Coral reefs of Guatemala

Coral reefs of Guatemala

Ana C. Fonseca E.1 and Alejandro Arrivillaga2

1Centro de Investigación en Ciencias del Mar y Limnología (CIMAR),

Universidad de Costa Rica. San Pedro, San José 2060, Costa Rica 2Johnson Controls World Services. USGS National Wetlands Research Center,

700 Cajundome Blvd. Lafayette, Louisiana 70506, USA

ABSTRACT: Coral reef research in Guatemala has a very short history. Hard-bottom biotopes are sparse along both Guatemala’s coasts; however, there is no evidence of coral reefs presence on its Pa-cific coast. Reef development exists on the Caribbean coast, mainly around Punta de Manabique, and most reefs are protected by the Refugio de Vida Silvestre de Punta de Manabique (Punta de Manabique Wildlife Refuge). This is the sole marine park in Guatemala. Coral reefs at Punta de Manabique consist of a series of continental carbonated banks. Currently they are highly deteriorated, live coral cover is low (8.75%) and non-coralline macroalgae cover is high (65%). There are 29 species of scleractinian corals. The main impact to coral reefs at Punta de Manabique is terrestrial sedimentation coming from deforested lands, mainly from the Motagua river basin. High sediment loads are causing coral death and algae overgrowth. Apparently, current herbivorous populations are not able to control the macroalgae. Coral composition is atypical to most Caribbean reefs, which are dominated by Montastraea annularis: Guatemalan reefs are dominated by macroalgae and coral species resistant to sediments, such as Siderastrea siderea. In addition, natural events such as hurricanes, temperature increases and the massive mortality of Diadema in 1983, may have contributed to the deterioration of Guatemalan reefs. Coral reefs at Punta de Manabique constitute one of the most important resources of the coastal-marine territory of Guatemala. The economic activity in Punta de Manabique is focused on artisanal fisheries around these reefs and developing tourism. Promotion of environmental education and research, reinforcement of fishing and tourism regulations, and an adequate management plan for the Refuge and the regional river basins are recommended.

1. INTRODUCTION

Guatemala is located in the northern part of Central America and has borders with

Mexico to the north and west, and Belize, Honduras, and El Salvador to the east (Fig. 1). Its territory occupies 108,780 km2. The total littoral length is 405 km (Foer and Olsen 1992). The largest coastal zone is located on the Pacific side (255 km), in the southern portion of the country, and it consists of black volcanic sand beaches. Several small-to-medium sized rivers discharge on this side of the country and there are a series of coastal lagoons and mangrove forests.

The Caribbean coast of Guatemala extends for 150 km and is located on the Gulf of Honduras; despite its small size, it supports important commercial and artisanal fisheries.

Latin American Coral Reefs, Edited by Jorge Cortés © 2003 Elsevier Science B.V. All rights reserved.

A.C. Fonseca & A. Arrivillaga 160

Belize

Guatemala

Honduras

AmatiqueBay

Gulf ofHonduras

La GraciosaBay

Canal

Ingles

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Dulce

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rSarstoon River

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85.75ºW 88.50ºW 88.75ºW

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Fig. 1. Location of Guatemala (inset) and the maps of the country’s Atlantic coast with main rivers and geo-graphic features identified.

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La Guaira

Guinea Bank

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Chatarra Manglar Bank

16.0 N, 88.9 W

15.7 N, 88.7 Wo o

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N0 5 10 15 Kilometers

Fig. 2. Location of known reefs and hard bottom biotopes on the Atlantic coast of Guatemala.

Coral reefs of Guatemala 161

Punta de Manabique occupies a large section of this coast which is generally low with forested areas overlooking the coast, dissected with rivers. Most sea beds bottoms at the rivers mouths and bays consist of silty sands (PROARCA 1996). Three main rivers discharge their water into the Caribbean: Sarstoon, Motagua and Dulce (Fig. 1). The primary freshwater influence in the region is from Río Motagua located on the eastern side of Punta de Manabique (Gulf of Honduras). An artificial channel, Canal de los Ingleses, connects Graciosa Bay to the eastern shore of Punta de Manabique (Yáñez-Arancibia et al. 1999).

1.1. Reef research history

Coral-reef research in Guatemala is scarce and recent. Coral reefs of the Caribbean coast have been incidentally mentioned in studies conducted with other objectives in mind. One of the earliest written reports was that of Bortone et al. (1988), who studied fish communities in an artificial reef established in 1982 in the Amatique Bay, about 15 km NW of Puerto Barrios (Fig. 1). Similarly, the presence of reef formations was indicated by Cazali (1988) in her study of bivalves and Prado (1990) in her study of gastropods.

Near-shore coral reef and hard-bottom biotopes are sparse along both Guatemala’s coasts, but there is reef development on the Caribbean coast of Guatemala, mainly around Punta de Manabique (Fonseca 2000) and on Heredia Shoal, 15 km west of Punta Moreno (Cazali 1988). Hard bottoms and coralline algae are present at Punta Cocoly, Punta Herreria and La Guaira, and on the littoral from Punta Herreria to Tapon Creek (Cazali 1988). Other rocky and reef beds are present near the Sarstoon River mouth and at Palo Blanco Shoal. Other reef formations know to local fishermen are Chatarra, Hamilton and Satuye (Fig. 2). There is no evidence of the existence of coral reefs on the Pacific coast maybe only isolated corals on the few hard bottoms (Cortés and Hatziolos 1998; Kramer et al. 2000).

Fonseca (2000) made a rapid reef assessment on those Caribbean carbonated banks known by fishermen around Punta de Manabique in order to describe reef structure and the composition of benthic organisms. The Point Intercept Transects Method was used to determine the relative substrate cover; using a 10 m long linear transect and recording what was located at each point every 50 cm. Chain transects were used to calculate the spatial relief (Rogers et al. 1994). A collection of corals, other invertebrates, algae and seagrass samples were deposited at the Natural History Museum of the University of Guatemala. There were no previous descriptions or evaluations of these reefs. The results of this report (Fonseca 2000) are published in this chapter. Further research must be undertaken to determine the history of formation, extension, and diversity of these reefs.

Other coastal ecosystems have been studied, including mud flats (Salaverria and Rosales 1993) and seagrass beds (Arrivillaga and Baltz 1999; Yáñez-Arancibia et al. 1999; Arrivillaga 2000). Specifically, seagrass beds of La Graciosa Bay in Punta de Manabique have been the object of recent extensive studies (Arrivillaga 2000). In addition, the utilization of seagrass meadows by fishes and decapod crustaceans has been described for exposed (beach) and protected (bay) estuaries near the mouth and inside of La Graciosa Bay (Arrivillaga and Baltz 1999; Arrivillaga 2000). 1.2. Description of Punta de Manabique shoreline

Punta de Manabique is located southwest of the Gulf of Honduras on the northeast side of Amatique Bay (Fig. 1). This peninsula is 23 km long, and is separated from the


continent at its base by the Canal de los Ingleses. The coast of Punta de Manabique con-sists of a series of sandy bars and peat swamps. Sandy bars are old carbonated banks, and the swampy sections between these banks have been filled with sand and organic matter. Rain forest is the dominant vegetation (Centro de Estudios Conservacionistas 1995).

The northeastern section of Punta de Manabique, facing the Gulf of Honduras, has on the windward side narrow, fine gray sandy beaches exposed to high surges for most of the year. There is a large accumulation of trunks, branches, Sargassum, Thalassia, Syringodium, and garbage. These materials mainly come from the mouth of the Motagua river transported by the northwest current. The southwest section facing Amatique Bay, consists of leeward narrow, and fine gray sandy beaches with low surge.

On the southeast, Punta de Manabique has sandy beaches, with good nesting sites for the marine turtles Dermochelys coriaceae, Eretmochelys imbricata, Caretta caretta and Chelonia mydas. Graciosa Bay is a shallow coastal lagoon located on the east side of Amatique Bay. Graciosa Bay is surrounded by red mangrove forests although large sections of these mangroves were destroyed by Hurricane Mitch in 1998. It also has extensive seagrass meadows of Thalassia testudinum, Syringodium filiforme and Halodule beaudettei (Arrivillaga 2000). At La Graciosa and Amatique bays, the manatee Trichechus manatus, an endangered species worldwide, is frequently observed (Centro de Estudios Conservacionistas 1995; Fonseca 2000).

2. CORAL REEFS OF PUNTA DE MANABIQUE

2.1. Reef descriptions

The reefs at Punta de Manabique are low carbonated banks (Fig. 3), with moderate coral richness, low coral cover, high noncoralline algae cover (Fig. 4), and small coral colonies (diameter and height < 1 m). A list of species found in these reefs, 29 corals, 44 other invertebrates, 24 algae and 4 seagrasses is given in Tables 1, 2 and 3. Visibility is low (7 m) and the mean temperature is 29ºC. A general description of four of the main banks (Graciosa, Manglar, Guinea, and Cabo Tres Puntas banks) (Fig. 2) is presented below.

Graciosa bank is located 3 km west from La Graciosa Bay mouth (Fig. 1). It is 6 to 8 m deep, with an area of 25 x 50 m, and a low complexity index (i=1.37 ± 0.2; Fig. 3). Mean live coral cover is 10.1 ± 4.6%, and algae cover is 73.7 ± 7.6% (Fig. 4). Mean density of corals is 4.0 ± 1.0 colonies m-2 (Fig. 5) and 8 species were found. The most common coral species were Siderastrea siderea, Porites astreoides and Montastraea cavernosa. Sus-pended sediment concentrations are high and coral colonies are covered by layers of mucous. The sediment seen at this bank may come from the Canal de los Ingleses.

Manglar bank is located southwest of Punta de Manabique and 1.5 km of shore (Fig. 2). It is 14 to 17 m deep, has an area of 25 x 25 m, and has a low complexity index (i=1.25 ± 0.1; Fig. 3). Mean live coral cover is 13.6 ± 6.4%, and algae cover is 33.4 ± 4.3% (Fig. 4). Mean coral density is 4.0 ± 1.4 colonies m-2 (Fig. 5), and 8 species were found. The most common coral species were Siderastrea siderea, Siderastrea radians, Porites astreoides, Montastraea cavernosa, Madracis decactis, Stephanocoenia michelinii, Agaricia tenuifolia, Meandrina meandrites and Helioseris cucullata.

Guinea bank is composed of many reef patches of low relief (i=1.2 ± 0.0; Fig. 3), with a mean depth of 24 m and located northeast of Guinea creek in San Francisco del Mar (Fig. 2). Mean live coral cover is 1.5 ± 2.1%, and algae cover is 81.8 ± 8.6% (Fig. 4). Mean


1.1

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Fig. 3. Complexity index (i) by site, Punta de Manabique, Guatemala.

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Algae Sponges Mud Other sediments Otherinvertebrates

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Fig. 4. Substrate cover by site, Punta de Manabique, Guatemala.

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loni

es m

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Fig. 5. Coral colonies density by site, Punta de Manabique, Guatemala.


TABLE 1 Stony corals (Class Hydrozoa and Anthozoa) found in reefs around Punta de Manabique.

Millepora complanata Montastraea annularis Mycetophyllia aliciae Porites astreoides Montastraea franksi Mycetophyllia danaana Porites porites forma divaricata Montastraea faveolata Scolymia cubensis Siderastrea siderea Montastraea cavernosa Mussa angulosa Siderastrea radians Solenastrea hyades Eusmilia fastigiata Leptoseris cucullata Diploria strigosa Madracis decactis Agaricia tenuifolia Meandrina meandrites Madracis mirabilis Agaricia agaricites Manicina areolata Stephanocoenia michelinii Agaricia grahamae Colpophyllia natans Phyllangia americana Oculina difusa Mycetophyllia ferox

TABLE 2 Other marine invertebrates of Punta de Manabique (*Included in CITES list).

Phylum Porifera Phylum Ctenophora Class Demospongiae Class Tentaculata

Callyspongia plicifera Mnemiopsis mccradyi Callyspongia vaginalis Phylum Annelida Cibrochalina vasculum Class Polychaeta Verongula gigantea Bispira brunnea Xetospongia muta Bispira variegata Niphates erecta Phylum Molusca Niphates digitalis Class Gastropoda Amphimedon compressa Strombus raninus Cinachyra sp. Cassis madagascariensis Cliona delitrix Class Bivalvia

Phylum Cnidaria Spondylus americanus Class Hydrozoa Pinna carnea

Order Hydroida Phylum Arthropoda Dentitheca dentritica Class Crustacea

Order Stylasterina Order Decapoda Stylaster roseus * Periclimenes pedersoni

Class Anthozoa Stenorhynchus seticornis Subclass Octocorallia Phylum Ectoprocta

Order Gorgonacea Class Gymnolaemata Erythropodium caribaeorum Order Cheilostomata Pseudoplexaura sp. Trematooecia aviculifera Eunicea sp. Phylum Echinodermata Plexaurella sp. Class Asteroidea Muricea ¿pendula? Oreaster reticulatus Pseudopterogorgia sp. Class Holothuroidea

Pterogorgia guadalupensis Actinopygia agassizii Gorgonia ventalina Eostichopus amesoni

Order Alcyonacea Holothuria mexicana Carijoa riisei Class Echinoidea

Subclass Hexacorallia Lytechinus variegatus Order Antipatharia Diadema antillarum

Cirrhipathes (Stichopathes) leutkeni Echinometra virides Order Zoanthidea Eucidaris tribuloides

Palythoa caribaeorum Phylum Chordata Order Actinaria Class Ascidiacea

Condylactis gigantea Ascidia sydneiensis Bartholomea annulata


TABLE 3 Marine algae and seagrasses of Punta de Manabique.

MARINE ALGAE Valonia utricularis Phylum Phaeophyta Udotea sp.

Sargassum fluitans Acetabularia sp. Sargassum sp. Phylum Rhodophyta Dictyota ciliolata Galaxaura sp. Dictyota sp. Jania adherens Lobophora variegata Amphiroa tribulus Turbinaria turbinata Porolithon pachydermum

Phylum Chlorophyta Peyssonnelia sp. Halimeda incrassata Phylum Cyanophyta Halimeda discoidea Schizotrix sp. Penicillus pyriformis Penicillus dumetosus SEAGRASSES Caulerpa sertularoides Class Angiosperma Caulerpa racemosa Thalassia testudinum Caulerpa mexicana Syringodium filiforme Codium isthmocladum Halodule beaudettei Ventricaria ventricosa Halophila baillonis

density of corals is 0.5 ± 0.7 colonies m-2 (Fig. 5), and 9 species were found. The most common coral species were Siderastrea siderea, S. radians, Porites astreoides, Montastraea cavernosa, Madracis decactis, Stephanocoenia michelinii and Leptoseris cucullata. Several vase sponges (Niphates digitalis and Callyspongia plicifera) have been found in this site.

Northeast of Cabo Tres Puntas (“Three Point Cape”), between the second and the third points, 1 to 5 km from the coast, a number of patch reefs can be found, which are well known to fishermen (Fig. 2). The bottom is sandy except for the nearest section to the first point where there is a large accumulation of mud, apparently coming from the Motagua River and other small effluents such as the San Francisco and Canal de los Ingleses. At 9 to 12 m depths, there are dense octocoral gardens (11.1 octocorals m-2). From 12 to 18 m deep, there are several reef patches of low relief (i=1.19 ± 0.2; Fig. 3), of around 15 x 15 m, which are separated from 50 to 100 m. There are many fleshy macroalgae attached to the substrate over the reef patches and floating over the surrounding sandy plains. Some algae reach heights of up to 1 m. Mean live coral cover is 9.1 ± 3.0%, and algae cover is 55.6 ± 21.5% (Fig. 4). Mean density of corals is 2.0 ± 1.0 colonies m-2 (Fig. 5), and 8 species were found. The most common coral species were Montastraea faveolata, M. franksi, Siderastrea siderea, Stephanocoenia michelinii, Agaricia agaricites, Agaricia tenuifolia, Porites astreoides and Eusmilia fastigiata.

Offshore from the sandy bank where these reef patches are located, there is a deep channel parallel to the shoreline and is used by large ships. Behind this channel, 12 km offshore, there is another sandy bank with reef patches that have never been evaluated.

The substrate is mostly covered by non-coralline macroalgae, especially on Guinea and Graciosa banks (Fig. 3). Macroalgae on these banks are known to be harmful to corals. The most common species are: Caulerpa sp. and Dictyota sp., Lobophora variegata, Ventricaria ventricosa, Valonia utricularis, Codium isthmocladium, Sargassum sp. and the blue-green algae Schizotrix sp. Frequently encountered coral species on these banks


are Siderastrea radians, S. siderea, Madracis decactis, Montastraea cavernosa, Stephanocoenia michelini and Porites astreoides. This is not the typical coral composition of most Caribbean reefs, which are usually dominated by Montastraea annularis (CARICOMP Data Base 1999). Particularly, Siderastrea siderea, Siderastrea radians and Porites astreoides are known to be very resistant to sediments and are relatively abundant in sites with a high sedimentation (Cortés and Risk 1985). The density of coral colonies is higher on the southwestern banks of Punta de Manabique (La Graciosa and Manglar) than on the northeastern banks (Guinea and Cabo Tres Puntas) (Fig. 5). The sea urchin Diadema has not been found (Fonseca 2000).

2.2. Present condition

Reef banks within Punta de Manabique show fleshy macroalgae typical of sandy plains where herbivory is low. Apparently, the herbivorous are not able to control macroalgae abundance as on other Caribbean reefs (Hay 1984). The Manglar bank could be considered the area in best condition, as it shows the highest levels of live coral and sponge cover (Fig. 4). However, the Cabo Tres Puntas bank has the largest reef area and coral richness (Fonseca 2000).

Reefs in Guatemala have degraded mainly as the result of siltation stress. Since infor-mation on these reefs is scarce, it is hard to determine changes that have occurred over time. However, live coral cover is similar to what is found on other reefs on the Central American Caribbean coast that are also affected by terrestrial sediments (e.g. 13% in Cahuita, Costa Rica) and Belize (10-16%) (CARICOMP Data Base 1999). In addition, as on other Caribbean reefs, algae cover is greater than coral cover (CARICOMP Data Base 1999).

3. NATURAL DISTURBANCES

Some natural events that have affected the entire Caribbean region could have con-tributed to the deterioration of Guatemalan reefs. Several hurricanes that affected the Guatemalan coast (e.g. hurricanes between 1945 and 1949, Hurricane Fifi in 1972, and Hurricane Mitch in 1998) could have caused some reef destruction. Local fishermen and diving instructors have observed several bleaching events in 1983 and 1998, possibly due to temperature increases, and the Diadema massive mortality (1983) that resulted in widespread non-coralline algae overgrowth (Hughes et al. 1987). There was also an earth-quake on February 4, 1976 that extended east and west from the Motagua fault (Young et al. 1989). The effects of this event on the coral reefs of Punta de Manabique were not quantified, but reefs could have been affected by sediment increase and substrate fracture, as observed in Costa Rica (Cortés et al. 1992).

4. ANTHROPOGENIC IMPACTS

It could be inferred from high-water turbidity that the biggest problem at Punta de

Manabique is the chronic influx of terrestrial sediments coming from upstream rivers. This is a major long-term threat for reefs at Punta de Manabique and for continental reefs elsewhere (Wilkinson 1992). Sediments are transported by the main current that flows northwest, causing corals death and algae overgrowth. On the northeastern section


of Punta de Manabique, the influence of the Motagua river, and other small effluents, which flow over deforested and eroded watersheds is evident. Amatique Bay also receives terrestrial sediments from the Dulce, Machacas and Sarstoon rivers and Canal de los Ingleses (Centro de Estudios Conservacionistas 1995).

Human population is relatively low (850 inhabitants) on Punta de Manabique. A narrow coastal band has been used mostly for small house construction, subsistence agriculture, and cattle ranching. Artisanal fishing is the main economic activity, primarily along the eastern section of Amatique Bay and the Gulf of Honduras (Echeverría de León 1994). Fishermen like to fish and dive over coralline banks where fishes, lobsters (Panulirus spp.) and snails (Strombus spp.) are concentrated. Some of the common fishing tech-niques (e.g. netting) are not recommended since they are neither size nor species selective. Capture levels of commercially important species from Punta de Manabique are unknown. Moreover, inhabitants from Punta de Manabique prefer to dive and extract marine resources from nearby Belizean Cays. Turtle nesting is common on the beaches of Punta de Manabique; but, eggs extraction is occasional and for domestic consumption (Centro de Estudios Conservacionistas 1995).

Tourist visitation to Punta de Manabique is sporadic, and although it has increased in the last decade and the region has a high potential for greater increase (Centro de Estudios Conservacionistas 1995). Recently, a dive store was established at Amatique Bay, and it is promoting tourism. This could be positive if a good regulation plan is implemented.

5. PROTECTION AND MANAGEMENT

Reef banks on the Caribbean coast of Guatemala are within the Punta de Manabique

Wildlife Refuge (15º40’-16º00’N; 88º13’- 88º44’W). This refuge protects an area of 1393 km2 (449 terrestrial and 944 marine). It comprises the peninsula, 27 km east to the Motagua river, 12 km offshore in the Honduras Gulf, and the Amatique and Graciosa Bays (Centro de Estudios Conservacionistas 1995). This is the only marine park in Guatemala. However, for protection a management plan for the refuge and regional river basins must be implemented. Particularly, fishing and tourism must be regulated. Guardianship at the refuge needs to be reinforced and the environmental consciousness of the population should be promoted.

Reefs from the Caribbean coast of Guatemala are within the Mesoamerican Reef System (“Sistema Arrecifal Mesoamericano”, SAM) which is considered a priority for conservation worldwide. This reef system extends from the northern end of the Yucatán Peninsula in Mexico, to the Bay Islands in Honduras, including the Barrier Reef of Belize and the Caribbean coast of Guatemala. It has been recognized by national governments at the meeting of Tulum in 1997, where they committed their countries to support its management and conservation (Marín 2000).

From the species found on Punta de Manabique reefs (Tables 1, 2 and 3), all hard corals, the lace coral Stylaster roseus, and the fire coral Millepora alcicornis are included in the second category of the CITES list. This means that these species cannot be extracted from their natural environments. At present they are not in the Red List of the IUCN.

Today, the most pressing research need for Guatemala’s coastal ecosystems is to de-termine the distribution, abundance, and status of the main coral species. The location,


composition and extent of these coastal ecosystems may be a crucial indicator of water quality and overall health of coastal resources. Unless these resources are identified and research is begun on their ecosystems, there will be few objective criteria for the for-mulation of management policies to foster scientifically based resource conservation.

ACKNOWLEDGMENTS

Mario Yon, Luisa Paredes and Eddy helped during the exploration to the reefs of Punta

de Manabique. PANADIVERS store provided logistical support and valuable information. Mario Dary Foundation (FUNDARY) and The Nature Conservancy (TNC) provided coordination and funding.

REFERENCES Arrivillaga, A. 2000. Ecology of seagrass fishes and macroinvertebrates on Guatemala’s

Atlantic coast. Ph.D. dissert., Louisiana State Univ., Baton Rouge, Louisiana. 163 p. Arrivillaga, A. & D.M. Baltz. 1999. Comparison of fishes and macroinvertebrates on sea-

grass and bare-sand sites on Guatemala’s Atlantic coast. Bull. Mar. Sci. 65: 301-319. Bortone, S.A., R.L. Shipp, W.P. Davis & R.D. Nester. 1988. Artificial reef

development along the Atlantic coast of Guatemala. Northeast Gulf Sci. 10: 45-48. Cazali, G.M. 1988. Inventario de pelecípodos de la costa Atlántica de Guatemala con

enfasis en especies comestibles. Thesis, Universidad de San Carlos, Ciudad de Guatemala. 134 p.

Centro de Estudios Conservacionistas. 1995. Estudio técnico del área de protección es-pecial “Punta de Manabique”. Propuesto Biotopo Protegido. Facultad de Ciencias Químicas y Farmacia, Universidad de San Carlos de Guatemala, Ciudad de Guate-mala. 82 p.

Cortés, J. & M.E. Hatziolos. 1998. Status of coral reefs of Central America: Pacific and Caribbean coasts. 32-37. In: C. Wilkinson (ed.), Status of Coral Reefs of the World: 1998. GCRMN, Australian Institute of Marine Science.

Cortés, J. & M.J. Risk. 1985. A reef under siltation stress: Cahuita, Costa Rica. Bull. Mar. Sci. 36: 339-356.

Cortés, J., R. Soto, C. Jiménez & A. Astorga. 1992. Earthquake associated mortality of intertidal and coral reef organisms (Caribbean of Costa Rica). Proc. 7th Int. Coral Reef Symp., Guam 1: 235-240.

Echeverría de León, A.G. 1994. Caracterización de la actividad pesquera artesanal en la Península de Manabique, Puerto Barrios, Izabal. Facultad de Agronomía, Instituto de Investigaciones Agronómicas, Universidad de San Carlos de Guatemala, Ciudad de Guatemala. 89 p.

Foer, G. & S. Olsen. 1992. Las costas de Centro América: diagnósticos y agenda para la acción. ROCAP-USAID-University of Rhode Island. 290 p.

Fonseca E., A.C. 2000. Evaluación ecológica rápida de los arrecifes coralinos de Punta de Manabique, costa Caribe de Guatemala. Report for The Nature Conservancy (TNC), Washington D.C. 23 p.

Hay, M.E. 1984. Patterns of fish and urchin grazing on Caribbean coral reefs; are previous results typical? Ecology 65: 446-454.


Hughes, T.P., D.C. Reed & M.-J. Boyle. 1987. Herbivory on coral reefs: community structure following mass mortalities of sea urchins. J. Exp. Mar. Biol. Ecol. 113: 39-59.

Kramer, P., P.R. Kramer, E. Arias-González & M. McField. 2000. Status of coral reefs of northern Central America: México, Belize, Guatemala, Honduras, Nicaragua and El Salvador. 287-313. In: C. Wilkinson (ed.), Status of Coral Reefs of the World: 2000. GCRMN and Australian Institute of Marine Science.

Marín, S. 2000. Unidos por un tesoro mundial en Mesoamérica. WWF, Centroamérica, 3 (2): 7-12.

Prado, L.M. 1990. Colecta, clasificación y distribución de las especies de gasterópodos en la costa Atlántica de Guatemala. Thesis, Universidad de San Carlos, Ciudad de Guatemala. 120 p.

PROARCA 1996. Gulf of Honduras: preliminary site overview. PROARCA/Costas, TNC, WWF and University of Rhode Island. 22 p.

Rogers, C.S., G. Garrison, R. Grober, Z.M. Hillis & M.A. Franke. 1994. Coral reef monitoring manual for the Caribbean and Western Atlantic. Natl. Park Serv., Virgin Islands.

Salaverria, A. & F. Rosales. 1993. Ecología pesquera de la costa Atlántica de Guatemala: evaluación inicial, Bahía de Amatique, Izabal. Informe de Avance. Universidad de San Carlos, Ciudad de Guatemala. 105 p.

Wilkinson, C.R. 1992. Coral reefs of the world are facing widespread devastation: can we prevent this through sustainable management practices? Proc. 7th Int. Coral Reef Symp., Guam 1: 11-21.

Yáñez-Arancibia, A., D.J Zarate-Lomeli, M. Gomez Cruz, R. Godinez Orantes & V. Santiago Fandino. 1999. The ecosystem framework for planning and management the Atlantic coast of Guatemala. Ocean Coastal Manag. 42: 283-317.

Young, C.J., T. Lay & C.S. Lynnes. 1989. Rupture of the 4 February 1976 Guatemalan earthquake. Bull. Seism. Soc. Amer. 79: 670-689.

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Coral reefs of Guatemala