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Abstract

Key words: coral reefs, Costa Rica, monitoring, GCRMN, conservation, management.

Resumen

Costa Rica tiene comunidades coralinas y arrecifes en la costa Caribe y del lado Pacífico a lo largo de la costa y en islas mar afuera). Arrecifes de franja y de parche, bancos carbonatados y una cresta de algas coralinas incipiente están presentes en la sección sur de la costa Caribe. Comunidades coralinas, arrecifes y colonias de coral aisladas se encuentran a lo largo de todo el Pacífico de Costa Rica. Los arrecifes coralinos han sido impactados seriamente en los últimos 30 años, principalmente por sedimentos en el Caribe y algunos arrecifes del Pacífico, y por el calentamiento durante el Fenómeno de El Niño-Oscilación Sureña en el Pacífico. Tres sitios en el Caribe y otros tres en el Pacífico están siendo monitoreados. Después de reducciones significativas en la cobertura de coral vivo en la década de 1980, tanto en el Caribe como en el Pacífico, la mayoría de los arrecifes se está recuperando. El gobierno de Costa Rica es consciente de la importancia de los arrecifes coralinos y de otros ecosistemas marinos, y en años recientes han implementado (o están en proceso de aprobación) varios decretos para la protección de los arrecifes. Pero, hay limitación de fondos y personal para su adecuado manejo y protección. Se necesita más divulgación de información sobre los arrecifes coralinos al público en general y usuarios de los mismos; de nuevo, la limitación de fondos restringe lo que se puede hacer.

Palabras clave: arrecifes coralinos, Costa Rica, monitoreo, GCRMN, conservación, manejo.

Costa Rica has coral reefs on the Caribbean coast and on the Pacific side (coast and off-shore islands). The Caribbean coast is 212km long and consists mainly of high-energy sandy beaches that on the Southern section are interrupted by carbonate promontories (Fig. 1). These promontories are made up primarily of fossil reefs (Pleistocene, Holocene), and beachrock in some sections. Extant reefs develop on top of these rocky outcrops (Cortés & Guzman 1985, Cortés & Jiménez 2003a). Three coral reef areas are recognized on the Caribbean coast: 1) fringing reefs between Moín and Limón, 2) fringing and patch reefs, and carbonate banks at Cahuita National Park, and 3) fringing and patch reefs, carbonate banks, and incipient algal ridge between Puerto Viejo and Punta Mona (Fig. 1). Seasonal variability is minimum along this coast, with rains year round and no upwelling. Most of the touristic activity on the Southern Caribbean coast of Costa Rica is oriented toward the reefs, as are also the fisheries.

The Pacific coast of Costa Rica is 1 160km long, and has a high diversity of habitats: rocky shores of a wide variety of rock types, sandy beaches of several compositions and grain size, mangrove forests, estuaries, a tropical fjord, islands of various sizes, and several gulfs and bays (Fig. 2). The northern section of the coast (Gulf of Papagayo) is characterized by a dry tropical forest, with a very dry season from December to April, and a rainy season from May to November. This part of the coast is exposed to seasonal upwelling (McCreary et al. 1989) during the dry season due to the crossing the isthmus of Trade Winds. The central and southern coast is covered with tropical rain forest, with a transition zone in between in the Central Pacific coast. Coral communities, reefs and isolated coral colonies can be found along the Pacific of Costa Rica, which has been divided into eight regions (Cortés & Jiménez 2003b): 1) Santa Elena, in the Northern section of the coast next to the border with Nicaragua; 2) Bahía Culebra, to the South of Santa Elena and North of Península de Nicoya; 3) Península de Nicoya, mainly the seaward side of the peninsula, since the inner section is an estuarine environment with very little coral growth; 4) Pacífico Central, extends from the East end of Golfo de Nicoya to the largest mangrove area of Costa Rica, the Sierpe-Térraba complex; 5) Península de Osa, the outer section and part of the entrance to Golfo Dulce; 6) Golfo Dulce, includes the reefs within the gulf; and offshore islands: 7) Isla del Caño, 15km from the coast, and 8) Isla del Coco, more than 500km from the coast (Fig. 2).

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Moín-Limón: Fringing reefs and carbonate platforms are found in this section of the coast which is subject to heavy wave action, with the largest reefs growing on the leeward side of the islands. This area is exposed to freshwater runoff, sewage, petroleum pollution and solid wastes from the main ports on the Caribbean (Mata et al. 1987, Guzman & Jiménez 1992, Guzman & García 2002, Acuña-González et al. 2004). The coast between Moín and the Port of Limón is bordered by a carbonate platform that extends from above the upper tidal zone down to 15m depth (Cortés & Guzman 1985). The submerged platform is covered with algae, hydroids, sponges and isolated corals. The most abundant corals in 1983, when the area was surveyed, were: P. astreoides and A. agaricites forma purpurea (Cortés & Guzman 1985). Beyond 15m depth the bottom consisted of soft mud (Cortés 1998). Isla Uvita is off the Port of Limón (Fig. 1), but a strong current separates it from the port and its pollution. So, it is possible to observe well-developed reefs around the island. The east side had an extensive carbonate platform, with microatolls of Siderastrea siderea in the lagoon that was further exposed with the 1991 Limón Earthquake (Cortés et al. 1992). A monitoring station was established in 2005 in the west side, coral cover was very low and sponges were dominant (Fonseca et al. 2006a).

Cahuita: The largest fringing reef of the Caribbean in the country is located at Parque Nacional Cahuita (Fig. 1). This reef consists of three barriers; the outer barrier, stretching for 5km from the Western side of Punta Cahuita, to the Eastern side, it is separated from the coast between 100m and 2km. Between this barrier and the coast there is a small fringing reef, about 500m long and within 100m from the shore, called the inner crest. Finally, a 100m barrier is located of the Western end of the reef (Cortés & Risk 1984, 1985, Cortés & Guzman 1985). Partch reefs are found within the lagoon and carbonate banks off shore. The coral reef at Cahuita had been degrading during the last 30 years due to siltation, other anthropogenic impacts and natural causes (some of them exacerbated by human activity): massive death of organisms, earthquake, and warming events. Live coral coverage decreased from 40% in the early 1980's to 10% in the early 1990's (Cortés 1994). Using the AGRRA surveying protocol, live coral was 2-3% in the late 1990's (Fonseca 2003). The reef of Cahuita has been monitored since 1999 using the CARICOMP monitoring protocol, and in recent years with funding from the Southern Tropical America-Global Coral Reef Monitoring Network (STA-GCRMN) node. At the CARICOMP site, live coral increased slightly from around 15% to 17% in 2004, however turf algal cover has increased significantly (Fonseca et al. 2006a, Fig. 3).

The populations of the black sea urchin, Diadema antillarum, which were decimated in the early 1980’s in Costa Rica and wider Caribbean (Lessios et al. 1984, Murillo & Cortés 1984) are recovering. Before the 1983 mass mortality densities were 3.6 to 8.8 ind/m2 (Valdez & Villalobos 1978), after the mortality densities were 0.25 to 2 ind/m2. By 1992 it had reached 0.001 ind/m2 (Cortés 1994) and in 2004, 0.3 to 0.7 ind/m2 (Alvarado et al. 2004, Fonseca et al. 2006a). The increase in Diadema may be contributing to the recovery of the reef at Cahuita (Wyckoff & Cortés unpubl. data). Similar observations were done by Myhre & Acevedo-Gutiérrez (2007) in Manzanillo, in which live coral cover has increased. Mean while, Echinometra viridis decreased significantly between 1999 and 2004, from 2 to 0.07 ind/m2 at the CARICOMP site (Fonseca et al. 2006a).

Bahía Salinas: Located on the North Pacific extreme of Costa Rica and exposed to the seasonal upwelling, the area around Salinas has isolated, but relatively large coral communities made up by Pavona gigantea. Other zooxanthellate corals found were Porites panamensis and a few colonies of Pavona varians. Many azooxanthellate corals have been found: Tubastrea coccinea, Cladopsammia eguchi, Oulangia bradleyi, several species of Astrangia and one species of Phyllangia (Cortés unpubl. data). Many species of algae, octocorals, mollusks and polychaetes have been collected and are now being studied.

Santa Elena: Coral reefs and communities North and South of Península de Santa Elena (mostly within the Área de Conservación Guanacaste, Fig. 2) were described by Cortés (1996/1997). Corals found in low densities in other parts of the Eastern Pacific, formed small patch reefs there, for example Pocillopora eydouxi and Pavona gigantea. Relatively large reefs frameworks built up by Pocillopora damicornis have been found on the North side of Península de Santa Elena. Coral communities formed by mixed assemblages of species are located on the South of Península de Santa Elena, protected from the full impact of the seasonal upwelling in the region, and from strong wave action. Live coral cover ranged from 47.5 to 95.2% in 1993-94. Pocillopora damicornis predominated in the shallow sections, while Psammocora stellata predominated the deeper sections.

Bahía Culebra: The coral reefs and communities of this area (Fig. 2) were succinctly described by Glynn et al. (1983) and by Cortés & Murillo (1985), and in more detail by Jiménez (1997, 1998, 2001a), Jiménez et al. (2001), and Jiménez & Cortés (2003a). Coral reefs and communities have been variably disturbed by natural (El Niño) and anthropogenic impacts (siltation, divers, fishers). Several coral species that were found abundantly in Bahía Culebra in the early 1980’s, have become rare because of extraction for aquarium trade (Cortés & Murillo 1985). Bahía Culebra had rare and unique coral communities and reefs. One in particular, a Leptoseris papyracea reef suffered a high mortality associated to the 1997/98 El Niño (Jiménez et al. 2001), but it is recovering (Jiménez unpubl. data).

One of the reefs in Bahía Culebra has been monitored since 1994 (Fig. 5). Average live coral cover at three depths was generally higher before the warming and bleaching cycle of 1997/98 and didn’t change significantly in the following years. The impact of the warming event was also evident with increasing dead coral and macroalgae covers, with the only exception at 12m depth.

Península de Nicoya: This large section of the coast has coral communities, and a few coral reefs (Cortés & Murillo 1985). Most of the coast is exposed to strong wave action that may preclude the development of coral reefs. However, sheltered from the SW swells, there is an extensive reef, Matapalo (about 1.7km long) at the vicinities of Punta Gorda, 12km south from Bahía Culebra. Matapalo reef is dominated by branching coral species (Pocillopora spp.), which on average have a high live cover (>50%), similar to that of Bahía Culebra and Islas Murciélagos (Jiménez in prep.).

In another section of the coast, at Sámara, small Pocillopora, and Porites lobata patch reefs had been observed. Octocorals were abundant on rocky outcrops, and on islands and islets off Península de Nicoya. Patch reefs and coral communities have been observed between Cabo Blanco and the Islas Negritos (Fig. 2), in some areas growing over very extensive dead reefs. Octocorals are abundant on the Islas Negritos. Freshwater runoff in Golfo de Nicoya (Fig. 2) prevents the development of corals in its inner section of the gulf (Cortés & Jiménez 2003b). Islands in the middle section of the gulf, that many years ago had corals don't have any more because of the increase in freshwater due to water deviation projects in the watershed.

Pacífico Central: The central section of the Pacific coast of Costa Rica has few reefs (Cortés & Murillo 1985). Isolated corals and small coral communities can be found on rocky outcrops along the coast, including Parque Nacional Manuel Antonio (Fig. 2). Patch reefs of Pavona clavus have been observed to the South.

Rich coral communities have been observed and studied at Parque Nacional Marino Ballena (Alvarado 2004, 2006, Alvarado et al. 2005, 2009). The main reef-builder species was Porites lobata, with a high live coral cover (40%) in some sites, follow by Pavona clavus (Fig. 6). Since 1992, the coral communities have been monitored (Fig. 7), showing a decrease in live coral cover due mainly to the El Niño events during the 1990's (Jiménez & Cortés 2001, 2003b). After 2001, a slow recovery in coral cover has been observed but also a decrease or absence of species that are less resistant to sedimentation (Alvarado unpubl. data). The large fresh water lens between Parque Nacional Marino Ballena and Península de Osa from the Térraba-Sierpe mangrove complex (Fig. 2) inhibits the development of corals in that area. ]]>

Península de Osa: The second largest peninsula on the Pacific coast of Costa Rica is Península de Osa, on the Southeastern part of the country (Fig. 2). The coral communities and reefs on Península de Osa were described by Cortés & Jiménez (1996). A 250m2 Pocillopora patch-reef was studied at one site and it consisted of close to 100% live coral cover of Pocillopora damicornis in 1994. Exposed rocky substrates were covered by octocorals, and isolated corals were found on all rocky outcrops. The area has not been visited since the mid 1990's.

Golfo Dulce: This gulf is located on the Southern part of Costa Rica, and it is about 50km long by 10 to 15km wide, oriented NW to SE (Fig. 2). It is an enclosed embayment of tectonic origin, with a 60m deep sill and depths of 200m in the inner section (Hebbeln et al. 1996) that are anoxic. For this reason it is described as a tropical fjord (Hebbeln & Cortés 2001). Several types of coral reefs and coral communities have been found in the gulf, and they can be divided into two groups, the ones of the inner section of the gulf, and the ones from the outer section (Cortés 1990a, b). The inner Gulf reefs are characterized by live and dead Porites lobata on the reef front and dead Pocillopora damicornis and Psammocora stellata on the reef flat. Coral diversity is low and topographic relief high, with steep reef-fronts and sides. Live coral cover ranged from less than 1% to 8% in 1987. The outer Gulf reefs are characterized by a relatively high live coral coverage (range from 29 to 46%), high coral diversity, and low topographic relief (Cortés 1990b, Cortés & Guzman 1998).

The reef structure at Punta Islotes started growing 5 500 years ago over a basaltic substrate. Between 2 500 and 500 years ago reef growth was high, accumulation most of the structure. Around 500 years ago, due to tectonic activity, two rivers started flowing into the inner part of Golfo Dulce, retarding and in some areas stopping reef accretion. In the last 50 years, reef growth has ceased due to the increase in terrigenous sedimentation product of coastal and inland alteration (Cortés et al. 1994).

Between 1992 and 1996 live coral cover at Sándalo (outer reef) decreased from 29 to 17%, while at Punta Islotes (inner reef) it increased from 2 to 10%. Dead coral dominates the substrate. The reef at Sándalo was greatly affected by the sedimentation due to Hurricane Caesar in 1996 (Fonseca et al. 2006b). The structural balance of these reefs was negative, since bioerosion rates were higher, 0.71kg/m2.y1 in Punta Islotes and 2.61kg/m2.y in Sándalo, than construction rates, 0.41kg/m2.y in Punta Islotes and 0.56kg/m2.y in Sándalo, resulting in a net reef production of -0.3kg/m2.y in Punta Islotes and -2.05kg/m2.y in Sándalo. If prevailing conditions continue the reef of Punta Islotes will be completely destroyed in 6 400 years (Fonseca 1999).

Isla del Caño: This island is located 15km offshore from Península de Osa (Fig. 2), and the coral reefs were studied by Guzman (1986). The island had five coral reef flats, ranging in size from 0.8 to 4.2hectares. These fringing reef flats were mainly built by dead pocilloporid corals, covered by crustose coralline algae, isolated live colonies of pocilloporids and poritids, and microatolls of Porites lobata. The reef slope and base was dominated by the massive coral Porites lobata, which was the predominant species of the island. The shallow sections of the reef were structured mainly by physical factors: wave action, temperature and salinity fluctuations, and low tide exposure. While the deeper sections were controlled by biological interactions: bioerosion, damselfish algal lawns, and corallivores (Guzman 1988, Guzman & Cortés 1989). These reefs were impacted by the 1982-83 El Niño, with lost of up to 50% of the live coral coverage (Guzman et al. 1987) followed by phytoplankton blooms that killed shallow pocilloporid reefs (Guzman et al. 1990). Again, during the 1992 and the 1997-1998 El Niño there was extensive bleaching but lower mortality, less than 5%. Pulses of recruitment have been observed after the 1982-83, 1992 and 1997-98 El Niño events (Guzman & Cortés 2001).

Isla del Coco: This island is approximately 500km southwest of Costa Rica mainland (Cortés 2008). It had the highest number of zooxanthellate corals (17 spp.) on the Pacific of Costa Rica (Cortés & Guzman 1998). Isla del Coco also had the highest number of azooxanthellate corals, but more than half of this corals were collected in deep waters (Cairns 1991, Cortés unpubl. data). Fringing reefs ranging in size from less than one hectare to more than 50 hectares grew around Isla del Coco. Most of the reefs were constructed by Porites lobata (Bakus 1975, reported as Porites californica) but there were also extensive zones of agariciids: Pavona spp. and Gardineroseris planulata. Macintyre et al. (1992) found that the largest colonies of Porites lobata were between 250±50 and 430±80 years old, but it is probaly an underestimation of the age the bases of the large colonies bioeroded. Most of the coral died during the 1982-1983 El Niño, in 1987 live coral cover was between 2.6 and 3.5% (Guzman & Cortés 1992). The feeding and erosional activity of the sea urchin Diadema mexicanum was concentrated on surviving individuals of the 1982-83 El Niño.

In 2002, Isla del Coco was visited again. Reefs that previously had high densities of D. mexicanum were almost complete bioeroded, while reefs that in the 1980's had a high number of coral recruits and had high coral cover in 2002. Live coral cover of several species has increased to 23% (Fig. 8) and one species, Leptoseris scabra, not seen before, was present and very abundant in some reefs (Guzman & Cortés 2007).

Populations of corallivores were evaluated in the late 1980's and again in 2002. The density of A. planci did not exhibit a significant change between surveys while the density of Arothron meleagris increased significantly overall between 1987 and 2002. The overall average of D. mexicanum significantly decreased at all reefs, from 14.5 ind/ha in 1987 to 0.84 ind/ha in 2002 (Guzman & Cortés 2007) (Fig. 9).

Status of coral reef fishes

Pacific: In a study of the reef fishes of Bahía Culebra (Fig. 2), Dominici-Arosemena found 75 species in 28 families (Dominici-Arosemena 1999, Dominici-Arosemena et al. 2005). The most abundant species was Chromis atrilobata, followed by Thalassoma lucasanum, Abudefduf troschelii and Halichoeres dispilus in shallow waters, and Haemulon steichdachnederi, H. maculicauda and Stegastes flavilatus in deeper waters. Most of the species found showed a positive correlation in their abundance with live coral cover. But, there are signs of problems for the reef fish at Bahía Culebra. On the one hand, work by Jiménez (1998) indicates a progressive degradation of the reef areas between 1996 and 2005. And on the other hand, Dominici-Arosemena (1999) and Dominici-Arosemena et al. (2005) found that because of fish extraction, there is a reduction of terminal males of Thalassoma lucasanum, and some individuals with the terminal male coloration are relatively small. Another problem with this species is that the authorities consider the juveniles and the adults as separate species, wrongly assigning fishing quotas for each. Other species, Pomacanthus zonipectus and Holocanthus passer show signs of over exploitation. Catch has gone down and fishing is taking place farther and deeper than a few years ago. Other commercial species are also being overexploited.

Anthropogenic threats to coral reef bioversity caribbean

Siltation: The main impact on Caribbean coral reefs of Costa Rica is increased terrigenous sediment loads (Risk et al. 1980, Cortés 1981, 1992, 1994, Cortés & Risk 1984, 1985, Cortés et al. 1998). At Cahuita, live coral coverage is low and average colony size is large (Fonseca 2003). The amounts of suspended and resuspended sediments are very high, as well as sediments trapped in the massive coral skeletons. Coral growth rates are low, and are significantly inversely correlated with sediment resuspension rates. The increase in terrigenous sediments in this reef is due to deforestation of nearby watersheds (Cortés 1981, Cortés & Risk 1984, 1985). Terrigenous sediment concentrations had continued to increased and live coral coverage to drop (Cortés 1994, Fonseca et al. 2006a). This, together with other anthropogenic disturbances, e.g. increased tourism and pollution, and natural impacts (but probably exacerbated by human activity) such as bleaching and massive death of organisms, have degraded the reef at Cahuita even more (Cortés 1994, Fonseca & Nielsen unpubl. data). In 2004 the inner reef crest was studied again, and it was found that live coral has increased at the same time that siltation rates have increased slightly. The rise in both sediment levels and live coral coverage over the last ten years may indicate that other factors besides sedimentation, e.g. D. antillarum die-offs and the 1991 earthquake, may have had a bigger impact than previously thought in the drop in coral cover from 1980 to 1993 (Wyckoff & Cortés unpubl. data).

Pollution: The coastal zone between Portete and Limón (Fig. 1) are polluted with fecal wastes and garbage, mainly from banana plantations and coastal towns. Analysis between 1981 and 1985 of petroleum pollution indicated that the levels were relatively low, with sporadic events of high concentrations. The highest concentrations were found in the port areas (Mata et al. 1987). In a study done between 2000 and 2002 in the Limón area, the level of petroleum pollution was still low (Acuña-González et al. 2004). High levels of heavy metal pollution in coral skeletons and sediments from reefs, even in "pristine" reefs were found by Guzman & Jiménez (1992). The sources of the heavy metals may be natural, but mainly they are of anthropogenic origin, including sewage discharges, oil spills (from refineries and tankers), the misuse of agricultural chemicals and fertilizers, and topsoil erosion (Guzman & García 2002). García-Céspedes et al. (2004) studying trace metals in several coastal areas of Costa Rica, found highest levels of lead in Limón, and intermediate levels of iron, zinc, and copper. Concentrations of PBCs are low in the Limón area (Spongberg 2004).

Impact of tourism: Tourist visitation to the Caribbean coast of Costa Rica, as well as coastal development has increased enormously in the last two decades. This activity has had an impact on the coral reefs of the region, due to trampling of corals and other organisms on the lagoon, and kicking of corals during diving (Cortés 1994, Gove & Cortés unpubl. data).

Siltation: Terrigenous sediments have affected several reefs on the Pacific coast (Cortés & Murillo 1985), at Bahía Culebra (Jiménez 1998), at Parque Nacional Marino Ballena (Alvarado 2006) and at Golfo Dulce (Cortés 1990a, b, 1991, Cortés et al. 1994, Fonseca 1999, Fonseca et al. 2006b). Sedimentation has increased because of deforestation of the watersheds, road and tourist complexes constructions, and inappropriate agricultural practices (Cortés 1990a, b, Jiménez 1998, Alvarado 2004, 2006). A significant reduction in reef accretion can be observed in reef cores from Golfo Dulce (Fig. 2), and it has been attributed to recent terrigenous sedimentation (last 50 years) as the area was deforested for banana plantation (Cortés et al. 1994, Fonseca 1999).

Extraction of reef organisms: Some species of corals had been extracted for the local curio trade. The populations of Pocillopora eydouxi and of P. meandrina, both very attractive branching coral have been reduced because of exploitation (Glynn 1979, Cortés & Murillo 1985). Extraction of reef fish has severely impacted the populations of some regions (Dominici-Arosemena 1999, Dominici-Arosemena et al. 2005).

Tourism: Coral reefs on the Pacific side of Costa Rica have become important for diving tourism. Touristic activity has direct and indirect effects on the coral reefs (Cortés & Murillo 1985, Jiménez 1997, 2001a). Direct effects included: coral extraction and damage to colonies by divers and boat anchors. Indirect effects included: over-exploitation of resources (fish, lobster, mollusks) increased sewage and increased sediment loads from coastal alteration (urbanization, road construction).

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