and parallel to it there is a rubble and sand channel; upon reaching 7 m depth, a high relief spur and groove system begins with relatively healthy coral growth covering 33% of the base. This is one of the few places in the country where coral cover surpasses algal cover (26%). Villas del Mar- Juan Dolio reefs. The breaker zone of this area is composed of A. palmata, D. strigosa, the M. annularis complex, M. complanata, P. porites and P. astreoides. Most are dead colonies covered by turf algae. They are not cemented, but are placed in situ. They are heavily used by beach goers and are usually covered by free sediments and detritus, together with solid waste and garbage. Encrusting and boring sponges, such as Anthosigmella varians and C. langae, thrive here. The deterioration of this reef crest has created a stronger shore current, which has increasing beach erosion and The coral reefs of the Dominican Republic 99 sand transport inside the reef lagoon. The lower Palmata zone has started to receive large amounts of sediment coming from the lagoon region, where the presence of encrusting and boring organisms has also increased. Towards the deeper region (12 m), the reef has not yet suffered, due to the buffering effect of an 80 rn sand channel. A high relief reef begins with the following basic benthic composition: coral 33%, 8% octocorals, 2% sponges, and 26% algae. Stresses are evident due to bleaching and black band disease being present in some of these colonies. Punta Garza. This system is a unique reef front formed by a Porites-Montastraea association, without acroporids. It grows towards the shoreline and connects with it by a very shallow and often exposed (at low tides) sand accumulation, where Syringodium dominates. Eight species of algae, 10 species of octocorals, 7 species of sponges, and 21 corals represent the most common benthic organisms there. This region is being severely impacted by dredging and jetties deployed to sustain the development of tourism. This has also increased the turbidity of the coastal waters, and it is now common to find large amounts of sediments covering coral colonies, with discoloration and death of organisms found in some cases. Guayaeanes Reef. The conditions of the shore near this reef are similar to the ones described for the other reefs in this coastline" they are severely impacted, due to urban and tourist development. The reef is divided into two portions. The eastern portion is a very shallow (0.2 - 1 rn deep) reef fiat hard platform (10 - 40 m wide), that connects to the beach. The cemented remains of Porites and Montastraea are the main components of the base of this reef crest. Seaward, this platform drops abruptly (2 m) to a hard substrate, which continues into a shallow sand channel; there, MiHepora and octocorals appear. Further east, at Punta Cadillo, this reef type changes back to a typical A. palmata breaker, although it is equally highly impacted and covered by algae such as Mycrodictyon sp. and A canthophora sp., in addition to zoanthids (Zoanthus pulchellus).

2.2.11. Boca Chica reef, Bahia de AndrOs, Santo Domingo (Fig. 8) This reef has the longest history in the country, as a study case. The Boca Chica reef is located in the center of a bay, and has a south-southeasterly orientation, facing dominant winds and swells. The coast is a dissolution basin of the carbonate terraces that are characteristic of the south coast. At this site, several underground springs emerge from the Rio Brujuelas, creating a shallow sand deposit suitable for a f~inging- barrier reef establishment. In the mid-1930s the lagoonal area of this reef was dredged, and the material used as landfill in the coastal mangrove swamps, to reclaim the land and convert this into a resort town. Since then, hotels, sugar mills, and port facilities have been installed, sheltered by the reef barrier. In this process, Isla la Piedra was created using the by-products of port construction and dredging. There is also a small natural mangrove key called La Matica. Today, the region hosts 150,000 permanent residents and 5,000 hotel rooms, in a stretch of less than 3 km of coast, thus creating intense pressure and impacting the coastal and marine resources and habitats found here. The reef of Boca Chica may be described as an eroded spur and groove formation, with a clearly defined zonation pattern as regards depth: sandy beach, back reef lagoon, reef crest, breaker zone, lower palmata zone, spurs and grooves, sand channel, buttress zone, and drop-off. All these reef regions are impacted either by natural stresses (3 hurri canes in 25 years) or by anthropogenic activities (overfishing, coral extraction, dredging, 1O0 F.X. Geraldes

Boca Chica ~:,.:.,:,:,,:,,:::,,:,,:,,:,:.,:,,:,,:,,:,:,:.,:,,:,,:,,:,,~--; ~'*'~' ',',', ', ',',',','| ',',

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69o37 ' I Fig. 8. Boca Chica, Bahia de Andr6s. pollution, divers, etc.). The regions where this is most evident are: La Matica key, the backreef, reef fiat, the breaker zone, and the spurs, which in some places are well eroded and have a slow cementation. At approximately 10 m depth, there are large concentrations of loose rocks and reef debris, with rubble covering 20% of the base. Nevertheless, this reef's benthic composition still has a predominance of coral coverage with 20%. Sponges are also present, with a 5% cover. Algal cover is high (56%). The major groups of benthic organisms found here are represented by: 12 species of algae, 6 species of octocorals, 22 sponges and 30 corals. Agaricia spp. and the M. annularis complex are the most common species found. Algae such as Laurencia sp., and encrusting sponges (Cliona sp. and A. varians) are also common. In the shallows, at the breaker zone (1-4 rn deep), large colonies of A. palmata, the M. annularis complex surrounded by P. astreoides, P. divaricata, P. porites, M. complanata, and G. flavellum, are still present. Nevertheless, the seascape seems catastrophic. Pieces of coral lie all around, and degradation by physical with bioerosional processes is obvious. Diadema antillarum and tunicates, boring sponges, and large amounts of turf algae cover all available basal surfaces. Amongst all this, small colonies of A. palmata appear. At the breaker zone, the Acropora barrier is dead, but still in place and functional. Encrusting algae (Porolithon pachydermun) and zoanthids fill the surface area of this zone. The reef crest and reef fiat contains large amounts of reef debris. The intense erosional process found here produce sand for the Boca Chica beach. In deeper waters, at about 30 m, plate forms of Agaricia spp., the M. annularis complex, C. natans, M. meandrites, M. angulosa and Mycetophyllia spp. accompany a The coral reefs of the Dominican Republic 101 massive growth of Halimeda sp. and Amphiroa sp. At 20 m, Erythopodium spp. appear with the encrusting sponges C. langae and Ectyoplsia ferox. Some large Xetospongia muta are also found. Corals are not the dominant features in this seascape, but there are well-developed colonies of A. cervicornis, D. cylindrus, P. astreoides, P. furcata, M. alcicornis and Agaricia spp.

2.2.12. Parque Nacional Submarino La Caleta, Santo Domingo. This is an 18 krn2 marine protected area located on the leeward side of the Caucedo Peninsula. Rocky shores surround the area. Nearshore, there is a sandy base with patchy corals. This continues into seagrasses, and at 10 m spurs and grooves appear. At greater depths (18 m) there is an abrupt drop to 25 - 40 rn, where there are low and medium relief spurs and grooves. Here, there are hard base reef-fiat carbonate platforms, reef walls, and sunken structures deployed there to serve as artificial reefs and fish aggregation devices. The main benthic organisms recorded for this area are: 32 species of coral, (A. agaricites, D. strigosa, L. cucullata, M. decactis, M. meandrites, M. alcicornis, M. cavernosa, P. astreoides, S. siderea and Stylaster roseus, amongst others), 20 species of octocorals (G. flabellum, is the most common, followed by E. caribaeorum and P. bipinnata, together with P. homomalla and P. americana), 50 species of sponges (Amphimedon compressa, Aplysina cauliformis, Ircinia strobilina and Pseudoceratina crassa are found at almost all depths, followed by Agelas conifer, Callyospongia vaginalis, Ectyoplasia ferox, Iotrochota birotulata and Xetospongia muta), and 45 species of algae. The benthic coverage is as follows: algae 41%, sponges 13%, octocorals 13%, and corals 28% (Geraldes 1994a; Geraldes and Vega 1995a).

2.2.13. Najayo-Palenque Reef. On the southem coast, a large open bay is found between Punta Caucedo to the East (La Caleta) and Punta Najayo to the West, where the Najayo- Palenque Reef is located. The site sits after the Ozama - Haina- Nigua river complex. These three rivers discharge in Santo Domingo's (Capital City) coastline, which is composed mostly of reef terraces until reaching the Nigua River. These rivers do not form a tree estuary, due to the great ocean depth found nearshore (>300 m). The water discharged is polluted, and enriched with sediments from the cities of Santo Domingo and San Crist6bal and the surrounding sugar cane and farm areas. The coastal currents flow westward towards the Najayo-Palenque Reef. These reefs are typical of high energy conditions, with low profiles dominated by a hard, current swept base, and sparse octocoral growth. It is assumed that the substrate is non-uniform due to tectonics or erosional processes caused by catastrophic events that have altered it, leaving boulders and uplifted tile-like structures where corals attach in the leeward domain.

2.2.14. El Derrumbao Wall, Las Salinas, Bani. From Najayo and Palenque westward, the climate changes to very dry. The Derrumbao Wall is located near a coastal desert with large sand dunes. The shore is composed of dark sand and coarse gravel of terrigenous origin. Through bio-cementation processes, sandstone is found close to the shore, serving as appropriate substrata for corals and other life forms to establish themselves. This shallow, seagrass-dominated feature terminates some 30 m from shore, where there is an abrupt drop towards a submarine fault (> 1,500 rn deep). Cemented sand rectangular blocks, measuring 3-5 m by 2 m, form the wall. Madracis sp., P. porites, P. 102 F.X. Geraldes

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Fig. 9. Puerto Viejo, Azua_ the M. annularis complex, M. cavernosa, C. natans and Siderastrea siderea are common species found in the first 10 m. Then at this depth, an overhang occurs and cryptic species abound. Several large sponges, such as Cliona sp., E. ferox and X. muta can also be found. After the overhang, the wall turns into sand, with a large angle of slope, which is usually covered with f'me sediments that are easily stirred by divers.

2.2.15. Puerto Viejo, Azua, Bahia de Oeoa (Fig. 9). This reef site lies in a dry region on the southwestern coast. The reef forms a small barrier with an 8 km2 lagoon. Due to its sheltering features, it has been used as a natural port since the 1500s. In 1957, a modem port was built which altered the reef. At the breaker zone there are several keys formed by reef deposits that are now colonized by mangroves. Near the shore tidal fiats with Halodule and Syringodium are common. Corals are found in the sand depressions or erosional pits, the most common being M. aerolata, P. porites, P. divaricata, S. radians and D. strigosa. The lagoon has a variable depth of 0.5 to 14 m, and T. testudinum is dominant. Corals grow in patches, and Diploria spp. and Siderastrea spp. dominate. Others, such as A. cervicornis, M. complanata, P. astreoides, A. agaricites, P. furcata and the rare The coral reefs of the Dominican RepubBc 103

Acropora prolifera are sparsely found. Strong tidal currents flow and influence the circulation of this lagoon, enhancing coral growth especially in the back reef, where A. cervicornis, P. astreoides, P. divaricata, D. strigosa, S. siderea, M. areolata, S. radians and M. alcicornis are the common species. The reef fiat is dominated by zoanthids growing on top of coral debris. Corals like P. astreoides, D. clivosa and very large colonies of Acropora palmata dominate in the breaker zone. In deeper waters (10 m), a hard basal carbonate platform is found, with sand pockets. Large tree-like colonies of A. palmata, as well as large boulders of C. natans, are found here creating intricate structures where other reef creatures seek refuge and nourishment. These coral patches tend to grow closer together in deeper water. At 13 m deep, they usually merge to form low relief spurs and grooves.

2.2.16. Barahona reef. The reef at Barahona is a fringing reef that has not been thoroughly studied. It consists of a well developed spur and groove system, breaker zone, reef fiat, and islets that surround a lagoon, which has been converted into a port. The reef barrier was cut, dredged, and enlarged to allow for port transit and ship movement. On the protected coast lies the town of Barahona. Nearby and upstream there is the Yaque del Sur River estuary with considerable sediment loads that sometimes reach this reef and affect it.

2.2.17. Parque Nacional Jaragua. Parque Nacional Jaragua is located at the southwestern end of the Dominican Republic. This is a dry region, where cacti and desert-like vegetation dominate. No rivers or surface runoff is found in these Pleistocene carbonate reef terraces. On its windward side strong seas, medium-sized cliffs, and high energy pebble and pocket beaches with fringing reefs are found. On its leeward coast, protected by high cliffs, sheltered, long, white sandy beaches are common, with consolidated hard carbonate offshore, where coral cover and density is high. There is not a well developed fringing or bank reef in most of the zone, except near Cabo Rojo (Weil 1997). At the southern tip Beata Island is found, and further out is Alto Velo Island. Moving towards the southwest, at the edge of the continental shelf, there is an elevation of the seafloor called Los Frailes Shoals. Other descriptive studies of this area are included in Borrell (1981) and Vega (1981). Beata Island reefs. This island is formed by reef terraces and has a low relief. On its windward side at its northeastern end, coral reefs are located offshore forming bank reefs (Weil 1997). On the leeward side there are seagrass beds, followed by a hard carbonate base with sparse coral, sponge, and octocoral growth, and with a high cover of algae. The most abundant corals are Porites, Undaria (Agaricia), Diploria, S. siderea and M. cavernosa. Alto Velo Island. This small island, located 20 km. away from the mainland, is an oceanic island of volcanic origin. Its slopes are bare and drop steeply into the ocean depths. In the shallows on the leeward side (west), one can find large boulders and caves/crevices formed by land slides. Here, cryptic reef creatures, mostly of crustose and flattened forms, are sparsely located. Los Frailes Shoals. Located 12 miles southwest of Cabo Rojo, Los Frailes shoals is a seafloor intrusive elevation, that rises from the seafloor, and surfaces in a similar manner to Alto Velo Island. These rock outcrops receive clean oceanic waters, which allow the establishment of a diverse community. There are boulders and submerged walls 10 m 104 F.X. Geraldes high, covered by Tubastrea aurea. Large Montastraea, Diploria and Colpophyllia colonies, as well as sponges, are present on the other side (Weil 1997). Cabo Falso to Bahia de Las Aguilas. In the Beata Canal, where there is protection from the incoming ocean lies a platform that reaches a depth of 18 m. It is covered with algae, hydrozoans, gorgonians and Cliona, as well as some A. palmata colonies (Weil 1997). The deeper areas have a highly diverse community associated with large boulders, where octocorals, zoanthids, and sponges predominate. At Lanza Zo to the north, in 15-25 m of water, there is a bank reef located some 800 m offshore which runs parallel to the shoreline. Here algae such as Dyctiota, Lobophora, and Halimeda are abundant. Sponge diversity and abundance is also significant, but corals are sparse. At Bahia de Las Aguilas the reef has a higher coral diversity and coverage, with Montastraea, Porites, Undaria (Agaricia) and Agaricia being dominant. Rare species such as Mycethophyllia reesi are uncharacteristically abundant (Weil 1997). Cabo Rojo. This reef is located a few kilometres northwest of Cabo Rojo, where the continental shelf forms a submarine canyon. The reef begins at 18 m on a wide platform covered by seagrasses. The reef then drops to 45 m, to a sandy base. There is a large degree of cover by the plate-like Montastraea, Agaricia, and Undaria, and large Colpophyllia colonies can be found. The blue-green algae Schyzothrix is also abundant between 13 and 18 m. Octocorals are scarce and scattered along the slopes. Sponges are abundant and diverse. Millepora is common in the shallow areas, where A. cervicornis patches are common. Signs of white band disease are present. M. franksi and A. lamarcki are the dominant coral species in the deep area of the reef (Weft 1997).

3. NATURAL DISTURBANCES

3.1. Sedimentation The Dominican Republic occupies a fairly large land mass. There are large rivers and streams washing extensive watersheds, and there are usually no coral formations directly downstream from them. The Caribbean coast is basically composed of carbonate reef terraces, allowing shallow fringing reefs to develop. On the northeastern region there are mountainous terraines close to the shore, associated with increased rainfall, which in turn cause short torrential streams that drain into the adjacent sea, loading it with sediments, and limiting reef growth. This occurs for approximately one third of the coastline. Along the rest of the coast, reef growth is of the fringing or barrier type. These usually occur in association with the dry regions of the country, where waters are clear. Never- theless, even in these dry regions there are three places that have natural sediment inputs and restrict reef settlement: Punta Martin Garcia in Barahona, Punta Salinas in Peravia, and El Morro in Montecristi.

3.2. Coral bleaching Coral bleaching events have been associated with abnormal water temperatures and other undetermined stresses. There has not been a country-wide study of bleaching for the Dominican Republic. There are, however, reports of its occurrence at most reefs, the most evident being the reef sites near major urban settlements which are more heavily visited or overfished, such as Puerto Plata, Sos~a, Las Terrenas, Macao, B~ivaro, Gua- yacanes, Boca Chica, and La Caleta. The coral reefs of the Dominican Republic 105

3.3. Mass mortalities and other stresses The mass mortalities of A. palmata and D. antillarum have triggered major ecological changes in the coral reef ecosystem. These no longer resembles the classical description of coral reefs by Goreau and Goreau (1973). The breaker and lower palmata zones have been specially affected by the mortality of this coral. This causes, the urchin to die off, and this in association with overfishing and increased nutrient pollution has enabled algae to flourish and cover the eroding reef environment, which perpetuates reef erosion. In some reefs Millepora sp. and Zoanthus sp. are seen at the breaker zone, substituting for A cropora.

3.4. Hurricanes and tropical storms Hurricanes and tropical storms are common in Hispaniola. There have been more than 200 of these events recorded since the fifteenth century. These phenomena are more common for the Caribbean southern coast, rarely affecting the Atlantic coast. Neverthe- less, all major reef sites have been affected by at least one of these events. The major direct effect is the initial impact and there is massive destruction of the coral stands. Large portions of living material are usually transferred and deposited on top of the reefs. This debris accumulation later turns into suitable substrate for furore colonization of corals, and may form islets on the reef fiat, to be colonized by a succession of mangroves and other coastal vegetation. In the deeper portions the debris is accumulated in two regions of the reef: at the deepest portion of the spur and groove in the sand channel, and at the end and deep portion of the buttress zone. This debris is either consolidated, serving as base for future growth, or bioeroded and turned into sand and pebbles. Other effects are those caused by the increment in water turbidity and reduced salinity, due to the large volume of downpour rains and excessive runoff due to deforestation on the nearby watersheds. This increases river flooding as well as sediment loads, which reach distant reef sites. In these locations, it is common to f'md stresses in corals, where bleaching and other stress-related symptoms are obvious amongst the reef inhabitants. These reefs usually remain in a delicate condition. For their mitigation, there is a need for ample reforestation programs of the watersheds, as well as an integrated approach to coastal zone management. Reefs that are in this situation and described in this work are: Juan Dolio-Guayacanes, Boca Chica, E1 Portillo, Playa Dorada, and Puerto Viejo reefs.

4. ANTHROPOGENIC IMPACTS

Destruction and degradation of coral reefs now appears near most human settlements, due to the increase in negative environmental impacts to the aquatic and coastal environments. There is also clear evidence that some of these impacts are caused by remote human activities such as agriculture, animal husbandry, and industrial development. In general, land-based fertilizers, pesticides, domestic and industrial trash, and wastes from the mining industries are all reaching the river mouths, estuaries, and adjacent marine ecosystems like coral reefs (Puerto Plata, Sosua, Las Terrenas, Miches, Juan Dolio-Guayacanes, Boca Chica, La Caleta, Palenque) via runoff. Coastal development has led to the destruction of wetlands and mangroves for landfills, coastal construction, and dredging. Ports and shipping activities, recreational boats and marinas, some localized and rare reef-based coral mining, can also be found in Montecristi, Luper6n, Puerto Plata, Sosua, 106 F.X. Geraldes

Las Terrenas, Macao, B~ivaro, Punta Cana, Bayahfbe, Guayacanes-Juan Dolio, Boca Chica, La Caleta, Palenque, and Barahona. Overfishing is also heavily affecting Dominican reefs. Overharvesting of commer- cially important species such as Strombus gigas, S. pugilis, Panulirus argus, and fish of the Serranidae, Lutjanidae, and Scaridae families, is a problem. Lately, there has been an increase in the harvesting of other reef creatures such as black corals, hermit crabs, and omarnental reef fishes, starfish, sea urchins, and live-rocks for the souvenir industry. Most of these activities are prohibited or regulated by Dominican laws, but there is a lack of political and financial support to achieve the proper enforcement and personnel training. In order to minimize the negative effects on reefs, non-government organiza- tions have contributed with conservation actions.

5. CONSERVATION AND MANAGEMENT

Actions in this regard have been implemented since 1962, due to efforts of the Centro de Investigaciones de Biologia Marina of the Universidad Aut6noma de Santo Domingo, which is the pioneer institution on marine and coastal issues and studies in the country. Later, in 1984, the Fundaci6n Dominicana Pro-Investigaci6n y Conservaci6n de los Recursos Marinos (MAMMA), promoted conservation actions on behalf of coastal and marine resources, especially issues regarding coral reefs and fisheries. All these activities motivated the Government to decree the presidential act (No. 112/96) that specifically protects coral reefs in the Dominican Republic. The Governmental bodies in charge of protecting this natural resource are: the Dominican Navy and its Coast Guard unit, the Subsecretaria de Recursos Naturales of the Ministry of Agriculture, and the Instituto Nacional de Protecci6n Ambiental (INPRA). As part of the country's conservation efforts, there exist on its coasts three relatively large representative coastal-marine protected areas, with big tracks of coral reefs, in good natural conditions, however they are under fishing pressure. These parks are: Montecristi in the North shore, del Este in the Eastern portion, and Jaragua in the South West; all shelter important biodiversity resources, and create nurseries for economically species of interest as well as unique tropical marine and coastal habitats and representative ecosystems.

ACKNOWLEDGMENTS

I am particularly grateful to my wife and colleague M6nica B. Vega for her help in review and editing of this manuscript, as for her companionship on field expeditions. I would also like to thank Enrique Pugibet, Rub6n Torres, Yira Rodrfguez, Kathleen Sullivan, John Tschirky, Mark Chiappone, as well as the staff of CIBIMA and Fundaci6n MAMMA, Inc. for their support.

REFERENCES

Bamwell, F.H. 1983. Impacto del Hurac~ David (1979) sobre los corales de la cresta del arrecife de Boca Chica, Bahia de Andr6s, Repfiblica Dominicana. Contribuciones del CIBIMA, Santo Domingo, Dominican Republic 15: 1-10. The coral reefs of the Dominican Republic 107

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TAXONOMIC LIST OF STONY CORALS OF THE DOMINICAN REPUBLIC

Class Hydrozoa Order Athecatae Family Stylasteridae Stylaster roseus Family Milleporidae Millepora alcicornis (Linnaeus) M. complanata Lamarck M. squarrosa Lamarck Class Anthozoa Order Scleractinia Family Astrocoeniidae Stephanocoenia michelinii Milne Edwards and Haime S. intersepta Family Pocilloporidae Madracis decactis (Lyman) M. formosa Wells M. mirabilis Wells M. pharensis (Heller) M. pharensis (Heller) forma luciphilia Wells M. senaris Wells Family Acroporidae Acropora cervicornis (Lamarck) A. palmata (Lamarck) A. prolifera (Lamarck) Family Agariciidae ~ Agaricia agaricites forma agaricites (Linnaeus) forma carinata Wells forma danai Milne Edwards and Haime formapurpurea (Lesueur) A. fragilis forma fragilis Dana A. humilus Verrill A. lamarcki Milne Edwards and Haime A. grahamae Wells A. tenuifolia Dana Leptoseris cuculIata (Ellis and Solander) Family Siderastreidae Siderastrea radians (Pallas) S. siderea (Ellis and Solander) Family Poritidae Porites astreoides Lamarck P. branneri Rathbun P. porites (Pallas) P. furcata Lamarck P. divaricata Lesueur Family Faviidae Colpophyllia natans (Houttuyn) C. breviserialis C. amaranthus Diploria clivosa (Ellis and Solander) D. labyrinthiformis (Linnaeus) D. strigosa (Dana) Favia fragum (Esper) Manicina areolata forma areolata (Linnaeus) 110 F.X Geraldes

forma mayori Wells The Montastraea annularis complex (Ellis and Solander) M. cavernosa Linnaeus M. franksi Solenastrea bourmoni Cladocora arbuscula (Leseur) Family Rhizangiidae Astrangia solitaria (Lesueur) Phyllangia americana Milne Edwards and Haime Family Oculinidae Oculina difussa Lamarck Family Meandrinidae Dendrogyra cylindrus Ehrenberg Dichocoenia stellaris Milne Edwards and Haime D. stokesi Milne Edwards and Haime Meandrina meandrites forma meandrites (Linnaeus) M. memorialis M. braziliensis Milne Edwards and Haime Family Mussidae Isophyllastrea rigida (Dana) Isophyllia sinuosa (Ellis and Solander) Mussa angulosa (Pallas) Mycetophyllia aliciae Wells M. danaana Milne Edwards and Haime M. ferox Wells M. lamarckiana Milne Edwards and Haime M. ressi Wells Scolymia cubensis Milne Edwards and Haime S. lacera (Pallas) S. wellsi Family Caryophylliidae Eusmilia fastigiata (Pallas) Family Dendrophylliidae Tubastrea coccinea Lesson T. aura Rhyzosmilia maculata